全 文 ：第 49 卷 第 5 期
2 0 1 3 年 5 月
林 业 科 学
SCIENTIA SILVAE SINICAE
Vol. 49，No. 5
May，2 0 1 3
doi:10．11707 / j．1001-7488．20130517
Received date: 2012 － 02 － 21; Revised date: 2012 － 10 － 10。
Foundation project: Special Fund for Forest Scientific Research in the Public Interest ( 201004077 ) ; National Natural Science Foundation
(31170607) ;Program for Changjiang Scholars and Innovative Research Team in University of China( IRT1035) ．
* Corresponding author: Chen Hui． We thank Lian Xihong ( Northwest A＆F University) for his comments and discussion for some of the ideas
presented in this work，and grateful to Qinling National Forest Ecosystem Research Station，Northwest A＆F University for supporting the necessary
experiment condition．
华山松 16 种小蠹(鞘翅目: 象虫科: 小蠹亚科)
前胃结构的形态学*
卜书海 陈 辉
(1. 西北农林科技大学生命科学学院 杨凌 712100; 2. 西北农林科技大学林学院 杨凌 712100)
摘 要: 通过光学显微和扫描电镜技术，对危害华山松的 16 种小蠹前胃形态结构进行观察。根据前胃板的结构
特征可以将 16 种小蠹前胃分成 3 个类型，这些前胃形态结构的差异可以作为小蠹分类鉴定的辅助手段，且前胃类
型与韧皮部小蠹虫在华山松树干上的空间分布相吻合，华山松韧皮部小蠹前胃结构的特化是适应食物类型(营养
的累积和质地)的结果。
关键词: 华山松; 前胃形态; 小蠹
中图分类号: S718. 7 文献标识码: A 文章编号: 1001 － 7488(2013)05 － 0128 － 07
A Morphological Study on the Proventriculi of Sixteen Species of Bark Beetles
(Coleoptera: Curculionidae: Scolytinae) Bred in Pinus armandi
Bu Shuhai1 Chen Hui2
(1. College of Life Sciences，Northwest A＆F University Yangling 712100; 2. College of Forestry，Northwest A＆F University Yangling 712100)
Abstract: Proventriculus morphology of 16 bark beetle species ( Curculionidae: Scolytinae ) from the host tree—
Chinese white pine ( Pinus armandii) was studied by using a light microscopy and a scanning electron microscopy．
Obvious differences were observed in the sclerotization degree，the shape of the anterior plate and type of sclerotized
appendices among the different bark beetle groups． Three types of proventriculi were identified according to the basic
morphological comparison of the chitinous plates，and their morphology may supply auxiliary characters for taxonomy of
these bark beetle species． The spatial distribution of proventricular types was consistent with the occurrence of spatial
sequence for 16 species of bark beetles in Chinese white pine trees． Obvious morphological variation and the sclerotization
degree of proventriculus may reflect an adaptation to variations in diet．
Key words: Pinus armandii; proventricular morphology; bark beetles
Scolytinae is a large and highly adapted group of
weevils of considerable economic significance because
of their impact on trees and forestry industry
(Rudinsky，1966; Wood，1982; Miller et al．，1989;
Klepzig et al．，1991; Wallin et al．，2002) ． In Pinus
armandii ( Chinese white pine ) stands in Qinling，
Shaanxi，China，there are about 22 species of bark and
ambrosia beetles forming a stable ecosystem by complex
interactions among spatial and trophic competition and
cooperation (Chen et al．，2007) ．
The proventriculus is ectodermic in origin; it is
lined by a sclerotized cuticular intima． The structure of
the proventriculus in bark beetles was briefly studied
by several authors such as Balogun (1969)，Nobuchi
(1969 )，Yin et al． ( 1984 )，Lopez-Buenfil et al．
(2001) and Díaz et al． (2000; 2003) ． Their studies
concentrated on the anatomy and histology of the
proventriculus and formed preliminary work on the
physiology of its nutrition． In all species studied，
display similar general pattern with each other，as well
第 5 期 卜书海等: 华山松 16 种小蠹(鞘翅目: 象虫科: 小蠹亚科)前胃结构的形态学
as between male and female． Taking the proventriculus
of Dendroctonus armandi as an example，it looks like a
small lantern or bulb from an outer view (Fig． 1) (Bu
et al．，2009) ． It is formed by eight thick folds of the
proventricular wall，which is composed of circular and /
or longitudinal muscles，an epithelial cell layer and
chitinous plates projecting to the proventriculus lumen
(Fig． 1) ． Each chitinous plate can be divided in two
