Avian Community Response to Construction of a Naturalistic Golf Course in Tallgrass Prairie in Kansas

Transcription

1 183 Avian Community Response to Construction of a Naturalistic Golf Course in Tallgrass Prairie in Kansas ROBERT J. ROBEL 1, SAMANTHA L. BYE, KENNETH E. KEMP, and STEPHEN J. THIEN Division of Biology, Kansas State University, Manhattan, KS (RJR, SLB) Department of Statistics, Kansas State University, Manhattan, KS (KEK) Department of Agronomy, Kansas State University, Manhattan, KS (SJT) ABSTRACT -- We monitored the impacts on avian communities of constructing a naturalistic golf in tallgrass prairie from summer 1997 to winter Avian data were gathered during one year before construction, two years during construction, and one year after construction in grasslands on the golf area, on a nearby control grassland area, and in woody draws and woodland habitats on the golf area. Point count surveys were used to estimate avian abundance, species composition, and species richness; Jaccard s Coefficient of Community and Morisita s Index of Community Similarity were used to compare structures of avian communities. Construction activity reduced bird abundance in the grasslands of the golf more during winter than during summer. Jaccard s Coefficient of Community increased slightly from preconstruction to postconstruction years in the golf grassland during winter, whereas Morisita s Index of Similarity declined. Point count summer surveys detected no significant changes in total bird abundance in the grassland of the golf between preconstruction and postconstruction years, but more species common to both areas were more abundant in the control grassland area than on the golf grassland after the first year of construction. Jaccard s Coefficient of Community indicated greater likeness of avian communities in grasslands of the control and golf areas during summer of the preconstruction year than later, whereas Morisita s Index of Similarity detected little difference in avian community structure in the two grassland areas during summer from preconstruction to postconstruction years. After golf construction began, avian species 1 Corresponding author. address: rjrobel@ksu.edu

2 184 The Prairie Naturalist 37(4): December 2005 richness generally was less in the golf grassland than in the control grassland during winter and summer. Changes in numbers of the most and least abundant avian species in the golf grassland before construction did not differ from differences between preconstruction and postconstruction numbers of those species on the golf and control grassland areas. Golf related effects on bird populations in woody draws and woodlands of the golf area could not be assessed because neither had control areas with which data could be compared, but several avian species decreased by more than 85% in these areas between pre-construction and post-construction years. Key words: avian abundance, avian communities, avian populations, golf construction, Kansas, naturalistic golf construction, tallgrass prairie. Golf is a fast growing recreational industry in the United States, but its environmental impacts have received little attention (Thien et al. 2001). In the United States, more than 150 new golf s were established in 2004, and golf s now occupy more than 1.2 million ha of land surface. The 16,057 golf s (18 hole equivalents) in use in the United States in 2004 exceeded those in use in 1986 by approximately 25% (National Golf Foundation 2005). Golf s generally protect land from intensive residential and industrial development, but most also alter limited-use habitats to areas managed intensively for turfgrass. Reduced adverse ecological impacts are projected for naturalistic golf s where habitat alterations and toxic chemical uses are minimized (Audubon International 2005). In the Great Plains, golf s commonly are established in grassland areas, a once extensive habitat of which less than 1% remains (Samson and Knopf 1994). The loss of grassland habitat has contributed to the decline of grassland bird numbers (Herkert 1995, Herkert et al. 1996, Best et al. 1997) and caused Knopf (1994) to consider the grasslands of the Great Plains one of the most endangered ecosystems in North America. The extensive tallgrass native prairie areas of the Flint Hills of Kansas are strongholds for endemic grassland birds such as the dickcissel (Spiza americana), greater prairie-chicken (Tympanuchus cupido), grasshopper sparrow (Ammodramus savannarum), and Henslow s sparrow (A. henslowii) (Herkert 1994). Recent land-use changes in the Flint Hills are typified by urban development and some rangeland to cropland conversions (Kansas Agricultural Statistics 1999). When plans were announced in 1996 to construct a naturalistic 129 ha World Class PGA golf (Colbert Hills) in 1999 to 2000 on a 486 ha tract of tallgrass prairie in the Flint Hills, it created an opportunity to monitor the impact of this action on the avian community associated with the construction site. We initiated research to monitor the avian community at the site during the year before

