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The Occupation of Small Islands by Passerine Birds

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The Occupation of Small Islands by Passerine Birds THE OCCUPATION OF SMALL ISLANDS BY PASSERINE BIRDS DOUGLASS H. MORSE Comparisons of island and mainland faunas STUDY AREA AND METHODS have provided great insight into how com- Most of the 12 islands censused are partially or totally munities are structured (see MacArthur 1972). forested by white spruces (Picea g&x), with small In particular, islands permit natural compari- but varying amounts of balsam fir ( Abies balsumeu), sons of how animals under varying degrees of red oak (Quercus rubru), and white birch (Betulu fauna1 impoverishment exploit their resources. pupytifem). Two of the islands, however, contain To date, most islands studied from this point substantial amounts of mountain maple (Acer spicu- turn), striped maple (A. pensyltxznicum), and yellow of view are sufficiently large to support size- birch (Bet&z Idea). In addition, some have sizeable able populations of many or all of the species open areas sparsely covered by various grasses, rasp- that occupy them. Further, island popula- berry ( Rubus sp. ), bayberry ( Myricu pennsyluunicu), tions typically studied are either of unknown or goldenrod (Solidago spp. ). Since my primary concern lay with the forest-dwelling birds, I have genetic relationship to their mainland counter- included typically open-country species only when parts, or they are easily distinguished from they used the forested areas of these islands. These them, often being considered separate sub- islands are described in detail elsewhere (Morse species or even species. 1971a, 1973a). No visible successional change oc- In this paper I report upon the patterns curred during the period of the study. Records of breeding and visiting species were made by which passerine birds occupy extremely during intensive studies upon the warblers (Paruli- small forested islands and adjacent large main- dae) of the islands (Morse 1971a, 1973a). These land habitats in and around Muscongus Bay, studies provided information on the fledging suc- Maine, thus expanding upon earlier observa- cess of the warblers during the 1967-1970 seasons. During that period, several of these birds were either tions (Morse 1971a). I attempt to answer individually marked or identified by their songs. Ob- the following questions: From which main- servations of other species were designed to give an land habitats do the inhabitants of these accurate measure of presence or absence, even though islands come? Which characteristics of the abundances are in some cases estimates where many pairs were present (e.g., Song Sparrow, Melospizu islands best predict the number of species melodiu). After 1970, I made at least six censuses that will occupy them? What factors best each year between 10 June (after the last spring account for the presence or absence of any migrants) and 15 July (after the latest establishment of breeding individuals) (Morse 1971a, 1973a). given species on the islands? How stable are These censuses ranged from 15-20 min per visit on the island assemblages in time? How does the smallest islands (under 0.2 ha) to an hour or the total density of birds and the densities of more on the largest islands. Minimum criteria for colonization were satisfied if a species was seen on individual species compare with the densities half or more of the censuses and if both a male and on the mainland? Then, where appropriate, I a female were detected (i.e., settlement of a mini- relate my findings to island biogeographic mum breeding unit). Year-by-year lists of breeding species are available on request. theory (MacArthur and Wilson 1963, 1967, Several aquatic species (herons, ducks, gulls) Diamond 1975). nested on these islands but foraged elsewhere. Since Small islands of this sort are excellent sys- I have no evidence that any of these birds competed for nest sites or food resources with fully resident tems for answering such questions. By virtue species or that they preyed upon these residents of their small size (0.14.0 ha), it is often pos- regularly, I will not consider them further. sible to determine the minimal conditions To determine food supplies available to the foliage- inhabiting species, I measured the biomass of foliage- (space, etc.) required for a species to estab- inhabiting insects on these islands durinn the 1968- lish itself. Potentially key variables (physical 1970 seasons, using the same technique as in my characteristics of the forest, food supply, etc.) other studies (Morse 1973a, 1976). Weekly or bi- weekly I collected 20 live branches at random (ap- can be monitored readily. A variety of species proximately 0.2 m” and 6.0 m3) with their associated compositions provides the opportunity to eval- arthropods on the islands and mainland, attempting uate the effects of certain species upon others. to lose as few arthropods as possible. The branches were placed in plastic bags, fumigated, and then Lastly, the birds of any given species inhabit- beaten over a screen of hardware cloth. Data on ing these islands almost certainly belong to arthropods so obtained were pooled for each two- the same gene pool as those on the adjacent week period, permitting a direct comparison of island mainland (Morse 1971a), so one can eliminate and mainland insect stocks. Although this technique extracts over 99% of the arthropods on the branches evolutionary change as the basis for such dif- at the time that they are placed in the bags (Morse ferences. 