Biodiversity and Conservation 11: 1919–1937, 2002.

 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Species diversity of longicorn beetles in humid warm-

temperate forests: the impact of forest management
practices on old-growth forest species in southwestern
Japan

KAORU MAETO 1,3, *, SHIGEHO SATO 1 and HIROAKI MIYATA2

1
Shikoku Research Center, Forestry and Forest Products Research Institute, Asakura-Nishimachi, Kochi
780 -8077, Japan; 2 Kochi Prefectural Forest Technology Center, Ouhira, Tosayamada 782 -0078,
Japan; 3 Current address: Entomological Laboratory, Faculty of Agriculture, Kobe University, Nada-ku,
Kobe 657 -8501, Japan; * Author for correspondence (e-mail: maeto@ kobe-u.ac.jp; fax: 181 -78 -803 -
5871)

Received 22 June 2001; accepted in revised form 26 November 2001

Key words: Attractant trap, Bio-indicator, Chronosequence, Coarse woody debris, Conifer plantation,
Insect biodiversity, Pidonia, Pollinator, Saproxylic Coleoptera

Abstract. In the humid warm-temperate zone of southwestern Japan, old-growth forests have been
seriously fragmented to small remnants due to traditional agriculture and coppicing as well as recent
rapid plantation with conifers. Assemblages of longicorn beetles (Coleoptera: Disteniidae and
Cerambycidae) were compared among old-growth forests, second-growth forests and conifer plantations
using collision traps baited with chemical attractants. Species richness of longicorn beetles was poorer in
second-growth forests and conifer plantations than in old-growth forests. It was proved by multi-
dimensional scaling (MDS) that the beetle assemblages of old-growth forests were distinct from those of
conifer plantations, while those of second-growth forests were intermediate between them. Further
analysis showed that a number of species, including many Pidonia spp., were specific to or closely
associated with old-growth forests, and the results were largely supported by the indicator value (IndVal)
approach. It is likely that many of such old-growth forest species in the larval and pupal stages require
large broad-leaved trees standing or fallen with thick bark. At the same time, the flower-visiting adults
would play an important role in pollinating various herbaceous and woody plants. Regional forest
management for the conservation of insect biodiversity is also discussed.

Introduction

Southwestern Japan was formerly widely covered by humid warm-temperate

evergreen or deciduous broad-leaved forests often mixed with evergreen conifers,
but such forests have been extensively exploited since ancient times (Shidei 1974).
Almost all lowland forests have been converted to rice paddies, farmland and
residential areas. In hilly and mountain areas, the forests have been severely altered
through shifting cultivation, coppicing for manure and fuel wood production, and
plantation with conifers. After World War II, remaining old-growth forests were
rapidly cleared and usually converted to plantations of Japanese cedar and cypress
(Japan FAO Association 1997). As a result, relatively unchanged forests cover less
than 1% of the potential area of this forest type in Japan (Sasse 1998). NACS-J and
1920

WWF Japan (1996) reported that warm-temperate forests have been seriously
affected by deforestation and plantation with conifers, and the remnants need urgent
conservation. Although arthropod diversity may be deteriorated due to the forest
conversion, only a little is known about the importance of old-growth forests in the
maintenance of insect diversity in Japan (Maeto and Makihara 1999).
For sustainable forest management in consideration of regional biodiversity, it is
necessary not only to know the effects of forest conversion on insect species
richness but also to understand the ecological requirements and functional interac-
tions of the insects that depend on old-growth forests (e.g. Fisher 1998). Quite a few
studies have been conducted on the influence of forest management practices on
arthropod communities or assemblages in temperate and boreal forests, e.g., canopy
arthropods (Schowalter 1989), ground beetles (Niemela¨ et al. 1993; Magura et al.
2000), boreal forest beetles (Martikainen et al. 2000). However, the implications of
forest treatments for longicorn beetles are largely unknown, although they are
principal components of forest ecosystems as herbivores and detritivores of woody
plants, pollinators of herbaceous and woody plants, and prey of insectivorous
animals (Linsley 1959; Ohbayashi et al. 1992; Hanks 1999).
The purpose of this study was to compare the assemblage of longicorn beetles
among old-growth forests without logging records, second-growth forests and
conifer plantations in the humid warm-temperate zone of Japan, in order to identify
the characteristic species of old-growth forests and understand their ecological
requirements. Collision traps baited with chemical attractants (Maeto et al. 1995;
Shibata et al. 1996) were used for a quantitative comparison of beetle assemblages.

