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Cerambycidae of Northern Asia
Volume 2, Part I

Ag TT 85-1-0754

536

£ j3 ? :6 AKADEMIIA NAUK SSSR

^ 2^ y SiBIRSKOE OtDELENIE

V I Biologicheskii Tnstitut

f '■ ACADEMY OF SCIENCES OF THE USSR

Siberian Division

Biological Institute

CERAMBYCIDAE OF
NORTHERN ASIA

VOLUME 2
Cerambycinae
Part I

[Usachi Severnoi Azii {Cerambycinae)]

A.I. CHEREPANOV

Translated from Russian

Amerind Publishing Co. Pvt. Ltd., New Delhi

1988

Nauka Publishers, Siberian Division
Novosibirsk, 1981

Translated and published under an agreement for the United States

Department of Agriculture, Washington, D C., by

Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001

Translator: P.M. Rao

General Editor: Dr. V.S. Kothekar

Printed at Gidson Printing Works, New Delhi, India

UDC 595.768.1

This monograph presents keys for the identification of the tribes, genera,
and species of the subfamily Cerambycinae (tribes Hesperophanini, Ceram-
bycini, Callidiopini, Graciliini, Obriini, Nathriini, Molorchini, Dilusini,
Callichromini, Rosaliini, and Callidiini). The morphology, biology, charac-
teristics of development, host relationships, and other specific features are
described and the role of each species in the conservation of nature assessed.
The system of ecological concomitants of cerambycid beetles inhabiting
widely separated regions is analyzed and the evolutionary processes in the
subfamily Cerambycinae of northern Asia during the post-Tertiary period
are traced.

This monograph is intended for entomologists, ecologists, and workers
engaged in the control of forest pests. It can also be used as a field guide by
biology students in universities, forestry and educational institutes, and specia-
lized technical schools.

Editor-in-Chief

N.A. ViOLOVICH
Doctor of Biological Sciences

CONTENTS

FOREWORD ix

SYSTEMATIC LIST OF SPECIES ............ ........ xi

V. Subfamily Cerambycinae 1

12. Tribe Hesperophanini ............................ 15

13. Tribe Cerambycini .............................. 20

14. Tribe Callidiopini 24

15. Tribe Graciliini . 27

16. Tribe Obriini 31

17. Tribe Nathriini (= Psebiini) 54

18. Tribe Molorchini 61

19. Tribe Dilusini 105

20. Tribe Callichromini 113

21. Tribe Rosaliini . 148

22. Tribe Callidiini . 155

SUPPLEMENT 279

INDEX OF LATIN NAMES OF CERAMBYCID BEETLES 288

FOREWORD

This book is a continuation of the study of the species composition,
geographic distribution, morphological identification of preimaginal
stages, and biology of northern Asian cerambycid beetles (Ceramby-
cidae). In 1979 the first volume of Usachi Severnoi Azii (Cerambycidae
of Northern Asia) was published and covered four subfamilies (Prioni-
nae, Disteniinae, Lepturinae, and Aseminae). This volume included 124
species, of which the preimaginal stages of 99 were identified, the charac-
teristic features of biology described, general features of phenology elu-
cidated, and host specificity determined. The taxonomic position of some
species was clarified and some genera were revised (Evodinus and
Brachyta).

The present monograph is devoted to the subfamily Cerambycinae.

This book includes the results of investigations conducted at various
times over vast territories — from the Urals (inclusive) to the coast of the
Pacific Ocean, and from northern Kazakhstan, Altai, Amur, and Lake
Khasan in the south to the forest tundra in the north. I have presented
descriptions of 105 species, identified the preimaginal stages of 86, deter-
mined the calendar periods of population growth under field conditions,
assessed many ecological aspects, elucidated characteristics of develop-
ment, recorded variations in weight indices of insects during metamor-
phosis, presented the duration of the life cycle of each species, given
information on the role of each in the biocenose, and other aspects.

Of the 19 species whose biology was not studied by me, only the
type specimens are known for some (Xylotrechus pavlovskii Plav., X.
mixtus Plav., Clytus hypocrita Plav., Chlorophorus ubsanurensis Tsher.,
and Amarysius grallator Baeckm.) and stray finds for others {Stenygrinum
quadrinotatum Bat., Xylotrechus rufilius Bat., X. capricornis Gebl., X.
villioni Vill., Clytus nigritulus Kr., C. venustulus Plav., Chlorophorus
sartor Mull., Purpuricenus petasifer Fairm.). Thus all these species are
very rarely found in northern Asia. Information on the range of some
species in northern Asia {Xylotrechus chinensis Chevr., Chlorophorus
figuratus Scop.) needs to be confirmed.

The biology of some species [Gracilia minuta (F.), Nathrius brevipen-
nis (Muls.), Molorchus umbellatarum (Schreb.), and Clytus arietis (L.)],
distributed only from Europe up to the southern Urals, has been traced
by me into the forests of northern Caucasus (Kislovodsk and Sochi).

I also succeeded in identifying monotypic development among some
proximate species. For example, Clytus arietoides Reitt., growing on
coniferous species, is widely distributed in Siberia. Clytus arietis (L.),
colonizing the shoots of deciduous species, is more often found in the
forests of northern Caucasus. However, pupation is completed in both
these species by the end of summer, the pupae enter diapause and
remain in hibernation, and adults emerge in May-June.

4 This monograph is a continuation of the first book, devoted to the
subfamilies Prioninae, Disteniinae, Lepturinae, and Aseminae. The classi-
fication adopted here is the same as that by several present-day resear-
chers (Gressit, 1951; Kojima and Hayashi, 1960; Linsley, 1962-1964).
In describing the genera and species, publications after 1940 are cited.
1 did not consider it necessary to cite all the references given in the
major compilation of N.N. Plavil’shchikov (1940).

I will consider my task fulfilled if this book provides relatively com-
plete and up-to-date information not only on the species composition
and geographic distribution, but also the bionomics of the subfamily
Cerambycinae inhabiting northern Asia as a whole, and if this mono-
graph long serves as a reference work for naturalists and in solving the
problems of conservation of nature.

N.E. Cherepanova conducted laboratory and field investigations with
extraordinary zeal and scrupulous precision and made significant contri-
butions to the biology of long-horned beetles Without her assistance,
this monograph could not have assumed its present comprehensive
form.

V.K. Stroganova, V.I. Ostanin, A.Yu. Kharitonov, A.V. Barkalov,
A.L. Pakhotskaya, and other colleagues at the Biological Institute parti-
cipated in the collection of field material from time to time.

I am sincerely grateful to artist A.Z. Ermolenko who not only drew
the illustrations for this monograph, but also assisted in the collection
of field material. I likewise sincerely thank all those who rendered assis-
tance and extended cooperation in the completion of this work.

All the illustrations are original; some have been published earlier in
various journals.

This monograph, devoted to the subfamily Cerambycinae, has been
published in two parts. Part I provides a list of long-horned beetles, keys
to the tribes based on different stages of development, and information
on the taxonomy, geography, and biology of the tribes Hesperophanini
to Callidiini. Part II gives keys to the genera and species of the tribes
Clytini and Stenaspini based on the adult, larval, and pupal stages, and
provides information on the morphology, geographic distribution, and
biology of each species.

SYSTEMATIC LIST OF SPECIES

Family CERAMBYCIDAE

V. Subfamily Cerambycinae

12. Tribe Hesperophanini

1. Genus Trichoferus Woll.

1. T. campestr is (Fsild.) 15

13. Tribe Cerambycini

1 . Genus Mallamhyx Bat.

1. M. raddei (Bless, and Sols.) 21

14. Tribe Callidiopini

1. Genus Stenygrinum Bat.

1. S. quadrinotatum Bat. 25

15. Tribe Graciliini
1. Genus Gracilia Serv.

1. G, minuta (F.) 28

16. Tribe Obriini
1. Genus Obrium Curt.

1. O. cantharinum (L.) 34

2. O. brevicorne Plav 39

3. O. gracileVldiN 44

2. Genus Stenhomalus White

1. S. vulcanus Tsher 49

17. Tribe Nathriini ( = Psebiini)

1 . Genus Nathrius Breth. ( = Leptidea Muls.)

1. N. brevipennis (Muls.) 55

Xll

18. Tribe Molorchini

1. Genus Molorchus F.

1. M. minor (L.) 65

2. M. ussuriensis Plav 71

3. M. umbellatarum (Schreb.) 75

4. M. kiesenwetteri Muls. and Rey 80

5. M. heptapotamicus Plav 86

6. M. kohotokensis Ohb 91

7. M. incognitus Tsher 95

2. Genus Nadezhdiana Tsher.

1. N. villosa Tsher 100

19. Tribe Dilusini
1. Genus Deilus Serv.

1. D.fugax (Oliv.) 106

20. Tribe Callichromini
1. Genus Aromia Serv.

1. A. moschata (L.) 116

2. Genus Chloridolum Thoms.

1. C. sieversi Ganglb 122

3. Genus Chelidonium Thoms.

1. C. Plav 128

4. Genus Leontium Thoms.

1. L. viride Thoms 134

5. Genus Polyzonus Cast.

1. P. fasciatus (Jt ) 140

21. Tribe Rosaliini
1. Genus Rosalia Serv.

1. JR. coelestis Sem 149

22. Tribe Callidiini

1 . Genus Hylotrupes Serv.

1. H. bajulus (L.) 160

xiii

2. Genus Rhopalopus Muls.

1. R. clavipes (F.) 166

2. R. signaticollis Sols 173

3. R. speciosus Plav 178

4. JR. aurantiicollis Plav 183

5. R. ruficoUis Mats. 1^4

3. Genus Pronocera Motsch.

1. P, brevicollis (Gehl.) 185

4. Genus Semanotus Muls.

1. S. undatus (L.) 192

2. S. bifasciatus (Motsch.). 199

5. Genus Oupyrrhidium Pic

1. O. cinnabarinum (Bless.) 204

6. Genus Callidium F.

1. C. violaceum (L.) 212

2. C. aeneum Deg 217

3. C. coriaceum Payk 222

4. C. chlorizans (Sols.) 225

7. Genus Phy mat odes Muls.

1. P. testaceus (L.) 236

2. P. zemlinae Plav. and Anufr 242

3. P. ussuricus Plav 245

4. P. vandykei Gress 249

5. P. abietinus Plav. and Lur : 254

6. P. mediofasciatus Pic 258

7. P. maaki (Kr.) 263

8. P. «/m(L.) 268

9. P. ermolenkoi Tsher. . . 273

V. Subfamily Cerambycinae

7 Adult insect characterized by elongate body, usually with parallel
sides, rather large (Cerambycini) or very small (Graciliini). Head narrows
gradually but insignificantly posterior to eyes, without sharp cervical con-
striction. Temples taper gently, do not project, and well distinguished in
this respect from adult insects of the subfamily Lepturinae. Frons broad
or narrow, with median longitudinal suture (Hesperophanini, Callidiini,
many Clytini, Stenaspini) or with longitudinal carinae (Xylotrechus)
dipping markedly or almost upright (some Clytini). Genae long, broad
(Clytini, Rosaliini, Callichromini) or extremely short (Hesperophanini).
Eyes large, with large (Hesperophanini, Cerambycini, Obriini) or minute
(Molorchini, Callidiini, Clytini), sometimes rather sharp or smoothened
facets, very intensely and deeply (Callidiini, Rosaliini, Callichromini,
Molorchini) or moderately (many Clytini, Obriini) emarginate, or almost
not emarginate but highly convex and reniform (Nathriini). Antennae
long, considerably (especially among males) longer than body (Ceram-
bycini, many Callichromini, Rosaliini, some Clytini) or short, with apices
not reaching (especially among females), or just reaching middle of
elytra (Dilusini, many Clytini) or their hind clivus.

Pronotum barely longer, or not longer, or even shorter than wide,
rounded on sides (Hesperophanini, Callidiini, Clytini), more rarely ex-
tended tubercularly, or even spinelike (Callichromini and some Stenas-
pini), or with conical tubercle dorsally receding inward from lateral
margin (Rosaliini), broadly stretched anteriorly and posteriorly (Obriini
and Molorchini), or with distinct recurved margins (Cerambycini), narrow
transverse groove posteriorly with uniform, deep (many Clytini, Stenas-
pini) or rugose (some Clytini) punctation; in some (Callidiini, Molorchini)
with smooth prominent shields or with large transverse sinuous grooves
on disk (Cerambycini— genus Mallamhyx).

Elytra markedly (Dilusini, Clytini— genus Rhaphuma) or moderately
(Clytini — genus Xylotrechus) elongate, or considerably shortened (Molor-
chini, Nathriini), individually or jointly rounded apically (Hesperopha-
nini, Callichromini, Rosaliini, Callidiini, Obriini), obtuse or truncate
(many Clytini), monochromatic, with dark brown or black (Ceramby-
cini, some Callidiini), metallic blue or bronze (some Callichromini, Calli-
diini), or rusty or rusty-brown (Obriini and Graciliini) tone, often with
pattern or spots and transverse bands (Clytini, Rosaliini, some Callidiini,

Callichromini), more rarely red with black pattern on suture (Stena-
spini).

Legs moderately or very long. Forecoxae elongate, contiguous (Molor-
8 chini and Nathriini), or not elongate and not contiguous but wide set
(Dilusini, Callichromini). Femora thicken gradually toward apex, not
clavate (Cerambycini, Callichromini, Clytini, Stenaspini), or thicken
abruptly and distinctly clavate (Callidiopini, Obriini, Nathriini, Molor-
chini, Callidiini). Hind femora short, extend only beyond middle of
elytra (Dilusini), or comparatively long and reach or just reach elytra!
apex (many Clytini, Callidiini, Rosaliini, Callichromini) or even, espe-
cially among males, extend beyond it (some Clytini, Callichromini).

Metasternum with fine dense punctation and median longitudinal
groove, with aromatic pores (Callichromini, Rosaliini, Stenaspini) or
without them (Molorchini, Clytini). Mes- and metepisterna with sparse
hairs, almost glabrous (Callidiopini, Callichromini, Obriini) or with very
dense discontinuous or continuous (Rosaliini, many Clytini) hairy cover.
Abdomen comparatively elongate; abdominal sternites convex, with fine,
rather dense or sometimes sparse punctation, with sparse adherent or
semierect setaceous hairs, or sometimes very dense tomentose to dense
hairy cover (Rosaliini, some Clytini).

Egg: Usually white, rarely greenish (Plagionotus floralis Pall.), or
yellowish [Polyzonus fasciatus (F.)], highly (Dilusini), moderately (Cly-
tini, Callidiini), or slightly elongate or oval (Stenaspini). Chorion smooth,
lustrous, semihyaline (majority of Clytini, Callidiini), more rarely with
fine reticulate sculpture and matte (Dilusini).

Larva: Unlike larvae of other subfamilies, readily recognized by
structure of maxillae, epistoma. and hypostoma. Half or more of head
retracted into prothorax. Anterior margin of epistoma with brownish or
rusty-brown border, lustrous or with transverse narrow grooves here; fuses
laterally with parietals or slightly demarcated from them by faint frontal
sutures; posterior to transverse brownish border stray piliform setae
form transverse row, or innumerable setae form transverse band. Median
epistomal suture distinct or, more often (Clytini), discernible only at
apex, or not visible. Hypostoma divided medially by gula into two trian-
gular sclerites, its anterior margin smooth or with minute transverse
grooves (Clytini, Callidiini) or deep longitudinal ones (Stenaspini, some
Clytini). Anterior half of parietals with short or long hairs (piliform
setae), anterior margin with rusty-brown or brownish border partially
or entirely covering ocellar-antennal area. One (Clytini, Obriini, Calli-
chromini, Stenaspini), two (Dilusini), or three (Rosaliini, Cerambycini,
some Callidiini — Rhopalopus) ocelli located around each antennal base.
Clypeus small, trapezoid, in some species flattened basally, almost in form
of strip. Labrum very small, convex, overlies only mandibular joint, with

short light-colored setae in anterior half or along margins. Mandibles
short, convex outside, concave inside (as though hollowed), broadly
rounded apically, apposed with cultrate edge end to end. Labial sub-
mentum broad, usually tetragonal, with two troughlike grooves, rarely
(genus Teratoclytus) hexagonal, longitudinally striate in posterior half.

Pronotum moderately or insignificantly slopes toward head, in ante-
rior third with transverse yellow or rusty band usually separated medially
and laterally by light-colored gaps into four spots, of which two median
9 on anterior margin (among many Clytini) with white alveolate notch.
Sides and disk of pronotum anterior to scutum with dense or sparse
rusty or light rusty hairs. Pronotal scutum coriaceous, white, squarrose,
coarsely rugose or rugulose (Callichromini), or lustrous, with fine longi-
tudinal, sometimes very dense streaks, matte silvery only at base, with-
out streaks (genus Phymatodes), or yellowish and sclerotized, with
minute spinules (some species of the genus Xylotrechus), bound laterally
by short deep longitudinal grooves. Prothoracic presternum laterally with
long, on disk short or uniform hairs. Eusternum glabrous, lustrous,
sometimes notably rugose, in anterior half with hairy gap or without it,
usually without lateral grooves, merging there with general surface of
presternum, rarely demarcated from it by shallow grooves. Thoracic legs
comparatively long, distinctly (Callichromini, Rosaliini) or poorly (some
genera of Clytini) developed or lacking iXylotrechus, some species of
Clytini).

Abdomen moderately elongate, usually narrows insignificantly toward
posterior end, with dense or sparse rusty or light-colored hairs along
sides. Locomotory ampullae developed on abdominal segments I to VII,
compratively uniform, identically convex, only in some (Obriini, Nathriini)
protrude notably on abdominal segments III to VI in form of knobs or
lumpy bulges, usually coriaceous, rarely (some species of Xylotrechus)
sclerotized, with barely visible spinules, even more rarely (Dilusini) with
granulations in two transverse rows. Abdominal tergite IX longitudinal or
transverse, rounded posteriorly, without urogomphi, and without spinules
here; innumerable spinules on disk form an extensive field only in one
genus (Nadezhdiana). Anal pore triradial. Interstadial variations among
larvae of different tribes or even genera manifested variously. Among I-
instar larvae, head and prothorax relatively large, some species with long
sharp spinules laterally along abdomen. For example, in larvae of
moschata (L.) and Leontium viride Thoms, such spinules seen laterally on
segments III to V and in Rhopalopus clavipes (F.) and Phymatodes ermo-
lenkoi Tsher.* on VI and VII. These spinules disappear after molt.

*Spelling of author names in taxonomic divisions sometimes differs from spelling
in text and bibliography since the Israeli orthography has been followed in this
translation — General Editor.

Pupa: Characterized by comparatively convex pronotum, arrangement
of spinules on abdominal tergites, and other features. Head moderately
bent under, eyes shifted toward base of jaws (Hesperophanini) or nota-
bly removed from jaws. Antennae long, directed forward and looplike
(Callichromini) or arcuate (some Clytini, Rosaliini), or short and pressed
to sides, with apices directed backward or bent ventrad (Clytini).

Pronotum convex, distinctly longitudinal, with flanges on anterior
and posterior margins (Obriini), or slightly elongate, sometimes even
somewhat transverse, without anterior flange, posteriorly with narrow
transverse groove (Clytini, Stenaspini), rounded laterally (Hesperopha-
nini, Clytini), or with conical lateral tubercle (Callichromini). Mesono-
tum bulges forward, posterior to middle with transverse, saddle-shaped
dent, with angularly extended or rounded scutellum on posterior margin.
Metanotum insignificantly convex, medially with or without longitudinal
troughlike groove, sometimes flattened laterally, with minute dent, usually
broadly rounded at posterior margin.

Abdomen moderately or markedly elongate, usually tapers slightly
toward anterior end and notably toward posterior end. Abdominal tergi-
tes convex, with common median longitudinal groove, with minute spinu-
10 les forming distinct transverse row or band in posterior half. Abdominal
tergite VII more elongate, triangular (Obriini, Callichromini, Clytini) or
tetragonal, transversely truncate along posterior margin, upright (Stenas-
pini), with large spinules in posterior half close to posterior margin bent
forward and forming posterior transverse row, with two pairs of large in-
curved spinules in front of this row forming two medial transverse rows,
and minute spinules in anterior half forming anterior transverse row; in
some (Moiorchini) anterior row of spinules lacking, and only posterior
row comprising two to four curved spinules directed forward present;
more often (Clytini) posterior row with four to six, rarely (Stenaspini)
eight to 12 spinules; sometimes (Callichromini) spinules on tergite VII
smaller than those on anterior tergites. Abdominal tergite VIII short,
with four to six small erect spinules on posterior margin, rarely without
them. Tip of abdomen (ventral view) obtuse, without lateral spinules.
Valvifers of female large, rarely minute, hemispherical, with small apical
sometimes conical, tubercle. Hind femora pressed to abdomen laterally,
long, slender (Callichromini), or distinctly clavate (Moiorchini, some
Clytini, Callidiini).

Biology: Most species of the subfamily Cerambycinae have a two-
year life cycle, with development from egg to adult requiring one or
three years in only a few species. Adult insects of the genera Chloropho-
rus, Clyfus, Leonlium, MoJorchus, Obrium, Polyzonus, and others feed on
pollen for gonadal maturation and then begin to reproduce. Adult in-
sects of many species of Xylotrechus, Rhopalopus, Plagionotus, and other

genera do not require supplementary feeding; beetles emerge with devel-
oped gonads, mate immediately, and oviposit. In some species (some
Clytini, Callidiini) beetles commence flight in May and stop in My,
while in others (Callichromini, many Clytini, Stenaspini) flight occurs
from early July through August or even the end of September
fasciatus (F.)]

The fertility of beetles varies widely. For example, among Obriini
(Obrium gracile Plav., Stenhomalus vulcanus Tsher.) a single female lays
1 8 to 20 eggs during her lifetime, while some Clytini [Xylotrechus altai-
cus (Gebl.), X. rusticus (L.)] lay 30 to 108 eggs. Eggs are laid on woody
and shrub species and only a few beetles {Xylotrechus arnoldi Kost.
and Plagionotus floralis (Pall.)] oviposit on grass, Compositae, Legumi-
nosae (Medicago), and other vegetation. Most beetles (Callidiini, Clytini)
lay eggs singly in bark crevices, spacing them out. Rarely (Obriini,
Stenaspini) the female initially glues the egg to the bark and covers it
later with small scales scraped earlier from the bark surface by means
of a special brush on the ovipositor (Stenaspini) or on the posterior mar-
gin of abdominal sternite II (Obriini). Some species [Polyzonus fasciatus
(F-)] glue the egg to the bark and cover the top with secretions that
harden on exposure to air and form a stable shell. Larvae hatch in the same
summer eggs are laid. Embryonic development is completed in nature in
two to four weeks in various species. Newly hatched larvae bore shoots
and make galleries under bark or in wood, plugging them compactly
with frass. Larvae of some genera {Polyzonus, Chelidonium, Purpurice-
rms, Amarysius) nibble ventilation holes in the shoot surface through
which frass is discarded. As a result, the galleries are cleared of frass
and the larvae able to move rather rapidly from one end to the other.

11 The larval period varies in different species. Some live no more than
one year and some two to three years. Mature larvae make cells and
pupate. Based on the prepupal period and pupal development, the sub-
family Cerambycinae can be divided into four groups. The first, more
abundant group comprises species (many Clytini, Callidiini, Stenaspini)
whose larvae pupate at the end of May and in June. Beetles emerge from
pupae in the last 10 days of June and in July. The second group includes
Molorchus minor (L.),,M. kobotokensis Ohb., Semanotus undatus {\^.), Dei-
lusfugax (Oliv.), Aglaophis colobotheoides Bat., and others whose larvae
pupate in the latter half of July and in early August; young bettles emerge
from pupae in the latter half of August and in early September. The third,
very small group comprises species [Clytus arietoides Reitt., C. arietis (L.)]
in which pupation occurs at the end of July and in August; the pupae
enter diapause, hibernate, and beetles emerge after hibernation. The fourth
group comprises species (Leontium viride Thoms.) whose larvae make
pupal cells at the end of August, enter diapause, hibernate in that state.

later molt, and pupate the next spring when the weather warms up.

In some species (Callichromini, Obriini, Callidiini, most Clytini,
Stenaspini) the pupal stage continues for two to four weeks and beetles
emerge the same summer; in others {Clytus arietoides Reitt., C. arietis (L.),
and possibly some others] pupae diapause for a long time and adults
appear after nine to ten months of pupation. In many species (Obriini,
Callichromini, Callidiini, Clytini, Stenaspini) emerging adults abandon
the pupal cell the same summer and soon commence reproduction. Only
in some species (Deilus, some Molorchus, Semanotus) do the beetles'
emerging from pupae enter diapause and remain in hibernation, usually
in pupal cells, abandoning the latter only in the following spring.

Most species (Callidiini, Clytini) develop on physiologically weak-
ened, drying, and newly fallen trees; rarely [Rosalia coelestis Sem., Rha-
phuma acutivittis (Kr.)] development occurs on desiccated trees. A goodly
number of species [Chelidonium zaitzevi Plav., Aromia moschata (L.),
Xylotrechus altaicus (Gebl.)] inhabit growing trees. These species are
generally monophagous in the larval stage. For example, Xylotrechus
altaicus (Gebl.) develops only on larch {Larix), Chelidonium zaitzevi
Plav. on maple (Acer), Aromia moschata (L.) on willow (Salix), and
Chloridolum sieversi Ganglb, on walnut (Juglans), Species of Ceramby-
cinae, unlike those of Lepturinae, usually do not inhabit dead fallen
trees with decomposing wood.

Long-horned beetles of the subfamily Cerambycinae are ecologically
associated mainly with deciduous trees; only an insignificant number
colonize coniferous vegetation. During forest inspections we collected
over 6,000 species of this subfamily from various trees, shrubs, semi-
shrubs, and herbaceous plants. A large number of beetles were raised
in the laboratory from larvae collected in nature. It was established that,
of the 105 species (Table 1) inhabiting northern Asia, 14 habitually
develop on coniferous species, 83 on deciduous, and two on herbaceous.
Chlorophorus gracilipes (Paid.) usually inhabits deciduous trees, but
sometimes migrates to coniferous; Asias tuvensis Tsher. inhabits semi-
arid steppes and develops on semishrubs (Nanophiton erinaceum). These
facts indicate that Cerambycinae fauna originated mainly on deciduous
vegetation, with only a few species adapting later in the course of
evolution to coniferous and vary rarely to herbaceous plants.

12 The cerambycid fauna of broad-leaved forests revealed a series of
species forming a complex of ecological concomitants characterized by
monotypic ecological characteristics, discontinuity of geographic range,
and common phylogenesis. For example, south of the Urals broad-leaved
forests are inhabited by Phymatodes alni (L.), Xylotrechus antilope
(Schonh.), X. arvicola (Oliv.), Chlorophorus herbsti (Brahm), C. varius
(Mull.), Plagionotus detritus (L.), P, arcuatus (L.), and other species

Table 1. Dlstribatioa of Cerambycioae larvae based oo host plants

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leterminei

ecologically associated to some extent or the other with oak. Their con-
comitants living in broad-leaved forests of Ussuri-Primor’e region are
Phymatodes ermolenkoiTshcx., Xylotrechus cuneipennisKi., Chlorophorus
sexmaculatus (Motsch.), Plagionotiis christophi Kr., P. pulcher Bless.,
and others. Some inhabit trunks and others crowns of thin shoots. This
tendency for occupying niches has been preserved in several recently
formed species (within the genus) even with a change in host plants.
For example, in northern Asian fauna the genus Phymatodes comprises
nine species, of which five develop on grapevine (Vitis), three on oak
(Quercus), and one on Siberian fir {Abies sibiricus). But all of the sespe-
cies generally inhabit crowns of thin shoots. Such a conservatively topo-
logical (levelwise) distribution is observed even in other beetles, reflect-
ing the general system of ecological concomitants inhabiting different
regions, as can be seen from the following scheme:

f X. antilope (Schonh.) — mainly oak trunks.

I Southern Urals, Europe.

J X. altaicus (Gebl.) — larch trunks. .

^ Siberia.

I X. cuneipennis Kr. — oak trunks.

L Ussuri-Primor’e region.

f P. detritus (L.) — oak trunks.

J Southern Urals, Europe.

1 P. pulcher Bless. — oak trunks.

[_ Ussuri-Primor’e region

f P. alni (L.) — thin shoots of oak.

Southern Urals, Europe.

P. ermolenkoi Tsher. — thin shoots of oak.
Ussuri-Primor’e region.

P. abietinus andLur. — thin shoots of Siberian

fir. Salair range.

P. ussuricus Plav. — thin shoots of Amur vine.
Ussuri-Primor’e region.

f P. kaehleri (L.) — oak shoots in crown zone.

I Southern Urals, Europe.

J P. tsherepanovae Tsher.

I Ob’ region.

I P. petasifer Fairm. — shoots of fruit trees in crown
L zone. Eastern Asia.

The fact that ecological concomitants inhabit widely separated regions
in nature indicates that many ecological groups of long-horned beetles
in forest formations evolved in the Tertiary period in broad-leaved forests
and have preserved their form to date. Their evolution proceeded gene-
rally along the course of renewal (changeover) of species composition.

1. Xylotrechus

2. Plagionotus

3. Phymatodes

4. Purpuricenus

This could explain the fact that most species of Phymatodes, Xylotrechus,
Plagionotus, and others are trophically associated in the larval stage with
deciduous trees, and only some [Xylotrechus altaicus (Gebl.) and P.
abietinus Plav. and Lur.] have adapted to growth on conifers.

Economic importance: Long-horned beetles of the subfamily Ceram-
bycinae constitute a vital link in forest biocenoses. Some of them (Obriini,
Molorchini, Chelidonium zaitzevi Plav., Phymatodes ermolenkoi Tsher.)
colonize thin shoots of growing trees, weaken them physiologically, and
affect their resistance to secondary pests and fungal diseases. Others
[Aromia moschata (L.), Chloridoliim sieversi Ganglb., Semanotus undatus
(L.), Xylotrechus altaicus (Gebl.), X. adspersus (Gebl.), X. pantherinus
(Sav.), Plagionotus christophi Kr., P. pulcher Bless., P. detritus (L.)]
colonize trunks of physiologically weakened or healthy trees, damage
the bast, and transport fungal spores which cause wood damage. Such
trees consequently die. In recent years shoots of goat willow in strip
forests on river banks in southern Tuva have been severely damaged by
Xylotrechus pantherinus (Sav.), maple plantations by Chelidonium zaitzevi
Plav., Manchurian walnut stocks near Partizansk in the Far East by
Chloridolum sieversi Ganglb., and spruce plantations in the Salair foot-
hills by Semanotus undatus (L.) and Pronocera brevicorne (Gebl.).

In the 1930’s deciduous plantations in many regions of Trans-
Baikal (Undinsk, Baleisk, Aleksandrov-Zavodsk, Gazimuro-Zavodsk, and
others) were severely attacked by the Altai long-horned beetle {Xylo-
trechus altaicus (Gebl.)]. Wood damaged by larvae is totally useless for
commercial purposes and hence preparing marketable timber is fraught
with difficulties. In the southern Urals oak forests are greatly damaged
by Rhopalopus clavipes {¥.), Purpuricenus kaehleri (L.), Plagionotus detri-
tus (L.) and others. Beetles damaging Amur vine in the Far East fall in-
to a special group comprising species of the genus {Phymatodes, Bra-
chyclytus singularis Kr., Teratoclytus plavilstshikovi Zaitz.,* and others.
From 1971 to 1973 these pests destroyed 30% or more of the vine in
some forests (Cherepanov and Cherepanova, 1974).

Long-horned beetles cause great losses in forest lumber. Plagionotus
14 pulcher Bless., P. arcuatus (L.), P. detritus (L.), Chlorophorus sexmacu-
latus (Motsch.), Xylotrechus rusticus (L.), X, cuneipennis Kr., and others
inhabit freshly prepared deciduous species of wood. Clytus arietoides
Reitt., more rarely Semanotus undatus (L.) and other species often infest
logs of conifers. Forests littered with wind-fallen trees or damaged by
fire serve as sites of en masse proliferation for many species of beetles.

^These species should be placed under strict surveillance by the quarantine
services. Their accidental import into Central Asia, the Caucasus, Crimea, and
Moldavia could result in devastating consequences for viniculture.

Overmature deciduous wood stocks serve as latent reproduction centers
for Xylotrechus altaicus (GebL). With the onset of favorable conditions,
this beetle infests extensive mature deciduous plantation. A significant
increase in reproduction centers of this pest was recorded in Trans-
Baikal from 1932 to 1946.

KEY TO TRIBES
Adult Insects

1 ( 2). Eyes notably proximate to mandibular base; genae very short,
their least length equal to 2.0 times diameter of ocular sclerite.
Hind femora thicken gradually toward apex, not clavate. Body

length 11 to 20 mm 12. Hesperophanini.

2(1). Eyes not proximate to mandibular base; genae long, at least
several times diameter of ocular sclerite. If genae short, hind
femora clavate.

3 ( 4). Anterior margin of pronotum with broad distinct flange, about

0.66 width at base; disk with large sinuous grooves [Mallam-

byx raddei (Bless, and Sols.)]. Body length 33 to 54 mm

13. Cerambycini.

4 ( 3). Anterior margin of pronotum without broad distinct flange, in

any case not narrower or only slightly narrower than base; disk
without large sinuous grooves, at most with coarse punctation
forming transverse wrinkles (some species of Xylotrechus).

5 ( 6). Pronotum markedly elongate, 0.25 times longer than maximum

width, with almost parallel sides, and broadly but indistinctly
rounded laterally. Hind femora clavate. Body length 8.0 to
13.5 mm 14. Callidlopini.

6 ( 5). Pronotum less elongate, rounded or with tubercle projecting

laterally.

7 ( 8). Body small. Elytra elongate, markedly flattened in anterior

half, with deep, not very dense punctation, matte; hind clivus
more convex, smooth, without punctation. Hind femora with
elongate clava. Body length 4.0 to 6.0 mm [Gracilia miuuta
(F.)] .15. Graciliini.

8 ( 7). Body very large. If small, elytra lustrous not matte, distinctly

shorter, and hind femora sharply clavate.

9 (10). Abdominal sternite I highly elongate, almost not shorter or

even longer than all sternites together 16. Obriini.

10 ( 9). Abdominal sternite I moderately elongate, shorter than remain-
ing sternites together.

11 (14). Elytra short, do not cover top of abdomen. Membranous wings

fall open on abdomen. Forecoxae elongate, cylindrical or coni-
cal, contiguous on inner side.

12 (13). Elytra cover most of abdominal segment I. Cavities of hind

coxae open from behind. Body length 3.0 to 6.0 mm

17. Nathriini (= Psebiini).

13 (12). Elytra much shorter, cover only base of abdominal segment I.

Cavities of hind coxae closed from behind. Body length 4.0 to

16.0 mm 18. Molorchini.

14 (11). Elytra long, cover top of abdomen. Membranous wings fold

on abdomen under elytra. Forecoxae not elongate, not conti-
guous on inner side.

15 (16). Legs very short; apices of hind femora barely extend beyond

0.66 of elytra. Elytral disk with longitudinal groove near suture
extending from base almost up to hind clivus. Body length

6.0 to 11.5 mm 19. Dilusini.

16 (15). Legs very long; hind femora reach hind clivus of elytra or ex-

tend beyond their apex. Elytral disk convex or flat.

17 (18). Pronotum laterally with sharp spinelike tubercle. Elytra with

metallic bronze, bluish or greenish iridescence, or with two

transverse yellow bands (genus Polyzonus)

20. Callichromini.

18 (17). Pronotum laterally without sharp spinelike tubercle; if tubercle

present, elytra not metallic toned (some species of Stenaspini),
or tubercle conical, extends upward, its edge dorsal (Rosaliini).

19 (20). Elytra with bright turquoise-blue and velvet-black dense hairy

cover that forms pattern of broad transverse bands. Antennal

segments apically with dense black brush of hairs

21. Rosaliini.

20 (19). Elytra with varied hairy cover or without it. Antennal segments

apically without dense brush of hairs.

21 (24). Metasternum without aromatic pores. Pronotum laterally with-

out tubercle, rounded, at most broadens angularly.

22 (23). Antennae more wide set; distance between them at base more

than gap between upper lobes of eyes on vertex. Femora cla-
vate 22. Callidiini.

23 (22). Antennae proximate; distance between them at base less than

gap between upper lobes of eyes on vertex. Femora not cla-
vate 23 Clytini.

24 (21). Metasternum with aromatm pores. Pronotum laterally with

minute or large sharp tubercle or without it. Elytra usually red
with black pattern on suture or without it, or black with red-
dish base 24. Stenaspini.

Larvae

1 ( 2). Posterior half of abdominal sternite VIII laterally with yellow-
ish oval spot obliquely disposed in depression surrounded by

stray hairs 12. Hesperophanini.

2(1). Posterior half of abdominal sternite VIII laterally without
yellowish spot.

3 ( 4). Body very large (length of mature larvae up to 65 mm and

width of head 8.0 mm). Dorsal locomotory ampullae separated
by two transverse grooves converging laterally, with deep later-
al rugose folds. Ventral locomotory ampullae coarsely rugose.
Thoracic legs developed. Three ocelli on each side of head. . . .
13. Cerambycini.

4 ( 3). Body less large or small, short (length of mature larvae not

more than 6.0 to 40.0 mm and width of head I.O to 5.0 mm).
Structure of dorsal and ventral locomotory ampullae different.
5(6). Sides of head (dorsal view) with minute round black spot re-
sembling an ocellus shifted far away from brownish margin

(long genus Gracilia Serv.) 15. Graciliini.

6(5). Sides of head without minute round black spot resembling an
ocellus, at most with one to three hyaline or pigmented ocelli
near antennal bases ventrally.

7 (10). Abdominal segments III to VI with highly protuberant loco-

motory ampullae in form of nodular or lumpy bulges.

8 ( 9). Parietals (sides of head) in anterior half with long dense hairs

forming tufts bent down and backward sharply. Prothoracic
eusternum coriaceous, without silvery reticulate sculpture. Hairs
on pronotal disk directed backward or upright. ... 16. Obriini.
9(8). Parietals (sides of head) in anterior half with sparse hairs that
do not form dense tuft. Prothoracic eusternum with minute
silvery reticulate sculpture. Hairs on pronotal disk anterior to
scutum directed forward 17. Nathriini (= Psebiini).

10 ( 7). Abdominal segments III to VI with ordinary locomotory am-

pullae, not in form of nodular or lumpy bulges.

11 (12). Parietals (sides of head) in anterior half with long dense seta-

ceous hairs bent backward (as though broken)

18. Molorchini.

12 (11). Parietals (sides of head) in anterior half with usual hairs, not

bent backward (as though not broken).

13 (14). Anterior third of pronotum with narrow transverse yellowish-

rust band extending laterally and interrupted medially by nar-
row indistinct gap. Locomotory ampullae separated by trans-
verse groove from which short grooves originate and form

granular prominences in two transverse rows. Two hyaline or
pigmented ocelli on each side of head. Thoracic legs well devel-
oped .19. Dilusini.

14 (13). Anterior third of pronotum with two yellow or rusty trans-

verse spots on disk interrupted medially by broad gap.

15 (16). Body of live larvae yellow. Dorsal locomotory ampullae sepa-

rated by two transverse grooves, with shallow grooves between
them in form of cells. One transparent ocellus on each side of
head. Thoracic legs long, well developed. ... 20. Callichromini.

16 (15). Body of live larvae white. Structure of dorsal locomotory am-

pullae different; if same in structure (Rosaliini), then three
ocelli on each side of head. Thoracic legs either developed or
lacking.

17 (18). Dorsal locomotory ampullae with two transverse grooves, with

fine grooves forming network. Three transparent ocelli on each
side of head. Thoracic legs well developed 21. Rosaliini.

18 (17). Structure of dorsal locomotory ampullae different, with grooves

not forming fine network or without grooves.

19 (22). Anterior margin of hypostoma smooth, without longitudinal

striation.

20 (21). Pronotal scutum lustrous, with fine longitudinal striation, matte

silvery only at base. One (Phymatodes and others) or three
(Rhopalopus) ocelli on each side of head. Thoracic legs devel-
oped. 22. Callidiini.

21 (20). Pronotal scutum sclerotized (many Xylotrechus) or coriaceous

(Clytus, Chlorophorus, and others), matte or lustrous, at most
with large longitudinal striations. One {Xylotrechus, Clytus,
Chlorophorus) or two {Epiclytus) ocelli on each side of head.
Thoracic legs developed (many Clytus, Chlorophorus) or lack-
ing {Xylotrechus, some Clytus, and others) 23. Clytini.

22 (19). Anterior margin of hypostoma not smooth, with distinct longi-

tudinal striation. One ocellus on each side of head. Thoracic
legs developed 24. Stenaspini.

Pupae

1 ( 2). Head short, eyes proximate to mandibular base. Posterior

third of abdominal tergite VII lustrous, glabrous, without spi-
nules; bulges notably in front, with prominent spinules on
protuberant coriaceous base. Antennae bent ventrad, semicir-
cular. Body length up to 14 mm 12. Hesperophanini.

2 ( 1). Head usually more elongate, eyes not proximate to mandibular

base. If short (Nathriini), pronotum glabrous, without setae.

Posterior third of abdominal tergite VII with spinules, only
sometimes (Obriini male) projects conically and without spi-
nules.

3 ( 6). Pronotum elongate, basally (sometimes at anterior margin) with

flange; long acicular setae anterior to base form indistinct
transverse row or two independent tufts. Posterior margin of
abdominal tergite VII conically produced in male, rounded in
female. Second half of antennae bent ventrad, in form of ring
(female) or loop (male). Hind femora clavate, with long shaft.

4 ( 5). Anterior margin of pronotum with setaceous spinules

15. Gradliini.

5 ( 4). Anterior margin of pronotum glabrous, without setaceous spi-

nules 16. Obriini.

6 ( 3). Pronotum barely elongate or transverse, without flange at ante-

rior margin, glabrous anterior to base, without setae (Nath-
riini), only sometimes with setae or spinules that do not form
transverse row from inner side of posterior angles. Posterior
margin of abdominal tergite VII not conically produced in
male; if produced (Nathriini), pronotum without setae or spi-
nules.