3 Robel et al.: Birds and golf construction 185 construction, the two years of construction, and for one year after construction. We specifically studied the effects of golf construction on avian community composition, abundance of individual avian species, and numbers of bird species (species richness) by comparing data collected from the construction area with similar data gathered from a nearby 400-ha grassland control area. METHODS The golf construction site was in Riley County, approximately 5 km northwest of Manhattan, Kansas. The area was typical tallgrass prairie consisting of 80% grassland, 5% woody draws, and 15% riparian woodland. Dominant grasses included big bluestem (Andropogon gerardii), Indian grass (Sorghastrum nutans), and little bluestem (Schizachyrium scoparium) occasionally interspersed with small stands of smooth sumac (Rhus glabra). Woody draws consisted of smooth sumac, rough-leaved dogwood (Cornus drummondii), and plum (Prunus spp.). Woody draws lead to lowland riparian areas dominated by eastern red cedar (Juniperus virginiana), redbud (Cercis canadensis), bur oak (Quercus macrocarpa), and honey locust (Gleditsia triacanthos). The control study area was located approximately 7 km northwest of Manhattan and consisted of 95% tallgrass prairie with the balance a mix of woody draws and riparian woodland. The grassland of the control area was similar in topography and vegetative composition to that of the golf construction area. Construction of the golf began in the summer of We collected data on avian communities for one year before construction activity (1997 summer, winter), two years during construction (1998 and 1999 summers, and winters), and one year after construction (2000 summer, winter). Construction of the golf progressed in a systematic manner, with trees removed first to make way for heavy earth-moving equipment. The area was landscaped to the architect s specifications and covered with topsoil to the ground surveyor s specifications, and altered areas were covered with grass sod within 10 days of initial disturbance. Lawngrass (Zoysia spp.) sod was used to cover the tees and fairways, L93 creeping bentgrass (Agrostis stolonifera) sod was laid on greens, and tall fescue (Festuca spp.) and bluegrass (Poa spp.) sod was laid in cart paths and rough areas. Other disturbed areas were seeded to a blend of native grasses including big bluestem, little bluestem, Indian grass, sideoats grama (Bouteloua curtipendula), blue grama (B. gracilis), pitcher sage (Salvia azurea), purple prairie clover (Dalea purpurea), and Maximillian sunflower (Helianthus maximiliani) and overlaid with erosion-preventing nets. Of the 129 ha golf area (27 holes, clubhouse, practice range, parking areas, and so on), 45 ha was turfed (18 ha = greens, tees, and fairways; and 27 ha = primary rough), and 84 ha was seeded to native grasses. The figure 8 shaped

4 186 The Prairie Naturalist 37(4): December 2005 golf was surrounded by 357 ha of undisturbed native tallgrass prairie. Approximately 50 ha (32%) of the 129 ha golf site had been altered by the end of summer 1998, 80 ha (62%) by the end of winter , and all 129 ha had been altered by the end of summer Road work, pond development, construction of buildings, and final landscaping work continued through winter , with most work being completed by late spring Avian communities on the site of the golf and the control area were monitored by using unlimited distance point counts (Reynolds et al. 1980, Verner 1985). In spring 1998, 12 permanent survey locations were established randomly in the grasslands of each of the two study areas. In addition, six permanent survey locations were established in typical woody draws and six in the riparian woodlands of the golf area (sufficient woody draw and riparian woodland habitat was not available for randomization of survey locations in those habitats on the golf site). Lack of sufficient woody draw or riparian woodland habitat precluded establishing survey locations in those habitats on the control study area. Avian point counts were conducted at each survey location in both June and July (summer) and in both January and February (winter) of each year of the study. At least one point count survey was conducted at each survey location during each designated month of each year; additional point count surveys were conducted during summers (1998, 1999, and 2000) and winters ( , , and ) when funds and personnel permitted. Each point count consisted of a 5 min quiet waiting period, followed by a second 5 min period during which all birds heard or seen (except those flying over) were counted. Point counts were conducted between sunrise and 3 hr after when winds were less than 16 kmph and no precipitation was falling. Care was taken not to double count individual birds. Differences between the abundances of individual avian species counted in the golf site and the control area were assessed by using the Wilcoxon signed rank test (Ott 1993). Additionally, counts of the 10 most abundant avian species on the golf site during the summer of 1997 (before construction) were compared to counts of those same species during the summer of 2000 (after construction) by using the Wilcoxon signed rank test to compare the changes in the counts of the respective species in the control area to the changes in the counts in the golf site. A similar comparison was done for the 10 least abundant avian species counted during summers. Similar comparisons were made for winter avian count data, but because there were fewer than 20 avian species (only 17) observed during the winter at the golf site, the comparisons were based on the eight most abundant and the nine least abundant avian species counted. The similarities of grassland avian community composition between the golf and control sites within years and between years were compared by using