regions: an anterior plate covered by cuticle with
denticles， spines and a masticatory plate with the
movable brushes carrying bristles．
Fig． 1 Longitudinal section of the proventriculus of Dendroctonus armandi
c:Cuticles; cm:Circular muscle; e:Epithelium; f:Fold; fd:Food;
lm:Longitudinal muscle; Pv:Proventriculus; sb:Stopping bristles;
Sv:Stomodeal valve; td:Transverse denticle． Light microscope．
In scolytid insects，the structures of masticatory
plates are similar． Their cuticles are highly
sclerotizated and in general have many bristles or hairs
to form the comb-like movable brushes． These brushes
and long bristles can stir and filter solid particles from
the anterior plate contents down to the midgut as results
of muscular action，while stopping bristles function
controlling the descending rate and quantity of food． At
the anterior plates，their cuticles are sclerotized at the
different level，and form grindstone with many spines
or denticles，which can crush and triturate foods from
the crop together with circle muscular action．
In our research， different types of the
proventriculi are focused on，with comparision of the
spatial distribution of bark beetles in the Chinese white
pine． Basically，as an inlet of midgut，proventriculus
plays a protective function and primary digestion on
solid food with potential poisonous material．
Meanwhile， microhabitat variations could strongly
indicate the microdifference of proventriculi． Thus we
examined the ultrastructures of the proventriculi from
16 species of bark beetles in the Chinese white pine by
light microscopy and scanning electron microscopy
( SEM ) in this study． We attempted to find
differences， by comparative morphology of
proventriculi， to reveal the morpho-functional
adaptations to the subcortical environment in the host
tree P． armandii．
1 Materials and methods
The adult specimens of bark beetles for this study
were collected directly from the bark of infested P．
armandii over 30 years old on the south side of the
Qinling Mountains (33° 36 N，108° 28 E，1 800 －
2 000 m a． s． l． ) between Huoditang and Xunyangba of
Ningshan County，Shaanxi Province，China，from April
to June 2009． In detail，twenty white pine trees under
attack by bark beetles were random logged from Chinese
white pine pure stand and mixed stands in three
months． Two hundred forty square sample phloem
pieces，sampling unit the size of the diameter at the
location on the tree trunk，were dissected from the
logged trees at 1 m intervals from the root-collar of the
trunk to upper trunk ＜ 5 cm diameter upper of trunk．
Each species of bark beetles was identified by male
seminal rods and external characters according to Yin et
al． ( 1984 ) ． We examined 16 species of 10 genera
belonging to six tribes: Hylastini， Hylurgini，
Polygraphini，Dryocoetini，Ipini and Cryphalini (Wang
et al．，2012; Bouchard et al．，2011; Yin et al．，1984)
(Tab． 1) ．
Ten specimens of each species were dissected，
and the chitinous plates of proventriculi were isolated
from the alimentary canal． Five proventriculi of each
species were transferred to Bouins fixative solution for
24 h，dehydrated in ethanol series，and embedded in
paraffin． Serial sections of 10 μm were cut on a rotary
microtome， de-paraffinized and stained with
hematoxylin-eosin． Five proventriculi of each species
for SEM ( scanning electron microscopy) were digested
in 10% NaOH for 30 min before cleaning in Ultrasonic
cleaners for 1h to washout food particles from the endo-
surface of chitinous plates． The chitinous plates were
dried for 72 h at 4 ℃ and mounted on aluminium
921
林 业 科 学 49 卷
stubs， then sputter-coated with gold / palladium
(40∶ 60) in a polaron E5400 high-resolution sputter
coater． Specimens were examinated in a JEOL T330
SEM operated at either 15 kV． Comparing proventriculi
among bark beetles， six characters were used for
taxonomic considerations (Tab． 1) ．
2 Results
