Diversity and stability in garrigue ecosystems after fire

Summary Vegetation dynamics after fire was studied in six communities in Bas-Languedoc (Southern France). 47 plots were observed by means of a permanent transect for ten years.In the first part, we describe floristic richness, species fugacity and the way by which, the terminal community (as defined by the last observation) appears. The dynamics of all these communities follows a simple and general model: floristic richness reaches its maximum during the first two years after a fire, then decreases and becomes stable. Fugacity follows a similar model, whereas the mergence of the terminal community is rapid: one year after fire 70% of the plots have already acquired 75% of the species of the terminal community. There is no succession (in the general sense of the word), but a progressive reappearance of the species belonging to the original community.In the second part, we study floristic similarities between our plots and corresponding associations as described in literature. It appears that after a fire the floristic diversity of the landscape remains high; while the communities rapidly reach a relative maturity.In the study area fire seems to be a rather superficial phenomenon; it does not lead to an important modification of the community dynamics, because probably the most frequent species in Bas-Languedoc developed adaptations to withstand fire.     

Plant species diversity in abandoned coppice forests in a temperate deciduous forest area of central Japan

We investigated plant species diversity as it related to stand structure and landscape parameters in abandoned coppice forests in a temperate, deciduous forest area of central Japan, where Fagus crenata was originally dominant. The species occurring in the study plots were classified into habitat types based on a statistical analysis of their occurrence bias in particular habitats (e.g., primary forest, coniferous plantation) in the landscape studied. The relationships between stand structure, which reflected the gradient of management, and forest floor plant species diversity (H and J) and richness (number of species per unit area) were not significant. However, these factors did influence the forest floor plant composition of the different types of habitat. According to the multiple regression analysis, species diversity and the richness of forest floor plants was affected by landscape parameters rather than by stand structure. For trees, species richness was mainly affected by the relative dominance of F. crenata, which is one of the stand structure parameters that decreases with intensive management. This is probably because many of the tree species that are characteristic of coppice forests increase after F. crenata have been eliminated by management; these species are not dominant in the original forest, where they are suppressed by F. crenata, the shade-tolerant dominant species. The species diversity (H and J) of trees was positively correlated with some landscape parameters, including the road density around the study plot, which may be associated with the intensity of management activity. The number of disturbance-tolerant species increased with increasing road density. Stand structure mainly affected disturbance-intolerant forest floor plant species and disturbance-tolerant tree species. Thus, the species diversity responses differed between forest floor plants and trees. The impact of forest management on species diversity was more prominent for forest floor plants.     

Epiphytic succession on tree trunks in a mixed oak-cedar forest,Kumaun Himalaya

The epiphytes present at about breast height on trunks of different size were studied for three major tree species in a seasonally wet forest at 2050 m altitude in the Kumaun Himalaya: Cedrus deodora, Quercus floribunda and Q. leucotrichophora. The total biomass and species number per unit trunk area, were found to increase with trunk size. It was supposed that the results indicated a succession in the type of epiphytic cover from young trunks to older trunks.The amount of loose material (plant remains and soil) per unit area of trunk increased with increasing girth. The C:N ratio in this material was initially very high on the oaks (129–197) and declined with increasing trunk size (to 73–78); the ratio was constant across girth classes in the cedar (86–87).Bryophytes produced most biomass on most trunks; next to them were lichens on the smallest trunks, and flowering plants on the largest. The number of species of epiphytes was similar on all three host species. The results are discussed in relation to contemporary ideas on diversity and strategies.Nomenclature follows: Ganguli (1972, 1977 & 1980); Kashyap (1929); Nayar (1970); Osmaston (1927).Financial support from the Council of Scientific and Industrial Research, Delhi and University Grants Commission, New Delhi is gratefully acknowledged.     

