Linking carbon balance to establishment patterns: comparison of whitebark pine and Engelmann spruce seedlings along an herb cover exposure gradient at treeline

There is increasing evidence that landscape vegetation patterns near species?? range limits are associated with positive biotic interactions, such as in the alpine-treeline ecotone. In the northern Rocky Mountains, whitebark pine (Pinus albicaulis) is considered an early-successional species, able to establish in exposed microsites, while late-successional species such as Engelmann spruce (Picea engelmannii) are more dependent on neighboring vegetation to facilitate establishment. We compared ecophysiological traits associated with carbon balance of newly germinated seedlings of whitebark pine and Engelmann spruce along an herb cover gradient to (1) infer which ecophysiological properties explain the establishment success of seedlings, and (2) to assess differences in establishment patterns with respect to distance from neighboring vegetation. We measured survival over 2 years, and concurrently measured gas exchange and water relations (photosynthesis, respiration, and transpiration), morphology [specific leaf area (SLA)], and biochemistry [chlorophyll fluorescence (F _v /F _m) and nonstructural carbohydrates]. Both species initially established in the most exposed microsites away from vegetation during their first growing season, but only pine persisted in exposed microsites to the end of the second growing season. Pine exhibited phenotypic traits to increase stress tolerance (e.g., higher soluble sugar concentrations, lower SLA) and improve carbon balance (e.g., greater water use efficiency, lower respiration, higher F _v /F _m) compared to spruce in exposed sites, but had lower carbon balance under herb cover. Superior establishment success of pine in exposed microsites at treeline could thus be attributed to a suite of intrinsic physiological advantages that are apparent at the earliest stage of development.     

植物群落类型和围栏封育对高山林线岷江冷杉幼苗成活的影响

林线树种幼苗的生长紧密关系着高山林线的形成。通过室内发芽实验测定了岷江冷杉种子萌发能力。2015年春,在川西巴郎山岷江冷杉林线之上的高山林线交错带,通过播种处理(播种和不播种),研究了不同植物群落类型(灌木和草地群落)和围栏处理(围栏和不围栏)对岷江冷杉幼苗成活率的影响,以期为进一步探讨高山林线形成机制提供科学参考。结果表明:(1)岷江冷杉室内发芽率31.4%,成活率随时间下降。(2)灌木群落的全年日平均空气温湿度显著高于草本群落,而两群落的全年日平均土壤温度无显著差异。(3)未播种样地没有发现岷江冷杉幼苗,说明林线交错带缺乏岷江冷杉种子。(4)与不围栏样地相比,围栏封育分别显著提高了岷江冷杉幼苗成活率2.0%(2015年)和2.2%(2016年);与灌木群落相比,草本群落中岷江冷杉幼苗成活率显著提高0.8%(2015年)和1.2%(2016年),说明灌木对林线交错带幼苗更新具有更强的竞争作用。围栏处理下,草本群落中岷江冷杉幼苗成活率显著高于灌木群落;不围栏处理下,草本群落中岷江冷杉幼苗成活率显著低于灌木群落,2017年在不围栏样地没有幼苗存活,说明动物干扰对林线交错带森林更新有阻碍作用。研究表明,高山林线交错带森林更新,受种子、植物竞争和动物干扰(例如,放牧)等多方面因素的共同影响,可以通过播种、植物剔除(去除竞争)和围栏等促进幼苗更新和成活。     

Conifer seedling recruitment across a gradient from forest to alpine tundra: effects of species,provenance, and site

Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine–treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated.

Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA.

Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt.

Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings.

Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance – a factor that has rarely been considered in previous studies of tree recruitment and species range shifts.     