Materials and methods

Study sites

The study focused on the Shimanto River Basin of Kochi Prefecture in Shikoku, the
fourth largest island of the Japanese archipelago. The river basin is covered with
lowland and hilly forests up to 1400 m in elevation, principally composed of conifer
plantations (about 60% of total forest area), second-growth forests (about 40%), and
remnants of old-growth forests without historical records of clearance (less than
1%). Old-growth forests are dominated by evergreen conifers (Abies firma, Tsuga
sieboldii, Chamaecyparis obtusa) as well as by diverse evergreen broad-leaved trees
(Quercus spp., Castanopsis cuspidata, Machilis japonica, Cleyera japonica, etc.).
Starting at 800 m a.s.l., evergreen broad-leaved trees are gradually replaced by
deciduous trees (Carpinus spp., Fagus spp., Betula grossa, Acer spp., etc.). Second-
growth forests have been repeatedly cut at intervals of 30–80 years; coppices
dominated by evergreen broad-leaved trees (C. cuspidata, Quercus glauca, etc.) are
developed in lowlands, and mixed forests of red pine (Pinus densiflora) and
deciduous broad-leaved trees (Q. serrata, Carpinus spp., etc.) are widespread in the
hills. Conifer plantations are monocultures of Japanese cedar (Cryptomeria
japonica) or Japanese cypress (C. obtusa); in appropriately managed plantations
 1921

over 30 years old, forest floors are covered with rich species of shrubs after thinning,
and diverse tree species grow along forest edges, paths and streams. Annual mean
temperature and annual precipitation in 1991–2000 at Yusuhara (415 m a.s.l.,
Figure 1) averaged 13.4 8C and 2722 mm, respectively.
Seven old-growth forests (without any records of clearance at least for 120 years),
five second-growth forests (30–70 years old) and two appropriately managed
conifer plantations (30–40 years old) were selected for sampling sites within a 50 3
50 km area of the river basin (338059–338309 N, 1328359–1338059 E; Figure 1). For
each site, woody plant species observed in a plot of about 0.05 ha were recorded,
and the diameter at breast height (DBH) of the trees (DBH .5 cm) with the canopy
hanging over a 20 m randomly placed straight line was measured. Site location,
forest type, approximate forest age, altitude, direction, slope, the number of woody
plant species, and the maximum DBH of conifers and broad-leaved trees are listed
in Table 1.

Sampling and identification

Specimens were collected with yellow or white collision traps, each baited with

Figure 1. Location of the study sites in Kochi Prefecture, Shikoku Island, Japan.
 1922

Table 1. Location, forest type, approximate forest age and stand variables of the study sites.
c
Site Location a Forest type b Age of forest (years) Approx. area (ha) Altitude (m) Direction Slope (8) Number of woody Maximum DBH (cm)
plant species d of trees e

Conifers Broad-leaved trees

A Irazu, H OLD .190 150 1050 N 15 23 99.8 (2) 33.1 (20)

B Tengu, H OLD .120 100 1250 S 40 8f ,5.0 (0) 26.8 (11)
C Yokogai, Y SEC 40–50 .200 800 N 20 24 42.0 (6) 13.5 (15)
D Tarohgawa, Y SEC 30–40 .200 700 W 40 11 12.4 (5) 25.3 (10)
E Kohnoshi, Y CPL 30–35 .200 700 S 25 17 23.0 (11) ,5.0 (0)
F Takatori, Y OLD .170 90 350 N 20 27 58.2 (2) 78.6 (19)
G Kubotani, Y OLD .200 80 700 W 45 24 76.3 (4) 27.9 (5)
H Tsuzuragawa, T SEC 40–50 20 400 W 40 36 43.2 (2) 14.3 (14)
I Ichinomata, T OLD .180 50 500 N 40 22 51.0 (2) 42.4 (12)
J Ichinomata, T CPL 30–35 .200 450 N 35 23 16.9 (9) ,5.0 (0)
K Nakaba, N SEC 20–30 .200 100 N 45 21 ,5.0 (0) 21.7 (30)
L Kuroson, N SEC 70 30 350 S 40 15 37.6 (2) 30.6 (12)
M Kuroson, N OLD .120 250 700 E 35 15 116.6 (1) 34.4 (16)
N Haraikawa, N OLD .140 50 500 N 40 16 ,5.0 (0) 39.4 (17)
a b
H – Higashitsuno-mura; Y – Yusuhara-cho; T – Taishoh-cho; N – Nishitosa-mura. OLD – old-growth forest; SEC – second-growth forest; CPL – conifer plantation.
c d e
Approximately estimated area of the same type, continuous forests. Number of woody plant species observed in a plot of about 0.05 ha. For all trees hanging over a
f
straight line of 20 m; numbers in parentheses indicate the number of trees with DBH .5 cm. Forest floor was poor in species, being covered with dwarf bamboo.
 1923