5 Robel et al.: Birds and golf construction 187 Jaccard s Coefficient of Community (Brower et al. 1989) and Morisita s Index of Community Similarity (Brower et al. 1989, Krebs 1989). We used both indices because of their intrinsic differences. Jaccard s Coefficient of Community (JCC) is a simple ratio of species unique within and common between communities whereas Morisita s Index of Community Similarity (MIS) factors in species abundance and sample size, as well. Both indices provide values of 0 to 1 with 0 indicating no similarity and 1 reflecting perfect similarity of communities. The Wilcoxon signed rank test and community similarity indices also were used to determine year to year changes in point count survey data from woody draw and riparian woodland habitats between years on the golf. Winter Survey Data RESULTS Twenty four grassland, six woodland, and six woody draw point count locations were monitored in January and in February (once each month in ; twice each month in , , and ) for 5 min increments, totaling 474 unlimited distance point count surveys. During the surveys, 7,773 birds of 53 avian species were counted in the control and golf areas. Surveys in the grassland areas recorded 4,493 birds (42 species) [2,565 birds (38 species) in the control area; 1,928 (40 species) in the golf area], whereas surveys in woodland and woody draw areas in the golf tallied 1,997 birds (29 species) and 1,283 birds (30 species), respectively. No avian species listed in Kansas as Threatened and Endangered (Kansas Department of Wildlife and Parks 2005a) or Species in Need of Conservation (Kansas Department of Wildlife and Parks 2005b) were observed on the control grassland or golf study areas during our study. During winter (before construction), mean numbers of birds counted during point count surveys in grasslands, woody draws, and woodland sites on the golf were similar (Table 1). The mean number of birds counted in the grassland of the control area was very similar to that in the grasslands of the golf during that first winter. The number of avian species with larger counts in the grassland of the golf or the grassland of the control area were very similar (13 and 11, respectively) during the winter (Table 2). Avian species richness was 20 and 19 in the grassland of the golf and control areas, respectively, during the winter (Table 3). Bird counts were less in all areas during the winters of , , and than during the winter, except for the woodland area in During the and winters (construction years), mean numbers of birds counted during point count surveys in grassland areas of the golf were fewer (P < 0.05) than on the control grasslands (Table 1).

6 188 The Prairie Naturalist 37(4): December 2005 Table 1. Mean number of birds counted during 5 min point count surveys on the golf and control areas during summer and winters, 1997 to Control area Golf area Period Grassland Grassland Woody draw Woodland Winter Summer Likewise, numbers of avian species counted in greater numbers in the grasslands of the golf were significantly fewer (P < 0.05) than on the grasslands of the control area during the winters of and (Table 2). Avian species richness was higher in the control grassland during the winters of and (25 and 23, respectively) than in the golf grassland during those winters (15 and 22, respectively) (Table 3). During the winter (first year after construction) the mean number of birds tallied during point count surveys in grassland, woody draws, and woodland areas of the golf increased by 50 to 90% over the counts made during the construction winters of and (Table 1). The mean number of birds counted in grasslands of the control area decreased slightly from the previous two winters, and did not differ from counts on the grassland areas of the golf during the winter (Table 1). In winter , the number of bird species with higher numbers was greater in the grasslands of the control area than in the golf (21 and 4, respectively; P = 0.007) (Table 2). Avian species richness was 26 in the control grassland during the winter whereas it was 20 in the grassland of the golf area (Table 3). The 10 most frequently tallied birds during point count surveys in grasslands constituted 86.9% of the total birds counted (3,905 of 4,493) during the winters of

7 Robel et al.: Birds and golf construction 189 Table 2. Number of avian species counted, and those with greater numbers, during point count surveys in grasslands on the golf and control areas during summer and winter, 1997 to Period Winter Species counted Species with greater counts at 1 Control area Golf area P 2 Coefficient of community < Summer Species counted on both areas that were in significantly (P < 0.05) greater numbers on one of the two areas. 2 Probability that species in greater numbers differed between areas (Wilcoxon signed rank tests). 3 Jaccard s Coefficient of Community: 0.0 = no similarity, 1.0 = complete similarity through The American crow (Corvus brachyrhynchos) was the most frequently counted bird (39.7% of top 10 total), followed by the American robin (Turdus migratorius) (25.1%), European starling (Sturnus vulgaris) (7.8%), eastern bluebird (Sialia sialis) (7.2%), blue jay (Cyanocitta cristata) (6.6%), northern cardinal (Cardinalis cardinalis) (4.5%), and American goldfinch (Carduelis tristis) (2.8%). The meadowlarks (Sturnella magna and S. neglecta), tufted titmouse (Baeolophus bicolor), and northern flicker (Colaptes auratus) each contributed greater than 2.5% to the total counts of the 10 most abundant avian species. The point count survey data reflected considerable year to year and area to area variation in abundances of different avian species during the winters on the two study areas (Table 3). The 10 most frequently tallied birds during point counts in woody draws of the golf contributed 85.8% of the total birds counted in woody draws (1,713

