Benefits of Using Shrubs as Nurse Plants for Reforestation in Mediterranean Mountains: A 4-Year Study

Shrubs are commonly considered competitors of planted seedlings in reforestation programs. However, shrubs can facilitate the establishment of understory seedlings in environments that, like Mediterranean‐type ecosystems, are characterized by harsh environmental conditions. In 1997, an experiment was set up in the Sierra Nevada mountains (southeast Spain) to test the use of shrubs as nurse plants for an alternative reforestation technique. Two‐year‐old seedlings of Pinus sylvestris and Pinus nigra were planted in four microhabitats: (1) open interspaces without vegetation (which is the usual method employed in reforestation programs), (2) under individuals of the shrub Salvia lavandulifolia, (3) under the north side of spiny shrubs, and (4) under the south side of spiny shrubs. Seedlings were also distributed in plots with and without ungulates to test the effect of herbivore damage. We report here the results of survival and growth after four growing seasons, a time span long enough to draw robust conclusions concerning the suitability of this technique. Pine survival was remarkably higher when planted under individuals of S. lavandulifolia as compared with open areas (2.6 times for P. sylvestris and 1.8 for P. nigra). The survival of both pine species was also higher when planted on the north side of spiny shrubs, while mortality on the south side was similar to that found in open areas. The reduction of solar radiation by the canopy of shrubs was likely the main factor determining shrub facilitation. The growth of the pines differed among years. However, growth was not inhibited when planted with shrubs as compared with open areas in any of the years. Herbivore damage was low but was mostly concentrated in the leader shoot, exacerbating the deleterious effect of ungulate herbivores on pine growth. We conclude that the use of shrubs as nurse plants for reforestation is a viable technique to increase establishment success of reforestation in Mediterranean‐type ecosystems and that it might be similarly useful in other water‐stressed environments. In addition, this technique offers the advantage of following natural succession, thus minimizing the impact in the community.     

Restoration of Dry Afromontane Forest Using Pioneer Shrubs as Nurse-Plants for Olea europaea ssp. cuspidata

Shrubs are often considered competitive barriers for seedlings planted in reforestation programs, although they can facilitate tree recruitment, especially in ecosystems under high abiotic stress. An alternative reforestation technique using pioneer shrubs as nurse‐plants for Olea europaea ssp. cuspidata was tested in exclosures in northern Ethiopia. Seedlings were planted in three different microhabitats, and their survival was monitored. The microhabitats were bare soil patches between shrubs, patches under the dominant shrub Acacia etbaica, and patches under Euclea racemosa, an evergreen shrub, which supports the majority of naturally established Olea recruits. The ability of shrubs to offer protection against browsing goats was tested experimentally. Controlled shading was used to determine whether solar irradiation causes seedling mortality in environments without water stress. Data were analyzed using Kaplan–Meier survival analysis, Kruskal–Wallis analysis of variance (ANOVA), and one‐way ANOVA. Olea survival was significantly higher and shoot damage by goats was lower when planted under shrub cover compared to bare soil patches, particularly under Euclea canopies, although high shade levels reduced seedling performance. Reduction of solar radiation by shrub canopies and thus control of soil–water evaporation and seedling transpiration most likely controlled the observed facilitation. Planting under shrubs may increase seedling survival and assist regeneration of dry Afromontane vegetation. Preserving pioneers also reduces soil erosion and conserves biodiversity. Excluding livestock is essential for Olea woodland restoration and allows persistent but morphologically modified Olea shrubs to develop vigorous regrowth. Facilitative processes are guiding principles for assisted forest restoration, but above‐average rains may be critical to restore higher biomass levels in semiarid areas.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.     

Effects of pioneer shrubs on the recruitment of the fleshy‐fruited tree Olea europaea ssp. cuspidata in Afromontane savanna

