Alternatives for Reintroducing a Rare Ecotone Species: Manually Thinned Forest Edge versus Restored Habitat Remnant

Species native to ecotones are often overlooked in restoration efforts despite the increasing rarity of ecotone habitat. In fragmented, fire-suppressed landscapes, true ecotone may no longer exist. Restoration biologists interested in reintroducing ecotone species must decide whether to plant them in historic ecotones maintained by manual thinning or whether to opt for discrete restoration areas that are easier to maintain. We investigated these two alternatives with Lantana canescens , a rare tropical shrub native to the ecotone between pine and hardwood forests of Miami-Dade County, Florida, U.S.A. Our short-term findings show that after 15 and 18 months, survival of transplants was 69% in a restored site and 65% and 84% in two historic ecotone sites. The restored site had significantly higher photosynthetically active radiation (PAR) (75%) than the historic ecotones (25–39%). Correspondingly, 267 seedlings have recruited at the restored site, whereas only 8 have emerged at both historic ecotone sites. Seedling establishment was associated with higher PAR at the restored site. We found that overall population sustainability was higher at the restored site where there is the additional benefit of less maintenance. Our work suggests that, by reducing succession, a discrete restoration area can approach the historic conditions of hardwood/pine forest ecotone more closely than degraded historic ecotones themselves. We present a viable solution for conserving rare ecotone species when their natural habitat and the processes that maintained it no longer exist.     

Dark septate endophyte and arbuscular mycorrhizal status of vegetation colonizing a bottomland hardwood forest after a 100 year flood

Levels of arbuscular mycorrhizal (AM) colonization and dark septate endophyte (DSE) colonization were assessed in the vegetation recolonizing a remnant bottomland hardwood forest in north central Texas following a 100 year flood. Thirty seven plant species representing 21 dicotyledonous and 2 montocotyledonous families established following floodwater recession. AM and/or DSE were found in all species. AM colonization was found in 31 out of the 37 species assessed including both monocotyledonous families (Poaceae and Cyperaceae) and 17 out of 21 dicotyledonous families (Acanthaceae, Asteraceae, Boraginaceae, Cucurbitaceae, Euphorbiaceae, Lamiaceae, Loganiaceae, Lythraceae, Malvaceae, Onagraceae, Pedaliaceae, Ranunculaceae, Sapindaceae, Scrophulariaceae, Solanaceae, Verbanaceae and Violaceae). DSE were found in 31 out of 37 species assessed including both monocotyledonous families and 15 out of 21 dicotyledonous families (Amaranthaceae, Asteraceae, Boraginaceae, Brassicaceae, Cucurbitaceae, Euphorbiaceae, Lamiaceae, Lythraceae, Malvaceae, Pedaliaceae, Phytolaccaceae, Polygonaceae, Ranunculaceae, Sapindaceae, Scrophulariaceae and Violaceae). There were no detectable differences in AM or DSE colonization levels among wetland indicator groups (p > 0.05). Levels of DSE colonization were negatively correlated with vesicular colonization and hyphal colonization for the obligate wetland species. There were no other significant relationships between AM and DSE colonization detected. Spearman rank order correlation coefficients did not differ significantly among wetland indicatory category for any level of AM or DSE colonization.     

Phenology of a northern hardwood forest canopy

While commonplace in other parts of the world, long‐term and ongoing observations of the phenology of native tree species are rare in North America. We use 14 years of field survey data from the Hubbard Brook Experimental Forest to fit simple models of canopy phenology for three northern hardwood species, sugar maple (Acer saccharum), American beech (Fagus grandifolia), and yellow birch (Betula alleghaniensis). These models are then run with historical meteorological data to investigate potential climate change effects on phenology. Development and senescence are quantified using an index that ranges from 0 (dormant, no leaves) to 4 (full, green canopy). Sugar maple is the first species to leaf out in the spring, whereas American beech is the last species to drop its leaves in the fall. Across an elevational range from 250 to 825 m ASL, the onset of spring is delayed by 2.7±0.4 days for every 100 m increase in elevation, which is in reasonable agreement with Hopkin's law. More than 90% of the variation in spring canopy development, and just slightly less than 90% of the variation in autumn canopy senescence, is accounted for by a logistic model based on accumulated degree‐days. However, degree‐day based models fit to Hubbard Brook data appear to overestimate the rate at which spring development occurs at the more southerly Harvard Forest. Autumn senescence at the Harvard Forest can be predicted with reasonable accuracy in sugar maple but not American beech. Retrospective modeling using five decades (1957–2004) of Hubbard Brook daily mean temperature data suggests significant trends (P≤0.05) towards an earlier spring (e.g. sugar maple, rate of change=0.18 days earlier/yr), consistent with other studies documenting measurable climate change effects on the onset of spring in both North America and Europe. Our results also suggest that green canopy duration has increased by about 10 days (e.g. sugar maple, rate of change=0.21 days longer/yr) over the period of study.     

