Impact of vertebrate acorn- and seedling-predators on a Mediterranean Quercus pyrenaica forest

We have experimentally investigated the impact of biotic factors, acting at the seed and seedling stages, on a Quercus pyrenaica forest in the Sierra Nevada mountains (SE Spain). We monitored the natural establishment of the oak for 3 years in two forest plots and two shrubland plots, by counting seedlings and juveniles. In addition, we established several experiments in these plots to examine acorn and seedling survival, while also considering the microhabitat effect on survival probability. Dispersed acorns were quickly consumed by several species of predators, particularly wild boar (Sus scrofa) and woodmouse (Apodemus sylvaticus). Less than 4% of the experimental acorns survived to produce seedlings, even when they were buried 4 cm in soil, simulating caches. No effect of microhabitat was found on predation, and thus no safe site appears to exist for Q. pyrenaica acorns in the study area. Some 98% of the 1000 experimental seedlings were killed by herbivores, notably woodmice, wild boar, and domestic and wild ungulates. Seedling survival varied spatially, being significantly higher under shrubs (4%) than in any other microhabitat (less than 0.5%). Both acorn and seedling survival were much lower in the shrublands than in the forests. In shrubland plots, the main agent of seedling mortality was trampling by domestic ungulates foraging in herds. This study suggests that the regeneration of Q. pyrenaica forests in Mediterranean mountains can be limited by herbivores acting at several life-history stages.     

Germination and seedling growth of Quercus (section Erythrobalanus) across openings in a mixed-deciduous forest of southern New England, USA

In this study three species of the genus Quercus section Erythrobalanus (Quercus coccinea, Quercus rubra, Quercus velutina) were investigated. All occur together as canopy trees in forests of southern New England. Acorns of each Quercus species were planted in plots located in five zones that represent a range of forest gap/canopy conditions that can occur within a southern New England forest. These five zones were demarcated adjacent to and across large openings of two physiographic sites—valley and ridgetop. Experiments were designed to monitor germination and initial growth of seedlings for the first three growing seasons. During the start of the first growing season germination was monitored. At the end of the first growing season measurements of height and number of flushes were taken and destructive samples of seedlings made for dry mass of root, stem and leaves. At the end of the third growing season height was recorded for surviving seedlings. Comparisons were made of germination and growth of seedlings located in the different gap/canopy conditions.

Results demonstrated clear differences in patterns of germination and early growth among species and among gap/canopy conditions of the sites. All species showed an increased lag in germination with reduced amounts of light. Highest growth and flushing rate were in the center conditions of the openings for all species during the first growing season. Quercus rubra had the greatest height growth the first growing season but a lower number of flushes than Q. velutina and Q. coccinea. In comparison with the other species, Q. rubra had the greatest total dry mass in most gap/canopy conditions after the first growing season. However, Q. velutina had the greatest total dry mass in the center of the ridgetop opening. In almost all gap/canopy conditions Q. velutina had greater proportions of dry mass allocated to roots compared with the other species.

After 3 years, greatest height growth in any of the gap/canopy conditions was recorded for all three species in the center of the valley site. Under this condition Q. rubra had significantly greater growth than Q. velutina and Q. coccinea. Quercus rubra also had significantly greater height growth and survival beneath the canopy conditions of the valley site than the other species. On the ridgetop site regeneration failed to establish beneath canopy conditions that provided low amounts of light. Quercus velutina showed greatest height growth after three years in the center and edge conditions of the ridgetop opening compared with the other species. Environmental influences that determine species germination and growth performance are suggested.     

Dispersal of Quercus mongolica acorns in a broadleaved deciduous forest

Effects of two sets of factors on the disappearance of acorns of Q. mongolica var. grosserrata were investigated. Acorns were distributed on (a) the floor of a forest with its canopy intact, (b) the floor of a forest clearing, (c) the vegetated sloping bank of a forest road, and (d) a bare forest road. They disappeared fastest from the forest with intact canopy and slowest from the bare road, an effect thought to be directly related to amounts of cover.When the acorns were covered with frames having netting with different-sized apertures, a mesh of 3.0×4.0 cm did not appreciably alter the rate of acorn disappearance compared with the uncovered controls: a mesh of 0.5×0.5 cm totally inhibited the disappearance of acorns.Five species of small mammal were live-trapped. Of these, Clethrionomys rufocanus, Apodemus argentinus and A. speciosus are thought to be implicated in the disappearance of acorns. The rate of acorn disappearance seems to be directly related to total numbers of small mammals.Acorns disappeared more rapidly when distributed on the ground than when buried.     

