Soil seed bank composition along a forest chronosequence in seasonally moist tropical forest,Panama

Abstract. We used a forest chronosequence at the Barro Colorado Nature Monument (BCNM) to examine changes in the abundance and species composition of seeds in the soil during forest succession. At each of eight sites varying from 20 yr to 100 yr since abandonment, and at two old-growth (> 500 yr) forest stands, we established two 160-m transects and sampled the surface 0–3 cm of soil in cores collected at each 5 m interval. Seed densities were estimated from the number of seedlings germinated from the soil over a six-week period. Contrary to expectation, neither the density of the soil seed bank, nor species richness or diversity were directly related to age since abandonment, but the density of the soil seed bank was correlated with the abundance of seed-bank-forming species in the standing vegetation. In marked contrast to published studies, herbaceous taxa were rare even in the youngest stands, and the common tree species, which accounted for most seeds in the soil, were present in all stands. The pioneer tree Miconia argentea (Melastomataceae) was the single most common species in the seed bank, accounting for 62% of seeds and present in 92% of soil samples. Rapid recovery of the vegetation of young regrowth stands on BCNM, when compared to sites elsewhere may be partly due to allochthonous seed rain from nearby mature forest stands and the lack of seed inputs of weeds and grasses from agricultural and pasture lands which may inhibit forest succession.     

The soil seed bank and understorey regeneration in Eucalyptus regnans forest,Victoria

Abstract The soil seed bank and its relation to the extant vegetation in a Eucalyptus regnans F. Muell. forest in the Central Highlands of Victoria were examined. The average seed density was 430 germinable seeds m^?2 to a depth of 2 cm. There was a polynomial regression relationship between the density and species richness of seeds in soil and forest age (0. 6–54 years). Species richness was not significantly different among soil depths (0- 2 , 2- 5 , 5–10 and 10–20 cm) in the forest stand of 54 years old. More seeds germinated from the 5–10 cm depth than from the other depths. Forbs accounted for 73% of the total germinable seeds and there was no germination of E. regnans. The number of species, particularly woody plant species, germinating from the soil seed bank were significantly lower than in the extant vegetation. However, almost all species present in the soil seed bank were present in the vegetation. The soil seed bank provides an important source for the rapid regeneration of understorey vegetation following clear-cutting and slash-burning in the E. regnans forest. The rapid understorey establishment may play an important role in protecting soil from erosion, in nutrient conservation, replacement and redistribution. The soil seed bank may also be a necessary source of maintaining genetic diversity in the forest over the long term.     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

Seed dynamics during forest succession in Costa Rica

Soil seed banks and current seed inputs each play a role in tropical succession. We compared the abundance and floristic composition of seeds from these two sources at a Costa Rican site by germinating seeds from the soil, measuring seed inputs for 3 yr, and monitoring the earliest colonists in a forest clearing.There were an estimated 6800 viable seeds/m² in the soil of 3.3-yr-old vegetation, 9500 seeds/m² in 11-yr-old vegetation, and 7000 seeds/m² in a 75-yr-old forest. An estimated 10100 seeds/m² fell on the soil surface of the young successional vegetation during 3 yr and 3700 seeds/m² fell during that same time in the forest.Locally produced seeds accounted for about 75% of the seed input to the soil surface early in succession. Seeds dispersed out of young successional vegetation increased the quantity and species richness of the seed input and storage in an adjacent forest. Much of the species richness of the young successional vegetation resulted from seeds dispersed there from other communities by animals.Deforestation stimulated germination of most seeds in the surface soil of the old forest, including seeds of the dominant canopy tree. The recruitment of seedlings from the soil seed bank numerically overwhelmed that from post-disturbance seed rain and sprouts.We evaluated patterns of soil seed storage during succession and predicted the ability of vegetation of differing ages to respond to disturbance. Immediately after disturbance the number of seeds in the soil plummeted due to mortality, low inputs, and germination. As the vegetation regrew, the soil seed bank increased to a peak after 4 to 7 yr, then gradually decreased to its pre-disturbance size. High-frequency pulses of disturbance should result in reduced species richness, dominance by species with long-lived seeds, and fast recovery by seedling recruitment from the soil seed bank.Journal series number 6459 from the Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida 32611, USA.Reprint requests to J. J. E. at Florida.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Frugivoría por Aves en un Paisaje Fragmentado: Consecuencias en la Dispersión de Semillas1

