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Breeding Habitats of the Wood Thrush and Veery

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Breeding Habitats of the Wood Thrush and Veery BREEDING HABITATS OF THE WOOD THRUSH AND VEERY ROBERT I. BERTIN Habitat-related differences in the abundance wise, the direction was taken from a table of random of bird species are appreciated by every bird- numbers. Vegetation cover data were taken at each watcher and have received their share of of 100 points (even,. 2 m ). The kinds and numbers of plants touching a 10 mm diameter peg between scientific study. In very few cases, however, 0 and 0.2 m above the ground (C layer) were re- do we understand why a species occurs in a corded. Similar data were recorded between 0.2 and 1.0 m (B laver) and between 1.0 and 3.0 m (A particular habitat, either in an ultimate (evo- .I lutionary) sense or a proximate (behavioral) layer) using a 25 mm diameter pole. These cover sense. Before these questions can be ade- figures are slightly greater than true percent cover because determination of the latter requires dimen- quately approached, a knowledge of the quan- sionless sampling devices. I noted the type of sub- titative aspects of species ’ habitats is necessary. strate at the same 100 points. Tree size and Most information on this subject comes from distribution were evaluated by point quarter sampling comparative habitat analyses (e.g., Sturman at 25 points (every 8 m). Moisture regimes were recorded according to a scale used by Damman 1968, Wiens 1969, James 1971, Anderson and ( 1964). The scale runs from zero (here converted Shugart 1974). Vegetation structure has been to -1) for loose sand, rock piles, and gravel, to 8 emphasized heavily in such studies and often for permanently inundated soils. It is based on soil has been adequate to measure habitat dif- features such as texture, state of oxidation, and ferences among species. Floristic and physical water table, and is independent of short term weather conditions. For example, sandy loams typically have factors have received less attention although moisture regimes of 2, whereas anaerobic soils with MacArthur (1964) suggested that the latter maximum water level close to the soil surface are may be important for at least some species. classified as 5. I estimated the territory percentage This study examines quantitatively the habi- occupied by different successional stages and plant communities. Successional stages were classified as tats of two woodland thrushes, the Wood “early”: maximum tree height < 14 m; “late”: Thrush (Hylocichla mustelina) and the Veery maximum tree height > 14 m, but with many early (Catharus fuscescens). My purposes were to successional plants remaining (and often a thicker determine how and to what extent their shrub layer); and “mature”: tree height > 14 m habitats differed and to indicate factors which and with a mature flora. Plant communities are described elsewhere ( Bertin 1975, Kershner 1975 ) seemed to be favored by each species in its according to techniques of the Zurich-Montpelier selection of habitats. Some physical and school (e.g., Shimwell 1971: Ch. 6). Communities floristic, as well as structural, habitat features were delimited on the basis of floristic similarities were included to examine their effects on the and differences which corresponded to differences in soil type, moisture regime, topography, exposure, etc. distribution of the two species. Territories of the two thrush species were compared using the Mann-Whitney U-test on parameters taken METHODS from the raw data. Structural parameters included percent vegetation cover in A, B, C, and tree layers, I studied thrush territories at two localities in Litch- tree height and diameter, number of stems > 25 mm field Co., western Connecticut. The Pond Mountain dbl 1 per ha, and percentage of territorial area in area in Kent is a 320-ha deciduous woodland (max. different successional stages. Physical factors in- relief = 230 m), with some open fields and thickets. cluded moisture regime, substrate type, and slope. The mature lowland woods are dominated by Sugar Floristic parameters included importance values for Maple (Acer saccharum) and White Ash (Fraxinus the 40 commonest tree species and abundance values americana), and the uplands by Red and Chestnut for the 100 commonest plant species. Each importance Oaks (Quercus r&a and Q. p&us). The Catlin value combines density, dominance, and frequency, Wood area in Litchfield is a 20-ha mature lowland with 300 as the maximum possible value (for a forest (max. relief = 10 m) dominated by Eastern monotypic stand) (Phillips 1959:44). Plant abun- Hemlock ( Tsuga canadensis), White Pine (Pinus dance values refer to total hits on the sampling pole strobus) and Red Maple ( Acer rubrum). or peg at all 100 sampling points in a territory for Territories of 49 Wood Thrush and 55 Veery pairs the particular plant species. Initially, I compared at Pond Mountain. and those of 6 Wood Thrush and the amounts of each plant community in territories 6 Veery pairs at Catlin Wood were mapped between of the two thrush species, but they did not prove 10 May and 15 June 1974. I visited each territory useful in differentiating territories and are not dis- an average of 6 times; each visit averaged about cussed further. 