'gin ,vada. Some Effects of a Campground l and on Breeding in

Stewart W. Aitchison:;

Abstract.--Over a three year period, breeding den­ sities were found to be similar between a constructed camp­ ground and a relatively natural area when the campground was closed to campers. However, bird species composition differ­ ed between sites, the campground having relatively heavier bodied birds (x = 48.5 g) than the control area (x = 38.2 g). Once the campground was opened for human use, the breeding bird population decreased in density and diversity. On the he control site the population either remained the same or -423. increased. ultures. Vol. INTRODUCTION Financial support for this study was par­ tially provided by the u.S. Forest Service. It has been well documented that the Coconino National Forest,and to this end I human manipulation of Southwestern habitats wish to thank B. Burbridge and W. Finley. Also, greatly affects the configuration of the avian the Sedona Ranger District graciously allowed community that will continue to utilize the us to set up the two study plots in Creek area (e.g., Carothers et al. 1974. Carothers Canyon. and Johnson 1975). These studies have prima­ rily concerned themselves with phreatophyte control, channelization, and other water STUDY AREA management practices. Very little research has dealt with the impact caused by the con­ Two sites were chosen which appeared sim­ struction of permanent structures and human ilar in vegetative structure, each being com­ occupation of these areas (e.g., subdivisions, posed primarily of ponderosa with some trailer parks, and campgrounds). The present cottonwood, Arizona walnut and other deciduous study examines the effects of a U.S. Forest trees and shrubs. (Table 1 summarizes the Service improved campground upon breeding birds. vegetation analysis of the study areas.) Both sites are within Oak Creek Canyon, Sec. 27, T19N, R6E, Coconino County, Arizona, at an ACKNOWLEDGEMENTS elevation of 1,646 m. One plot was located within the Cave Springs Campground; the other The author wishes to thank M.E. Theroux plot, control, was slightly north and across for collecting and processing the vegetation the Oak Creek. data, S.W. Carothers and O.J. Reichman for The Cave Springs Campground study site critically proofing the manuscript, and many is 4.0 ha. Extensive timber and shrub removal, other members of the Museum of Northern Arizona construction of roads, pit toilets, and erection staff for their help and encouragement. Special of tables has occurred here. The campground is thanks goes to B. Johnston who did nearly all open for public use from approximately Memorial the field work in 1974 and a substantial amount Day to Labor Day of each year. in 1975. Control is approximately 2.0 ha. Relative to the campground, this area is undisturbed by l/Contributed paper, Symposium on the human activities. Imp0lrtance, Preservation and Management of the Though the two sites are small in area, thus Riparian Habitat, July 9, 1977, Tucson, Arizona. making extrapolation of figures somewhat mis­ 2/ Research Assistant, Museum of Northern leading, they encompass as much homogeneous Arizona, Route 4, Box 720, Flagstaff, Arizona habitat as possible. Further, the small size 86001. enabled the investigators to know the avian components .intimately, and therefore we feel a very accurate count of species was achieved.

175 Table l.--Vegetation Analysis

Tree Basal Average Study Dens~ty Area (m2) Tree Area Per Per Height (m) Hectare Hectare Ponderosa Pine Pinus Ponderosa 437.1 2153.6 12.4 CAMPGROUND *All other species 316.5 415.0 7.3

Ponderosa Pine 347.8 2118.4 14.0 CONTROL **All other species 1010.7 447.8 5.0

* Acer negundo, Alnus oblongifolia, Juniperus scopulorum, Quercus gambelli, Populus lanceolata, Salix gooddingii.

** Acer negundo, Alnus oblongifolia, Fraxinus.pennsylvanica var. velutina, Juniperus scopulorum, ~. monosperma, Pinus edulis, Populus lanceolata, Quercus spp.