Few data exist on seed dispersal by frugivorous birds in fragmented landscapes, originating from tropical dry forests, in contrast to more abundant data from tropical rain forests. In this study, we assessed the effect of frugivorous birds in a fragmented landscape of Veracruz, Mexico, now occupied by remnant fragments of tropical semi‐deciduous forest and dry deciduous forest, grassland, and shrubby patches on sand dunes. We determined four characteristics related to seed dispersal by birds: the interacting species of plants and birds, the characteristics of these species, spatio‐temporal variation in the dispersal system, and the outcome of the process. During one year, we recorded 54 frugivorous bird species and 33 ornithochorous plant species, which engaged in 176 different bird‐plant species interactions. Similarity (Sorensen index) of frugivorous bird communities using different vegetation types was high (>70%), suggesting that many bird species used all of the vegetation types. In contrast, the similarity of ornithochorous plant communities among vegetation types commonly was low (<37%), suggesting that most plant species were restricted to particular sites in this landscape. At the landscape level, as well as for tropical deciduous forest, we detected a significant positive relationship (Spearman's correlation of rank coefficient >0.65, P <0.05) among richness per month of frugivorous birds and plant species bearing fleshy fruits. Seeds of many plant species previously detected in studies of seed rain at the site were eaten by birds during this study. Most seeds of zoochorous species, which are deposited in the dry and decidous tropical forests patches, are produced within these vegetation types (i.e., they are autochthonous species), whereas bird‐dispersed seeds arriving in grassland and shrubby patches are produced outside (i.e., allochthonous) and are mostly woody species. Birds are important seed dispersers among vegetation types in this landscape but they have different effects in each one. The four characteristics studied, as well as the landscape approach of this research, allowed us to detect spatial and temporal patterns that otherwise would have remained undetected.     

Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Restoration prospects of abandoned species‐rich sandy grassland in Hungary

Abstract. Secondary succession and seed bank formation was studied in a formerly grazed, abandoned, eastern Hungarian sandy steppe‐meadow (Pulsatillo‐Festucetum). The vegetation was sampled at different elevations of a sand dune which became partly invaded by the tree Robinia pseudo‐acacia ca. 10 yr ago. Pre‐abandonment vegetation records were used as historic references. Though composition of the non‐invaded grassland only changed moderately, dominance of tall grasses (Elymus hispidus, Poa angustifolia) increased significantly at the cost of annuals and low stature perennials. In the stand invaded by Robinia most grassland species were lost and replaced by nitrophytes. Vertical position influenced species abundance, but affected the composition only moderately. Fine‐scale zonation of the vegetation also changed with time. Species richness of the above‐ground vegetation and the seed density of soil samples at the lower elevation were slightly greater than at the higher sites. Seed banks of sensitive grassland specialists (e.g. Pulsatilla pratensis subsp. hungarica) disappeared during grass encroachment. Following extinction from above‐ground vegetation, restoration must rely on dispersal from adjacent areas. In contrast, several annuals and perennials, which survived this degradation stage in the above‐ground vegetation, possessed seed banks. Many of these species became extinct from the vegetation during the Robinia invasion but left viable persistent seeds. This fact is promising for restoration of the Potentillo‐Festucetum sandy pasture. Competitive weedy species and sprouting Robinia can, however, limit seedling establishment.     