The effect of weevil and fungal attacks on the germination ofQuercus robur acorns

Oak,Quercus robur, acorns are sometines attacked by a weevil or infested by fungi (Murray, 1974). The number of acorns produced varies greatly from year to year. Acorns produced in the years between mast years may escape these weevil or fungal attacks, be dispersed and germinate.

Acorns were collected annually from 1983 to 1989 from five oak trees in southern Sweden. The percentage of acorns attacked by a weevil or fungi, or both, were monitored and the acorns' ability to germinate was tested.

Not all acorns were damaged in low crop years; the proportion of undamaged acorns varied between 33 and 69%. The proportion of weevil-attacked acorns differed significantly between the years. The proportion of acorns that were able to germinate in each category did not vary significantly between years. The germination ability was significantly higher in undamaged or fungi-infested acorns.     

Seed dispersal by small rodents favors oak over pine regeneration in the pine-oak forests of the Qinling mountains,China

Abstract

Natural regeneration differences between oak and pine are common in the Qinling Mountains of central China. Thus, we examined whether this difference could be explained by seed dispersal. We investigated the seed dispersal of Quercus aliena var. acuteserrata Maxim. and Pinus tabulaeformis Carr. and the regeneration characteristics of both species were investigated in the pine-oak forest belt (oak stands, pine-oak stands, and pine stands) of Huoditang Forest in the Qinling Mountains in central China. We found that Q. aliena var. acuteserrata acorns were usually scatterhoarded, while seeds of P. tabulaeformis were generally consumed immediately. We found that 55% of the released acorns were transferred to the P. tabulaeformis stands. Moreover, Q. aliena var. acuteserrata regenerated successfully and was recruited in the three forest stands. However, the natural regeneration of P. tabulaeformis was very low in all stands because of high predation by small rodents. We propose, therefore, that the seed dispersal advantage of Q. aliena var. acuteserrata compared with P. tabulaeformis contributed greatly to the regeneration difference between these two species in the pine-oak forest belt of the Qinling Mountains in China.     

Edge Effect on Acorn Removal and Oak Seedling Survival in Mexican Lower Montane Forest Fragments

Vegetation structure of forest edges and type of adjacent field can affect seed and seedling survival patterns of tree species. We investigated acorn removal and seedling survival of Quercus germana and Q. xalapensis in relation to woody plant density across old field-forest gradients with soft and abrupt edges. Experiments were established along four parallel bands located at 40–50 m in the forest, forest and field edges (0–10 m to each side of the border), and 20–30 m in the old field. Within each band, woody plant and acorn density was measured, and four points for acorn and seedling removal experiments were randomly positioned. In each position, 20 acorns or four seedlings were placed. Survival was monitored during one month. In abrupt edges, acorn density was higher along edges than in forest interior. Also, higher acorn removal and seedling mortality were observed in the adjacent old-field whereas acorn and seedling survival was the highest at the edges. Acorn and seedling survival was positively correlated to woody vegetation density. As the vegetation density of field and forest edges grows, abrupt edges develop into soft edges, and they become more hospitable to oak recruitment and then to forest cover expansion.     

Structure and regeneration dynamics of dominant tree species along altitudinal gradient in a dry valley slopes of the Bhutan Himalaya

The aim of our study was to characterize the structure and regeneration dynamics of dominant tree species along altitudinal gradient in dry valley slopes of the Bhutan Himalaya. In the typical dry valley slopes of west-central Bhutan, we analyzed forest stratification, tree crown projection, and population structure from lower dry valley bottom (1520 m a.s.l.) to upper humid ridge top (3370 m a.s.l.) across five major forest types: i.e. (1) dry pine forest with Pinus roxburghii as a dominant species; (2) mixed broad-leaved forest with Quercus lanata (evergreen), Quercus griffithii, (deciduous), and Rhododendron arboreum (evergreen) as major canopy species; (3) evergreen broad-leaved forest dominated by Quercus semecarpifolia, Quercus glauca and Quercus oxyodon; (4) cool conifer forest dominated by Abies densa and Tsuga dumosa; (5) cold conifer forest with A. densa and Juniperus recurva as dominants. In general, regeneration pattern of major dominant species shifted from inverse-J (lower altitudes), to sporadic (mid-altitudes), and to uni-modal type (upper altitudes) corresponding to three regeneration trends: (1) invasive at the lower, warm, dry forest under relatively strong human disturbances; (2) stable/balanced at the mid-altitude, in a relatively stable, mature moist evergreen broad-leaved forest with gap regeneration; and (3) poor/low regeneration at the upper, cool, humid conifer forest with a continuously cattle-grazed understory. Overall, regeneration patterns were balanced in all forest types, however care should be taken to prevent excessive exploitation of dry pine forest of lower valley bottom, and to control cattle grazing at higher altitudes. Our findings can be incorporated into management plans for sustainable management and conservation of mountain forests.     