1973a), it underestimates the abundance of strongly- K3991 The Condor 79:399-412, 1977 400 DOUGLASS H. MORSE flying insects. However, since most of the foliage- TABLE 1. Contributions of spruce and deciduous- foraging birds were primarily gleaners, this potential edge components to island populations of birds dur- problem should not bias the results seriously. ing 1967-1975 period. Total pairs RESULTS Species-year@ of individuals THE SPECIES POOL Deciduous- DeCi~iUS- Island Spruce edge Spruce One can construct a mainland species pool Strawberry 0 8 0 8 (Appendix I) with considerable accuracy from Byers’ several sources of information. Mainland is Crotch I: 118 I: 1116 here defined as that land physically connected Crow 15 11 15 38 25 to the continental shoreline, plus islands of Jim’s 12 25 42 Ram 18 12 18 26 These large islands are over 25 ha in size. Crane 20 31 20 93 partly or totally covered with spruces. Earlier Indian 28 12 28 27 (Morse 1966) I compiled lists of species nest- Thief 41 20 45 92 ing in spruce forests, deciduous forests, hur- Franklin 0 31 0 174 Haddock 15 26 116 ricane-damaged areas, and edge situations on 15 Wreck 22 29 28 163 Hog Island, a large island (132 ha) separated at points from the mainland by not over 200 TOTALS 191 223 201 806 m. This compilation was based upon personal PERCENTAGE 46.1 53.9 20.0 80.0 field work and the breeding-bird censuses of n One species-year represents the occupation of an island Cadbury and Cruickshank from 1937 to 1958. for one year by <,neor nxxe breeding pairs of a species. Subsequent intensive field work has added but two more species to these lists. Thus, the forest group. This characteristic was espe- updated lists presented as Appendix I ap- cially striking in the case of the Song Sparrow, pear to be largely complete. which made up 44.8% of the total number Another group of species breeds on the of birds. Not suprisingly, the Song Sparrow mainland within 2 km of the shoreline, but is an extremely successful colonizer in other does not occur on the Hog Island lists (Ap- areas as well (Beer et al. 1956, Yeaton and pendix I). I compiled this list from personal Cody 1974). The spruce-forest component observations made since 1957 and from Cruick- was much more evenly distributed than that shank (no date). Over half of these species of the deciduous-edge component, because occur in open-country or brushy areas, and the former was comprised of species that most of the remaining species typically occur seldom attained a density greater than one in deciduous forests. pair per island (Fig. 1). CONTRIBUTIONS OF THE DIFFERENT FOOD SUPPLY COMPONENTS The foliage sampling for arthropods indicated Seventeen species of small passerine birds a rather constant average biomass of insects bred on forested parts of the small islands through the season (Fig. 2). During the first during the 1967-1975 period. They came en- part of the season, the average biomass was tirely from the first two components of the similar to that of a white spruce forest on the mainland species pool: (1) the spruce forest mainland, while later it exceeded that of the and (2) the adjacent deciduous, disturbed, mainIand. Therefore, the abundance of food and edge areas (henceforth referred to as on the foliage does not appear to account for deciduous-edge areas). These two compo- the absence of many species from these is- nents made similar contributions to the island lands. Other possible food sources (from species pool (nine from the spruce forest ground, litter, etc.) were not systematically group, eight from the deciduous-edge group). measured. The two components also contributed rather similar numbers of species-years (defined as NUMBER OF SPECIES PRESENT the occupation of an island by a species dur- ing one breeding season) on the islands (Ta- Using stepwise multiple regression, I tested ble 1). However, the relative abundance of six potentially important factors (the inde- the deciduous-edge species was much greater pendent variables of Table 2) for their rela- than that of the coniferous species (Table 1)) tionship to the number of species that oc- because several members of the deciduous- cupied an island. I used three different edge group established considerably more measures of species numbers-cumulative list dense populations than did those of the spruce of species over 1967-1975 period, maximum BIRDS ON SMALL ISLANDS 401 TABLE 2.
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