Soil seed bank and regeneration potential on eroded hill slopes in the Kondoa Irangi Hills,central Tanzania

Abstract. The soil seed bank of the severely eroded Kondoa Irangi Hills, Tanzania was studied in order to determine the seed density and composition and to establish the relationship between seed bank and standing vegetation. The area had not been grazed for 15 yr prior to the study. A recently grazed area was used for comparison. The seed bank density (at 0 - 5 cm depth) ranged from 344 to 915 8 seeds/m² in the dry season and 172 to 5107 seeds/m² in the wet season. The seed bank was very heterogeneous, both spatially and temporally, and it showed significant variation in size and composition in both sampling periods. The species similarity between the seed bank and the above-ground vegetation in all plots was low (Sørensen's index = 0.00–0.44). The highest similarity was found in the recently grazed area. The seed bank was dominated by annuals and by early successional species. It is concluded that re-vegetating the hill slopes with woody vegetation by using the seed bank will be difficult because seeds of woody species were not found in the soil.     

Seed banks of a rare wetland plant community: distribution patterns and effects of human-induced disturbance

Seed banks in rare shoreline communities (Nova Scotia, Canada), were examined in order to determine their role in the persistence of rare plant species. A pristine shoreline which supports globally and nationally significant Atlantic coastal plain species averaged 8500 seeds/m². Seedsweremost abundant high on the shore and where standing crop was greatest. Rare species, which made up 22% of the standing crop and litter of the adult vegetation, comprised only 4% of the seed bank. Common rushes such as Juncus canadensis and J. filiformis, were not well represented in the adult vegetation (5% of the standing crop and litter) but were abundant in the seed bank (52% of all seeds). Seed densities were on average, much lower on an intensely disturbed shoreline (1000 seeds/m²), where heavy all-terrain vehicle traffic had destroyed the adult vegetation. These results suggest that intense disturbances will destroy both existing vegetation and the seed bank and moderate disturbances will alter community composition so that common rushes will replace already endangered Atlantic coastal plain species.     

Seed bank dynamics and tree‐fall gaps in a northwestern Mexican Quercus‐Pinus forest

Questions: How does the seed bank respond to different types of tree‐fall gaps and seasonal variations? How does the soil seed bank influence recovery of the standing vegetation in the mature forest and tree‐fall gaps? Location: 1800 — 2020 m a.s.l., Quercus‐Pinus forest, Baja California Sur, Mexico. Methods: Seed size, species composition and germination were estimated under different environmental conditions during dry and rainy seasons: a mature forest plot and gaps created by dead standing trees, snapped‐of f trees and uprooted trees. The soil seed bank was investigated using direct propagule emergence under laboratory conditions, from soil cores obtained during both seasons. Results: 21 species, 20 genera and 14 families constitute the seed bank of this forest community. Fabaceae, Asteraceae, Euphorbiaceae and Lamiaceae were the most frequently represented families in the seed bank. Floristic composition and species richness varied according to the different modes of tree death. Species composition of seed banks and standing vegetation had very low similarity coefficients and were statistically different. Seed bank sizes varied between 164 and 362 ind.m^‐2 in the mature forest plot for the dry and rainy seasons, respectively, while soil seed bank sizes for gaps ranged between 23–208 ind.m^‐2 forthe dry season and between 81–282 ind.m^‐2 for the rainy season. Conclusions: Seed bank sizes and germination response were always higher in the rainy season under all the environmental conditions analysed. Results suggest that timing responses to gap formation of the soil seed bank could be more delayed in this temperate forest than expected.     

Seed Availability as a Limiting Factor in Forest Recovery Processes in Costa Rica

