Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Insular patterns of calicioid lichens in a boreal old-growth forest-wetland mosaic

Fragmentation of the forested landscape poses a threat to many aspects of biodiversity associated with old-growth forests Studies of the effects of forest fragmentation are often complicated by the variation in composition and age of patches and the matrix This study used a system of isolated stands where patch age and composition were similar and the matrix variability negligible The patches were composed of old-growth Picea abies stands of varying size and shape in a wetland matrix The study organisms were epiphytic crustose calicioid lichens (also known as Caliciales), many of which are very substrate-specific and restricted to old-growth stands The aim of the study was to measure the effect of patch size, patch isolation, habitat and substrate quality on the species riochness and composition of epiphytic calicioids Twenty-four patches ranging from 0 4 to 15 9 ha in size were studied All species of calicioid lichens were registered in 0 1 ha plots in each patch Isolation was measured as the percentage of available habitat within 400 m of a patch Twenty-two species were found with an average of 9 48 ± 0 26 (SE) species per patch and 292 ± 0 18 (SE) species per tree Species richness at patch level correlated with stand structure, primarily tree density, while number of species per tree (reflecting population size) was strongly correlated with island size and several stand variables There was no effect of isolation on species richness Species composition was influenced by both substrate variables and patch size The species composition on the islands showed a significant nestedness, i e species composition on species-poor islands constituted a non-random subset of the species composition on species-rich islands We propose that the explanation for the strong relationship between species richness at tree level and stand size is an edge effect which implies that unaffected interior areas only occur on large islands The different microclimate of the patch edge enables only the hardiest species to establish large populations there whilst shade and moisture demanding species are restricted to the interiors of larger islands     

Viability of ecological processes in small Afromontane forest patches in South Africa

Abstract The conservation of biodiversity is dependent on protecting ecosystem‐level processes. We investigated the effects of fragment size and habitat edge on the relative functioning of three ecological processes – decomposition, predation and regeneration of trees – in small Afromontane forests in KwaZulu‐Natal, South Africa. Ten sampling stations were placed in each of four forest categories: the interior of three large indigenous forest fragments (100 m from the edge), the edges of these large fragments, 10 small indigenous fragments (<1 ha) and 10 small exotic woodlands (<0.5 ha). Fragment size and edge effects did not affect the abundance of the amphipod Talitriator africana, a litter decomposer, and overall dung beetle abundance and species richness significantly. Bird egg predation was marginally greater at large patch edges compared with the other forest categories, while seed predation did not differ among forest categories. Tree seedling assemblage composition did not differ significantly among large patch interiors and edges, and small indigenous fragments. Sapling and canopy assemblage composition each differed significantly among these three indigenous forest categories. Thus, while tree recruitment was not negatively affected by patch size or distance from the edge, conditions in small fragments and at edges appear to affect the composition of advanced tree regeneration. These ecological processes in Afromontane forests appear to be resilient to fragmentation effects. We speculate that this is because the organisms in these forests have evolved under fragmented conditions. Repeated extreme changes in climate and vegetation over the Pleistocene have acted as significant distribution and ecological extinction filters on these southern hemisphere forest biota, resulting in fauna and flora that are potentially resilient to contemporary fragmentation effects. We argue that because small patches and habitat edges appear to be ecologically viable they should be included in future conservation decisions.     

Structures determining bryophyte species richness in a managed forest landscape in boreo-nemoral Europe

Forest patches with high biological value are protected as woodland key habitats (WKH), which are identified by the presence of forest structures and indicator species. However, management for conservation needs to consider also managed forests as habitats for species. In this respect, there is a need to set quantitative targets for species and structures at different landscape scales. Due to non-intensive methods of forest management used prior to 1940 in Latvia, it might be expected that large areas of forest have developed structures that can support many species characteristic of natural forests. The aim of the study was to create a model that best described the richness of bryophyte species that are characteristic of natural forests, using forest structures as explanatory factors. The structures and bryophyte communities on living trees and coarse woody debris (CWD) were described in plots along transects blindly placed in areas dominated by State forests under commercial management. Explanatory variables related to tree species composition and tree size explained 54% of the variation in WKH indicator species richness on living trees. The best explanatory factors were maximum diameter of deciduous tree species and CWD. Low richness of total bryophyte and indicator species was found on dead wood, and the amount of variation in bryophyte species richness on CWD explained by explanatory variables was low. The study indicates the importance of deciduous tree substrate in managed forests in maintaining the spatial continuity of epiphytic species diversity. However, the forests in the managed forest landscape did not support high diversity of epixylic species, even in the WKHs, due to low diversity of suitable dead wood substrate.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