Within the basic morphological pattern described
in Dendroctonus armandi， some variation can be
considered among the different bark beetle species
analyzed． Furthermore，the sclerotizated appendices of
each chitinous plate，which are more or less different
in size，shape and number among these bark beetles
include transverse tooth lines，anterior margin，median
suture， median denticles， basic denticles and
substitute stopping bristles in anterior plate and
declivity，masticatory brushes and stopping bristles in
the masticatory plate (Tab． 1) ．
Tab． 1 List of scolytinae taxa used in the analysis and annotated list of proventriculi characters
Tribes Species
Character
A B C D E F G
Hylastini Hylastes parallelus Chapuis，1875 0 0 0 0 0 0 1
Hylurgops longipillus Reitter，1894 0 0 0 0 0 0 1
Hylurgini Tomicus piniperda (Linnaeus，1758) 0 0 0 0 0 0 0
Dendroctonus armandi Tsai et Li，1959 0 0 0 0 0 0 1
Polygraphini Polygraphus polygraphus(Linnaeus，1758) 1 1 0 0 0 0 1
Polygraphus sinensis Eggers，1933 1 1 0 0 0 0 1
Dryocoetini Dryocoetes hectographus Reitter，1913 1 1 1 2 0 0 2
Dryocoetes autographus (Ratzeburg，1837) 1 1 1 2 0 0 2
Ipini Pityogenes japonicus Nobuchi，1974 1 2 1 2 2 0 2
Ips acuminatus(Gyllenhal，1827) 1 2 1 1 1 2 2
Ips sexdentatus(Boerner，1767) 1 2 1 1 1 2 1
Ips mannsfeldi(Wachtl，1880) 1 2 1 1 0 2 1
Orthotomicus laricis(Fabricius，1792) 1 2 1 2 0 1 2
Cryphalini Cryphalus lipingensis Tsai et Li，1959 1 2 2 2 0 0 3
Cryphalus chinlingensis Tsai et Li，1963 1 2 2 2 0 0 3
Cryphalus pseudochinlingensis Tsai et Li，1963 1 2 2 2 0 0 3
A． Types of denticles: 0． Transverse tooth; 1． Median denticles． B． Shapes of median suture: 0． No median suture; 1． Wide; 2． Narrow． C．
Arranges of basic dentucle: 0． No basic dentucle; 1． 1 row; 2． 3 rows． D． The space between median denticles and basic dentucle: 0． None; 1． Far;
2． Close． E． Shape of anteriorior plate: 0． Rectangle; 1． Long columnar; 2． Short columnar． F． Number of substitute stopping bristles: 0． Degeneration
and deletion; 1． A few; 2． Many． G． Shape of stopping bristles: 0． Long columnar-shaped; 1． Thread-shaped; 2． Thread-shaped with spine-like
projections; 3． Finger-like．
Hylastes parallelus， Hylurgops longipillus，
Tomicus piniperda and Dendroctonus armandi present
the high density of tooth-like projections that form
plates with 12 － 21 rows of transverse tooth lines (Fig．
2 ) in the proventriculi． Unlike 4 species above，
Polygraphus polygraphus and Polygraphus sinensis have
sharp median denticles and wide median suture without
transverse tooth lines ( Fig． 3A， 3B; Tab． 1 ) ．
Dryocoetes hectographus and Dryocoetes autographus
have median denticles and wide median suture，and
basic denticles，which are arranged like a converse
“V” shape，are located at the end of the column of
median suture (Fig． 3C，3D; Tab． 1) ．
Among Pityogenes japonicus，Ips acuminatus，Ips
sexdentatus，Ips mannsfeldi and Orthotomicus laricis，2
columns of median denticles are linked with each other
by narrow median suture of the proventricular plate
(Fig． 4A，4B，4C，4D; Tab． 1)，contrasting to wide
median suture and without median suture found in the
other bark beetles analyzed． Cryphalus lipingensis，
Cryphalus chinlingensis and Cryphalus
pseudochinlingensis have 3 rows of basic denticles like
3 rows of palisades，each row of which has long basic
denticles in the middle ( Fig． 4F，4G，4H; Tab． 1) ．
In Orthotomicus laricis， the appearance of median
suture is sharper than those of Pityogenes japonicus，Ips
acuminatus，Ips sexdentatus and Ips mannsfeldi，and
the demarcation line between its basic denticles and
median sutures is not distinguished because of their
similar appearance (Fig． 4E) ．
031
第 5 期 卜书海等: 华山松 16 种小蠹(鞘翅目: 象虫科: 小蠹亚科)前胃结构的形态学
Fig． 2 SEM micrographs，chitinous plates of the proventriculi of stype A
A． Dendroctonus armandi ; B． Tomicus piniperda ; C． Hylurgops longipilis ; D． Hylastes parallelus ． d: Denticles;