Abandoned pastures and secondary forests are increasingly prominent features of tropical landscapes. Forest regrowth on abandoned pastures is generally slow and virtually limited to regeneration from seeds from external sources, since agricultural activities alter site conditions. We hypothesize that seed availability is a major limiting factor in forest recovery on abandoned pastures. This hypothesis was tested by studying the seed bank, seed rain, and seed predation in a small pasture (1 ha) situated in a forest‐pasture mosaic in northwestern Costa Rica. The tree seed density in the pasture seed bank was much lower (21/m²) than the density in the seed bank of a neighboring secondary forest (402/m²). Within a period of five weeks, 23 tree seeds entered the pasture by seed rain. This number is low compared to densities found in closed forests but higher than densities reported in other studies where virtually no seeds were found beyond 20 m from the forest edge. Possibly the small size of the pasture with seed sources nearby and the small‐scale landscape mosaic enhance seed dispersal. Predation limits the seed density in pastures, with 42% of the woody species consumed by predators. The low seed density in the seed bank, and hampered recruitment combined with significant losses, pose severe restrictions to forest recovery on abandoned pastures. Moderate land use, and small sized clearings with seed sources nearby may increase the pace of recovery. Nevertheless, forest establishment may still take a considerable time. Thus, enlarging the available pool of species may be a worthwhile management strategy.     

The role of the soil seed bank in vegetation recovery on an oceanic island severely damaged by introduced goats

Question: Are the seed banks of an isolated subtropical oceanic island capable of naturally regenerating vegetation either with species of the historical forest community or with the existing grassland community after severe damage to the vegetation by goats? Location: Nakoudojima Island, Bonin Archipelago (Ogasawara Shoto), Japan. Methods: Soil samples were collected at 0–5 cm and 5–10 cm depths from seven plots in forests, grasslands, artificially matted areas and bare land. Soil seed banks were assessed using the seedling emergence method followed by the hand‐sorting of ungerminated seeds. We determined the size and composition of the seed banks in upper soil layers of plots and compared the seed banks to the standing vegetation. Results: A total of 12 220 seedlings belonging to 42 species from 20 families germinated. Total mean seed density (0–5 cm depth) was low in all plots within forest, grassland, and heavily degraded vegetation types (34.7 ± 8.6 to 693.5 ± 123.6, 58.6 ± 7.8 to 107.1 ± 10.0, and 1.1 ± 0.5 to 7.2 ± 2.3 seeds/m², respectively). Forbs and graminoids dominated the seed banks of grassland and forest plots including Cyperus brevifolius, Gnaphalium pensylvanicum, Oxalis corniculata and Solanum nigrum, and these alien species comprised 90% of the density of the seed bank. There was little correlation between seed banks and standing vegetation of the island (Sørensen similarity coefficient values 0.26 to 0.45). Conclusions: If natural regeneration occurs from the seed bank of the island, future vegetation will not move toward the original forest community, because the seed bank is dominated by non‐native herbaceous grassland species. Though isolated, a few forest remnants with low species richness could be an important source for the natural re‐establishment of forest on the island; however, seed availability may be limited by either poor dispersal or pollination so that woody species will probably recover very slowly on this goat‐impacted island.     

Viable soil seed banks in disturbed lowland tropical rainforest sites in North Queensland

Four 0.125 m² soil samples, 50 mm deep, from each of eight tropical lowland rainforest sites ranging from primary and secondary forests to degraded Imperata grasslands were exposed to shade-house conditions. Germinations of soil seeds were monitored over 28 weeks in April-October. The viable seed banks in the primary forest (592 m^-2) and the secondary forest sites resulting from one cycle of clearing and abandonment (1300–1400 m^-2) were composed predominantly of secondary species. The viable seed banks were small in Imperata grassland sites that had been burnt regularly (200–400 m^-2) and very large (2000–3000 m^-2) in sites that had not been burnt for five years or more. The seed banks of the burnt and unburnt Imperata grassland sites were composed mostly of agricultural weeds whilst seeds of secondary rainforest trees were rare or absent. Similarities between the species composition of soil seed banks and regrowth vegetation types suggest that soil seed composition is important in determining the initial floristic composition following disturbance and that lack of seed of secondary rainforest trees is one factor which restricts re-growth and secondary succession on degraded Imperata grasslands. Species such as Eucalyptus intermedia R. T. Bak., E. pellita F. Muell. and Imperata cylindrica (L.)Beauv. which can establish and deflect the normal rainforest secondary succession were not present in the soil seed bank, and these species appear to rely on fresh seed inputs, lignotubers or rhizomes to colonize and regenerate after disturbance.     