Figure 2. Collision trap baited with benzyl acetate and ethyl alcohol.

benzyl acetate and ethyl alcohol. The collision traps consisted of a roof, two
collision plates (about 25 3 20 cm) intersecting each other, and a bucket (Figure 2;
Sankei Chemical Co., Ltd.; Maeto et al. 1995; Shibata et al. 1996). Benzyl acetate is
one of the main components of floral fragrance, and it lures various flower-visiting
beetles (Ikeda et al. 1993; Sakakibara et al. 1997b). Ethyl alcohol is known to attract
xylophagous beetles for oviposition (Ikeda et al. 1980). Dispensers of the chemicals,
each about 25 ml, were placed under the roof. Water containing a surface-active
agent and sorbic acid was poured into the bucket to preserve the specimens. The
traps were set at about 1.5 m above the ground.
At each site, two white and two yellow traps were placed alternatively at intervals
of about 50 m in line. Trapping was conducted from early April to mid-September in
1998. Every 2 weeks, chemicals were renewed and materials in the buckets were
taken to the laboratory. Trap sampling was repeated in the same manner at five sites
(A, C–F) in 1999, to confirm the invariability of species assemblages between years.
Longicorn beetles (Coleoptera: Disteniidae and Cerambycidae) were pinned for
identification. They were identified with Ohbayashi et al. (1992).Voucher specimens
will be deposited in the Shikoku Research Center, Forestry and Forest Products
Research Institute, Kochi, Japan.
To evaluate the taxonomic bias in the trap sampling, the number of species
collected in this study was compared with the number of species collected by
general methods in the same region for each family and subfamily (Nakayama et al.
1994). They reported 148 species of longicorn beetles collected by hand, in beating,
1924

Table 2. Number of longicorn beetle species collected in white and yellow collision traps attached with
benzyl acetate and ethanol in 1998, and of those collected with general methods (i.e. by hand, in beating,
in light traps and in collision traps) over 6 years (1987–1993) in the same region.
Family Subfamily Number of species Number of species T/O
collected in the collected with
traps in 1998 a general methods b (O)
WT YT Total (T)
Disteniidae 0 2 2 2 1.00
Cerambycidae Prioninae 2 2 3 4 0.75
Spondylinae 1 1 1 4 0.25
Lepturinae 34 13 34 36 0.94
Necydalinae 0 0 0 1 0.00
Cerambycinae 17 17 19 36 0.53
Lamiinae 16 18 23 65 0.35
Total number of species 70 53 82 148 0.55
a
WT – white traps; YT – yellow traps; both were baited with benzyl acetate and ethanol. b At Mt.
Ohnakayama (Figure 1), Yusuhara-cho, Kochi Prefecture (Nakayama et al. 1994).

in light traps and in various bait traps for 7 years on Mt. Ohnakayama, Kochi
Prefecture, up to about 800 m in elevation (Figure 1). Vegetation was composed of
second-growth forests, plantations of Japanese cypress, plantations of Q. acutissima,
and old-growth remnants.

Data analyses

All the specimens captured in the two white and two yellow traps at each site every
year were pooled together for the following analyses. Two species richness indices,
i.e. the number of species and the Margalef index (Magurran 1988), were compared
among three forest types and among four directions by the Kruskal–Wallis test
based on the samples of 1998. Kendall’s coefficient of rank correlation (Kendall’s
tau, t ) between the species richness indices and the stand variables (altitude, slope,
the number of woody plant species, and the maximum DBH of conifers and
broad-leaved trees) were tested. The Margalef index was calculated as

Margalef index5(S21) / lnN

where S is the number of species and N the number of individuals.

Similarity of assemblages between each pair of samples was computed from
abundance data using the rank correlation coefficient (Kendall’s tau, t ) (Ghent
1963; Huhta 1979). From the matrix of similarity (t ), the samples were plotted in a
two-dimensional space by multidimensional scaling (MDS). First, we performed
two-dimensional MDS for all samples collected in 1998 (14 sites) and 1999 (5
sites). The samples were clustered with the group average method based on the
Euclidean distance between each pair of them in the MDS space. After we
confirmed that the samples of 1998 and 1999 collected at every site were fairly close
to each other in the dendrogram, we performed two-dimensional MDS for the
 1925

samples of 1998 (14 sites) again. For each of the two MDS dimensions, the
differences in site score among forest types and among directions were tested by the
Kruskal–Wallis test. Kendall’s coefficient of rank correlation was tested between
the MDS site scores and the stand variables. For the species collected at three or
more sites, Kendall’s coefficient was tested between the MDS scores and the
number of individuals. All statistical analyses were performed with STATISTICA
(StatSoft Inc. 1998).
We also applied the indicator value method proposed by Dufrene ˆ and Legendre
(1997) to identify the most characteristic species of each forest type for the samples
of 1998. This method combines a species relative abundance with its relative
frequency of occurrence in a particular habitat type to obtain the indicator value for
the habitat type. The species indicator value (IndVal) is the maximum indicator
value over all habitat types (three forest types in our study). Statistical significance
of IndVal was evaluated with a Monte Carlo test (McCune and Mefford 1997).