Edge type effect on germination of oak tree species in the Highlands of Chiapas,Mexico

The purpose of this study was to examine the effect of habitat edges on the probability of acorn germination of three oak species (Quercus crassifolia Humb. and Bonpl., Quercus rugosa Née and Quercus laurina Humb. and Bonpl.). The effects of edge type (hard and soft), habitat type (grassland, edges and forest) and leaf litter cover (covered or uncovered acorns) on acorn germination was evaluated by the experimental establishment of acorns along transects crossing habitat edges. More acorns developed into seedlings in grasslands (38%) than in the forest edge (18%) or the forest interior (15%). In sites with soft edges, a higher number of seedlings emerged from acorns covered by litter compared with acorns sowed in the adjacent forest edge and forest interior (P < 0.05). In sites with hard edges, fewer seedlings emerged in the edge (14%) compared with the adjacent grassland (38%), and the adjacent forest (20%) presented intermediate values. However, in sites with soft edges significant differences in seedling emergence were recorded between the grassland (38%) and the forest (10%), whereas the edge presented intermediate values (23%). The effect of leaf litter cover on acorn germination was only significant in grasslands in sites with soft edges (P < 0.05). Acorns in grasslands received relatively little insect damage (10%) compared with the edge (19%) and the forest (30%, P < 0.05), emphasising the importance of acorn dispersal for successful oak establishment. The implications for regeneration of these oak species and the dynamics of montane oak forests in Mexico are discussed.     

Effect of drought on diameter increment of Quercus ilex, Phillyrea latifolia, and Arbutus unedo in a holm oak forest of NE Spain

The present study was carried out to elucidate the drought growth responses of Quercus ilex L., Phillyrea latifolia L., Arbutus unedo L., and other accompanying woody species of the Mediterranean holm oak forest. We submitted holm oak forest stands in Prades mountains (NE Spain) to a 2-year experimental drought. We reduced soil water availability about 15% by plastic strips and funnels that partially excluded rain throughfall and by ditch interception of water runoff. Mean stem diameter increment showed a great variation depending on the species. A. unedo had larger growth rates than Q. ilex and P. latifolia, but it was also the species that experimented the highest growth reduction in the drought plots (77%), suggesting a higher drought sensitivity than Q. ilex (55%) and P. latifolia (no drought effect). The growth reduction was specially marked in the larger trees. Aboveground stand biomass increment, estimated from stem diameter by allometric relationships, was 1.9 Mg ha⁻¹ per year in the control plots. The 15% reduction in the upper soil moisture produced 42% reduction in this biomass increment. In the drier conditions predicted in this Mediterranean area in the frame of climate change, an important reduction of growth rates can be hence expected, accompanied by a gain of dominance of drought-tolerant species such as P. latifolia in detriment of more mesic species such as Q. ilex.     

The increasing scarcity of red oaks in Mississippi River floodplain forests: Influence of the residual overstory

Red oaks – cherrybark oak (Quercus pagoda Raf.), willow oak (Quercus phellos L.), water oak (Quercus nigra L.), and Nuttall oak (Quercus texana Buckley; aka: Quercus nuttallii Palmer) – are not regrowing in Mississippi Delta river floodplain forests in the southeastern United States in sufficient numbers to sustain the former species composition and timber and wildlife values. Even if vigorous red oak reproduction becomes established, partial harvesting that does not remove the taller trees will suppress understory red oak height growth more than it will suppress height growth of such other species as sugarberry (Celtis laevigata Willd.), American elm (Ulmus americana L.), cedar elm (Ulmus crassifolia Nutt.), swamp dogwood (Cornus foemina Mill.), green ash (Fraxinus pennsylvanica Marshall), and sweetgum (Liquidambar styraciflua L.). Consequently, the red oaks in these partially harvested stands become increasingly suppressed and probably die; and there is a shift in species composition to the other species. In addition to ensuring vigorous oak reproduction, silvicultural clearcutting or rapid removal of the residual trees following shelterwood or seed tree harvesting to provide full sunlight is needed to ensure red oaks become a dominant part of these future river floodplain stands.     

