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Researches on

Res. Popul. Ecol. 39(1), 1997, pp. 23-35.
9 1997by the Societyof Population Ecology OLOGY pap ]

An Island Biogeographicai Approach to the Analysis of Butterfly

Community Patterns in Newly Designed Parks

Masahiko KITAHARAl) and Koichi FUJII*

Graduate School of Environmental Sciences, The University of Tsukuba, Tsukuba, Ibaraki 305, Japan
*Institute of Biological Sciences, The University of Tsukuba, Tsukuba, Ibaraki 305, Japan

Abstract. We analyzed the butterfly communities in the newly designed city parks (area C), "newly
opened habitat islands", of Tsukuba City, central Japan. The area constituted a natural ecological
experiment on the mainland for clarifying the pattern and process of faunal immigration. We com-
pared butterfly communities in area C with those in two other areas in the light of the theory of island
biogeography and the concept of generalist/specialist. Our results showed the following: (1) Fewer
species were found in area C than in other areas, due largely to the absence of many specialist types,
restricted and habitat specialists, and/or low density species in the area. Generalist types, widespread
and habitat generalists, and/or high density species predominated in area C. (2) The difference in the
species numbers among the three sections within area C could be explained by the habitat structure
in and around the respective sections. (3) The densities of many species were low in area C, pro-
bably due to its man-modified habitat structure. In particular, several species occurred at extremely
low densities in area C, but at high densities in other areas. (4) The internal structure of the habitat
island butterfly community in area C was almost perfectly consistent with that of "quasi-equilibrium"
communities that appear during the colonization of an island. Our results demonstrate that the
synergetic application of the generalist/specialist concept and the island biogeography theory is
effective for the understanding of the patterns and structures of habitat island communities.

Key words: island biogeography theory, island patterns, generalist/specialist, habitat island, newly
designed city parks, butterfly communities.

Introduction forest canopy) can also be regarded as habitat islands

(Wilson and Bossert 1971; MacArthur 1972; Diamond and
The studies of island plant and animal communities have May 1981; Giller 1984; Pianka 1988; Begon et al. 1990).
played an important role in the development of evolu- As Wilson (1992) noted, habitat islands are subject to
tionary and ecological thought (e.g., MacArthur and the same ecological and evolutionary processes as real
Wilson 1963, 1967; MacArthur 1972; Simberloff 1974; islands. Up to the present, some progress has been made
Cody and Diamond 1975; Gorman 1979; Diamond and in applying the equilibrium theory of island biogeography
May 1981; Pianka 1988). Islands have definite boundaries (MacArthur and Wilson 1967) to such habitat islands (e.g.,
and come in many different sizes, heights, and degrees of Vuilleumier 1970; Brown 1971; Gorman 1979). However,
isolation, and therefore constitute some of the finest of the mainland habitats usually form very complex
natural ecological experiments (Diamond and May 1981; mosaics. As a result, diversity patterns on the habitat
Pianka 1988). islands cannot so easily be studied and our understanding
Some mainland habitats which are effectively isolated is much less advanced than those on real islands (Cody
from each other (e.g., a patch of forest, isolated lakes, 1975; Gorman 1979; Giller 1984).
alpine mountaintops, recent fire burns, and gaps in a On the other hand, the usefulness of the island
biogeography theory in conservation biology has been
1) Present address: Laboratory of Animal Ecology, Yamanashi Institute of claimed (e.g., SpeUerberg 1991; Primack 1993; Hunter
Environmental Sciences, Ken-marubi, Fujiyoshida, Yamanashi 403, Japan 1996). However, little attempt has yet been made to
E-mail: evaluate the simultaneous applicability of the generalist/