Results

Taxonomic bias in trap sampling

A total of 25 115 individuals belonging to 82 species of longicorn beetles were

collected in 1998. As compared with the fauna of longicorn beetles in the same
region (Nakayama et al. 1994), our trapping system collected all species of the
family Disteniidae, most of the subfamily Lepturinae, but a rather small portion of
the subfamilies Cerambycinae and Lamiinae (Table 2). The lepturine species were
all captured in the white traps, while the disteniid species and some cerambycine
and lamiine species were captured only in the yellow traps.

Figure 3. Species richness indices of longicorn beetles in relation to the number of individuals collected
in 1998. For forest type symbols, see Figure 1.
1926

Figure 4. Margalef index in relation to the number of woody plant species (a), and to the maximum DBH
of broad-leaved trees (b) (cf. Table 4). For forest type symbols, see Figure 1.

Table 3. Median and range (in parentheses) of the species richness indices and MDS site scores for
longicorn beetle assemblages of three forest types in 1998.
Type of forest H
Old-growth Second-growth Conifer
forest n 5 7 forest n 5 5 plantation
n 52
Species richness
Number of species 27 (15–33) 22 (9–24) 21 (20–22) 2.61
Margalef index 3.79 (3.03–4.27) 2.97 (1.67–3.32) 2.41 (2.03–2.79) 8.14 *
MDS site scores
First dimension 20.60 (21.05 to 20.04) 0.33 (20.24–0.82) 1.45 (1.35–1.55) 10.13 **
Second dimension 0.03 (21.34–0.74) 20.07 (20.44–1.21) 20.38 (20.98–0.22) 0.56
Differences were examined by the Kruskal–Wallis test. * P , 0.05; **
P , 0.01.

Table 4. Rank correlation coefficient (t) between the species richness indices and MDS site scores for
longicorn beetle assemblages of 1998 and the stand variables of study sites (n 5 14 sites).
Altitude Slope Number of woody Maximum DBH Maximum DBH of
plant species of conifers broad-leaved trees
Species richness indices
Number of species 20.036 20.217 0.268 0.430* 0.380 ms
Margalef index 0.265 20.061 0.156 0.380 ms 0.420*
MDS site scores
First dimension 20.219 0.012 0.045 20.313 20.508*
Second dimension 20.357 ms 0.209 20.022 20.067 0.331
ms
*P , 0.05; 0.05 , P , 0.1.
 1927

Figure 5. Cluster analysis of the longicorn beetle assemblages from 1998 to 1999. For site abbreviations,
see Figure 1. Boldfaced letters and italic letters indicate the samples of 1998 and 1999, respectively.

Species richness

The number of longicorn beetle species rose as the number of individuals collected
increased (Figure 3a; t 5 0.412, n 5 14, P , 0.05). It was not significantly
different among forest types, whereas the median value was largest in old-growth
forests (Table 3). The Margalef index was independent of the number of individuals
(Figure 3b; t 5 20.121, n 5 14, P . 0.5). It was significantly different among
forest types, and the median value decreased from old-growth forests to second-
growth forests to conifer plantations (Table 3). Neither value was significantly
different among directions (Kruskal–Wallis test; H 5 1.36, P . 0.5, for the number
of species; H 5 0.79, P . 0.5, for the Margalef index). While not correlated with
altitude, slope or the number of woody plant species, they were positively correlated
with the maximum DBH of conifers and broad-leaved trees, either significantly or
marginally (Table 4). Figure 4 shows the relationship of the Margalef index to the
number of woody plant species, and to the maximum DBH of broad-leaved trees.

Site ordination

As shown in the dendrogram in Figure 5, the longicorn beetle samples of 1998 and
1999 collected at every site (A, C–F) were close to each other, indicating that the
beetle assemblages were stable and did not change considerably from 1998 to 1999.
Figure 6 shows the final two-dimensional configuration of study sites by MDS
based on the rank correlation coefficients between them calculated from the samples
of 1998. The site score for the first dimension was significantly different among
forest types, while the score for the second dimension was not (Table 3). The former
1928

Figure 6. Two-dimensional ordination of study sites by the MDS based on the degree of similarity
between each pair of sites measured by means of the rank correlation coefficient (Kendall’s tau, t ) on the
abundance of longicorn beetle species collected in 1998. For site abbreviations, see Figure 1.