Distribution patterns of acorns after primary dispersion in a fragmented oak forest and their consequences on predators and dispersers

Forest fragmentation affects seed production, but little is known about how seeds are distributed in fragmented forests after primary dispersion and how this affects their interactions with other organisms. This study addressed these issues in a fragmented oak forest of Quercus laeta where four habitat types were recognized: interior and edge of patches, deforested matrix around patches, and an abandoned field. Areas and distances to the abandoned field were measured for 40 patches, and acorns were sampled in all habitats. Only samples from the interior and edge of patches had acorns, which were classified as aborted, viable, and parasitized by insects or fungi. Acorns in all these classes were regressed against patch area and distance to the abandoned field. Field experiments were also conducted to determine whether acorn removal rates varied across habitats. Aborted acorns within patches were not related to area or distance, but they decreased with increasing values of both variables at patch edges. Within patches, viable acorns were positively related to increases in area and distance, while the converse occurred for insect-damaged acorns; fungal-infested acorns were not related to patch variables in this habitat. Complex interactions between area and distance explained how acorns in these classes were distributed across patch edges. Acorn removal was higher within patches and their edges when compared to human-disturbed habitats; both mice and birds were identified as potential acorn dispersers. These results suggest that forest fragmentation affects acorn-mediated processes and that this may influence the recovery of patch connectivity in the landscape.     

章古台固沙林更新造林中天然更新的结合与利用

辽宁省章古台固沙林经过经营疏伐与块状采伐以后,通风透光条件改善,促进了天然下种更新。天然更新强度,在采伐迹地为690株·hm^-2,人工更新造林地为575株·hm^-2,油松和赤松林下松树为181株·hm^-2,其他树种295株·hm^-2。通过采取保护1年生樟子松幼树越冬,抚育油松、赤松幼树等措施,促进针叶树天然更新。在更新造林中,沙丘下部与丘间低地营造杨树,利用行间天然更新树木形成阔叶树混交林,沙丘中上部营造针叶树,利用行间天然更新灌木,形成乔灌混交林。     

Dispersal of Quercus mongolica acorns in a broadleaved deciduous forest 2. Scatterhoarding by mice

Acorn transporting and hoarding, distribution, and utilization by mice of hoarded acorns were studied. Apodemus speciosus and A. argentus were observed to transport acorns. An individual A. speciosus observed scatterhoarded at least 49% of acorns transported. The rate of acorn utilization was higher in locations where two or more acorns were buried than where a single acorn was buried. Forty-seven percent of scatterhoarded acorns disappeared before snow cover and 99% before the next spring. Based on the extent of home ranges for mice, dispersal distance is presumed to be from 30 to 40 m.     

立地因子和林分因子对牡丹江地区蒙古栎林橡实产量的影响

通过调查牡丹江地区的26块蒙古栎林标准地橡实产量,研究立地因子和林分因子对蒙古栎林橡实产量影响的结果表明,立地因子和林分因子对蒙古栎橡实产量影响不同,按影响大小排序为:密度>胸径>坡度>海拔>树高>树龄>土壤厚度>坡位>坡向>郁闭度,处于低海拔、阳坡半阳坡的低密度中龄林橡实产量较高。     

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factors controlling arboreal seed removal in oak-beech forests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.     

Pre- and post-dispersal damage to the acorns of two oak species (Quercus serrata Thunb. andQ. mongolica Fisher) in a species-rich deciduous forest

A comparative study of pre- and post-dispersal damage to the acrons of two oaks,Quercus serrata Thunb. andQ. mongolica Fischer, was done over 3 years in a species-rich deciduous forest in central Japan. The numbers of acorns bored into by moth larvae and byKobuzo rectirostris (Roelofs) was low and relatively uniform each year for both oak species, but the numbers bored into byCurculio spp. varied from year to year. The proportions of acorns attacked were independent of the density of acorns produced for both oak species. The proportions of the dispersed acorns of both oak species on the forest floor that disappeared between autumn and spring were less than 27%, and were lower for species in the stand that belong to family Fagaceae. However, by the next summer, most dispersed sound acorns had been infested by moths that specialized on post-dispersal acorns.     