7 (10). Hind femora sharply clavate, markedly dilated apically, with

long shaft.

8(9). Abdomen with parallel sides, does not narrow toward base.

Pronotum without setae or spinules 17. Nathriini.

9(8). Abdomen not laterally parallel, narrows sharply toward base,
appears petiolate. Pronotum usually with setae or spinules. . .
18. Molorchini.

10 ( 7). Hind femora not clavate, broaden gradually from base toward

apex; if notably clavate (Callidiini), with short shaft.

11 (12). Spinules on pronotum and abdominal tergites bent down and

forward. Antennae bent ventrad, semicircular. Body length up
to 10 mm 19. Dilusini.

12 (11). Spinules on pronotum and abdominal tergites bent down and

mostly backward; if bent forward, body dimensions considerably
larger.

13 (16). Pronotum with produced large or small lateral tubercle. Abdo-

minal tergite VII with minute spinules, smaller than tergites in
front. Antennae bent ventrad, looplike. Hind femora long,
slender, not clavate.

18 14 (15). Lateral tubercle on pronotum large projects conically sideways.

20. Callichromini.

15 (14). Lateral tubercle on pronotum small, projects tubercularly up-
ward, slightly recessed from lateral margin 21. Rosaliini.

16 (13). Pronotum without lateral tubercle. Abdominal tergite VII at

least partly with large spinules; if with minute spinules, femora
clavate.

17 (18). Femora clavate. Abdominal tergite VII with minute spinules.

Antennae bent ventrad, looplike 22. Callidiini.

18 (17). Femora not clavate, thicken gradually toward apex. Abdomi-

nal tergite VII with large spinules on posterior margin bent
forward, forming transverse row.

19 (20). Abdominal tergite VII triangular, rounded posteriorly, poste-

rior margin not upright; spinules on disk form three or four
transverse rows; spinules of posterior row bent forward, of two
medial rows (each consisting of two spinules) bent inward to-
ward middle. Antennae short, pressed to sides, with apices
directed backward or bent ventrad, or long, bent ventrad or
arcuate and directed forward 23. Clytini.

20 (19). Abdominal tergite VII tetragonal, transversely truncate poste-

riorly, posterior margin upright, with large spinules there bent
forward (eight to 12 spinules each), forming distinct transverse
rows; minute spinules on disk in posterior half form tuft.

Antennal apices bent ventrad and directed forward

24. Stenaspini.

12. Tribe HESPEROPHANINI

Adult insect with elongate body, with parallel sides, rounded at
elytral apex and on sides of pronotum; antennae long, with numerous
hairs.

Larva characterized by poorly developed thoracic legs, and rather
convex, coriaceous (not sclerotized) locomotory ampullae on abdominal
segments I to VII.

Pupa recognized by presence of transverse wrinkles (striation) on
vertex and pronotal disk, and minute spinules on abdominal tergites I
to VI and large spinules on tergite VII.

The tribe Hesperophanini is abundantly represented in the warm
tropical zone. Represented in northern Asia by a single genus with a
lone species.

1. Genus 7>/c/?o/erM5 Woll.

Wollaston, 1854, Yns. Maderens, p. 427; Plavil’shchikov, 1940, Fauna
SSSR, 22, 2, 62-64.

Adult: Distinguished from adult insects of other genera of the tribe
Hesperophanini by poorly produced, rounded tubercles near articulate

antennal sockets, less spherical pronotum, absence of hairy band on 3rd
antennal segment, and spotty pubescence on elytra.

Larva: Head slightly compressed dorsoventrally, with three trans-
parent ocelli laterally near antennal bases forming transverse band.
Anterior third of pronotum with transverse rusty band in which innu-
merable whitish dots occur in posterior half.

19 Pupa: Head glabrous, without setae, transversely rugose on vertex.
Antennae arcuate. Pronotum laterally rounded, transversely rugose on
disk, with spinules here forming transverse band, and small tuft in
anterior half. Abdominal tergites I to VI with sharp minute spinules,
VII with large spinules, VIII with few fine setae.

Only one species known in the fauna of northern Asia, three in
China, and no more than five in Europe, including the Mediterranean
region.

Type species: Cerambyx cinereus Villers, 1789.

1. Trichoferus campestris (Paid )

Faldermann, 1825, Mem. Acad. St. Petersb., vol. 2, p. 435 {Hespero'
phanes); Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 69-71; Kostin, 1973,
Zhuki-dendrofagi Kazakhstana, p. 102.

Adult (Figure 1): Readily recognized by uneven, spotty hairy cover
on elytra. Head with dense punctation and tightly adherent hairs direct-
ed forward and inward posterior to eyes, and sideways from midline
between antennae. Frons bulges gently, tubercularly near antennal bases,
with deep median longitudinal suture. Genae short, barely visible. Eyes
highly convex, coarsely and sharply faceted, deeply emarginate. Anten-
nae comparatively long, with apices reaching posterior third of elytra
(female) or extending beyond 0.75 their length (male), with short tight-
ly adherent hairs and on inner side (basally) (especially on 1st to 6th
segments) with long erect setaceous hairs. Third antennal segment longer
than 4th, equal to 5th; 11th segment short, obtusely rounded apically
(female) or elongate, slender, pointed (male). Pronotum narrows abrupt-
ly anteriorly, gently posteriorly, broadens roundly in anterior third,
with dense large punctation, tightly adherent light-colored hairs direct-
ed toward middle of disk, sometimes with smooth wartlike tubercles, of
which two anterior to middle and one medial in posterior half. Scutel-
lum with dense gray hairs, broadly or narrowly gently rounded apically.
Elytra elongate, with parallel sides, convex, somewhat flattened on disk
along suture, rounded apically, with coarse punctation and tightly adhe-
rent gray hairs forming dense, more or less distinct spots, with stray
20 erect setaceous hairs. Hind femora markedly fall short of elytral apex;
hind tarsi shorter than tibiae, their 1st segment not longer than two
successive segments together. Abdominal sternite V broad and broadly

19 Figure 1. Trichoferus campestris (FM.).

rounded posteriorly (female) or comparatively elongate and narrowly
rounded posteriorly (male). Entire body, elytra, antennae, and legs
rusty-brown. Length 13 to 20 mm.

Egg: White, slightly elongate, narrows notably toward poles, nar-
rowly rounded at poles. Chorion finely sculptured, matte. Length 1.9
mm, width 0.6 mm.

Larva (Figure 2): Head highly retracted into prothorax. Anterior
margin of epistoma distinctly emarginate at level of clypeus. Hypostoma
slightly convex, almost flat. Gula broad between sclerites of hypostoma.
Parietals (sides of head) narrow anteriorly; three distinct transparent
ocelli near antennal bases form transverse band. Antennae long, with

Figure 2. Larva of Trichoferus campestris (Paid.),
a — ^head and pronotum; b— abdominal tergite IV
with dorsal locomotory ampulla.

four segments; 1st antennal segment thick, narrows markedly toward
apex, longer than almost all successive segments. Clypeus short, base
broadens markedly. Labrum convex, narrows sharply anteriorly, with
short lateral setae. Inner masticatory lobes of maxillae digitate, apically
with thin piliform setae. Maxillary palps long, project forward marked-
ly beyond apex of inner lobes. Mandibles massive, broadly rounded
apically; transverse basal ridge demarcated from rest of surface by nar-
row groove, with faint median longitudinal groove.

Prothorax barely longer than meso- and metathorax together. Pro-
notum 2.0 times wider than long, narrowly rounded anteriorly, with
21 rounded anterior margin, in anterior third with transverse rusty band
interrupted medially and laterally by broad white gap, with long hairs
forming two transverse fields, of which one on anterior margin of rusty
band (hairs here bound by sclerotized ringlet) and second before scutum
(hairs not bound). Posterior margin of transverse rusty band glabrous,
with whitish dots. Pronotal scutum bulges slightly, without hairs, demar-
cated laterally by longitudinal grooves. Prothoracic presternum with
uniformly dense rusty hairs; eusternum glabrous, lustrous. Thoracic legs
short, claws poorly developed, setaceous.

Abdomen narrows posteriorly, with fine light-colored hairs laterally.
Abdominal segments I to VI with lateral transverse flange in anterior
third. Dorsal locomotory ampullae moderately convex, coriaceous, ante-
rolaterally with oblique pitlike depression, with short transverse posterior
grooves originating from it. Ventral locomotory ampullae slightly con-
vex, smooth, coriaceous, laterally with pitlike depression demarcated by
faint transverse grooves. Abdominal sternite VIII laterally with dull spot
in posterior half that looks like oblique dent. Body length up to 25 mm,
width of head 3.0 mm.

Pupa (Figure 3): Body moderately elongate. Head narrows uniformly
posterior to eyes, glabrous, bulges transversely between antennae, with
longitudinal, transversely rugose band between upper lobes of eyes, short
longitudinal dent medially on anterior margin adjoining two sutures,
and laterally arcuate. Antennae pressed to sides, bend round midfemora,
with apices touching midtibiae.

Pronotum narrows gradually toward base, rounded anteriorly, slight-
ly convex on disk, with transverse grooves and short obtuse spinules
forming median (in rugose field) transverse band and small tuft in

Figure 3. Pupa of Trichoferus campestris (Paid.).

anterior third. Mesonotum smooth, bulges (rhomboidal) on posterior
margin in region of scutellum. Metanotum with median, longitudinal,
transversely slightly rugose band, with three inconspicuous setae later-
ally.

Abdomen narrows slightly toward base and more toward tip. Abdo-
minal tergite I short, with transverse rugose dent laterally, with incons-
picuous stray spinules. Abdominal tergites II to VI slightly convex, with
minute sharp spinules that usually form three transverse rows; posterior
row with four to eight paramedial spinules, middle row two spinules,
and anterior row three to five spinules. Abdominal tergite VII elongate,
rounded posteriorly, with large spinules on disk, falcate toward middle
of disk. Tergite VIII with barely discernible stray setae. Valvifers of
female large, adjacent, apically obtuse. Body length up to 20 mm, width
of abdomen 4.5 mm.

Material: Collected in Ussuri-Primor’e region. Adult insects six,
larvae three, pupa—one female.

Distribution: Tadzhikistan, Uzbekistan, southern Kazakhstan, Ussuri-
Primor’e region; northern Mongolia, northern China, North Korea.

Biology: Inhabits broad-leaved and mixed vegetation. Flight of bee-
tles from end of June to early August; insects sometimes fly to light.
Under laboratory conditions female laid eggs singly on birch shoots 2.5
to 3.0 cm in diameter under scaly remnants of bark, rarely two eggs
simultaneously. Newly hatched larvae bore bark and plug chorion with
frass. Larvae make galleries along shoot under bark and plug them with
frass. Galleries usually faintly, rarely significantly impressed on sapwood.
Due to larval activity much of bark, except outer layer, damaged. Width
22 of gallery made by mature larvae 5.0 to 12.0 mm. Mature larva raised
in laboratory weighed 171.5 mg. Pupae formed on April 29, 1974 from
larvae that had hatched in August, 1972, suggesting that one generation
of Trichoferus campestris (Paid.) lives for not less than two years. Accord-
ing to observations by Samoilov (1938), this species colonizes Micro-
meles alnifolia in nature.

13. Tribe CERAMBYCINI

Adult insect differs from those of other tribes of the subfamily in
large massive body. Pronotum narrows more anteriorly, with distinct
flange near anterior margin, transverse groove at base, convex on disk,
coarsely rugose (quite often grooves transverse, sinuous), and rounded
{Mallambyx) or with acute spine {Plocaederus, Cerambyx) laterally.
Elytra elongate, smooth {Mallambyx) or coarsely rugose, especially in
anterior half {Cerambyx).

Larva characterized by large body, inconspicuous hairy cover on
abdominal pleura, large spiracles laterally on abdominal segment I,
which are not smaller than on mesothorax.

The greatest diversity of the tribe Cerambycini is seen in Southeast
Asia. Comparatively large number of species known in Mediterranean
fauna. Represented in northern Asia by a single genus with a lone
species.

1 . Genus Mallambyx Bat.

Bates, 1873, Ann. Mag. Nat. Hist., 4, 12, 152; Plavil’shchikov, 1940,
Fauna SSSR, 22, 2, 79; Gressit, 1951, Longicorn Beetles of China, vol.
2, p. 135; Cherepanov and Cherepanova, 1973, Nov. imaloizv. vidyfauny
Sibiri, 6th ed., p. 52

Adult'. Body large. Antennae longer (male) or shorter (female) than
body, thin markedly toward apex. Pronotum transversely coarsely rugose,
rounded laterally, narrows more anteriorly, less posteriorly. Elytra elon-
gate, smooth, narrowly rounded apically and there, on side of suture,
with acutely produced small spinule.

Larva'. Body of mature larva massive, thick. Half of head retracted
into prothorax. Pronotal scutum, prothoracic basisternum, and locomo-
tory ampullae coriaceous, not sclerotized. Pronotum with sparse large
setaceous hairs forming two transverse bands in anterior third and be-
hind middle before scutum; bands interlinked laterally and along median
longitudinal suture.

Pupa'. Not known.

Only one species known in the fauna of northern Asia; inhabits
southeastern forests.

Type species'. N eocer ambyx raddei and Solsky, 1872 (=japo-

WCW5 Bates, 1873).

1 . Mallambyx raddei (Bless, and Sols.)

Blessig and Solsky, I SI 2, Moral. Soc. Entom. Ross., vol. 9, p. 170
(Neocerambyx)', =japonicus. Bates, 1873, Ann. Mag. Nat. Hist., 4, 12,
152; Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 80-82; Kupyanskaya,
1968, Fauna ekologiya nasekomykh D. Vostoka, p. 101; Kojima and
Hayashi, 1969, Insect Life in Japan, vol. 1, pp. 47-48; Cherepanov and
Cherepanova, 1973, Nov. i maloizv.vidy fauny Sibiri, 6th ed., pp. 52-53.

23 Adult (Figure 4): Well distinguished from other species of the sub-
family Cerambycinae by large body size, elongate smooth elytra, and
transversely rugose pronotum. Head narrower than pronotum, barely
elongate, with tightly adherent hairs directed forward, genae shorter
than wide, frons near antennal bases clavately convex, with narrow

Figure 4. Mallambyx raddei (Bless, and Sols.).

median suture extending into parietal area, and occiput with fine dense
punctation. Eyes large, broad, very deeply emarginate; upper lobes
minute, less than half size of lower lobes. Antennae considerably longer
(male) or slightly shorter (female) than body; 3rd segment 2.0 times
longer than 1st, slightly shorter than 4th and 5th together.

Pronotum narrows sharply in anterior half, slightly in posterior half,
with distinct flange near anterior margin, transverse groove at base,
and deep transverse furrows, tightly adherent grayish-yellow pubescence
directed forward in posterior half and backward in anterior half, and
sparse erect setae laterally. Scutellum flat, narrowly rounded apically,
with fine adherent hairs.

24 Elytra long, smooth, with parallel sides (female) or narrow slightly
toward apex, narrowly rounded apically, with sharp projecting spinule

on inner margin of suture, and minute tightly adherent grayish-yellow
hairs. Abdominal sternite V posteriorly narrowly emarginate (male) or
broadly rounded (female). Legs relatively long; femora with parallel
sides, flat; hind tarsi shorter than tibiae. Body, antennae, elytra, and
legs brownish, with rusty tone. Body length 35 to 52 mm.

Larva (Figure 5): Mature larvae characterized by large body, loco-
motory ampullae on abdominal segments I to VII, and poorly develop-
ed thoracic legs. Head markedly retracted into prothorax, almost 0.50
width of pro thorax. Epistoma divided by well-developed, median, longi-
tudinal, dark brown suture, in middle third with numerous short seta-
ceous hairs forming transverse band, weakly demarcated laterally, fron-
tal sutures inconspicuous, almost imperceptible. Hypostoma with stray
short hairs on inner anterior margin of sclerites, demarcated laterally
by sutures diverging forward. Gula trapezoid, flat or slightly convex.
Sides of head (parietals) narrowly rounded anteriorly, in anterior half
with sparse thick setaceous hairs. Antennae short, project slightly be-
yond anterior margin of parietals; 1st segment notably thicker than 2nd.
Clypeus lustrous, white, trapezoid or almost rectangular, 2.0 times wider
than long. Labrum convex, narrows angularly toward front, with nar-
rowly rounded anterior margin, dense setae laterally and anteriorly, gla-
brous and smooth at base and in middle. Mandibles massive, rounded
apically, with extended cultrate edge, smooth on outer side. Inner lobes
of maxillae and labial ligula with dense setae. Maxillary palps with three
segments and labial palps two segments, short.

Figure 5. Larva of Mallambyx raddei (Bless, and Sols.).
a~“head and pronotum; b— abdominal tergite IV with
dorsal locomotory ampulla.

Prothorax notably wider than mesothorax. Pronotum slopes toward
head, with short setaceous hairs in anterior half and laterally. Pronotal
scutum convex, demarcated laterally by straight longitudinal groove,
with thin longitudinal furrows, sometimes with minute sparse hairs later-
ally near posterior angles. Prothoracic presternum in young larvae with
dense thin, in mature larvae with sparse thick setaceous hairs bound
25 basally by sclerotized rusty ringlet; eusternum bulges insignificantly,
coriaceous, broadly rounded apically, with sparse setaceous hairs later-
ally; base of prosternum (basisternum*) coriaceous, rugulose. Thoracic
legs short, poorly developed, with short pointed claw.

Abdomen thick, narrows insignificantly toward tip, with sparse
short thin hairs laterally. Dorsal locomotory ampullae slightly convex,
coriaceous, with fairly developed furrows (sometimes forming reticulate
granulation, especially in mature larvae) separated by median longitu-
dinal groove and two transverse grooves joined laterally, with lateral
longitudinal groove diverging posteriorly. Ventral locomotory ampullae
in young larvae slightly rugose, in mature larvae distinctly granular,
demarcated by transverse groove adjoining lateral longitudinal groove,
which in mature larvae looks like deep dent with furrows diverging
from it.

Body white. Anterior margin of head dark brown, almost black,
with rusty tinge. Mandibles black. Anterior third of pronotum with
yellowish-rust band interrupted medially and laterally by white gap,
sometimes with whitish notch on anterior margin close to lateral gap.
Body length of mature larvae up to 65 mm, width of head up to 8.0 mm.

Pupa: Not known.

Material: Collected in Ussuri-Primor’e region. Adult insects four,
larvae seven.

Distribution: Ussuri-Primor’e region; northern China, Korea, Japan.

Biology: Inhabits broad-leaved forests. Flight of beetles end of July
and in August; insects sometimes fly to light at night. Lives in trunks of
stunted oak trees up to 52 cm in diameter (Kupyanskaya, 1968). Larvae
live initially under bark, then in wood, usually make longitudinal gal-
leries in latter. Mature larva makes pupal cell along trunk (Kojima and
Hayashi, 1969) and pupates in it. Pupae evidently appear end of June
and in July. Beetles mainly emerge in July.

14. Tribe CALLIDIOPINI

Adult insect characterized by elongate body with parallel sides.
Head not wider or narrower than pronotum. Eyes highly emarginate.

*Strictly speaking, the sternellum — General Editor.

Antennae do not reach (female) beyond elytral apex. Pronotum elongate,
slightly oval. Femora clavate. Outer side of tibiae with sharp edge.

This tribe consists of a lone genus in the fauna of northern Asia.

1 . Genus Stenygrinum Bat.

Bates, 1873, Ann. Mag. Nat. Hist., 4, 17, 154; Plavil’shchikov,
Fauna SSSR, 22, 2, 120; Gressit, 1951, Longicorn Beetles of China, vol.
2, p. 158.

Adult Characterized by the following features in addition to those
listed above. Apices of maxillary palps markedly broad, dolabriform.
Third antennal segment shorter than 5th, equal to 1st, distinctly longer
than 4th. Pronotum slightly rounded laterally, and slightly narrower or
not narrower anteriorly than at base. Femora apically sharply convex,
with thin shaft at base.

This genus consists of a lone species, widely distributed in Southeast
Asia.

Type species: Stenygrinum quadrinotaturn Bates, 1873.

26 1 . Stenygrinum quadrinotaturn Bat.

Bates, 1873, Ann. Mag. Nat. Hist., 41, 17, 154; PlaviPshchikov, 1940,
Fauna SSSR, 22, 2, 121-122; Kojima and Hayashi, 1969, Insect Life in
Japan, vol. 1, p. 54.

Adult (Figure 6): Body elongate. Head not wider than pronotum,
appears elongate posterior to eyes. Frons with large deep punctation,
with narrow median longitudinal suture, with sharp tubercle lateral to
antennal bases protruding upward, spinelike. Vertex with uneven, some-
times large deep, sometimes smooth punctation. Occiput uniformly
rounded, with transversely rugulose punctation. Eyes moderately con-
vex, coarsely faceted, broadly emarginate on inner side, bordered here
with dense golden hairs. Antennae narrow toward apex, reach elytral
apex or notably shorter than elytra. First antennal segment with fine
punctation, laterally smooth at apex, lustrous; 3rd segment almost equal
to or slightly longer than, 5th, 1.5 times longer than 4th. Pronotum

27 almost 2.0 times longer than width at base, broadens insignificantly and
ovally anterior to middle or in middle, uniformly convex on disk, with
sometimes fine deep, sometimes smoothened rugose punctation, medi-
ally with prominent smooth longitudinal line. Scutellum very small,
narrowly rounded posteriorly, with minute inconspicuous hairs. Elytra
4.0 times longer than wide, with gabled suture, posterior to scutellum
with broad longitudinal dent, jointly rounded apically, with almost
straight, slightly rounded inner angle, and lustrous; deep punctation in
anterior half, especially at base, coarse punctation in posterior half, fine

Figure 6. Stenygrinum quadrinotatum Bat.

punctation otherwise, especially at apex, with delicate light-colored
pubescence and sparse stray erect setae. Femora slender at base, clava-
tely convex at apex. Tarsi short, with distinctly broadened segments; 1st
segment of hind tarsi not longer than two successive segments. Body
chestnut-reddish-rust, pronotum usually red, antennae and legs light
rust or reddish-rust. Elytra on disk dark brown, at base (narrow band)
and apex (in last third) light reddish or light rust with two light yellow-

ish spots; one spot anterior to middle and second posterior to it (f.
typica), sometimes spots fuse longitudinally (ab. conjunctum Matsusch.),
or only posterior spot present, anterior one lacking (ab. binotatum Plav.).
Body length 8.0 to 13.5 mm.

Distribution: All of Southeast Asia, including southern part of Ussuri-
Primor’e region, Korea, China, Indochina, and Japan. Described from
specimens from Korea and Japan (collection of Moscow State Univer-
sity). We did not find it»

15. Tribe GRACILIINI

Adult insect characterized by small body, elongate elytra with paral-
lel sides, comparatively long slender antennae, and distinctly emarginate
eyes.

Larva with white, moderately elongate body, short thoracic legs,
anterior half of pronotal scutum with sharp longitudinal striation, and
locomotory ampullae developed on abdominal segments I to VII.

Three genera of this tribe reported in USSR fauna. Of them, only
one species of the genus Gracilia Serv. found in the southern Urals. Not
reported elsewhere in northern Asia.

1 . Genus Gracilia Serv.

Serville, 1834, Ann. Soc. Entom. France, vol. l,p. 81; Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 125; Gressit, 1951, Longicorn Beetles of China,
vol. 2, p. 160; Mamaev and Danilevskii, \91 5, Lichinki zhukov-drovosekov,

pp. 181-182.

Adult: Body comparatively narrow, slightly flattened. Head short,
with short adherent hairs and stray long setae. Genae short. Antennae
comparatively widely separated, antennal sockets in ocular notch; dis-
tance between antennal bases more than gap between upper lobes of
eyes. Pronotum elongate, flattened dorsally, angularly broadened later-
ally. Elytra with parallel sides, matte, with short pubescence, sharply
rounded apically, matte. Femoral clava highly flattened from sides.

Larva: Head markedly retracted into prothorax, narrowly rounded
anteriorly, with distinct minute black spot laterally. Prosternum with re-
ticulate sculpture.

Pupa: Abdominal tergites with fine setae forming transverse row.

28 This genus consists of a lone species inhabiting Europe, western Asia,
northern Africa, North America, and Japan.

Type species: Callidium pygmaeum Fabricius, 1 834 [=Callidium minuta
(F.)].

1. Gracilia minuta (F.)

Fabricius, 1781, Spec, Insect, voL 1, 235 (Saperda); Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 127; Duffy, 1952, Monograph Beetles, pp.
194-195; Linsley, 1962, Cerambycidae of North America, 20, 3, 49-50;
Demelt, 1966, Tierwelt Deutschlands, vol. 2, pp. 52-64; Kojima and
Hayashi, 1969, Insect Life in Japan, vol. 1, pp. 56-57.

Adult (Figure 7): Body small, elongate. Head short. Frons broad,
with minute punctation, very sparse light-colored hairs, median longi-
tudinal suture, and convex around antennal bases. Eyes coarsely faceted,
very deeply notched, with narrow gap between lobes. Antennae widely

Figure 7. Gracilia minuta (F.).

separated at base, gap between them more than between upper lobes of
eyes, longer than body. First antennal segment shorter than 5th, barely
longer or not longer than 4th.

Pronotum slightly elongate, angularly broadened mediolaterally,
somewhat narrower toward base and less anteriorly, with less distinct
flange on anterior margin, more distinct one on posterior margin, flat-
tened on disk, with fine dense punctation and highly minute light-
colored pubescence not forming dense cover. Scutellum elongate, with
parallel sides, broadly rounded apically, medially with broad groove.

Elytra elongate, with parallel sides, individually rounded apically,
with longitudinal, not very long dent on suture behind scutellum, broad
depression on anterior third of disk, with large punctation in anterior
half and indistinct punctation in posterior half (especially on hind clivus),
with minute light-colored adherent hairs directed laterally from suture.
Femora clavate, markedly flattened on sides. Hind tarsi 0.66 length of
tibiae; 1st segment of hind tarsi distinctly longer than two successive
together. Body, antennae, elytra, and legs monochromatic, rusty. Body
length 4.0 to 6.0 mm.

Larva (Figure 8): Characterized by small body, presence of minute
29 black spot on each side of head, and longitudinal striation on pronotal
scutum. Body white, moderately elongate. Head narrowly rounded an-
teriorly, with broadly rounded anterior margin, highly retracted into
prothorax. Epistoma bulges slightly, lustrous. Hypostoma narrows in-
significantly toward front. Parietals with broad rusty-brown border on
anterior margin that does not encompass antennal sockets from behind,
in anterior half with short setaceous light-colored hairs, on sides (dorsal
view), behind rusty border, with sharply projecting small black spot
resembling ocellus. Clypeus short, flattened, translucent. Labrum con-
vex, lustrous, with rounded anterior margin, short light-colored setae.

Figure 8. Larva of Gracilia mimta (F.)
a — head and pronotum; b — tip of abdomen, dorsal view.

Mandibles thick, reddish-brown, in second half almost black, broadly
rounded apically, with cultrate mesal surface.

Pronotum narrows anteriorly, with very minute (just visible under
high magnification) light-colored hairs anterior to scutum and on sides.
Pronotal scutum white, moderately convex, bound laterally by short
longitudinal grooves, in anterior half with longitudinal parallel streaks
forming fairly distinct striation. Prothoracic presternum bulges, with just
visible sparse light-colored hairs; eusternum glabrous, indistinctly bound
by shallow groove with reticulate sculpture. Thoracic legs short.

Abdomen moderately elongate, narrows insignificantly from thorax
toward tip, with very sparse minute light-colored hairs laterally. Abdo-
minal tergites transverse. Dorsal locomotory ampullae on tergites I to VII
insignificantly convex, coriaceous, with shagreen sculpture, separated
by common median longitudinal groove. Ventral locomotory ampullae
divided by transverse groove. Body length up to 6.0 mm.

Pupa: Body elongate. Head moderately bent under, with stray setae
near antennal bases. Antennae pressed to sides, bent ventrad in second
half. Pronotum elongate, gently rounded laterally, with short setaceous
spinules on anterior margin forming transverse band, setae anterior to
middle forming transverse row, and thin minute setae on hind clivus
forming one small tuft on each side. Posterior margin of mesonotum
with fully extended scutellum. Abdominal tergites with four to six setae
forming transverse row on posterior margin. Tip of abdomen rounded,
with long setae forming transverse row. Body length 3.5 to 7.0 mm
(Dulfy, 1953).

Material: Collected in the Caucasus. Adult insects (collection of the
Zoological Museum, Moscow State University) several specimens, larva
one.

30 Distribution: West from Atlantic Ocean coast east to the Urals, north
from Sweden and Finland south to northern Africa, Japan, North Ame-
rica. Reported from the southern Urals, but we did not find it there.

Biology: According to information available in literature (Plavil’sh-
chikov, 1940; Romadina, 1954; and others), ecologically associated with
deciduous species. Larvae live under bark and make longitudinal sinuous
galleries deeply imprinted in wood. Mature larva bores wood, makes cell
there, and pupates in it. Beetles emerge from pupae the same summer,
nibble an opening in shoot surface, and exit through it. One generation
per year. Gracilia minuta (F.) inhabits slender dried shoots of willow
(Salix), blackthorn {Prunus spinosa), hawthorn {Crataegus), dog rose
{Rosa), oak {Quercus), birch {Betula), and other deciduous species. Dam-
ages hoops of wine barrels. According to some investigators, imported
from Europe into North America and Japan with wine barrels.

16. Tribe OBRIINI

Adult insect characterized by elongate body, long slender antennae,
highly convex, coarsely faceted, and highly emarginate eyes. Pronotum
elongate, with laterally produced tubercle. Elytra with parallel sides,
notably convex on disk (Obrium) or flat {Stenhomalus). Femora clavate.
Ventral side of abdomen with dense setaceous brush performing function
of scraper during oviposition.

Larva characterized by slender elongate body. Half of head retract-
ed into prothorax; anterior half of parietals with long dense hairs on
sides, bent backward; epistoma with barely perceptible median longi-
tudinal suture merging laterally with parietals; frontal sutures not visi-
ble. Pronotal scutum coriaceous, not sclerotized, white. Thoracic legs
lacking, rudiment faintly visible just before pupation. Abdominal seg-
ments I, II, and VII with poorly developed locomotory ampullae; seg-
ments III to VI with highly convex, widely separated locomotory am-
pullae projecting nodularly; anterior and posterior to ampullae deep
transverse constrictions form secondary pseudoannuli, as a result of
which abdomen appears multiannulate.

Pupa with short head; antennae long, bent ringlike {Obrium) or loop-
like (Stenhomalus) in posterior half. Pronotum elongate, with more
(Obrium) or less (Stenhomalus) projecting tubercle laterally, setae or aci-
cular spinules in posterior half forming one or two transverse bands.
Abdomen elongate, narrows sharply posteriorly and notably toward ante-
rior end; abdominal tergites with minute spinules; tip of abdomen with
(Stenhomalus) or without (Obrium) urogomphi.

Two genera of the tribe Obriini found in the USSR, of which Obrium
Curt, inhabits almost the entire Holarctic while Stenhomalus White is
distributed mainly in Southeast Asia. Species of these genera are ecolo-
gically associated with deciduous vegetation. Inhabit trunks and branches
of trees with thin smooth bark. Characteristic behavior of female seen
during oviposition. First, by means of a very small brush on ventral
side of abdomen, she scrapes minute fibrous scales from bark, lays a
sticky egg (which adheres to shoot surface), then covers top of egg with
scales scraped earlier. A laid egg looks like a tiny mound and merges
with the general background of bark.

KEY TO GENERA

Adult Insects

1 (2). Episterna of metathorax with deep longitudinal groove. Forecoxae
elongate, conical ...... ... ...... 1. Obrium Curt.

2 (1). Episterna of metathorax uniform, without longitudinal groove.
Forecoxae not elongate, spherical 2. Stenhomalus White.

Larvae

1 (2). Prothoracic eusternum convex, distinctly demarcated from pre-

sternum by groove. Anterior margin of hypostoma around inner

angles of sclerites with spinelike outgrowth or deep notch

1 . Obrium Curt.

2 (1). Prothoracic eusternum not convex, not demarcated from prester-

num by groove, merges with it to form common surface. Anterior
margin of hypostoma smooth around inner angles of sclerites,
without spinelike outgrowth, with only inconspicuous notch. . . .
2. Stenhomalus White

Pupae

1 (2). Posterior end of abdominal tergite VII narrowly rounded coni-

cally, not produced; abdominal tergite VIII not bent dorsad ....
1 . Obrium Curt.

2 (2). Posterior end of abdominal tergite VII conically produced to a

point; abdominal tergite VIII bent dorsad (lateral view)

2. Stenhomalus White.

1. Genus Obrium Curt.

Curtis, 1825, Brit. Entom., vol. 2, p. 91; Plavil’shchikov, 1940, Fauna
SSSR, 22, 2, 133-134; Linsley, 1963, Cerambycidae of North America,
vol. 21, pp. 136-137.

Adult: Readily recognized by structure of forecoxae, long abdominal
sternite I, presence of deep longitudinal groove on metepisternum, and
other features. Body elongate. Head short, with deep or indistinct punc-
tation and sparse long thin hairs. Eyes large, convex, coarsely faceted,
highly angularly emarginate or saccate. Antennae long, slender; 10th or
11th (female) or even 9th (male) segment generally extends beyond ely-
tral apex. Pronotum elongate, with distinct tubercle laterally, broad
flange anteriorly and especially at base, with long light-colored hairs.
Elytra parallel, apically with rounded inner and significantly truncate
outer angle, with more (O. gracile Plav.) or less [O. cantharinum (L.)]
large punctation. Forecoxae elongate, not spherical. Femora clavate.
Hind tarsi significantly shorter than tibiae. Abdominal sternite I long,
equal to rest of sternites together (female) or markedly shorter (male).
Sternite II posteriorly emarginate in female and here setae broaden ter-
minally and form dense brush.

Larva: Body elongate, slender. Pronotal scutum coriaceous, demarcat-
ed laterally by deep longitudinal grooves, glabrous, without hairs. Pro-
thoracic eusternum convex, coriaceous, usually glabrous, without hairs.
Locomotory ampullae on abdominal segments 1, 11, and VII poorly deve-
loped, markedly produced nodularly on segments III to VI, widely sepa-
rated. Abdominal segments anterior and posterior to latter ampullae
with deep transverse grooves forming secondary pseudoannuli.

32 Pupa: Antennae slender, bent angularly. Head narrowly rounded on
occiput, without setae, glabrous. Pronotum longitudinal, with distinct
tubercle on each side, with setae or acicular spinules forming one (at
base) or two (at base and middle) transverse bands in posterior half,
glabrous in anterior half. Abdominal tergites with well-developed or
weak spinules.

Four species are known in the fauna of the USSR; of these, one is
distributed in Europe up to the Urals, one widespread in the Palearctic,
and two in the Far East, mainly in Ussuri-Primor’e region. Up to six
species are known in Southeast Asia and 10 in North America. All spe-
cies of this genus found in northern Asia inhabit forests and are ecolo-
gically associated with only deciduous woody species.

Type species: Ceramhyx cantharinum Linnaeus, 1767.

KEY TO SPECIES

Adult Insects

1 (4). Body and elytra rusty, entire abdomen light or dark rust.

2 (3). Pronotum smooth, without large deep punctation. Europe and

Asia 1.0. cantharinum (L.).

3 (2). Pronotum not smooth, with large deep punctation. Eastern Asia.

2. O. brevicorne Plav.

4 (1). Body and elytra blackish-brown, abdomen rusty-red only at tip.

Eastern Asia 3. O. gracile Plav.

Larvae

1 (4). Ocelli near antennal bases distinct, pigmented, black.

2 (3). Anterior margin of hypostoma near inner angles of sclerites with

sharply produced spinelike projections, posterior to which lie arti-
culate spinelike maxillary processes. Found on poplar and asp.
I. O. cantharinum (L.).

3 (2). Anterior margin of hypostoma near inner angles of sclerites with

notch accommodating articulate maxillary processes. Found on
ash 2. O. brevicorne Plav.

4 (1). Ocelli not perceptible near antennal bases. Pigmentation lacking.
Found on ash 3. O. gracile Plav.

Pupae

1 (4). Abdominal tergites with large spinules that usually form trans-

verse band in posterior half.

2 (3). Pronotum with fine setae forming one or two transverse bands.

Spinules on basal adominal tergites broaden markedly

1.0. cantharinum (L.).

3 (2). Pronotum with long acicular spinules forming single transverse

band. Spinules on basal abdominal tergites broaden insignificantly.
2. O. brevicorne Plav.

4 (1). Abdominal tergites with minute, barely discernible spinules form-

ing transverse row 3. O. gracile Plav.

1. Obrium cantharinum (L.).

Linnaeus, 1767, Syst. Nat., 12th ed., vols. 1-2, p. 637 (Cerambyx);
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 134-135; Cherepanov and
Cherepanova, 1973, Nov. i. maloizv. vidy fanny Sibiri, 6th ed., pp. 47-49.
33 Adult (Figure 9): Body comparatively large, light rust, with conspi-
cuous black, highly convex eyes, and faintly punctate, lustrous pronotum.
Head short, retracted into prothorax almost up to eyes, tubercularly
convex around antennal bases from inner side, with broad median longi-
tudinal suture, transversely impressed anteriorly, lustrous, with barely
perceptible, minute, very sparse, obliterated punctation. Eyes large,
broadly emarginate, coarsely faceted . Antennae widely separated, slender;
10th segment (male) or only 11th (female) extends beyond elytral apex,
with minute tightly adherent hairs; 1st to 4th segments (especially in
male) with semiadherent long hairs; 3rd segment longer than 4th (male)
or equal to it (female).

Pronotum elongate, distinctly narrower than head, with conical tu-
bercle anterolateral to middle, basally and apically with slight transverse
flange (first, at base, usually more prominent than one at apex); pro-
notal disk convex, lustrous, with fine uneven punctation, and long erect
rusty hairs directed backward in anterior half, and forward in posterior
half. Scutellum elongate, with almost parallel sides, narrows slightly an-
teriorly, broadly rounded posteriorly (mainly in male) or triangular,
pointed anteriorly (mainly in female), with barely perceptible, obliterated
punctation, with very minute hairs visible only under high magnification.
Elytra with parallel sides, convex, apically with narrowly rounded inner
and slightly truncate outer angle, with projecting shoulders; inward to
latter with distinct longitudinal impression, with punctation large and

Figure 9. Obrium cantharinum (L.).

more distinct in anterior half, less distinct in posterior half, and minute
semiadherent hairs. Legs moderately long; hind femora dilate gently to-
ward apex. Hind tibiae 1.5 times longer than tarsi; 1st segment of hind
tarsi equal to two successive together.

Adomen lustrous, with minute, barely perceptible punctation, ^and
fine rusty hairs; sternite II in female with emarginate posterior margin
and here with deep golden setae. Body, antennae, and legs monochro-
matic, rusty-black. Body length 5.5 to 10.0 mm.

Egg: White, moderately elongate, rounded at poles. Chorion" with
flat reticulate sculpture, gaps between cells resemble septa, thin. Length
1.0, width 0.4 mm.

Larva (Figure 10): Body elongate, narrows markedly from thorax
to posterior end. Half of head retracted into prothorax; head ’narrows
slightly anteriorly. Epistoma with smooth lustrous dark brown border

along anterior margin; fuses laterally with parietals; frontal sutures not
visible, median suture dark brown, streaklike. Hypostoma divided by
broad gula into two triangular sclerites with one sharp spinelike pro-
jection on anterior margin near each inner angle, posterior to which
lie posterior articulate maxillary tubercles. Parietals in anterior half
laterally with long dense hairs bent backward. Antennae comparatively
long, with four segments. Pigmented ocelli present, one on underside of
each segment. Clypeus short, white, sometimes masked up to apex by
anterior margin of epistoma. Labrum round, convex, glabrous on disk,
with tender setae along margins in anterior half. Mandibles narrow in-
significantly toward apex, broadly rounded there, with cultrate mesal
surface.

Pro thorax thick, markedly broader than meso- and metathorax. Pro-
notum slopes moderately toward head, with coarse light-colored hairs
in anterior half and laterally, without perceptible yellow transverse band.
Pronotal scutum slightly convex, white, demarcated laterally by deep
34 longitudinal grooves, with thin longitudinal striation. Alar lobes with
long light-colored hairs. Prothoracic presternum densely hairy; eusternum
convex and coriaceous, glabrous at base, with sparse hairs at apex.
Thoracic legs lacking.

a — head and pronotum; b— abdominal tergite IV with dorsal locomotory ampulla.

Abdomen with thin light-colored hairs laterally; abdominal segments
with additional pseudoannuli demarcated by narrow constrictions; loco-
motory ampullae coriaceous, developed on abdominal segments I to VII.
Dorsal locomotory ampullae on tergites III to VI highly convex, widely
separated; semicircular groove anterior to sides. Ventral locomotory am-
pullae with pitlike depression, with radial grooves. In prepupa rudimen-
tary thoracic legs and sclerotized lobes appear laterally on meso- and
metathorax. Body length of mature larvae 12 to 15 mm, width of head
1.8 mm.

Pupa (Figure 1 1 ): Characterized by sharp spinules on abdominal tergi-
tes and long setae on pronotum. Head rounded on occiput, distinctly
broadens between antennae, usually with three setae anterior to anten-
nal bases forming transverse row. Antennae slender, bent angularly pos-
terior to midfemora, pressed to legs ventrally, with apices adjoining
sides of head.

Pronotum slightly longer than width at base, convex and smooth on
disk, with projecting conical tubercle laterally; narrows somewhat toward
anterior end, with small flange at base; in some specimens fine setae
form transverse row at base anterior to flange; in others an additional
transverse band occurs in middle; in still others only transverse bands
present in posterior third; and in yet others one sparse tuft occurs on
each flank basally. Meso- and metanota convex, with or without stray
setae. Hind femora clavate, with apices bent somewhat dorsad.

Abdomen broadens in region of sements III and IV, narrows toward
base and tip. Abdominal tergites convex, in posterior half with sharp,
usually setaceous spinules that are very broad at base and form trans-
verse row. Spinules generally in pairs on disk, single on sides. Tergite
VII elongate, narrowly rounded posteriorly, with three to five spinules
forming tuft. Tip of abdomen (ventral view) slightly obtuse, almost
rounded. Valvifers of female small, insignificantly separated, project
laterally at apex. Body length 7.0 to 10.0 mm, width of abdomen 2.8 to
3.2 mm.