Figure 7. Relationships between the site score on the first MDS dimension and the maximum DBH of
broad-leaved trees (a), and between that on the second MDS dimension and altitude (b) (cf. Table 4). For
forest type symbols, see Figure 1.

increased from old-growth forests to second-growth forests to conifer plantations

(Figure 6). The site score for the first dimension was significantly correlated with
the maximum DBH of broad-leaved trees, whereas the score for the second
dimension was marginally correlated with altitude (Table 4, Figure 7). The site
scores were not different among directions (Kruskal–Wallis test; H 5 0.99, P . 0.5,
for the first dimension; H 5 4.61, P . 0.2, for the second dimension). Neither were
 1929

Figure 8. Number of longicorn beetle species collected in 1998 in relation to the site score on the first
MDS dimension.

they correlated with the slope, the number of woody plants and the maximum DBH
of conifers (Table 4).
Table 6 shows the rank correlation coefficients between the abundance and the
MDS site scores for 40 species collected at three or more sites. Negative correlation
with the site score for the first dimension was significant in 12 species, including six
species of the genus Pidonia. This shows that they are specific to or closely
associated with old-growth forests. On the other hand, positive correlation was
significant in four species, e.g. Parastrangalis spp., suggesting that they are absent
or uncommon in old-growth forests. For the other 24 species, no significant
correlation with the site score was shown for the first dimension. Three species
showed a positive correlation with the site score for the second MDS dimension,
which was somewhat correlated with altitude (Table 4).
For all eight species of Pidonia listed in Table 5, the correlation between the
abundance and the site score for the first MDS dimension was negative if not
significant at the 5% level. Eleven species of Pidonia, including three infrequent
species (P. yamato, P. neglecta, P. chujoi), were collected in 1998. The number of
Pidonia species was highly and negatively correlated with the site score for the first
MDS dimension (Figure 8; t 5 20.716, n 5 14, P , 0.001), although the number
of the other species was not correlated with it (Figure 8; t 5 0.012, n 5 14, P .
0.5). Also the number of Pidonia species was significantly different among three
forest types (Kruskal–Wallis test; H 5 6.65, P , 0.05).

Species indicator values

Species indicator values (IndVal) were computed for 40 species collected at three or
more sites in 1998. The values over 40% are shown in Table 5. Large IndVal for
old-growth forests were indicated in Dinoptera minuta, many Pidonia species, and
Pseudalosterna misella. Many species, including Parastrangalia spp., exhibited
1930

Table 5. Rank correlation coefficient (t ) between the number of individuals and the site scores in the
MDS ordination (n 5 14 sites) for the species collected at three or more sites in 1998, and the species
indicator value (IndVal .40%) for the species.
Family or Correlation coefficient IndVal Total number of
subfamily species (t) between abundance (%) individuals/sites of
and MDS site score occurrence a
First Second Old-growth Second-growth Conifer
dimension dimension forests forests plantations
(7 sites) (5 sites) (2 sites)
Disteniidae
D. gracilis 20.42* 0.13 42.9 3/3
Prioninae
Psephactus remiger 0.04 0.01 8/5 8/3 3/1
Lepturinae
Lemula japonica 0.02 0.37 3/2 1/1
Di. minuta 20.66** 20.17 57.1 4/4
Pidonia mutata 20.54** 20.01 54.3 49/5 11/2
P. signifera 20.46* 0.04 48.6 8/4 1/1
P. grallatrix 20.51* 0.31 83.0* 43/6 1/1
P. aegrota 20.45* 0.22 47.8 30/6 12/3 2/1
P. puziloi 20.64** 20.06 92.2* 422/7 18/3 3/1
P. approximata 20.38 0.36 56.5 23/6 6/3 1/1
P. amentata 20.12 0.53** 19/3 2/1
P. simillima 20.76** 20.09 82.3* 34/6 1/1
Ps. misella 20.44* 0.23 71.4* 12/5
Anoploderomorpha excavata 20.49* 0.24 42.9 3/3
Anastrangalia sequensi 0.19 0.27 4/2 3/2 2/1
Leptura ochraceofasciata 0.15 20.02 70.8* 86/6 86/5 143/2
Parastrangalis lesnei 0.43* 20.34 62.5 3/2 2/2
Pa. shikokensis 0.62** 0.05 90.9** 1/1 4/2
Idiostrangalia contracta 0.43* 20.11 81.4* 1/1 1/1 3/2
Cerambycinae
Allotraeus sphaerioninus 20.41* 0.00 18/5 6/3 5/1
Stenodryas clavigera 20.08 20.03 17/3 25/3 2/1
Ceresium holophaeum 20.02 0.41* 4/2 1/1
Cleomenes takiguchii 0.04 0.19 4/3 2/2
Chloridolum viride 20.37 0.40* 42.5 58/4 11/2
Callidiellum rufipenne 0.44* 20.02 93.3* 1/1 4/2
Xylotrechus emaciatus 0.11 20.24 6/4 2/2 5/1
X. pyrrhoderus 20.07 0.31 6/3 5/2
X. cuneipennis 0.12 20.31 5/4 3/3 4/1
X. grayii 0.19 0.09 70.0* 3/1 2/2
Demonax transilis 0.16 0.05 80.6* 5612/7 4118/5 13488/2
Lamiinae
Asaperda rufipes 20.03 0.36 48.3 42/6 35/5 3/1
Pterolophia tsurugiana 20.48* 20.34 42.9 4/3
Acalolepta fraudatrix 0.06 20.04 61.7* 13/6 19/5 1/1
A. sejuncta 20.12 20.35 9/4 7/4 1/1
Uraecha bimaculata 20.26 0.28 190/7 111/5 38/2
Xenicotela pardalina 0.07 0.27 4/3 3/3
Rhodopina integripennis 20.30 20.13 5/4 2/2
Pareutetrapha simulans 0.21 0.08 4/1 2/2 1/1
Praolia citrinipes 20.36 0.13 62.9 31/5 3/2
Glenea relicta 20.31 0.16 2/2 1/1
a
Boldfaced numbers show the data set for the IndVal. *P , 0.05; **P , 0.01.
 1931