云南拉沙山黑白仰鼻猴对种子传播潜力的评估

灵长类是森林生态系统中植物种子的主要传播者,有助于森林植被的更新,然而受研究方法的限制,灵长类种子传播潜力常被低估。为全面评估温带灵长类动物的种子传播潜力,采用直接观察法和粪便分析法评估珍稀濒危灵长类动物黑白仰鼻猴的种子传播潜力。于2018年11月—2019年10月采用直接观察法(瞬时扫描取样法)收集云岭省级自然保护区拉沙山黑白仰鼻猴的活动时间分配数据,获取每月取食果实的比例;同时每月收集黑白仰鼻猴的粪便,采用粪便分析法分拣猴粪中残留的植物种子,统计有完整种子残留的月份和粪便比例,应用这两种方法评估黑白仰鼻猴种子传播潜力及其差异。结果表明：直接观察法收集到黑白仰鼻猴取食果实的月份数为6个月(7—12月),月均取食果实的比例为(15.31±20.15)%,共取食13种果实;而粪便分析法发现黑白仰鼻猴粪粒内全年都有完整种子残留,粪便中月均完整种子残留比例(35.19±35.43)%,其中9月至第二年1月粪便中种子残留比例都大于50%,共取食18种果实;综合两种方法发现云南拉沙山黑白仰鼻猴共取食20种植物果实,具有较高的种子传播潜力。直接观察法可确定黑白仰鼻猴取食果实的物种数,而粪便分析法能...     

Persistent soil seed banks in old-growth Hyrcanian Box tree (Buxus hyrcana) stands in northern Iran

Little is known about the soil seed bank and the influence of plant communities on the interaction between the seed bank and aboveground vegetation in the Hyrcanian temperate deciduous forest. We surveyed species composition and diversity of the persistent soil seed bank and the aboveground vegetation in six community types in old-growth Hyrcanian Box tree (Buxus hyrcana) stands in northern Iran. Fifty-two species with an average of 3,808 seeds/spores m⁻² germinated; forbs accounted for 64% of the seed bank flora. Thirty-four species in the aboveground vegetation were not presented in the seed bank, 32 species in the seed bank were not found in the vegetation, and 20 species were in both. The dominant tree species were Diospyros lotus and Alnus subcordata with an average of 17 and 4.6 seeds m⁻², respectively. Our results suggest that (1) vernal geophytes and shade-tolerant perennials are not incorporated in the seed bank, (2) early successional species are well represented in the seed bank, (3) plant community type has significant impacts on seed bank densities, and seed bank richness and diversity were significantly related to presence/absence of Box tree in the aboveground vegetation. The persistent seed bank contained species that potentially have a negative impact on the regeneration of forests, thus forest managers should retain old-growth Hyrcanian Box tree stands to conserve disturbance-sensitive indicator forest species.     

Persistent soil seed bank and standing vegetation at a high alpine site in the central Chilean Andes

The persistent soil seed bank (viable seeds >1 year) and standing vegetation were investigated in the upper alpine belt (3250 m) in the Andes of central Chile, 33° S. Nine species (eight in standing vegetation) were found in a total persistent seed bank of 899 seeds m⁻². Seven additional species were represented by physically intact, non-viable seeds. Over 90% of the persistent seed bank was concentrated in Montiopsis sericea (Portulacaceae), Pozoa coriacea (Umbelliferae), Phacelia secunda (Hydrophyllaceae) and Oxalis compacta (Oxalidaceae). Examination of the seed/cover ratio revealed different propensities for persistent seed bank formation among species, and annuals formed persistent seed banks more frequently than perennial species. Abundance in the standing vegetation had predictive value for abundance in the persistent seed bank only when non-persistent seed bank species in the standing vegetation were discarded from the analysis. At the local scale, species diversity in the persistent seed bank and standing vegetation were correlated, but compositional similarity was low. Secondary down-slope dispersal promoted by frost heaving in combination with runoff, and life-form correlates are discussed as possible factors accounting for poor correspondence between the persistent seed bank and the standing vegetation at a local scale. The high Andean seed bank is similar to or larger than that reported for two Arctic tundra sites, but smaller than for a northern hemisphere subalpine site. If seed bank size is considered in relation to plant cover, the Andean seed bank greatly exceeds that of one Arctic site. Our study constitutes the first demonstration of a sizable persistent seed bank at an alpine site in the South American Andes and in southern hemisphere temperate mountains in general. Received: 26 May 1998 / Accepted: 21 November 1998     