35 Material: Collected in Ussuri-Primor’e region, Tuva, Altai, Ob’ region,
and the southern Urals. Adult insects 26, larvae 49, pupae— 10 males
and seven females, larval exuviae with beetles from cells 11.

Distribution: West and eastern Europe, northern Asia. More abun-
dant in south, especially in foothills of western Siberia (Tuva, Altai).

Biology: Inhabits forests with viable asp and poplar. Beetles begin to
fly from end of June through August. Maximum numbers found in July.
Female lays eggs singly on branches and trunks of Populus. Colonize
sections with smooth, usually thin bark. Embryonic development conti-
nues up to three weeks. For example, larvae began hatching on August
28 from eggs laid July 30 through August 4. The atmospheric tempera-

Figure 11. Pupa of Obrium cantharinum (L.), female.

ture during this period varied from 10.8 to 28.1°C (average 18.5 +
0.6°C). Newly hatched larvae bore bark and plug eggshell remnant with
frass. They make longitudinal galleries under bark from the top down-
ward, which are deeply impressed in sap wood. Galleries are densely
sealed with fine frass consisting of wood and partly bark. Mature larva
burrows gradually into wood and makes cell there in upper layer along
trunk and plugs inlet with frass. Larva then turns around, facing inlet,
and pupates with head upward. Length of larval gallery under bark up
to 23 cm and width 2.0 to 3.0 mm. Width of inlet 2.5 to 3.0 mm, length
of pupal cell up to 13 mm, width up to 4.0 mm, length of plug sealing
cell 7.0 mm.

36 Pupation commences in first 10 days of June and ceases by end of the
month. Maximum number of pupae seen in middle of last 10 days of
June. Duration of pupal stage 2.5 to 3.0 weeks. For example, in Altai
beetles began emerging in middle of first 10 days of July from pupae
appearing in last 10 days of June, with the first beetles seen on July 4th.
Emerging beetles break inlet plug, push back frass, nibble an oval (flight)
opening in bark and exit. Flight opening 2.0 mm x 3.0 mm. Emergence
of adults from wood commences end of June and ceases in last 10 days
of July. Mass emergence takes place in middle 10 days of this month.
Life cycle completed in two years (Table 2).

Weight of larvae before pupation 12.1 to 36.0 mg, of pupae 11 to
33 mg, and of newly developed beetles before emerging from wood 9.0
to 27.5 mg. Females considerably larger. For example, records of 13
specimens (six males and seven females) revealed that male larvae before
pupation weighed 13.1 to 21.0 mg, pupae 11 to 19 mg, and adult in-
sects 8.5 to 15.0 mg; respective values for females were 20.8 to 36.0, 19 to
33, and 16.0 to 27.5 mg, i.e., nearly double those of males.

Long-horned beetles of this species colonize mainly mature, often
overmature trees. They first colonize the apex, then central section of the
crown, and sometimes significant parts of the apex and trunk simulta-
neously. In 1975 we found a focal point of this species near Lake Telets
in Altai. It colonized not only branches but also many trunks of asp up
to 30 to 36 cm in diameter. Colonization of the trunk occurred to a
height of 2.0 m to the apex. Population density during pupation (June 18)
six per dm^. Viable trees were colonized. Initially such trees became stag
headed, after which desiccation set in. This species was soon followed by
Acanthoderes clavipes Schr., Saperda perforata Pall., S, scalar is (L.), and
others which colonized the same trees.

2. Obrium brevicorne Plav.

Plavilstshikov [Plavil’shchikov], 1940, Fauna SSSRy 22, 2, 138-139.

Adult (Figure 12): Proximate to O. cantharinum (L.) but differs in
more elongate, distinctly punctate pronotum, and deep coarse puncta-
tion of elytra. Head with sparse deep punctation, long yellowish hairs.

Table 2. Periods of development of Obrium cantharinum (L.)

Year of
development

May

June

July

August

September

October

1st

LPA

PAE

AEL

2nd

3rd

LPA

PAE

AEL

For explanation of abbreviations used here and elsewhere in the text, see p. 29
of Volume I.

Figure 12. Obrium brevicorne Plav.

and smooth longitudinal streaklike suture. Antennae in male and female
longer than body; 10th segment (male) or 11th (female) extends beyond
elytral apex; 3rd segment longer than 4th, considerably shorter than 5th.
Eyes deep, usually angularly emarginate.

Pronotum elongate, longitudinal, 2.0 times longer than width at base:
anterior to middle with projecting obtuse tubercle laterally; broad, well-
demarcated flange basally and distinct but less demarcated flange anteri-
orly; moderately convex on disk; with deep distinct punctation and long
37 light rust erect hairs. Scutellum lustrous, smooth, narrows significantly
toward apex, sometimes almost triangular, narrowly rounded or almost
pointed at tip. Elytra elongate, with parallel sides, convex, with rounded
humeri, with dense deep punctation. Hind tarsi 0.66 length of tibiae; 1st
segment slightly longer than two successive together or equal to them.

posterior margin of abdominal sternite II deeply notched, with more ex-
tensive setaceous brush in front, with acicular setae and from flanks long
piliform setae bent inward. Body, elytra, legs, and antennae monochro-
matic, brownish or reddish-rust, eyes black. Body length 6.0 to 9.0 mm.

Egg: White, matte, oval, rounded at one pole, extended collarlike at
the other, with flat reticulate sculpture. Length 1.2 mm, width 0.45 mm.

Larva (Figure 13); Differs from O. canthariimm (L.) in smoothened
projections on anterior margin of hypostoma, markedly elongate abdo-
men, and presence of yellow transverse band on anterior margin of pro-
notum. Head insignificantly retracted into prothorax, notably narrows
anteriorly. Epistoma with narrow lustrous brownish-rust border along
anterior margin, divided by faint median longitudinal suture. Sclerite of
hypostoma triangular, with sharp inner angle; sharp subulate articulate
maxillary process set in notch on anterior margin near sharp inner angle.
Parietals with long dense hairs directed backward in anterior half. An-
tennae short, conical. One pigmented ocellus near each antennal base.
Clypeus transparent, hyaline, broadens markedly at base. Labrum very
small, transversely oval, with broadly rounded anterior margin, barely
perceptible setae along margins. Mandibles black and thick, short, with
transverse groove on outer side at base.

Figure 13. Larva of Obrium brevicorne Plav.
a — head and pronotum; b— abdominal tergite IV
with dorsal locomotory ampulla.

Prothorax broadens in posterior half, narrows gently in anterior half.
Pronotum slopes slightly toward head, with thin tender hairs in anterior
half and laterally, transverse yellow band near anterior margin interrupt-
ed medially by white longitudinal field. Pronotal scutum white, slightly
convex, bound on sides by longitudinal grooves, slightly produced in
middle of anterior margin and appears bifurcate here. Prothoracic pre-
sternum with dense yellow hairs; eusternum glabrous, coriaceous.

Abdomen finely elongate, with short indistinct light-colored hairs
laterally. Abdominal segments III to VI markedly elongate, with sharply
(nodular) projecting dorsal and ventral locomotory ampullae, with trans-
verse grooves in front and behind giving impression of pseudoannuli.
Body length of mature larvae 13 to 14 mm, width of head 1.6 mm.

Pupa (Figure 14): Readily recognized by acicular spinules on prono-
tum and large spinules at posterior margin of abdominal tergite VII.

39 Figure 14. Pupa of Obrium brevicorne Plav., female.

Head flat between upper lobes of eyes, spherically convex on occiput,
with or without acicular spinules near antennal bases. Antennae bent
angularly ventrad, with apices adjoining sides of head (female) or extend-
ing beyond it (male).

Pronotum elongate, laterally with medially projecting obtuse tubercle,
basally with more (male) or less (female) distinct transverse flange, with
paired acicular spinules forming transverse band in front of it; disk con-
vex, smooth, narrows somewhat anteriorly. Mesonotum convex, with
acute projecting tubercle laterally. Scutellum elongate, rounded posteri-
orly, with fine transverse striation. Metanotum with median longitudinal
groove and one to three spinules laterally.

38 Abdomen elongate, narrows toward anterior and posterior ends.
Abdominal tergites with short spinules directed backward, forming com-
mon transverse band of individual tufts (two to five spinules each) in
posterior half. Posterior end of tergite VII with group of spinules, of
which two large ones bent down and forward, forming transverse row,
the rest fine, usually erect. Valvifers of female widely separated, coni-
cal, apically resemble mastoid projection, and slightly shifted laterally.

Material: Collected in Ussuri-Primor’e region. Adult insects 195,
larvae 46, pupae — four males and two females, larval exuviae with beetles
from cells five.

Distribution: Ussuri-Primor’e region, Komarovka and Suvorovka
Rivers, Osinovka, Vladivostok.

Biology: Inhabits broad-leaved forests. Ecologically associated with
ash {Fraxinus). Flight of beetles from end of June to mid-August. In 1971
first beetles sighted on June 25 in Ussuri forest region. Maximum num-
ber seen in first half of July. Beetles emerge from cells with developed
gonads. Ovaries of one female dissected on emerging from wood contain-
ed 32 mature eggs, of another female 36. Beetles mate soon after emer-
gence from cells and oviposit. Larvae hatch from eggs in July and August.
They bore bark, make longitudinal, often meandering galleries under it,
and plug them with fine frass. Galleries sharply impressed on sapwood.
Larva makes cell in upper layer of wood at end of gallery and pupates
in it.

Pupation commences from end of May. Pupae mainly found in June.
Young beetles emerge in second half of June. Weight of larvae 13.4 to
25.6 mg, pupae 12.5 to 24.0 mg, and beetles emerging from cells 10 to
19 mg. Average weight of beetles 13.8 mg.

This species generally inhabits tops and branches of drying and phy-
siologically weakened trees. Mainly damages ash. For example, of 169
beetles collected from wood cuttings gathered in a forest, 168 were found
in ash and one in maple. Obrium gracile Plav. and Rhopaloscelis bifasci-
atus Kr. quite often colonize ash together with O. brevicorne Plav.

39 3. Obrium gracile Plav.

Plavilstshikov [Plavil’shchikov], 1933, Entom. Anzeig., vol. 13, p. 167;
=graciliforme, Lipp, 1939, Entom. Bldtt., voL 35, p. 255; Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 137-138.

Adult (Figure 15): Differs from other species of this genus in dark
brown body and small size. Body moderately elongate. Head broader
than pronotum, with large, sometimes obliterated punctation, with
smooth median longitudinal suture between antennae, and comparatively
long light-colored (gray) hairs. Eyes highly convex, coarsely faceted,
deeply emarginate, saccate. Antennae slender, longer than body; 9th seg-
ment (male) or llth (female) extends beyond elytral apex, with minute
adherent hairs; underside of 1st to 5th segments with several semi-
adherent hairs.

Figure 15. Obrium gracile Plav.

Pronotum 1.5 times longer than width at base, with large conical
tubercle laterally in middle, well-developed anterior and posterior flanges,
more rarely flanges poorly developed, uniformly convex on disk, some-
times with longitudinal carina, with distinct, rather sparse punctation
(gaps between punctures lustrous, 3.0 to 5.0 times larger than punctures
themselves), with long light-colored adherent hairs behind anterior half
and in front of posterior half. Scutellum elongate, triangular, smooth,
narrowly rounded or pointed posteriorly. Elytra convex, with parallel
sides, well-developed dents around humeni, large, dense, deep puncta-
tion (distance between punctures not more than, or 1.5 times larger than
punctures themselves), with semiadherent light-colored fine hairs. Hind
femora apically with gradually thickening clava. First segment of hind
tarsi almost 1.5 times longer than two successive together.

40 Abdomen narrows slightly toward tip. Abdominal sternite I equal to
II and HI sternites (male) or II to V (female) sternites together; sternite
II in female highly emarginate posteriorly and here with unusually dense
setae, thickened and scraperlike at apices, forming triangular brush
bound in front by short acicular setae, with long setae incurved from
flanks that seem to brace brush. Sternite III very short, only sides per-
ceptible, with long light-colored setae. Entire body, antennae, elytra,
and legs monochromatic dark brown, with rusty tinge. Body length 5.0
to 6.0 mm.

Egg: White, elongate, pointed at one pole, rounded at the other.
Chorion with indistinct sculpture. Length 1.0 mm, width 0.4 mm.

Larva (Figure 16): Very similar to larva of O. brevicorne Plav. but
differs in faint nonpigmented ocelli, denser setae on abdomen, and
smaller mature larva. Half of head retracted into prothorax; head narrow-
ly rounded anteriorly. Epistoma fuses laterally with parietals, divided
by barely perceptible median longitudinal suture, with smooth lustrous
brownish-rust border along anterior margin, and behind border several
tender setae forming transverse row. Anterior margin of hypostoma
near inner angles of sclerites with slight notch for articulate maxillary
processes. Parietals with dense tuft of long hairs bent backward. Ocelli
not seen near antennal bases, pigmentation lacking.

Pronotum broadly rounded anteriorly, with erect light-colored hairs
laterally; hairs on disk in anterior half bent down forward and sideways.
Pronotal scutum white, slightly convex, bound laterally by deep grooves,
outwardly convex, with two slight emarginations along anterior margin,
insignificantly produced forward medially but not bifurcate, with fine
longitudinal striation. Prosternum with short light- colored hairs bent
down and slightly forward.

Abdomen slender, elongate, with sparse short hairs larerally. Abdo-
minal segments III to VI with elongate lateral tubercles and protuberant

locomotory ampullae. Body length of mature larvae 11 to 1 2 mm, width
of head 1.0 mm.

41 Pupa (Figure 17): Differs from other species of this genus in barely
developed spinules on abdominal tergites. Head between upper lobes of
eyes not impressed, merges gradually posteriorly with common hemi-
spherically convex occipital surface, and with pair of acicular spinules
or without them anterior to antennae. Antennae bent angularly, with
apices pressed to sides of head ventrally.

Pronotum with parallel sides, laterally with large conically project-
ing tubercle; base, especially laterally, with narrow transverse constric-
tion, convex and lustrous on disk, with seven or eight long acicular
spinules in posterior third forming transversely elongate tuft. Meso- and
metanota with one or two setae laterally.

Abdomen moderately elongate, broadens slightly medially, narrows
markedly toward tip. Posterior half of abdominal tergites with barely
discernible spinules forming indistinct, at places widely interrupted row.
Tergite VII narrowly rounded posteriorly, with pair of widely separated

Figure 16. Larva of Obrium gracile Plav.
a — head and pronotum; b— abdominal tergite IV
with dorsal locomotory ampulla.

/^/

'W'

'W #«1^4 Ilf'

iV-S.' '.-iK^ ^'*-i" ’X vV:^-'

mlA,: ::Mm.

Figure 17. Pupa of Obrium gracile Plav.

spinules on posterior margin bent forward. These spinules lacking in
some specimens. Valvifers of female very small, hemispherical, slightly
separated. Body length 4.0 to 6.5 mm, width of abdomen 1.1 to 1.4 mm.

Material: Collected in Ussuri-Primor’e region and in Sakhalin. Adult
insects 294, larvae 113, pupae — eight males and one female, larval
exuviae with beetles from cells eight. Two generations of beetles were
raised in the laboratory.

Distribution: Ussuri-Primor’e region, Sakhalin. We found large num-
bers in forests in the Komarovka River basin.

42 Biology: Lives in broad-leaved forests. Emergence of beetles from
wood commences mid-June and ends in July. Flight of beetles continues
up to August inclusive. They avidly feed on flowers of spiraea and
other plants and frequently mate while feeding. Gonads mature during

this period. Ovaries of one female dissected eight days after emergence
from v^ood contained 18 mature eggs, of another female 19.

Female lays eggs on thin shoots 0.6 to 3.0 cm in diameter on matur-
ing or mature drying trees. Sometimes undergrowth colonized. Larvae
hatch from eggs in three to four weeks by rupturing shell. They immedi-
ately bore bark and make longitudinal sinuous or straight galleries
underneath it, faintly impressed on alburnum, and plug them with fine
frass. Mature larva bores wood and makes cell there along axis of shoot.
Sometimes cells are made under bark, in which case impression in
alburnum deep. Length of gallery under bark 10,0 to 26.2 cm, in wood
up to 1.6 cm; width of gallery 3.0 to 6.0 mm. Length of cell 7.0 to 10.0
mm, width 2.0 to 4,0 mm.

Larvae pupate in May and June after second hibernation. Maximum
number of pupae seen in nature from middle of first 10 days to begin-
ning of last 10 days in June. Pupae develop for about two weeks in
nature. Beetles nibble oval opening (1.0 mm x 2.0 mm) in bark surface
and exit from cell through it. First beetles recorded in cells on June 3.
Found in large numbers in middle 10 days of June, Emergence of
beetles from wood commences mid- June. For example, in 1971 field
observations recorded 67 beetles emerging from cuttings of ash shoots
colonized by this species; of these, 39 (58.2%) emerged in second half
of June and 28 (41.8%) in July. First beetles emerged June 15 and last
July 26, i.e., emergence of beetles from wood continued for more than
a month.

According to records of 16 specimens, larvae before pupation weigh-
ed 4.0 to 13.1 (5.8+0.5) mg, pupae 3.0 to 12.0 (5.1 +0.5) mg, and
beetles 1.6 to 9.8 (4.2+ 0.4) mg. Population density of this species is
comparatively high. For example, a shoot 1.5 cm in diameter and
30 cm long contained four beetles. Similar density recorded for other
shoots.

Obrium gracile Plav. is among the monophagous insects inhabiting
ash {Fraxinus rhynchophylla and F. mandschurica). In a three-year period
shoots collected from a forest contained 233 beetles, of which 230 were
found in ash and three in unidentified shoots. Obrium brevicorne Plav.
and Rhopaloscelis bifasciatus Kr. colonize ash together with this species.

2. Genus Stenhomalus White

White, 1855, Catal. Coleopt, Brit, Mus., vol. 8, p. 243; PlaviPshchi-
kov, 1940, Fauna SSSR, 22, 2, 139-140; Gressit, 1951, Longicorn Beetles
of China, vol. 2, p. 163; Kojima and Hayashi, 1969, Insect Life in Japan,
vol. 1, p. 57.

Adult: Body elongate, flat. Head short. Genae barely perceptible.
Antennae slender, longer than body. Pronotum elongate, with small
obtuse tubercle laterally. Elytra parallel, flat on disk, individually round-
ed apically. Metepisternum uniform, without longitudinal groove. Abdo-
minal sternite I in male considerably shorter, in female only slightly
shorter than rest of sternites together. Sternite II in female posteriorly
emarginate, with setae forming dense brush; sternite III with long in-
curved setae on posterior margin.

43 Larva: Very similar to larvae of the genus Obrium. Differs in ab-
sence of pigmented ocelli, more uniform anterior margin of hypostoma,
absence of grooves demarcating pro thoracic eusternum, with latter
merging into general surface of presternum.

Pupa: In structure of head and pronotum very similar to the genus
Obrium. Differs markedly in structure of tip of abdomen. Abdominal
tergite VII produced conically at posterior margin, projects far beyond
tergite VIII (lateral view), with four large spinules forming transverse
row in posterior half, of which middle ones bent down, forward, and
inward and lateral ones backward and sideways. Tip of abdomen with
pair of urogomphi terminating in setaceous sclerotized spinule.

Only one species found in the fauna of northern Asia. Most species
of this genus occur in Southeast Asia and the islands of Japan.

Type species: Stenhomolusfenestratusy^hiXQ, 1855.

1. Stenhomalus vulcanus Tsher.

Tsherepanov [Cherepanov and Cherepanova], 1976, Novosti fauny
Sibiri {Nov. i maloizv. vidy fauny Sibiri, 10th ed.), pp. 79-83.

Adult (Figure 18): Body elongate. Head short, directed forward,
with uneven, coarse punctation. Genae barely perceptible. Frons slightly
elevated around antennal bases, with median longitudinal suture.
Antennae long, slender; 9th segment extends beyond elytral apex; 1st to
6th segments with long setaceous hairs on inner side; 3rd segment
longer than 4th, shorter than 5th. Eyes convex, coarsely faceted, deeply
emarginate on inner side.

Pronotum elongate, 2.0 times longer than wide, anterior margin
laterally with projecting tubercle, disk at level of lateral tubercles broadly
(but insignificantly) impressed, with coarse punctation, minute white
adherent and large brownish erect hairs, posteriorly with more distinct,
and anteriorly less distinct flange. Scutellum flat, very small, broadly
rounded apically. Elytra parallel, apically broadly rounded individually,
with more truncate outer angles, somewhat flat on disk, with sparse
deep punctation, minute gray adherent hairs, and sparse, uniformly
disposed, semierect, acicular setae. Legs comparatively long; femora
clavate. Hind tarsi distinctly shorter than tibiae. First segment of hind

tarsi markedly shorter than all segments together.

Abdomen of female comparatively broad, Sternite I considerably
shorter than remaining sternites together; sternite II broadly emarginate,
in posterior half with dense reddish-rust brush of setae broadening api-
cally. Posterior margin of sternite III with row of long piliform setae
directed inward, framing brush from behind. Abdomen of male convex,
smooth, almost glabrous, with minute hairs not forming compact cover.
Tergite II without notches and without brush in posterior half. Body
black. Elytra dark brown, with rusty, light-colored band extending
obliquely from humeral tubercle to middle. Antennae and legs rusty,
hind femora darkened. Sometimes antennae dark brown, with rusty,
light-colored ringlets at base of segments. Body length 8.0 to 9.0 mm.

Egg: White, narrowly rounded at poles. Chorion hyaline, without
cellular sculpture. Length 0.6 to 0.8 mm, width 0.3 to 0.4 mm.

Larva (Figure 19): Recognized by structure of prothoracic eusternum,
absence of pigmented ocelli, and anterior margin of hypostoma. Head
44 markedly retracted into prothorax, narrowly rounded anteriorly. Epis-
toma along anterior margin with lustrous brownish-chestnut border,

divided by median longitudinal streaklike suture, fuses with parietals
laterally. Frontal suture not visible. Hypostoma narrows anteriorly,
with straight lateral sutures, smooth anterior margin, with barely dis-
cernible notch near inner angles of sclerites. Parietals laterally in ante-
rior half with numerous hairs bent backward. Antennae with four seg-
ments. Ocelli not visible. Clypeus hyaline, transparent. Labrum small,

Figure 19. Larva of Stenhomalus vulcanus Tsher.
a— head and pronotum; b— abdominal tergites IV and V
with dorsal locomotory ampullae.

with pointed or narrowly rounded anterior margin, with short setae
along margins. Mandibles broad, black, basally light rust, as if with frill.

Pronotum narrows slightly anteriorly, with rounded anterior margin,
dense light rust hairs on disk anterior to scutum and laterally, with
smooth, lustrous, yellowish-rust, transverse band in anterior third in-
terrupted medially by longitudinal white gap. Pronotal scutum white,
bulges slightly, demarcated laterally by longitudinal sutures, Prothoracic
presternum with uniform, numerous light rust hairs; eusternum coria-
ceous, glabrous, merges with presternal surface, not demarcated from it
by groove.

Abdomen slender, elongate, with barely perceptible light-colored
hairs laterally. Locomotory ampullae on segments I, II, and VII poorly
developed, on III to VI highly convex, nodular, and project sideways,
with barely perceptible longitudinal groove. Tergite VIII hyaline, with -
parallel sides, elongate, distinctly longer than VII. Tergite IX slightly
wider at base, or not wider than long, broadly rounded posteriorly, with
sparse minute hairs. Body length of mature larvae 10 to 11 mm, width
of head 1.0 mm.

Pupa (Figure 20): Characterized by straight projecting urogomphi
on tip of abdomen and four large spinules on posterior margin of ter-
gite VII. Head with uniformly rounded occiput, two or three setae near
antennal bases, and with or without minute setae on anterior margin in
front of clypeus. Antennae pressed to sides of body, bent ventrad, loop-
like, with apices adjoining sides of head.

Pronotum longitudinal, bulges, smooth, lustrous, with barely per-
ceptible or very distinct obtuse tubercle laterally (sometimes looks round-
ed on sides), tapers somewhat more posteriorly than anteriorly, with
long setae forming transverse band or row anterior to hind clivus; in
some specimens setae cover entire posterior half. Anterior half of pro-
notum usually glabrous, rarely with stray setae. Meso- and metanota
convex, each with pair of widely separated setae.

Abdomen elongate, narrows markedly toward base. Abdominal ter-
gites slightly convex, with four small sharp setaceous spinules in pos-
45 terior half forming transverse row. Posterior margin of tergite VII pro-
jects conically, rises above tergite VII (lateral view), with four large
spinules, of which middle ones directed forward and inward, lateral
backward and sideways. Tip of abdomen with pair of straight urogomphi
projecting posteriorly and terminating in sharp sclerotized setaceous
spinule. Valvifers of female small, tubercular, widely separated. Body
length 7.0 to 8.0 mm, width of abdomen up to 1.8 mm.

Material: Collected in Kunashir (Golovnina volcano). Adult insects
32, larvae nine, pupae 10, larval exuviae with beetles from cells eight.

Distribution: Kuril’ Islands, Kunashir Island.

Biology'. First found in broad-leaved forests of Golovnina volcano.
Beetles begin flying in first half of summer. Colonize thin shoots of
Phellodendron sachalinense. Single female can lay up to 20 eggs.

Newly hatched larvae first make galleries in bark, then under bark
along shoot, plugging them with fine frass. Galleries under bark deeply
46 impressed on alburnum. Sometimes gallery looplike, longitudinally elon-
gate. Mature larva bores wood at end of gallery, makes cell along shoot
at a depth of 4.0 to 10.0 mm, plugs inlet with frass, turns head toward
inlet, and pupates. Length of larval gallery under bark up to 16 cm,
width 1.0 to 4.0 mm. Size of inlet 1.5 mm x 2.0 mm. Length of pupal
cell up to 9.0 mm, width 3.0 mm.

Pupation commences early August, terminates by September. Young
beetles emerge end of August and in September, but hibernate in pupal

cells, abandoning them the following spring. Thus the life cycle is com-
pleted in two years (Table 3). Weight of larvae before pupation 5.5 to

18.0 mg, pupae 5.0 to 13.5 mg, and young beetles before hibernation

4.0 to 10.0 mg. Females generally larger than males.

Table 3. Periods of development of StenhomaJus vulcams Tsher.

Year of

development

May

June

July

August

September

October

1st

AEL

2nd

LPA

3rd

AEL

Population density on some trees comparatively high. For example,
on a shoot 52 cm long and 3.4 cm in diameter 19 specimens were found,
and on another shoot 24 cm long and 2.3 to 3.1 cm in diameter 11
specimens. Colonize physiologically weakened trees. Sydonia divaricata
Bat. appears on the same trees a year or two later.

17. Tribe NATHRIINI ( = PSEBIINI)

Adult insect similar to beetles of Molorchini in general appearance,
but differs in posteriorly open forecoxal cavities. Head short; genae very
short, in form of narrow strip anterior to lower lobes of eyes. Elytra
short, cover part of segment I or first half of abdomen.

Larva similar to larvae of Obriini, but characterized by locomotory
ampullae protruding markedly laterally and tubercularly on abdominal
segments III to VI, and silvery reticulate sculpture of prothoracic
eusternum and pronotal scutum.

Pupa without setae on pronotum. Abdominal tergite VII with four
spinules. Frons near base of antennae with or without large spinule.

Many genera of this tribe are represented in African fauna. Only one
genus inhabits the southern regions of Europe and spread east, apparently
up to the southern Urals.

1. Genus Nathrius Breth. ( = Leptidea Muls.)

Brethes, 1916, Rev. Chilena de Hist. Nat., vol. 20, p. 76; = Leptidea,
Mulsant, 1839, Hist. Nat. Coleopt. France Longicornica, 1st ed, p. 105;
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 141 {Leptidea); Linsley, 1963,
Cerambycidae of North America, 21, 4, 153 {Nathrius).

Adult’. Body very small, elytra short, Frons short, with median longi-
tudinal suture. Eyes highly convex, reniform, without distinct notches.

Antennae long, slender. Abdominal segment I long, equal to all other
segments together.

47 Larva: Body elongate, very small. Thoracic legs lacking, locomotory
ampullae well developed on abdominal segments III to VI, hairs on
pronotal disk not very dense, bent down and forward. Monotypic
genus.

Type species: Nathrius porteri Brethes, 1916.

1. Nathriusbrevipeniiis(Muls.)

Mulsant, 1839, Hist. Nat, Coleopt. France Longicornica, 1st ed.,
p. 105 (Leptidea); Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 142-143
(Leptidea); Duffy, 1953, Monograph Immat. Stages of British and Im-
ported Timber Beetles, pp. 197-198 (Leptideella); Demelt, 1966, Tier-
welt Deutschlands, vol. 52, pp. 60-61 (Nathrius); Linsley, 1963, Ceramby-
cidae of North America, 21, 4, 154-156 (Nathrius).

Adult (Figure 21): Characterized by small body, short elytra. Head
short, broad, with fine punctation, and stray long yellowish hairs. Frons
bound laterally by smooth brownish riblike fold, with median longitudi-
nal deep suture. Genae very short. Eyes large, highly convex, coarsely
faceted. In antennae 9th or 10th segment extends beyond elytral apex;
1st segment markedly thickens toward apex; 5th longer than 3rd and
4th segments together. Pronotum slightly longer or almost not longer
than width in middle, insignificantly narrowly rounded anteriorly and
posteriorly, with deep, very broad groove posteriorly and narrow trans-
verse groove anteriorly and with double edge at these sites; flat on disk,
with fine, uneven, widely spaced punctation, long erect yellowish, hairs,
smooth longitudinal prominence laterally in anterior half. Scutellum
elongate, smooth, broadly impressed, glabrous, rounded posteriorly.
Elytra comparatively short, extend just beyond abdominal base, with
gaping, narrowly rounded inner angle and more truncate outer angle at
apex in posterior third, flat on disk, with fine sparse punctation, smooth-
ened apically, thin yellowish (very long at base) hairs. Femora broaden
notably in distal half, highly flattened laterally. Hind tibiae with long
setaceous hairs. Hind tarsi 0.50 length of tibiae. First segment of hind
tarsi slightly curved, 2.0 times longer than two successive segments
together.

Abdomen of male narrow; abdominal sternites with sparse long
hairs, with steeply truncate posterior margin. Abdomen in female broad
(in specimens not yet ovipositing appears bulged); sternite II broadly
emarginate posteriorly, with dense short setae in notch forming golden-
orange, and on anterior margin rounded brush; latter on posterior
margin fringed with dense, long, light-colored, piliform setae; sternite
III short, emarginate posteriorly, with dense long setae bent down and

Figure 21. Nathrius brevipennis (Muls.),

backward on posterior margin. Abdominal tergite V emarginate (male)
or narrowly rounded (female) posteriorly. Body and elytra chestnut-
brown, darker in male, almost black, light colored with rusty tinge in
female. Antennae and legs rusty-brown. Body length 3.0 to 6.0 mm.

Egg: Orangish-red, oval, tapers markedly toward poles, narrowly
rounded at one pole, pointed at the other. Chorion smooth, lustrous.
Length 0.8 mm, width 0.4 mm.

Larva (Figure 22): Very similar to larvae of Obriini in general ap-
pearance and structure of abdominal locomotory ampullae. Differs in
sparse (stray) hairs laterally on head, shagreen sculpture on base of pro-
thoracic eusternum and pronotal scutum. Body elongate. Head mark-
edly retracted into prothorax. Epistoma slightly convex, with broadly
emarginate anterior margin, its brownish border broadening laterally
here and reaching laterally up to antennal bases. Frontal sutures not
visible, median suture likewise indistinct. Hypostoma lustrous, tapers
48 anteriorly, with deep notch on anterior margin near inner angles for

Figure 22. Larva of Nathrius brevipennis {yLxjXs.).

a— head and pronotum; b — abdominal tergite IV with dorsal
locomotory ampulla.

articulate, spinelike, acutely sclerotized maxillae. Parietals ventrally on
anterior margin with broad brownish border that does not cover antennal
sockets from behind, in anterior half with stray light-colored hairs that
do not form dense tuft (in larvae of Obriini they form tuft bent down
and sharply backward). Antennae short; 1st segment thick, whitish;
apical segment slender, brownish, more sclerotized. Clypeus broad, tra-
pezoid, convex, narrows markedly anteriorly, lustrous. Labrum convex
on disk, broadly rounded apically, with short light-colored setae along
margins. Mandibles broad, almost semicircular on mesal surface. Maxillae
at base of cardo with sclerotized brown band, and on inner margin of
band with sharp basal spinules in marginal notch of hypostoma.

Pronotum transverse, slopes significantly toward head, moderately
convex on disk, anterior to scutum with innumerable uniformly spaced
hairs bent down and forward, laterally in anterior half with dense long
hairs directed backward. Pronotal scutum white and convex, glabrous
(without hairs), silvery shagreen, laterally bound by deep longitudinal
grooves extending forward almost up to anterior margin of pronotum.
Prothoracic presternum convex, with sparse short hairs; eusternum gla-
brous, conveXj with minute silvery reticulate sculpture, broadly rounded
apically, demarcated by deep groove. Thoracic legs lacking.

Abdomen elongate, with very small, barely perceptible, light-colored
hairs laterally. Dorsal and ventral locomotory ampullae poorly developed
or almost not developed on segments I, II, and VII, and well developed
(project laterally as tubercles) on segments III to VI, as a result of which
abdomen appears nodular. Tergites VIII and IX with deep punctation
49 on disk, gaps between punctures not larger than punctures themselves.
Segment X projects conically, with sparse minute light-colored hairs.
Anal opening triradial. Body white. Body length 5.5 to 7.0 mm, width
of head 0.6 to 0.8. mm.

Pupa (Figure 23): Readily recognized by slender, very small, highly
elongate body, absence of setae and spinules on pronotum, presence of
four spinules bent down and forward at posterior margin of abdominal
tergite VII. Head short, round (ventral view); occiput lustrous, narrowly
rounded. Frons broad, with faint median longitudinal groove, laterally
near antennal bases with large, sharp (conical), sclerotized spinule or
without it. Antennae long, in second half bent ventrad, looplike, with
apices adjoining foretibiae.

Pronotum elongate, almost 1.5 times longer than width at base, uni-
form, convex, and lustrous on disk, with broadly rounded sides, and an-
terior margin produced angularly in middle. Mesonotum moderately
convex, with slightly protruding scutellum on posterior margin, lustrous,
with notably rounded posterior margin.

Abdomen elongate, with parallel sides. Abdominal tergites bulge.

lustrous, medially with faint narrow groove; sides of segments IV to VI
50 just slightly rounded. Tergite VII narrows markedly in posterior half,
posterior margin narrowly (angularly) rounded, and near it four sclero-
tized spinules bent down and forward form transverse row. Tergite VIII
convex, lustrous, projects conically posteriorly, matte laterally and on
posterior projection. Pro-, meso-, and metanota, and abdominal tergites
I to VI and VIII without spinules or setae. Tip of abdomen (ventral
view) slightly obtuse, laterally without perceptible carina. Valvifers of
female project, notably separated. Body length 4.6 to 5.1 mm, width of
abdomen 0.8 mm.

Material: Larvae collected in the Caucasus (Sochi), and pupae and
beetles raised from them. Adult insects 15, larvae 18, pupae — 14 males
and females, larval exuviae from cells with pupae and beetles six.

Distribution: Southern Europe from Atlantic coast to the southern
Urals (Ural’sk and Orenburg), northern Africa. Imported into North
America (New York, California) and South America (Chile, Argentina).
Sightings sporadic. Abundant in the Caucasus. We found them in Sochi
region.

Biology: Inhabits deciduous forests, wooded parks, and gardens.
Ecologically associated with large number of deciduous woody species.
Flight of beetles from June through August. After mating, female ovi-
posits on thin shoots of apple, willow, ash, hazelnut, alder, and other
woody and shrub species. In the laboratory, where shoots of apple,
lilac, and bird cherry were available, only apple was selected for ovi-
position. Female first scrapes minute particles from bark surface by means
of abdominal setae, lays a sticky egg on shoot, then covers it with minute
particles (dust) collected earlier. Egg consequently looks like a small
tubercle merging with bark surface. Sometimes eggs are not covered with
epidermal bark particles and then are distinctly visible against general
background of bark as orange dots. Oviposition is rather intensive. For
example, one female laid 38 eggs in one day on an apple shoot. Ovaries
of another female just emerging from pupal cell contained 38 eggs. Larvae
hatched in the laboratory at 24°C after 15 to 18 days (average 16.1 +
0.3 days). We had kept 42 eggs under observation.

Newly hatched larvae make longitudinal galleries under bark, deeply
impressed in wood, and plug them with fine frass. Walls of galleries
steep, acute. Sometimes parallel galleries fuse to form broad striplike
niche, which is compactly plugged with frass. Mature larva bores deep
into wood (on thin shoots up to pith), makes longitudinal gallery there,
then cell at end of gallery in which it pupates with its head toward inlet.
Length of gallery under bark up to 17 cm, in wood up to 6.0 cm, width
2.0 to 3.0 mm. Cells often disposed directly around inlet and some-
times under bark. Length of cell 8.0 to 10.0 mm, width 1.8 to 3.0 mm.
Pupae develop in two to three weeks. For example, in the laboratory at
16 to 23° C (18 + 0.5° C) young beetles emerged 14 to 16 days after pupa-
tion. They exited from cells one week later with developed gonads, made
an oval flight opening (up to 1.0 mm X 0.8 mm) on surface of shoot,
and emerged. Males emerge before females. Beetles begin to reproduce
soon after emergence from wood. Weight of larvae before pupation 1.4
to 3.0 mg (male) to 3.1 to 8,2 mg (female), pupae 1.2 to 2.8 mg (male)
to 2.7 to 7.1 mg (female), and beetles before emerging from wood 0.9
to 2.4 mg (male) to 2.3 to 6.0 mg (female).

Nathrius brevipennis (Muls.) inhabits many woody and shrub species,
forming very dense populations. Six larvae were once detected on a thin
51 alder shoot 45 cm long and 4.0 to 8.0 mm in diameter. On another
shoot, 17 cm long and 12 mm in diameter, 15 specimens (larvae, pupae,
and adults) were found. In Sochi region (Caucasus) larvae were found on
alder {Aims), ash {Fraxinus), and pear {Pyrus), In the laboratory they
avidly sought out shoots of apple {Malus). According to data available
in literature (Plavil’shchikov, 1940; Duffy, 1953; Linsley, 1963; Demelt,
1966), this species develops on willow (Salix), dog rose (Rosa), chestnut
(Castanea), fig (Ficus carica), hazelnut (Corylus), walnut (Juglans), haw-
thorn (Crategus), and other woody and shrub species.

18. Tribe MOLORCHINI

Adult insect distinguished by short (Molorchus, Nadezhdiana, Epania)
or markedly elongate elytra that fall open at the back (Stenopterus, CaF
limellum). Legs comparatively long, femora often clavate, petiolate.
Mesosternal process tapers toward apex (Molorchus), or flat, broad, al-
most square (Stenopterus, Nadezhdiand).

Larva with short transverse head highly retracted into prothorax.
Epistoma barely demarcated, frontal suture almost imperceptible. Parie-
tals in anterior half with long replicated hairs.

Pupa characterized by narrow elongate body. Disk of pronotum with
long setae, usually forming two transverse bands, of which one anterior
to middle and second posterior to it (anterior to posterior flange). Abdo-
minal tergites with minute spinules forming transverse row. Abdominal
tergite VII with very large spinules bent forward.

Two genera of this tribe (Molorchus and Nadezhdiand) are known in
northern Asia, of which Molorchus is distributed in the Holarctic. The
existence of the genus Molorchus (Shabliovskii, 1936) needs to be con-
firmed since the type specimens are lost.

KEY TO GENERA
Adult Insects

1 (2). Mesosternal process narrow, sides not parallel, tapers toward apex,

usually triangular 1 . Molorchus F.

2 (1). Mesosternal process broad, with parallel sides, almost square . . . .

. 2. Nadezhdiana Tsher.

Larvae

1 (2). Abdominal tergite IX without spinules, smooth, with sparse fine
hairs on apex 1. Molorchus F.

2 (1). Abdominal tergite IX with spinules forming sizable tuft on disk.
2. Nadezhdiana Tsher.

Pupae

1 (2). Pronotum without anterior flange. Setae on abdominal tergite VIII

without sclerotized border at base 1 . Molorchus F.

2 (1). Pronotum with distinct anterior flange. Setae on abdominal tergite

VIII with sclerotized border at base. 2. Nadezhdiana Tsher.

1 . Genus Molorchus F.

Fabricius, 1792, Entom. Syst., 1, 2, 366; = Caenoptera, Thomson,
1859, Sknd. Coleopt., vol. 1, p. 150; = Mulsant, 1862, Co/.

52 France Longicornica, 2nd ed., p. 226; PlaviPshchikov, 1940, Fauna SSSR,
22, 2, 152-155; Gressit, 1951, Longicorn Beetles of China, vol. 2, p. 169;
Linsley, 1963, Cerambycidae of North America, 21, 4, 156-157; Kojima
and Hayashi, 1969, Insect Life in Japan, vol. 1, p. 60; Mamaev and Dani-
levskii, 1975, Lichinki zhukov-drovosekov, pp. 186-188.

Adult: Body elongate. Intercoxal mesosternal process tapers toward
apex. Elytra short. Hind (membranous) wings fall open on abdomen.
Femora clavate. Antennae usually long, with 12 segments in male (sub-
genus Molorchus s. str.) or 1 1 segments (subgenus Linomius Muls.).

Larva: Head highly retracted into prothorax. Parietals laterally in
anterior half with long hairs bent backward. Epistoma poorly demar-
cated. Frontal suture imperceptible. Hypostomal sclerites widely sepa-
rated by gula, pointed or rounded at inner angles. Labial submentum
uniform, fairly convex, smooth {Molorchus s. str.) or longitudinally striate
[M. ussuriensis Plav., M. umbellatarum (Schreb.)]. Dorsal and ventral
locomotory ampullae on abdominal segments I to VII slightly convex,
shagreen, matte. Thoracic legs lacking.