Long-term changes in bryophyte diversity of central European managed forests depending on site environmental features

Cryptogamic diversity is a reliable indicator of the state of forest ecosystems. In this study we analysed the variations in both bryophyte species richness overall and number of hemerophobic bryophyte species in Central European managed forests over a 20-year time span, based on data collected in 132 plots scattered across Poland. We tested differences in species richness among five temporal replicates, as well as among site types grouped based on elevation a.s.l., dominant tree species and stand age. The analyses revealed no significant trend in species richness across years. Meanwhile, species richness significantly increased along with elevation a.s.l., especially in broadleaved forests. No significant difference in species number between spruce and pine dominated stands emerged for mature stands, while there was a strong difference for young stands, with spruce forest hosting a much higher number of species. Species richness exhibited a slight, but not significant, increase over time in broadleaved forests, no significant variations in pine dominated stands and significant fluctuations in spruce dominated stands, yet without a significant trend. Out of the tested drivers, dominant tree species exhibited the strongest impact on species community composition. Number of hemerophobic and strongly hemerophobic species did not undergo significant variations across years either. The lack of bryophyte diversity trends highlighted in this study suggests Central European managed forests are in an equilibrium sate, maintained by the opposing effects of climate changes, on one side and of more sustainable forest management and pollutant deposition decline, from the other.

     

Edge effect in different groups of organisms: vascular plant, bryophyte and fungi species richness across a forest-grassland border

Six transects were established on the edges of two deciduous forests near Krosno (Carpathian Foothills) to compare the species richness pattern of vascular plants, bryophytes and fungi. The transects had the shape of a cross with one arm 10 m along the forest edge and the other across the edge, 50 m into the forest interior and 12 m into the grassland. They consisted of 2×2 quadrats. The strongest edge effects were recorded for bryophyte, shrub and tree species richness, the weakest for herbaceous plant species richness. Overall vascular plant species richness and herbaceous species richness were higher in grassland than in the forest and peaked in grassland, 3 m from the forest edge. The shrub species richness was highest 1 m from the edge (in the forest) and the tree species richness 3 m from edge. Bryophyte species richness had roughly the same level across the grassland and within the first several meters of the forest, except for the 2 m zone on the edge itself where species richnes was as low as in the forest interior. Fungi species richness was low in the grassland and on the forest edge and rose dramaticaly a few meters from the edge, ramaining at the same level within the forest. The species composition across the forest-grassland border was analysed using detrended correspondence analysis. It revealed that in the case of bryophytes the increase in species richness did not correspond to a change in species composition, such as might have been caused by a general increase in bryophyte density.     

Are post-fire residual forest patches refugia for boreal bryophyte species? Implications for ecosystem based management and conservation

Residual patches of forest remaining after natural or anthropogenic disturbance may facilitate regeneration of fragmented forest. However, residual patch function remains unclear, especially after natural wildfire. We investigate the role of residual boreal forest patches as refugia for bryophytes and ask the question, do they house bryophyte communities similar to those encountered in undisturbed forests? Bryophytes were sampled in three habitat types in black spruce boreal forests illustrating a gradient of disturbance severity: undisturbed forests, residual patches and burned matrices. Temporal, disturbance severity, spatial and structural variables of habitats were also recorded. Bryophyte community composition differed among habitat types with residual patches characterized by higher species richness, the loss of forest specialists and the addition of disturbance-prone species. The bryophyte community found in residual patches is at the interface between the communities of undisturbed forests and burned matrices. As residual patches did not conserve all species, particularly forest specialists, they were not refugia. However, we identify temporal, spatial and structural characteristics that can maintain bryophyte communities most similar to undisturbed forests and enhance residual patch “refugia potential”. Residual patches enhance bryophyte diversity of the landscape housing species that cannot survive in the burned matrix. As conclusion we discuss the use of retention patches in harvested stands, together with the preservation of undisturbed stands that house singular bryophyte communities and especially sensitive forest specialists.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

Predictors of moss and liverwort species diversity of microsites in conifer‐dominated boreal forest