Effects of microsite light availability on the survival and growth of oak seedlings within a grassland

Effects of microsite light availability on the growth and survival of transplantedQuercus serrata Thunb. seedlings in aMiscanthus sinensis Anderss. grass canopy were investigated by a “plant's eye view” approach. Diffuse site factors, i.e., the fractional transmissions of diffuse photosynthetic photon flux density, were estimated at 15 cm aboveground in 108 microsites where the seedlings grew. Microsite diffuse site factors were significantly different between surviving and dead seedlings during the experiment period from April to October (F_[1,14]=10.9, P<0.01). Relative growth rate of dry weight for individual seedlings positively depended on the diffuse site factors (r²=0.482, P<0.001 in May; r²=0.312, P<0.001 in October). Only 16 seedlings produced their second stem flush within the grass canopy. The ratio of height to dry weight of the second stem flush was significantly higher for the seedlings grew in shady microsites than for those in less shady microsites (r²=0,471, P<0.01 in May). This study suggests that the microsite heterogeneity of light availability is one of the important factors affecting the establishment of tree seedlings in patchy grasslands.     

Whitebark pine facilitation at treeline: potential interactions for disruption by an invasive pathogen

In stressful environments, facilitation often aids plant establishment, but invasive plant pathogens may potentially disrupt these interactions. In many treeline communities in the northern Rocky Mountains of the U.S. and Canada, Pinus albicaulis, a stress‐tolerant pine, initiates tree islands at higher frequencies than other conifers – that is, leads to leeward tree establishment more frequently. The facilitation provided by a solitary (isolated) P. albicaulis leading to tree island initiation may be important for different life‐history stages for leeward conifers, but it is not known which life‐history stages are influenced and protection provided. However, P. albicaulis mortality from the non‐native pathogen Cronartium ribicola potentially disrupts these facilitative interactions, reducing tree island initiation. In two Rocky Mountain eastern slope study areas, we experimentally examined fundamental plant–plant interactions which might facilitate tree island formation: the protection offered by P. albicaulis to leeward seed and seedling life‐history stages, and to leeward krummholz conifers. In the latter case, we simulated mortality from C. ribicola for windward P. albicaulis to determine whether loss of P. albicaulis from C. ribicola impacts leeward conifers. Relative to other common solitary conifers at treeline, solitary P. albicaulis had higher abundance. More seeds germinated in leeward rock microsites than in conifer or exposed microsites, but the odds of cotyledon seedling survival during the growing season were highest in P. albicaulis microsites. Planted seedling survival was low among all microsites examined. Simulating death of windward P. albicaulis by C. ribicola reduced shoot growth of leeward trees. Loss of P. albicaulis to exotic disease may limit facilitation interactions and conifer community development at treeline and potentially impede upward movement as climate warms.     

Microsites for seedling establishment of subalpine conifers in a forest with moss-type undergrowth on Mt. Fuji, central Honshu, Japan

We examined microsites for the seedling establishment of Tsuga diversifolia, Abies veitchii and Abies mariesii in a subalpine coniferous forest with moss-type undergrowth dominated by Hylocomium splendens and Pleurozium schreberi on Mt. Fuji, a less-snowy mountain located on the Pacific Ocean side of the central part of Honshu Island, Japan, and compared these microsites with cases in other regions of Japan with different undergrowth vegetation (herb type, dwarf-bamboo type) under snowier conditions. L-shaped size structures with abundant seedlings and small trees were identified for all three species. Established seedlings of T. diversifolia showed remarkable preference for elevated microsites such as fallen logs and convex ground (i.e. ground raised around trunk bases), while A. veitchii and A. mariesii indicated no significant bias toward such elevated microsites in the large seedling stage. However, the relative frequency of T. diversifolia seedlings established on flat ground reached high values of 54–77%, since the relative occupation area of the flat ground was large (85–89%). Such a large share suggests that flat ground should be regarded as the most important microsite for the seedling establishment of T. diversifolia in forests with moss-type undergrowth. This preference trait for T. diversifolia observed with moss-type undergrowth has not been reported in previous studies examining forests with herb- and dwarf bamboo-type undergrowth, where T. diversifolia seedlings are scarce on the ground and restricted to elevated microsites. We conclude that the major microsite for T. diversifolia seedling establishment changes according to the undergrowth vegetation type: a large ground share for moss-type undergrowth and a small ground share for herb- and dwarf bamboo-type undergrowth.     