Impact of management intensity on non-vascular epiphyte diversity in cacao plantations in western Ecuador

A first study on the biodiversity of non-vascular epiphytes in cacao (Theobroma cacao L.) plantations in western Ecuador yielded 112 species (51 bryophytes, 61 lichens). Epiphyte assemblages of cacao plantations resembled those of tropical rain forests but species richness was usually lower and individual species were found at lower heights on the trunks. The vast majority of the species are widespread neotropical or pantropical species; one species, Spruceanthus theobromae (Spruce) Gradst., is endemic to cacao plantations of western Ecuador. Differences in management intensity, by manual removal of epiphytes from tree trunks (limpia), had a significant impact on epiphyte species diversity. Total species richness was significantly reduced in plantations with high management intensity, due mainly to the decreased diversity of lichens and liverworts; moss diversity was not affected by management regime. Total percentage cover of bryophytes was highest in plantations with low management intensity, while lichen cover was greatest in plantations with high management intensity. Crustose lichens and smooth mats growing closely appressed to the substrate were little affected by the limpia and their growth may have been promoted when larger-sized species were removed, by reducing competition. Cacao plantations with low and moderate management intensity serve as an important substitute habitat for ecological specialists (sun epiphytes, shade epiphytes) of the rain forest and are of considerable significance for their conservation.     

Macrofaunal diversity and habitat structure in intertidal boulder fields

Low-midshore boulder fields in Europe are increasingly subject to degradation related to beach fishing for crabs and abalones. The aim of the study was to better understand the structure and species diversity of macrofaunal assemblages living in a low-midshore boulder field in order to define conservation strategies for this biotope. Sampling strategy involved different spatial scales (macro- and microstrata) relative to the complexity and heterogeneity of the habitat. Assemblages of species and the corresponding habitats were identified by multifactorial analysis and compared by ANOVAs. The results show a hierarchical organization of the macrofauna within the boulder field, corresponding to three spatial levels of habitat structure: (i) eight microhabitats at the lowest level of spatial organization, each defined by a specific assemblage (subcommunity); (ii) three habitats at a middle level combining these microhabitats, each associated with a specific community (open rock, protected rock and sediment); and (iii) three macrohabitats at the uppermost level (corresponding to the scale of the entire boulder field and including the main geomorphological features of the beach), each defined by a specific assemblage of species (boulders on boulders, boulders on bedrock, and boulders on sediment). Two microhabitats with particularly high species diversity were regarded as the most important ecological units of the field in terms of conservation of specific and functional biodiversity. Comparison of habitat/community parameters showed (i) that habitat heterogeneity was not an accurate indicator of faunal diversity, and (ii) that habitat complexity enhanced the species richness of the fauna, but only above a threshold value. This enhancement was due to semi-sheltered microhabitats, which were found only in the most complex areas of the boulder field. It is likely that this complexity affects species richness qualitatively more than by the diversity of microhabitats. In other words, a cross-scale effect is apparent in which high habitat complexity on the middle spatial scale creates microhabitats on the lowest spatial scale that are characterized by stable semi-sheltered environmental conditions conducive to a maximum of species.     

Breeding-habitat use by five Amazonian frogs at forest edge

Understanding patterns of speciesl habitat use and their response to dramatic changes in habitat constitutes a basis for sound conservation practice. This study examined use of breeding habitat by two rainforest-breeding frog species near primary forest edge. Nine artificial breeding pools were established along each of three transects perpendicular to continuous primary forest near Manaus, Brazil. Five frog species bred in the pools over the six-month monitoring period. For the two most abundant species, patterns of habitat use were independent of proximity to forest edge. Also,Epipedobates femoralis used pools outside the forest, in secondary growth, at the same frequency as pools inside the forest.Osteocephalus taurinus did not breed outside primary forest. These results have clear implications for conservation biology: (i) buffer zones around isolated reserves, to counter possible negative edge effects on habitat, are unnecessary for these frog species, and (ii) secondary growth habitat, which is not natural breeding habitat forE. femoralis, could serve as a source of colonists for emigration into new areas and does not represent a barrier to dispersal for this species. These results argue against generalization of edge effects for different taxa.     