Question: Is seedling recruitment of a fleshy‐fruited tree in degraded Afromontane savanna dependent on shelter from pioneer shrubs, and is shelter availability related to shrub traits? Location: Degraded montane savanna in northern Ethiopia (13°36′N, 39°21′E). Method: Nurse plants of Olea europaea ssp. cuspidata seedlings were recorded using T‐square plotless sampling and clustered according to shrub traits, using Ward's method after Principal Components Analysis. Facilitation was further examined through experimental planting and Kaplan‐Meier survival analysis. Results: Both in grazed and protected areas, Olea recruits were found exclusively under shrubs, primarily under Euclea racemosa although Acacia etbaica was more abundant. Olea recruitment is distributed randomly at landscape scale, but depends on shelter at patch scale. Shelter ability is related to shrub shape and species identity. Dense multi‐stemmed shrubs with a wide base and crown on a mulch‐rich mound are key recruitment foci. Euclea shrubs have these favoured traits and probably act as preferential perching sites for avian seed dispersers. Soil and organic matter accumulation under Euclea shrubs may also create favourable conditions for Olea germination and survival. Experimentally planted seedlings had a better chance for survival under Euclea. Conclusions: Olea regeneration is probably subject to both passive (disperser‐mediated) and active facilitation. Small changes of shrub traits can alter the suitability of a patch for Olea recruitment. Protection of shrubs can increase facilitation for seedlings, while pruning may reduce competition for saplings and thus enhance forest succession. Planting of raised Olea seedlings under Euclea shrubs in years with a good rainy season may further assist forest restoration.     

Does shelter enhance early seedling survival in dry environments? A test with eight Mediterranean species

Question: Do solid‐walled polyethylene tubes and mesh fabric tubes improve the short‐term survival of eight Mediterranean tree and shrub species often used in the restoration of arid environments? Location: We conducted two experimental plantations in degraded field sites in the province of Almería (SE Spain), under arid Mediterranean conditions. Methods: One‐year‐old seedlings of Ceratonia siliqua, Juniperus phoenicea, Olea europaea, Pinus halepensis, Pinus pinaster, Quercus coccifera, Quercus ilex and Tetraclinis articulata were planted either sheltered by one of the above shelter tubes, or by being left unsheltered. Survival was recorded the first growing season after planting, which was a very dry season. Results: Overall, seedling survival ranged from as little as 0% to 24%, and tree shelters consistently enhanced survival in Quercus species only, ranging from 16% in walled shelters to 8% in mesh shelters. Shelters failed to boost survival in the six remaining species. Conclusion: The results of this study suggest that both walled and mesh shelters were mostly ineffective at increasing seedling survival for the Mediterranean species used in this experiment; these species coincide with those used in restoration programs. The use of shelters in restoration programs conducted in arid environments should be reconsidered, while walled shelters might be advisable for Mediterranean Quercus species only. Further research is necessary to develop and assess improved types of shelters for arid environments.     

Use of Shrubs as Nurse Plants: A New Technique for Reforestation in Mediterranean Mountains

Common techniques currently used for afforestation in the Mediterranean basin consider the pre‐existing vegetation (mainly shrubs) as a source of competition for trees, and consequently it is generally eliminated before planting. Nevertheless, it has been demonstrated that woody plants can facilitate the establishment of understory seedlings in environments that, like the Mediterranean area, are characterized by a pronounced dry season. In this study, we experimentally analyze the usefulness of shrubs as nurse plants for afforestation of two native conifers, Pinus sylvestris L. (Scots pine) and Pinus nigra Arnold (black pine). Two‐year‐old seedlings were planted in four microhabitats: (1) open interspaces without vegetation (which is the usual method used in afforestation programs), (2) under individuals of Salvia lavandulifolia, (3) under the north side of spiny shrubs, and (4) under the south side of spiny shrubs. Pine survival was remarkably higher when planted under individuals of the shrub S. lavandulifolia (54.8% for Scots pine, 81.9% for black pine) compared with open areas (21.5% for Scots pine, 56.8% for black pine; chi square, p < 0.05). The survival of both pines was also higher when planted on the north side of spiny shrubs, although the survival on the south side was similar to that found in open areas. In addition, pine growth was not inhibited when planted in association with shrubs. This pattern appears to result from the combination of abiotic conditions imposed by the presence of a nurse shrub, which leads to improvement in seedling water status and therefore reduced summer mortality by drought. The results show that the use of shrubs as nurse plants is a technique that offers both economic and ecological advantages, in terms of savings in labor and plant material and reduced and even negligible impact on the pre‐existing vegetation.     