0.5 h. To map, I plotted positions of males sincrincr- - and reacting aggressively to recorded songs of their RESULTS species. IDENTIFICATION OF IMPORANT Habitat data were recorded in each territory along HABITAT FEATURES 200 m of transect, which was divided according to territory size. In heterogeneous territories the transects Territories of both thrush species at Pond were aligned along the most obvious gradient; other- Mountain were clustered along streams and [3031 The Condor 79:303-311, 1977 304 ROBERT I. BERTIN LEGEND summit complex Dryopteris group Eupatorium group ] Viburnum group Skiff Mountain Road open areas early successional areas 0 Wood Thrush territories Av eery territories roads H streams / boundaries of bnd Mountain Natural Area -- limits of orea in which Veery territories were mapped FIGURE 1. Distribution of Wood Thrush and Veery territories at Pond Mountain with respect to plant com- munity groups. Average moisture regimes: summit complex -1.0, CUWX group 1.0, Gaylussacia group 0.5, Viburnum group 2.0, Dryopteris group 3.5, Euputorium group 4.0, Iris group 6.0. Map based on Kershner 1975. in wetter places (Fig. 1). No similar pattern areas and the drier unoccupied areas. In was observed at Catlin Wood, as this area analyzing them, the availability of each factor contained no permanent water courses, and in the entire study area had to be considered the range in moisture conditions was less as this affects the range of habitats occupied. than at Pond Mountain. Many environmental For example, average moisture regimes in factors differed between the wet occupied most territories of both thrushes was 2 to 3 WOOD THRUSH AND VEERY HABITATS 305 A 16 3 1; n w a 2 8 0 0 t I B 0 80- w K wet s dry o 60- MOISTURE REGIME 0 FIGURE 3. Percentage occupancy of plant com- munities with different moisture regimes in mature woodland at Pond Mountain. Squares denote Wood Thrushes, circles denote Veeries. Correlation of per- cent occupancy with moisture regime = 0.88 for Wood Thrushes and 0.90 for Veeries, both significant at .05 level. 1 2 3 4 6 7 dry wet lated to these variables in a stepwise multiple MOISTURE REGIME regression, This technique is not ideally suited FIGURE 2. (A) Area of mature woodland at Pond to the present situation because the results Mountain occupied by Wood Thrush territories at dif- can be adversely affected by high correlation ferent moisture regimes. (B ) Percentage of available of independent variables (e.g., Neter and area at different moisture regimes occupied by Wood Wasserman 1974), and, like other statistical Thrush territories in mature woodland at Pond Moun- tain. No territory included sites with a moisture techniques, it does not address causation. regime less than 0. Category 5-6 was excluded, as no However, it does indicate which of the in- plant community had this moisture regime. dependent variables account for the greatest amount of variability in habitat occupancy. For the Wood Thrush, four variables made (Fig. 2A) although regimes between 6 and 7 significant contributions to the regression: were preferred (Fig. 2B) ; this difference re- moisture regime (P < .OOl), tree diameter flects the greater amount of available habitat (P < .Ol), herb (A + B) cover (P < .05) with intermediate moisture regimes. and shrub (C) cover (P < .05). Moisture Habitat availability was taken into account regime alone accounted for 76.0% of the for seven properties of the plant communities. variability in habitat occupancy. The next The relative homogeneity of vegetation struc- highest single variable (percentage of bare ture in each mature plant community allowed soil exposed) accounted for only 37.4% of me to calculate average values of these pa- this variability. The four significant variables rameters applicable to each community as a together accounted for 98.3% of the variability whole. Percentage occupancy of each com- in the dependent variable. For the Veery, munity was calculated from territory size and two variables made significant contributions composition measurements from this study, to the regression: moisture regime (P < .OOl) and plant community area measurements from and herb (A + B) cover (P < .05). Moisture Kershner (1975). Average values of the fol- regime alone accounted for 78.4% of the vari- lowing parameters were calculated for each ability in habitat occupancy, and the next (mature) plant community: tree, herb (A + highest single variable (herb cover) ac- B ), and shrub ( C) layer cover, tree diameter counted for only 38.2% of this variability. and density, percentage of bare soil exposed, These two significant variables together ac- and moisture regime.
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