de: Declivity; mb: Masticatory brushes; fd: Food; s: Spines; sb: Stopping bristles; tl: Transverse tooth lines．
Fig． 3 SEM micrographs，chitinous plates of the proventriculi of stype B
A． Polygraphus sinensis ; B． Polygraphus polygraphus; C． Dryocoetes hectographus; D． Dryocoetes autographus． am: Anterior
margin; bd: Basic denticles; de: Declivity; mb: Masticatory brushes; md: Median denticles; ms: Median satures; fd: Food; s:
Spines; sb: Stopping bristles．
131
林 业 科 学 49 卷
Fig． 4 SEM micrographs，chitinous plates of the proventriculi of stype C
A． Ips sexdentatus ; B． Ips acuminatus; C． Ips mannsfeldi; D． Pityogenes japonicus; E． Orthotomicus laricis; F． Cryphalus pseudochinlingensis; G．
Cryphalus lipingensis; H． Cryphalus chinlingensis． am: Anterior margin; bd: Basic denticles; de: Declivity; mb: Masticatory brushes; md: Median
denticles; ms: Median satures; fd: Food; s: Spines; sb: Stopping bristles; ss: Substitute stopping bristles．
Similarly， three types of proventriculi are
identified according to the basic morphological
comparison of the chitinous plates． ( 1 ) Type A．
Dendroctonus armandi as a representative share many
characters with Tomicus piniperda，Hylastes parallelus，
and Hylurgops longipilis． The obvious features in this
type are the developed transverse tooth lines without
median denticles， basic denticles and substitute
stopping bristles for weak degree of sclerotization ( Fig
2) ． (2) Type B． This type appears as a medial degree
of sclerotization among Polygraphus polygraphus，P．
sinensis， Dryocoetes hectographus， and D．
autographus． Apparently，this type differs from Type A
in having the wide median sutures and developed
median denticles，whereas the proventricular plates in
Dryocoetes have developed basic denticles ( Fig． 3 ) ．
(3) Type C． The remaining 8 species belong to this
type in which the degree of sclerotization is high． Eight
beetles above all have the developed basic denticles，
apart from developed median denticles． In Type C，the
median suture is very narrow，compared with Type B
(Fig． 4) ．
3 Discussion
For bark beetles，carbohydrates，vitamins，fiber
and minerals are the most important nutritional
constituents from pine cambium． When they invaded
the host trees the bark beetles with the assisting of their
associated fungus can break the physical structure of
phloem and xylem tissue，block resin ducts of xylem
tissue，and cause resin and water metabolic confusion，
which will cause the host trees weaken， then，
secondary bark beetles were induced to attack the
suppressed and dying phase of the pine ( Xie et al．，
2008; Christiansen et al．，1990; Ren et al．，1959) ． Li
et al． (2009) thought that the content of total protein，
231
第 5 期 卜书海等: 华山松 16 种小蠹(鞘翅目: 象虫科: 小蠹亚科)前胃结构的形态学
lipid，and glycogen of the barks were related linearly to
physical life (healthy，suppressed and dying phase) of
Chinese white pine． The degree of sclerotization of the
proventriculi， to some extent， could possibly be
correlated with major types ( the hardness and sappy
degree) and nutrition of food from different part of
healthy， suppressed， diseased， dying and decaying
trees( Pu et al．，2007) ． Our studies showed that the
types of the proventriculi were consistent with the
spatial distributions among sixteen bark beetles for a
shared host (Chinese white pine) resource，generally．
In spatial distribution of bark beetles in trunks，the
first group ( Hylastes parallelus， Dendroctonus
armandi，I． sexdentatus and Hylurgops longipilis) were
common species at the low trunk ( 1 － 6 m ); The
second group consisted of Dryocoetes hectographus，D．
autographus， Polygraphus sinensis， P． polygraphus
and I． mannsfeldi， I． acuminatus， O． larici and
Pityogenes japonicus，were concentrated at the middle
trunks (7 － 12 m); The third group (C． lipingensis，
C． chinlingensis， C． pseudochinlingensis and T．
piniperda) were predominant species at the upper trunk