Landscape patterns and plant species diversity of forest reserves in the Kanto region, Japan

Plant species richness of twenty old-growth forest reserves in the cool-temperate zone in the Kanto region, Japan were investigated to detect the effect of forest fragmentation. The species richness of trees and forest floor plants were analyzed by multiple regression models relating to nine variables on the characteristics of landscape, local habitat and forest stand. The total species diversity did not have a significant correlation with any variables of landscape patterns. In this study, single large reserve in the SLOSS discussion did not seem very effective to preserve more species. However, forest reserves in large patches tend to have relatively infrequent species. Large patches of natural forests were regarded as one of the important factors to preserve infrequent species.     

Stand structure and woody species diversity in relation to stand stratification in a subtropical evergreen broadleaf forest, Okinawa Island

Stand structure and woody species diversity in a subtropical evergreen broadleaf forest grown in a silicate habitat, Okinawa Island, have been investigated on the basis of stand stratification. The forest stand consisted of four layers. The floristic composition of the top and the lower three layers was only slightly similar, although approximately one-third of the species were common to them. Mean tree weight decreased from the top toward the bottom layer whereas tree density increased from the top downward. This trend resembled the mean weight–density trajectory of self-thinning plant populations. The relationship between mean tree height and tree density for the upper two layers supported Yamakuras quasi –1/2 power law of tree height. The values of the Shannon–Wiener index, H, and the equitability index, J, tended to increase from the top layer downward except for the bottom layer. The values of H and J were, respectively, 4.83 bit and 0.82 for trees taller than 0.10 m. The lower layers contained many species of smaller height. High species diversity of the forest depended on small trees in the lower layers. Conservation of small trees in the lower layers, especially the bottom layer, is indispensable for sound maintenance of Okinawan evergreen broadleaf forests.     

Population structure and dynamics of a neotropical palm in fire-impacted fragments of the Brazilian Atlantic Forest

The demography of the acaulescent palm Attalea humilis was studied from 1996 to 1999 in three forest fragments (1.6, 6.4, and 9.9ha) in southeastern Brazil (22°30–22°33 S, 42°15–42°19 W). A human-induced fire damaged the populations in August–September 1997. Palm density was higher in the medium-sized fragment, where it was concentrated in a single disturbed sector. Population ontogenetic stage structure was dependent on fragment and year, but in general later stages were more abundant than initial ones. Population finite growth rates projected from linear, stage-structured demographic models indicated that the populations were increasing before fire. Population growth rate in the smallest fragment was larger due to a single recruitment event, being most sensitive to changes in survival and growth of seedlings, virgins and the immature, and relatively insensitive to changes in shrinkage and fecundity. Observed growth rates for the entire study period showed that populations remained in equilibrium after fire, but the population in the medium-sized fragment showed a declining tendency. Population structure was resilient to fire, which caused a transitory concentration of individuals in the medium-sized stage. The self-sustainability of the population in the largest, most preserved fragment suggests that habitat-protection measures may suffice to ensure the persistence of populations in small fragments. However, population fluctuations and exclusion from hyper-disturbed sectors in the smallest fragment suggest that reduced fragment area, habitat degradation and human-induced fires can interact to threaten the persistence of even abundant, preadapted to fire, species like A. humilis.     

Carbon and nitrogen isotope ratios in different compartments of a healthy and a declining Picea abies forest in the Fichtelgebirge,NE Bavaria