Pupa: Body elongate. Head short, moderately bent under. Antennae
pressed to sides of body, in male bent forward at level of tip of abdomen
to form common ellipsoidal ring framing periphery of body from below
[M. umbellatarum (Schreb.)], or form independent narrow loops with
apices adjoining head [M. minor (L.)]. Pronotum convex, with fine setae
on disk forming two transverse bands separated into individual tufts
{M. ussuriensis Plav.), or with scutiform spinules on hind clivus (M. kiesen-
wetteri Muls. and Key). Abdominal tergites with minute setaceous spi-
nules directed backward. Tergite VII rounded posteriorly, with two to
six large spinules bent forward. Valvifers of female with thick setae an-
teriorly {M. ussuriensis Plav.) or without setae [M. umbellatarum (Schreb.)]

More than ten species of the genus Molorchus are known in USSR
fauna. Of these, seven inhabit northern Asia, including one species {M.
minor (L.)] widely distributed in the Palearctic, two [M. umbellatarum
(Schreb.), M. kiesenwetteri Muls. and Rey] from Europe entering the
southern Urals and partly northern Asia, one {M. heptapotamicum Plav.)
distributed in the southern Urals and northern Asia, two (M. ussuriensis
Plav. and M. incognitus Tsher.) in Ussuri-Primor’e region, and one spe-
cies (M, kobotokensis Ohb.) inhabits Kunashir Island and northern Japan.
No less than 15 species are known in Southeast Asia, 12 in Japan, and
three in North America. Larvae live under bark of trees, mainly in thin
shoots. Two species [(Af. minor (L.), M. kobotokensis Ohb.] inhabit coni-
ferous trees, while the other species [M. ussuriensis Plav., M. incognitus
Tsher., M. umbellatarum (Schreb.) and so forth] inhabit deciduous species.
Mature larva bores wood, makes cell along shoot, and pupates in it with
its head toward inlet.

Type species: Necydalis minor Linnaeus, 1758.

KEY TO SPECIES
Adult Insects

1(2). Antennae with 12 segments in male, 1 1 segments in female (sub-
genus Molorchus s. str.). Elytra posterior to middle with white
oblique band. Eurasia. 1. M. minor (L.).

2(1). Antennae with 11 segments in both male and female (subgenus
Linomius Muls). Elytra posterior to middle without white oblique
band, on disk posterior to scutellum with broad white spot or
without it.

53 3 (12). Pronotum with simple, deep, indistinct punctation.

4 (11). Elytra on disk posterior to scutellum monochromatic, rusty or

brownish, or with whitish diffuse spot lacking sharp outline.

5 ( 6). Elytra on disk posterior to scutellum which large light-colored

(whitish) spot. Antennae of male longer than body. Ussuri-
Primor’e region 2. M. ussuriensis Plav.

6 ( 5). Elytra on disk posterior to scutellum monochromatic, brownish

or rusty, sometimes with blackened apex (M. kiesenwetteri Muls.
and Rey). Antennae of male not longer than body.

7 (10). Pronotum with deep distinct punctation; spaces between punc-

tures distinct.

8 ( 9). Elytra elongate, distinctly longer than pronotum. Europe and

northern Kazakhstan. 3. M. umbellatarum (Schreb.).

9 ( 8). Elytra broaden at shoulders, not longer or only slightly longer

than pronotum. Southern and central Europe, southern Urals,
Central Asia . . ........ 4. M. kiesenwetteri Muls. and Rey.

10 ( 7). Pro no turn with fine, indistinct, very dense punctation forming

shagreen sculpture; spaces between punctures barely visible.

Northern Kazakhstan, southern Urals

5. M. heptapotamicus Plav.

11 ( 4). Elytra on disk posterior to scutellum with sharp while spot,

emarginate at anterior outer angles. Kunashir and Islands of
Japan 6. M. kobotokensis Ohb.

12 ( 3). Pronotum with flat cellular punctation; spaces between cells

narrow, resemble septa. Ussuri-Primor’e region

7. M. incognitus Tsher.

Larvae

1 ( 2). Gula apically broad, 0.66 hypostomal sclerites. Found on coni-

ferous species 1 . M. minor (L.).

2 ( 1). Gula apically narrow, less than 0.50 hypostomal sclerites.

3 ( 6). Labial submentum with minute longitudinal streaks.

4 ( 5). Pronotal scutum without longitudinal striation, basally matte.

Found mainly on maple 2. M* ussuriensis Plav.

5 ( 4). Pronotal scutum with longitudinal striation, not matte basally.

Found on pear and other deciduous species

3. M. umbellatarum (Schreb.).

6 ( 3). Labial submentum without minute longitudinal streaks; at most

with three longitudinal smoothened grooves.

7 (12). Pronotal scutum with two emarginations on anterior margin,

produced forward medially and at anterior angles.

8 (11). Hypostomal sclerites pointed at inner angles.

9 (10). Pronotum anterior to scutum paramedially roundly impressed,

with sparse short hairs there. Found mainly on apple

4. M* kiesenwetteri Muls. and Rey.

10 ( 9). Pronotum anterior to scutum uniform paramedially, almost in

same plane as scutum, with dense long hairs here. Found mainly
on dog rose . 5. M. heptapotamicus Plav.

11 ( 8). Hypostomal sclerites rounded at inner angles, not pointed.

Found on thin shoots of spruce 6. M. kobotokensis Ohb.

12 ( 7). Pronotal scutum not emarginate along anterior margin but

transversely truncate, almost straight, and does not project for-
ward at anterior angles. Found on deciduous species

7. M. incognitus Tsher.

Pupae

1 ( 2). Antennae long, bent forward, looplike in second half in male,
with apices pressed to head; in female antennae arcuate, with
apices pressed to middle of elytra 1 . M. minor (L.).

2(1). Antennae shorter, bent only at apex in male, with apices short
of reaching head; in female antennae bent slightly ventrad, with
apices pressed to sides of body.

3 (12). Abdominal tergite VII with four spinules bent forward and

forming transverse row along posterior margin.

4 (11). Abdominal tergite VIII long, not shorter than VII.

5 ( 8). Tip of abdomen (tergites VIII and IX) with long setae.

6 ( 7). Abdominal tergite VII with six sharp spinules bent forward and

forming transverse row ........... 2. M. ussuriensis Plav.

7 ( 6). Abdominal tergite VII with four spinules bent forward and

forming transverse row ...... 3. M* umbellatarum (Schreb.).

8 ( 5). Tip of abdomen (tergites VIII and IX) glabrous, without setae.

9 (10). Pronotum on hind clivus laterally with innumerable scutiform

spinules bearing one seta each, forming two large tufts anterior
to posterior angles. ..... 4. M* kiesenwetteri Muls. and Rey.

10 ( 9). Pronotum on hind clivus laterally without scutiform spinules,

with only stray setae not forming tufts .................

......................... 5. M. heptapotamkus Plav.

11 (4). Abdominal tergite VIII short, 0.66 length of VII, with dense

setae forming two tufts in posterior half .

.......... ................ 6. M* kobotokensis Ohb.

12 ( 3). Abdominal tergite VII with two spinules bent forward on pos-

terior margin .................. 7. M. incognitus Tsher.

1. Molorchus minor (L.)

Linnaeus, 1758. Syst. Nat., 10th ed., p. 421 (Necydalis); PlaviFshchi-
kov, 1940, Fauna SSSR, 22, 2, 155-158; Cherepanov and Cherepanova,
1973, Nov. i maloizv, vidy fanny Sibiri, 6th ed., pp. 46-47.

Adult (Figure 24): Well distinguished from other species of the genus
Molorchus by oblique white band in posterior half of elytra. Head com-
55 paratively broad, slightly narrower than pronotum. Frons broad and
flat, with large deep punctation. Antennae slender, with II segments
(female) or 12 segments (male); 10th segment (female) or 6th (male) ex-
tends beyond posterior end of body. Eyes finely faceted, broadly emargi-
nate, with very narrow upper lobes. Pronotum notably elongate, angu-
larly broadens medially, with transverse flanges near anterior and poste-
rior margins, dense, minute, longitudinally elongate punctation, smooth
lustrous wartlike elevation on disk paramedially, with long erect, some-
times sessile gray hairs. Scutellum small, narrowly rounded posteriorly,
sometimes with adherent gray hairs. Elytra narrow posteriorly, indivi-
dually rounded apically, taper more from inside, with humeri projecting
forward, sparse deep punctation, and white oblique band in posterior
half extending diagonally from suture forward. Legs long, with long seta-

ceous hairs. Femora clavate; clava of hind femora less than 0.50 length
of shaft (slender part of femora). Hind tarsi shorter than tibiae; 1st seg-
ment considerably longer than two successive together. Body blackish-
brown, antennae rusty, elytra chestnut with rusty tinge and white oblique
band in posterior half; femoral clava dark brown, femoral bases (base of
shaft) and tibiae light rust (f. typica). Sometimes oblique band on elytra
yellow, not white (ab. incarinatus Plav.); rarely body reddish-rust, oblique
band on elytra lacking (ab. rufescens Kiesw.), sometimes elytra blackened
apically (ab. apicalis Plav.). Body length 6.0 to 12.0 mm.

Figure 25. Larva of Molorchus minor (L.).
a — head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla.

56 Egg: White, oval, elongate, rounded at one pole, slightly pointed at
the other, lustrous, without cellular sculpture. Length 1.2 mm, width
0.5 mm.

Larva (Figure 25): Differs in structure of hypostoma, mandibular
features, and other characteristics. Body tapers from prothorax back-
ward. Head transverse, rounded anteriorly. Epistoma slightly flattened,
whitish, at anterior angles with dentate projection, along anterior mar-
gin with dark brown smooth border, behind which lie short piliform
setae, divided by median longitudinal suture, fuses laterally with parietals.
Frontal suture not visible. Hypostoma slightly convex, broadens toward
base, with rounded sclerites on inner margin, and widely separated by
gula. Parietals rounded anteriorly, in anterior half laterally with dense
setaceous hairs bent backward. Antennae conical; 1st segment 2.0 times
thicker than 2nd. Clypeus very small, broadly flattened at base. Labrum
rounded, with sparse short setae. Mandibles thick, short, broadly rounded
apically from inside along cultrate surface, deeply hollowed, transversely
sinuous at base, convex on outer side, smooth, with median transverse
groove. Inner masticatory lobes of maxillae elongate, digitate, with stray
apical setae.

Pronotum in anterior half and laterally with minute setaceous rusty
hairs, with narrow white glabrous border on anterior margin. Pronotal
scutum convex, coriaceous, laterally demarcated by deep longitudinal
groove. Meso- and metanota matte, white, coriaceous, divided by median
transverse groove; prothoracic presternum with uniform rusty hairs;
eusternum coriaceous, glabrous. Meso- and metasterna coriaceous, sha-
green, with stray setaceous, barely visible hairs, divided by median trans-
verse groove.

Abdomen elongate, with sparse tender hairs laterally. Dorsal loco-
motory ampullae moderately convex, shagreen divided in anterior half
by fairly distinct transverse groove joining radial dent on sides. Ventral
locomotory ampullae with short longitudinal groove laterally, and short
transverse groove extending inward from it. Abdominal tergite IX on
posterior margin with setaceous hairs forming indistinct transverse row.
Body length 11 to 12 mm, width of head 1.8 to 2.0 mm.

Pupa (Figure 26): Readily recognized by markedly broadened seg-
ments IV and V of abdomen, sharp apical thickening of femora, and
microscopic setaceous spinules on abdominal tergites. Head glabrous,
without setae, spherically rounded on occiput, with faint broad longitu-
dinal groove between antennae, slightly elevated from inside around an-
tennal bases. Antennae of male long, pressed to sides of body, bent for-
ward in middle, looplike at level of abdominal tergite VI, with apices
pressed against head in region of frons. Antennae of female short, bent
forward in middle at level of tergite III, with apices pressed against
underside of forelegs.

Pronotum convex, longer than wide, insignificantly rounded on sides,
with narrow flange at base, slopes gradually toward anterior margin,
without anterior flange; disk with fine setae forming transverse band
before middle and sometimes two small lateral tufts behind middle. Meso-
notum faintly convex, glabrous, with posteriorly extended and rounded
scutellum. Metanotum with median longitudinal groove, laterally with
three to five setae forming one row extending from anterior angles and
57 sloping toward longitudinal groove posteriorly. Apices of femora sharply
clavate, dilated. Hind femora extend beyond middle of abdominal ter-
gite VI (male) or only beyond posterior margin of V (female).

Abdominal segments IV and V very broad, narrow sharply ante-
riorly and posteriorly. Abdominal tergites slightly convex, with faint
common median longitudinal groove, on posterior margin (tergites II
to VI) with five to seven minute setaceous paramedial spinules forming
small tuft (base of each spinule with one seta from posterolateral side).
Abdominal tergite VII apically with five or six spinules forming tuft.
Tergite VIII with stray setae, sometimes posterior margin with pair of
setaceous spinules. Valvifers of female large, almost cylindrical, directed

sideways, with round flat tubercle apically. Body length 9.0 to 13.0 mm,
width of abdomen 3.0 to 4.0 mm.

Material: Collected in eastern Ural region, Altai, Ob’ region, Tuva,
taiga of Yenisey region, Trans-Baikal, Ussuri-Primor’e region, and Kuna-
shir. Adult insects 321, larvae 118, pupae 13, larval exuviae with beetles
from cells four.

Distribution: From Atlantic Ocean coast to Pacific Ocean coast.
South from Altai, Tuva, Sayan, northern China, and Korea, north to
Polar Circle; Sakhalin, Kunashir, Islands of Japan (Honshu and Hok-
kaido).

Table 4. Periods of development of Molorchus minor (L.)

Year of

development

May

June

July

August

September

October

1st

AEL

2nd

LPA

3rd

AEL

Biology: Ecologically associated with coniferous vegetation, mainly
spruce. Rises in hills to 2,000 m, found in large numbers at Ayukol’
(Lake Telets) at a height of 1 ,000 m. Beetles fly from end of May to
July. For example, in Altai during the flight period 102 specimens were
collected, with 14 (13.7%) in middle 10 days of June, 81 (79.4%) last
10 days, and seven (6.9%) in July. Flight ceases in second half of July.
Beetles found on flowers of Umbelliferae, Rosales, and other plants.
After mating, female lays eggs in bark crevices on thin shoots of spruce,
fir, and other coniferous species. Colonizes weakened but not yet dried
shoots ranging in diameter from 2.0 to 10.0 cm. Drying undergrowth
often infested. Ovaries of one female contained 10 mature eggs. Em-
bryonic development completed in about three weeks.

Newly hatched larvae bore bark, then make sinuous logitudinal or
transverse galleries underneath it, which are deeply impressed on al-
burnum. Walls of galleries acute, upright; galleries compactly plugged
with fine frass. Sometimes galleries so close together wood looks en-
graved.

Larva of the first year hibernates under bark (Table 4), the next year
bores wood, makes a cell in upper layer along trunk, and pupates inside
it with its head toward inlet. Both inlet and pupal cell sealed off from
rest of gallery by plug of frass. Gallery between cell and inlet remains
58 hollow, free of frass. Inlet transversely elongate, ellipsoidal (3.5 mm X
1.5 mm). Length of hollow gallery up to 15 mm, length of cell 14 mm,
width up to 4.0 mm.

Pupation completed by end of July and in August. Young beetles
emerge in second half of August and in September, enter diapause, and
hibernate. In spring (May-June) emerging beetles rupture plug sealing cell,
nibble oval opening (2 5 mm X 3.0 mm), on shoot surface, and emerge.
Emergence of adults from cells commences in first 10 days of June and
concludes by end of that month.

Weight of larvae before pupation 16 to 80 mg, pupae 10 to 80 mg,
and adult insects 7.5 to 40 mg. One larva weighed 32 mg before pupa-
tion, its pupa 31 mg, and beetle 23 mg, i.e., during metamorphosis the
insect lost 28.1% of its weight.

Population density of this species is very high on some trees. For
example, a fir shoot 25 cm long and 2.0 cm thick contained four larvae,
one pupa, and four young beetles. Generally found on spruce shoots,
rarely on others. For example, while inspecting various woody species
162 insects comprising larval, pupal, and adult stages were found; of
these, 87 were collected from spruce shoots, 55 from fir, six from pine,
four from maple, one from larch, and nine from unidentified shoots.
Pogonocherus fasciculatus Deg., Super da interrupt a Gebl., and others
sometimes colonize the same shoots as this species, M. kobotokensis
Ohb. is found in Kunashir.

2. Molorchus iissuriensis Plav.

(Plavilstshikov) PlaviPshchikov, 1940, Fauna SSSR, 22, 2, 160-161;
Cherepanov and Cherepanova, 1976, Novostifauny Sihiri (Nov, i maloizy.
vidyfauny Sihiri, 10th ed.), pp. 71 ”74.

Adult (Figure 27): Differs from the proximate species Molorchus
umbellatarum (Schreb.) in elongate, densely punctate pronotum, very
fine antennae in male, and other features. Head with dense large deep
punctation (distance between punctures usually not more than punctures
themselves), with broad median longitudinal suture between antennae.
Genae short, length 0.50 width. Eyes large, bulge markedly, with broad
notch; upper lobes narrow, striplike, narrower than 3rd antennal segment,
and markedly fall short of anterior margin of pronotum. Antennae of fe-
male notably shorter than body, thicken slightly toward apex commenc-
59 ing from 5th segment, with short adherent hairs, long and numerous on
first five segments; 11th segment equal to 5th, tapers conically in posterior
half, not bent apically. Antennae of male longer than body, with lOth
segment extending beyond tip of abdomen, slender, do not thicken to-
ward apex, with minute adherent hairs, but sparse long hairs at base; 11th
antennal segment elongate, markedly longer than 5th, apically falcate.

Pronotum elongate, 2.0 times (male) or almost 2.0 times (female)
longer than width at base, posterior margin with sharp broad flange,
anterior margin with distinct transverse groove, notably broadens in
posterior third, and laterally here in male with distinct, in female barely
perceptible smoothened tubercle, with dense punctation, long erect light-
colored hairs, with fairly distinct lustrous smooth paramedial ampullae or
almost without them. Scutellum broadly rounded posteriorly, with minute
gray adherent hairs. Elytra moderate elongately, narrowly rounded
apically, with dense deep large punctation, with erect light-colored (long at
base and on sides, short on disk) hairs, with rounded, poorly visible
humeri. Legs with long light-colored semiadherent hairs. Hind femora
gradually thicken apically, clavate; hind tibiae 2.0 times longer than tarsi.
First segment of hind tarsi longer than 2nd and 3rd segments together.

Abdomen moderately elongate, narrows slightly toward base but more
toward tip. Abdominal sternites lustrous, with sparse punctation, later-
ally with denser long light-colored hairs. Sternite V broadly rounded,
shorter than width at base. Body black, elytra straw-yellow, dark brown
laterally and at apex. Antennae and legs dark brown, with rusty tinge.
Base of femora lighter colored, apex (at clava) black or blackish-brown.
Body length 7.0 to 8.0 mm

Egg: White, elongate, rounded at poles, matte. Length about 1.0 mm,
width 0.4 mm.

Larva (Figure 28): Proximate to larva of Motor chus umhellatarum
(Schreb.) in labial structure. Differs in sparse pubescence on pronotum.

60 Body moderately elongate, narrows from prothorax to abdominal segment
IV. Head narrowly rounded anteriorly; highly retracted into prothorax.

Anterior margin of epistoma with broad lustrous brownish border,
with four or five widely separated setae behind it, at anterior angles with
gently rounded dentate projection slightly produced anteriorly. Median
longitudinal suture and frontal sutures smoothened, not perceptible.
Hypostoma narrows anteriorly, with straight sutures laterally, without
perceptible brownish tentorial pits along sides of gula. Parietals narrowly
rounded anteriorly, in anterior half laterally with long setaceous hairs
directed backward. Antennae short, with apices not projecting beyond
mandibular bases. Clypeus short, broadly flattened. Labrum very small,
white, broadly rounded anteriorly, with sparse long setae on anterior
margin. Mandibles short, thick, smooth on outer surface in anterior half.
Labial submentum transverse, with longitudinal striation and seven
longitudinal carinae. Mentum elongate, rectangular. Maxillary palps
long, project forward beyond apex of inner masticatory lobe.

Pronotum rounded anteriorly, with sparse short hairs in anterior
half on disk and sparse long hairs directed backward on sides. Pronotal
scutum slightly convex, white, glabrous, without hairs on anterior margin,
extends forward medially and at anterior angles, bound laterally by deep
longitudinal grooves. Alar lobes lustrous, with "sparse hairs. Prothoracic
presternum laterally with very dense hairs and disk with very sparse short
hairs; eusternum glabrous, lustrous, bulges moderately; basisternum*
narrow, fold like, with not very dense lateral hairs.

Abdomen laterally with short light-colored hairs. Dorsal locomotory
ampullae moderately convex, with common median longitudinal groove
and short, outwardly convex, longitudinal folds. Vental locomotory am-
pullae separated by two transverse grooves joined laterally, demarcating

Figure 28. Larva of Molorchus ussuriensis Plav.
head and pronotum.

*Strictly speaking, the sternellum — General Editor,

ellipsoidal carina between them. Body length 6.0 to 8.0 mm, width of
head 0.8 mm.

Pupa (Figure 29): Readily recognized by number of spinules on ab-
dominal tergite VII. Body more (male) or less (female) elongate. Head
short, hemispherically rounded on occiput, with three thick setae from
inner side of eyes. Antennae pressed to sides of body, with five or six
spinelike outer projections, in female medially bent ventrad with apices
pressed to hind tibiae; in male bent forward, looplike, with apices
pressed to sides of body, forming elongate ellipse (ventral view). Eyes with
or without setae in middle.

Pronotum elongate, broader in female, broadly rounded anteriorly,
slopes somewhat toward anterior margin, basally with more (female) or
less (male) sharp flange, lustrous, convex on disk, with well-developed
groups of setae forming two transverse bands, one anterior to middle,
the other in posterior third. Mesonotum slightly convex, glabrous. Meta-

Figure 29. Pupa of Molorchus ussuriensis Plav., female.

notum with median longitudinal groove, with one or two barely percep-
tible paramedial setae. Apices of hind femora just reach posterior margin
of abdominal segment VI.

Abdomen of female broader, narrows anteriorly and posteriorly.
Abdominal tergites I and II glabrous, without spinules. Abdominal ter-
gites III to VI posteriorly with minute sharp spinules forming somewhat
regular or indistinct row. Abdominal tergite VII with six large spinules
directed forward, forming transverse row, with stray long setae along
61 sides of row. Apex of tergite VIII with long fine setae forming trans-
verse row. Valvifers of female hemispherical, apically contiguous, with
lateral tubercles and thick seta in front. Body length 6.0 to 6.5 mm, width
of abdomen 1.3 to 1.5 mm.

Material: Collected in Ussuri-Primor’e region. Adult insects 19, larvae
three, pupae five, larval exuviae with beetles from cells 10.

Distribution: Ussuri-Primor’e region (Sedanka station, Komarovka
River, Kedrovaya Pad’ sanctuary, Suvorovka River).

Biology: Beetles fly from end of May, early June. Rarely seen on
flowers. Eggs laid on thin shoots of Manchurian striped maple (Acer
tegmentosum). Larvae live under bark, make longitudinal galleries, gener-
ally sinuous initially, then straight, well impressed on alburnum, and
plug them with fine frass. Mature larva bores wood, extends gallery along
shoot, plugs inlet with fine frass, and at end of gallery makes pupal cell
14 mm long, 2.0 mm wide, in which it pupates with head toward inlet.
Length of gallery under bark 9.0 cm, maximum width 3.5 mm. Length of
gallery in wood with cell up to 3.0 cm. Larvae pupate at end of summer.
Young beetles appear A.ugust-September, hibernate in cells, and with
the onset of warm weather in spring push frass out, nibble round or
slightly oval opening in bark, 1.8 mm in diameter, and exit. Emergence
of beetles from cells ceases early June. Weight of pupae 5.9 to 8.7 mg,
of young beetles on emergence from cells 3.4 to 7.0 mg.

Beetles colonize shoots 2.5 to 4.8 cm in diameter. Population density
of shoots varies widely. Two beetles were found on a shoot 56 cm long
and 4.7 cm in diameter, while on a shoot 34 cm long and 4.0 cm in dia-
meter five specimens were found; including four pupae and one beetle
(Cherepanov and Cherepanova, 1976).

3. Molorchus umbellatarum (Schreb.)

Schreber, 1759, Nov. Spec. Ins., p. 9 (Necydalis); Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 159--160.

Adult (Figure 30): Very similar to Molorchus ussuriensis Plav. Differs
in much shorter, less punctate pronotum, short 1st segment of hind tarsi,
and other features. Head not wider or somewhat narrower than prono-
tum, frons flat, with sparse deep punctation, vertex with median longi-

tudinal smooth band, sinciput with dense punctation. Genae short, coar-
sely punctate. Eyes broadly emarginate, upper lobe not narrower than
basal diameter of 3rd antennal segment. Antennae of male longer than
body, with 9th segment extending beyond tip of abdomen; 1st to 4th
segments lustrous at base, with numerous long erect hairs, matte from
5th segment, with minute adherent hairs, and 1 1th segment equal to 5th,

Figure 30. Molorchus umbellatarum (Schreb.).

somewhat bent down at apex. Antennae of female shorter than body,
reach only posterior margin of abdominal sternite IV, and 11th segment
considerably shorter than 5th, equal to 4th.

Pronotum 1.5 times longer than width at base, broadens angularly
behind middle where laterally small, sometimes smoothened, indistinct
tubercle occurs, with broad sharp posterior flange, anteriorly with trans-
verse groove, with uneven large, sometimes minute punctation, laterally
and sometimes medially with smooth, longitudinally elongate, lustrous
ampullae, with not very dense erect light-colored hairs. Scutellum nar-
rows apically, with dense punctation, narrowly rounded posteriorly.
Elytra moderately elongate, project forward at humeri, individually
rounded, sometimes angularly, at apex, somewhat impressed subapically,

62 with uneven punctation; hairs on sides long, on disk short, light-colored
and erect. Hind femora almost extend beyond abdominal sternite IV,
thicken gradually, clavate apically. First segment of hind tarsi longer
(female) than two successive together, or equal to them (male).

Abdomen elongate. Abdominal sternites lustrous, with sparse fine
punctation, with dense semierect hairs. Body black or dark brown with
rusty tinge, antennae and legs dark brown, bases of femora (up to clava)
usually light rust. Elytra straw-yellow, dark brown apically, laterally,
and basally. Body length 6.0 to 8.0 mm.

Egg: White, narrows gradually and sharply pointed at one pole,
obtusely rounded at the other. Chorion with fine sculpture, imparting
matte appearance. Length 1.0 mm, width 0.4 mm.

Larva (Figure 31): Differs from larva of Motor chus ussuriensis Plav.
in very narrow (not transverse) labial submentum and other features.
Body elongate, narrows insignificantly from thorax backward. Head
short, transverse, retracted into prothorax up to lateral hairy field on
parietals. Epistoma bulges slightly, divided by well-developed, brownish
median longitudinal suture, faintly demarcated laterally. Frontal sutures
barely visible. Hypostoma flat, narrows sharply anteriorly, with straight
lateral sutures. Hypostomal sclerites widely separated by gula. Parietals
in anterior half with long hairs directed backward, forming common
lateral hairy field with hairs on sides of thorax (ventral view). Clypeus
broad, notably flattened at base. Labrum transversely oval or slightly
elongate, narrows basally, broadly rounded apically, convex, in anterior
half with short setae. Mandibles thick, light red at base, black in second
half. Labial submentum rectangular, slightly elongate, with longitudinal
striation. Striation more distinct in mature larvae. Labial palps project
forward almost up to apex of maxillary palps.

63 Pronotum narrows anteriorly from base, rounded at anterior margin,
with dense long hairs laterally, sparse short hairs on disk in anterior
half. Pronotal scutum slightly convex, longitudinally rugulose, insigni-

Figure 31. Larva of Molorchus umbellatarum (Schreb.)
a~head and pronotum; b — tip of abdomen.

ficantly produced medially at anterior margin, laterally straight, bound
laterally by deep longitudinal grooves. Alar lobes glabrous and lustrous,
or with stray hairs. Pro thoracic presternum with somewhat dense long
hairs laterally and sparse short hairs on disk; eusternum insignificantly
convex, coriaceous, with distinct furrows.

Abdomen moderately elongate, with sparse short hairs laterally.
Dorsal locomotory ampullae divided by common median longitudinal
groove, with convex lateral longitudinal fold, with grooves radiating
inwardly from fold forming common elongate ellipse. Ventral locomo-
tory ampullae with transverse groove in posterior half, transverse furrows
curved forward in anterior half, and lateral dent. Abdominal tergite IX
broadly rounded apically, with sparse long erect hairs in posterior half.
Body length 8.0 to 11.0 mm, width of head 1.2 mm.

Pupa (Figure 32): Readily recognized by chaetotaxy on frons and
pronotum. Head between antennae flat (female) or longitudinally impress-
ed, with 10 setae between lower lobes of eyes, usually arranged in pairs
and forming irregular transverse band. Eyes without setae in middle.

Figure 32. Pupa of Molorchus umbellatarum (Schreb.).

Antennnae of male pressed to sides of body, smoothened on outer side,
without discernible tubercles, bentventrad, looplike near abdomen, with
64 apices pressed against each other forming common elliptical ring. Anten-
nae of female pressed to sides of body, with apices somewhat bent ven-
trad; 5th to 8th segments with barely perceptible spinelike tubercle at
apex (in Molorchus ussuriensis Plav. tubercles better developed, project
notably).

Pronotum bulges, notably longer (male) or almost not longer (female)
than width in middle; disk anterior to middle with setae forming trans-

verse continuous band, posterior to middle with setae forming broad
interrupted band, in anterior third near anterior margin with stray setae,
and basally with narrow transverse flange. Mesonotum slightly convex,
glabrous, without setae, with roundly extended scutellum behind. Meta-
notum with faint median longitudinal groove, laterally with long setae
(male) or without them (female).

Abdomen elongate, narrows anteriorly, more so posteriorly (from
segment V). Abdominal tergites bulge, lustrous. Tergites I and II without
spinules, III to VI with minute setaceous spinules on posterior margin
forming transverse row and insignificantly interrupted medially. Tergite
VII apically with four large spinules bent down and forward, with two
long large setae lateral to them. Tergite VIII in posterior third with six
to eight long setae forming transverse row. Tip of abdomen with long
thin setae. Valvifers of female small, contiguous, with elongate apical
tubercle. Body length 7.0 to 8.0 mm, width of abdomen 1.5 to 1.8 mm.

Material: Collected in northern Caucasus. Adult insects seven, larvae
17, pupae — one male, three females.

Distribution: Europe, northern Turkey, the Caucasus, and the southern
Urals.

Biology: Inhabits deciduous forests. Often found on fruit trees in
gardens. Ecologically associated with deciduous vegetation. Beetles fly
from early May through June. Quite often found on flowers of Rosales
and other plants. Female oviposits on shoots 0.4 to 2.0 cm in diameter
of drying as well as healthy trees of pear, apple, and other species.

Larvae initially live in bark, later penetrate underneath it, make lon-
gitudinal sinuous galleries, and plug them with fine frass consisting mainly
of bark particles. Galleries faintly impressed on alburnum. Mature larva
bores wood, makes inlet on surface across shoot, and plugs it with fine
frass. Width of inlet 2.0 mm. Thereafter it makes a gallery along axis of
shoot, scoops out cell at end of it, and isolates cell from rest of gallery
with plug consisting of frass. Gallery between cell plug and inlet hollow,
free of frass. Length of cell 12 mm, width 1.7 to 1.8 mm. Length of plug
2.0 mm, of hollow gallery before plug 4.0 to 6.0 mm. Width of gallery
under bark up to 4.0 mm. Larvae hibernate second time in cells. Pupa-
tion occurs in April-May. In the Caucasus (Kislovodsk) pupae seen end
of March. Beetles seen in cell from April 4th. Mass emergence of bettles
occurs mid- April. Weight of larvae before pupation 5.5 to 12.9 mg,
pupae 5.0 to 11.8 mg, and beetles emerging from cells 4.0 to 9.9 mg.
Tetrops praeusta (L.), T. gilvipes Paid., Acmaeops collar is {L.),a,nd others
found together with this species on shoots of pear.

4. Molorchus kiesenwetteri Muls. and Rey

Mulsant and Rey, 1861 Ann, Soc. Linn. Lyon, 8, 2, 173; Plavil’shchi-
kov, 1940, Fauna SSSR, 22. 2. 164-166.

65 Adult (Figure 33): Characterized by dark brown or black body, coarse
punctation, more elongate clava of hind femora. Head short, appears
round laterally, with light brown hairs, uneven, coarse punctation, broad-
ly impressed between antennae, slightly convex around antennal bases.
Genae very short. Eyes deeply emarginate, saccate, finely faceted. Anten-
nae thicken apically, matte from 5th segment, with short, tightly adhe-
rent hairs; 3rd antennal segment equal to 4th or slightly shorter, usually
shorter than 5th.

Pronotum notably elongate, highly produced basally but slightly
apically, somewhat convex on disk, rounded laterally, narrows marked-
ly posteriorly but gradually anteriorly, with coarse deep punctation.

Figure 33. Molorchus kiesenwetteri Muls. and Rey.

long dark brown hairs, small smooth ampullae or without them, and
lustrous in between punctures. Scutellum small, rounded posteriorly, with
fine punctation and sparse adherent hairs.

Elytra short, slightly longer than total width at base, with convex
humeri projecting forward, apically individually broadly or somewhat
narrowly rounded, faintly or deeply impressed on suture posterior to scu-
tellum, with broad subapical dent on disk, notably convex in posterior
quarter, with uniform punctation and short light-colored hairs. Legs
with dense light-colored setaceous hairs. Clava of femora moderately
dilated, in hind femora markedly shorter than shaft. Hind tibiae 1.5 times
longer than tarsi. First segment of hind tarsi equal to two successive
together. Body ventrally with long light-colored semiadherent or erect
hairs. Entire body black with brown or rusty tinge. Antennae dark or
light rust. Legs brownish-rust with chestunt hue. Elytra straw-yellow or
light brown, apically usually darkened (f. typica) or monochromatic,
rusty-brown (ab. castanipennis Reitt.) or brownish-yellow (ab. brunnei-
pennis Plav.). Body length 5.0 to 7.0 mm.

Egg: White (matte), elongate, rounded at anterior pole, pointed at
posterior and narrower. Length l.I mm, width 0.4 mm.

Larva (Figure 34): Very similar to that of Molorchus heptapotamicus
Plav. Differs in more convex pronotal scutum and other minor fea-
tures. Body minute, elongate. Head rounded anteriorly, thick. Epi-
stoma slightly convex, flat and impressed apically, with broad smooth
brownish border on anterior margin, with six minute setae behind it
forming transverse row, faint or barely perceptible median longitudinal
suture in posterior half, and fuses with parietals laterally. Frontal sutures
not visible. Hypostoma flat, markedly narrows anteriorly, with narrow
rusty-brown border on anterior margin and around suture; inner angles
of sclerites pointed. Apex of gula narrow, whitish (light-colored), without
brownish border, markedly broadens toward base. Parietals on anterior
margin with broad (sometimes indistinct posteriorly) rusty-brown border
that does not cover articulate antennal sockets from behind; numerous
long hairs behind border form extensive field. Antennae short, apices
barely extend beyond anterior margin of cephalic capsule; 1st segment
whitish (light-colored), rest brownish. Clypeus brownish, short, barely
visible from behind epistoma. Labrum very small, convex, round, lus-
trous, whitish or brownish, with very sparse light-colored setae. Mandibles
66 on outer side flattened basally and here with narrow transverse groove,
back in apical half, reddish toward base.

Pronotum transversely oval; flat paramedial dents anterior to scutum
rounded posteriorly (hence pronotal scutum appears more convex on
anterior margin, with two emarginations), laterally and in anterior half
in front of scutum with dense thin rusty hairs, in anterior third with

Figure 34. Larva of Molorchus kiesenwetteri
Muls. and Rey.

a — head and pronotum; b — ^abdominal tergite IV with
dorsal locomotory ampulla; c — tip of abdomen.

faint diffuse yellowish-rust transverse band. In some specimens band al-
most imperceptible. Pronotal scutum convex, white, demarcated laterally
by deep longitudinal grooves, often with faint longitudinal smoothened
streaks, anterior margin medially rounded, anterior angles sharply pro-
duced. Prothoracic presternum convex, laterally with sparse long, on disk
short rusty hairs; eusternum glabrous, lustrous. Thoracic legs lacking.

Abdomen elongate, laterally with sparse short light-colored hairs.
Dorsal locomotory ampullae moderately convex, with narrow median
longitudinal groove, with fine shagreen sculpture, in anterior half with
transverse grooves, laterally with short longitudinal distinct or faint
grooves. Ventral locomotory ampullae distinct or slightly convex, medi-
ally separated by common longitudinal groove; disk with rugulose or

shagreen sculpture and narrow transverse groove. Abdominal tergite IX
same length as width at base, narrowly rounded posteriorly, basally
glabrous in anterior half, and with minute light-colored hairs in posterior
half. Body length of mature larvae up to 7.0 to 8.0 mm, width of head
about 1.0 mm.

Pupa (Figure 35): Readily recognized by presence of two tufts of spi-
nules on pronotal base. Body elongate, comparatively narrow. Head
short, between antennae flat or with faint longitudinal groove, on vertex

Figure 35. Pupa of Molorchus kiesenwetteri
Muls. and Rey.

and occiput slopes almost in same plane, very slightly convex, lustrous.
Antennae pressed to sides of body, apices set backward (female) or curve
gently ventrad (male).

Pronotum transverse (female) or not wider than long (male), broad-
ens roundly or angularly posterior to middle, disk insignificantly con-
vex, with dense fine longitudinal striation, and posterolaterally in front

67 of posterior angles with short, scutiform, setaceous, sclerotized spinules
forming two dense, transversely elongate tufts. Mesonotum slightly con-
vex, insignificantly produced angularly or even rounded posteriorly. Meta-
notum insignificantly convex or flattened, with narrow median longitudinal
groove, broadly rounded posteriorly.

Abdomen with parallel sides, narrows posteriorly from segment V,
sometimes broadens slightly in region of segments III and IV. Abdomi-
nal tergites convex, rounded laterally, with faint median longitudinal
groove or without it, without perceptible spinules. Apex of abdominal
tergite VII broadly rounded, lustrous, convex; posterior margin with two
to four minute spinules, visible only under high magnification. Tergite
VIII not shorter than VII, not longer than its own basal width, lustrous,
rounded apically. Valvifers of female elongate, proximate, rounded api-
cally. Hind femora thicken gradually toward apex, with apices reaching
abdominal tergite II. Body length 5.0 to 5.7 mm.

Material: Collected in broad-leaved forests in the southern Urals.
Adult insects 23, larvae 82, pupae — six males and six females, larval
exuviae with beetles from cells three.

68 Distribution: Southern Europe (from Atlantic Ocean coast), Iran, Asia
Minor, Syria, the Caucasus, the southern Urals, Central Asia. Found in
large numbers in the southern Urals (from Orenburg to Ural’sk).

Biology: Inhabits deciduous vegetation. Ecologically associated with
apple, oleaster, willow, and buckthorn. Beetles fly from May to July.
Female oviposits under bark scales on thin drying shoots 4.0 to 12.0 mm
in diameter, comparatively very fertile. Ovaries of one female dissected
before oviposition contained 30 mature eggs.

Larvae bore bark, later make longitudinal galleries under bark im-
printed on alburnum, and plug them with fine frass. Galleries more often
made from top downward, rarely otherwise. Length of gallery under
bark 6.5 to 13.2 cm, width 1.5 to 3.0 mm, sometimes up to 4.0 mm at
end. Mature larva bores deeper into wood, extends gallery there along
shoot, plugs inlet densely with frass, makes pupal cell at end of gallery,
and seals it with plug of frass. Rarely, cells without plug. Larva hiber-
nates in cell. Length of pupal cell 10 to 16 mm, width up to 2.0 mm.
Length of plug sealing cell 3.0 to 4.0 mm, length of hollow gallery be-
tween plug and inlet up to 7.0 mm. Width of inlet 1.8 to 2.0 mm.

Pupation completed end of April or in May (Table 5). Pupa lies in
cell with head toward inlet and develops for about three weeks. In the
laboratory one pupa completed development in 20 days at 15.6 to 20.0°C
(17.6 + 0.3°C), and another in 18 days at 15.6 to 24.0°C (19.9 + 0.6®C).
Young adults emerge from pupae in May. Gonads mature w'hile in pupal
cell. Young beetles rupture seal, push frass aside, nibble oval flight open-
ing in bark (1.0 mm X 1.5 mm) and emerge. Mating and oviposition
occur soon after emergence from pupal cells. Weight of insects highly
variable. For example, records of 28 insects showed: larvae before pupa-
tion weighed 3.1 to 11.5 mg, pupae 2.3 to 10.3 mg, and young beetles
before emergence from wood 1.8 to 8,4 mg.

Table 5. Periods of development of Molorchus kiesenwetteri
Muls. and Rey

Year of
development

April

May

June

July

August

September

1st

LPAE

AEL

2nd

3rd

LPAE

AEL

Molorchus kiesenwetteri Muls. and Rey damages mainly apple shoots,
rarely other species. Twenty- three beetles developed from larvae collected
in nature in the southern Urals: 14 from apple, four from oleaster, three
from willow, and two from buckthorn. Moreover, while inspecting
forests, 87 insects were found (larvae, pupae, and adults), of which 70
came from apple shoots, six from blackthorn, five from willow, and two
from oleaster,

69 5. Molorchus heptapotamicus Plav.

PlaviFshchikov, 1940, Fauna SSSR, 22, 2, 163-164.