and branches of Chinese white pines ( 11 － 16 m) ．
Similarly，the species of the first group have the weak
degree of sclerotization plates except for I． sexdentatus，
and fed on the thicker texture of phloem with the high
content of total protein and lipid from healthy or
suppressed trees． The species at the middle trunks is a
complex group with the middle and high degree of
sclerotization plates，who ate thick texture with low
content of total protein and lipid from suppressed trees．
The third group which ate thin texture of phloem and
xylem with lower content of total protein and lipid from
dying tree，have the high degree of sclerotization plates
except T． piniperda． So the morphological variation
and the degree of sclerotization of anterior plate
reflected the subtle feeding habits of these bark
beetles．
Our results indicated various types of proventriculi
might undertake different detoxification in digestive
process of bark beetles． The action and structure of
anterior plate can prolong retention time of food，which
improve the effective rate of neutrolizing toxic materials
mixed with digestive enzyme ( e． g． PAL，PPO，SOD)
from saliva，midgut and symbiotic fungus． To type A
such as D． armandi，the bark beetles had the complex
structures and showed the highest detoxifying degree，
than weaker in type B such as Polygraphus species，
and the weakest in Type C such as Ips species． The
differences of proventricular types reflected their
defense resistance to special physical phase of Chinese
white pine．
The structure of the proventriculus usually has
considerable variations among different insects that feed
on solid food with chewing mouthparts such as in the
cockroach ( Blattodea )， cricket ( Orthoptera )，
Rhynchophorinae and Hydrophilidae beetles
( Boudreaux， 1980; Chapman， 1998; Fontanetti
et al．，2002) ． The diversity in the morphology of the
proventriculus of insects has led several authors to use
it as an auxiliary character in systematics and
phylogeny， with special emphasis on the
proventriculus， whose morphological variability is
related to the diversity of insect feeding habits (Rentz
et al．，1990; Bland et al．，1991; Fontanetti et al．，
2002; Szinwelski et al．，2009; Serro，2001; 2007) ．
In the present study， these results indicate that the
morphological structure of the proventriculus exhibited
some divergence，and may supply auxiliary taxonomy
characteristics for the bark beetles in Chinese white
pine，especially useful for groups， since for many
species the external morphology is insufficient for
characterization of genera and species． We also believe
that these differences may reflect an adaption to
variations in diet to which these populations attacked
and brood developed in Chinese white pine．
参 考 文 献
Balogun R A． 1969． The anatomy and histology of the proventriculus of
Ips cembrae Heer ( Coleoptera: Scolytidae) ． Proc Royal Ent Soc
Lond，44 (10 － 12) : 158 － 161．
Bland T G，Rentz D． 1991． Studies in Australian Gryllacrididae: the
proventriculus as a taxonomic character． Invertebr Taxon，5 (2 ) :
443 － 455．
Boudreaux H B． 1980． Proventricular acanthae and their phylogenetic
implications． Ann Entomol Soc Am，73 (2) : 189 － 196．
Bouchard P，Bousquet Y，Davies A E，et al． 2011． Family-group names
in Coleoptera ( Insecta) ． ZooKeys，88: 1 － 972．
Bu S H，Chen H． 2009． The Alimentary Canal of Dendroctonus armandi
Tsai and Li ( Coleoptera: Curculionidae: Scolytinae ) ． The
Coleopterists Bulletin，63 (4) : 485 － 496．
Chapman R F． 1998． The Insects: Structure and Function 4 th ed．
331
林 业 科 学 49 卷
Cambridge university Press，Cambridge，UK．
Chen H，Tang M． 2007． Spatial and temporal dynamics of bark beetles
in Chinese white pine in Qinling Mountains of Shaanxi Province，
China． Environ Entomol，42 (5) : 1124 － 1130．
Christiansen E，Solheim H． 1990． The bark beetle associated blue-stain
fungus Ophiostoma polonicum can kill various spruces and Douglas