METHODS (see Shannon and Weaver 1963) as it applies to biological parameters (MacArthur and MacArthur The populations of breeding birds were 1961; MacArthur 1965; Pielou 1966a,b; Lloyd determined by the spot-map method (Williams et al. 1968). 1936, Kendeigh 1944). On each census the Tree density, species composition and basal location of singing males, song posts, and area were determined by the plotless point­ nest sites was recorded for each census and quarter method of Cottam and Curtis (1956). information on every species was later recorded Tree heights were determined by use of a clino­ onto species maps. Censusing was carried out meter. Samplings witha diameter at breast from 4 April to 6 July 1973, 18 February to height (DBH) of less than 7.6 cm were treated 1 July 1974, and 9 May to 10 July 1975; these as shrubs. periods included the entire observable breeding Avian standing crop biomass (SCB) was season. Densities were determined for each determined by taking the average adult weight area before and after the date the campground (W) times the number of adults per unit area. was opened (e.g., 29 May 1973, 17 May 1974, Existence Energy (EE) or the amount of kcal and 16 May 1975). [Note: All densities were consumed per ha per 24 hours was calculated from extrapolated to 40 ha to make inter-area these two formulae: comparisons easier.] LogEE 0.3581 + 0.5876 Log W (for ) Foliage height diversity (FHD) was sampled Log EE = 0.0649 + 0.6722 Log W (for non-passer­ along ten 100 m transects established at random ines) • throughout the study areas. Presence or absence These formulae give the energy requirements of vegetation at 2 m intervals along the trans­ to maintain a constant weight at rest. To ects was noted at three layers chosen to determine actual community energetics it would approximate foliage stratification into be necessary to include energy requirements of herbaceous (0-0.6 m), shrub (0.6-4.49 m) and the immature birds, and the various energy canopy (:>4.5 m) layers. A 4. 5 m rod marked demanding activities of breeding birds (e.g., at 0.6 m from one end was used to record the singing, displaying, nest building). The presence or absence of foliage for the herba­ limitations of this procedure notwithstanding, ceous and shrub layers, and the ocular tube it is instructive to make inter- and intra­ method (Winkworth and Goodall 1962) was used community comparisons with these low estimates for the canopy layer. For recording the pres­ of avian community energetics (see Karr 1968 sence of vegetation in the herbaceous and and 1971). shrub layers, it was necessary for green foliage to touch the vertically held rod. All vegetative and avian diversity indices RESULTS AND DISCUSSION are computed as H' .. - ~ P. log P. based on the Shannon-Wiener model of inloimation theory A total bf 58 species (Table 2) of birds

176 .-;. ;:t . j.:. • 11 .q en 6 - ill P,":"ni.o I o.r+1Il 6>/ii IF· (I) .... I!J I!" P,

Table 3.--Avian density and species richness

1973 1974 1975 SPECIES CONTROL CAMPGROUND CONTROL CAMPGROUND CONTROL CAMPGROUND. Before After Before After Before After Before After Before After Before After Mourning Dove 9.9 9.9 Broad-tailed Hummingbird 19.8 19.8 24.7 19.8 9.9 9.9 19.8 19.8 9.9 9.9 Red-shafted Flicker 9.9 9.9 9.9 9.9 9.9 19.8 Hairy Woodpecker 9.9 19.8 9.9 19.8 9.9 Yellow-bellied Sapsucker 9.9 9.9 Cassin's Kingbird 19.8 Black Phoebe 19.8 19.8 Western Flycatcher 19.8 19.8 Western Wood Pewee 39.5 9.9 19.8 19.8 39.5 Steller's Jay 39.5 19.8 49.5 24.7 19.8 19.8 19.8 19.8 19.8 19.8 19.8 White-breasted Nuthatch 9.9 9.9 29.7 19.8 Pygmy Nuthatch 19.8 19.8 19.8 9.9 19.8 19.8 9.9 19.8 House Wren 59.3 59.3 39.6 39.6 19.8 39.5 39.6 29.7 59.3 59.3 19.8 39.6 Robin 39.5 39.5 44.5 29.7 19.8 19.8 29.7 19.8 39.5 29.7 39.6 Solitary Vireo 49.4 49.4 9.9 19.8 9.9 9.9 19.8 19.8 Warbling Vireo 9.9 19.8 Virginia's Warbler 39.5 .­-.. -.. Grace's Warbler 9.9 19.8 19.8 9.9 29.7 19.8 Painted Redstart 9.9 19.8 39.5 9.9 Red-faced Warbler 19.8 19.8 4.9 Bullock's Oriole 19.8 19.8 14.8 14.8 19.8 19.8 19.8 19.8 19.8 Western 9.9 9.9 Hepatic Tanager 19.8 19.8 19.8 9.9 9.9 19.8 Black-headed Grosbeak 59.3 39.5 64.3 39.6 19.8 39.5 39.6 39.6 39.5 19.8 29.7 39.6 Lesser Goldfinch 19.8 Rufous-headed Towhee 19.8 19.8 Total Density 326.0 365.6 297.0 178.2 158.4 296.7 297.0 222.7 178.1 445.0 178.2 257.4 Species Richness 9 12 12 8 8 12 16 13 7 17 10 12 Weight/Individual 40.0 31. 7 52.7 54.2 46.8 23.4 39.4 41.6 27.7 32.3 53.3 52.3 SCB 26078.4 31315.7 14830.2 23397.6 9854.2 19013.8 23178.5 19316.9 13858.6 18509.0 28710.0 26918.0