8 190 The Prairie Naturalist 37(4): December 2005 Table 3. Numbers of avian species counted (species richness) and individual numbers of the 10 most abundant species during point count surveys on grasslands of the control and golf areas during winters of to , and Morasita s Index of Similarity (MIS) for those 10 species communities. Species 1 Control Golf Control Golf Control Golf Control Golf AMCR AMGO AMRO BLJA EABL EUST MEAD NOCA NOFL TUTI Richness MIS American Bird Banding codes (see Appendix A for common names), scientific names in text. 2 Includes eastern and western meadowlarks. 3 Total numbers of avian species counted during point count surveys. of 1,997) during the winters of through The American crow contributed the most (20.2%) to the total numbers of the top 10 species, followed by the American robin (14.1%), American goldfinch (12.4%), eastern bluebird (10.0%), black-capped chickadee (Poecile atricapillus) (8.5%), dark-eyed junco (Junco hyemalis) (8.3%), meadowlarks (8.3%), cedar waxwing (Bombycilla cedrorum) (8.0%), American tree sparrow (Spizella arborea) (5.9%), and European starling (4.3%). The 10 most frequently recorded birds during winter point count surveys in woodlands of the golf contributed 97.2% of the total birds counted in woodlands (1,940 of 1,997). The American robin was the most

9 Robel et al.: Birds and golf construction 191 frequently counted bird in the woodlands of the golf, contributing 42.1% of the top 10 total, followed by the American crow (13.9%), blue jay (10.4%), cedar waxwing (7.7%), black-capped chickadee (7.5%), northern cardinal (5.7%), darkeyed junco (5.6%), eastern bluebird (3.2%), American tree sparrow (2.2%), and American goldfinch (1.8%). Winter point count survey data detected large year to year variation in abundances of different avian species in woody draws and woodlands of the golf (see Bye 2001 for details). Of the eight most abundant avian species counted during winter in the golf grassland before construction ( ), seven were less abundant during the winter after construction ( ) (Table 5). Of the same eight species in the control grassland area during the winter, five were less abundant during the winter. Analysis of the changes in numbers of the eight most abundant avian species detected no difference (P = 0.922) between the before and after construction differences in avian numbers in the control and golf grassland areas. Of the nine least abundant avian species counted during winter in the golf grassland before construction ( ), numbers of the dark-eyed junco, downy woodpecker (Picoides pubescens), Harris s sparrow (Zonotrichia querula), meadowlarks, and tufted titmouse counted were lower during the winter after construction ( ); American tree sparrow, black-capped chickadee, and house finch (Carpodacus mexicanus) were higher; and the number of red-tailed hawk (Buteo jamaicensis) counted was unchanged (Table 5). Analysis of the changes in numbers of the nine least abundant avian species detected no difference (P = 0.461) between the before and after construction differences in avian numbers in the grasslands of the control and golf areas. The bird communities of the grasslands of the two study areas were less similar during winters of and (JCC values of and 0.538, respectively) than during the winters of and (JCC values of and 0.643, respectively) (Table 2). Avian assemblages composed of the 10 most commonly counted birds in the grasslands of the golf and control areas were more similar during the winters of and (MIS = 0.938) than in the winters of (MIS = 0.850) and (MIS = 0.458) (Table 3). Summer Survey Data The 24 grassland, 6 woodland, and 6 woody draw point count locations were monitored in June and in July (once each month in 1997, four times each month in 1998, and twice each month in 1999 and 2000) for 5 min increments, providing 648 point count surveys and tallying a total of 11,307 birds of 78 species in the control and golf areas. Surveys of the grassland areas recorded 7,980 birds (67 species) [4,162 birds (60 species) in the control area; 3,818 birds (56 species) in the