fir． Eur J For Path，20 (6 － 7) : 436 － 446．
Díaz E， Arciniega O， Sánchez L， et al． 2003． Anatomical and
histological comparison of the alimentary canal of Dendroctonus
micans， D． ponderosae， D． pseudotsugae pseudotsugae， D．
rupifenis，and D． terebrans (Coleoptera: Scolytidae) ． Ann Entomol
Soc Am，96 (2) : 144 － 152．
Díaz E， Cisneros R， Zúiga G． 2000． Comparative anatomical and
histological study of the alimentary canal of Dendroctonus frontalis
(Coleoptera: Scolytidae) ． Ann Entomol Soc Am，93 (2) : 303 －
311．
Fontanetti C S， Zefa E， Passeti F， et al． 2002． Morphological
characterization and comparative analysis of the proventriculus from
three species of Endecous saussure，1878 ( Orthoptera: Gryllidae:
Phalangopsinae) ． Entomotropica，17 (1) : 15 － 23．
Klepzig K D，Smalley E B，Raffa K F． 1991． Combined chemical
defenses against insects and fungi associated with a forest decline
disease． J Chem Ecol，22: 1376 － 1388．
Li Z (李臻)，Chen H (陈辉)，Wang S J (王胜军) ． 2009． Utilization
of nutritients and mineral elements in Pinus armandii by Chinese
white pine beetle (Dendroctonus armandi) ． Scientia Silvae Sinicae
(林业科学)，45 (11) : 98 － 103．
Lopez-Buenfil J A， Valdez-Carrasco J， Equihua-Martinez A， et al．
2001． El proventriculo como estructura para identificar generos
Mexicanos de Scolytidae ( Coleoptera ) ． Folia Entomologica
Mexicana，40:325 － 372．
Miller D R，Borden J H，Slessor K N． 1989． Inter- and intrapopulation
variation of the pheromone， ipsdienol produced by male pine
engravers，Ips pini ( Say) (Coleoptera: Scolytidae) ． J Chem Ecol，
15: 233 － 247．
Szinwelski N，Rodrigues M S，Pereira M R，et al． 2009． Proventriculus
of Three Nemobiinae Crickets ( Orthoptera: Grylloidea:
Trigonidiidae) ． J Orthop Res，18 (1) : 59 － 63．
Nobuchi A． 1969． A comparative morphological study of the
proventriculus in the adult of the superfamily Scolytoidea
(Coleoptera) ． Gov For Exp Sta Bull，224: 39 － 110．
Pu X J (蒲晓娟)，Chen H (陈辉) ． 2007． Relations between attacking
of Dendroctonus armandi and nutrition and resistance materials of
host trees ( Pinus armandi) ． Journal of Northwest A＆F University
(西北农林科技大学学报)，35 (3) : 106 － 110．
Ren Z F (任作佛 )，Dang X D (党心德 ) ． 1959． Investigation and
control of Dendroctonus armandi Tsai et Li in Qinling Mountain． J
Northwest Agric Coll (西北农学院学报)，(2) : 59 － 89．
Rentz D C F， John B． 1990． Stuclies in Australian Gryllacrididae:
Taxonomy，biology ecology and cytology． Invertebrate Taxonomy，3:
1053 － 1210．
Rudinsky J A． 1966． Host selection and invasion by the Douglas-fir
beetle，Dendroctonus pseudotsugae Hopkins，in coastal Douglas-fir
forests． Can Entomol，98: 98 － 111．
Serro J E． 2001． A comparative study of the proventricular structure in
corbiculate Apinae (Hymenoptera: Apidae) ． Micron，32 (4) : 379
－ 385．
Serro J E． 2007． Proventricular structure in the bee tribe Augochlorini
(Hymenoptera: Halictidae) ． Org Divers Evol，7 (3) : 175 － 180．
Wallin K F，Raffa K F． 2002． Density-mediated responses of bark beetles
to host allelochemicals: A link between individual behavior and
population dynamics． Ecol Entomol，27 (4) : 484 － 492．
Wang Z L (王志良)，Zhang R Z(张润志) ． 2012． Taxonomic status of
bark beetles ( Coleoptera，Curculionidae ) ． Acta Zootaxonomica
Sinica(动物分类学报)，37 (2) : 291 － 295．
Wood S L． 1982． The bark and ambrosia beetles of north and central
America ( Coleoptera: Scolytidae)，a taxonomic monograph． Great
Basin Nat Mem，6: 1 － 1359．
Xie S A (谢寿安)，Lu S J(吕淑杰)，Axel S，et al． 2008． Anatomical
characteristics in xylem tissue of Pinus armandi infected by the bark
beetle Dendroctonus armandi ( Coleoptera: Scolytidae ) and its
associated blue-stain fungus Ceratocystis polonica． Acta
Entomologica Sinica (昆虫学报)，51 (12) : 1327 － 1333．
Yin H F (殷慧芬)，Huang F S (黄复生)，Li Z L (李兆麟) ． 1984．
Economic insect fauna of China． Fasc． 29． Coleoptera． Scolytidae．
Science Press (科学出版社)，Beijing，China．
(责任编辑 朱乾坤)
431