11697.7 12302.4 6088.0 10166.4 4984.0 7134.0 EE 11438.6 7698.6 7785.7 8079.5 13783.8 10203.0 Table 2.--Species recorded on or immediately were seen on or immediately adjacent to the ripar:~a adjacent to the study area. [Note; Bird study areas. Of these, 23 species nested on cotton\< scientific names according to A.O.U. Checklist, one or both of the s.tudy areas at least once apparer 1957; and 32nd A.O.U. Supplement, Auk 90(2): during the three years of censusing. The On the 411-419.] density and species' richness (Whittaker 1970) c are summarized in Table 3. The changes in in 197: Red-tailed Hawk Buteo jamaicensis these values and other resultant calculations differE Turkey Vulture Cathartes fasciata prior to and after the opening of the camp­ that dE Band-tailed Pigeon Columba fasciata ground are discussed below as indicators of in wha1 Mourning Dove Zenaidura macroura human impact upon the breeding bird community. for ei1 Great Horned OWl Bubo virginianus A Flammulated OWl Otus flammeolus Avian Density and Species Richness on the White-throated Swift Aeronautes saxatalis 87.3 pI Broad-tailed Hummingbird Selasphorus playcercus Every species is apparently adjusted to was oF' Belted Kingfisher Megaceryle alcyon breed at the time of year at which it can raise may ac. Red-shafted Flicker Colaptes cafer its young most efficiently (Immelmann 1971). contro Hairy Woodpecker Dendrocopos villosus For most northern temperate birds this nesting specie Yellow-bellied Sapsucker Sphyrapicus varius period extends from late spring to mid-summer on the Western Kingbird Tyrannus verticalis (Lack 1950). This is certainly true for the satisf Cassin's Kingbird Tyrannus vociferans Cave Springs area of Oak Creek Canyon, where S Black Phoebe Sayornis nigricans breeding begins about mid-April and lasts on the Weste~n Flycatcher Empidonax difficilis through July. The campground opening date contro Western Wood Pewee Contopus sordidulus falls within this period. Violet-green Swallow Tachycineta thaIassina 1975.­ Steller's Jay Cyanocitta stelleri 1973.--In 1973 a total of 17 species nested on on one Common Raven Corvus corax one or both of the study areas (Table 3). A New br Mountain Chickadee Parus garnbeli 40 percent decrease in density occurred on Kingbi Dipper Cinclus mexicanus the campground after the opening day. Part Lesse:r White-breasted Nuthatch Sitta carolinensi~ of the losses incurred were through direct In pre Pygmy Nuthatch Sitta pYgrnaea human manipulation of the nest site. Forest appea:r Brown Creeper Certhia familiaris Service employees, by removing trees and slash, canyor House Wren Troglodytes aedon destroyed 20 percent of the Steller's Jay nests. ] Canyon Wren Catherpes mexicanus Campers destroyed 30 percent more of the increc Mockingbird Mimus polyglottos Steller's Jay nests and 20 percent of the Robin grounc Robin Turdus migratorius nests by removing branches for firewood, making was WE Catharus guttata room for tents, and other reasons. the 01 Townsend's Solitaire Myadestes townsendi The parulid warblers, Solitary Vireos, tempe] Ruby-crowned Kinglet Regulus calendula Broad-tailed Hummingbirds, and Hairy Woodpeckers preci] Solitary Vireo Vireo solitarius abandoned their nests but occasionally foraged speciE Warbling Vireo Vireo gilvus within the area. No losses can be attributed peratl Virginia's Warbler Verrnivora virginiae to adults leaving with fledged young prior to or egl Yellow Warbler Dendroica petechia the opening date. Of those actually nesting WilsOl Audubon's Warbler Dendroica auduboni on 20 May 1973, breeding had not proceeded two WI Grace's Warbler Dendroica graciae beyond the incubation stage. the s, MacGillivray's Warbler Oporonis tolmiei The density on the control site increased and 6 American Redstart Setophaga ruticilla 12.1 percent after the opening date. This was 1975 Painted Redstart Setophaga picta not due to individuals emigrating from the activ Red-faced Warbler Cardellina rubrifrons campground but rather to the arrival of mid­ build Wilson's Warbler Wilsonia pusilla summer breeders, namely, the Red-faced Warbler, Hooded Oriole Icterus cucullatus Western Wood Pewee, and Hepatic Tanager (Bent ing d Bullock's Oriole Icterus galbula 1968) • was s Brown-headed Cowbird Moluthrus ater After the opening of the campground, species 1973 ludoviciana richness went from 12 to 8 on the campground Hepatic Tanager Piranga flava and increased from 9 to 12 on control. perce Summer Tanager Piranga rubra doubt Rose-breasted Grosbeak Pheucticus ludovicianus 1974.--In 1974 a total of 17 species nested on Befo:r Black-headed Grosbeak Hesperiphona vespertina one or both of the study areas (Table 3). were Indigo Bunting Passerina cyanea However, these were not the same 17 of 1973. addit Pine Siskin Spinus pinus There was a change of three species, with nest American Goldfinch Spinus tristis Black Phoebes, Western Flycatchers and Hepatic usual Lesser Goldfinch Spinus psaltria being replaced by Pygmy Nuthatches, canyc Rufous-sided Towhee Pipilo erythrophthalmus Warbling Vireos and Summer Tanagers. This abOVE Gray-headed Junco Junco caniceps area in Oak Creek is an ecotonal situation Perh, between confierous forest and a deciduous less