10 192 The Prairie Naturalist 37(4): December 2005 golf area], whereas surveys in woodland and woody-draw sites on the golf area tallied 1,546 birds (57 species) and 1,781 birds (65 species), respectively. During summer 1997 (before construction), mean numbers of birds counted during point count surveys in grasslands, woody draws, and woodland sites on the golf area ranged from to 23.05, and the mean number of birds counted in the grasslands of the golf and control areas were very similar (Table 1). The number of avian species having larger counts on the grasslands exceeded that for the golf area but the difference was not significant (P = 0.129). (Table 2). Avian species richness was 32 in the grassland of the control area compared to 27 in the golf grassland (Table 4). Numbers of birds counted decreased in all areas of the golf in summer of 1998 (first year after construction), and in the grassland control area (Table 1). The numbers of avian species with larger numbers in grasslands were significantly greater (P = 0.028) on the control area than on the golf area during summer 1998 (Table 2). Species richness in the control grassland was 56 during the summer of 1998, considerably greater than the 38 in the grassland of the golf. The number of birds counted during 1999 summer point count surveys increased on both study areas, with the greatest number being recorded on the grasslands of the control area (Table 1). The numbers of avian species with greater numbers on grasslands of the control or golf area were almost equal in summer 1999 (Table 2), and species richness was very similar as well (Table 4). During summer 2000 (first year after construction), numbers of birds counted during point count surveys approximated preconstruction (summer 1997) counts (Table 1) and numbers of avian species with larger counts did not differ (P = 0.132) between the control grasslands and the golf area (Table 2). Species richness in the control grassland was 48 compared to 43 in the grassland of the golf area during summer 2000 (Table 4). The 10 most frequently tallied birds during point count surveys in grasslands constituted 75.3% of the total birds counted (6,012 of 7,980) during summers of 1997 through Meadowlarks were the most frequently counted (16.1% of top- 10 total), followed by the dickcissel (15.1%), northern bobwhite (Colinus virginianus) (14.2%), grasshopper sparrow (13.2%), brown-headed cowbird (Molothrus ater) (12,9%), American crow (10.9%), northern cardinal (7.1%), field sparrow (Spizella pusilla) (5.0%), blue jay (3.0%), and upland sandpiper (Bartramia longicauda) (2.5%). The point count survey data reflected considerable year to year and area to area variation in abundances of different avian species during the summers on the two study areas (Table 4). The 10 most frequently tallied birds during the point count surveys in woody draws of the golf contributed 65.8% of the total birds counted in woody draws (1,172 of 1,781) during the summers of 1997 through The northern

11 Robel et al.: Birds and golf construction 193 Table 4. Total numbers of avian species counted (species richness) and numbers of the 10 most abundant species during point count surveys of grasslands of the control and golf areas during summers of 1997 through 2000, and Morasita s Index of Similarity (MIS) for those 10 species communities. Species 1 Control Golf Control Golf Control Golf Control Golf AMCR BHCO BLJA DICK FISP GRSP MEAD NOBO NOCA UPSA Richness MIS American Bird Banding codes (see Appendix A for common names), scientific names in text. 2 Includes eastern and western meadowlarks. 3 Total numbers of avian species counted during point count surveys. bobwhite contributed the most (15.9%) to the total numbers of the top 10 species, followed by the dickcissel (14.1%), brown-headed cowbird (11.5%), meadowlarks (10.3%), northern cardinal (10.1%), American crow (9.1%), field sparrow (8.8%), grasshopper sparrow (8.8%), black-capped chickadee (6.0%), and American goldfinch (5.4%). The 10 most frequently recorded birds during point count surveys in woodlands of the golf contributed 62.4% of the total birds counted in woodlands (965 of 1,546). The northern cardinal, contributing 20.0% of the top 10 total, was the most frequently counted bird in the woodlands of the golf, followed by the American crow (15.9%), northern bobwhite (13.4%), field sparrow

12 194 The Prairie Naturalist 37(4): December 2005 Table 5. Numbers of the most and least abundant avian species counted during winter point count surveys in the grasslands of the control and golf areas before ( ) and after ( ) construction of a naturalistic golf in tallgrass prairie in Kansas. Species 1 Control Golf Most abundant species Before construction After construction Differences Control Golf Control Golf AMCR AMGO AMRO BLJA EABL EUST NOCA NOFL Least abundant species ATSP BCCH DEJU DOWO HASP HOFI MEAD RTHA TUTI American Bird Banding codes (see Appendix A for common names), scientific names in text. 2 Includes eastern and western meadowlarks. (10.4%), brown-headed cowbird (9.7%), black-capped chickadee (8.8%), dickcissel (6.7%), American robin (5.6%), mourning dove (Zenaida macroura) (5.1%), and American goldfinch (4.8%). Summer point count survey data detected large year-