Spatial arrangement of branches in relation to slope and neighbourhood competition

To gain a better understanding of the effects of spatial structure on patterns of neighbourhood competition among hardwood trees, the three-dimensional extension of primary branches was surveyed for ten community-grown Castanea crenata (Fagaceae) trees with respect to the positioning of neighbouring branches and the slope of the forest floor. There were significantly more branches extending towards the lower side of the slope than towards the upper side, but structural properties such as branch length and vertical angle were not affected by slope. When horizontal extension of a branch towards its neighbour was compared for a C. crenata branch and a neighbouring heterospecific, the former was significantly narrower than the latter when the inter-branch distance (horizontal distance between the base positions of two neighbouring branches) was short (< approx. 5 m). Castanea crenata branches tended to extend in a direction avoiding neighbouring branches of heterospecifics when the inter-branch distance was short. Furthermore, for an inter-branch distance <3 m, the horizontal extension of a C. crenata branch was less when it was neighbouring a heterospecific branch than when neighbouring a conspecific branch. These results suggest that horizontal extension of C. crenata branches is more prone to spatial invasion by nearby neighbouring branches of heterospecifics, and that the invasion can be lessened when C. crenata trees are spatially aggregated. The reason why such an arrangement occurs is discussed in relation to the later leaf-flush of C. crenata compared with that of other species in the forest.     

Vegetative Characteristics of Recently Reforested Bottomlands in the Lower Cache River Watershed, Illinois, U.S.A.

Abstract Interest in restoring native ecosystems is resulting in conversion of marginal agricultural lands to bottomland hardwood‐dominated forests in the midwestern and midsouthern United States. Growing stock for these efforts typically consists of planted oak (Quercus spp.) and volunteer vegetation. Reports of mixed reforestation success and the lack of post‐establishment tree growth data prompted this evaluation of vegetation characteristics of 5‐ to 7‐year‐old operational restorations in the Lower Cache River Watershed in southernmost Illinois, U.S.A. Fraxinus pennsylvanica (green ash), Acer negundo (box‐elder), and Liquidambar styraciflua (sweetgum) together comprised 77% of all tree stems observed. Full stocking of overstory tree species can be expected to produce a closed canopy stand within 160 m of a forested edge, due primarily to the abundance of rapidly growing volunteer‐origin trees. Planted oaks contributed minimally to total tree stocking but were present in sufficient numbers to eventually improve wildlife habitat, and therefore satisfied restoration objectives. Oak height was 23% greater when in the presence of a non‐oak tree species. Herbaceous cover was dominated by Solidago gigantea (late goldenrod) and Juncus spp. (rushes). Solidago gigantea was associated with poor growth and low density of non‐oak stems, whereas Juncus dudleyi (Dudley's rush) was associated with taller non‐oak stems. These results suggest that the presence of volunteer‐origin trees is crucial for the creation of full stand stocking that will result in rapid development of a closed canopy forest. Improved success of future reforestation efforts will require more intensive methods to establish adequate stocking beyond 160 m of a forest edge. Methods described here could be adapted for agricultural field to forest restorations in other regions to predict critical distances from volunteer seed sources within which supplemental planting would be unnecessary to meet tree stocking objectives.     