Factors Limiting the Survival of Native Tree Seedlings Used in Conservation Efforts at the Edges of Forest Fragments in Upland Madagascar

We tested four reforestation techniques in tropical forest fragments that were damaged by fire in upland Madagascar. We conducted a full‐factorial experiment on the survival of transplanted seedlings of five native tree species in grassland plots adjacent to the forest fragments in the Ambohitantely Forest Reserve. The species studied were Dodonaea madagascariensis, Filicium decipiens, Olea lancea, Podocarpus madagascariensis, and Rhus taratana. A total of 480 seedlings were planted; 207 survived the 15 months of the experiment. The factors examined were distance of the reforestation plots from the forest, mixing of forest soil into the plots, application of chemical fertilizers, experimental shading of plots, and the cover of naturally establishing shrubs. Both increasing the distance of plots from the forest edge and adding chemical fertilizers significantly reduced the survival of all seedlings. The surprising negative effects of fertilization may be partly due to increased competition from naturally establishing shrubs that are adapted to exploit high nitrogen levels. Mixing soil from the forest areas into the plots did not change seedling survival. Shading reduced the survival of D. madagascariensis seedlings and did not increase the survival of any species. These findings suggest that the success of reforestation projects can be increased by planting seedlings close to the existing forest fragments. Reforestation of similar tropical forests is likely to be more successful if efforts are focused on expanding the size of existing fragments of tropical forest, rather than on establishing new fragments in grassland openings.     

Facilitation and interference underlying the association between the woody invaders Pyracantha angustifolia and Ligustrum lucidum

Questions: 1. Is there any post‐dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage‐protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse‐plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well.     

Shrub influences on seedling performance when restoring the slow‐growing conifer Pilgerodendron uviferum in southern bog forests

Forest restoration is most efficient if it can take advantage of facilitative interactions between established vegetation and planted trees. However, positive and negative interactions have been identified in a number of plant communities. After centuries of anthropogenic fires, forest recovery has been extremely slow in southern bog forests previously dominated by the slow‐growing and vulnerable conifer Pilgerodendron uviferum on Chiloé Island, Chile. Today, the landscape is dominated by secondary shrublands with scattered patches of Sphagnum moss and limited natural tree regeneration. We hypothesized that the retention of secondary shrubs facilitates the early performance of P. uviferum restoration plantings by providing better microsite conditions. To test this hypothesis, we compared the response of seedlings planted on sites prepared at two levels of intervention: after shrubs had been removed or where shrubs were retained. Shrub retention showed a nurse‐plant effect on P. uviferum seedlings 4 years after planting, which resulted in reduced physiological stress (measured as Fv/Fm) for seedlings, as well as reduced browsing. Consequently, the seedlings growing in areas with shrub retention had larger height increment and higher vitality than those in areas where shrubs had been removed. Thus, the more open micro‐site conditions created by shrub removal resulted in generally poorer seedling performance, although seedling mortality—which was low overall (approximately 2–4%)—showed no significant difference between the two levels of intervention. These findings have direct implications for the restoration of slow‐growing conifers that can tolerate extreme wet conditions in highly degraded forests.     

Avian Response to Bottomland Hardwood Reforestation: The First 10 Years

Bottomland hardwood forests were planted on agricultural fields in Mississippi and Louisiana predominantly using either Quercus species (oaks) or Populus deltoides (eastern cottonwood). We assessed avian colonization of these reforested sites between 2 and 10 years after planting. Rapid vertical growth of cottonwoods (circa 2–3 m/year) resulted in sites with forest structure that supported greater species richness of breeding birds, increased Shannon diversity indices, and supported greater territory densities than on sites planted with slower‐growing oak species. Grassland birds (Spiza americana[Dickcissel] and Sturnella magna[Eastern Meadowlark]) were indicative of species breeding on oak‐dominated reforestation no more than 10 years old. Agelaius phoeniceus (Red‐winged Blackbird) and Colinus virginianus (Northern Bobwhite) characterized cottonwood reforestation no more than 4 years old, whereas 14 species of shrub‐scrub birds (e.g., Passerina cyanea[Indigo Bunting]) and early‐successional forest birds (e.g., Vireo gilvus[Warbling vireo]) typified cottonwood reforestation 5 to 9 years after planting. Rates of daily nest survival did not differ between reforestation strategies. Nest parasitism increased markedly in older cottonwood stands but was overwhelmed by predation as a cause of nest failure. Based on Partners in Flight prioritization scores and territory densities, the value of cottonwood reforestation for avian conservation was significantly greater than that of oak reforestation during their first 10 years. Because of benefits conferred on breeding birds, we recommend reforestation of bottomland hardwoods should include a high proportion of fast‐growing early successional species such as cottonwood.     