Difference in microhabitat-related regeneration patterns between two subalpine conifers, Tsuga diversifolia and Abies mariesii, on Mount Hayachine, northern Honshu, Japan

Seedling and tree-establishment microhabitats of Tsuga diversifolia and Abies mariesii were examined on the herb- and dwarf bamboo-dominated forest floor on Mount Hayachine, a mountain under intermediate snow conditions in northern Honshu, Japan. The four microsite types were fallen logs, buttresses, rocks and ground. The ground substratum was further divided into four subtypes by dominant undergrowth species: Lycopodium, Pteridophyllum, Carex and dwarf bamboo. The establishment of T. diversifolia seedlings on the ground was scarce, and depended mostly on non-ground microsites (i.e. fallen logs, buttresses and rocks). The seedling establishment of A. mariesii was not dependent on specific substrata, although on the ground, establishment sites were limited to the Lycopodium subtype situated on convex sites. Among the microhabitats for seedling establishment, larger trees of A. mariesii rarely occurred on higher portions of the non-ground microsites. In contrast, T. diversifolia could grow up to the height of canopy-layer trees in such microsites. Thus, non-ground microsites seem to be unsuitable for tree establishment in A. mariesii, and are probably useful microhabitats for regenerating T. diversifolia to avoid competition with A. mariesii. We also compared seedling-establishment microhabitats for the two conifers between Mount Hayachine and two other mountain regions under different undergrowth conditions (moss-dominated and dense dwarf bamboo-dominated). Our findings suggested that seedling recruitment in non-ground microsites was primarily determined by undergrowth conditions; T. diversifolia preferred such microsites where the moss-covered area was low, and A. mariesii preferred where dwarf bamboo-covered area was high.     

Seedling occurrence in alpine treeline conifers: A case study from the central Rocky Mountains,USA

Information concerning the occurrence of very young (1- to 10-year-old) tree seedlings in the alpine treeline ecotone is rare. Seedling occurrence of the dominant conifers Picea engelmannii and Abies lasiocarpa was measured in the treeline ecotone of the Medicine Bow Mountains, Wyoming (central Rocky Mountains, USA), an area composed of elongated tree islands separated by open meadows (ribbon forest) that grade into the closed forest. No seedlings were found on the windward sides of tree islands, while a mean of 0.6 seedlings.m^–2 occurred on the leeward (downward) sides. These values compared to the 4.2 seedlings.m^–2 in the closed forest. In addition, a strong correspondence was found between snowpack depth and seedling abundance, with depths that were either too shallow (< 0.5 m) or too deep (> 1.5 m) associated with fewer or no seedlings. A. lasiocarpa seedlings made up much less of the overall seedling population in the ribbon forest (6 %) than in the closed forest (22 %). Seedling establishment in this portion of the alpine treeline ecotone appears to be occurring at a low rate that differs between the two dominant species and may be strongly influenced by wind-driven snow accumulation patterns.     

Seedling establishment ofPolygonum cuspidatum andPolygonum weyrichii var.alpinum at high altitudes of Mt Fuji

Seedling establishment ofPolygonum cuspidatum (Seib. et Zucc.) andPolygonum weyrichii (F. Schmit) var.alpinum (Maxim.), dominant early-successional species in a volcanic gravel area on Mt Fuji, Japan, was compared for current-year seedlings at 2500 m a.s.l. The rate of survival in the first winter after germination at 2500 m was 2% and 24% forP. cuspidatum andP. weyrichii seedlings, respectively, and correlated with the upper distribution limit of each (2500–2600 m forP. cuspidatum; 3200–3300 m forP. weyrichii). The critical size for survival was 12 mg dry weight for overwintering current-year seedlings of both species. At 2500 m, the percentage of seedlings that attained the critical size for survival at the end of the first growing season was 19% and 85% forP. cuspidatum andP. weyrichii, respectively. Growth curves indicated that the larger-seeded characteristic ofP. weyrichii contributed to larger end-of-season size and resulted in higher rate of seedling survival at 2500 m, compared with smaller-seededP. cuspidatum. Current-year seedlings ofP. weyrichii, at least up to 3250 m, attained the first year annual growth critical for overwintering and therefore for successful seedling establishment, due to the large-seeded characteristic. At the top (3770 m) beyond the upper distribution limit ofP. weyrichii, most seedlings raised artificially failed to attain the critical size for survival in the first growing period due to the reduced growing period. Failure of seedling establishment would thus appear to restrict the distribution of the species at altitudes higher than 3300 m.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Limitations to seedling establishment in a mesic Hawaiian forest