Patterns of moss and liverwort species diversity — species richness and species turnover (β‐diversity) — in three conifer‐dominated boreal forest stands of northern Alberta, Canada are described. We examined the relationship between bryophyte species diversity and micro‐environment at two sample grains, the microsite — substrate types for moss colonization: logs, stumps, tree bases, undisturbed patches of forest floor (dominated by feather moss species), and disturbed patches of forest floor — and the mesosite (25 m × 25 m plots). Microsite type and properties (e.g. decay class, hardwood vs softwood, pH) were the principal predictors of bryophyte species diversity and not micro‐environment variation among mesosites. Microsite type was the strongest predictor of microsite species richness and β‐diversity was higher among microsites and types and within microsites than among mesosites or stands. Microsite properties were significant predictors of species richness for all microsite types. Log and stump decay classes, influenced also by hardwood vs softwood predicted species richness of woody microsite types and soil pH and moisture predicted species richness of forest floor microsites. β‐diversity was highest for tree bases and disturbed patches of forest floor and lowest for logs. Mesosite β‐diversity was lower than that among microsites, and mesosite species richness was not well explained by measured environmental parameters. Results suggest that in conifer‐dominated boreal stands, species richness of microsites is only negligibly influenced by within‐stand variation at the mesosite grain and that substrate characteristics are the most important predictors of bryophyte species diversity in this ecosystem.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

Bryophytes and their microhabitats in coniferous forest pastures: should they be considered in the pasture management?

Forest pastures, like many other semi-natural (traditional) rural biotopes, have undergone a drastic decline both in area and quality during the last century in many areas. We explored the bryophyte flora of Finnish coniferous forest pastures on acidic soil and aimed to recognize the most important microsites (rocks, coarse woody debris, tree bases, mineral soil patches and closed vegetation) for bryophyte diversity. The effects of microhabitat heterogeneity (microsite entropy) on bryophytes was also examined. We found altogether 83 bryophyte species. The only red-listed species, Tayloria tenuis, was frequently found on dung patches and a few rare ruderal species grew exclusively on bare mineral soil. Rocks comprised the most species rich microsite and many common forest floor species showed preference for this microsite. Microhabitat heterogeneity explained bryophyte species richness on both alpha (plot average) and gamma (pasture total) scales. The results suggest that certain individual bryophyte species and their microsites should be taken into account in the management of this biotope, rather than guiding the management solely on the basis of the overall bryophyte diversity.     

Determinants of stream bryophyte community structure: bringing ecology into conservation

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Identifying indicator species for bryophyte conservation in fragmented forests

It is important to conserve bryophyte diversity in fragmented forests, due to the vulnerability of this group to environmental change. In this study, the utility of bryophyte functional groups (taxonomic classes, substrate classes, and life-forms) was examined as indicators for planning urban area conservation of bryophyte diversity. The study sites comprised 27 fragmented forests in Kyoto City, Japan. Using linear regression models, it was found that the species richness of almost all functional groups was positively correlated with patch size. Furthermore, two types of bryophyte life-form (fans and thalloid mats) were significantly correlated with environmental factors considered important for conservation planning. The species richness of fan bryophytes was positively correlated with the presence of natural forest patches and was negatively correlated with distance from mountainous areas. Similarly, the species richness of thalloid mat bryophytes was negatively correlated with maintenance practices. These results may be explained by the vulnerability of these two bryophyte groups to environmentally caused drought stress, accompanied by decreasing patch size, maintenance practices, disturbance, and/or loss of natural vegetation. Considering that drought stress represents a major threat to bryophyte diversity in fragmented forests, the species richness of hygrophilous life-forms (e.g., fans and thalloid mats) may be used as an indicator of fragmented forests that are less affected by drought stress, and these species should be preferentially conserved to maintain high levels of bryophyte diversity.     

Conserving a variety of ancient forest patches maintains historic arthropod diversity

Forests are naturally extensive tracts. However, in South Africa natural fires over many millennia have reduced forested areas into small remnants spread throughout a grassland matrix. Small patches, especially distant patches, are generally considered to be adverse for forest specialists, owing to decreased forest interior and increased edge. Here we test this assumption by determining the impact of forest interpatch distance and patch size on epigaeic arthropod diversity in this globally rare vegetation type. Forty patches were selected: ten large (100–435 m diameter) that are distant (500–645 m) from other patches, ten large that are close to other patches (38–97 m), ten small (30–42 m) that are distant, and ten small-close patches. Each patch had two plots: edge and interior. Arthropods were sampled using pitfall traps, Berlese-Tullgren funnels and active searches. Interiors and edges had similar species richness and composition, excluding spiders, which were richer in interiors. Patch size significantly influenced species richness of predatory beetles and arthropod assemblages, excluding spiders. Effect of the interaction between patch size and interpatch distance on species richness and composition varied among taxa. Furthermore, large patches supported similar assemblages regardless of interpatch distance. Arthropod response, particularly ants to patch size and interpatch distance, was partly shaped by the matrix type. The percentage of surrounding grassland had little effect on arthropod diversity. We can conclude that large and close patches are important for arthropod conservation. Nevertheless, it is also important to conserve a variety of patch sizes at various distances to maximize overall arthropod composition.     