178 e riparian habitat. The Summer Tanager prefers and these species chose to accept marginal on cottonwoods along streams for nest sites and habitat under these limitations. ce apparently found conditions suitable in 1974. Species richness went from 10 to 12 on the On the other hand, the Hepatic Tanager prefers campground and 7 to 17 on control. 970) pines and and found Cave Springs acceptable Yearly fluctuations of density on each in 1973. It is probably subtle environmental area are difficult to explain because of so ons differences (i.e., temperature, rainfall, etc.) many determining factors. Not only local weather that determine which tanager will be present but events on the wintering grounds can play an of in what might be considered marginal habitat important role in predicting a particular year's ity. for either. breeding population. Attempts to explain avian A 25 percent decrease in densities occurred population fluctuations have so far led to only on the campground, whereas there was a dramatic ambiguous conclusions (Von Haartman 1971). It 87.3 percent increase on control. The campground is pertinent to note, though, that over the to was opened 12 days earlier than in 1973 and three-year period there was nearly twice the raise may account for the initially low density on range of densities on control as the campground. ) . control. It was simply too early for many ting species to be breeding. Yet initial densities Avian Diversity and Habitat Diversity Iller on the campground matched 1973 figures. No :le satisfactory explanation has been found. MacArthur (1964) found a correlation between ere Species richness dropped from 16 to 13 BSD and FHD in "tall forests of sycamores and on the campground and climbed from 8 to 12 on cottonwoods" in southeastern Arizona. The control. relationship in this study between BSD and FHD was almost identical to what he and others 1975.--During 1975 a total of 21 species nested found in earlier studies in eastern deciduous 1 on on one or both of the study areas (Table 3). forests (MacArthur and MacArthur 1961, MacArthur A New breeders included Mourning Doves, Cassin's et al. 1962). Austin (1970), working in "desert Kingbirds, Virginia's Warblers, Western Tanagers, riparian" habitats in Nevada, plotted his data Lesser Goldfinches, and Rufous-sided Towhees. against MacArthur's (1964) regression line for In previous years all of these had either BSD vs. FHD and found similar results. Carothers ;t appeared as transients or nested within the et al. (1974) found that in "desert riparian" .ash, canyon but off the study areas. habitats immediately adjacent to areas of Lests. In 1975 for the first time there was in relatively higher productivity but low avian increase in density (44.4 percent) on camp­ densities, the BSD and FHD correlation no tobin ground. In 1973 and 1974 breeding activity longer held. Yet, Carothers found that in !king was well underway on the campground prior to "desert riparian" habitats immediately adjacent the opening date. In 1975, however, colder to areas of relatively the same productivity temperatures, higher winds, and increased and having a compliment avian community, the 'ckers precipitation postponed breeding. In many BSD and FHD relationships did come close to ,ged species a positive correlation between tem­ MacArthur's regression line. ed perature and the rate of testicular development The BSD's and FHD's obtained in Oak Creek to or egg production has been found (Farner and are summarized in Table 4 and graphed in Figure g Wilson 1957). In 1975 the average temperature 1. Although the points do cluster around two weeks prior to opening was 57.7°F and for MacArthur's line, there is enough deviation to the same period in 1973 and 1974 was 63.7°F suggest other forces at work besides foliage sed and 62.5°F, respectively. Perusal of the height diversity. was 1975 censuses indicates almost no breeding As in Carother's study plots, this is a activity (i.e., singing, displaying, nest riparian system and MacArthur's line fails to building) before 16 May. take into account the added dimension of ler, It is interesting to note that once breed­ permanent water. Also, human disturbance is nt ing did commence, the maximum density reached not considered. An additional downfall of was still less than the maximum measured in FHD is that there has been no stipulation by past 1973 and 1974. investigators when to measure FHD. As we see On control there was in increase of 149.8 here, BSD and FHD vary through time (or sampling percent. This phenomenal climb is also no error) • BS 0 was measured from the first signs doubt related to the later breeding period. of breeding to the opening date and then from on Before the opening day, weather conditions that date to the end of breeding activity. On were too severe for breeding to commence. In the other hand, FHD was measured once before addition, several species that would normally and once after. It is possible that a day nest elsewhere (e.g., Rufous-sided Towhees could be found during the vegetative growing :ic usually nest in the chaparrel found on the season when the FHD would be such that BSD canyon walls and Lesser Goldfinches usually nest for the entire period matched MacArthur's line. above the rim) were found on the control. This leads me to question the value of FHD as Perhaps environmental conditions were relatively .a predictor of BSD except in those specific less severe within the canyon than elsewhere cases studied by MacArthur and the need for