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.     

Demographic structure of California chaparral in the long-term absence of fire

Abstract. Demographic structure of 12 chaparral sites unburned for 56 to 120 years was investigated. All sites were dominated by vigorous shrub populations and, although there was colonization by seedlings of woodland tree species in several stands, successional replacement of chaparral was not imminent. Although successional changes in community composition were evident, there was no indication of a decline in species diversity. Non-sprouting species of Ceanothus suffered the greatest mortality at most, but not all, sites. Sprouting shrubs, such as Quercus and Heteromeles had very little mortality, even in stands more than a century old. All postfire resprouting species had multiple stems of different ages indicating these shrubs were capable of continuously regenerating their canopy from basal sprouts. Ceanothus populations were highly clumped and there was a significant correlation across all sites between variance/mean ratio and percentage mortality. As Ceanothus populations thinned, they became less clumped. In mixed chaparral stands, Quercus and Heteromeles were significantly taller than associated Ceanothus shrubs and overtopped the Ceanothus; at two sites, the density of live Quercus per plot was correlated with the density of dead Ceanothus. Thus, mortality of Ceanothus stems is likely related to both intra and interspecific interations. Seedling recruitment was observed for most shrub species that regenerate after fire by resprouting; seedling and sapling densities ranging from 1000–36 500 ha^-1 were recorded for Quercus dumosa, Rhamnus crocea, Prunus ilicifolia, Heteromeles arbutifolia and Cercocarpus betuloides. For all but the last species, seedlings and saplings were most abundant beneath the canopy cover and not in gaps. Across all sites, recruitment was significantly correlated with depth and bio-mass of the litter layer. Cercocarpus betuloides was present in several stands, but seedling establishment was found only in one very open, disturbed stand. Regardless of stand age, taxa such as Adenostoma, Arctostaphylos and Ceanothus, which recruit seedlings after fire, had no significant seedling production.     

Tissue water relations of four co-occurring chaparral shrubs

Summary Chaparral shrubs of California have a suite of morphological and physiological adaptations to withstand the prolonged summer droughts of a mediterranean climate. Not all species of chaparral have the same rooting depth and there is some evidence that those with shallow roots have tissue that is most tolerant to water stress. We tested this notion by comparing the tissue water relations of four co-occurring chaparral shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a pressure-volume technique and a dew-point hygrometer to metsure seasonal changes in osmotic potential when plant tissue was fully hydrated and osmotic potential at predawn, midday, and the turgor loss point. We also calculated seasonal changes in the minimum daily turgor potential, saturated weight/dry weight ratio of leaf tissue, and the bulk modulus of elasticity. We had information on the seasonal water use patterns and apparent rooting depths of these same four shrubs from a previous study (Davis and Mooney 1986). All evidence indicated that Rhamnus had shallow roots and Quercus deep roots. Our results indicated that the tissue water relations of our four co-occurring chaparral shrubs were not alike. Even though Rhamnus had shallow roots, it had the least xerophytic tissue. Seasonal osmotic potential and saturated weight/dry weight ratios were relatively high and bulk modulus of elasticity and minimum daily turgor potentials were low. Furthermore, even though Quercus had deep roots and experienced no seasonal water stress at our study site, its tissue water relations indicated relatively high tolerance to water stress. We conclude that seasonal drought tolerance of stem and leaf tissue of co-occurring chaparral shrubs does not necessarily correspond to rooting depth, to soil moisture resources available to the shrub, or to the degree of seasonal water stress experienced by the shrub.     

