Researches on

~_F~cULATIONO~ginal
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

The studies of island plant and animal communities have
played an important role in the development of evolu-
tionary and ecological thought (e.g., MacArthur and
Wilson 1963, 1967; MacArthur 1972; Simberloff 1974;
Cody and Diamond 1975; Gorman 1979; Diamond and
May 1981; Pianka 1988). Islands have definite boundaries
and come in many different sizes, heights, and degrees of
isolation, and therefore constitute some of the finest of
natural ecological experiments (Diamond and May 1981;
Pianka 1988).
Some mainland habitats which are effectively isolated
from each other (e.g., a patch of forest, isolated lakes,
alpine mountaintops, recent fire burns, and gaps in a
1) Present address: Laboratory of Animal Ecology, Yamanashi Institute of
Environmental Sciences, Ken-marubi, Fujiyoshida, Yamanashi 403, Japan
E-mail: mkita@yies.pref.yamanashi.jp
(Wilson and Bossert 1971; MacArthur 1972; Diamond and
May 1981; Giller 1984; Pianka 1988; Begon et al. 1990).
As Wilson (1992) noted, habitat islands are subject to
the same ecological and evolutionary processes as real
islands. Up to the present, some progress has been made
in applying the equilibrium theory of island biogeography
(MacArthur and Wilson 1967) to such habitat islands (e.g.,
Vuilleumier 1970; Brown 1971; Gorman 1979). However,
the mainland habitats usually form very complex
mosaics. As a result, diversity patterns on the habitat
islands cannot so easily be studied and our understanding
is much less advanced than those on real islands (Cody
1975; Gorman 1979; Giller 1984).
On the other hand, the usefulness of the island
biogeography theory in conservation biology has been
claimed (e.g., SpeUerberg 1991; Primack 1993; Hunter
1996). However, little attempt has yet been made to
evaluate the simultaneous applicability of the generalist/
 24 KITABARA&FuJn
specialist concept and the island biogeography theory in
order to analyze habitat island ecological communities.
In particular, we do not know any attempt of applying
the generatist/specialist concept directly to the actual
component species of habitat island communities.
In our previous paper (Kitahara and Fujii 1994), we
treated the newly designed parks in Tsukuba City, central
Japan, mainly as the most human-disturbed site of all
the study areas. In addition to this high degree of human
disturbance, these parks were also newly established after
intensive ground preparation, and most of the vegetation
was transplanted from other areas. Thus, their environ-
mental structures are very different from the surrounding
habitats. Most animal species present there seem to have
immigrated from adjacent habitats after the birth of the
parks. Thus, we can think of these parks as "habitat
islands" with newly established faunas, surrounded by
habitats with relatively old faunas. These parks can con-
stitute one of the finest of natural ecological experiments
on the mainland for clarifying the patterns and processes
of faunal immigration into the newly established habitat
islands.
Schaefer (1982) demonstrated that, in studying the struc-
ture and function of green urban ecosystems, it is impor-
tant to compare these ecosystems with semi-naturai ones
in the surroundings of the urban area (also see Frankie
and Ehler 1978). Thus, in the present study, to analyze the
patterns of the habitat island communities, we used data
from not only the butterfly communities in these parks,
but also from two other butterfly communities in more
natural habitats in the surrounding areas, and compared
them in the light of the theory of island biogeography and
the concept of generalist/specialist. Butterflies are the most
suitable organisms for the study of the structure and
dynamics of biological populations and communities
(Ehrlich 1992). The adults can be identified easily in the
field and their life-history information (e.g., hostplants,
voltinism) are already well known in Japan (e.g., Fukuda
et al. 1972).
In our previous paper (Kitahara and Fujii 1994), we
analyzed butterfly community structures based on the
classification of their component species into generalist,
specialist, and other species. In this paper, we analyze
butterfly community structures in more detail based on the
more exact classification of the component species as detail-
ed in the section "Definitions". Our goals in this paper
were (1) to search for butterfly community patterns in
newly designed parks by comparing them with those in two
other areas with relatively old faunas, (2) to understand
the habitat island community patterns detected in the light
of the generalist/specialist concept and the island
biogeography theory, and (3) to evaluate the synergetic
effectiveness of both the concept and the theory for the
analysis of habitat island community patterns.
Study areas and methods
In this study we used the data collected in our previous
study, and the details of the census areas and sections
within were described elsewhere (Kitahara and Fujii
1994). Only short descriptions follow. We established
three census routes (in areas A, B, and C) in and around
Tsukuba City, Ibaraki Pref., central Japan. Route A was
established in area A located in secondary natural forest
on the gentle south slope of Mt. Tsukuba (alt. 875.9 m)
at an altitude of 150-200m (36~ 140~ It was
860 m in length and divided into 2 sections (A-1 and A-2).
Route B was established in area B located in cultivated
lands and villages from 50 to 100 m in altitude at the foot
of Mt. Tsukuba. It was 1280 m in length and divided
into 4 sections (B-l, B-2, B-3-1 and B-3-2).
Route C, which was 2610 m in length, was established
in area C located in newly designed city parks with
transplanted trees and a connecting pedestrian road at
an altitude of 25 m in Tsukuba City (36~ 140~
Area C was occupied mainly by patchy forests, lawns,
paved roads, and man-made ponds. Most forest trees
were transplanted when the parks were designed. These
forests were mixtures of various tree species belonging
to deciduous and evergreen broad-leaved trees and con-
iferous trees. Most of the forest floors were covered by
lawns and grasses, and the rest was bare ground. Human
activities (mowing, pruning, and insecticide spraying) by
park management were frequently conducted aperiodical-
ly. Thus, succession was almost prevented by human ac-
tivities, and the environmental structure was very different
from those of the surrounding areas. When we studied
the butterfly community of area C in the year 1980, several
years had passed since the birth of the parks.
Route C was divided into three census sections, C-1
(510m), C-2 (900m) and C-3 (1200m) based on the
different vegetation and environmental structures within
and around each section. C-1 was located in Akatsuka
Park in which some patches of secondary forest remained.
The park was surrounded mainly by cultivated fields and
residential area. C-2 was located along a pedestrian road
about 20m wide, connecting C-1 and C-3. It was situated
next to remaining forests dominated mainly by deciduous
broad-leaved trees and Pinus densiflora. C-3 was located
in Doho Park in which most trees had been transplanted.
There was no forest in the surrounding area.
We censused butterfly communities twice a month dur-
ing the adult flight season (from March to November,
1980) by line transect method (Pollard 1977; Thomas 1983;
Gall 1985; Pollard and Yates 1993). Details of the census
method are described elsewhere (Kitahara and Fujii 1994).
We analyzed the butterfly community structure using
the following ecological indices; monthly mean popula-