179 Table 4.--Bird species diversity and habitat diversity. specific are to bE CAMPGROUND CONTROL Exan Spec~es Foliage Height Bird Species Foliage Height Bird 1974 thel Diversity Diversity Diversity Diversity campgrour (BSD) (FHD) (BSD) (FHD) occurred After Before After Before After Before After Before again inc 1973 2.19 1.95 .98 1.00 2.08 2.34 1.04 1.08 The reasc breeding 1974 2.62 2.42 .96 .98 2.08 2.43 1.06 LOS

1975 2.19 2.34 .97 1.01 1.83 2.71 .99 .97 In c and orgaJ is impor1 (SCB) anc (Salt 19~ total we: communit~ metaboli! BSD Table 5.--Individual bird weightsl and the diff, 3 individual existence energy. in body \ energy (c Weight in Existence communit: Species Grams Energy limitatic

1973.--Tl after 29 This is J per indio 31. 7 g. bodied b 2 faced Wa Table 5 Steller' moved of The however, remained The decr to a gen The the aver the cont Before After each oth control Cl 1973 Comp ground. Control 0 • • The Camp 6. the same 1974 Control <> • BE showe • We 19715 Comp V Control 0 • campers • weight p the . same the aver dissimil I BE was n FHD and collections of the Museum of Northern Arizona. 1974.--1 Figure l.--Bird species diversity (SD) slightly as a function of foliage height diver­ increase sity (FHD) before and after occupation in the a of the campground by campers. Regress­ (46.8 g ion line from MacArthur et al. 1966. smaller­

180 .c .

. ...''~....•','.'.•.. '.....,:.. :~; -"<") .~ .','

;.,~ specific time limitations when these parameters The campground SCB decreased greatly, as in are to be measured. 1973, and the average weight per individual Examining BSD, we see that in 1973 and remained fairly constant. I 1974 there was a decrease in diversity on the In 1974, larger-bodied birds occupied the campground after it was opened. An increase control initially, but this changed sharply after occurred on control. In 1975, contol's diversity the opening of the campground. again increased but so did the campground's. EE values on control changed upwardly 27.8 The reason for this is, onr.e again, the late percent, whereas the campground's was decreased breeding season in 1975 (see previous section). by 22.3 percent. Once again, the opening appears to be Bioenergetics detrimental to the birds in the campground.