Adult (Figure 36): Differs from other species of the genus in mono-
chromatic light or dark rusty body, deeply punctate matte elytra, marked-
ly clavate (dilated) femora, and comparatively long slender antennae.
Head not narrower or slightly narrower than prothorax, with erect light-
colored hairs. Frons flat, nearly upright, with narrow median longitu-
dinal suture, laterally with narrow parallel longitudinal folds, with flat,
barely perceptible reticulate sculpture, tubercularly convex around anten-
nal bases, with broad depression between elevations. Vertex with flat
faint punctation. Eyes sharply faceted, broadly notched in upper part,
with very narrow upper lobes bearing just three rows of minute facets.
Antennae comparatively long, slender; 8th or 9th segment extends beyond

Figure 36. Molorchus kept apot amicus Plav.

elytral apex, with dense light-colored hairs; 3rd antennal segment equal
to 4th, notably shorter than 5th; 11th segment slender, with parallel
sides, pointed apically.

Pronotum elongate, notably longer than wide, broadens angularly
on sides, flattened on disk, with basal flange, near anterior margin with
or without transverse groove, with median longitudinal impression, elon-
gate paramedial, smooth or coarsely punctate ampullae, light-colored
erect hairs, and minute coarse or large flat punctation, appearing almost
cellular in some specimens. Scutellum very small, with median longitu-
dinal furrow or without it, broadly rounded posteriorly.

Elytra comparatively elongate, extend far beyond base of hind fe-
mora (lateral view), broaden at humeri, narrow somewhat posteriorly
and there with parallel sides, apically broadly rounded individually, with
notably convex disk, on suture posterior to scutellum with fairly distinct

small impression, matte, with deep, not very dense punctation, and short
light-colored semiadherent or erect hairs. Legs with long setaceous hairs.
Femora apically markedly dilated, clavate; clava of hind femora of male
short, highly dilated, of female less dilated. Body, elytra, antennae, and
legs monochromatic, rusty, rusty-brown, or rusty-chestnut. Antennae
and legs (especially forelegs) sometimes lighter in color. Body length 4.5
to 7.0 mm.

Egg: White, highly elongate, pointed at poles. Chorion smooth, lus-
trous. Length 1.0 mm, width 0.2 mm.

Larva (Figure 37): Body elongate. Head markedly narrows anteriorly,
and half of it retracted into prothorax. Epistoma without notches on
anterior margin, with broad smooth brownish border, produced anterior
angles, in posterior half with distinct median longitudinal suture, and
fuses laterally with parietals. Frontal sutures not visible. Hypostoma

Figure 37. Larva of Molorchus kept apot amicus Plav,
a-— head and pronotum; b — abdominal tergite IV with dorsal
locomotory ampulla; c—abdominal tergite IX.

narrows anteriorly, with rusty-brown border on anterior margin that
broadens toward sides and is medially interrupted by narrow gap. Gula
narrows apically, light-colored, without border. Parietals with broad
rusty-brown border that does not cover antennal sockets, posterior to
antennae with long rusty hairs directed backward and forming extensive
lateral fields on head. Clypeus short, brown, broadly flattened laterally,
resembles transverse band. Labrum slightly transverse, small, broadly
rounded apically, brownish and with short setae. Mandibles with outer
70 transverse groove basally, elongate anteromedially, black, dark or light
red basally.

Pronotum slopes markedly toward head, narrows anteriorly from
base, broadly rounded on anterior margin, with sparse short rusty hairs
on sides and in anterior half. Pronotal scutum white, notably convex at
base as though elevated, bound laterally by longitudinal grooves, rugose
or smooth in anterior half. Sides of prosternum with dense, disk with
sparse setaceous hairs; prothoracic eusternum without distinct lateral
grooves, merges with presternum, glabrous basally, with stray hairs an-
teriorly. Thoracic legs lacking.

Abdomen elongate, with short thin light-colored hairs laterally. Dor-
sal locomotory ampullae slightly convex, transversely elongate, shagreen,
laterally with barely perceptible, obliquely longitudinal dent. Ventral
locomotory ampullae matte, insignificantly convex, separated by trans-
verse shallow groove. Abdominal tergite IX with uniform light-colored
hairs, glabrous only at base. Body length of mature larva 7.0 to 8.0 mm,
width of head 1.1 mm.

Pupa (Figure 38): Characterized by glabrous pronotum, presence of
apical spinules on abdominal tergite VII, and short, markedly dilated
clava of hind femora. Head slightly narrower than pronotum, with broad
longitudinal groove between antennae, flatly impressed on vertex, lus-
trous on occiput, rounded hemispherically. Antennae slender, pressed to
sides of body, apically falcate.

Pronotum glabrous, without spinules, flattened on disk, with thin
transverse striation, rounded laterally, with narrow flange basally; nar-
rows gradually toward apex, not longer than width in middle, with stray
light-colored setae barely visible under high magnification. Mesonotum
transversely impressed, with raised, obtusely projecting scutellum on
posterior margin. Metanotum flat in posterior half, with median longi-
tudinal groove in anterior half, rounded on posterior margin, with
narrow lustrous fringe.

Abdomen narrowly elongate, narrows gradually from segment IV
toward tip. Abdominal tergites convex, with narrow median longitudinal
groove, in posterior third with one or two faint paramedial spinules or
(in region of anterior segments) without them. Abdominal tergite VII

narrows posteriorly, rounded on posterior margin, dilated in posterior
71 third and here with pair of sharp spinules set independently on pro-
duced coriaceous base. Tergite VIII elongate, not shorter than VII,
narrows toward apex, apically 0.50 width at base, smooth, without
spinules. Valvifers of female large, short, cylindrical, obtuse apically.
Hind femora pressed to sides of body, with long thin shaft and short,
highly convex (dilated) clava. Body length up to 6.5 mm, width of
abdomen 1.2 mm.

Material: Collected in the southern Urals. Adult insects eight, larvae
35, pupae — two females. Series of adult insects in collections of the
Zoological Museum, Moscow State University and the Zoological Insti-
tute, USSR Academy of Sciences, Leningrad also studied.

Figure 38. Pupa of Molorchus kept apot amicus Plav., female.

Distribution: From the southern Urals to Balkhash, Hi River, and
Tien Shan foothills. We found it in Orenburg region, Utvinsk village
(Krasnokholmsk forestry).

Biology: Inhabits deciduous woody and shrub vegetation. Develops
on buckthorn and dog rose. Beetles fly from May through June. Female
oviposits on shoots 0.6 to 1.5 cm in diameter. One female can lay more
than 10 eggs in her lifetime.

Young larvae generally appear end of May and in June, live under
bark, make longitudinal galleries impressed in wood, and plug them with
72 frass. Walls of galleries gentle or steep. Sometimes larval galleries parallel
and merge; as a result the upper layer of wood is so badly damaged
that only frass remains under bark. Mature larva bores wood, makes
longitudinal pupal cell there along shoot, and seals inlet densely with
frass. Length of larval gallery under bark up to 7.5 cm, width 2.0 to
3.0 mm. Length of pupal cell 6.0 to 14.0 mm, width 1.5 to 2.0 mm.
Width of inlet up to 2.0 mm. Length of frass plug at inlet 2.5 mm.

Larvae pupate early spring after hibernation. Pupa lies in cell with
head toward inlet. Young beetles appear in May, possibly early June,
nibble flight opening (1.0 mm X 1.5 mm to 1.5 mm X 2.0 mm) on
shoot surface through which they exit. Mating and oviposition occur
soon after emergence. Ovaries of one female dissected six days after
emergence, from wood contained 12 mature eggs. Records of 11 speci-
mens revealed: weight of larvae before pupation 2.8 to 8.2 mg, pupae
2.6 to 8.1 mg, and young beetles before emergence from wood 2.0 to
6.5 mg.

While inspecting vegetation in the southern Urals, 47 specimens
were found; of these, 22 came from dog rose and 13 from buckthorn.*
Five beetles and two pupae developed from the larvae collected in
nature. Population density comparatively high. For example, on a shoot
of buckthorn 17 cm long and 6.0 to 9.0 mm in diameter 12 larvae were
found.

6. Molorchus kobotokensis Ohb. kunashiricus Tsher., ssp. nov.

Ohbayashi, \%?>,Syst. Nat, Descript. New Forms Ceramb. of Japan^
Fragm, Col,yo\. 2, p. 710; vol. 3, pp. 11-12; Kojima and Hayashi,
1969, Insect Life in Japan, vol. 1, p. 60; Nakano, 1970, Beetles of Japan,
vol. 30, Cerambycidae, no. 25, p. 4; Cherepanov and Cherepanova,
1976, Novosti Fauny Sibiri {Nov. i maloizv. vidy fanny Sibiri, 10th ed.),
pp. 74-77; Hayashi and Matsuda, 1976, Bull. Osaka J. Women's Jr.
College, vol. 11, p. 27.

Adult (Figure 39): Differs from nominal form (M. kobotokensis
Ohb.) in dark rusty (brown) legs and antennae, and more distinct punc-

* Source of remaining 12 omitted in Russian original — General Editor.

Figure 39. Molorchus kobotokensis Ohb.

tation on elytra and pronotum. Head short, retracted into prothorax
almost up to eyes. Frons flat and narrow (male) or slightly convex and
broad (female), with dense deep punctation, slightly impressed between
antennae. Genae very short, look like bands. Eyes deeply and broadly
emarginate, upper lobes narrow, cover antennal bases from behind.

73 Antennae long in male, distinctly shorter than body in female; 4th
antennal segment distinctly longer than 3rd, shorter than 5th.

Pronotum highly convex, rounded laterally, notably longer than width
in middle, with broad basal flange, narrow transverse groove near apex,
dense large punctation, semiadherent long light-colored hairs directed
forward on disk and backward on anterior angles of sides, with small
lustrous ampullae (two in anterior half and one elongate medially in
posterior half) or without them. Scutellum comparatively large, rounded
posteriorly, slightly convex, with dense adherent white hairs. Elytra
short and flat, only 1.5 times longer than maximum width, with straight
shoulders, individually rounded posteriorly, with fine, not very dense

Figure 40. Larva of Molorchus kobotokensis Ohb.,
head and pronotum.

punctation and sparse short erect hairs. Hind femora gradually thicken
clavately in second half, with apices reaching posterior margin of ster-
nite III. First segment of hind tarsi equal to two successive segments to-
gether or slightly longer.

Abdomen comparatively broad. Abdominal sternites with dense fine
punctation, matte, lustrous only on posterior margin, with dense ad-
herent light-colored hairs and stray long erect hairs. Body black. Antennae
and legs rusty or rusty-brown. Femoral clava usually blackish-brown,
darkened. Elytra light colored (straw-yellow), dark brown basally and
apically. Body length 7.0 to 8.0 mm.

Larva (Figure 40): Differs from larvae of other species of this genus
in shagreen (matte) locomotory ampullae of abdomen, and presence of
only three longitudinal streaks or grooves on labial submentum. Head
narrow, slightly flattened dorsoventrally. Epistoma with distinct dark
brown median suture, anterior lustrous brownish border with short
setaceous hairs forming transverse row. Frontal suture not visible. Hypo-
74 stoma narrows anteriorly, anterior margin with rusty border, divided
into two triangular, widely separated sclerites by gula; inner angles of
sclerites obtusely rounded. Parietals in anterior half with long setaceous
hairs bent down and backward or straight. Antennae slender, project
markedly from antennal sockets. Clypeus small, trapezoid, brownish-rust
basally. Labrum slightly elongate, convex, broadly rounded anteriorly,
with sparse short setae in anterior half. Mandibles broaden basally, gently
rounded apically. Labial submentum not wider or only slightly wider
than long, with three longitudinally grooved distinct streaks.

Pronotum slightly convex on disk, almost flat, slopes anteriorly, ante-
rior margin broadly rounded, in anterior third with transverse yellow band
with narrow white strip in middle, in anterior half and on disk with sparse
short hairs and laterally with dense very long replicate hairs. Pronotal
scutum slightly convex, white, laterally with longitudinal grooves. Alar
lobes with sparse thin hairs. Prothoracic presternum with regular rusty
hairs on disk and laterally; eusternum slightly convex, without hairs.

Abdomen moderately elongate, laterally with sparse, tender, barely
perceptible hairs. Dorsal and ventral locomotory ampullae slightly con-
vex, separated by common median longitudinal groove, shagreen, matte.
Apex of tergite IX obtusely rounded, with sparse rusty hairs in second
half. Body length before pupation up to 11 mm, width of head 1.5 mm.

Pupa (Figure 41): Characterized by short antennae and presence of
small number of setae on pronotum. Head short, moderately bent under,
broadly impressed between antennae, spherically rounded on occiput.

Figure 41. Pupa of Molorchus kobotokensis Ohb.

with paired or single sessile setae before antennae between lower ocular
lobes. Eyes without setae. Antennae comparatively short, pressed to sides
of body, in male bent down and forward at level of hind femora, in
female slightly dov/nward, with apices touching hind tibiae.

Pronotum convex, almost square (female) or slightly elongate (male),
rounded laterally, with narrow basal flange, slopes gradually toward
apex, without transverse grooves on anterior margin, with thin setae on
disk forming diffuse transverse tuft anterior to middle and two trans-
verse, slightly oblique tufts posterior to middle displaced laterally. Meso-
notum glabrous, without setae, convex, with narrow extended scutellum.
Metanotum with median longitudinal groove with pair of contiguous
projecting setae along each side.

Abdomen elongate. Abdominal tergites convex, on segments III to
VI in posterior half with minute setaceous spinules forming transverse
row, usually interrupted medially by fairly wide space. Tergite VII
broadly rounded apically, posterior margin with four large and two
minute spinules bent down and forward, forming transverse row. Ter-
gite VIII obtusely conical in male, with long hairs bent backward form-
ing semicircle apically, elongate in female, cylindrical, with medial setae
forming transverse row. Valvifers of female large, apically with short
projecting tubercle. Body length 9.5 to 10.0 mm, width of abdomen
2.1 mm.

Material: Collected on Kunashir Island. Adult insects four, larvae
two, and pupae two (male and female).

Distribution: Islands of Japan and Kunashir (Sernovodsk, Alekhino).

75 Biology: Inhabits coniferous vegetation. Beetles fly in June. Female
oviposits on thin drying shoots of growing spruce. Larvae make sinuous
irregular galleries under bark and plug them densely with fine frass.
Galleries impressed on alburnum. Mature larva bores deep into wood,
leaving surface inlet about 2.0 to 3.0 mm wide, and plugging it with
frass. Cell at end of gallery 16 mm long and 2.0 to 3.0 mm wide; larva
lies in cell with head toward inlet and pupates. Length of gallery with
cell in wood about 4.5 cm. Pupation and pupal development completed
by October. Beetles hibernate, abandon wood only in the following
spring (end of May or in June). Weight of larvae before pupation 10.5
to 19.0 mg, pupae up to 17.0 mg, and beetles after hibernation up to
9.0 to 12.5 mg.

Evidently found rarely. Inhabits shoots 2.0 to 4.0 cm in diameter at
lower levels (Cherepanov and Cherepanova, 1976).

7. Molorchus incognitus Tsher.

(Tsherepanov) Cherepanov and Cherepanova, 1975, Zhuki-drovoseki
ivovykh lesov Sibiri, pp. 83-86.

Adult (Figure 42): Readily distinguished from other species of the
genus Motor chus by flat reticulate punctation on pronotum. Body narrow,
elongate. Head retracted into prothorax up to eyes, with large dense
punctation and erect hairs. Frons broad and flat, raised tubercularly
along margins around antennal bases. Eyes markedly and broadly emargi-
nate, distinctly faceted. Antennae slender, with 1 1 segments, shorter
than body, do not reach tip of abdomen; apices of 3rd to 6th segments
thickened nodularly, with long hairs slanting slightly backward; 3rd
antennal segment longer than 4th, equal to 5th.

Pronotum elongate, 2.0 times longer than width at base, broadens
angularly, in posterior third, narrows gradually anteriorly and sharply
posteriorly, with straight basal flange, barely perceptible transverse groove
or without it near anterior margin, with dense erect rusty hairs, dense
reticulate punctation (punctures resemble facets separated by narrow
septa). Scutellum very small and narrowly rounded.

Elytra notably shorter than pronotum, in any case not longer, with
projecting shoulders, individually rounded apically, flat on disk, with
large punctation, and short semiadherent light-colored hairs. Legs with

Figure 42. Molorchus incognitus Tsher,

long erect or semiadherent hairs. Fore- and midfemora markedly dilat-
ed and hind femora moderately clavate. Hind tarsi about 0.50 length of
tibiae; 1st segment slightly shorter than two successive together.

Body ventrally with erect light-colored hairs. Abdomen narrows
slightly anteriorly, more so posteriorly. Abdominal tergites with large
punctation. Tergite V in female more elongate, rectangular, in male not
longer than width at base. Body black or blackish-brown. Antennae
and legs light rusty. Elytra light colored, pale yellow in anterior half,
dark brown, almost black at base and in posterior half. Body length
4.5 to 7.5 mm.

Egg\ White, elongate, narrows toward one pole. Chorion matte,
with fine sculpture. Length 1.0 mm, width 0.4 mm.

Larva (Figure 43): Characterized by sharp inner angle of hypostomal
sclerites and uniform anterior margin of epistoma. Head narrowly round-
ed anteriorly, insignificantly flattened. Epistoma fuses with parietals,
frontal sutures barely perceptible. Median suture slightly visible only in
76 posterior half. Anterior margin of epistoma smooth, uniform, with
barely visible lateral projection. Hypostoma narrows anteriorly, widely
separated medially by trapezoid gula into two sclerites with pointed
inner angles. Parietals in anterior half with long setaceous hairs forming

Figure 43. Larva of Molorchus incognitas Tsher.

a — head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla.

dense replicated tuft. Antennae comparatively long, with apices extend-
ing beyond 1st segment of maxillary palps. Clypeus very small, not
flattened, rectangular, convex. Labrum convex, broadly rounded apically,
and with sparse long setae on anterior margin. Mandibles comparatively
short, thick, smooth on outer side, convex, deeply hollowed inside, and
broadly rounded apically. Labial submentum with faint longitudinal
streaks.

Prothorax broad, 2.0 times wider than long, longer than meso- and
metathorax together, Pronotum markedly narrows anteriorly, broadly
rounded on anterior margin, with thin, not very dense hairs in anterior
half and laterally. Pronotal scutum moderately convex, white, glabrous,
demarcated laterally by deep longitudinal grooves. Prothoracic presternum
with sparse short hairs; eusternum slightly convex, without perceptible
hairs; base of prosternum (basisternum s. sternellum) in form of narrow
transverse fold, broadens slightly laterally and narrows medially.

Abdomen narrows posteriorly, broadens in region of segments VII
and VIII, with dense short light-colored hairs laterally. Dorsal loco-
motory ampullae convex and coriaceous, not sclerotized, separated by
common broad longitudinal groove, with small longitudinal dent later-
ally in anterior half. Ventral locomotory ampullae slightly convex, with
transverse groove in anterior half joining lateral longitudinal dent.
Anal (X) segment with dense long hairs. Body length of mature larvae
6.0 to 8.0 mm, width of head 0.7 to 0.8 mm.

Pupa (Figure 44): Body elongate. Head short, slightly bent under;
frons broadly impressed medially. Antennae tightly pressed to sides of
body, with apices slightly bent ventrad (toward base of hind femora).
Pronotum elongate, 2.0 times longer than wide, smooth, uniformly con-
vex, basally with narrow transverse groove, disk medially with group
of short hairs. Meso- and metanota glabrous, lustrous. Abdomen elon-
gate, broadens in region of segments IV and V, narrows anteriorly and
77 posteriorly, appears stretched anteriorly. Abdominal tergites moderately
convex, without spinules, only apex of tergite VII with pair of proximate
spinules bent forward. Valvifers of female small, slightly elongate. Body
length 6.0 mm (Cherepanov and Cherepanova, 1975).

Material: Collected in Ussuri-Primor’e region. Adult insects 24,
larvae three, pupa—one female (which developed into adult), larval
exuviae with beetles from cells 1 3.

Distribution: Ussuri-Primor’e region (Kovarovka and Suvorovka
Rivers).

Biology: Inhabits deciduous vegetation. Beetles fly in June and July,
disappear by mid-August. Rarely seen on flowers. Female oviposits in
bark crevices of thin shoots 0.6 to 1.5 cm in diameter. Larvae hatch
from eggs in two to three weeks and immediately bore into bark. They

Figure 44. Pupa of Molorchus incognitus Tsher.

initially live under bark, make longitudinal sinuous galleries, and plug
them with fine frass. Mature larva bores wood, makes longitudinal
gallery 6.0 cm long, and at end of gallery scoops out cell along trunk.
Length of cell 8.0 to 14.0 mm, width up to 3.0 mm. Pupation of larvae
in cells mainly observed in June. Young beetles emerge from pupae in
second half of June and early July. Beetles exit from wood end of June
and in July through oval openings in shoot surface up to 1.5 mm in
diameter. Pupal cell usually oblique to surface of trunk. Weight of
larvae before pupation 1.8 to 8.2 mg, pupae 1.5 to 7.2 mg, young beet-
les 1.2 to 6.0 mg.

Molorchus incognitusT&htt. develops on desiccated (sometimes viable)
shoots of various woody species. We raised 14 beetles from willow, one

100

78 from Euonymus, one from oak shoots, three from elm, and five larvae
and one pupa from Amur maple. Exocentrus stierlini Ganglb. was also
found with this species on thin willow branches (Cherepanov and
Cherepanova, 1975).

2. Genus Nadezhdiana Tsher.

(Tsherepanov) Cherepanov and Cherepanova, 1976, Novosti fanny
Sibiri {Nov. i maloizy. vidy founy Sibiri, lOth ed.), pp. 66-61.

Adult: Proximate to the genus Molorchus. Well distinguished by broad
mesosternal process and usually dense pubescence on ventral side of body.
Head not broader than pronotum. Antennae with 1 1 segments, shorter
than body in female, reaching only abdominal segment II; longer than
body in male, with 9th segment extending beyond tip of abdomen.
Forecoxae transversely elongate, forecoxal cavities open, prosternal pro-
cess long, narrow, cuneiform apically, curved at end. Midcoxae widely
separated, mesosternal process between them broad, flat, not narrowing
toward apex, almost square. Prosternum with large dense, metasternum
with minute, less dense punctation, and medially (slightly closer to
posterior margin) with smooth lustrous plate. Body on ventral surface
with minute, dense, adherent, silvery pubescence and dense erect light-
colored hairs.

Larva: In general appearance and presence of long replicated hairs
laterally on parietals of head similar to larvae of the genus Molorchus.
Readily recognized by tuft of sharp spinules on abdominal tergite IX.

Pupa: Body elongate. Pronotum with anterior and posterior flanges,
disk with setae forming two transverse bands interrupted medially by
small gap. Abdominal tergites II to VI with minute spinules forming
transverse row. Tergite VII with large (five or six) spinules directed
forward. Tergite VIII with large setae, framed at base by sclerotized
ringlet.

This genus, named after Nadezhda Epifanovna Cherepanova, com-
prises only a single species to date, and is ecologically associated with
broad-leaved forests.

Type species: Nadezhdiana villosa Tsherepanov, 1976.

1. Nadezhdiana villosa Tsher.

(Tsherepanov) Cherepanov and Cherepanova, 1976, Novosti fauny
Sibiri {Nov. i maloizv. vidy fauny Sibiri, 10th ed.), pp. 67-71.

79 Adult (Figure 45): Body elongate, dorsally flat. Head directed for-
ward, slopes slightly downward, barely narrower than pronotum, with
dense large punctation and long erect light-colored hairs, Frons broad,
flat, with median longitudinal suture, laterally with barely perceptible

101

longitudinal carina. Eyes large, highly convex, finely faceted, broadly
emarginate, lower lobes 3.0 times larger than upper. Antennae of male
long, slender, perceptibly longer than body, with elongate cylindrical
segments, angularly rounded, not extended on distal outer margin; 3rd
segment shorter than 5th, apex of 11th falcate. Antennae of female
markedly shorter than body, barely extend beyond abdomen, compara-
tively thick, with short adherent hairs not forming compact cover; inner
side of segments (especially 3rd to 6th) with long hairs; 3rd segment
slightly longer than 4th, but definitely shorter than 5th; 5th to 10th

Figure 45. Nadezhdiana villosa Tsher.

102

matte, extended, spinelike, on distal outer margin; 11th segment apically
pointed conically.

Pronotum elongate, more so in male, broadens medially in female,
flattened on disk, near anterior and posterior margins with broad trans-
verse flange, curves angularly on sides, with deep large mammilary
punctation; spaces between punctures smaller than punctures; anterior
80 half laterally with dense adherent squamiform hairs forming two whitish-
silver (female) or yellowish (male) spots; basally on flange with tender
adherent white hairs forming light-colored transverse band; disk and
sides with long, erect, setaceous, brownish hairs; anterior half laterally
in male with faint longitudinal lustrous elevation. Scutellum elongate,
with parallel sides, broadly rounded apically, with dense adherent sil-
very hairs. Elytra not longer or slightly longer than pronotum, with
humeri projecting forward, flat lustrous disk, sparse fine punctation and
sparse, short, light-colored, semiadherent hairs, individually rounded
apically, with distinct narrow border. Legs long, with long thin setaceous
hairs; femora clavate, forefemora apically highly and abruptly dilated,
hind ones dilate gradually. Hind tibiae slender, curve slightly, not shorter
than hind femora. Hind tarsi 0.40 to 0.50 length of tibiae; 1st segment
barely longer than two successive together.

Body ventrally with dense, minute, adherent, silvery and long erect
light-colored hairs. Abdomen elongate, with parallel sides, narrows
posteriorly. Abdominal sternites bulge, lustrous, with fine punctation.
Sternite V short, broad, narrows posteriorly, rounded at end, without
squamiform hairs in female. Body black. Antennae rusty, somewhat
darkened apically. Elytra black, disk with whitish translucent spot,
anterior margin angularly produced, posterior margin slightly rounded.
Length 9.0 mm.

Larva (Figure 46): Differs from larvae of all other species in presence
of area on tip of abdomen covered with spinules (dorsal view). Body
slightly elongate, with almost parallel sides. Half of head retracted into
prothorax; head markedly narrows anteriorly. Anterior margin of parie-
tals with short hairs directed backward. Epistoma lustrous, laterally
with indistinct frontal sutures, medially divided by distinct longitudinal
suture. Hypostoma divided into two triangular sclerites displaced pos-
teriorly by gular plates. Clypeus small. Labrum very small, elongate,
with longitudinal streaks on disk, sparse short setae on anterior margin.
Mandibles thick, short, broadly rounded apically, hollowed from inside,
cultrate, horseshoe shaped. Pronotum narrows slightly (roundly) anterior-
ly, with long thin hairs forming transverse row on anterior margin close
to sides, and tender uneven hairs laterally. Pronotal scutum convex,
coriaceous, with minute furrows, bound laterally by long longitudinal
folds reaching almost to anterior margin. Thoracic legs lacking.

103

79 Figure 46. Larva of Nadezhdiana villosa Tsher.

a— head and pronotum; b— abdominal tergite IX.

Abdomen with parallel sides, narrows somewhat on segments VIII
and IX. Dorsal locomotory ampullae slightly convex, medially divided
by common longitudinal groove, laterally with longitudinal fold, on
anterior angles with longitudinal groove. Ventral locomotory ampullae
almost identical in structure. Tergite IX convex, laterally with long thin
hairs, smooth lustrous disk on which sharp acicular erect spinules (about
50) form broad field. Body length 9.0 mm, width of head 1.4 mm.

Pupa (Figure 47): Body elongate, slender. Head bent under. Genae
short, 0.33 width of lower ocular lobes. Antennae pressed to sides, bent
ventrad; 11th segment adjoins elytral apex. Frons flat, with median
longitudinal suture, near antennal bases and medially with three pairs
of setae. Occiput smooth, glabrous, spherical. Pronotum elongate, with
broad flange on anterior margin and sharp narrow one at base; disk
81 flat, anterior and posterior to middle near base with dense setae forming
two transverse bands interrupted medially. Mesonotum medially and
metanotum apically with stray minute setae.

104

79 Figure 47. Pupa of Nadezhdiana villosa Tsher.

Abdomen broadens in region of segments IV and V and narrows
toward base and apex. Abdominal tergites moderately convex, in poste-
rior half with minute acute spinules forming transverse row (four to six
paramedial spinules). Tergite I glabrous, without spinules; II with faint
' spinules; VII apically with much larger spinules (up to six) bent down
and forward and forniing transverse row. Tergite VIII narrow, elongate
in posterior half, with long setae directed backward forming two tufts
(up to 10 setae per tuft) that diverge at an acute angle. Valvifers of
female hemispherical, with projecting apical tubercle. Body length 8.5
mm, width of abdomen 1.8 mm.

Material: Collected in Ussuri-Primor’e region (Novomoskovka village).
Adult insects three, larvae two, pupa — one female, larval exuviae with
beetles from cells two. . '

105

Distribution: Ussuri-Primor’e region.

Biology: Inhabits broad-leaved forests. Ecologically associated with
Manchurian walnut (Juglans manshurica). Female oviposits on decompos-
ing shoots 1.8 to 3.0 cm in diameter. Larvae make longitudinal sinuous
galleries under bark, deeply impressed on sapwood, and plug them
densely with frass. Larva later bores wood up to 5.0 mm, makes cell
there along shoot, turns head toward inlet, and pupates. Length of
gallery under bark up to 8.0 cm, width before boring wood 3.5 mm.
Width of inlet up to 3.0 mm. Length of pupal cell 16 to 18 mm, width
3.0 to 4.0 mm. Pupae develop at room temperature in about 20 days.
Young beetles nibble oval opening (2.0 mm X 3.0 mm to 3.0 mm X 3.5
mm) on shoot surface and exit from pupal cell through it. Weight of
larvae before pupation 13 to 18 mg, pupae up to 15 mg, and beetles
after exiting from wood 7.0 mg or more. Pupation observed in spring.
Young beetles seen in June, fly in July (Cherepanov and Cherepanova,
1976).

19. Tribe DILUSINI

Adult insect characterized by minute narrow body. Differs from
proximate tribes (Gracilini, Obriini, Nathriini, and Molorchini) in these
features. Eyes highly emarginate. Pronotum elongate, with broad flange
along posterior margin. Elytra elongate, cover entire abdomen dorsally,
flat on disk. Abdominal sternite I moderately elongate, shorter than rest
together.

Larva small, body elongate. Parietals laterally in anterior half with
usual hairs, not bent backward, with pair of hyaline or pigmented
ocelli near antennae ventrally. Thoracic legs well developed.

Pupa recognized by minute elongate body. Head narrow. Antennae
bent ventrad behind midfemora, semicircular there. Spinules on pronotum
and abdominal tergites mainly bent forward. Tergite VII narrowly round-
ed apically, in posterior half with two pairs of spinules forming
transverse row.

This tribe is represented in USSR fauna, including northern Asia,
by lone monotypic genus.

1 . Genus Deilus Serv.

Serville, 1834, Ann.Soc. Entom. France,vo\, 13; Plavifshchikov,
1940, Fauna SSSR, 22, 2, 190 (Dilus); Mamaev and Danilevskii, 1975,
Lichinki zhukov-drovosekov, pp. 188-189 (Dilus).

Adult: Differs in very narrow, comparatively flat body. Head elong-
ate anteriorly, flat between antennae and upper ocular lobes, with large

106

longitudinally elongate punctation. Antennae short, thick, just reach
(male) or notably fall short of (female) middle of elytra, from 5th
segment with distinctly produced outer angle. Pronotum elongate, with
narrow flange at base and broad one at apex. Elytra narrow, almost
with parallel sides, flat on disk, with lateral carinate streaks extending
from ventral side of humeral tubercle to hind clivus. Legs short, femora
clavate.

Larva: Characterized by elongate slender body. Head narrows ante-
riorly, with smooth rusty border on anterior margin of epistoma, narrow
notch near clypeus medially. Gula projects beyond anterior margin of
hypostoma, broadens here, with anterior angles produced laterally to
form, so to speak, common gular-labial complex with labium. Pronotal
scutum short, transversely straightly truncate on posterior margin. Pro-
thoracic presternum with sparse setaceous hairs, sharply ringed at base
with sclerotized ringlet. Legs short, minute. Locomotory ampullae deve-
loped on abdominal segments I to VII, separated by transverse groove,
with uneven, finely granular or granulate sculpture forming transverse
elongate ellipse with silvery sheen.

Pupa: Readily recognized by a combination of characters. Head
narrow, extends narrowly and roundly on occiput. Antennae short,
apices bent ventrad, Pronotum medially, hind clivus, and abdominal
tergites I to III with several spinules forming transverse row. Meso-
and metanota glabrous, lustrous, without spinules. Legs short; femora
clavate; apices of hind femora barely reach posterior margin of tergite
III.

This genus consists of a single species, distributed sporadically in
central and southern Europe.

Type species: Callidium fugax Olivier, 1790.

1. Deilus fugax (Oliv.)

Olivier, 1790, Enc. Meth. Ins., vol. 5, p. 253 (Callidium); PlaviT-
shchikov, 1940, Fauna SSSR, 22, 2, 190-192 (Dilus); Demelt, 1966,
Tierwelt Deutschlands, vol. 2, pp. 63-64.

Adult (Figure 48): Frons slightly tubercularly elevated laterally near
antennal bases, with narrow median longitudinal groove extending onto
vertex. Eyes sharply faceted, emarginate up to posterior margin; gap
between ocular lobes very narrow, only with stray facets in one row.

83 First segment of antennae thick, with large flat punctation; remaining
segments with fine punctation. Second antennal segment nodular, slightly
wider than long; 4th segment barely longer than 3rd, equal to 5th or
insignificantly shorter.

Pronotum broadly rounded laterally, 1.5 times longer than width at
base, with deep coarse punctation, spaces between punctures markedly

107

Figure 48. Deilusfugax (Oliv.).

smaller than punctures, with gray adherent and erect hairs. Scutellum
broad, broadly rounded posteriorly, with dense, light-colored, compactly
adherent hairs.

Elytra with round projecting humeri, inner angle apically produced
spinelike or angularly rounded, and outer angle broadly rounded (hence
84 appears slightly emarginate near inner angle), with dense large deep
punctation, shagreen coarse sculpture in spaces between punctures, with
tender gray hairs combed sideways on disk and from base to apex later-
ally. Prosternum with dense punctation; prosternal process long, with
parallel sides, extends beyond posterior margin of forecoxae and bent
down there, with large punctation. Femora clavate, in second half highly
thickened as though dilated, with sharp flange apically (ventral view).
Ventral surface of body with dense gray adherent hairs. Body black with
bronze iridescence. Antennae variegated; 1st antennal segment black,
rest reddish-rust, at apex (especially from 5th segment) black. Legs
reddish-rust, femoral clava and apex of tibiae black, tarsal segments
darkened apically. Body length 7.0 to 10.0 mm.

108

Egg: White with green tinge, elongate, narrowly rounded at poles, nar-
rower toward posterior pole. Chorion with minute reticulate sculpture,
cells deep, flat, with spaces between them somewhat larger than cells.
Length 1.0 mm, width 0.3 mm.

Larva (Figure 49): Head slightly retracted into prothorax; general
contour appears rounded anteriorly. Epistoma laterally fused with parie-
tals (frontal sutures not visible), divided by faint median longitudinal
suture. Hypostoma with rusty border on anterior margin, narrows anter-
iorly, laterally with straight sutures, obtusely rounded at anterior outer
angles, and narrows markedly posteriorly along inner margin of sclerites.
Gula broadens markedly toward base, emarginate anterolaterally. Parie-
tals in anterior half with innumerable setaceous hairs, near antennal
bases with pair of small proximate pigmented or hyaline ocelli. Antennae
short, narrow gradually toward apex; 1st segment of antennae not longer
than two successive together. Clypeus very small, trapezoid, brownish-
rust. Labrum convex, rounded anteriorly, narrows slightly toward base,
with light rust setae in anterior half Mandibles apically broadly rounded,
black, basally rusty-red, with transverse groove on outer side. Inner mas-
ticatory lobes of maxillae thick, apically obtuse, with short thin setae.
Maxillary palps insignificantly longer than inner lobes, laterally with
long setae, with digitate apical segment rounded terminally.

Pronotum almost 3.0 times wider than long, in anterior third with
narrow transverse yellow band that is separated medially and laterally by
narrow white clearance into four spots, which are either distinct or faint,
sometimes quite diffuse. Sides and disk of pronotum anterior to scutum
with dense long light rust hairs, anterior margin of yellow band with stray
hairs somewhat clustered at base and forming transverse row. Pronotal
scutum short, white, 4.0 times wider than long, bound laterally by long
longitudinal grooves, straightly truncate on anterior margin, medially and
at anterior angles amost not produced forward, basally with minute reti-
culate squamose sculpture imparting silvery sheen, elsewhere with longi-
tudinal striations; spaces between striations lustrous. Prothoracic prester-
num with rather sparse setaceous hairs ringed at base with sclerotized
ringlet; eusternum glabrous or with stray short hairs, merges with prester-
num. Grooves between sterna imperceptible, rarely visible. Thoracic legs
short, with completely or poorly sclerotized minute apical claw.

Abdomen elongate; segments VI and VII broaden slightly, laterally
85 with short thin light-colored hairs. Abdominal tergites laterally in anterior
half with deep flange. Dorsal locomotory ampullae moderately convex,
with transverse groove, uneven granular sculpture; some granules very
large, lustrous, and form two rather distinct rows separated by transverse
groove. Ventral locomotory ampullae similar in structure. Body length of
mature larvae 10 to 12 mm, width of head 1.8 mm.

109

Figure 49. Larva of Deilus fugax (Oliv.).
a — head and pronotum; b — abdominal tergite IV with locomotory ampulla;
c — tip of abdomen (ventral view).

Pupa (Figure 50): Body elongate, narrow. Head small, markedly
narrower than pronotum, near antennal bases slightly convex from inner
side, with median longitudinal groove, narrowly (as though produced)
rounded on occiput, lustrous, without hairs and without spinules.
Antennae short, apices bent ventrad posterior to midfemora.

Pronotum 1.5 times longer than width at base, posterior to middle
broadly rounded laterally, narrows more anteriorly, less posteriorly,
basally with narrow flange; disk convex, with faint tender transverse
streaks, medially with more, sometimes less distinct spinules forming two

Figure 50. Pupa of Deilusfugax (Oliv.).

Ill

paramedial tufts; hind clivus with very large sharp spinules forming in-
distinct transverse row broadly interrupted medially (usually four para-
medial spinules). Mesonotum convex, lustrous, glabrous, with slightly
hollow scutellum projecting posteriorly. Metanotum broad, with in-
distinct median longitudinal groove, with faint transverse streaks, poste-
rior margin broadly rounded, glabrous, without spinules. Legs short;
hind femora just reach or do not reach posterior margin of tergite III.

Abdomen markedly elongate, broadens slightly in region of segment
IV, narrows from there gently anteriorly, more distinctly posteriorly.
Abdominal tergites uniformly convex, in posterior half with well- develop-
ed minute or barely perceptible acute spinules directed backward; these
spinules form broad transverse row interrupted medially (three paramedial
spinules). Tergite VII not longer than width at base, tapers angularly in
posterior quarter, narrowly rounded apically, convex and lustrous on
disk; posterior to middle with two pairs of very large sharp spinules
directed forward and forming transverse row. Tergite VIII lustrous,
semitransparent, rounded apically, near posterior margin with thin trans-
verse furrows and deep longitudinal furrows posterolaterally. Valvifers
of female hemispherical, small, apically resemble lateral mastoid projec-
tions. Body length 7.0 to 10.0 mm, width of abdomen 2.1 mm.

Material. Collected in the southern Urals, north of Ural’sk (Embu-
latovka River). Adult insects 39, of which 26 raised in laboratory from
larvae, larvae 41, pupae — five males and six females, larval exuviae with
pupae from cells two.

Distribution: From Sverdlov, Upper Volga to the Caucasus, from the
Urals to the Ukraine inclusive, central and southern Europe, Turkey,
northern Africa. Sporadic everywhere.

Biology: Inhabits arid sections of forest-steppe zone. Ecologically as-
sociated with shrub (mainly legumes) vegetation. Beetles fly from May
to mid-July, seen on flowers of Umbelliferae, spiraea, and other plants.
Later colonize shoots of Cytisus, Saratharnnus, and other shrubs. We
found them in the steppe zone 80 km north of Ural’sk (Embulatovka
Rivers) on Russian broom (Cytisus ruthenicus).

Larvae live under bark of thin shoots 0.3 to 2,5 cm in diameter, make
longitudinal sinuous galleries, plug them densely with frass, which re-
86 mains covered externally by thin bark film. Galleries under bark on thin
shoots up to 1 .0 cm in diameter, deeply impressed in wood; walls steep,
sometimes acute. On very thick shoots up to 2.5 cm in diameter,
especially near root zone, galleries faintly impressed in alburnum, with
smooth walls. Length of gallery under bark 15.4 to 23.5 cm, width 0.2 to
0.5 cm. Mature larva gradually bores deeper into wood and makes hollow
gallery 2.5 to 4.5 cm long in upper layer along shoot, with cell at end;
gallery sealed with plug of coarse fibrous frass. Width of inlet 0.2 to 0.3

112

cm, length of smooth gallery anterior to cell up to 2.4 cm, length of plug
sealing cell from gallery 0.2 to 0.7 cm; length of cell 1.2 to 2.0 cm,
width 0.25 to 0.30 cm. In a shoot 0.9 to 1.0 cm in diameter, one larva
made a gallery 23.5 cm long under bark from top downward, widened
it at end from 0.3 to 0.6 cm, turned back, bored deeper gradually from
bottom upward in wood, made there a hollow gallery along shoot, pre-
pared cell at its end, and sealed it from smooth gallery with plug of
fibrous frass. Larva pupates with head toward inlet, facilitating beetle’s
exit from wood later.

Pupation commences after first hibernation in middle 10 days of July
and ends in first half of August. On July 19 broom shoots cut open
yielded 24 insects, of which nine (37.5%) were larvae before pupation
and 15 (62.5%) pupae. Duration of pupal stage in nature at 14.4 to
31.4°C (23.0 + 0.7®C) 17 to 19 days, average 18.1 + 0.1 days. Young
beetles sighted in first 10 days of August, enter diapause, hibernate in
cell. Emergence of young adults from wood completed in spring with the
onset of warm weather. By this time gonads have matured. Ovaries of
one female dissected after emergence from wood contained 36 mature
eggs. Weight of insects in larval stage before pupation 10.0 to 26.5 mg
(19.0 + 0.8), pupae 9.0 to 24.0 mg (16.9 + 0.7), and adult insects be-
fore hibernation 8.0 to 20.0 mg (13.7 + 0.6). Weight of insects decreased
by an average of 27.8% during metamorphosis, more in some indivi-
duals. Generation completed in two years (Table 6). Hibernates first
time in second larval instar under bark or in wood (weight of larvae at
this time 8.4 to 14.0 mg, average 12.0 + 0.6 mg) and second time as
adult in wood. In 1978 we collected a large number of dead beetles from
pupal cells subsequent to hibernation. This indicates that during hiber-
nation the population is considerably reduced in the adult stage.