Summary Natural carbon and nitrogen isotope ratios were measured in different compartments (needles and twigs of different ages and crown positions, litter, understorey vegetation, roots and soils of different horizons) on 5 plots of a healthy and on 8 plots of a declining Norway spruce (Picea abies (L.) Karst.) forest in the Fichtelgebirge (NE Bavaria, Germany), which has recently been described in detail (Oren et al. 1988a; Schulze et al. 1989). The ¹³C values of needles did not differ between sites or change consistently with needle age, but did decrease from the sun-to the shade-crown. This result confirms earlier conclusions from gas exchange measurements that gaseous air pollutants did no long-lasting damage in an area where such damage was expected. Twigs (¹³C between-25.3 and-27.8) were significantly less depleted in ¹³C than needles (¹³C between-27.3 and-29.1), and ¹³C in twigs increased consistently with age. The ¹⁵N values of needles ranged between-2.5 and-4.1 and varied according to stand and age. In young needles ¹⁵N decreased with needle age, but remained constant or increased in needles that were 2 or 3 years old. Needles from the healthy site were more depleted in ¹⁵N than those from the declining site. The difference between sites was greater in old needles than in young ones. This differentiation presumably reflects an earlier onset of nitrogen reallocation in needles of the declining stand. ¹⁵N values in twigs were more negative than in needles (-3.5 to-5.2) and showed age- and stand-dependent trends that were similar to the needles. ¹⁵N values of roots and soil samples increased at both stands with soil depth from-3.5 in the organic layer to +4 in the mineral soil. The ¹⁵N values of roots from the mineral soil were different from those of twigs and needles. Roots from the shallower organic layer had values similar to twigs and needles. Thus, the bulk of the assimilated nitrogen was presumably taken up by the roots from the organic layer. The problem of separation of ammonium or nitrate use by roots from different soil horizons is discussed.     

Snake communities of moist rainforest and derived savanna sites of Nigeria: biodiversity patterns and conservation priorities

The taxonomic composition and the abundance of two communities of snakes were studied in two different areas of southern Nigeria. One community was studied in a derived savanna area (environs of Ejule, 06°54N, 07°23E), and one community was studied in a moist rainforest area (environs of Eket, 04°50N, 07°59E). Both the specific diversity and the mean frequency of observation of snakes were significantly higher in the forest area than in the savanna area, and the dominance index was higher in the savanna than in the forest site. However, most of the species were found in only one of the two areas, depending on their habitat requirements (e.g. Gastropyxis smaragdina, Dispholidus typus, Thelotornis kirtlandii, Dendroaspis jamesoni, Bitis nasicornis, Causus maculatus, etc). The forest community housed a significant number of arboreal and semiaquatic taxa, but in both sites most of the species were terrestrial. The commonest species in the forest area was an oviparous, semiaquatic, batracophagous natricine snake (Afronatrix anoscopus), whereas the commonest species in the savanna area was an oviparous terrestrial lacertophagous psammophine snake (Psammophis phillipsi). Some conservation implications of our biodiversity analyses are presented. It is suggested that the moist rainforest represents a critically endangered habitat, and should deserve special attention by the international scientific community. Oil industry activity is especially dangerous for snake communities, especially in the southernmost regions of Nigeria.     

Diversity,metabolic types and δ13C carbon isotope ratios in the grass flora of Namibia in relation to growth form,precipitation and habitat conditions

The grass flora of Namibia (374 species in 110 genera) shows surprisingly little variation in ¹³C values along a rainfall gradient (50–600 mm) and in different habitat conditions. However, there are significant differences in the ¹³C values between the metabolic types of the C4 photosynthetic pathway. NADP-ME-type C4 species exhibit the highest ¹³C values (–11.7 ) and occur mainly in regions with high rainfall. NAD-ME-type C4 species have significantly lower ¹³C values (–13.4 ) and dominate in the most arid part of the precipitation regime. PCK-type C4 species play an intermediate role (–12.5 ) and reach a maximum abundance in areas of intermediate precipitation. This pattern is also evident in genera containing species of different metabolic types. Within the same genus NAD species reach more negative ¹³C values than PCK species and ¹³C values decreased with rainfall. Also in Aristida, with NADP-ME-type photosynthesis, ¹³C values decreased from –11 in the inland region (600 mm precipitation) to –15 near the coast (150 mm precipitation), which is a change in discrimination which is otherwise associated by a change in metabolism. The exceptional C3 species Eragrostis walteri and Panicum heterostachyum are coastal species experiencing 50 mm precipitation only. Many of the rare species and monotypic genera grow in moist habitats rather than in the desert, and they are not different in their carbon isotope ratios from the more common flora. The role of species diversity with respect to habitat occupation and carbon metabolism is discussed.     