Legacy microsite effect on the survival of bitterbrush outplantings after prescribed fire: capitalizing on spatial variability to improve restoration

Restoration of shrubs in arid and semi‐arid rangelands is hampered by low success rates. Planting shrub seedlings is a method used to improve success in these rangelands; however, it is expensive and labor intensive. The efficiency of shrub restoration could be improved by identifying microsites where shrub survival is greater. Bitterbrush (Purshia tridentata Pursh DC) is an important shrub to wildlife that has declined because of conifer encroachment, excessive defoliation, wildfires, and low recruitment. We investigated planting bitterbrush seedlings in western juniper (Juniperus occidentalis ssp. occidentalis Hook) encroached shrublands after prescribed fire was used to control trees. Bitterbrush seedlings were planted in under (canopy) and between (interspace) former juniper canopies at five blocks and evaluated for three growing seasons. Bitterbrush survival was greater than 50% in the former canopy, but only 5% in the interspace microsite by the third growing season. Growth of bitterbrush was also greater in the former canopy compared with the interspace, potentially due to markedly less perennial vegetation in this microsite. Exotic annual grasses and annual forbs became prevalent in the former canopy in the second and third growing season, suggesting that soil resource availability was greater in this microsite. These results suggest that restoration success will vary by specific locations within a burned landscape and that this variability can be used to improve restoration efficiency. In this situation, bitterbrush restoration can be improved by planting seedlings in former canopy compared with interspace microsites.     

The effectiveness of different planting frameworks for recruitment of tropical rainforest species on ex‐rainforest land

A long‐term rainforest restoration experiment was established on abandoned pasture in northeastern Queensland in 1993 to examine the effectiveness of five different restoration planting methods: (T1) control (no plantings); (T2) pioneer monoculture (planting seedlings of one pioneer species, Homalanthus novoguineensis, Euphorbiaceae); (T3) Homalanthus group framework method (H. novoguineensis and eight other pioneer species); (T4) Alphitonia group framework method (Alphitonia petriei, Rhamnaceae, with eight other pioneer species); and (T5) maximum diversity method (planting pioneers, middle‐phase species, and mature‐phase species). We investigated temporal patterns in the (1) fate of seedlings originally planted in 1993; (2) natural recruitment of native plant species; and (3) current habitat structure (canopy cover and ground cover of grasses and invasive plants) within each restoration treatment. A total of 97% of seedlings planted in T2 died within the first 13 years and all had died by 2014. A total of 72% of seedlings planted in T3, 55.5% of seedlings planted in T4, and 55% of seedlings planted in T5 also died by 2014. By 2014, 42 species from 21 families had recruited across the experimental site, and the abundance of recruits was almost twice that recorded in 2001 and 2006. Overall, T3, T4, and T5 had the greatest diversity and abundance of recruits. By 2014, canopy cover was greatest in T3, T4, and T5 but grass cover was least in T5. It is concluded that some restoration success measures increase with planting diversity, but overall the rate of recovery is similar in framework species and maximum diversity method.     

Controls on Shumard Oak (Quercus shumardii) Establishment into the Cross Timbers Ecotone of Oklahoma: Implications for Restoration

To better understand how the Cross Timbers ecotone between U.S. eastern deciduous forest and tallgrass prairie is organized and maintained, I set out seedlings of Shumard oak (Quercus shumardii) for 1 year with treatments of water addition, trenching in pots to reduce belowground competition, burning, and patch type. I found that (1) seedlings in pots survived twice as much as those without pots and seedlings in unburned areas survived up to four times as much as those in burned areas; (2) seedlings in pots grew twice as fast as those planted directly in the soil without a pot; and (3) whereas adding water, planting in a pot, and planting under shrubs all increased leaf chlorophyll absorption, it was when the seedling was watered, under a shrub, or in a pot that chlorophyll increases were 2‐fold. Results suggest these management and restoration strategies to increase Cross Timbers oak establishment: first reduce belowground competition by the resident grasses; second, do not burn; and third, use shrubs as a facilitating tool. Adding water to tree seedlings should be used as a management tool only after others have been tried and failed. Finally, an ecotone conceptual vegetation model is presented both to incorporate the results and to explain how ecotones may be managed, restored, and maintained.     

Species Identity and Water Availability Determine Establishment Success Under the Canopy of Retama sphaerocarpa Shrubs in a Dry Environment