Effects of silvicultural disturbance on acorn infestation and removal

Oak (Quercus spp.) regeneration is an important forest management goal in the central hardwood forest region of the United States. Silvicultural methods that target oak regeneration, such as the creation of canopy openings and removal of shade-tolerant midstory trees, are complicated by the impact of pre- and post-dispersal predators of oak acorns like acorn weevils (Curculio spp.) and granivorous small mammals. Understanding the effects of forest disturbance created by silviculture on these predators is important to promote successful oak regeneration. We conducted a 6-year study of acorn production, weevil infestation, and acorn removal at 113 black (Q. velutina) and white oak (Q. alba) trees impacted by three types of silvicultural treatments: adjacent to harvest openings, midstory removal (first-stage shelterwood harvest), and control (no harvest) in south-central Indiana, USA. We observed highly variable acorn production across years, but minimal impact of silvicultural treatments. Levels of weevil infestation and acorn removal by small mammal seed predators also varied greatly, and predation pressure was highest in years when acorns were scarce. Weevil infestation was reduced following midstory removal in shelterwood harvests, but probability of acorn removal by small mammals was unchanged following harvest. Damaged, germinated, and weevil-infested acorns were less likely to be removed by seed predators, suggesting additive effects of pre- and post-dispersal predators. This study emphasizes the importance of considering acorn predators in oak regeneration silviculture, and timing harvests to follow large acorn crops in order to reduce predation pressure and generate a high number of seedlings.     

Effects of flood duration and season on germination of black,cherrybark, northern red,and water oak acorns

Effects of flood duration (0, 10, 20, and 30 days) and season (winter and spring) on acorn germination were tested for two upland oaks [black and northern red oak (Quercus velutina Lam. and Q. rubra L.)] and two bottomland oaks [cherrybark and water oak (Q. pagoda Raf. and Q. nigra L.)]. Acorns were stratified for 30 days before flooding at a depth of 15 cm along the edge of a small pond. After flooding, acorns were sowed in sand-filled plastic cups and germinated for 40 days. Flood duration and season strongly affected radicle and epicotyl emergence of the upland oaks, but effects were generally limited to spring flooding. Embryo axes of the upland oak acorns were severely damaged with as little as 10 days of spring flooding. Almost no epicotyls developed, but radicles developed from the connective tissues between embryo axes and the cotyledons of many acorns. Spring flooding also significantly increased the percentage of decayed acorns for the upland oaks. In contrast, germination of the bottomland oaks was slightly improved by flooding during both seasons. Results demonstrated that the effects of flooding on the distribution of species within bottomlands can begin with seed storage and germination.     

栓皮栎种子虫蛀率的年际变化及其对幼苗发育的影响

为了解栓皮栎种子虫蛀率的年际变化,及虫蛀率对林木种子出苗率及幼苗生长的影响,于2010年和2011年秋季,在国有济源市愚公林场,收集成熟栓皮栎种子,调查两年的虫蛀率;于2010年9月,分别种植完好和虫蛀的栓皮栎种子各50枚,定期观察记录其发芽和生长状况。结果表明:1)栓皮栎的虫蛀率在2010年为47%,在2011为50%,二者差异不显著;2)虫蛀栓皮栎种子的发芽率为52%,完好栓皮栎种子的发芽率为88%,二者差异较为显著;3)虫蛀栓皮栎苗的生物量为1 968±400 mg,而完好栓皮栎苗的生物量为2 920±300 mg,二者差异较为显著。由此可见,栓皮栎种子虫蛀率的年际变化不大,但虫蛀对种子的发芽和出苗率有直接影响,因此,在采用种子直播造林时,应尽可能选择完好种子,以减少虫蛀对建苗的影响,提高造林成活率。     

Changes in the stand structure of a pine forest after rapid growth of Quercus serrata Thunb.

We studied the species composition and stand structure of secondary pine forests affected by pine wilt disease, caused by epidemic attacks of the nematode Bursaphelenchus xylophilus, in western Japan. Based on the species composition, the vegetation was divided into four types: (1) Cladonia rangiferina, (2) Wikstroemia sikokiana, (3) Quercus glauca, and (4) Gleichenia japonica. The sequence of types from (1) to (3) represents a successional series in accordance with topographical position. The relative basal areas of Rhododendron reticulatum and Juniperus rigida in the C. rangiferina and W. sikokiana types were larger than in the Q. glauca type. The relative basal areas of Q. serrata, Clethra barbinervis, and Eurya japonica in the C. rangiferina and W. sikokiana types were less than in the Q. glauca type. The mortality of light-demanding trees was higher in the Q. glauca type than in the C. rangiferina and W. sikokiana types. In stands that were severely affected by pine wilt disease, light was reduced and soil became moist after rapid growth of Q. serrata and other species which are not affected by the disease. The numbers of species and components of evergreen broad-leaved natural forests were decreased, because pine forests in the study area had been cut frequently and the land had degraded. Pine wilt disease has shifted the dominance of pine forests from Pinus densiflora to deciduous oaks such as Q. serrata and physical conditions seemed to provide a safe site for shade-tolerant plant species, which are components of evergreen broad-leaved forests. Moreover, the seed dispersal of evergreen oaks appears to be an important factor in the succession to evergreen broad-leaved forest in degraded areas.