13 Robel et al.: Birds and golf construction 195 to-year variation in abundance in different avian species in woody draws and woodlands of the golf (see Bye 2001 for details). Of the 10 most abundant avian species counted during summer in the golf grassland before construction (1997), the brown thrasher (Toxostoma rufum) and grasshopper sparrow were less abundant during the summer (2000) after construction (Table 6). Of the same 10 species in the control grassland area, none were less abundant during summer 2000 than summer Analysis of the numbers of the 10 most abundant avian species detected no difference (P = 0.846) between the before- and after-construction differences in avian numbers in the control and golf grassland areas. Of the 10 least abundant avian species counted during summer in the golf grassland before construction (1997), the Carolina wren (Thryothorus ludovicianus), red-bellied woodpecker (Melanerpes carolinus), northern mockingbird (Mimus polyglottos), and red-tailed hawk exhibited little to no change in numbers, whereas the barn swallow (Hirundo rustica), blue grosbeak (Guiraca caerulea), common grackle (Quiscalus quiscula), eastern bluebird, mourning dove, and ring-necked pheasant (Phasianus colchicus) increased (Table 6). Of the same 10 species in the control grassland area; counts of the blue grosbeak and mourning dove decreased; barn swallow, eastern bluebird, northern mockingbird, and ringnecked pheasant increased; and Carolina wren, common grackle, red-bellied woodpecker, and red-tailed hawk essentially were unchanged. Analysis of the numbers of the 10 least abundant avian species detected no difference (P = 0.398) between the before- and after-construction differences in avian numbers in the control and golf grassland areas. Bird communities of the grasslands of the golf and control area during summer 1997 were more similar (JCC = 0.794) than during other summers, with similarities being least in summer 1998 (JCC = 0.541) (Table 2). Avian assemblages composed of the 10 most commonly counted birds on the grasslands of the golf and control area varied little during the summers of 1997 through 2000 with MIS values ranging from (summer 2000) to (summer 1999) (Table 4). DISCUSSION We acknowledge that our study was an observational study, not truly experimental. Because replications of disturbance were not performed, no causal relationships can be established. Additionally, because of the lack of replications of study areas and because our point count survey data were not adjusted for bird detection probabilities, we were reluctant to attempt species specific statistical analysis of our data, i.e., trying to determine impacts of golf construction on populations of individual avian species. However, the changes we observed in

14 196 The Prairie Naturalist 37(4): December 2005 Table 6. Numbers of the most and least abundant avian species counted during summer point count surveys in the grasslands of the control and golf areas before (1997) and after (2000) construction of a naturalistic golf in tallgrass prairie in Kansas. Species 1 Control Golf Most abundant species Before construction After construction Differences Control Golf Control Golf AMCR BHCO BRTH DICK FISP GRSP MEAD NOBO UPSA Least abundant species BARS BLGR CAWR COGR EABL MODO NOMO RBWO RNPH RTHA American Bird Banding codes (see Appendix A for common names), scientific names in text. 2 Includes eastern and western meadowlarks.

15 Robel et al.: Birds and golf construction 197 avian communities provided an insight into the impacts associated with the establishment of a naturalistic golf in native tallgrass prairie. Grassland Survey Sites - Winter Point count surveys detected decreases in avian abundance in grasslands of both study areas during the second winter of the study. The decreased avian abundance on our two study areas was consistent with a 50 to 60% regional decline in avian populations detected in Christmas Bird Counts and Breeding Bird Surveys. Such year to year changes are not unexpected (Wiens and Dyer 1975). But the decreases were greater on the golf area where construction was ongoing. Reduced avian abundance continued in the golf grassland through the third year (second year of construction). During the final year of the study (after construction), point count surveys detected similar numbers of birds in the grasslands of the control and golf areas. However, species richness was lower in the golf grassland than the control grassland after construction of the golf. We were not surprised to detect smaller numbers of birds on the golf grassland area during construction (second and third years of the study) than on the control area because of the extensive disturbance and habitat alteration caused by heavy earth moving equipment on the golf. Six of the 12 point count survey sites in the grasslands of the golf were physically altered (or land was bulldozed within 25 m of a survey site) during construction. Postconstruction point count surveys reflected similar avian abundance on grasslands of the undisturbed control area and the altered grassland of the golf area. The recovery of bird numbers on the altered golf grassland was unexpected. Approximately 30% of the grassland of the golf area was modified during construction, and most of that was covered quickly with turf grass sod or reseeded to native grasses. Our postconstruction point count data from the golf could have represented a recovery of avian abundance evenly across the altered and unaltered grasslands, reflected an increase of bird numbers in the remaining undisturbed areas and masked reduced numbers in altered areas, or have been a combination of both. During the preconstruction year, the numbers of avian species with greater abundance on the grasslands of the two study areas was approximately equal. During the construction and postconstruction years, more avian species were more abundant in the grasslands of the control area than on the golf grasslands. Relative to their preconstruction abundances, eight avian species (American crow, American goldfinch, American tree sparrow, dark-eyed junco, eastern bluebird, European starling, greater prairie-chicken, and northern cardinal) declined more during the construction years on the golf grassland than on the control area; the abundance of the killdeer (Charadrius vociferus) increased on the