Snow depth, soil freezing, and fluxes of carbon dioxide, nitrous oxide and methane in a northern hardwood forest

Soil–atmosphere fluxes of trace gases (especially nitrous oxide (N₂O)) can be significant during winter and at snowmelt. We investigated the effects of decreases in snow cover on soil freezing and trace gas fluxes at the Hubbard Brook Experimental Forest, a northern hardwood forest in New Hampshire, USA. We manipulated snow depth by shoveling to induce soil freezing, and measured fluxes of N₂O, methane (CH₄) and carbon dioxide (CO₂) in field chambers monthly (bi-weekly at snowmelt) in stands dominated by sugar maple or yellow birch. The snow manipulation and measurements were carried out in two winters (1997/1998 and 1998/1999) and measurements continued through 2000. Fluxes of CO₂ and CH₄ showed a strong seasonal pattern, with low rates in winter, but N₂O fluxes did not show strong seasonal variation. The snow manipulation induced soil freezing, increased N₂O flux and decreased CH₄ uptake in both treatment years, especially during winter. Annual N₂O fluxes in sugar maple treatment plots were 207 and 99 mg N m⁻² yr⁻¹ in 1998 and 1999 vs. 105 and 42 in reference plots. Tree species had no effect on N₂O or CO₂ fluxes, but CH₄ uptake was higher in plots dominated by yellow birch than in plots dominated by sugar maple. Our results suggest that winter fluxes of N₂O are important and that winter climate change that decreases snow cover will increase soil:atmosphere N₂O fluxes from northern hardwood forests.     

Seven-Year Survival of Perennial Herbaceous Transplants in Temperate Woodland Restoration

Little is known about restoring the perennial herbaceous understory of Midwestern deciduous woodlands, despite the significant and widespread degradation of remnants due to human activities. Because many woodland understory species have reproductive characters that make reestablishment from seed slow or difficult, we investigated transplanting as a strategy for introducing 24 species to a degraded early-successional woodland in central Iowa, U.S.A. Plants were planted in single-species groups of generally four individuals, and then monitored for survival five times over a 7-year period, and for flowering during the first year. After 7 years, persistence of these groups was 57% averaged across species. Survival in years 5–7 does not reflect individuals that spread beyond the original planting units by self-sowing or vegetative spread and is therefore a minimum estimate of the abundance of many species at the site. Mean percent flowering was 72% across single-species groups for 15 species monitored. We consider these survival and flowering rates acceptable indicators of establishment success, especially given drought conditions at our site in the first few years and lack of weed control beyond the first year, and evidence that transplanted species were establishing outside the original planting locations. Additional work is needed to investigate regional differences in transplant success, and methods for sustainable production of species are not suitable for introduction by seed. We caution that our results do not necessarily apply to the restoration of rare species.     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Removal of Nonnative Vines and Post‐Hurricane Recruitment in Tropical Hardwood Forests of Florida1

In hardwood subtropical forests of southern Florida, nonnative vines have been hypothesized to be detrimental, as many species form dense “vine blankets” that shroud the forest. To investigate the effects of nonnative vines in post‐hurricane regeneration, we set up four large (two pairs of 30 X 60 m) study areas in each of three study sites. One of each pair was unmanaged and the other was managed by removal of nonnative plants, predominantly vines. Within these areas, we sampled vegetation in 5 X 5 m plots for stems 2 cm DBH (diameter at breast height) or greater and in 2 X 0.5 m plots for stems of all sizes. For five years, at annual censuses, we tagged and measured stems of vines, trees, shrubs and herbs in these plots. For each 5 X 5 m plot, we estimated percent coverage by individual vine species, using native and nonnative vines as classes. We investigated the hypotheses that: (1) plot coverage, occurrence and recruitment of nonnative vines were greater than that of native vines in unmanaged plots; (2) the management program was effective at reducing cover by nonnative vines; and (3) reduction of cover by nonnative vines improved recruitment of seedlings and saplings of native trees, shrubs, and herbs. In unmanaged plots, nonnative vines recruited more seedlings and had a significantly higher plot‐cover index, but not a higher frequency of occurrence. Management significantly reduced cover by nonnative vines and had a significant overall positive effect on recruitment of seedlings and saplings of native trees, shrubs and herbs. Management also affected the seedling community (which included vines, trees, shrubs, and herbs) in some unanticipated ways, favoring early successional species for a longer period of time. The vine species with the greatest potential to “strangle” gaps were those that rapidly formed dense cover, had shade tolerant seedling recruitment, and were animal‐dispersed. This suite of traits was more common in the nonnative vines than in the native vines. Our results suggest that some vines may alter the spatiotemporal pattern of recruitment sites in a forest ecosystem following a natural disturbance by creating many very shady spots very quickly.