Table 6. Host condition, larval host tissue and host trees for the indicator species with IndVal.
Host Larval host Host
condition tissue a trees b
Indicator species for old-growth forests
D. gracilis Dying or humid dead tree IB / SW BL / C
Di. minuta Dead twig BL
P. mutata
P. signifera Dead branch IB BL
P. grallatrix BL
P. aegrota Humid dead tree and branch IB BL
P. puziloi
P. approximata
P. simillima Dead branch IB BL
Ps. misella Standing tree IB / SW/ R BL
An. excavata Dead tree BL
Ch. viride Dead branch W BL / C
Pt. tsurugiana Dead branch IB / SW BL
Pr. citrinipes Dead tree and branch BL
Indicator species for second-growth forests
As. rufipes Dead twig W BL
Ac. fraudatrix Dead tree and branch IB / W BL / C
Indicator species for conifer plantations
L. ochraceofasciata Dead tree W C / BL
Pa. lesnei Dried dead branch IB / SW C / BL
Pa. shikokensis Dead tree BL
I. contracta Dead vine BL
Ca. rufipenne Dead or dying tree IB / SW C
X. grayii Dead tree BL
De. transilis Dead tree and branch W BL / C
Data sources are Kiyosawa et al. (1981), Kojima and Nakamura (1986), Kuboki (1987). a IB – inner bark;
SW – sap wood; W – wood; R – root. b BL – broad-leaved tree; C – conifer.

high indicator values for conifer plantations. Acalolepta fraudatrix was also
characteristic of second-growth forests. The results of the IndVal approach agreed
essentially with those from the MDS ordination analyses. According to literature
(Kiyosawa et al. 1981; Kojima and Nakamura 1986; Kuboki 1987), host condition,
larval host tissue and host trees for the indicator species with IndVal .40% are
compiled in Table 6.

Discussion

Limitations of trap sampling

As previously reported by Ikeda et al. (1993), Shibata et al. (1996) and Sakakibara
et al (1997a, b, 1998), the white collision trap baited with benzyl acetate to mimic
wild flowers is efficient for collecting flower-visiting species of the subfamilies
Lepturinae and Cerambycinae. On the other hand, the species without flower-
1932

