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Landscapes Forest and Global Change - ESA - Escola Superior ...

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K. L. Martin & P.C. Goebel 2010. Impact of hemlock decline on successional pathways <strong>and</strong> ecosystem function<br />

149<br />

1. Virginia (Jefferson <strong>and</strong> George Washington National <strong>Forest</strong>s), where HWA has been present<br />

for 25-30 years <strong>and</strong> hemlocks in invaded st<strong>and</strong>s are likely dead (based on the 4-15 year<br />

mortality timeframe; McClure 1991);<br />

2. West Virginia (New River <strong>and</strong> Gauley River National Recreation Areas, Monongahela<br />

National <strong>Forest</strong>), where HWA has recently invaded (2-5 years) <strong>and</strong> trees are declining; <strong>and</strong>,<br />

3. Ohio (Lake Katherine State Nature Preserve, Sheick Hollow State Nature Preserve, Hocking<br />

State <strong>Forest</strong>) where HWA is not yet present.<br />

In this paper, we focus on data collected in the Unglaciated Allegheny Plateau physiographic<br />

province of southeastern Ohio (Brockman 1998). This province of deep cliffs <strong>and</strong> valleys cut<br />

into the s<strong>and</strong>stone bedrock supports the majority of T. canadensis within Ohio (Black <strong>and</strong> Mack<br />

1976). Three of our steam reaches were located at Lake Katharine, an 817-ha State Nature<br />

Preserve, four were located within the 3,924-ha Hocking State <strong>Forest</strong>, <strong>and</strong> one at Sheick<br />

Hollow, a 61-ha State Nature Preserve. The climate in the area is continental with cold, snowy<br />

winters (mean = 0 o C) <strong>and</strong> warm, humid summers (mean = 21.6 o C) with an average rainfall of<br />

102 cm distributed evenly throughout the year (Kerr 1985; Lemaster <strong>and</strong> Glimore 1989). All<br />

areas were second-growth forest ravines with hemlock approximately 50% or more of the basal<br />

area. Any areas with evidence of recent human disturbance (logging roads, stumps) were<br />

excluded.<br />

We sampled vegetation using a series of five 100-m 2 circular plots (5.64 m radius) centered 10,<br />

30, <strong>and</strong> 50-m from small streams <strong>and</strong> 15 m apart. The species <strong>and</strong> d.b.h. (diameter at breast<br />

height, 1.67 m) of all woody vegetation was recorded. Stems 2.5 - 10 cm d.b.h. were classified<br />

as saplings. All stems > 10 were coded as dominant (full canopy in sun), co-dominant (portions<br />

of canopy in sun), intermediate (top of canopy not overtopped), or sub-canopy (overtopped). For<br />

analyses, the five subplots in each transect were added together for analyses for a 500-m 2 scale.<br />

To underst<strong>and</strong> community characteristics at different slope positions across the ravines, we also<br />

analyzed functional diversity using species guilds developed for the Central Hardwood <strong>Forest</strong> of<br />

the USA by Sutherl<strong>and</strong> et al. (2000). As we are focused on T. canadensis as a foundation<br />

species currently subjected to species-specific mortality, we separated it into its own functional<br />

group for analyses.<br />

Mean species <strong>and</strong> functional richness <strong>and</strong> diversity were compared by transect using one-way<br />

analysis of variance (ANOVA) tests. In cases where means differed, the three transects were<br />

subjected to Tukey’s mean separation test. All analyses were calculated using PASW statistics<br />

18 software (SPSS Inc, Chicago, IL)<br />

3. Results<br />

Our results indicate that in ravines of the Unglaciated Allegheny Plateau of Ohio, T. canadensis<br />

is highly dominant with few other species in the canopy, sub-canopy, or sapling layers (Table<br />

1). Hemlock basal area did not vary by transect, <strong>and</strong> neither did overall basal area. Although<br />

there was not a strong separation (Canopy species richness F = 2.57, P = 0.100, d.f. = 2), Betula<br />

lenta L. <strong>and</strong> Liriodendron tulipifera L., as well as Fagus gr<strong>and</strong>ifolia Ehrh., were more common<br />

adjacent to the stream. Sutherl<strong>and</strong> et al. (2000) categorize B. lenta <strong>and</strong> L. tulipifera as a part of a<br />

long-lived, shade intolerant functional group, indicating they are gap-dependent in hemlock<br />

ravines. On the other h<strong>and</strong>, F. gr<strong>and</strong>ifolia is part of the same extremely shade tolerant, slowgrowing,<br />

<strong>and</strong> long-lived functional group that includes T. canadensis (Sutherl<strong>and</strong> et al. 2000).<br />

Moving from the stream upslope, there was also a slight increase in canopy functional richness<br />

(F= 3.95, P = 0.035, d.f. = 2). This was largely due to increases in Quercus <strong>and</strong> Carya species,<br />

which Sutherl<strong>and</strong> et al. (2000) group into the red oak- hickory (i.e. Quercus rubra L., Q.<br />

coccinea Münchh. <strong>and</strong> Carya spp.) <strong>and</strong> the white oak (Q. alba L., Q. prinus L. ) groups due to<br />

differing germination requirements. Along the 30-m transect, the shade-tolerant Acer-Ulmus<br />

<strong>Forest</strong> <strong>L<strong>and</strong>scapes</strong> <strong>and</strong> <strong>Global</strong> <strong>Change</strong>-New Frontiers in Management, Conservation <strong>and</strong> Restoration. Proceedings of the IUFRO L<strong>and</strong>scape Ecology<br />

Working Group International Conference, September 21-27, 2010, Bragança, Portugal. J.C. Azevedo, M. Feliciano, J. Castro & M.A. Pinto (eds.)<br />

2010, Instituto Politécnico de Bragança, Bragança, Portugal.

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