Table 6. Periods of development of Deilus fugax (Oliv.)

Year of

development

April

May

June

July

August

September

1st

AEL

2nd

LPA

3rd

AEL

Population density in broom shoots was comparatively high. We
found four hibernating beetles in shoots 4.5 cm long, 1.0 cm in diameter.
Wood between pupal cells was preserved in form of narrow septae. Broom
shoots damaged by larvae of Deilus fugax (Oliv.) have withered by the
time the second or even first hibernation takes place. Chlorophorus varius

113

87 (Miill.) is often found together with this species on the same shoots; the
latter species colonizes the upper and lower parts of broom and the
former only the lower parts near the roots. The larvae of C. varius live
mainly under bark, penetrating wood only for pupation, while D. fugax
lives in wood and makes long galleries there. For example, in autumn
seven larvae of Deilus fugax (Oliv.) and four larvae of Chlorophorus
varius (Mull.) were recovered from a section of broom root (length 24.0
cm, diameter 2.25 cm). Larvae of the latter were found under bark and
in wood and of the former only in wood.

20. Tribe CALLICHROMINI

Adult insect distinguished by elongate, notably flat (Aromia, Cheli-
donium) or cylindrical (Polyzonus) body. Antennae long, extend beyond
tip of abdomen {Aromia, Chelidonium) or slightly shorter than body
(Chloridolum). Pronotum laterally with sharp {Aromia, Chelidonium) or
obtusely rounded {Chloridolum) tubercle; disk with obliterated {Aromia)
or sharp and deep {Polyzonus) punctation, or thin transverse furrows
{Leontium). Metasternum posterolaterally with projecting ellipsoidal
aromatic pore. Elytra monochromatic, metallic green, blue, or violet,
with partly bronze iridescence {Chloridolum, Aromia) or blue with broad
transverse bands {Polyzonus).

Larva with half or more of head retracted into prothorax. Epistoma
medially divided by narrow longitudinal suture, merges laterally with
parietals, frontal sutures imperceptible or faint. Pronotum with dense
{Aromia, Chelidonium, Leontium) or sparse setaceous {Chloridolum) hairs,
forming two transverse fields, one just anterior to scutum, the other in
anterior third. Pronotal scutum convex, coriaceous, demarcated laterally
by deep groove. Prothoracic eusternum glabrous, coriaceous merges with
general surface of presternum, not demarcated from it by groove. Thor-
acic legs small, with minute claw. Locomotory ampullae coriaceous, not
sclerotized, developed on abdominal segments I to VII, separated dorsally
by two and ventrally by one transverse groove.

Pupa with glabrous, lustrous head, with longitudinal groove between
antennae. Antennae in second half looplike, pressed to sides ventrally.
Pronotum laterally with large {Aromia, Chloridolum) or minute {Leontium)
tubercle, in posterior half with paramedial setae set on protuberances
and forming transverse band, medially on disk with setae arranged in
transverse row or narrow transverse band, on anterior margin with seta-
ceous tubercle {Chelidonium), transverse glabrous carina {Chloridolum),
or flat and without setae {Aromia). Meso- and metanota glabrous, lustrous
{Aromia, Chloridolum, Leontium), or with large setae forming two tufts
on each {Chelidonium). Abdominal tergites with very small spinules
generally bent down and forward.

114

Callichromini are abundantly represented in Southeast Asia but im-
poverished in northern Asia. They occur here only in the southern
regions, with five genera represented by one species each. Most species
are ecologically associated with deciduous species; only one species
(Leontium viride Thoms.) lives on spruce shoots and preferentially colo-
nizes viable trees.

KEY TO GENERA

Adult Insects

88 1 (8). Body slightly flat. Pronotum broad, uneven on disk, with sparse
smoothened or dense highly minute punctation, or fine transverse
furrows sometimes forming concentric circles. Elytra monochro-
matic, metallic blue, green, or violet.

2 (3). Hind femora short, far from reaching elytral apices, maximally

extend up to 0.66 their length. Palearctic 1 . Aromia Serv.

3 (2). Hind femora long, extend beyond elytral apices or slightly short

of them (Chelidonium Thoms.).

4 (7). Body large, 22 to 35 mm long.

5 (6). Pronotum red, laterally with large, smooth, lustrous, conical

tubercle pointed at end. Mainly Southeast Asia

2. Chloridolum Thoms.

6 (5). Pronotum green, laterally with obtuse, densely punctate tubercle;

disk with dense, highly minute punctation. Southeast Asia

3. Chelidonium Thoms.

7 (4). Body 14 to 18 mm long. Pronotum with fine transverse furrows.

Body and elytra bright green. Southeast Asia

4. Leontium Thoms.

8 (1). Body cylindrical. Pronotum comparatively narrow, distinctly elon-

gate, evenly and highly convex on disk, with dense large deep
punctation. Elytra with broad transverse yellow bands. East Asia.
5. Polyzonus Cast.

Larvae

1 (8). Abdominal sternite X entirely covered with hairs. Apex of gula

narrow, about 0.25 width of hypostomal sclerite on anterior
margin.

2 (7). Dorsal locomotory ampullae reticulately rugose, not matte. Found

on deciduous species.

3 (6). Hairs on pronotum fine, not dispersed; distance between hairs less

than their length. Hairs on pronotum equally dense posterolater-
ally and in anterior half.

115

4 (5). Abdominal tergite IX glabrous on disk, laterally and on posterior

margin with sparse hairs. Found on willow. ..... 1. Aromia Serv.

5 (4). Abdominal tergite IX with sparse hairs on disk but dense hairs

laterally and on posterior margin. Found on Manchurian walnut.
.............................. 2. Chloridolum Thoms.

6 (3). Hairs on pronotum setaceous, thick, dispersed; distance between

them not less than or even more than their length. Hairs on pro-
notum dense posterolaterally, sparse in anterior half. Found on
maple shoots at crown of thick-trunked viable trees. .........

.. ............................ 3. Chelidonium Thoms.

7 (2). Dorsal locomotory ampullae not reticulately rugose, smooth,

matte. Found on coniferous species. ...... 4. Leontium Thoms.

8 (1). Abdominal sternite X glabrous, with thick acicular setae only on

posterior margin forming transverse row. Apex of gula broad,
0.33 anterior width of hypostomal sclerite. Found on dog rose.
5. Polyzonus Cast.

Pupae

1 (8). Abdominal tergites with spinules forming transverse band that

does not curve forward laterally and is not circular.

2 (7). Hind femora reach large abdominal tergite VII (Male Chloridolum

sieversi Ganglb.), do not extend beyond its apex.

3 (4). Pronotum without tubercle and without carina on anterior margin,

uniform or sometimes slightly elevated there, glabrous, without

setae 1. Aromia Serv.

89 4 (3). Pronotum with tubercle or carina on anterior margin.

5 (6). Pronotum with well-developed carina on anterior margin. Meso-

and metanota glabrous, without setae

............................... 2. Chloridolum Thoms.

6 (5). Pronotum with sharply projecting setaceous tubercle on anterior

margin. Meso- and metanota laterally with large acicular setae
forming tuft. .................... 3. Chelidonium Thoms,

7 (2). Hind femora reach abdominal tergite VII (female) or extend

beyond its apex (male). ............... 4. Leontium Thoms.

8 (1) Abdominal tergites with spinules forming transverse band that

curves forward laterally or is circular ...... 5. Polyzonus Cast.

1. Genus Aromia Serv.

Serville, 1833, Ann. Soc. Entom. France, vol. 2, p. 359; PlaviFshchikov,
1940, Fauna SSSR, 22, 2, 196-”197; Gressit, 1952* Longicorn Beetles of
China, vol. 2, p. 200; Mamaev and Danilevskii, 1975, Lichinki zhukov-
drovosekov, pp. 189-191.

116

Adiilv. Body large, somewhat flat. Antennae not longer (female) or
markedly longer than body. Pronotum laterally with large conically pro-
duced tubercle. Elytra elongate, with parallel sides, narrowly rounded
apically, glabrous (subgenus Aromia s. str.) or with short hairs (subgenus
Tomentaromia Plav.). Apex of sternite V emarginate (male) or rounded
(female). « ' ■ ^ :

Larva: Characterized by presence of hyaline, nonpigmented ocellus
at antennal bases, short rusty hairs on pronotum forming two transverse
fields anterior to scutum, and developed nonsclerotized locomotory
ampullae on abdominal segments I to VII.

Pi4pa: Body large, flat. Antennae long, looplike, curving forward at
level of middle or beyond apex of hind femora. Abdominal tergites
with innumerable short spinules directed mainly anteromedially. Apex
of abdominal tergite VII with minute spinules forming tuft.

Genus widely distributed in Eurasia. One species inhabits northern
Asia, its areal covering almost the entire Palearctic.

Type species: Cerambyx moscholifs Linnaeus, 1758.

1. Aromia moschata (L.)

Linnaeus, 1758, Syst. Nat., 10th ed., p 39 (Cerambyx); PlaviPshchi-
kov, 1940, Fauna SSSR, 22, 2, 198-204; Cherepanov and Cherepanova,
1975, Zhuki-drovoseki ivovykh lesov Sibiri, pp. 87-90.

Adult (Figure 51): Characterized by large body and elytra long, with
parallel sides. Head glabrous, lustrous, without hairs, with uneven coarse
punctation, bulges transversely between antennae, and with median longi-
tudinal suture. Eyes finely faceted, broadly emarginate. Antennae slender;
9th segment extends beyond apex of elytra (male) or does not reach it
(female); longitudinal groove on outer side commences from 4th segment.
First antennal segment thick, considerably shorter than 3rd, with acutely
projecting outer distal angle.

Pronotum transverse, laterally with large tubercle that projects
acutely, spinelike, narrows to same degree anteriorly and posteriorly, near
anterior and posterior margins with faint transverse groove; disk convex,
uneven, with rugose punctation, glabrous, with stray, slightly obliterated,
minute hairs laterally. Scutellum triangular, acutely or narrowly round-
90 ed apically, with minute, sometimes rugose punctation. Elytra elongate,
almost 4.0 times longer than wide, with dense rugulose punctation; with
faint longitudinal carinae or without theni, disk slightly convex, almost
flat, apically with narrowly rounded inner angle and tapering outer,
glabrous, without hairs. Metasternum with sparse deep punctation, dense
adherent light- colored hairs and sparse semiadherent brownish hairs.
Aromatic pores open, metepisternum around them distinctly impressed.
Legs long, slender. Hind femora curve notably. Hind tarsi shorter than

117

Figure 51. Aromia moschata (L.).

tibiae; 1st segment not longer than two successive together, arolia with
broad median longitudinal groove. Abdominal sternite V emarginate
(male) or narrowly rounded (female; at apex. Entire body and elytra
bluish or greenish with bronze metallic iridescence [A. m. moschata (L.)]
or pronotum red, and dark transversely rugose border present at base
and apex {A. m. orientalis Plav.). Body length 23 to 34 mm.

Egg: White, comparatively thick, obtusely rounded at poles. Chorion
with fine cellular-reticulate sculpture. Cells flat, fringed with thin lines.
Length 2.8 mm, width 1.0 mm.

118

Larva (Figure 52): Readily recognized by short rusty hairs on pro-
notum, structure of dorsal locomotory ampullae, and abdominal tergite
IX. Head narrows moderately anteriorly. Anterior margin of epistoma
with broad rusty-brown border, behind which pair of long setae occur,
laterally with single long seta, medially with longitudinal sharp suture,
laterally with faint frontal sutures. Hypostoma slightly convex, on an-
terior margin around inner angles with sharp, highly extended projec-
tions, and inward to them deep notch accommodating spinelike articulate

91 Figure 52. Larva of Aromia moschata (L.).

a — head and pronotum; b — abdominal tergite IV with
dorsal locomotory ampulla; c — abdominal segments III
to V of I-instar larva.

119

process of maxillae. Clypeus small, narrowly trapezoid, white. Labrum
convex, narrowly or broadly rounded apically, white, along margins with
dense rusty setae. Mandibles massive, basally flat on outer side or with
longitudinal groove. Parietals in anterior half with sparse, near frontal

91 sutures short hairs forming single row. Antennae with four segments.
One hyaline ocellus near each antennal base.

Pronotum slopes markedly toward head, rounded laterally, with
white glabrous (hairless) border on anterior margin, behind which trans-
verse yellow band occurs (with two deep saccate white notches antero-
laterally and longitudinal white clearance medially), with short rusty
hairs on sides and disk forming two transverse fields, one anterior to
scutum, the other on anterior edge of yellow transverse band. Pronotal
scutum convex, with two emarginations on anterior margin, with short
lateral longitudinal groove, white, coriaceous, with stray setaceous
hairs. Alar lobes with dense rusty hairs. Prothoracic presternum and
disk with sparse hairs, laterally with dense very long hairs; eusternum
glabrous, merges with general surface of presternum. Legs fully developed,
with sharp claws.

Abdomen laterally with short rusty hairs. Dorsal locomotory am-
pullae slightly convex, coriaceous, with faint furrows, separated by two
transverse grooves (anterior one usually straight, posterior one notably

92 convex), with longitudinal groove medially and longitudinal folds later-
ally. Ventral locomotory ampullae demarcated laterally by short longi-
tudinal grooves joined by transverse replicate groove. Abdominal tergite
IX glabrous on disk. Body length of mature larva 30 to 43 mm, width
of head 3.2 mm. I-instar larva with one spinule laterally on each side
of abdominal segments III to V, which disappear after first molt.

Pupa (Figure 53): Readily recognized by large body and smooth
anterior pronotal margin devoid of setae. Body comparatively flat. Head
glabrous, between antennae transversely convex, with broad transverse
fold on vertex, and broadly rounded on occiput. Antennae pressed to
sides of body, bent ventrad, looplike in female at level of middle of hind
femora, beyond apex of hind femora in male.

Pronotum slightly convex on disk, uneven, on anterior margin with-
out tubercle or carina, laterally with large conically produced tubercle
and near it, closer to base, with tubercular elevation bearing spinules;
inward to latter spinules form small tuft (sometimes tuft dispersed into
band adjoining spine-bearing tubercular elevation). Mesonotum convex,
glabrous, with transversely rugose scutellum produced posteriorly. Meta-
notum broad, almost flat, glabrous.

Abdomen slightly convex, broadens in region of segments III and
IV, narrows insignificantly anteriorly and more abruptly posteriorly.
Abdominal tergites medially with longitudinal groove, laterally in pos-

Figure 53. Pupa of Aromia moschata (L.).

terior half with sharp short spinules bent anteromedially. Apex of ter-
gite VII narrowly rounded, with spinules usually forming broad trans-
verse field. Tip of abdomen (dorsal view) with perceptible fork (female)
or narrowly rounded, without fork (male). Valvifers of female hemi-
spherical, proximate. Body length 25 to 35 mm, width of abdomen 8.0
to 10.0 mm.

Material: Collected in Altai, Ob’ region, Tuva, Trans-Baikal, Ussuri-
Primor’e region, and Sakhalin. Adult insects 48, larvae 43, pupae — one
male, nine females, larval exuviae with beetles from cells three.

Distribution: From Atlantic to Pacific Ocean coasts, south from north-
ern Africa, northern Kazakhstan, Altai, northern Mongolia, northern
China, and Japan, north almost up to and inclusive of taiga zone. Two
subspecies inhabit northern Asia: Range of m. moschata (L.) extends
from Europe to Baikal, of A. m. orientalis Plav. expanse east of Baikal.

i2i

Biology: Inhabits willow and mixed forest plantations containing
willows. Covers mainly river valleys, terraces of hill rivers, hill sections
along rivers, etc. Beetles fly in July and first half of August. Sighted in
maximum numbers in second half of July on flowers of Umbelliferae, Ro-
sales, and other plants. In 1968 and 1969 in Salair 123 beetles were collect-
ed: 97(78.9%)in July and 26 (21.1%) in August. Beetles feed on flowers,
then fly to viable willows, mate there, and oviposit in bark crevices on
lower part of trunk. Ovaries of one female contained 25 eggs. Weight
of freshly laid eggs 1.5 mg. According to our observations, one female
can lay up to seven eggs in 25 minutes. Oviposition selective; in gardens
only freshly cut willows were chosen and all other plant species rejected.
Eggs are laid in nature from July to early October. Incubation period
at 18.5°C continued for 20 to 26 days, average 22.3 days; at much higher
temperatures incubation period reduced to two weeks.

Larvae rupture chorion to hatch, bore into bark, and live under it
93 for sometime. Later bore deeper into wood, make long longitudinal,
sometimes sinuous galleries from bottom upward. Here bark surface
moistened with dark brownish spots of exuding sap. Gallery length 40
cm, width up to 13 to 18 mm. Upper, larger part of gallery not plugged
with frass, remains hollow. Larvae live in moist wood of growing trees.
Later, not before third hibernation, each larva nibbles cell in wood along
trunk, demarcates it from hollow part of gallery by plug of fibrous frass,
and pupates with its head downward Length of pupal cell up to 50 mm,
width 10 to 15 mm.

Pupation mainly in June. Pupae found up to first few days of July.
Three weeks later young beetles emerge from pupae. Emergence of young
adults from wood commences in first 10 days of July and ends in early
August. For example, we raised 96 beetles in the forest. Of these, two
(2.0%) emerged from wood in first 10 days of July, 16 (16.7%) in second
10 days, 62 (64.6%) in last 10 days, and 16 (16.7%) in early August.
Weight of larvae before pupation 667 to 825 mg, pupae 516 to 750 mg,
and young beetles before emerging from cells 336 to 525 mg. One larva
(male) before pupation weighed 605 mg (100%), pupa developing from
it 526 mg (86.9%), and beetle before emerging from wood 362 mg

Table 7. Periods of development of Aromia moschata (L.)

Year of

development

May

June

July

August

September

October

1st

PAE

AEL

2nd-3rd

4th

PAE

AEL

122

(59.8%). Corresponding weights in another case (female): 766 mg (100%),
718 mg (93.7%), and 460 mg (60.0%). Generation completed in three
years (Table 7). Aromia moschata (L.) inhabits only trunks of viable
trees, mainly root section with diameter up to 10 cm and above. Not
seen on desiccated trees.

2. Genus Chloridolum Thoms.

Thomson, 1864, Syst. Ceramb., vol. 174, p. 420; Plavii’shchikov,
1940, Fauna SSSR,22, 2, 207-208.

Adult: Body large. Antennae markedly longer than body. Pronotum
laterally with large conical tubercle, in posterior half with pair of widely
separated, transversely elongate, smooth tubercles; disk with deep fur-
rows arranged in whorl. Elytra long, with parallel sides (female) or nar-
row somewhat anteriorly (male), with dense fine punctation. Legs long,
slender; hind femora extend beyond apex of elytra (male) or just reach
it (female).

Larva: Disk of pronotum with dense rusty hairs forming two trans-
verse fields. Distance between hairs less than length of hairs. Abdominal
tergite IX with sparse hairs on disk and denser hairs laterally.

Pupa: Distinguished from pupae of other genera by carina on anterior
margin of pronotum without hairs. Meso- and metanota glabrous, lus-
trous, without hairs.

Inhabits mainly regions of Southeast Asia within the Indo-Malayan
zone. Only one species in northern Asia.

Type species: Callichroma bivittatum White, 1853.

94 1. Chloridolum sieversi Ganglb.

Ganglbauer, 1886, Horae Soc.Entom. Ross., vol. 20, p. 135; PlaviP-
shchikov, 1940, Fauna SSSR, 22, 2, 209-211; Cherepanov and Chere-
panova, 1977, Taksony fauny Sibiri {Nov. i maloizv. vidy fauny Sibiri),
pp. 142-146.

Adult (Figure 54): Body elongate, large. Head small, distinctly narrow-
er than pronotum. Frons between antennae transversely convex, smooth,
with sparse fine punctation, with narrow longitudinal suture. Vertex slight-
ly impressed, with deep furrows diverging posteriorly. Occiput with
dense deep punctation. Eyes large, broadly emarginate. Antennae 2.0
times (male) or almost 1.5 times (female) longer than body; 6th segment
in male extends beyond apex of elytra, 8th segment in female; from 3rd
segment with sharp groove on outer side. First antennal segment thick,
matte, with sparse punctation, with light yellow articulate tubercle at
base; rest of segments with fine notchlike punctation.

Pronotum slightly longer than width at base, medially broadens
moderately, laterally with large smooth conical tubercle, apically with

123

Figure 54. Chloridolum sieversi Ganglb.

124

acutely produced, spinelike tubercle; disk convex, uneven; in anterior
half with two proximate rugose tubercles; in posterior half with two
widely spaced, transversely elongate, smooth tubercles; near apex and at
base with broad transverse flange and there with transverse furrows;
disk with deep whorled furrows. Scutellum triangular, broad, with dense
fine punctation, pointed apically. Elytra elongate, narrow slightly pos-
teriorly, individually rounded apically; disk insignificantly convex, smooth,
without visible carinae, with unusually dense fine punctation and minute
light-colored hairs not forming compact cover. Legs long, slender; hind
femora almost reach apex of elytra. First segment of hind tarsi notably
longer than two successive together. Body ventrally with iridescent golden
hairy cover. Abdominal sternites laterally with pitlike dents. Sternite V

95 transverse, notched apically (male), or elongate, narrowly rounded api-
cally (female). Ventral side of body and pronotum yellowish-red. Head
and elytra blue or bluish-green with metallic sheen. Antennae, scutellum,
and legs bluish-brown or almost black; articulate tubercle at base of 1st
antennal segment light yellow. Body length 24 to 32 mm.

Egg: White with greenish iridescence, narrower toward one pole,
broadly rounded at both poles. Chorion smooth, without cellular sculp-
ture, matte. Length 3.0 mm, width I.O mm.

Larva (Figure 55): Body massive. Half of head retracted into pro-
thorax. Epistoma slightly convex, rusty-yellow, on anterior margin with
rusty-brown border sloping roundly forward, medially with sharp, slight-
ly impressed longitudinal suture, bound laterally by faint frontal sutures;
disk with innumerable short hairs. Hypostoma convex, uniform along
anterior margin, without notches, around inner angles of sclerites with
spinelike projection. Parietals laterally in anterior half with several short
hairs. Clypeus white, broadly flattened- Labrum convex, along margins
with short coarse brown setae, glabrous and white on disk, with brownish
transverse band at base. Mandibles massive, outer side with broad dent,
black, basally with red border. Inner masticatory lobes apical, maxil-
lary stipes on outer side, and labial mentum posterolaterally with coarse
rusty setae. Labial submentum with large elongate rusty spot at base,
rounded apically.

96 Pronotum narrowly rounded anteriorly, slopes markedly toward head,
with pitted sculpture in anterior half, white border on anterior margin,
behind which broad rusty-yellow transverse band occurs (with white
longitudinal clearance medially and two paramedial alveolar white
notches on anterior margin); sides and disk with short thick hairs form-
ing two transverse fields interrupted medially, one on anterior margin of
transverse rusty-yellow band, the other anterior to scutum. Pronotal
scutum white, coriaceous, convex, with longitudinal furrows demarcated
laterally by longitudinal grooves; at base and on disk with stray setae or

125

Figure 55. Larva of Chloridolum sieversi Ganglb.
a — head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla.

without them, near posterior angles with innumerable minute hairs. Alar
lobes with dense rusty hairs. Apparently hairy field covering sides of
pronotum and alar lobes proceeds from there onto posterior angles of
scutum. Prothoracic presternum laterally with dense hairs and medially
with sparse rusty hairs, paramedially with brownish-rust streaks; euster-
num coriaceous, rugose, merges with general presternal surface. Thoracic
legs short, poorly developed, claws minute, spinelike.

Abdomen laterally with dense rusty hairs. Dorsal locomotory am-
pullae quite convex, rugose, separated by two transverse grooves merg-
ing laterally with longitudinal grooves. Ventral locomotory ampullae
with lateral longitudinal groove and transverse band diverging from it,
which often merges with transverse band on opposite side. Abdominal
sternites VIII and IX entirely covered with rusty hairs. Tergite IX with
sparse hairs on disk and denser hairs on sides.

Pupa (Figure 56): Readily recognized by large body, long looplike
antennae, and pronotum. Head glabrous, without setae, transversely
convex between antennae, with broad longitudinal groove, broadly

126

impressed on vertex, and broadly rounded on occiput. Antennae long,
posterior to apices of hind femora bent forward, looplike.

Pronotum transverse, wider at base than long, mediolaterally with
sharp protuberance, near anterior margin without flange but with medial
setaceous tubercle sloping forward uniformly, near base with narrow
transverse groove; disk convex, medially with minute setae forming trans-
verse row, laterally anterior to transverse groove with basal, transverse-
ly elongate, setaceous elevation (setae thin, with small reddish sclero-
tized tubercle at bases). Mesonotum convex, lustrous, posteriorly with
narrowly extended, transversely rugose scutellum. Metanotum glabrous,
convex, with transverse fine striation on disk, and median longitudinal
groove.

Abdomen elongate, barely broadens in middle, narrows gradually
posteriorly. Abdominal tergites quite convex, with common median

127

longitudinal groove, in posterior half with sharp paramedial spinules bent
down and forward forming transverse band of two or three rows. Tergite
VII not rounded apically; spinules in posterior third form small tuft.
Hind femora pressed to sides, almost reach middle of abdominal tergite
VII. Body length 25 to 36 mm, width of abdomen 7.0 to 8.0 mm.

Material: Collected in Ussuri-Primor’e region. Adult insects 11, lar-
vae 21, pupae— two males, larval exuviae with beetles from cells one.

Distribution: Ussuri-Primor’e region, northeast China, North Korea.

97 Biology: Inhabits broad-leaved forests. Ecologically associated with
Manchurian walnut {Juglans manshurica). Beetles fly in August and first
half of September. After mating, female oviposits in crevices of thick
bark (usually on bast); eggs laid singly or in pairs. In gardens up to eight
eggs have been laid in one batch on plywood planks. One female can lay
as many as 68 eggs in her lifetime.

Colonizes trunks of susceptible trees up to 36 cm or more in diameter.
Development of eggs takes two to three weeks. Larva nibbles opening in
bark, makes gallery along trunk under bark, and plugs it behind with
fine frass. Gallery weakly impressed on alburnum, quite often with step-
like flexure at right angle. Length of gallery under bark 17 to 20 cm,
width 1.2 to 2.8 cm. Mature larva bores wood up to 6.0 to 7.0 cm, later
makes longitudinal gallery 12 cm long, 1.1 cm wide. Inlet (1.1 cm X 0.6
cm) visible on 'sapwood surface, extends along trunk, and is plugged
with frass. Gallery in wood hollow, only anterior to pupal cell plugged
with coarse fibrous and fine frass; plug 2.0 to 2.5 cm long. Length of
pupal cell 3.2 to 4.0 cm, width 0.7 to 1.5 cm. Pupation of larvae occurs
after second hibernation in July and first half of August. Beetles sighted
end of July and in August. Flight openings (5.0 mmX 10.0 mm to 6.0
mm X 12.0 mm) of young beetles visible on surface of trees.

Weight of mature larvae prior to readying for pupation 370 to 1,520
mg. Weight of larvae before pupation 259.0 to 885.5 mg, pupae 218 to
805 mg, and beetles- 191.0 to 609.9 mg. Larval population density on
Manchurian walnut trunks comparatively low. For example, on one
trunk (length of populated zone 6.3 m, diameter of cutting 18.0 cm)
19 larvae were detected. We inspected habitats of Chloridolum sieversi
Ganglb. in 1971 near Partizansk in Primor’e region (First Kamenka
River). Manchurian walnut stock was damaged to the extent of 15 to
20%.

3. Genus Chelidonium Thoms.

Thomson, 1864, Syst.Cerambycidae,\o\, 175, p. 420; Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 213-215; Gressit, 1951, Longicorn Beetles of
China, vol. 2, p. 196.

128

Adult: Characterized by very short antennae not extending beyond
elytral apices. Disk of pronotum with dense, very minute punctation,
with rounded protuberances laterally.

Larva: Head highly retracted into prothorax. Disk of pronotum with
very sparse thick setae forming two transverse fields. Distance between
setae more than length of setae. Anal segment (X) of abdomen with
sparse thick setaceous hairs.

Pupa: Well distinguished from pupae of other genera of this tribe by
presence of setaceous tubercle on anterior margin of pronotum. Meso-
and metanota with long acicular setae forming paramedial tufts, one on
each side.

The genus Chelidomum belongs mainly to Southeast Asian fauna and
comprises over 10 species. Only one species penetrates Central Asia.

Type species: Cerambyx argent atus Dalman, 1817.

1 . Chelidonium zaitzevi Plav.

Plavilstshikov [PlaviPshchikov], 1933, Entom. Nachrbl, vol. 7, p. 107;
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 215-216; Cherepanov and
Cherepanova, 1976, Taksony fauny Sibiri {Nov. i maloizy. vidy fauny
5/Z?/n), pp. 138-142.

98 Adult (Figure 57): Readily recognized by its bright green body and
fine dense punctation on pronotal disk. Head short, highly retracted into
prothorax, bulges wartlike between antennae, with smooth median longi-
tudinal suture, deeply impressed between upper lobes of eyes, more finely
punctate on occiput. Genae broad, deep, sparsely punctate. Eyes broadly
emarginate, very small but distinctly faceted. Antennae shorter than
body, apices extend beyond 0.66 length of elytra; from 3rd segment with
outer troughlike longitudinal groove; 5th to 10th segments apically with
spinelike extended outer angles; 4th segment equal to 5th, about 0.50
length of 3rd.

Pronotum narrows more anteriorly, less posteriorly, not longer than
width at base, with obtusely extended rounded protuberance laterally,
very fine punctation on disk imparting matte appearance, laterally with
large punctation, medially with longitudinal uneven band, more distinct
on anterior and posterior margins, basally and apically with transverse
lustrous nonpunctate groove. Scutellum elongate, triangular, pointed
apically, with uneven punctation. Elytra markedly elongate, narrows
slightly posteriorly, individually rounded apically (inner angle narrowly
rounded and outer gently tapered), with dense very fine punctation, and
hairs barely perceptible under high magnification. Hind femora not long,
do not reach elytral apices. Hind tibiae straight, broaden gradually
toward apex. Hind tarsi somewhat shorter than tibiae; 1st segment con-
siderably longer than two successive. Body ventrally with compactly

129

Figure 57. Chelidonium zaitzevi Plav.

adherent minute silvery hairs not forming dense hairy cover. Abdominal
sternite V of female transverse, broadly rounded apically. Entire body
and elytra bright green with metallic sheen. Legs dark violet. Antennae
dark brown. Elytra laterally with golden-bronze border. Body length 17
to 25 mm.

130

Egg: Yellow, oval, thick. Chorion with minute cellular sculpture.
Septa between cells thin. Length 2.0 mm, width 1.1 mm.

99 Larva (Figure 58): Characterized by arrangement of coarse setaceous
hairs on pronotum and abdominal tergite IX. Head highly retracted into
prothorax. Epistoma slightly convex, almost flat, with narrow, median,
longitudinal, brownish suture, laterally with faint frontal sutures, on
anterior margin with dark brown border, with long and short setaceous
hairs posteriorly. Hypostoma smooth, uniform, without spinelike projec-

Figure 58. Larva of Chelidonium zaitzevi Plav.
a— head and pronotum; b— abdominal tergite IV
with dorsal locomotory ampulla; c — tip of abdo-
men (ventral view).

131

tions and notches along anterior margin, narrows slightly anteriorly,
without hairs. Anterior half of parietals with stray hairs, with one faint
pigmented ocellus below. Clypeus small, trapezoid, white. Labrum white,
notably elongate, slightly pointed apically, with minute light-colored
setae laterally. Mandibles massive, outside base with wide median longi-
tudinal groove. Maxillary stipes on outer side with four setae; mastica-
tory lobes of maxillae elongate, digitate, glabrous, with deep sparse setae
only at apex. Labial submentum with sharply prominent brownish-yellow
triangular spot pointed at apex and pair of thick setae; mentum with two
and ligula with eight thick setae forming transverse row.

100 Pronotum narrows anteriorly, broadly rounded on anterior margin,
in anterior third with yellowish-rust transverse band with narrow longi-
tudinal white clearance medially, white alveolar paramedial notch on
anterior margin, one white spot on each side; disk with sparse coarse
setaceous hairs forming two transverse fields, laterally closer to base
with dense setaceous hairs. Pronotal scutum convex, white, coriaceous,
with longitudinal striations (longitudinal streaks or furrows) extending
forward medially on anterior margin, bound laterally by deep straight or
flexed longitudinal grooves, at base with stray setae forming transverse
row. Setae more distinct in mature larvae. Prothoracic presternum later-
ally with dense thick hairs, on disk with sparse stray hairs, in anterior
half with two large spots. Eusternum coriaceous, glabrous, merges with
presternum. Thoracic legs short, with very short sclerotized claw.

Abdomen elongate; two anterior segments laterally with dense hairs,
remainder with very sparse coarse hairs. Dorsal locomotory ampullae on
tergites III to VII more convex, divided by common longitudinal groove,
two transverse grooves merging with short lateral ones. Ventral locomo-
tory ampullae divided by single transverse groove merging laterally with
lateral grooves. Abdominal segment X large, hemispherical, with sparse
coarse setaceous hairs. Body length of mature larvae up to 35 mm, width
of head 4.0 mm.

Pupa (Figure 59): Characterized by extended setaceous tubercle on
anterior margin of pronotum and development of setae on meso- and
metanota. Head glabrous, transversely convex between antennae, broadly
impressed between upper lobes of eyes, broadly rounded on occiput.
Antennae pressed to sides, in second third ventrally arcuate.

Pronotum bulges, broadens angularly on sides, narrows anteriorly
slightly more than posteriorly, with biapical protuberance medially on
anterior margin bearing two tufts of short setae, with short setae on disk
anterior to middle forming transversely elongate tuft, posterolaterally
with similar setae forming one tuft on each side. Mesonotum convex,
laterally with setae forming one tuft on each side reaching toward anteri-
or angles in anterior half, and one transverse row on each side in pos-

132

terior half. Scutellum triangularly produced on posterior margin of
mesonotum, transversely rugose, matte. Metanotum posteriorly with
rounded angles, medially with longitudinal groove, with fine transverse
striation, anterolaterally with setae forming one small tuft on each side
(ten setae per tuft).

Abdomen elongate, almost with parallel sides, narrows posteriorly
from segment VI. Abdominal tergites convex, medially with transverse
dent, divided by common longitudinal groove, paramedially posterior to
transverse dent with several large spinules forming transverse band, an-
terior to transverse dent with stray, sometimes paired, fine spinules.
Abdominal tergite VII broad, gently rounded apically, with minute spi-
nules, of which anterior ones form transverse row, posterior ones (usually
paired) dispersed tuft. Apex of tergite VIII narrowly rounded, with stray
minute acicular setae. Tip of abdomen (ventral view) obtuse, without
setae. Valvifers of female large, hemispherical, widely separated. Body
white, with yellowish tinge. Body length 25 mm, width of abdomen 6.0
mm.

Figure 59. Pupa of Chelidonium zaitzevi Plav.

133

101 Material: Collected in Ussuri-Primor’e region (Kamenka River near
Partizansk). Adult insects — four females, larvae five, pupa — one female,
larval exuviae from cells with beetles five.

Distribution: Ussuri-Primor’e region from Partizansk to Vladivostok.
Sporadic.

Biology: Inhabits broad-leaved forests of Ussuri-Primor’e region. Beet-
les fly in July and August. Female oviposits on thin adventitious shoots
of growing maple {Acer pictum, A. mono, A mandschuricum). Diameter
of shoots 3.0 to 5.0 mm. One female can layup to 36 eggs. Newly hatch-
ed larva bores shoot, makes longitudinal gallery toward base in pith,
and cuts ventilation opening through which frass is ejected. Migrates
from thin adventitious shoot into very thick main shoots 1.5 to 7.5 cm in
diameter, continuing to make gallery from top downward. As they move
larvae make lateral branches that terminate in round or oval ventila-
tion openings. Width (size) of ventilation openings in galleries made by
I-instar initially 1.0 to 3.0 mm and later 6.0 to 12.0 mm. Former usually
round, latter more elongate, oval. Distance between ventilation open-
ings at commencement of gallery 0.8 to 2.0 cm, at end 6.0 to 17.0 cm.
Quite often larval galleries penetrate shoots of tertiary and secondary
order, and terminate in shoots of primary order. Termination of larval
gallery in trunk zone has also been recorded. Total length of gallery 1.2
to 2.4 mm, width 8.0 to 12.0 mm. Gallery hollow throughout, free from
frass. Lavae move rapidly from one end of gallery to the other and thus
elude woodpeckers. Mature larva makes additional gallery from bottom
upward, parallel to main gallery, forms cell, and closes it off from rest of
gallery with parchmentlike stopper before pupating. Length of cell 38 to
54 mm, width 6.0 to 8.0 mm.

Pupa lies in celLwith head down (toward parchmentlike stopper). At
room temperature develops in about three weeks. Young beetle ruptures
stopper, penetrates larval gallery up to ventilation opening, and emerges
through it. Weight of mature larvae 232 to 639 mg, pupae 21 1.0 to 581.8
mg, and beetles 191 to 523 mg. Larvae pupate after third hibernation.
After second hibernation infested shoots shed leaves, wither in August,
their leaves turning reddish-yellow, and stand out distinctly against gen-
eral background of green crowns of trees. In 1973 we detected a small
zone sparsely infested with Chelidonium zaitzevi Plav. near Partizansk in
mature, growing woodstand on terraces of the First and Second Kamenka
Rivers. Lower and middle zones of shoots of mature woodstand were
damaged more often than upper. Usually the upper zone was left un-
touched.

134

4. Genus Leontium Thoms.

Thomson, 1864, Syst. Cerambycidae, vol. 175, p. 420; Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 212; Gressit, 1951, Longicorn Beetles of
China, vol. 2, p. 207.

Adult: Body elongate, bright green. Antennae barely longer than
body. Pronotum laterally with small sharp tubercle, transversely rugose
on disk. Elytra markedly elongate, with dense fine punctation. Legs very
long, slender; hind femora extend beyond elytral apices.

102 Larva: Body elongate, slender. Pronotum with dense hairs forming
two transverse bands on disk, one (broader) anterior to scutum and
second in interior third. Pronotal scutum white and convex, with longi-
tudinal striation.

Pupa: Distinguished from pupae of other genera of this tribe by elon-
gate body and long femora extending beyond tip of abdomen.

The genus Leontium is mainly found in the Indo-Malayan region;
about ten species inhabit Southeast Asia. One species is knowm in north-
ern Asia and represents an insular relict.

Type species: Leontium viride Thomson, 1864

1. Leontium viride Thoms.

Thomson, 1864, Syst. Cerambycidae, \o\. 175, p. 420; Plavi’lshchikov,
1940, Fauna SSSR, 22, 2, 213; Gressit, 1951, Longicorn Beetles of China,
vol. 2, p. 207; Kojima and Okabe, 1960, Food Plants of Japan. Ceram-
bycidae, p. 120; Cherepanov and Cherepanova, 1977, Taksony fauny
Sibiri (Nov. i maloizv. vidy fauny Sibiri), pp. 146-150.

Adult (Figure 60): Readily recognized by slender elongate body, long
slender legs, and bright green coloration. Head between antennae trans-
versely convex, with narrow median longitudinal suture, uneven, simple,
longitudinally rugose punctation, but medially smooth on vertex, without
punctation, with dense deep punctation on occiput. Eyes narrow but
deeply notched, very finely and sharply faceted. Antennae slender; 11th
segment extends beyond apex of elytra; outer side of 6th to 10th segments
produced apically, spinelike.

Pronotum slightly longer or even shorter than width at base, later-
ally with small sharp projecting tubercle, near anterior and posterior
margins with transverse flange, and here with transverse girdling furrows
(folds), disk convex, with short median grooves from which transverse
furrows radiate. Scutellum triangular, smooth, slightly impressed, with
fine uneven punctation. Elytra narrow, markedly elongate, with parallel

103 sides, 4.0 to 5.0 times longer than general width, moderately convex on
disk, finely and densely punctate (almost shagreen), apically with narrow-
ly rounded inner and gently tapered outer angle. Legs long, very slender.

135

102

Figure 60. Leonlium viride Thoms.

136

Hind femora thicken gently toward apex, extend beyond elytral apex.
Hind tarsi slender; 1st segment 2.0 times (male) or 1.5 times (female)
longer than two successive together.

Abdomen elongate, slender, with tender gray hairs not forming com-
pact cover. Abdominal sternite V broadly notched at apex (male) or
narrowly rounded, sometimes even angularly elongate, almost pointed
(female). Entire body, scutellum, and elytra bright green. Legs and
antennae dark brown. Body length 14 to 18 mm.

Egg: White, after oviposition acquires greenish hue, elongate, thicken-
ed in one half and narrows gradually in the other, broadly rounded at one
pole and narrowly at the other. Chorion smooth, lustrous, without per-
ceptible cellular sculpture. Length 1.4 mm, width 0.6 mm.

104 Larva (Figure 61): Distinguished from larvae of other species of the
tribe Callichromini by broad hairy field on pronotum and slender but not
very long body. Head narrows insignificantly anteriorly. Epistoma almost
flat, on anterior margin with lustrous brownish border, whitish on disk,
behind brownish border (especially laterally) with short thin hairs divid-
ed by distinct median longitudinal suture, laterally with barely percep-
tible frontal sutures. Hypostoma almost with parallel sides, with rounded
anterior outer angles, medially separated into two triangular sclerites by
broad flat gula on anterior margin uniform, smooth, without projection.
Parietals in anterior half with short hairs; hairs around frontal sutures
dense, laterally and ventrally sparse. Antennae long, slender, with four
segments. Ocelli not visible near antennal bases. Outer side ol mandibles
with longitudinal groove basally, sometimes here with stray setae form-
ing longitudinal row. Labial submentum with pair of widely separated
setae, faint brownish spot at base.