Frugivory and Seed Dispersal by Brown Lemurs in a Malagasy Tropical Dry Forest

In the Ankarafantsika tropical dry forest (northwestern Madagascar), the common brown lemur (Eulemur fulvus fulvus) is the largest frugivore and probably the sole disperser of large‐seeded plants (seed diameter > 10 mm). To investigate seed dispersal by this primate, I recorded the feeding activities of a troop; also conducted fecal analyses, germination trials on defecated seeds, and a vegetation survey over 1 yr (beginning Dec 2006). Brown lemurs mostly consumed fruit (68%). The fruit of Vitex beraviensis was the most exploited resource (21% of total feeding time). Among dung samples, 1126 contained intact seeds of 70 plant species, with a median of six seeds and two species per sample. These data indicate that the brown lemur population dispersed approximately 9854 seeds/km²/d. Although the number of annually defecated seeds was overwhelmingly the largest in Grewia triflora, many of the small seeds were often clumped in dung piles. In contrast, large seeds of V. beraviensis occurred in the largest number of dung samples. The rate and time of seed germination in V. beraviensis were improved by passage through brown lemur guts. Therefore, V. beraviensis may readily establish seedlings in sites of brown lemur fecal deposition. Vitex beraviensis and brown lemurs are probably involved in a strong mutualism. Twenty‐three large‐seeded plants were probably dependent on brown lemurs for seed dispersal and some of these species were common trees in the forest. Maintenance of these key plant–animal interactions will probably contribute to the conservation of species diversity and intact regeneration of the Ankarafantsika forest.     

Persistent soil seed banks and floristic diversity in Fagus orientalis forest communities in the Hyrcanian vegetation region of Iran

We assessed the size of seed bank, species diversity and similarity between seed bank and standing vegetation in four oriental beech (Fagus orientalis Lipsky) community types of the central Hyrcanian forests of northern Iran. For this purpose a total of 52 relevés was established in two associations and two subassociations of the beech forests, and six soil samples (20 × 20 cm square and to a depth of 10 cm) were collected in each relevé in mid-spring, after the germination season had ended. Soil seed bank was investigated using the seedling emergence method. A total of 63 species, 57 genera and 36 families was represented in the persistent soil seed bank of the forest communities. The seed bank contained 28 species not found as adult plants in the vegetation, but these were mostly early successional species. Size of the seed bank ranged from 3740 to 4676 individuals m⁻² in the Rusco hyrcani-Fagetum orientalis and Danae racemosae-Fagetum orientalis associations, respectively. Species composition of seed banks and aboveground vegetation had low similarity with an average of 24.3% in the four plant communities, because only 38% of the species were the same in the vegetation and the seed banks. Most seeds in the seed bank were from early successional species, and the only tree with a large persistent seed bank was the fast-growing pioneer Alnus subcordata. DCA ordination also demonstrated low similarity between soil seed bank and vegetation. The soil seed banks of the four beech communities did not differ significantly in size, composition, diversity and uniformity. Although above ground vegetation in the four community types is floristically distinct, there is considerable overlap among the soil seed banks because they contain in a similar way early successional species. Further, the absence of typical forest species in the soil seed bank indicates that restoration of forest tree species cannot rely on the soil seed bank.