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000–1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.     

The Restoration of Degraded Mountain Woodlands: Effects of Seed Provenance and Microsite Characteristics on Polylepis australis Seedling Survival and Growth in Central Argentina

South American high‐mountain ecosystems are greatly influenced by human disturbance. In the mountains of Córdoba, Argentina, Polylepis australis (Rosaceae) woodlands are currently highly fragmented and subject to extensive burning and livestock grazing, resulting in severe changes of habitat characteristics, which hamper natural regeneration. In order to find out how to achieve successful reforestation, we compared P. australis seedling survival and growth and the development of a shrubby habit for two seed provenances and different planting microsites. Survival of planted seedlings after 5 years was 70%, with most deaths (19%) in the first year and declining mortality with ongoing establishment. Survival did not show any relationship with seed provenance or microsite characteristics. Height growth averaged 34.6 ± 1.2 cm in 5 years. Seedlings produced from seeds collected in a well‐preserved woodland grew taller and showed a higher tendency for development of shrubby habit than those produced from seeds collected in a degraded woodland. Seedlings planted in more degraded microsites with exposed soil or rock due to past grazing pressure grew less and developed a more shrubby habit than those planted in better preserved microsites. Our results show that restoration of degraded areas with P. australis is possible and that there is potential to improve restoration success with a careful selection of seed provenance and planting microsites.     

Germination and early growth of <Emphasis Type="Italic">Ailanthus</Emphasis> and tulip poplar in three levels of forest disturbance

Increasing rates of forest disturbance may provide greater opportunity for invasion of nonnative species, thereby altering the successional trajectory of native plant communities. In the eastern U.S., invasive Ailanthus altissima and native Liriodendron tulipifera have similar life histories and niches and often co-occur. To examine how disturbance affects the establishment of these species, we performed field experiments to evaluate the response of sown seeds and transplanted seedlings to three levels of disturbance on north- and south-facing aspects. L.␣tulipifera germination was severely limited by low seed viability and had significantly lower germination than A. altissima in all sites. The effect of disturbance regime on A. altissima germination depended on aspect in the second growing season. In contrast, mean seedling survival, biomass, leaf area and leaf area ratio were greater for L. tulipifera in all field sites. Overall, the north-facing selective cut forest provided a disproportionately large number of suitable microsites for L. tulipifera establishment. Collectively, this study demonstrated that different timber harvest practices produce heterogeneous mosaics of suitable microsites for germination and establishment. Limited L. tulipifera germination may be a serious constraint to population establishment if seeds are deposited for the first time immediately after a disturbance event. However, if sufficient viable seeds of both species exist, L. tulipifera out-performs the invasive in the first two years following disturbance. This may explain why A. altissima has shown explosive population growth in a limited number of sites in the past century.     

Regeneration of beech (Fagus crenata) after the simultaneous death of undergrowing dwarf bamboo (Sasa kurilensis)

Seedling recruitment and survivorship of beech (Fagus crenata) were studied with special reference to the simultaneous death of undergrowing bamboo (Sasa kurilensis). The survival rate of beech seedlings on the floor whereSasa had withered was much higher than that on the floor whereSasa survived. Damping off caused the largest mortality among beech seedlings. However, the allocation pattern of matter to different parts of the seedlings indicated that their survival was greatly affected by production economy. The dense cover of dwarf bamboo prevented the establishment of beech seedling banks on the forest floor. The interval between the times when simultaneous death ofSasa occur and the length of its recovery period are thus important factors controlling the dynamics of beech forests in Japan.     