Tree species diversity of tropical forest vegetation in Xishuangbanna, SW China

Tree species diversity of four tropical forest vegetation types was investigated in Xishuangbanna, southwestern China. These are: tropical seasonal rain forest, tropical montane rain forest, evergreen broad-leaved forest and monsoon forest over limestone. A total of 17 samples were taken and four species diversity indices were calculated: Shannon-Wiener's H, the complement of Simpson's index, d, Fisher's and evenness index E. The results reveal the long-tailed rank/abundance diagrams of these forests. However, this feature is greatly reduced in the samples of monsoon forest over limestone. Tropical seasonal rain forest shows the highest tree species diversity of all four vegetation types. Owing to the variation of microenvironment, diversity values within the same vegetation type vary between the samples from different patches. The tree species diversity of single-dominant rain forest is not significantly lower than that of mixed rain forest, because the dominant species of some single-dominant rain forests are principally in the emergent layer. This is composed of sparse and huge trees of one species and, consequently, creates a unique canopy architecture and more heterogeneous microenvironments for the more diversified species composition under the emergent layer. The occurrence of tree species with small population sizes, particularly of species represented by only one individual, is highly correlated with the tree species diversity of the local forest vegetation. They are crucial elements in the richness of local biodiversity.     

Diversity and abundance of leafhopper species (Homoptera: Cicadellidae) among red maple clones

A survey was conducted in the 2001 growing season to examine the leafhopper diversity and abundance among trees of 17 red maple (Acer rubrum) clones. Yellow sticky traps were used to qualify and quantify the number of aerial leafhoppers from 1 May 2001 until 4 September 2001. A total of 45 species from eight different leafhopper subfamilies, for a total of 6055 individuals, were considered in this study. The mean number of leafhoppers collected, mean species richness, diversity and evenness were significantly lower on traps of trees for October Glory than for the other clones. Yet, none of the leafhopper species dominated the weekly samples. Species similarity among clones ranged from 56 to 90%. No two clones had complete similarity. Franksred and trees of a controlled cross between October Glory × Autumn Flame shared the highest degree of species similarity, while clones from PA, USA and RI, USA were the least similar. The development of new clones did not create new pest problems, but suppressed populations of damaging pests, and maintained the diversity of low abundance species.     

Testing for non-randomness in sizes and habitats of West Indian lizards: choice of species pool affects conclusions from null models

Summary Several null models are proposed for testing whether size or habitat differences in West IndianAnolis lizards are greater than expected by chance. The models differ primarily in choice of the pool from which species are sampled to form random communities. Regardless of choice of pool, size differences in the Lesser Antilles are greater than null models predict; the pool using species on the known source (Puerto Rico) gives a greater variance in ratios but about the same mean ratio (for males), or a greater mean ratio (for females), compared with the pool composed of species on the islands being tested (the Stronget al., 1979, Galápagos procedure). On satellite islands of the Greater Antilles, sizes do not differ more than expected from null models. Pools composed of mainland-source species give null communities with more small or more large ratios than those composed of island species, depending upon whether four-species islands are included or excluded, respectively. Colwell and Winkler's unmodified Narcissus hypothesis is contradicted by these results in procedures where species not likely to be able to occur on small islands are included in the species pool. Using the most biologically reasonable, but not other, choices of source pool, species on satellite islands of the Greater Antilles differ more in structural habitat then expected by chance. In contrast to some of the results on size, here mainland-source pools are more likely to produce a statistically significant difference between real and random communities, as predicted by the Narcissus hypothesis. However, exclusion of structural habitat categories not found on satellite islands is necessary to achieve this significance.     