In order to better understand the energetics 1975.--The overall trends remained the same in and organization of these avian communities, it 1975. Smaller-bodied birds made up a majority of is important to look at standing crop biomess the population on control. The increase in SCB (SCB) and existence energy (EE) of the birds and EE on the campground, of course, was related (Salt 1957, Karr 1968). The former is the to the density increase that year. total weight (in grams) of the entire avian community. In order to consider community metabolism, a conversion is made that reflects SUMMARY AND MANAGEMENT ALTERNATIVES the difference in metabolism due to differences in body weight. This is expressed as existence After three breeding seasons, several energy (or Kcal) consumed by the total avian phenomena were discerned: 1) although bird .ence community (see Carothers et al. 1974 for densities on the campground and control are rgy limitations of this measure) . similar before the campground opening date, the average weights of an individual bird is greater 1973.--The SCB of control decreased slightly on the campground (x = 48.5 g) than on control after 29 May, although density increased. (x = 38.2 g) and 2), population density and species 79 This is possible because the average weight diversity (H') decrease when the campground is 94 per individual bird decreased from 40.0 g to occupied by people. 90 31.7 g. Table 3 shows that several small­ In other words, the presence of the camp­ 15 bodied birds, Western Wood Pewees and Red­ ground produced a significant shift in the avian 00 faced Warblers, did move onto the area; see community to heavier bodied birds relative to the 47 Table 5 for weights. Two larger species, natural control area. This is probably a response 71 Steller's Jays and Black-headed Grosbeaks, to the "opening" of the habitat during camp­ 06 moved off the area. ground construction. Inhabitation of the camp­ 75 The campground had a drastic SeE decrease; ground by people causes a direct reduction in 14 however, the average weight per individual the numbers and kinds of breeding birds. 42 remained essentially constant (52.7 g to 54.2 g). Those in managerial posistions might 82 The decrease can therefore only be attributed consider the following suggestions: 08 to a general loss of birds of all sizes. 1. Locations for new campgrounds should 67 The initial and final differences between be carefully scrutinized in terms of usage by 53 the average weight per individual values on wildlife. In this specific case, riparian habi­ 48 the control and campground show that relative to tats are very important to birds and of such a 97 each other light-weight birds inhabited the limited extent in the Southwest that further 45 control and heavier birds inhabited the camp­ destruction of habitat needs to be discouraged. 67 ground. 2. Existing campgrounds should be period­ 67 The existence energy values were initially ically closed to allow regeneration of vegetation 54 the same but after the opening the campground and reduce stress on resident wildlife. This is 16 EE showed a decrease of 37.3 percent. being.done in Oak Creek Canyon, but.much too 33 We see then that before the intrusion of often the reason behind the closure is financial 58 campers the two areas differed in the average rather than ecological. 53 weight per individual by 12.7 g but the EE was 3. Opening the campground before or after l3 the same. Following campground occupation, the height of the breeding season may be better 50 the average weight per bird became more for the avifauna. If people are already present dissimilar (22.5 g), and the total community when birds arrive to nest, the birds may be able EE was nearly halved on the campground. to find suitable habitat elsewhere instead of 972, "wasting energy" by attempting to breed and then 1974.--The SCE of control, once again, decreased being disrupted during incubation•. Of course, slightly after the opening, although density not opening the campground until after breeding increased. Again, this was due to a decrease season would be ideal for the birds but probably in the average weight per individual bird very impractical for the campers. (46.8 g to 23.4 g) caused by an influx of 4. Hab).tat manipulation should be carefully smaller-bodied species (Table 3 and 4). controlled. This includes breaking off branches