visiting habits (Lamiinae, Prioninae and Spondylinae) are probably lured to ethyl
alcohol (Ikeda et al. 1980; Shibata et al. 1996; Sakakibara et al. 1997a). While
yellow traps are generally less attractive than white traps, certain species (e.g.
Disteniidae) have been largely collected in them (Sakakibara et al. 1997a). Thus, the
taxonomic bias observed in our trap sampling is fundamentally consistent with these
previous reports. The present combination of white and yellow collision traps baited
with benzyl acetate and ethyl alcohol appears to be useful for sampling diverse
longicorn beetles, although the collection is rather poor for non-flower-visiting
species.
Although the attractant traps lure the longicorn beetles moving just around them
(T. Ikeda, personal communication), they can catch those that fly from adjacent
stands as in other sampling methods. For example, adults of a lepturine species
(Anaglyptus subfasciatus) were caught in the traps located at a distance of 30–50 m
from the stands of emergence (Makihara 1992). Thus, the results may be biased by
the contamination with strong-flying species from other habitat types, especially
when the sampling site is small in area. However, such biases would be not serious
because our results show distinctive ordination of sampling sites corresponding to
forest types.
Malaise traps are also useful for the investigation of beetle assemblages (Maeto
and Makihara 1999). They can catch more non-flower-visiting species than the
collision traps with attractants, but they cost much more to operate than do collision
traps. To compare beetle assemblages among many sites at a time, collision traps
with attractants would be more practical than Malaise traps.
Although abundance data obtained from attractive traps are not direct measures of
real abundance of each species, the rank of abundance might change with the
alternation of species between sampling sites. Therefore, we used the rank correla-
tion coefficient (Kendall’s t ) as the similarity measure between assemblages (Ghent
1963). According to Huhta (1979), it is one of the best indices to measure the
species alternation of spiders and beetles in succession after clear-cutting. Our
results demonstrate that site ordination by MDS based on rank correlation co-
efficients is practical for analyzing the changes in insect assemblage using abun-
dance data obtained from attractant traps.
It is generally known that some herbivorous insects (moths, sawflies, bark beetles,
etc.) exhibit wide density fluctuations over years (e.g. Varley et al. 1973). If the
relative abundance of species varies greatly between years, 1-year sampling would
be insufficient to identify any indicator species being characteristic of certain habitat
types. However, the assemblages of longicorn beetles collected in two consecutive
years (1998 and 1999) were very close to each other for every site, so that the
general pattern of the beetle assemblages may be discussed based on a single year
sampling.

Decline of species richness

An obvious decrease in the Margalef index, the number of species adjusted for the
number of individuals, indicates that the conversion of old-growth forests into
 1933

second-growth forests and conifer plantations has diminished the species richness of
longicorn beetles. Shibata et al. (1996) mentioned that the number of longicorn
beetle species tended to increase in proportion to the number of woody plant
species. However, except for a cool-temperate beech forest, no correlation was
shown between the number of beetle species and that of woody plant species in the
warm-temperate forests in their study (Figure 3 of Shibata et al. 1996). This agrees
with our results, and it is not likely that forest conversion has reduced the species
richness of longicorn beetles through the decrease of woody plant species.
Our study suggests that the species richness of longicorn beetles increases with
the maximum diameter of trees. However, it is not surprising that old-growth forests
have large trees. Further examination will require more information about the
biology and natural history of the species closely associated with old-growth forests.

Ecological requirements and functions of the old-growth forest species

It was proved by the MDS ordination that the longicorn beetle assemblages of
old-growth forests were distinct from those of conifer plantations, while those of
second-growth forests were intermediate between them. Further analysis showed
that 12 species were specific to or closely associated with old-growth forests. Of the
12 species, six belong to the genus Pidonia of the subfamily Lepturinae. Further-
more, the total number of Pidonia species has definitely increased in old-growth
forests along the first MDS dimension. These findings indicate that most species of
Pidonia require some ecological conditions particular to old-growth forests, and
were supported by the results from the indicator value (IndVal) approach.
While it is widely distributed in the Holarctic Region, Pidonia is most highly
diversified in humid temperate forests of East Asia (Kuboki 1981). Larvae of
longicorn beetles feed on various parts (wood, sapwood or inner bark of trunks,
branches or roots) of woody plants in different conditions (decaying, dying or barely
living) (Hanks 1999). As shown in Table 6, specialization to inner bark is a peculiar
habit of Pidonia (Kuboki 1987). The larvae of this genus are found under the thick
bark of comparatively large, dead or living, trees and branches of various broad-
leaved species (Aceraceae, Araliaceae, Betulaceae, Cercidiphyllaceae, Fagaceae,
Rosaceae, Salicaceae) or, rarely, conifers (Pinaceae) (Kiyosawa et al. 1981; Kojima
and Nakamura 1986; Kuboki 1987). They pupate under the bark (never in wood) or
drop into humus for pupation. The humidity is unstable under thin bark, and
probably so they need thick bark under which they spend 1 or 2 years for growing
up. On the other hand, a thick, stable and moist humus layer of old-growth forests
may be also crucial to the larvae and pupae of Pidonia, which often live in humus or
under the bark of fallen logs lying on the forest floor (Kuboki 1987).
An additional species associated with old-growth forests, Distenia gracilis, also
prefers dead humid logs with thick bark to dried logs, having a small preference to
particular tree species (Kiyosawa et al. 1981). Another species, Ps. misella, is
known as a bark or root borer of standing trees of Salicaceae (Ohbayashi et al.
1992).
Therefore, it is most likely that the presence of large, dead or living, broad-leaved
1934