Food-web models that generate constant predator-prey ratios

Summary An approximately constant ratio of number of predator species/number of prey species is observed in several natural communities, although the exact value of the ratio may vary with habitat and the types of organisms in the food web. We test the hypothesis that a constant predator/prey ratio can be generated by what Holt (1977) terms apparent competition and what Jeffries and Lawton (1984) call competition for enemy-free space. We create simple, two trophic-level communities by drawing species of predators and prey at random from a species pool, simulating their interactions using Lotka-Volterra models. The simulated food webs converge over successive periods of invasion and extinction to locally stable systems with approximately constant ratios of number of predator species/number of prey species, despite varying initial conditions. As expected, predator/prey ratios take different values depending upon the biology of the simulated species. We conclude that apparent competition between prey species via shared enemies may be one mechanism whereby approximately constant predator/prey ratios are generated in natural communities.     

Forest responses to the large-scale east coast fires in Korea

The east coast forest fires of April 2000 were Koreas largest recorded fires. This, along with the fact that they took place in the region most frequently affected by fire, attracted a great deal of attention. Due to the variations in wind, topography and pre-fire forest stands, a heterogeneous landscape mosaic of burn severity was created across the region. It turned out to be an excellent opportunity to study various landscape-scale impacts of fires on forest dynamics. Therefore, we investigated stands in the 23794ha of burned forest region, in terms of burn severity, vegetation regeneration and forested landscape change as a measure of community stability. Using the geographic information system technique, we analyzed the differential severity and post-fire recovery of pre-fire forest types of different stand age both at stand and species level. Analysis showed that pre-fire vegetation was composed of mainly pine (Pinus densiflora) stands that occupied 70% of the whole forested area, while pine-hardwood and hardwood stands occupied only 28% and 3%, respectively. In addition, two-thirds of all stands were less than 30-years-old. Pine stands were the most severely burned, while conversely pine-hardwood and hardwood stands were less vulnerable. This implied that pine forests had fire-prone characteristics. Vegetation recovery went the opposite way; that is, the regenerating vegetation cover was 71% at pre-fire hardwood stands, and 65% and 53% at pine-hardwood and pine stands, respectively. However, these recovery rates were strikingly fast, considering that investigation took place about 3months after the fires. Fire did not initiate successional processes, but tended to accelerate the predicted successional changes by releasing pre-fire understory species that survived the fires and regenerated by sprouting. The dominant pre-fire tree species (P. densiflora) was susceptible to fire and not resilient enough to reestablish in competition with oak species. Contrary to pines, the abilities of oak species, mainly Quercus mongolica and Q. variabilis, to survive fires and to resprout vigorously made them dominant at most post-fire stands. These shifts in species abundance caused drastic changes to the landscape: from pine-dominated to oak-dominated stands without any notable change in species composition. The patterns in forest regeneration that we observed in Korea may be representative of forest responses to any long-term repeated disturbances, including fire.     

Species richness and composition of Curculionidae (Coleoptera) in a conifer plantation, secondary forest, and old-growth forest in the central mountainous region of Japan

Weevil (Curculionidae, Coleoptera) species richness and composition were investigated and compared among larch [Larix kaempferi (Lamb.) Carriére] plantations, secondary forests, and old-growth forests in the central mountainous region of Japan. In addition, to assess the effects of forest-management practices, namely thinning and long-rotation logging schedules (long rotation), the weevil assemblages of recently thinned middle-aged and long-rotated larch plantations were compared with those of middle-aged larch plantations. Malaise traps were set in 44 stands of these forest types, and weevils were separated and identified. Several environmental factors other than forest type were also examined. Weevil species richness and diversity indices [Shannon-Wiener diversity index (H) and Simpsons index of diversity (D)] were higher in the secondary forest than in the larch plantation. Because of its wide distribution and higher weevil species richness, the secondary forest contributed to maintaining weevil diversity in this region. Old-growth forest had higher diversity indices (H and D) than did the larch plantation. The secondary forest had the highest number of species in total. Though the number of individuals was the highest in larch plantation, species richness, H and D of the plantation were generally low. Weevil community structure and species composition differed among the three forest types, but the difference in weevil composition between the larch plantation and the other two forest types was the largest. Forest type is probably the most important factor for determining the differences in weevil assemblage, and further, both dominant tree type (coniferous trees versus broad-leaved trees) and the number of mature tree species seem to be important factors for weevil species composition. Among forest management practices, long rotation caused diversity indices (H and D) to increase while thinning appeared to cause only minor changes in the weevil assemblages. Because species richness and species composition of Curculionidae well reflected the differences in forest types and some other environmental factors investigated, this family seems suitable for diversity research in forests. Further research on biodiversity with the use of this family should, therefore, be expected.     