Alternative pathways of liana communities in the forests of northwestern Argentina

Liana dynamics in secondary and mature forests are well known in tropical areas dominated by native tree species. Outside the tropics and in secondary forests invaded by exotic species, knowledge is scarce. In this study, we compare liana communities between secondary and mature forests dominated by native species in a subtropical montane area of Sierra de San Javier, Tucuman, Argentina. Additionally, we evaluate changes of liana communities in secondary forests with increasing densities of Ligustrum lucidum and Morus alba, two of the most invasive exotic trees of the area. We surveyed liana species richness and density in three 30-year secondary patches, four 60-year secondary patches, and four mature patches dominated by native tree species, to analyze changes in liana communities with forest age. Within each patch, we sampled 10–25 20 × 20 m quadrats. Additionally, we surveyed liana density and species richness in secondary forest patches with different densities of L. lucidum and M. alba. In native-dominated forests, liana species richness increased and showed a tendency of increasing basal area from 30-year secondary forests to mature forests. Liana density was highly variable, and most of the species were shared between native-dominated secondary and mature forests. Liana density and species richness decreased with L. lucidum density, whereas in secondary forests highly dominated by M. alba, lianas increased in density. Overall, lianas followed different pathways influenced by native forest succession and exotic tree invasions.     

Invasion by native tree species prevents biotic homogenization in novel forests of Puerto Rico

There is concern that secondary forests dominated by introduced species, known as novel forests, increase taxonomical similarity between localities and lead to biotic homogenization in human-dominated landscapes. In Puerto Rico, agricultural abandonment has given way to novel forests dominated by the introduced African tulip tree Spathodea campanulata Beauv. (Bignoniaceae). In this study, I characterized the tree species composition of S. campanulata forests in Puerto Rico as means to evaluate if biotic homogenization is occurring. Non-metric multidimensional scaling was used to examine what variables were related to the large (≥10 cm diameter at breast height [DBH]), small (≥2.5 to <10 cm DBH), and juvenile (<2.5 cm DBH) tree species composition of 20 sites. Species composition was strongly related to substrate properties, less related to land use history, and unrelated to spatial attributes. The introduced species component was low (mean = 17%, S.E. = 1.8) and compositional differences were mostly due to native tree species of secondary to old growth forests on equivalent substrates. Animals appear to disperse most species (86%) into these forests yet because of this some introduced species will persist. Although uncommon species were largely absent, recent species establishment is shaped by substrate properties making biotic homogenization in these forests unlikely. The S. campanulata forests of Puerto Rico facilitate native tree species establishment in lands where poor management practices extirpated the original forest. These results highlight the importance of remnant old growth forests or trees that act as seed dispersal sources and facilitate native species recovery in novel forests.     

Spider diversity responds strongly to edge effects but weakly to vegetation structure in riparian forests of Southern Brazil

Riparian forests bordering open terrestrial environments may have three microhabitats differing in structure and conditions: a grassland/pasture-forest edge (GE), a forest interior (FI) and a river–forest edge. The influence of such edge effects and vegetation characteristics on spider diversity of riparian forests was evaluated in Southern Brazil. Four different rivers were sampled on the tree–shrub strata with a beating tray, twice per season for 2 years. There were six transects per river, two per microhabitat. We compared spider abundance, species richness and composition. Vegetation variables sampled were vertical structure and (horizontal) density, canopy height and cover. Overall 42,057 spiders were sampled, 28 spider families and 440 species. The FI had higher spider abundance than the edges. Average species richness differed among rivers. Microhabitats did not differ in average richness, although overall richness (from sample-based rarefaction) was higher for GE than FI. High abundances in FI may result from lowered stress due to abiotic conditions, while higher GE richness may result from a faunal superposition between forest species and those from the grassland/pasture. Only canopy cover returns a positive relationship with spider diversity (richness and adult abundance). This might result from more spider species preferring to build webs or hunt under low-light environments. Rivers had spider faunas differing in composition but among microhabitats species composition was the same. Vegetation structure has been hypothesized to affect spiders, but this impact might be best seen in specific subgroups or guilds within spiders, not in the whole assemblage.