Aerial seed storage in <Emphasis Type="Italic">Melaleuca ericifolia</Emphasis> Sm. (Swamp Paperbark): environmental triggers for seed release

Aerial seed banks are potentially the main source of sexual recruitment for woody wetland plants. Whilst the importance of soil seed banks for the persistence and recruitment of wetland plants has been examined in many studies, the role of aerial seed banks has been largely neglected. We used seed traps and the seedling emergence technique to quantify the seed rain from aerial seed banks of the Swamp Paperbark Melaleuca ericifolia Sm. (Myrtaceae) in Dowd Morass, a Ramsar-listed, brackish-water wetland in south-eastern Australia. Nine plant species germinated from material collected in seed traps over 2004–2005, but emergents were dominated (80–97%) by M. ericifolia. The mean number of M. ericifolia emergents ranged from <1 to <100 seedlings m⁻² day⁻¹, and showed a peak in the summer–autumn period. Regression analysis showed a significant negative correlation (r ² = 0.738) between the number of M. ericifolia emergents and water depth. Water depth and salinity were negatively correlated (r ² = 0.819), and increases in the number of M. ericifolia emergents as water levels fell were also associated with high salinities. Increasing air temperature and vapour pressure deficit also stimulated seed release during periods of drying. This study is one of the first to demonstrate the importance of aerial seed banks for sexual recruitment in woody wetland plants and the release of seed in relation to environmental factors. Aerial seed banks warrant consideration alongside soil seed banks for the establishment and long-term survival of woody plants in wetlands. Handling editor: Luis Mauricio Bini     

Comparative recruitment patterns of two non-pioneer canopy tree species in French Guiana

Summary A comparative study was conducted on the recruitment patterns of two non-pioneer tree species, one dispersed by arboreal mammals and birds (Virola michelii, Myristicaceae) and the other by rodents (Moronobea coccinea, Clusiaceae). These species differ in fruiting phenology, seed size, dispersal distance, germination time and seed nutrient exhaustion. In both species, establishment patterns were consistent with the escape hypothesis and the Janzen-Connell model. Virola seeds need not be buried to survive and germinate, and may produce a seedling carpet beneath the parent. Moronobea seedlings only establish from seeds buried by scatterhoarding rodents in the surrounding understory. One-year survival of Virola seedlings was 47.8% and was greater >10 m than < 10 m from the largest parent tree. In contrast, survival of Moronobea seedlings was 56% 3 years after seed dispersal. Survival of juveniles was greater in gaps than in the understory for Virola but not for Moronobea. Moronobea survival was greater than Virola survival in both microhabitats. Both species establish in the understory, yet both grew faster in gaps. Virola appeared to be more gap-dependent than Moronobea which may persist several years in the understory until a gap occurs. Virola and Moronobea illustrate two intermediate recruitment patterns along an hypothetical continuum of nonpioneer species replacement (Bazzaz and Pickett 1980; Swaine and Whitmore 1988).     