16 198 The Prairie Naturalist 37(4): December 2005 construction area, whereas it declined on the control area. The greater prairiechicken was observed on the golf area before construction of the golf began, but was not observed on the area once construction started, probably because it is very sensitive to human disturbance (Robel et al. 2004). Killdeer most likely became more abundant on the golf grassland because of the increased amount of open area created by construction activity (Bye 2001). Overall, differences in numbers of the most and least abundant avian species did not differ between point count surveys conducted in the golf grassland before and after construction of the grassland and the grassland of the control area. The number of avian species (species richness) recorded during winter point count surveys on the control and golf grassland areas were similar during the preconstruction winter (19 and 20 species on the areas, respectively). During construction and postconstruction winters, avian species richness was less on the golf grasslands than on the control grassland (0 = 19.0 and 24.7 species, respectively, for the three winters). The difference in species richness might be an artifact of our data because the unlimited distance point count method we used recorded birds beyond grassland habitat (e.g., European starling and various woodpeckers in adjacent woody draws and woodlands). Even so, the reduced species richness we detected on the golf grassland area could be symptomatic of human disturbance as observed by Van der Zande and Vos (1984), Gutzwiller et al. (1998), Fernandez-Juricic (2000) and others, but differs from Blair (1996), who reported avian species richness greatest on golf s in an urban ecological gradient in California. Grassland Survey Sites - Summer Point count surveys detected decreases in total avian abundance on grasslands of the control and golf areas between the first summer of the study (before construction) and the second summer (first year of construction on the golf area). Avian abundance returned to preconstruction numbers by the third year (second year of golf construction) on both areas, and remained at that level in the following year. Construction activity on the golf grassland had no measurable effect on total bird abundance, compared with that on the grassland of the control area. The limited avian species richness on the golf grassland in the second summer of the study might have been associated with construction activity on that area. More bird species also were more abundant on the control than the golf grassland that summer, again a possible effect of construction activity on the golf area. Recovery of the avian species richness and of the number of species with greater abundances on the golf during the second summer of construction might have reflected either habituation (Whittaker and Knight 1998)

17 Robel et al.: Birds and golf construction 199 or less tolerant individuals being replaced by more tolerant individuals (Gutzwiller et al. 1994, Riffell et al. 1996) by the second summer of construction. After the first year of construction on the golf, the species found most often in the grasslands of the two study areas became more similar over time. The similarity of avian communities on the grasslands of the control and golf areas remained high for most of the study, with a drop in the MIS index during the last year, primarily because of a shift, for unknown reasons, in numbers of American crow and American robin. Many of the most abundant avian species found in the grassland areas during summer were tolerant of human activity (DeGraaf and Wentworth 1986) and, therefore, would not be expected to decrease in numbers on the golf grassland during construction. Blair (1996) determined that several avian species were suburban adaptable, of those he listed that were counted in our golf grassland, the American robin, brown-headed cowbird, mourning dove, and house finch increased on our golf grassland between the time before and after construction but the European starling and northern mockingbird did not. Several of the least abundant species recorded on the golf grassland during point count surveys were tree or cavity nesters in woody draws and woodlands adjacent to the grasslands. The numbers of these birds (various woodpeckers, northern flicker, American robin, tufted titmouse, brown thrasher, etc.) declined in point count surveys on the golf grassland as woody draws and woodlands were destroyed by construction activity. Overall, differences in numbers of the most and least abundant avian species did not differ between point count surveys conducted on the golf grassland before and after construction of the naturalistic golf and the same species on the control grassland. Woody Draws and Woodlands - Winter Because we had no data from control areas to which point count data from the woody draws and woodlands of the golf could be compared, any construction related association inferences are weak at best. Point count surveys in woody draws detected similar avian abundance trends as did those in grassland sites of the golf area. Data from point count surveys in the woodland more closely resembled those from the control grassland except for higher avian abundance during the postconstruction year. Woody vegetation at three of the six point-count survey sites in woody draws was bulldozed out during construction of the golf. The sites were sodded or reseeded to grass, thus eliminating the woody vegetation at those sites. More than 60% of the woody draw habitat on the golf area was eliminated during construction. The abundances of the American tree sparrow, blue jay, tufted titmouse, meadowlark, and downy woodpecker each were less than 15% of the preconstruction numbers of those birds in the woody draws after construction began on the golf.