181 for firewood, trenching for tents, running of in selected Panama and Illinois habitats. noisy equipment, and even clearing of snags, Ecol. Monogr. 41:207-233. slash, and brush by USFS crews. Kendeigh, S.C. 1944. Measurement of bird 5. Educational programs may be the only populations. Ecol. Monogr. 14:67-106. effectual solution of human recreation and Lack, D. 1950. The breeding seasons of wildlife problems. Government agencies have European birds. Ibis 92:288-316. had very good results in some public educational Lloyd, M., J.H. Zar, and J.R. Karr. 1968. canpaigns (e.g., Smokey the Bear). There is On the calculation of information-theoretical no reason the general public could not be measures of diversity. Amer. MidI. Nat. 79: exposed to broad ecological concepts such as 257-272. camping with less impact. MacArthur, R.H. 1964. Environmental factors Finally, it is hoped that studies of this affecting birds species diversity. Amer. type and education of the public will lead to a Nat. 98:387-397. happy medium between preserving native wildlife MacArthur, R.H. 1965. Patterns of species and also allowing human enjoyment of an area. diversity. BioI. Rev. 40:510-533. MacArthur, R.H. and J.W. MacArthur. 1961. On bird species diversity. Ecology 42:594­ LITERATURE CITED 598. MacArthur, R.H., J.W. MacArthur, and J. Preer. American Ornithologist Union. 1957. Checklist 1962. On bird species diversity, II of North American Birds, Fifth Ed. Lord Prediction of birds census from habitat meas­ Baltimore Press, Baltimore. urements. Amer. Nat. 96:167-174. American Ornithologists Union, 32nd Supplement. MacArthur, R.H., H. Recher, and M. Cody. 1966. 1973. Auk 90:411-419. On the relation between habitat selection and Austin, G.T. 1970. Breeding birds of desert species diversity. Amer. Nat. 100:319-332. riparian habitat in southern Nevada. Marshall, J. and R.P. Balda. 1972. The breed­ Condor 72:431-436. ing biology of the Painted Redstart. Unpubl. Bent, A.C. 1968. Life histories of North manuscript, Northern Arizona Univ., Flagstaff. American birds. U.S. Govt. Print. Off., Morisita, M. 1959. Measuring of interspecific Washington, D.C. association and similarity between communities. Carothers, S.W. and R.R. Johnson. 1975. Water Mem. Fac. Sci.Kyushu Univ., Ser. E., 3:65-80. management practices and their effects on Odum, E.P. 1950. Bird populations of the nongame birds in range habitats. Proc. of highlands (North Carolina) plateau in relation the Sym. on Management of Forest and Range to plant succession and avian invasion. Po Habitat for Nongame Birds. U.S.D.A. Forest Ecology 31:587-605. they al Service General Tech. Report WO-l, July. Pielou, E.C. 1966a. Species-diversity and or less Carothers, S.W., J.R. Haldeman, and R.P. Balda pattern diversity in the study of ecological levels 1973. Breeding birds of the San Francisco succession. J. Theoret. BioI. 10:370-383. tions s Mountain Area and the White Mountains, Peilou, E.C. 1966b. Shannon's formulas as a respect Arizona. Museum of Northern Arizona Tech. measure of species diversity; its use and Cyclic Ser. Bull. No. 12. misuse. Amer. Nat. 100:463-465. four fu Carothers, S.W., R.R. Johnson, and S.W. Salt, G.W. 1957. An analysis of avifaunas in decline Aitchison. 1974. Population structure and the Teton Mountains and Jackson Hole, Wyoming. have in social organization of southwestern riparian Condor 50:373-393. microti birds. Amer. Zool. 14:97-108. Shannon, C.E. and W. Weaver. 1963. The mathe­ three-t Cottom, G. and J.T. Curtis. 1956. Use of matical theory of communication. Univ. of as well distance measurements in phytological Ill. Press, Urbana, Ill. Elton 1 sampling. Ecology 37:451-460. Von Haartman, L. 1971. Population dynamics. 1939, \oJ Farner, D.S. and A.C. Wilson. 1957. A quanti­ In D.S. Farner and J.R. King, eds., studied tative examination of testicular growth in the Avian Biology, Vol. 1. Academic Press, diversi Whi te-crowned Sparrow. BioI.Bull. 113.: 254-267. N.Y. and London. Immelmann, K. 1971. Ecological aspects of Whittaker, R.H. 1970. Communities and eco­ De periodic reproduction. In D.S. Farner and systems. MacMillan Co., N.Y. conside J.R. King, eds., Avian Biology, Vol. 1. Williams, A.B. 1936. The composition and of cyc! Academic Press, N.Y. and London. dynamics of a beech-maple climax community. Karr, J.R. 1968. Habitat and avian diversity Ecol. Monogr. 6:317-408. on strip-mined land in east-central Illinois. Winkworth, R.E. and D.W. Goodall. 1962. A 1/ Condor 70:348-357. crosswire sighting tube for point quadrat Impor Karr, J.R. 1971. Structure of avian communities analysis. Ecology 43:342. Ripar 2/ Associa ProfesE Dept. 2 StudieE

182