Lightning density and burning frequency in South African vegetation

This study investigates lightning density in relation to burning frequency in five South African biomes. Data from automatic lightning flash counters distributed throughout South Africa are used to estimate the range and mean values of annual lightning ground-strike density in each biome. The lowest average lightning frequencies were recorded in the karoo and fynbos biomes, while the highest values were obtained in the sour and inland mountain divisions of the grassland biome, with intermediate values computed for the savanna-woodland and forest biomes. These results were compared with published findings on the effects of different burning frequencies on plant communities in each biome. In general, plant communities occurring in areas experiencing the highest annual lightning densities (e.g., sour and mixed grassland, and moist savanna-woodland) tolerate and require more frequent burning than those typical of areas subject to lower lightning densities (e.g., fynbos, forest, sweet grassland, and arid savanna-woodland). These findings suggest that the constituent plant populations in each biome have adapted to burning frequency according to the local probability of ignition by lightning in the areas they inhabit, and that present day responses of plant communities to burning reflect their ancestral exposure to fire in the course of their evolutionary development under pre-existing, natural fire regimes.Research sponsered by the South African Council for Scientific and Industrial Research. We thank the Director, National Electrical Engineering Research Institute (NEERI), Pretoria, South Africa, and H. Kröninger, Lightning Research Division, NEERI, for providing data on lightning ground-strike densities and coordinates and elevations for counters in South Africa. We thank Drs J. H. Bock and M. T. Mentis for commenting on an early draft of this paper.     

The size and diversity of the soil seed banks and the light requirements of the species in sunny and shady natural communities of the Białowieża Primeval Forest

The research was conducted in two natural forest communities: Potentillo albae-Quercetum (oak forest) which allows much light to reach the forest floor and Tilio-Carpinetum typicum (hornbeam forest) which shades the herb layer heavily. The seed banks were estimated from numbers of seedlings emerging from soil samples over one growing season.(1) Our results confirm the hypothesis that persistent seed banks are mainly formed by species with high light requirements. Of the species found predominantly in the seed bank and absent from the herb layer or occurring there very rarely in both communities 83% of species and 70% of seedlings were strongly light-demanding (Ellenberg's light index 6–9). However, the results do not support the hypothesis that seed banks in natural deciduous forest communities are small, poor in species and do not reflect the species composition of herb layer.(2) The seed banks of both communities were rich in species and relatively large. Species richness in the oak forest turned out to be higher than in the hornbeam forest (51 vs 45 species/2.4 m₂), but size was smaller (2659 vs 5789 seedlings/2.4 m₂). In the oak forest the most abundant species in the seed bank was Galium boreale, but it constituted only 19% of the total number of seedlings, whereas in the hornbearn forest the dominant species, Urtica dioica, constituted 57% of the total.(3) In each community the species composition of the seed bank and the herb layer was very similar (>70%).(4) The seed bank was more diverse in the oak forest than in the hornbeam forest (H 2.34 vs 1.68).(5) The seed banks of both communities differed in the contribution of species with varied light requirements; in the sunny oak forest species with high light requirements dominated, whereas in the shady hornbeam forest both strongly and moderately light-demanding species had similar contributions.Nomenclature: Follows Ehrendorfer (1973) and Matuszkiewicz (1981).