Seedling establishment in harsh environments is often enhanced by the proximity of adult shrubs. This information has been used in restoration work by placing seedlings of species being restored under the canopy of some shrubs. However, monitoring this process is often restricted to a single species, and comparisons with practices that protect planted seedlings against harsh conditions are scant. Similarly, few studies have supplied seedlings with water in the summer to observe the effects of water availability on the interaction. We compared sapling survival of three woody species (Olea europaea, Pistacia lentiscus, and Ziziphus lotus) under the leguminous shrub Retama sphaerocarpa and in gaps covered with piled branches that mimicked a shrub canopy. After 3 years, survival of saplings planted under Retama differed depending on species identity and water supply. Survival of Olea saplings placed under Retama shrubs was twice that under piled branches if not watered (35 ± 8 vs. 17 ± 2 %, respectively), whereas survival of saplings under Retama, if watered, was less (48 ± 11 vs. 68 ± 8%, respectively). Retama shrubs had a negative effect on Ziziphus; most saplings died under its canopy, whereas survival in piled branches ranged 10–54%. Pistacia was neither facilitated nor outcompeted by Retama. Facilitation of Olea by Retama shrubs was more apparent under dry conditions where watering increased competition and decreased facilitation. Overall, we conclude that Retama shrubs can help dry land restoration to a greater extent than artificial shade for Olea when not watered. The critical role played by water supply in determining nursing success rates warrants further study.     

Short‐term bryoid and vascular vegetation response to reforestation alternatives following wildfire in conifer plantations

Question: How are dynamics of early‐seral post‐fire vascular plant and bryoid (terrestrial mosses, lichens, and fungi) vegetation impacted by reforestation activities, particularly manual vegetation removal and planting density? Does the relationship between vegetation dynamics and vegetation removal differ between harsh (west‐facing) and moderate (east‐facing) aspects? Location: Five high‐severity burn plantation forests of Pseudotsuga menziesii in southwestern Oregon, USA. Methods: Plantations severely burned in a recent wildfire were planted with conifer seedlings as a four‐species mixture or a monoculture, at two different densities, with and without manual vegetation removal. A subset of plots was also planted on a contrasting aspect within each plantation. The contrasting aspects differed in potential solar insolation and were indicative of moderate (eastern exposure) and harsh (western exposure) site conditions. Covers of shrub, herbaceous and bryoid vegetation layers were measured during reforestation activities 2–4 yr after the fire. Dynamics of structural layer cover and community composition were compared among treatments with analysis of variance and multivariate analyses (non‐metric multidimensional scaling and blocked multi‐response permutation procedure). Results: Structural layer cover and community composition differed between areas that received reforestation treatments and untreated areas. However, variability within treatments in a plantation was greater than variability within treatments across plantations. Effects of vegetation removal on composition and structure were more evident than effects of planting or altering planting density. Vegetation removal decreased cover of tall and low shrub and the bryoid layer, and increased herbaceous layer cover. Bryoid community and low shrub structural layer responses were more pronounced on moderate aspects than on harsh aspects. Vegetation removal shifted vascular plant community composition towards exotic and annual species. Conclusions: These reforestation treatments may be implemented without substantially altering early‐seral vegetation community composition dynamics, especially in areas with harsh site conditions. Site conditions, such as aspect, should be evaluated to determine need and potential effects of reforestation before implementation. Monitoring for exotic species establishment should follow reforestation activities.     

Population dynamics of four understorey shrub species in beech forest

Question: In the population dynamics of four understorey shrub species (Hp, Hydrangea paniculata Sieb, et Zucc.; Lu, Lindera umbellata Thunb. var. membranacea (Maxim.) Momiyama; Ms, Magnolia salicifolia (Sieb, et Zucc.) Maxim.; Vf, Viburnum furcatum Blume ex Maxim.), (1) What is the relative importance of seedling regeneration versus vegetative growth? (2) Can these shrubs persist stably for a long time in the understorey? (3) What kind of variation in demographic features is observed among these shrubs? Location: 780m a.s.l., north‐eastern Japan. Methods: Population dynamics were analyzed by using stage‐classified matrix models. Models were mainly constructed from five years stem‐census data, including current‐year seedlings and sprouts. Results: Current‐year sprouts emerged every year in every species. Current‐year seedlings emerged every year in Lu and Vf, but densities were very low. In every species, population growth rate (A) was close to the equilibrium value 1.0 and no statistical difference was found among species. The stable stage‐distribution predicted from the matrix model was similar to the observed distribution for Lu, Ms and Vf, but much different for Hp. Elasticity matrix was also similar among Lu, Ms and Vf, but was quite different for Hp. Conclusions: Lu, Ms and Vf were considered as climax shrubs that can regenerate and maintain their population stably in the understorey, even if canopy gaps form infrequently. Hp is a pioneer shrub that require more frequent formation of canopy gaps for long‐term persistence in the understorey.