trees with thick bark is essential for the longicorn beetles that depend on warm-
temperate old-growth forests. This is not inconsistent with our finding that the beetle
assemblages change with the increase of the trunk size of broad-leaved trees. We did
not measure the volume and size of dead wood, while both the size of trees and
amount of dead wood usually increase together with forest age (e.g. Spetich et al.
1999; Siitonen et al. 2000).
In addition to decomposing woody materials by the larvae, Pidonia and other
lepturine beetles may be playing an important role in pollinating herbaceous and
woody plants, since they are the most dominant beetles visiting wild flowers in
temperate forests (Kato et al. 1990; Sakakibara et al. 1997b). According to Kuboki
and Shimamoto (1979) and Kuboki (1980), Pidonia species have been found on
wild flowers of more than 15 families (e.g. Saxifragaceae, Rosaceae, Umbelliferae,
Caprifoliaceae). Moreover, some species-specific relationships have been observed
between Pidonia and flowers (Kuboki 1980). It must be noted that the old-growth
forest beetles, depending on large full-grown trees, will support the reproduction
and genetic diversity of other plant species. They might also be major prey for
predatory insects, spiders and birds visiting wild flowers during late spring and early
summer. At any rate, further investigations are necessary to understand effects of the
decline in insect diversity of old-growth forests upon the forest ecosystem.

Implications for the management of forest landscape

Conversion of old-growth forests into young forests results in considerable changes

in the diversity of various arthropod guilds, e.g., canopy arthropods (Schowalter
1989), ground beetles (Niemela¨ et al. 1993), and xeric insects (Lattin 1993), but
there is general agreement that saproxylic arthropods, depending on large pieces of
dead wood, are most threatened by the long-term reduction of temperate old-growth
forests (e.g. Warren and Key 1991; Lattin 1993; Maeto and Makihara 1999;
Martikainen et al. 2000; Thunes et al. 2000). While the larvae occasionally bore into
living tress, Pidonia species are also saproxylic in a broad sense since they depend
on large woody materials that are dead or dying.
To enhance the diversity of saproxylic insects, extended rotation as well as
leaving old trees, snags and dead wood in clear-cuts is recommended (Hansen et al.
1991; Martikainen et al. 2000). Our results suggest that size increase of living, and
thus dead or dying broad-leaved trees will augment the diversity of old-growth
forest beetles like Pidonia. Extended rotation or postponement of cutting is no doubt
most important to increase large trees in second-growth forests, and thinning may be
also effective in accelerating the growth of remaining trees. Thinning in second-
growth forests and plantations should be recommended since it also enhances the
diversity of ground insects (Magura et al. 2000) as well as woody and herbaceous
plants. After thinning, felled trees should be retained in the forests to be later
available for saproxylic species. As suggested by Siitonen et al. (2000), it would be
the most efficient short-term management strategy for the increase of structural
diversity and old-growth attributes in managed forests to retain the old-growth
 1935

characteristics (i.e. large living trees, snags and logs) that already exist in mature
stands.
In southwestern Japan, old-growth forests of the warm-temperate zone have been
seriously fragmented due to traditional agriculture and coppicing as well as recent
rapid plantation with conifers (e.g. Shidei 1974; Sasse 1998). On Shikoku Island,
only a dozen small remnants of old-growth forests (each at most 300 ha) are found in
lowlands and hills up to about 1000 m in elevation, and conifer plantations and
young second-growth forests distantly separate them. For the conservation of
regional forest biodiversity, restoration of old forests from young second-growth or
man-made forests surrounding old-growth remnants is necessary to secure the
habitat area of old-growth forest species. It is also essential to re-establish a belt or
stepping-stones of old second-growth forests connecting old-growth remnants
within the region. Pidonia and other longicorn beetles closely associated with
old-growth forests, which can be easily monitored with simple traps, may be
valuable indicators for the progress of such forest restoration.

Acknowledgements

We would like to thank Ryuichi Tabuchi, Takeshi Sakai, Shigeo Kuramoto and
Atsushi Sakai for the survey of site vegetation. Our thanks are also due to Toshihiko
Yamasaki for his help in the field, Tsuyoshi Yamada for showing us meteorological
data at Yusuhara, and Mariko Takeuchi for the preparation of insect specimens. We
also thank Shikoku Regional Forest Office, Kochi Prefecture and Yusuhara Town
for the permission to do field work in their forests. This work was partly funded by
the International Collaborative Research Programme of the Ministry of Agriculture,
Forestry and Fisheries.

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