18 200 The Prairie Naturalist 37(4): December 2005 Trees were removed from the vicinity of three of the six point count survey sites in the woodlands of the golf during the first year of construction. The number of birds detected during point count surveys of the woodlands after the removal of the trees did not seem to be impacted by the construction activity. Although woodland habitat in the vicinity of 50% of our survey sites was destroyed, this amounted to less than 20% of the total woodland habitat on the golf area. Point count surveys of woodland sites detected decreases of more than 85% from the preconstruction numbers for the American tree sparrow and blue jay during construction activity on the golf. Woody Draws and Woodlands - Summer Point count surveys in woody draw and woodland habitats during summer on the golf reflected decreased avian abundance during the construction years compared with preconstruction and postconstruction numbers. It is not clear if these depressed numbers were associated with construction activities per se, or simply represent normal annual fluctuations in bird numbers. Postconstruction abundances of the European starling, hairy woodpecker (Picoides villosus), house finch, house sparrow (Passer domesticus), and house wren (Troglodytes aedon) in woody draws were less than 15% of the preconstruction abundance, whereas the American goldfinch, brown-headed cowbird, and killdeer exhibited a threefold increase. Postconstruction abundances of the American redstart (Setophaga ruticilla), American robin, northern flicker, and white-breasted nuthatch (Sitta carolinensis) in the woodlands of the golf were less than 20% of their preconstruction abundance, whereas the American goldfinch, brown-headed cowbird, brown thrasher, dickcissel, killdeer, and meadowlarks exhibited a three- to tenfold increase in numbers. Some of these changes in abundance mght have been associated with construction activity on the golf, but we could not conclude such because of the lack of comparable control areas in our study. ACKNOWLEDGMENTS Our study was supported by the Kansas Agricultural Experiment Station (Contribution No J); the Kansas Department of Wildlife and Parks; and the Office of the President and the Division of Biology, Kansas State University. We acknowledge Kimberly Wolcott, Luke Westerman, Jethro Runco, Darel Zimmerman, and Christian Hagen for assistance in the field and Clenton Owensby for access to Donaldson Pastures.

19 Robel et al.: Birds and golf construction 201 Appendix A. American Bird Banding codes used in tables and common names of birds. Code Common name Code Common name AMCR American crow EUST European starling AMGO American goldfinch FISP Field sparrow AMRO American robin GRSP Grasshopper sparrow ATSP American tree sparrow HASP Harris s sparrow BARS Barn swallow HOFI House finch BCCH Black-capped chickadee MEAD Meadowlark (eastern and western) BHCO Brown-headed cowbird MODO Mourning dove BLGR Blue grosbeak NOBO Northern bobwhite BLJA Blue jay NOCA Northern cardinal BRTH Brown thrasher NOFL Northern flicker CAWR Carolina wren NOMO Northern mockingbird COGR Common grackle RBWO Red-breasted woodpecker DEJU Dark-eyed junco RNPH Ring-necked pheasant DICK Dickcissel RTHA Red-tailed hawk DOWO Downy woodpecker TUTI Tufted titmouse EABL Eastern bluebird UPSA Upland sandpiper LITERATURE CITED Audubon International Selkirk, New York (accessed 6 June 2005). Best, L. B., H. Campa, III, K. E. Kemp, R. J. Robel, J. A. Savage, H. P. Weeks, Jr., and S. R. Winterstein Bird abundance and nesting in CRP fields and cropland in the Midwest: a regional approach. Wildlife Society Bulletin 25: Blair, R. B Land use and avian species diversity along an urban gradient. Ecological Applications 6: Brower, J., J. Zar, and C. von Ende Field and laboratory methods for general ecology. Wm. C. Brown Publishers, Dubuque, Iowa.

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21 Robel et al.: Birds and golf construction 203 Robel, R. J., J. A. Harrington, Jr., C. A. Hagen, J. C. Pitman, and R. R. Reker Effects of energy development and human activity on the use of sand sagebrush habitat by lesser prairie-chickens in southwestern Kansas. Transactions of the North American Wildlife and Natural Resources Conference. 69: Samson, F. B., and F. L. Knopf Prairie conservation in North America. BioScience 44: Thien, S., S. Starrett, R. Robel, P. Shea, D. Gourlay, and C. Roth A multiple index environmental quality evaluation and management system. United States Golf Association Green Section Record 39: Van der Zande, A. N., and P. Vos Impact of a semi-experimental increase in recreation intensity on the densities of birds in groves and hedges on a lake shore in the Netherlands. Biological Conservation 30: Verner, J Assessment of counting techniques. Current Ornithology 2: Whittaker, D., and R. L. Knight Understanding wildlife responses to humans. Wildlife Society Bulletin 26: Wiens, J. A., and M. I. Dyer Rangeland avifaunas: their composition, energetics, and role in the ecosystem. Proceedings of the symposium on management of forest and range habitats for nongame birds. United States Department of Agriculture Forest Service, General Technical Report WO- 1: Received: 8 November 2004 Accepted: 6 November 2005 Associate Editor for Ornithology: Gregory A. Smith