Roles of cottony hairs in directed seed dispersal in riparian willows

Willows usually establish on wet substrates with fine sediments at sites that are created by large disturbances, but suitable microsites are spatially and temporally limited. Thus, we hypothesized that willow seeds are selectively dispersed to suitable microsites, such as those with a wet substrate, rather than unsuitable microsites, such as those with a dry substrate, with seedling establishment mediated by the cottony hairs attached to seeds (directed dispersal). To test our hypothesis, we compared several recruitment-related traits, including buoyancy, germination, and trapping at favorable microsites, in seeds of the riparian willows Salix sachalinensis and S. integra with and without cottony hairs in laboratory and field experiments. In both field and laboratory experiments, more seeds with cottony hairs were trapped in water and wet sand than in dry sand, in which no seeds of either species germinated. These results indicate that cottony hairs facilitate the recruitment of seeds to microsites favorable for seed germination and help seeds avoid unfavorable microsites. On the water surface, 17.6% of S. sachalinensis seeds and 68.0% S. integra seeds with cottony hairs floated for more than 6 days, whereas all seeds without cottony hairs sank immediately after being placed on the water surface. These results suggest that cottony hairs facilitate long-distance dispersal via flowing water and also help avoid germination under water, where willow seedlings fail to establish. Seeds of the two willow species were released from the cottony hairs and germinated immediately after the seeds were placed on wet sand, but not after placement on water or dry sand. These results suggest that the seeds are released from the cottony hairs when the hairs become wet and the seeds are striking to a suitable microsite for seedling establishment, such as wet sand. In riparian willows, the cottony hairs promote directed dispersal by moving seeds to discrete and predictable microsites where the seedling establishment is disproportionately high.     

Arbuscular mycorrhizal fungi increased early growth of two nontimber forest product species Dyera polyphylla and Aquilaria filaria under greenhouse conditions

Nontimber forest products (NTFPs) represent an important source of income to millions of people in tropical forest regions, but some NTFP species have decreased in number and become endangered due to overexploitation. There is increasing concern that the planting stocks of Dyera polyphylla and Aquilaria filaria are not sufficient to sustain the yield of NTFPs and promote forest conservation. The objective of this study was to determine the effect of two arbuscular mycorrhizal (AM) fungi, Glomus clarum and Gigaspora decipiens, on the early growth of two NTFP species, D. polyphylla and A. filaria, under greenhouse conditions. The seedlings of both species were inoculated with G. clarum or G. decipiens, or uninoculated (control) under greenhouse conditions. Percentage of AM colonization, plant growth, survival rate, and nitrogen (N) and phosphorus (P) concentrations were measured after 180 days of growth. The percentage of AM colonization of D. polyphylla and A. filaria ranged from 87 to 93% and from 22 to 39%, respectively. Colonization by G. clarum and G. decipiens increased plant height, diameter, and shoot and root dry weights. Shoot N and P concentrations of the seedlings were increased by AM colonization by as much as 70–153% and 135–360%, respectively. Survival rates were higher in the AM-colonized seedlings at 180 days after transplantation than in the control seedlings. The results suggest that AM fungi can accelerate the establishment of the planting stocks of D. polyphylla and A. filaria, thereby promoting their conservation ecologically and sustaining the production of these NTFPs economically.     

Seed-caching responses to substrate and rock cover by two <Emphasis Type="Italic">Peromyscus</Emphasis> species: implications for pinyon pine establishment

We examined whether pinyon mice (Peromyscus truei) and brush mice (P. boylii) could act as directed dispersal agents of pinyon pine (Pinus edulis) through differential responses to soil particle size and rock cover. In field experiments, we allowed mice to either cache pinyon seeds or recover artificially cached seeds (pilfer) from quadrats containing large- or small-particle soils. Both species placed most (70%) seed caches in small-particle soil. Pilfering was the same from both particle sizes in the first year, while more seeds were pilfered from large-particle soils in the second year. In separate experiments, rock cover interacted with soil particle size, with both species placing over 50% of their caches in small-particle soil with rock cover. Overall, we found greater seed-caching in small-particle soils near rocks, with equal or lower pilfering from small-particle soils, suggesting more seeds would survive in small-particle soils near rock cover. Three lines of evidence supported the hypothesis that mice could act as directed dispersers by moving pinyon seeds to beneficial microsites for germination and establishment. First, in greenhouse experiments, pinyon seed germination was 4 times greater in small-particle soil cores than in large-particle soil cores. Second, soils near rocks had significantly higher water content than areas of open soil at the driest time of the year, a critical factor for seedling survival in the arid southwestern USA. Third, 75% of juvenile pinyon trees were growing in small-particle soils, and 45% were growing near rock nurses.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.