specialist concept and the island biogeography theory in

order to analyze habitat island ecological communities. Study areas and methods
In particular, we do not know any attempt of applying
the generatist/specialist concept directly to the actual In this study we used the data collected in our previous
component species of habitat island communities. study, and the details of the census areas and sections
In our previous paper (Kitahara and Fujii 1994), we within were described elsewhere (Kitahara and Fujii
treated the newly designed parks in Tsukuba City, central 1994). Only short descriptions follow. We established
Japan, mainly as the most human-disturbed site of all three census routes (in areas A, B, and C) in and around
the study areas. In addition to this high degree of human Tsukuba City, Ibaraki Pref., central Japan. Route A was
disturbance, these parks were also newly established after established in area A located in secondary natural forest
intensive ground preparation, and most of the vegetation on the gentle south slope of Mt. Tsukuba (alt. 875.9 m)
was transplanted from other areas. Thus, their environ- at an altitude of 150-200m (36~ 140~ It was
mental structures are very different from the surrounding 860 m in length and divided into 2 sections (A-1 and A-2).
habitats. Most animal species present there seem to have Route B was established in area B located in cultivated
immigrated from adjacent habitats after the birth of the lands and villages from 50 to 100 m in altitude at the foot
parks. Thus, we can think of these parks as "habitat of Mt. Tsukuba. It was 1280 m in length and divided
islands" with newly established faunas, surrounded by into 4 sections (B-l, B-2, B-3-1 and B-3-2).
habitats with relatively old faunas. These parks can con- Route C, which was 2610 m in length, was established
stitute one of the finest of natural ecological experiments in area C located in newly designed city parks with
on the mainland for clarifying the patterns and processes transplanted trees and a connecting pedestrian road at
of faunal immigration into the newly established habitat an altitude of 25 m in Tsukuba City (36~ 140~
islands. Area C was occupied mainly by patchy forests, lawns,
Schaefer (1982) demonstrated that, in studying the struc- paved roads, and man-made ponds. Most forest trees
ture and function of green urban ecosystems, it is impor- were transplanted when the parks were designed. These
tant to compare these ecosystems with semi-naturai ones forests were mixtures of various tree species belonging
in the surroundings of the urban area (also see Frankie to deciduous and evergreen broad-leaved trees and con-
and Ehler 1978). Thus, in the present study, to analyze the iferous trees. Most of the forest floors were covered by
patterns of the habitat island communities, we used data lawns and grasses, and the rest was bare ground. Human
from not only the butterfly communities in these parks, activities (mowing, pruning, and insecticide spraying) by
but also from two other butterfly communities in more park management were frequently conducted aperiodical-
natural habitats in the surrounding areas, and compared ly. Thus, succession was almost prevented by human ac-
them in the light of the theory of island biogeography and tivities, and the environmental structure was very different
the concept of generalist/specialist. Butterflies are the most from those of the surrounding areas. When we studied
suitable organisms for the study of the structure and the butterfly community of area C in the year 1980, several
dynamics of biological populations and communities years had passed since the birth of the parks.
(Ehrlich 1992). The adults can be identified easily in the Route C was divided into three census sections, C-1
field and their life-history information (e.g., hostplants, (510m), C-2 (900m) and C-3 (1200m) based on the
voltinism) are already well known in Japan (e.g., Fukuda different vegetation and environmental structures within
et al. 1972). and around each section. C-1 was located in Akatsuka
In our previous paper (Kitahara and Fujii 1994), we Park in which some patches of secondary forest remained.
analyzed butterfly community structures based on the The park was surrounded mainly by cultivated fields and
classification of their component species into generalist, residential area. C-2 was located along a pedestrian road
specialist, and other species. In this paper, we analyze about 20m wide, connecting C-1 and C-3. It was situated
butterfly community structures in more detail based on the next to remaining forests dominated mainly by deciduous
more exact classification of the component species as detail- broad-leaved trees and Pinus densiflora. C-3 was located
ed in the section "Definitions". Our goals in this paper in Doho Park in which most trees had been transplanted.
were (1) to search for butterfly community patterns in There was no forest in the surrounding area.
newly designed parks by comparing them with those in two We censused butterfly communities twice a month dur-
other areas with relatively old faunas, (2) to understand ing the adult flight season (from March to November,
the habitat island community patterns detected in the light 1980) by line transect method (Pollard 1977; Thomas 1983;
of the generalist/specialist concept and the island Gall 1985; Pollard and Yates 1993). Details of the census
biogeography theory, and (3) to evaluate the synergetic method are described elsewhere (Kitahara and Fujii 1994).
effectiveness of both the concept and the theory for the We analyzed the butterfly community structure using
analysis of habitat island community patterns. the following ecological indices; monthly mean popula-