Pronotum slopes forward, in anterior half with broad rusty-yellow
transverse band, with longitudinal clearance medially and laterally, and
also with white alveolar paramedial notch on anterior margin; antero-
laterally with yellow transverse band and especially before scutum with
dense hairs forming two transverse fields. Prescutal hairy field broad,
with median longitudinally elongate clearance broadening posteriorly,
on posterior margin uniform, as though bound by a scale. Pronotal
scutum slightly convex, markedly transverse on anterior margin, projects
slightly anteromedially, with longitudinal groove laterally, and dense thin
longitudinal striation throughout surface. Prothoracic presternum later-
ally with dense hairs, anteriorly on disk with less dense (sometimes
sparse) hairs, in some specimens with brownish specks; eusternum cori-
aceous, lustrous, glabrous, merges with common surface of presternum.
Legs short, with small acute light-colored claw.

Abdomen markedly elongate, slender, laterally with short thin hairs.
Locomotory ampullae rather convex, coriaceous, dorsally with two trans-

137

103

Figure 61. Larva of Leontium viride Thoms,
a — head and pronotum; b — abdominal tergite IV with dorsal loco-
motory ampulla; c—abdominal segments III to V of 1-instar
larva.

verse grooves fusing laterally, and medially with ventral alveolar depres-
sion from which short furrows radiate; three to six short setae on inner
clivus form independent tuft. I-instar larvae on abdominal segments III
to V laterally with sharp spinules that slough after molt. Body length of
mature larvae 16 to 18 mm, width of head 2.5 mm.

Pupa (Figure 62): Characterized by slender elongate body and long
hind femora that extend beyond tip of abdomen. Head glabrous, without
setae, lustrous, around antennal bases tubercularly convex, uniformly
rounded hemispherically on occiput. Antennae long, pressed to sides,
in middle bent ventrad and forward, looplike, with second half pressed
to body ventrally.

138

Pronotum elongate, basally with narrow transverse flange, with pro-
jecting posterior angles, convex on disk, lustrous, narrows gradually
anteriorly, laterally in posterior half with distinctly produced protuber-
ance, anteromedially sometimes with minute setae forming very small
tuft on tubercular elevation, posterolaterally with transverse setaceous
band, medially sometimes with several thin setae forming transverse
row. Mesonotum convex, lustrous, with sharp scutellum posteriorly
cuneiform, laterally in posterior half with stray, barely visible setae.

105 Metanotum slightly convex, straightly truncate posteriorly, with distinct
posterior angles. Hind femora pressed to sides; extend beyond tip of

103

Figure 62. Pupa of Leontium viride Thoms.

139

abdomen. Abdomen quite elongate, in region of segments III and IV
slightly enlarged. Abdominal tergites posterior to middle or on poste-
rior margin with minute spinules directed forward. Abdominal tergite
VII elongate, narrowly produced posteriorly, glabrous in anterior half,
with minute spinules or setae forming faint tuft in posterior half. Valvifers
of female hemispherical, insignificantly separated. Body length 10 to 18
mm, width of abdomen 3.0 to 3.5 mm.

Material: Collected on Sakhalin and Kunashir. Adult insects 12,
larvae 81, pupae — two males and six females, larval exuviae with beetles
from cells two.

Distribution: In the USSR southernregionsof Sakhalin, Kuril’ Islands
(Kunashir and Shikotan); Japan, Taiwan.

Biology: Inhabits coniferous and coniferous-broad-leaved forests, and
ecologically associated with spruce. Beetles fly in second half of summer,
seen on flowers of Umbelliferae and other plants, usually in meadows.
Maximum number sighted in second half of July and in August. Beetles
feed on flowers, mate, and then fly to trees. Female lays eggs in bark
crevices singly or three to four in the same place. In gardens female
sometimes glues eggs to bark surface or to panes of greenhouses. Colo-
nizes shoots 2.5 to 7.2 cm in diameter on viable, drying, and felled trees
until resin supply exhausted. Does not inhabit shoots in which bast has
dried. Larvae appear two to three weeks after oviposition. Larvae
hatched from August 24 to 30 from eggs laid on July 20 to 28. Mass
hatching of larvae is observed in nature in last 10 days of August.

Newly hatched larva bores bark, makes sinuous gallery under bark,
deeply impressed on alburnum, and plugs it with fine frass. Width of
gallery 1.5 to 10.0 mm. Galleries fuse at places forming cells 1.5 to 2.0
cm wide. Mature larva bores wood up to 3.0 cm, makes longitudinal
gallery there, and does not close it with frass. Length of longitudinal
galleries in wood 5.0 to 9.0 cm, width up to 5.0 mm. Inlet plugged with
fine frass remains on bark surface, its size 1.8 mm X 3.0 mm to 3.5
mm X 6.0 mm. At end of longitudinal gallery pupal cell 2.2 to 2.5 cm
long, 4.0 to 5.0 mm wide. Cell sealed with small plug of fine frass. Larva
molts in ceil and enters diapause by autumn. Pupates after hibernation.
Pupa lies with head toward plug sealing cell from hollow larval gallery.
Pupation of larvae after hibernation in May and June. Adults develop
from pupae in July and August. Young beetle breaks frass plug, pushes
it aside, moves into hollow larval gallery toward shoot surface, nibbles
106 round opening in bark (3.5 to 4.0 mm), and emerges. Emergence of
beetles from wood commences early July and extends up to early August
inclusive. Weight of larvae before pupation 65 to 140 mg, pupae 52 to
112 mg, and beetles 39 to 92 mg. Generation completed in two years
(Table 8).

140

105 Table 8. Periods of development of Leontium viride Thoms.

Year of

development

May

June

July

August

September

October

1st

PAE

PAEL

AEL

2nd

3rd

PAE

PAEL

AEL

Population density comparatively high. For example, in a spruce
shoot 15.5 cm long, 5.0 cm in diameter from Kunashir Island (Alekhino)
three larvae were found under bark and seven in wood (10 specimens).
Similar larval population density of Leontium viride Thoms, observed
on other shoots. Massive reproduction zones of this species have been
found in unkept forests containing cut, wind-fallen, and snow-crushed
trees.

5. Genus Polyzomis Cast.

Castelnau, 1840, Hist. Nat., Vol. 2, p. 438; Plavil’shchikov, 1940,
Fauna SSSR, 22, 2, 216-217; Gressit, 1951, Longicorn Beetles of China,
vol. 2, p. 209.

Adult'. Characterized by elongate cylindrical body. Antennae extend
slightly beyond apex of elytra. Pronotum laterally with small sharp tuber-
cle, convex on disk, with large deep pun ctation. Elytra convex, with dense
fine punctation, and broad light-colored bands.

Larva'. Similar to larva of Chelidonium zaitzevi Plav. in structure of
hairy cover and yellow body; resembles larva of Leontium viride Thoms,
in structure of locomotory ampullae. Differs from both these species in
very broad gula and arrangement of corase setae on posterior margin of
abdominal sternite X, which form transverse row. Anterior half of ster-
nite X glabrous, without hairs.

Pupa'. Well distinguished from pupae of other genera of the tribe
Callichromini in semiannular or annular arrangement of spinules on
abdominal tergites.

Areal of genus extends toward Indo-Malayan region. Over ten species
known in Southeast Asia, one in southeastern part of northern Asia.

Type species'. Cerambyx fasciatus Fabricius, 1781.

107 1 . Polyzonus fasciatus (F.)

Fabricius, 1781, Syst. Entom., p. 232 {Cerambyx)', sibiricus, Gmelin
1790, Syst. Nat., 1, 4, 1840 (Cerambyx); Plavil’shchikov, 1940, Fauna
SSSR, 22, 2, 218-219.

141

Adult (Figure 63): Differs from other species of the tribe Callichro-
mini inhabiting the Palearctic in almost cylindrical body and presence
of broad transverse bands on elytra. Head retracted into prothorax almost
up to eyes, with coarse rugose punctation, and narrow median longitudi-
nal suture. Eyes large, finely faceted, inner side with broad deep notch,
often adjoins anterior margin of pronotum. Antennae slender; 6th to
108 10th segments with projecting outer apical angle, 10th segment extends
beyond apex of elytra (male) or just reaches it (female).

Pronotum distinctly (male) or slightly (female) elongate, convex, with
flange at base, barely perceptible broad transverse groove around anterior
margin or without it, sharp tubercle mediolaterally, with large fused
punctation on disk and minute uneven punctation laterally on anterior
and posterior fourth. Scutellum narrows posteriorly, pointed or narrowly
rounded apically, flat, sometimes with smooth median longitudinal groove.
Elytra elongate, convex, with parallel sides, apically with narrowly round-
ed inner angle, with very dense fine punctation and minute adherent
hairs. Legs slender, not very long. Hind femora just short of elytral apex
(male) or slightly shorter, reaching only 0.75 length of elytra (female).

Abdomen elongate, narrows insignificantly posteriorly, almost cylin-
drical, with minute light-colored hairs. Abdominal sternite V transverse,
apically truncate or slightly emarginate (male) or slightly elongate, nar-
rowly rounded (female). Body and legs dark blue or violet-blue. Anten-
nae, especially apically, brownish. Elytra dark blue with two broad, con-
tinuous, yellow transverse bands, one anterior, the other posterior to
middle (f. typica). Sometimes only anterior band widely interrupted on
suture by septum (ab. anticeinterniptus Plav.), or only posterior (ab. pos~
ticeinterruptus Plav.), or both anterior and posterior bands interrupted (ab.
biinterruptus Plav.); sometimes yellow bands slightly broader than dark
blue clearance between them (ab. latefasciatus Plav.). Body length 14 to
20 mm.

Egg\ Yellow, oval, rounded at poles, narrows more toward posterior
end. Chorion with minute noncellular sculpture, slightly matte. Length
2.8 mm, width 1.2 mm.

Larva (Figure 64): Body yellow. Head with parallel sides. Epistoma
lustrous, with coarse punctation subapically (especially laterally), smooth
posteriorly, with diffuse yellowish-rust border in front; behind border long
coarse setaceous hairs form indistinct transverse row, and behind this
row short setaceous hairs form extensive paramedial triangular field with
apex directed backward. Frontal sutures not visible. Epistoma fuses with
parietals, median longitudinal suture distinct in posterior half. Hypo-
stoma divided into two widely separated, slightly convex, triangular scleri-
tes with rounded anterolateral angles; near inner angles with stray short
setae, with very narrow brownish border on anterior margin, with

142

107

Figure 64. Larva of Polyzoms fasciatus (F.).
a-~head and pronotum; b — abdominal tergite IV with
dorsal locomotory ampulla; c — tip of abdomen
(ventral view).

144

supporting spinelike projection near inner angles. Gula broad, without
brownish border along anterior margin, Parietals in anterior half with
sparse setaceous hairs, near antennal bases with small black pigmented
ocellus. Antennae comparatively long, apices project notably beyond an-
terior margin of cephalic capsule; 1st antennal segment long, just slightly
thicker than 2nd, rusty-brown apically; base of 2nd and 3rd segments same
shade as apex of 1st. Clypeus narrow, trapezoid, semitransparent. Labrum
oval, narrows slightly toward base, narrowly rounded at apex, convex,
with hyaline sheen, and long acicular semierect setae. Apices of mandibles
broadly rounded, smooth on outer side, lustrous black, basally reddish-
rust or whitish, sometimes with rusty tinge on cultrate edge. Maxillary
109 stipes with nine or ten acicular setae on outer side, inner masticatory
lobes thick, obtuse apically, with innumerable short setae. Maxillary
palps slightly longer than inner lobes, their segments basally rusty-brown,
transversely oval. Labial submentum with pair of widely separated
acicular setae, mentum with squarrose setae forming tuft on each side,

Pronotum slopes markedly from base toward head, broadly rounded
on anterior margin, in anterior third with transverse yellow band inter-
rupted medially and laterally by longitudinal white strip into four trans-
versely elongate spots; two spots on disk and one on each side. Disk
anterior to scutum and pronotum laterally with coarse, uniform, seta-
ceous, rusty-red hairs forming common field interrupted by very narrow
clearance medially and in region of lateral longitudinal grooves. Prono-
tal scutum convex, coriaceous, bound laterally by long deep longitudinal
grooves, anterior margin almost straightly truncate, with thin longitudi-
nal furrows. Prothoracic presternum laterally and on disk with rusty
setaceous hairs; eusternum not demarcated from presternum, merges
with it, glabrous basally, lustrous. Thoracic legs well developed, with thin
acicular claw.

Abdomen elongate, laterally with several thin hairs. Dorsal locomo-
tory ampullae coriaceous, highly convex, divided by common median
longitudinal groove, laterally with oblique longitudinal grooves that
merge with transverse groove in anterior half. Ventral locomotory am-
pullae similar in structure. Abdominal tergites VIII and IX in anterior
half glabrous, lustrous, in posterior half with sparse rusty hairs. Sternite
VIII glabrous, only on posterior margin with stray setae; sternite X
in posterior half with 10 reddish-rust setae forming transverse row. Body
length of mature larvae 24 to 25 mm, width of head 2.5 to 3.0 mm. Body
of I-instar larva yellowish, comparatively thick, only 2.5 times longer than
width in prothoracic region. Abdomen without spinules laterally.

Pupa (Figure 65): Characterized by elongate body, antennae apically
falcate, tubercularly extended spiracles, presence of innumerable sharp
spinules on abdominal tergites, and other features. Head elongate, nar-

145

107

rows anteriorly, cuneiform, transversely convex between antennae, im-
pressed on anterior margin and vertex, uniformly rounded on occiput.
Antennae pressed to sides, apically falcate, turning forward from 9th to
11th segments.

Pronotum wider (female) or narrower (male) than long, laterally with
more (male) or less (female) projecting conical tubercle, narrows ante-
riorly and posteriorly, moderately convex on disk, with transverse streaks,
basally with narrow transverse groove, laterally on hind clivus with large

146

or minute setae forming one well-developed tuft on each side, sparse
(stray) short setae on foreclivus, which sometimes form small tuft only in
middle of anterior margin. Mesonotum in anterior half insignificantly
convex, transversely impressed posterior to middle, with triangularly
projecting scutellum elevated apically on posterior margin, laterally with
stray minute setae. Metanotum with median longitudinal groove, later-
ally gently impressed, with stray dispersed short setae or without them,
produced trapezoidally on posterior margin.

Abdomen elongate, with parallel sides or, in region of segments III
and IV broadens insignificantly, narrows posteriorly. Abdominal tergites
in posterior third convex, impressed like a facet anteriorly, with narrow,
sometimes sharp median longitudinal groove; innumerable sharp spinu-
les in posterior third form broad transverse band that curves laterally
110 around facetlike impression in form of semiannular or circular strip.
Tergite VII triangular or almost semicircular, broadly or, more often,
narrowly rounded angularly, convex on disk, in posterior half with small
(smaller than in preceding tergites) spinules forming one common or
several small tufts. Tergite VIII 0.50 fength of VII, angularly rounded
posteriorly, convex on disk, with sparse very minute spinules or without
them. Hind femora slender, pressed tightly to sides of abdomen, with
apices extending beyond abdominal tergite V. Vajvifers of female hemi-
spherical, widely separated, bent laterally, with small apical tubercle.
Body length 18 to 20 mm, width of abdomen 4.0 to 4.5 mm.

Material: Collected in eastern Siberia (Nerchinsk, Shakhtoma, and
Ussuri-Primor’e region). Adult insects 53, of which 46 raised in the
laboratory from larvae collected in nature, larvae 68, pupae — three males
and six females, larval exuviae more than 10.

Distribution: Southern regions of eastern Siberia from Baikal to Vla-
divostok, northern Mongolia, northern China, and Korea. Sporadic.

Biology: Inhabits forest-steppe zone and ecologically associated with
dog rose. Beetles fly end of June to last ten days of September. In 1978
numerous beetles were collected around Lake Khasan in the first few
days of August and in mid-September, but disappeared by September
24. Beetles require supplementary feeding and are seen on flowers of
Umbelliferae, Rosales, Compositae, and other plants. During this period
their gonads mature. Female lays eggs 10 to 1 4 days after mating on
shoots of dog rose and smooth root portion devoid of thorns. Oviposit-
ed eggs are covered with a sticky exudate that readily adheres to smooth
bark of shoots and hardens into rugose yellowish crust. Thus each egg
looks like a yellowish, slightly elongate seed. Eggs usually laid singly on
shoots. One female can lay more than 20 eggs in her lifetime.

Two females fed on honey in the laboratory lived for 28 and 35
days and laid 42 eggs. An unlaid egg was found in the ovaries of both

147

upon dissection. Egg development continued for not less than six weeks.
The laboratory females oviposited on April 13 to 14. Larvae hatched
by May 21 to 23^ i.e., 38 to 40 days after oviposition. Atmospheric tem-
perature during this period ranged from 19 to 33®C (average 23 +
0.4°C). Larvae bore bark without leaving the eggshell. Fine frass is dis-
carded through opening priorly cut in eggshell. Frass is observed on egg
surface for three to five days after commencement of larval activity.

Larvae live in shoots, make longitudinal galleries, plug them loosely
with frass or, more often, discard it through ventilation opening. Dam-
age almost entire tree so that sometimes only bark remains. Frass dis-
carded by larvae collects on soil around shoots. Shoots damaged by
larvae wither and break easily. Diameter of shoots colonized by larvae
0.3 to 1.0 cm. Length of gallery in aerial part of shoot 5.0 to 36.0 cm.
Mature larvae penetrate underground section of stalk or root, make lon-
gitudinal galleries there, and plug them with frass. Length of gallery in
underground section of stalk and root 10.5 cm, width 0.7 to 0.9 cm. Be-
fore or after hibernation in May-June mature larva makes cell in basal
part of shoot, demarcates it above and below with plugs of coarse fibro-
in us frass, and sometimes makes parchmentlike partition before exiting.
Pupa lies with head upward. Length of cell 2.8 to 3.0 cm, width 0.5 to
0,6 cm. Length of lower plug 1.5 to 3.5 cm, of upper one 0.6 to 0.7 cm.

Pupae develop in June-July for not less than three weeks. For ex-
ample, in the laboratory at 22.7°C (+ 0.2®C) pupae developed in 21.9
(± 0.3) days, and at 21.9®C ( + 0.6®C) in 23.8 (+ 0.4) days. In the first
case eight, in the second six insects were kept under observation. Young
beetles break the upper plug of fibrous frass, nibble round flight open-
ing (5.0 to 6.0 mm in diameter) in shoot, and emerge. Records of 29
insects revealed weight of larvae before pupation 101 to 314 mg (183.2
+ 7.4), pupae 80 to 277 mg (161 + 6.9), and adult insects 76 to 201
mg (126.1 + 4.3). Weight of insects during metamorphosis dropped by
an average of 30.7%. Generation completed in two years (Table 9).
Larvae of I- and possibly IT instar hibernate once and mature larvae a
second time before pupation.

Table 9. Periods of derelopment of Polyzonus fasciatuB (F.)

Year of

develop-

ment

April

May

June

July

August

September

October

1st

LPA

PAE

AEL

2nd

3rd

LPA

PAE

AEL

148

Polyzonus fasciatus (F.) inhabits viable shoots of dog rose {Rosa maxi-
mowicziam, R. ragosa, R. dahurica) usually growing in well-warmed
soil. One larva develops on each inhabited shoot. Basal section of shoots
damaged.

21. Tribe ROSALIINI

Adult insect, unlike adults of the tribe Callidiini, characterized by
more convex, cylindrical body and presence of piliform setae on inner
side of antennae. It differs from adults of the tribe Callichromini in
laterally rounded pronotum and absence of aromatic pores on metaster-
num.

Larva characterized by distinct transverse rusty-yellow band in
anterior third of pronotum and reticulate furrows on abdominal loco-
motory ampullae.

Pupa characterized by laterally rounded pronotum, with small tuber-
cle recessed from margin inward, uniformly convex on disk, with pili-
form setae forming medial tuft or transverse band. Abdominal tergites
with short, not very prominent spinules.

The tribe Rosaliini (Compsocerini) is rather small. In Europe,
northern Asia, and North America represented by a single genus
lia), and in southern Asia by two other genera {Pseudocallidium and
Mausaridaem).

1. Genus Rosalia Serv.

Serville, 1833, Ann. Soc. Entom. France, vol. 2,p. 561; Plavifshchi-
kov, 1940, Fauna SSSR, 22, 2, 220”222; Gressit, 1951, Longicorn Beet-
les of China, vol. 2, p. 212; Linsley, 1964, Cerambycidae of North
America, 5, 22, 4-5; Mamaev and Danilevskii, 1975, Lichinki zhukov-
drovosekov, pp, 191-193,

Adult: Antennae apically on 3rd to 6th or 3rd to 8th segments with
dense tuft of hairs on inner side forming brush. Pronotum laterally
112 rounded, recessed from lateral margin, with conical lateral tubercle ex-
tending upward (in Callichromini this tubercle extends sideways, form-
ing lateral margin of pronotum). Body and elytra densely covered with
flat adherent squamiform hairs.

Larva: Head short, markedly retracted into prothorax. Inner masti-
catory lobes of maxillae thick, cylindrical, with parallel sides, apically
almost straightly truncate, with short setae only at this place. Pronotum
with dense uniform hairs almost covering entire disk between anterior
white margin and scutum. Thoracic legs lacking.

Locomotory ampullae coriaceous, rugose-reticulate, developed on
abdominal segments I to VII.

149

Pupa: Head glabrous, without setae, gently rounded to occiput, flat
between antennae and upper lobes of eyes. Antennae curved, looplike.
Pronotum on disk broadly convex, laterally rounded, with very minute
piliform setae forming medial transverse band. Abdominal tergites with
not very large spinules forming transverse rows. Spinules on tergite VII
small, not different from spinules on other tergites.

The genus Rosalia is ecologically associated with broad-leaved forests
prevailing in the Tertiary period. It presently faces extinction. Some
stray species have been preserved: R. alpina (L.) in West and eastern
Europe, R. coelestis Sem. in Ussuri-Primor’e region, R.funebris Motsch.
on the Aleutian Islands and west coast of North America, R. batesi Har.
on Islands of Japan, and over 10 species {R. lameerei Br., R. later itia
Hope, R. decempunctata West., and others) in the south, especially
Southeast Asia.

Type species: Cerambyx alpina Linnaeus, 1758.

1. Rosalia coelestis Sem.

Semenov-Tjan-Shanski, 1911, Rev. Russ. d'Entom., vol. 11, p. 118;
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, [228-230; Cherepanov and

113 Cherepanova, 1977, Taksony fauny Sibiri (Nov. i maloizv. vidy fauny
Sibiri), pp. 150-155.

Adult (Figure 66): Characterized by blue or turquoise pubescence,
black piliform setae on inner side of antennae, and black transverse
bands on elytra. Head short, slightly longer than wide, transversely con-
vex between antennae, with narrow median longitudinal suture, broad
genae, and sparse punctation. Eyes slightly convex, finely faceted, broad-
ly emarginate, between upper and lower lobes with narrow septum.
Antennae longer than body, thin markedly toward apex, with 7th (male)
or 9th (female) segment extending beyond elytral apex, and 3rd to 6th
(male) or 3rd to 8th (female) segments with short dense black hairs at
apex forming brush on inner side.

Pronotum not longer than width at base, uniformly rounded later-
ally, uniformly convex on disk, with dense fine punctation, matte, close to
flanks dorsally with small pointed tubercle, its apex projecting upward.
Scutellum almost semicircular, flat, or slightly broadly impressed. Elytra
with parallel sides, slightly convex on disk, apically with narrowly round-
ed inner angle, gently tapering outer angle, with fine punctation and
dense hairs, and minute glabrous granular tubercles, better developed at
base. Legs slender, comparatively long; hind femora elongate, clavate,
reach hind clivus or almost elytral apex. Hind tarsi 0.50 length of tibiae;
1st segment equal to two successive together or somewhat shorter.

114 Abdomen convex, narrows gradually posteriorly. Abdominal sterni-
tes posteriorly with narrow black lustrous border. Sternite V transverse.

112

Figure 66. Rosalia coelestis Sem,

slightly impressed at posterior margin (male) or notably elongate, gently
rounded posteriorly (female). Body black, with dense compactly adherent
blue or turquoise hairs forming common background against contrasting
black pattern. Head turquoise-blue, genae and underside of head black.
Antennae variegate; 3rd to 11th segments with blue ringlets basally,

151

black apically. Pronotal disk with large tetragonal or rounded black
spot, often joining septum with lateral, minute, round black spot. Scu-
tellum entirely covered with turquoise-blue hairs. Elytra with three
transverse black bands, of which first in anterior third interrupted on
suture, rounded on inner margin, broadening laterally; second in middle
of elytra broad, quite often rounded or angularly produced posteriorly;
third in posterior third, its anterior margin on either side of suture us-
ually projects forward triangularly. Body ventrally entirely covered
with dense adherent hairs; legs with sparser bluish hairy cover. Coxae,
apices of femora, and bases of tibiae glabrous, black. Body length 1.5 to
2.0 mm.

Egg: White, markedly elongate, narrows slightly more toward one
pole than the other, rounded at poles. Chorion with minute coarsely
shagreen sculpture visible under high magnification. Sculpture at poles
prominent. Length 2.8 to 3.1 mm, width 0.7 to 1.0 mm.

Larva (Figure 67): Characterized by transverse rusty-yellow band
and dense hairy cover on pronotum, and reticulate-rugose locomotory
ampullae. Head markedly retracted into prothorax. Anterior margin of
epistoma broadly emarginate, with lustrous rusty-brown border; anterior
angles wartlike, project forward, and here (frontal view) with deep hori-
zontal groove; apex with longitudinal streaklike brownish suture, with
hairy setae in anterior half forming transverse band. Frontal sutures not
visible. Hypostoma consists of two triangular sclerites separated trape-
zoidly by gula, broadens anteriorly, lustrous on anterior margin and
here with short fine setae forming transverse row. Parietals in anterior
half with short erect hairs, behind antennae with three hyaline contigu-
ous ocelli. Clypeus white, hyaline, trapezoid. Labrum convex, narrowly
or broadly rounded apically, in anterior half and laterally with dense
short setae. Mandibles massive, smooth on outer side in anterior half,
rugose or with usual punctation in posterior half. Maxillary stipes later-
ally and cardo basally with dense long setae. Inner masticatory lobes of
maxillae thick, with short setae apically. Labial submentum with pair
of thick, widely separated setae; mentum with setae forming tuft on each
side.

Posterior half of pronotum broadens roundly, anterior margin with
white border behind which transverse rusty band medially and laterally
interrupted by white clearance; anterior margin with two deep longitu-
dinal notches; hairs on disk and laterally dense, uniform, and form
compact field between white border on anterior margin and scutum at
base. Pronotal scutum convex, white, bound laterally by long deep
grooves, with two emarginations on anterior margin, medially bifurcate
there, with longitudinally elongate dots sometimes looking like longitu-
dinal streaks. Alar lobes with dense hairs. Prothoracic presternum uni-

152

113 61 . 1^2iiv2L of Rosalia coelestis Sem.

a — head and pronotum; b — abdominal tergite IV with dorsal
locomotory ampulla.

formly convex, laterally and on disk with dense uniform rusty hairy
115 cover; eusternum glabrous, coriaceous, laterally and anteriorly inconspi-
cuously demarcated from presternum, almost merges with it. Thoracic
legs small, with rusty tinge, and slender claw.

Abdomen narrows insignificantly posteriorly, laterally with rusty
uneven hairs. Dorsal locomotory ampullae divided by common median
longitudinal groove, coriaceous, with wrinkles forming delicate reticu-
late pattern, with two transverse grooves joining laterally with lateral
longitudinal grooves. Tergite IX transverse, with sparse hairs; tergite X
small, rounded posteriorly, glabrous, with stray long hairs only along
posterior margin. Body length 30 to 35 mm, width of head 3.0 to 3.5
mm.

Pupa (Figure 68): Body large, moderately elongate. Head glabrous,
without setae, broadly rounded on occiput, flat between antennae. An-
tennae pressed to sides, in second half bent forward, looplike, with
apices at level of head (male) or level of elytral shoulder (female), and
pressed to underside of thorax.

Pronotum laterally rounded, uniformly convex on disk, medially
with minute piliform setae forming transverse band. Mesonotum slightly

113

Figure 68. Pupa of Rosalia coelestis Sem., female.

convex, almost flat, with stray faint setae or without them, on posterior
margin with gentle, slightly produced scutellum. Metanotum broad and
glabrous, slightly convex, with median longitudinal groove. Hind femora
extend beyond middle of abdominal tergite VII.

Abdomen elongate, narrows from segment IV posteriorly. Abdomi-
nal tergites with large spinules forming transverse row (six to ten para-
medial spinules), minute spinules anterior and posterior to these forming
independent tufts or transverse rows. Tergite VII elongate, rounded pos-
teriorly, in posterior half with dispersed minute spinules. Body length 21
to 28 mm, width of abdomen 6.0 to 7.0 mm.

Material. Collected in Ussuri- Primor’e region. Adult insects 140 (of
which 117 raised in laboratory), larvae 1 9, pupae — three males and four

154

females, larval exuviae with beetles from cells 20. Two generations raised
in the laboratory.

Distribution: Partizansk to Khasan in Ussuri-Primor’e region; north-
east China, North Korea.

Biology: Inhabits broad-leaved forests. Ecologically associated with
dried maple, mainly Acer tegmentosum. Beetles fly early July to early
August. Found in large numbers beginning of last 10 days of July. For
example, we collected 23 beetles in two hours on July 2 1st from trunks
of Manchurian striped maple. Beetles generally found only on trees, not
seen on flowers. Most active in clear warm weather from 12:00 noon to
2:00 p.m. During intense activity beetles respond to rustling caused
by their own movement as well as by extraneous objects. For example,
we attracted beetles resting at a height of up to 5.0 m or more by strok-
ing the tree trunk. They rapidly moved to the stroked area. With the
onset of overcast skies and especially during rains, beetles remain con-
cealed. Mating diurnal. Female oviposits in bark crevices and openings
made by beetles under intact bark. Colonizes only trunks of dry but
upright trees. Does not colonize felled or wind-fallen trees.

Eggs laid singly or in batches of two to five, one after the other.
Duration of egg development three to four weeks. Larvae hatched from
eggs laid in nature in last 10 days of July in second half of August.

116 Larvae live in wood, make longitudinal galleries, and plug them densely
with fine frass. Width of gallery made by mature larva 6.0 to 9.0 mm.
After third hibernation mature larva makes cell along trunk at a depth
of 5.0 cm, nibbles exit to trunk surface, plugs it with coarse fibrous frass,
and pupates with head toward exit. Often nibbles two openings on
trunk surface and plugs them with frass. Distance between two openings
0.5 to 1.0 cm. Size of each opening plugged with frass 4.0 mm X 6.0
mm to 6.0 mm x 10.0 mm.

Pupation of larvae occurs in June. Pupae at room temperature deve-
lop in 12 to 27 days, average 18 days (+ 1.3). Resultant adult insect
remains in cell for not more than one week. It then pushes out frass
from exit, widens opening, and emerges. Sometimes pupal cells are made
in upper layer of wood without exit, in which case beetle nibbles flight
opening in wood and bark (if it remains) and abandons cell through it.
Size of flight opening 4.0 mm x 7.0 mm to 6.0 mm X 12.0 mm. En
masse emergence of beetles from wood occurs in first half or middle of
July. Beetles do not feed, not seen on flowers. Begin to reproduce imme-
diately after emergence. Longevity of beetles, on the average, two weeks,
some living for 24 days or more. One female can lay up to 100 eggs.
Ovaries of three females raised in the laboratory contained 64, 84, and
84 eggs. Gonads of beetles mature in first week after pupal molt.

Weight of larvae before pupation 283 to 637 mg, pupae 194.9 to

155

613.3 mg, and adult insects 130.8 to 453.8 mg. Weight variation during
metamorphosis comparatively small. For example, seven larvae weighed
2,823.5 mg (100%) before pupation, their pupae 2,639.1 mg (93.5%);
another 16 pupae weighed 6,835.3 mg (100%) and their beetles imme-
diately after emerging from wood 4,812 mg (70.4%); i.e., total weight
reduction during metamorphosis 36.1%. Life cycle completed in two
years (Table 10). Young and mature larvae hibernate.

Table 10. Periods of development of Rosalia coelestis Sem.

Year of
development

May

June

July

August

September

October

1st

PAE

AEL

2nd

3rd

PAE

AEL

Rosalia coelestis Sem. develops in trunks of large diameter at a height
of 0.5 to 1 0.0 m. A single tree is often colonized for two to three con-
secutive years and hence populations of different generations found
simultaneously. Rhaphuma acutivittis (Kr.) and Necydalis morio Kr.
colonize with this species.

22. Tribe CALLIDIINI

Adult insect with elongate (Pronocera) or slightly broadened, rather
flat body (Callidium). Head short, between antennae with longitudinal
117 suture. Eyes broadly, often markedly emarginate, almost up to posterior
margin. Antennae longer (especially in male) or shorter than body. Pro-
notum laterally rounded, sometimes angularly produced, often trans-
verse, with dense large (Callidium) or minute (Phymatodes) punctation,
frequently with smooth shields (Rhopalopus, Semanotus). Elytra elongate,
with minute (Pronocera, Phymatodes) or large rugose punctation (Calli-
dium), sometimes longitudinal ribs (Oupyrrhidium). Forecoxae widely
separated (Hylotrupes), more often proximate (Rhopalopus, Pronocera)*
Prosternal process long, narrow, extends beyond middle or even beyond
posterior margin of forecoxae (Rhopalopus, Semanotus, Pronocera, Oupyr-
rhidium) or short, not extending beyond middle of forecoxae (Phyma-
todes), rarely broad and flat (Hylotrupes). Femora clavate; hind femora
thicken gradually in second half (Pronocera) or markedly as though
dilated (Oupyrrihidium, many Phymatodes).

Larva distinguished from larvae of some tribes by white body. Half
of head retracted into prothorax. Epistoma laterally fused with parietals,

156

frontal sutures not visible, median longitudinal suture faint only in
posterior half, anterior margin usually with sharply projecting dark
brown border and oblique longitudinal striation {Rhopalopus, Semanotus)
or with faint narrow indistinct border devoid of longitudinal striation
(Hylotrupes). Pronotum in anterior half with two transverse tetragonal
yellow, yellowish-rust, or bright rusty spots, alveolarly emarginate {Hy-
lotrupes) or not emarginate (remaining genera) on anterior margin,
laterally and on disk with rusty hairs usually forming two transverse
bands anterior to scutum. Pronotal scutum white, bound laterally by
deep longitudinal grooves, with minute sharp or faint longitudinal stria-
tion, rarely without it. Prothoracic presternum laterally with dense very
long hairs, on disk with short hairs; eusternum basally in form of two
round glabrous plates separated anteriorly by quite distinct hairy clear-
ance. Prothoracic eusternum distinct, demarcated laterally by deep groove
{Hylotrupes) or without groove and merges with presternum (remaining
genera). Thoracic legs short and poorly developed, often with faint claw;
legs in some species present only in Il-instar and mature larvae. Loco-
motory ampullae moderately convex, developed on abdominal segments
I to VII, with reticulate-squamiform or shagreen sculpture, matte, often
with silvery sheen.

Pupa characterized by moderately elongate w'hite, sometimes some-
what flat body. Head short and glabrous, without spinules, transversely
convex between antennae, rarely flat. Antennae pressed to sides, in
second half arcuate or bent angularly ventrad and forward. Pronotum
laterally rounded, on disk and base {Rhopalopus) or only base with spinules
{Pronocera, Oupyrrhidium), quite often altogether without spinules {Sema-
notus, Phymatodes). Meso- and metanota glabrous, lustrous, or with
minute spinules forming in posterior half one or two tufts each. Abdo-
minal tergites uniformly convex, with large {Oupyrrhidium) or minute
(some Phymatodes, Semanotus) spinules. Femora clavate.

In northern Asia this tribe is represented by seven genera, of which
six are Holarctic. This indicates that the taxa of the tribe Callidiini
were widely distributed in the pre-Glacial epoch and the evolution of
some {Callidium, Semanotus, Pronocera) proceeded in favor of adapta-
tion to coniferous vegetation, of others {Rhopalopus and most Phymato-
des) to deciduous woody formations. The ecological associations formed
118 then continued not only at the level of species but also of genera. Many
are serious forest pests. Rhopalopus clavipes (F.) severely damages oak
in the Urals, and Semanotus undatus (L.) and Pronocera brevicollis
(Gebl.) spruce-fir forests in the southern regions of Siberia. Many species
of the genus Phymatodes [P. maaki (Kr.), P. ussuricus Plav.] are serious
pests of grapes. For example, in 1971 and 1972, in some forest of
Ussuri-Primor’e region, 30% of more of the grape crop was damaged by

157

these species. Vines colonized by them usually wither while forming
secondary shoots (Cherepanov and Cherepanova, 1974). These species
should be placed under quarantine for regions of Central Asia, the
Caucasus, Crimea, and Moldavia engaged in viniculture.

KEY TO GENERA

Adult Insects

1 ( 2). Third segment of antennae almost 2.0 times longer than 4th,
1.5 times longer than 5th. Prosternal process broad, flat, equal

in width to forecoxae. Base of claw elongate, spinelike

1. Hylotrupes Serv.

2(1). Third segment of antennae less than 2.0 times longer than 4th
or even not longer, quite often equal to 5th or slightly longer.
Prosternal process narrow, narrows markedly toward apex,
significantly less in width than forecoxae. Base of claw smooth,
not spinelike.

3 ( 8). Prosternal process long, extends beyond posterior margin of

forecoxae.

4 ( 7). Elytra monochromatic, without white transverse bands.

5 ( 6). Antennae setaceous, basally thickened; apices of antennal seg-

ments with minute projecting denticle. Body dorsally glabrous.

2. Rhopalopus Muls.

6(5). Antennae slender, acicular, not thickened basally; apices of
antennal segments without projecting denticle. Body dorsally

with hairy cover 3. Pronocera Motsch.

7(4). Elytra with white transverse bands 4. Semanotus Muls.

8(3). Prosternal process short, does not extend beyond middle of
forecoxae; if it does, elytra red with longitudinal ribbing.

9 (10). Elytra vermilion. Third antennal segment not longer than 4th.

5. Oupyrrhidium Pic

10 ( 9). Elytra not vermilion. Third antennal segment not longer than

4th.

11 (12). Pronotal disk with large, rather uniform punctation. Elytra

monochromatic, with metallic sheen 6. Callidium F.

12 (11). Pronotal disk with dense minute (P. mediofasciatus Pic) or

sparse uneven [P. testaceus (L.)] punctation. Elytra monochro-
matic [P. testaceus (L.), P. abietinus Plav. and Lur.] or with
white transverse bands (P. mediofasciatus Pic and others) . . .
7. Phymatodes Muls.

158

1 ( 2).

2 ( 1).

3 ( 8).

4 ( 7).

5 ( 6).

6 ( 5).

7 ( 4).

8 ( 3).

9 (10).

10 ( 9).

11 (12).
12 (11).

Larvae

Prothoracic eusternum laterally demarcated from presternum
by distinct groove, triangular, rounded apically. Found on dry
shoots of dead trees and in buildings .... 1. Hylotrupes Serv.
Prothoracic eusternum not demarcated from presternum, merges
laterally with it, without distinct groove, basally in form of two
round glabrous plates separated anteriorly by hairy clearance.
Locomotory ampullae of abdomen with minute reticulate-
squamiform sculpture (imparting silvery sheen), with transverse
grooves,

Parietals medially without hairy transverse band, only with
sparse stray hairs near brown border on anterior margin.

Head near antennal bases without flat projection, rounded
here; ocelli (one on each side) oriented laterally. Found on deci-
duous trees 2. Rhopalopus Muls.

Head around antennal bases with flat projection, not rounded
here (lateral view), with parallel sides. Ocelli oriented ante-
riorly on flat projection. Found on coniferous trees

3. Pronocera Motsch.

Parietals medially with transverse hairy band between hypostoma
and epi stoma. Found on coniferous trees. ... 4. Semanotus Muls.
Locomotory ampullae of abdomen without minute reticulate-
squamiform sculpture, coriaceous, smooth, rugulose, or sha-
green; if reticulate-squamiform sculpture present, then without
transverse grooves.

Head laterally near antennal bases with flat projection (as in
Pronocera), with parallel sides here. Ocelli (one on each side)
oriented anteriorly. Pronotal scutum with sharp thin longitu-
dinal striation. Found on deciduous trees

5. Oupyrrhidium Pic.

Head laterally near antennal bases without flat projection,
narrows roundly here. Pronotal scutum without striation, or
with very distinct striation, in which case ocelli poorly devel-
oped and oriented laterally or totally imperceptible.

Dorsal locomotory ampullae of abdomen with transverse groove
on anterior margin that arcs posterolaterally. Found on coni-
ferous trees 6. Callidfum F.

Dorsal locomotory ampullae without transverse groove, or
with straight transverse groove that laterally joins lateral longi-
tudinal grooves. Found on grapes and other deciduous species,
rarely (P. abietinus Plav. and Lur.) on coniferous species. . . .
7. Phymatodes Muls.

159

Pupae

1 ( 2). Anterior half of abdominal tergite VII with spinules on disk.

1. Hylotrupes Serv.

2(1). Anterior half of abdominal tergite VII without spinules on
disk, with spinules only posterior to middle forming small tuft,
or without spinules.

3(4). Pronotum with innumerable spinules at base and on disk. . . .

2. Rhopalopus Muls.

4(3). Pronotum with several spinules at base or without them,
glabrous.

5(6). Spinules on pronotum sharp, acicular, form transverse row or

narrow transverse band at base 3. Pronocera Motsch.

120 6 ( 5). Spinules on pronotum acicular, short and thick, or lacking.

7 ( 8). Spinules lacking on pronotum. Found on conifers

4. Semanotus Muls.

8(7). Spinules present on pronotum; if lacking, these species mainly
develop on deciduous species, only some (P. abietimis Plav.
and Lur.) on conifers.

9 (10). Spinules on pronotum mono- or biapical, basally form trans-
verse band or two small tufts 5. Oupyrrhidium Pic

10 ( 9). Spinules on pronotum monoapical, minute, do not form tuft,

dispersed in posterior half, or lacking.

11 (12). Posterior half of pronotum with stray, sometimes barely visi-

ble spinules 6. Caliidium F.

12 (11). Pronotum in posterior half without spinules

7. Phymatodes Muls.

1 . Genus Hylotrupes Serv.

Serville, 1834, Ann, Soc. Entom. France, vol. 3, p. 77; Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 238-239; Linsley, 1964, Cerambycidae
of North America, 22, 5, 14-15; Gressit, 1951, Longicorn Beetles of
China, vol. 2, p. 217.

Adult: Well distinguished from those of other genera by widely
separated coxae, long 3rd antennal segment, and structure of claw.
Head small, frons transverse. Antennae short, slender; 3rd segment
almost 2.0 times longer than 4th, 1.5 times longer than 5th, equal to 1st.
Pronotum transversely oval, with smooth shields on disk. Elytra with uni-
formly rounded humeri, narrow slightly toward apex. Forecoxae widely
separated by broad flat prosternal process. Hind femora with clava, but
not very thickened. Hind tarsi about 0.50 length of tibiae; 1st segment
equal to two successive together. Claws with small denticle at base.

160

Larva: Characterized by distinct prothoracic basisternum, presence
of three ocelli near antennal bases, and other features. Head narrows
anteriorly, with rusty, barely visible border on anterior margin. Parie-
tals laterally with stray hairs. Anterior half of pronotum with tv/o trans-
verse yellow spots with one alveolar white notch each on anterior mar-
gin. Pronotal scutum convex and coriaceous, with longitudinal furrows.
Thoracic legs short. Locomotory ampullae well developed on abdominal
segments I to VII, coriaceous, coarsely rugose.

Pupa: Distinguished from other pupae of Callidiini by large number
of spinules on abdominal tergites. Disk of anterior half of abdominal
tergite VII with spinules forming extensive field (Duffy, 1953).

Genus monotypic, widely distributed in several continents.

Type species: Cerambyx bajulus Linnaeus, 1758.

1 . Hylotrupes bajulus (L.)

Linnaeus, 1758, Syst. Nat., 10th ed., p. 396 (Cerambyx); PlaviP-
shchikov, 1940, Fauna SSSR, 22, 2, 239-242; Duffy, 1953, Monograph
Immat. British Timb. Beetles, pp. 207-214; Durr, 1954, Entom. Memo-
tis, vol. 4, pp. 1-136; Becker, L. 1963, Z. Angewan. Entom., 52, 4,
368-390; Dominik, 1964, Sylwan, 108, 1, 47-52; Demelt, 1966, Tierwelt
Deutschlands, vol. 2, pp. 66-67; Becker, H., 1968, Z. Angewan. Entom.,
61, 3, 251-281; Cymorec, 1968, Z. Angewan. Entom., 62, 3,316-344;
Mamaev and Danilevskii, 1975, Lichinki zhukov-drovosekov,pp. 193-194.

121 Adult (Figure 69): Head slightly convex between antennae, with
median longitudinal suture, uneven punctation, adherent gray hairs bent
down and forward. Eyes broadly and not very deeply emarginate, space
between ocular lobes insignificantly narrower than upper lobe. Antennae
short, slender, do not reach middle of elytra; 1st segment slender, not
longer than 3rd.

Pronotum transversely oval, rounded laterally, with narrow, smooth,
slightly curved border at base, on disk with minute punctation and short
hairs, laterally with dense punctation and dense, long, erect, light-colored
hairs (therefore pronotum laterally appears densely setaceous), and
medially with pair of widely separated smooth shields. Scutellum short,
broadly rounded posteriorly. Elytra moderately elongate, convex, with
rounded humeri, without impression around humeri, with narrowly round-
ed inner angle and gently tapering outer angle at apex, with rugose,
highly uneven punctation, and tender gray hairs forming individual tufts
in form of uneven transverse spots. Pro, meso-, and metathorax with
comparatively dense fine punctation. Abdominal sternites with minute
notchlike punctation. Body ventrally with light-colored semiadherent or
erect hairs. Body dark brown, with brownish or chestnut hue; elytra
with two hairy, faintly spotted transverse bands (f. typica). Color of

161

Figure 69. Hylotrupes bajulus (L.).

elytra and arrangement of hairs on them variable. Sometimes elytra
light brown, legs reddish-rust (ab. puellus Villa), or elytra yellow at base,
122 dark brown otherwise (ab. theresae Pic), or entirely yellowish-brown
with indistinct gray bands, legs black (ab. scutifer Veet.). Body length
7.0 to 20.0 mm.

Egg: White, elongate, broadens more at one pole, broadly rounded
and pointed at the other. Chorion smooth, matte. Length about 2.0 mm,
width 0.5 mm.

Larva (Figure 70): Head narrowly rounded anteriorly, half retracted
into prothorax. Epistoma insignificantly convex, with indistinct rusty
border on anterior margin, slightly emarginate medially, in posterior half
with faint longitudinal suture. Frontal sutures not visible. Hypostoma
somewhat convex, white, on anterior margin without border or with
very narrow rusty border, broad, broadly rounded laterally, divided
medially by fiat gula and basally by slightly broadened gula. Parietals in

162

anterior half with stray hairs, on anterior margin with faint rusty border
not covering antennal bases, receding behind latter, with three unevenly
pigmented or nonpigmented ocelli forming transverse band. Antennae
long, project up to mid length beyond anterior margin of cephalic cap-
sule, with 1st segment not shorter than remainder together. Clypeus
slightly trapezoid, with rusty tinge. Labrum transversely oval, broadly
rounded on anterior margin, with short light-colored hairs. Mandibles
apically broadly rounded, black, basally red; outer side in posterior half
with deep median longitudinal groove. Inner masticatory lobes of maxil-
lae broadly rounded apically, with short setae here. Maxillary palps
short, only last segment projects forward beyond apex of inner lobes.

Pronotum in anterior half with two transverse tetragonal yellowish-
rust spots with one small notch each in anterior margin, laterally with
large yellow spot, on disk anterior to scutum and laterally with short rusty
hairs forming usually compact field. Pronotal scutum convex, coriaceous,
with longitudinal striation, with narrow median longitudinal groove,
laterally with long longitudinal grooves, and almost smooth (non-pro-
jecting) anterior margin and anterior angles. Prothoracic presternum

Figure 70. Larva of Hylotrupes bajulus (L.).
a — head and pronotum; b — abdominal tergite with dorsal locomotory

ampulla.

163

with uniform short rusty hairs; eusternum glabrous, bound laterally by
deep groove, with coarse furrows. Thoracic legs short, more developed
in some specimens, with sharp acicular claw.

Abdomen laterally with very thin light-colored hairs. Dorsal loco-
motory ampullae rather convex, coriaceous, rugose, divided by common
median longitudinal groove, sometimes dentate, sometimes radial and
joining anterior and posterior transverse grooves. Ventral locomotory
ampullae similar in structure, divided by just one transverse groove
merging laterally with lateral longitudinal grooves. Body length of mature
larva 25 mm, width of head 4.0 mm.

Pupa: Head square, rounded on occiput. Mesonotum longitudinal,
convex, with slightly extended scutellum. Metanotum with median longi-
tudinal groove. Abdominal tergites I to VI with minute setaceous spinules
forming transverse oval tuft interrupted medially. Tergite VII triangular,
elongate, narrowly rounded apically, with very large spinules directed
backward on anterior half of disk. Tergite VIII elongate, with almost
parallel sides, narrows slightly posteriorly, with minute paramedial
spinules. Hind femora extend beyond anterior margin of tergite IV. Body
length 14 to 25 mm, width of abdomen up to 7.0 mm (Duffy, 1953).

123 Material: Collected in the European part of the USSR. Adult insect
one, larvae four. Collection of the Zoological Museum, Moscow State
University used for description.

Distribution: Europe, northern Africa, Asia Minor, eastern China,
North America. Indicated for Siberia. We did not find it there, how-
ever.

Biology: Life cycle fairly well studied (Durr, 1954; Duffy, 1953;
Becker, 1968; Cymorec, 1968; and others). Under climatic conditions
of Europe adult insects seen in June and July. Female oviposits in wood
crevices of dead pine, spruce, fir, and other coniferous trees. Often colo-
nizes various structures made of timber, telegraph poles, etc. Female
can lay up to 100 eggs or more in her lifetime. Incubation period one to
three weeks. At 31.5°C larvae begin to hatch nine days after oviposition.

Larvae live in strong dried wood, make meandering galleries, and
plug them with fine white frass. Live for up to three, sometimes seven to
eight years. Mature larva makes cell along trunk in top layer of wood
and pupates in it. Larval pupation occurs in May. Beetles emerge from
pupae two to three weeks later and abandon wood. Life cycle three to
four years, rarely longer.

Causes considerable damage in Europe, destroying wooden structures.
Evidently very rare in Siberia. We did not find it there. Possibly import-
ed sporadically with transportation of dry timber. Assumed to be import-
ed from Europe into North America.

164

2. Genus Rhopalopus Muls.

Mulsant, 1839, Hist. Nat. Col. France Longic., pp. 39-40; PlaviP-
shchikov, 1940, 22, 2, 242-244; Gressit, 1951, Longicorn

Beetles of China, vol. 2, pp. 217-218; Linsley, 1964, Cerambycidae of
North America, 22, 5, 71-72.

Adulv. Characterized by comparatively long antennae extending be-
yond apex of elytra (male) or 11th segment reaching it (female). Pro-
notum uniformly convex, entirely covered with uniform dense punctation
(subgenus Rhopalopus s. str.) or with convex smooth shields, uneven
punctation (subgenus Prorrhopalopus Plav.). Elytra partly flat, almost
glabrous, with short, barely perceptible hairs not forming compact cover.
Legs comparatively long, femora clavate. First segment of hind tarsi equal
to two successive together; first three segments with dense hairy arolium
on underside.

Egg: Elongate, rounded at poles, more often pointed at one. Chorion
smooth, lustrous, transparent.

Larva: Body moderately thick. Head markedly retracted into pro-
thorax. Parietals in anterior half with stray hairs, laterally near antennal
bases with one round convex ocellus. Antennae long, with four segments;
apices project beyond anterior margin of cephalic capsule. Epistoma
only apically with barely visible longitudinal suture, merges laterally
with parietals. Frontal sutures not entirely visible. Anterior half of pro-
notum with yellowish or yellowish-rust spots, on disk with short hairs,
laterally much longer hairs. Pronotal scutum convex, with barely visible
longitudinal striation, basally with fine sculpture. Prosternum anterola-
terally with long reddish-rust setae forming distinct long tuft [Rhopalo-
pus clavipes (F.)] or with usual long light-colored hairs, not forming
prominent tuft. Thoracic legs very minute, with barely visible claw.

124 Locomotory ampullae moderately convex, with minute reticulate- squami-
form sculpture, matte. Abdominal sternite V in region of locomotory
ampullae with pair of round white paramedial ampullae {R. signaticollis
Sols.), or without them.

Pupa: Differs from pupae of other genera in innumerable large spinu-
les on dorsal side, including pro-, meso-, and metanota, which broaden
at base [R. clavipes (F.)], or comparatively minute spinules that do not
broaden at base. Pronotum laterally rounded or angularly produced.

In USSR fauna this genus comprises 12 species. Five inhabit northern
Asia, of which one spread to the central and southern Urals from the
west, four into eastern regions. Six are known in Southeast Asia and one
in North America. Almost all species of this genus are ecologically
associated with broad-leaved woody plants. Hence they are not known
in central Siberia. With the disappearance of broad-leaved forests there

165

in the post-Tertiary period, a discontinuity of geographic range occurred
for the genus Rhopalopus, which serves as an index of the history of
Tertiary fauna that survived the Glacial epoch in remote regions.

Type species: Callidium insubricum Germar, 1824.

KEY TO SPECIES

Adult Insects

1 (2). Pronotal disk entirely covered with dense large punctation,
without smooth clearances and without shields (subgenus Rho-
palopus s. str.). Body black, Europe up to the Urals inclusive. .

1 . R. clavipes (F).

-2 (I). Pronotal disk with sparse uneven, usually minute punctation,
with smooth clearances or with smooth convex shields.

3 (6), Pronotal disk with distinctly projecting smooth shields (subgenus

Prorrhopalopus Plav.).

4 (5). Pronotum black, rarely brownish-rusty (ab. mfithorax Plav.),

at places (between shields and laterally) with dense rugose
punctation. East Asia . . 2. R. signaticollis Sols.

5 (4), Pronotum red, lustrous, with sparse, very fine punctation.

Ussuri-Primor'e region and northeast China. .

3. R. speciosus Plav.

6 (3). Pronotal disk more uniform, without projecting shields (sub-

genus Pronocerodes Plav.).

7 (8). Antennal segments apically with distinctly projecting acute

spinule. Pronotum smooth, highly lustrous, with stray, very

fine punctation. Ussuri-Primor’e region

4. R. auranticollis Plav.

8 (7). Antennal segments apically with faint, barely projecting spinule.

Pronotum with large, distinctly projecting punctures, giving im-
pression of general background of punctation. Ussuri-Primor’e
region and Sakhalin. ............... 5. R. ruficollis Mats.

Larvae

1 (2). Anterior margin of prosternum laterally with very long reddish-

rust setae forming dense tuft. Found mainly on oak.

............................... 1 . R. clavipes (F.).

2 (1). Anterior margin of prosternum laterally without long reddish-

rust setae, with usual thin light-colored or rusty hairs not form-
ing very dense tuft.

125

166

3 (4). Abdominal sternite V with pair of sharply projecting, large,

white, paramedial ampullae in region of locomotory ampullae.

Found on maple 2. R. signaticollis Sols.

4 (3). Abdominal sternite V without round white ampullae. Found

on oak and maple 3. R. speciosus Plav.

Pupae

1 (2). Spinules on dorsal side of body large, broaden markedly at base.

1. R. clavipes (F).

2 (1). Spinules on dorsal side of body minute, do not broaden at

base.

3 (4). Pronotum laterally angularly produced

2. R. signaticollis Sols.

4 (3). Pronotum laterally rounded 3. R. specious Plav.

1. Rhopalopus clavipes (F.)

Fabricius, 1775, Syst. Entom., p. 188 {Callidium); Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 256-258; Demelt, 1966, Tierwelt Deutschlands,
vol. 2, pp. 67-68.

Adult (Figure 71): Characterized by densely punctate pronotum devoid
of smoothly convex shields and matte black elytra notably compressed
posterior to humeri. Head between antennae poorly convex, with faint
median longitudinal suture, dense coarse punctation, spaces between
punctures smaller than punctures. Eyes fairly convex, finely faceted,
widely emarginate, with space between lobes almost equal to 0.50 upper
lobe. Antennae extend beyond apex of elytra (male) or just reach it
(female). Apices of 3rd to 10th antennal segments project acutely, spine-
like on inner side; 5th segment shorter than 3rd, slightly longer than 4th;
1st to 7th segments with black hairs on inner side.

Pronotum narrows gently but significantly anteriorly and more
abruptly posteriorly, broadens angularly posterolateral to middle, near
anterior margin and at base without transverse groove, only on posterior
margin with narrow smooth fringe bent backward, bulges uniformly on
disk, without smooth shields, entirely covered with dense large coarse
punctation, with spaces between punctures lustrous, considerably narrow-
er than punctures. Scutellum not very broad, narrows slightly toward
apex, narrowly or broadly rounded posteriorly, almost not impressed in
middle.

Elytra convex, broaden slightly at humeri and posterior to middle,
notably compressed laterally beyond humeri, at apex with very narrowly
rounded inner angle and gently tapered outer angle, in anterior third
with large and elsewhere very minute rugose punctation imparting matte

167

126

Figure 71 . Rhopalopus clavlpes (F.).

appearance. Spaces between punctures with fine punctation visible under
high magnification. Pro-, meso-, and metathorax with dense deep punc-
tation, abdomen with more minute, less dense, notchlike punctation.
Body ventrally with short brownish hairs. Femora with short, more
sharply thickened clava. Entire body, antennae, and legs black. Elytra
black, lustrous at base, dull in middle and posterior third. Body length
12 to 18 mm.

168

Egg: White, oval, slightly elongate, narrows markedly toward poles
and narrowly rounded there. Chorion smooth, hyaline, transparent. Length
1.8 mm, width 0.9 mm.

Larva (Figure 72): Readily recognized by long setaceous rusty hairs
laterally on anterior margin of pronotum, forming dense tuft. Head

126 markedly retracted into prothorax. Epistoma insignificantly convex, with
brownish-rust border on anterior margin obliquely, not sharply striate
(streaks diverge from posterior margin laterally). Frontal sutures not
visible, median longitudinal suture of epistoma smooth, barely percep-
tible apically. Hypostoma broadens with narrow brownish border, sharp
(not rounded) outer anterior angles. Gula slightly rounded anteriorly,
does not extend beyond anterior margin of hypostoma, with transverse
groove at base. Parietals in anterior third with stray hairs. Near base of
antennae from underside convex round hyaline ocellus with black spot
recedes backward and deeply concealed in cuticle. Antennae compara-
tively long; 1 st segment slightly shorter or almost not shorter than two
successive together. Clypeus semitransparent, widely flattened basally.
Labrum broadly rounded anteriorly, narrowly and sharply rounded pos-
teriorly, convex and glabrous on disk, with sparse short setae along
margins. Mandibles narrow notably toward apex, rounded apically, on
outer side with narrow transverse groove, black, basally with rusty tinge.
Inner masticatory lobes of maxillae thick, taper slightly toward apex,
whitish, with sparse setae, lustrous on outer side, glabrous, brownish.
Maxillary palps distinctly longer than inner lobes.

Pronotum broadly rounded in posterior half, narrows anteriorly, in
anterior half with two sharp, tetragonal, transverse, rusty spots, laterally
with less distinct small spot, rusty hairs forming two transverse bands on
disk, one denser in region of rusty spots and second thinner anterior to
scutum. Hairs on disk (especially in region of rusty spots) comparatively
short, ringed with sclerotized ringlet, on sides of pronotum long. Prono-
tal scutum white, bound laterally by straight longitudinal grooves, with
very minute longitudinal striation. Prothoracic presternum with not very
dense short hairs, on anterior margin laterally with long dark rusty,
almost red setae forming two distinct tufts; eusternum glabrous, with

127 faint longitudinal furrows or striation. Meso- and metasterna rugulose,
with transverse groove interrupted broadly in middle. Thoracic legs small,
with minute claw.

Abdomen thick, narrows moderately from thorax toward posterior
end, with rusty hairs laterally. Dorsal locomotory ampullae convex, cori-
aceous, divided by common median longitudinal groove, transverse groove
in front merging laterally with lateral longitudinal grooves. Ventral loco-
motory ampullae laterally in anterior half with short bracketlike trans-
verse groove curved backward. Abdominal sternites, near ventral locomo-

169

Figure 72. Larva of Rhopalopus clavipes (F.).

a — ^head and pronotum; b — prosternum; c — abdominal
tergite IV with dorsal locomotory ampulla.

tory ampullae, with long rusty-red hairs forming tuft on each side. Body
length of mature larva 20 to 25 mm, width of head 2.8 mm. In I-instar
larvae abdominal segment VI laterally on posterior margin with short
spinule, VII with very long acute brownish spinule, its apex directed
backward. These spinules disappear after molt.

Pupa (Figure 73): Characterized by large flat dorsal spinules that
128 broaden basally, rounded or pointed apically. Head short, between anten-
nae insignificantly transversely convex, with median longitudinal suture
there, flat on vertex between upper ocular lobes, broadly rounded, gla-
brous, without setae on occiput. Antennae pressed to sides, bent forward,
looplike, in posterior third in male, bent ventrad in female.

170

127

Figure 73. Pupa of Rhopalopus clavipes (F.).

Pronotum transverse, broadens angularly posterior to middle, from
there narrows gradually anteriorly and more steeply posteriorly, bulges
gently on disk, with barely perceptible minute transverse striation, medi-
ally with minute spinules, on hind clivus with large, flat, triangular, some-
times biapical setaceous spinules forming two transversely elongate bands,
basally with narrow transverse groove.

Mesonotum transversely impressed in second third, extended convex
or elevated scutellum on posterior margin, minute sharp, usually paired
spinules medially in posterior half. Metanotum bulges slightly, on poste-
rior margin broadly rounded, on disk posterior to middle with flat spinu-
les bent forward and forming broad tuft.

Abdomen in region of segments III and IV broadens distinctly, nar-
rows markedly posteriorly. Abdominal tergites broad and convex, with
faint flat median longitudinal groove; abdomen laterally in anterior half

171

with minute paired spinules forming transverse row, inpo sterior third
with very large (tergites I to IV) or minute (tergites V to VII) flat spinu-
les forming transverse row or transverse band (three to nine paramedial
spinules). Spinules bent sideways, apically more sclerotized, rounded or
pointed, basally broaden, light yellow. Tergite VII 1.5 times wider at
base than long, convex on disk, with narrowly rounded posterior margin.
Tergite VIII with rows of small paired spinules on disk broadly rounded
apically. Valvifers of female hemispherical, proximate. Hind femora
clavate, pressed to sides of abdomen, with apices more (male) or less
(female) extending beyond abdominal tergite V. Body length 16 to 22 mm,
width of abdomen 5.0 to 7.0 mm.

Material: Collected in the southern Urals (from Orenburg to Ural’sk).
Adult insects 22, larvae 40, pupae — eight males and five females, larval
and pupal exuviae from cells with beetles six. Adults and pupae raised in
the laboratory from larvae collected in nature.

Distribution: From Atlantic Ocean coast to the Urals inclusive, from
Denmark, Sweden, Leningrad, Upper Volga to the Caucasus, Iran, Syria,
and the Mediterranean Sea. Sporadic everywhere. Common in broad-
leaved forests of the southern Urals.

Biology: Inhabits broad-leaved forests. Ecologically associated with
oak and other deciduous species. Beetles fly from end of May through
August. Found maximally (southern Urals) in early July. Lead cryptic
mode of life, not seen on flowers. Emerge from pupal cells VN^ith develop-
ed gonads. Ovaries of one female before emerging from pupal cell con-
tained 54 fully developed eggs. During reproduction period beetles found
on drying trees. After mating, female oviposits in bark crevices, mainly
on branches 3.0 to 5.0 cm in diameter at a height of up to 2.0 m or more
above ground.

Larvae hatch three weeks after eggs are laid. In the Urals larvae
began hatching in a forest (Krasnokholmsk reserve) on July 25th from
eggs laid on July 7 to 1 1 . Atmospheric temperature varied during this
period from 13.8 to 32.2°C (average 22.1+0.5®C). Newly hatched larva
129 immediately bores bark and makes a gallery initially in bark along shoot,
sometimes with lateral branches from it. 11- instar larvae make platform-
like galleries under bark imprinted on alburnum. These galleries are
usually hollow, not filled with frass, with gentle, at places steep walls.
Lateral branches (blind alleys), filled with frass, proceed from platform
gallery. Length of platform gallery under bark 7.0 to 11.0 cm, width 1.5
to 3.5 cm. However, configuration and length of gallery depend on dia-
meter of shoot colonized by larvae. Galleries in thin shoots are usually
long and narrow, in thick ones broader, platformlike. Quite often, many
larvae live side by side. Galleries made by them fuse and form an exten-
sive area fully or partly filled with frass. In one case three larvae made

172

a gallery 24 cm long in a blackthorn shoot 1.4 cm in diameter, with a
total area of 48 cm^. In another case one larva made a gallery 38 cm long
with a total area of 31.6 cm^ under the bark of a blackthorn shoot 3.0 cm
in diameter. In an apple tree trunk 10.5 cm in diameter a larva made a
gallery, 14 cm long with an area of 26 cm^.

Mature larvae bore wood and make a longitudinal gallery free of
frass which later serves as a pupal cell. Inlet plugged v/ith frass. Width
of inlet 0.7 to 1.0 cm. Length of cell (gallery free of frass) 7.5 cm, width
1.2 cm.

Pupation of larvae commences early May, ceasing in June. In the
laboratory at 1 8.3 +0.2°C pupae developed for 15 to 20 days (average
17.6+0.5 days). Young beetles mainly seen in June. Emergence of adults
from wood ceases toward end of June. In 1978 emergence had ceased
by last 10 days of June in forests of the southern Urals. Life cycle com-
pleted in two years (Table 11). Records of 22 insects revealed: weight of
larvae before pupation 128 to 530 mg (245.6 + 20.8), pupae 116 to 422
mg (211.3+18.9), and beetles before emerging from wood 94 to 335 mg
(169 + 15.8). However smaller insects were sometimes seen. For example,
one larva weighed 67 mg before pupation, pupa developing from it 58
mg, and beetle before em.erging from cell 47 mg.

Table 11. Periods of development of Rhopalopus clavipes (F.)

Year of

development

April

May

June

July

August

September

October

1st

LPA

PAE

AEL

2nd

3rd

LPA

PAE

AEL

While inspecting forests in the southern Urals we found 39 insects in
larval and adult stages. Of these seven were found on oak, 16 on haw-
thorn, four on apple, three on bird cherry, three on willow, one on cherry,
and five on blackthorn {Primus spinosa). Generally colonizes thin shoots
1.4 to 5.0 cm in diameter, more rarely 13 cm in diameter. Not found on
thicker trunks. Recently dried trees are usually colonized. Population
density comparatively high. On a cutting 20 cm long and 10.5 cm in dia-
meter seven larvae were found in wood; three mature larvae were recover-
ed from a blackthorn shoot 25 cm long and 1.4 cm in diameter. Phyma-
todes testaceus (L.) and Xylotrechus antilope (Schonh.) often colonize
oak together with this species, and Mesosa myops Dalm. and others
colonize apple simultaneously.

173

130 2. Rhopalopus signaticollis Sols.

Solsky, 1872, Horae Soc. Entom. Ross., vol. 9, p. 177; Plavirshchikov,
1940, Fauna SSSR, 22, 2, 246-247; Gressit, 1951, Longicorn Beetles of
China, vol. 2, p. 218.

Adult (Figure 74): Characterized by dense deep punctation and large
convex lustrous shields on pronotum and flat dull elytra. Head short,
appears transverse in dorsal view, bulges uniformly transversely between
antennae, with narrow median longitudinal suture, broad and moderate-
ly impressed between upper ocular lobes, with dense punctation. Eyes
finely and distinctly faceted, with broad deep notch, very narrow septum
between upper and lower lobes. Antennae longer than body; 8th or 9th
segment in male, 10th or 11th segment in female extends beyond apex of
elytra; 5th to 9th segments apically produced, spinelike; 10th and 11th
segments uniform; 5th segment longer than 4th, shorter than 3rd.

Pronotum transverse, medially notably wider than long, angularly
rounded or tubercularly produced laterally, with dark brown erect hairs,
narrows less anteriorly, more posteriorly, with narrow flange on posterior
margin, and five smooth lustrous convex shields on disk; of these, two
lateral in anterior half, two in posterior half, and one medial. Lateral
shields sometimes fuse into longitudinal smooth outcurved bands. Gaps
between shields and sides of pronotum with dense large punctation. Scu-
tellum broad, its length not more than width, broadly rounded posterior-
ly, with stray large punctation.

Elytra with parallel sides, flat, broadly rounded posteriorly, with
straight humeral tubercles projecting slightly laterally, more lustrous
basally, with large rugose punctation, matte farther away from scutellum,
with dense, very fine punctation, and short, faint, semiadherent black
hairs. Metathorax with dense punctation, abdomen with very sparse
notchlike punctures. Legs long. Second half of femora thickened, clavate.
Body ventrally with light-colored or brownish hairs. Body black or
brownishblack. Elytra with brownish tinge in second half (f. typica).
Sometimes pronotum brownish-rust (ab. riifithorax Plav.). Body length
9.0 to 14.0 mm.

131 Egg: White, elongate, narrows more toward one pole and pointed,
rounded at the other. Chorion smooth, hyaline, transparent. Length 1.6
mm, width 0.6 mm.

Larva (Figure 75): Unlike larvae of other species, readily recognized
by two round white ampullae on abdominal sternite V. Head narrowly
rounded anteriorly, markedly retracted into prothorax. Epistoma very
short, anteriorly with reddish-brown margin, darkened laterally on cly-
peus and here with faint, somewhat obliterated, oblique striation; median
longitudinal suture slightly translucent only at apex, frontal sutures
not visible. Hypostoma broadens anteriorly, rounded on anterior outer

174

130 Figure 74. Rhopalopus sigmticollis Sols.

angles, anteriorly with narrow rusty border, with small notch near inner
angles. Gula flat, does not project posterior to anterior margin of hypo-
stoma. Parietals in anterior half with sparse short hairs. Ocelli near anten-
nal bases ampullar, hyaline. Antennae long, slender, project forward
beyond margin of cephalic capsule. Clypeusvery short, broadens marked-
ly at base. Labrum small, convex, narrows anteriorly, narrowly rounded
apically, with dense piliform rusty setae along margins. Mandibles thick,
132 massive, uniformly rounded apically, matte on outer side, basally with
broad median longitudinal groove, black, reddish-rust at base. Maxillary
palps long, distinctly extend forward beyond apex of inner masticatory

175

131

Figure 75. Larva of Rhopalopus signaticollis Sols,
a — head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla; c — prosternum

d—abdominal sternite V.

lobe of maxillae. Labial mentum laterally with long setae forming tuft
on each side.

Pronotum broad, with not very sharp yellow spots in anterior half,
two on disk and one longitudinally elongate on each side, and also in
anterior half with comparatively uniform rusty hairs forming common
field, somewhat thinned in middle (hairs without basal ringlet). Pronotal

176

scutum white, convex, with dense, barely perceptible longitudinal stria-
tion, significantly produced medially on anterior margin and slightly so
on anterior angles, bound laterally by deep longitudinal grooves. Pro-
thoracic presternum with dense fine rusty hairs, laterally on anterior
margin [unlike larva of R. clavipes (F.)] with ordinary hairs not forming
long tuft; eusternum glabrous, with fine furrows, and shagreen matte tone.
Thoracic legs very minute, with weakly sclerotized claw.

Abdomen laterally, especially ventrally, with dense long light-colored
hairs. Dorsal locomotory ampullae moderately convex, shagreen, with
very fine reticulate- squamiform sculpture, with broad, common, median
longitudinal groove, laterally with short longitudinal grooves, on anterior
margin without transverse groove. Ventral locomotory ampullae similar
in structure, on sternite I laterally with transverse groove, on II to IV with
bracketlike replicate groove, on V and VI with short longitudinal groove,
and on VII with short longitudinal groove and transverse groove reced-
ing inward from it. In addition to locomotory ampullae, sternite V with
pair of round white paramedial ampullae. Body length of mature larvae
18 to 23 mm, width of head 2.9 mm.

Pupa (Figure 76): Characterized by innumerable spinules on dorsal
side of body and broad, slightly convex frons. Head short, narrows
anteriorly from antennal bases, slightly convex between antennae, barely
impressed between upper ocular lobes, uniformly rounded on occiput.
Antennae comparatively long, pressed to sides, with apices bent ventrad
and forward.

Pronotum transverse, angularly produced laterally, narrows toward
base, convex, transversely striate on disk, with innumerable spinules
forming distinct or less prominent transverse bands interrupted medially.
Spinules short, broaden basally, setaceous. Mesonotum convex, with
narrow median longitudinal groove, on posterior margin with elevated
scutellum, with minute spinules forming two paramedial tufts, each
anteriorly elongate and inclined. Metanotum broad, rounded posteriorly,
fairly convex, with median longitudinal groove, with well-developed spi-
nules forming two bands diverging from middle to anterior angles. Hind
femora with long, gradually thickening clava.

Abdomen elongate, with almost parallel sides, narrows from segment
V posteriorly. Abdominal tergites convex, with common median longi-
tudinal groove, with short pointed spinules forming characteristic un-
even transverse band of stray tufts, two (with three spinules per tuft)
posterior to middle nearer longitudinal groove, and two (with eight to
ten spinules per tuft) transversely elongate slightly posterior. Tergite VII
narrowly rounded posteriorly, with spinules on disk forming small com-
mon tuft immediately posterior to middle. Tergite VIII moderately elon-
gate, broadly rounded posteriorly, with stray minute spinules. Tip of

177

133 abdomen (ventral view) distinctly produced. Valvifers of female conti-
guous, slightly enlarged apically, lustrous at apex, with distinct minute
folds. Body length 12 to 16 mm, width of abdomen 4.0 mm.

Material. Collected in Ussuri-Primor’e region (Arsen’ev, Gornotae-
zhnoe, Komarovka River). Adult insects 35, larvae 20, pupae — two
males and three females, larval exuviae with beetles from cells nine.

Distribution: Amur region, Ussuri-Primor’e region, northeast China,
Korea, northern Japan.

Biology: Inhabits broad-leaved forests. Beetles seen in June up to
end of July. Inhabit dried and drying maple, creeping onto trunk to
mate. Female oviposits under bark scales or in wood crevices. Some-
times colonizes viable trees with cavities. Larvae hatched in 1.8 to 25
days from eggs laid in the laboratory at 13.0 to 19.4®C.

Newly hatched larvae bore bark, make longitudinal meandering
galleries imprinted on wood under bark, and plug them densely with
fine frass. Before second hibernation mature larva bores wood, makes

178

longitudinal gallery 2.0 to 4.0 cm long in upper layer to a depth of 0.2
to 2.0 cm, and plugs inlet with fine frass. Size of inlet 0.50 cm X 0.25
cm to 0.90 cm X 0.30 cm. Larvae remain in wood for second hiberna-
tion. Pupate in spring. Pupae oriented with head toward inlet.

Young beetles nibble oval flight opening (0.35 cm x 0.15 cm to 0.60
cm X 0.30 cm) in bark surface and emerge. Weight of larvae before
pupation 32.0 to 130.1 mg, pupae 29 to 117 mg, and young beetles
before emerging from cell 24 to 95 mg.

Rhopalopus signaticollis Sols, inhabits maple {Acer mono, A. tegmen-
tosum, and others). While inspecting forests in Ussuri-Primor’e we col-
lected 55 insects in larval, pupal, and adult stages. All of them came from
maple. Not found on other species. Necydalis morio Kr., N. ebenina Bat.,
and others sometimes colonize together with this species.

3. Rhopalopus speciosus Plav.

Plavilstshikov [Plavil’shchikov], 1915, Entom. Vestn., vol, 2, p. 108;
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 247-248.

Adult (Figure 77): Differs from other species of the genus Rhopa-
lopus in sparse fine punctation and red color of pronotum. Head be-
tween antennae transversely highly convex, with faint median longitudi-
nal suture, uneven punctation, smooth, not punctate near antennal
bases from inner side and on vertex. Eyes black, sharply faceted, broad-
ly emarginate, septum between lobes comparatively broad, barely narrow-
er than upper lobe. Antennae thin markedly toward apex, with 9th
(male) or 11th (female) segment extending beyond apex of elytra, long
black hairs on underside of 1st to 6th segments, with projecting spinules
apically on 3rd to 8th. Fifth antennal segment considerably shorter
than 3rd, equal to 4th.

Pronotum transverse, with transverse groove near anterior and pos-
terior margins, angularly or clavately produced laterally, with very fine,
smoothened, sometimes uneven punctation on disk, with convex smooth
shields, of which two on each side in anterior and posterior half, one
anterior to base medially. Sometimes shields greatly smoothened, faint.
Scutellum broad, smooth, with stray punctation or without, medially
impressed, troughlike.

134 Elytra with parallel sides, convex, with barely projecting humeral
tubercles, apically with narrowly rounded inner angles and broadly
rounded outer, dense large punctation in anterior third or only at base
and here comparatively lustrous, elsewhere with unusually fine puncta-
tion imparting matte tone. Metasternum and abdominal sternites with
dense fine punctation. Body ventrally with short, not very dense gray
hairs. Femora with long distinct clava. Body, antennae, and legs black
or blackish-brown, sometimes with chestnut tinge. Pronotum and pro-

179

sternum red or reddish-rust. Elytra dark violet or dark blue, in anterior
third with metallic sheen, in posterior half matte. Body length 13 to 15
mm.

Egg: White, elongate, rounded at one pole and pointed at the other.
Chorion smooth, hyaline, transparent. Length 2.1 mm, width 0.6 mm.

Larva (Figure 78): Similar to larva of R. signaticollis Sols., but well
distinguished from it in absence of round white ampullae on abdominal
sternite V and sparse short hairy cover laterally on abdomen. Head
markedly retracted into prothorax, glabrous laterally, with only stray
hairs around anterior brownish border. Base of antennae with single

180

convex, light-colored, hyaline, or brownish ocellus. Antennae long, slen-
der, project forward beyond anterior margin of cephalic capsule. Epi-
stoma insignificantly convex, on anterior margin with broad, reddish-
brown, smooth, nonstriate border, anteromedially with faint or almost
invisible longitudinal suture, merges laterally with parietals; frontal suture
not visible. Hypostoma flat, broadens anteriorly, on anterior margin
with narrow brownish-rust border. Anterior margin of gula at level of
anterior margin of hypostoma or slightly short of it. Clypeus short, barely
projects beyond anterior margin of hypostoma as transverse band. Labrum
white, narrows anteriorly, narrowly rounded apically, with short light-
colored setae along margin. Mandibles black, reddish-brown basally, with

135

Figure 78. Larva of Rhopalopus speciosus Plav.
a — head and pronotum; b-~abdominal tergite IV
with dorsal locomotory ampulla; c — abdominal
sternite V.

181

135 median longitudinal groove on outer side closer to base resembling deeply
impressed stria. Inner masticatory lobes of maxillae thick, insignificantly
narrower than long, apically obtuse and with stray setae here. Maxillary
palps distinctly longer than inner lobes. Labial mentum posterolaterally
with setae forming tuft.

Pronotum in anterior half on disk with two broad indistinct yellow
spots, laterally with broad longitudinally elongate yellow spot covering
almost entire lateral section, on disk with not very dense, short, light-
colored hairs forming two transverse bands, of which one more distinct
on anterior margin of yellow spots and second less distinct (diffuse)
anterior to scutum. Pronotal scutum basally convex and here with very
minute, reticulate-squamiform sculpture, flat anteriorly, sometimes with
minute, barely perceptible longitudinal striation, bound laterally by
short longitudinal grooves. Prothoracic presternum on disk with short
hairs, laterally with very long, thin, light-colored hairs not forming dis-
tinct tuft; eusternum glabrous, coriaceous, rugulose. Thoracic legs minute,
with sharp claw.

Abdomen laterally with very sparse short hairs, looks almost gla-
brous. Dorsal locomotory ampullae matte, convex, with common broad

136 median longitudinal groove, laterally with faint longitudinal grooves,
transverse groove lacking. Ventral locomotory ampullae on sternites I to
III and VII with transverse grooves, on sternites IV to VI with lateral
longitudinal grooves; abdominal sternite V without round white ampul-
lae in middle of locomotory ampullae. Body length of mature larvae 21
to 25 mm, width of head 3.0 mm.

Pupa (Figure 79): In arrangement of spinules on abdominal tergites
similar to pupa of R. signaticollis Sols. Differs in well-rounded, not
angularly produced pronotal flanks. Head moderately bent under, narrows
anterior to antennae, between upper ocular lobes flat or slightly impress-
ed, broadly rounded on occiput. Antennae pressed to sides, with apices
bent forward, looplike.

Pronotum laterally rounded, narrows almost identically anteriorly
and posteriorly, uniformly convex on disk, with faint minute transverse
striation, minute setaceous spinules forming two transverse bands, one
broader basally and second obscure in anterior half. Stray spinules
occur between these two bands. Mesonotum convex, lustrous, with al-
most straightly truncate posterior margin, and minute spinules forming
two paramedial tufts. Metanotum broad, slightly convex, with almost
straightly truncate posterior margin, median longitudinal groove, not
very large spinules forming extensive tuft extending longitudinally from
middle of base to anterior angles. Femora with long clava, pressed to
sides of body.

182

135

Figure 79. Pupa of Rhopalopus speciosus Plav.

Abdomen with parallel sides or broadens slightly in region of seg-
ment IV, narrowing posteriorly from segment V, and produced at tip.
Abdominal tergites moderately convex, medially with narrow, compara-
tively deep groove, with short spinules forming broad transverse band
(20 to 36 spinules per tergite). Abdominal tergite VII narrows conically
posteriorly, pointed there, with small spinules in middle of disk forming
small (up to six spinules) or large (over 20 spinules) tufts. Tergite VIII
insignificantly elongate, with hyaline sheen and stray, barely perceptible
spinules. Valvifers of female contiguous, almost hemispherical, do not
broaden apically but rounded. Body length 14 to 16 mm, width of
abdomen 4.0 mm.

183

Material: Collected in Ussuri-Primor’e region (Komarovka River).
Adult insects three, larvae seven, pupae— two females, larval exuviae
from cells with beetles three.

Distribution: Ussuri-Primor’e region and northeast China.

Biology: Inhabits broad-leaved forests. Beetles fly from end of June
to August. According to T.P. Samoilov (1936), beetles sighted on flowers
of Sorbaria sorbifolia. However, they emerge from wood with developed
gonads and are capable of reproducing without supplementary feeding.
For example, ovaries of one female dissected immediately after emer-
gence from wood contained 38 mature eggs. Female oviposits on trunks
of dried oak and maple. Larvae live under bark, make straight or,
more often, meandering galleries and plug them densely with fine frass.
Mature larva bores wood to a depth of up to 0.4 cm, makes cell along
trunk, and remains in it for second hibernation. Inlet to cell plugged
with fine frass. Length of pupal cell up to 2.5 cm, width 0.8 cm.

Larvae pupate in spring. Young beetles seen in June. They aban-
don pupal cell through inlet and nibble oval flight opening (5.0 mm X
3.5 mm to 6.0 mm x 3.5 mm) in bark and exit. Records of six insects
showed: weight of larvae before pupation 83.0 to 223.4 mg, pupae 72 to
137 201 mg, and young beetles before emerging from wood 58 to 161 mg.
One larva weighed 309.8 mg. While inspecting forests, larvae, pupae,
and adults of this species were found on maple.

4. Rhopalopus aurantiicollis Plav.

Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 250-251.

Adult: Unlike other species of this genus, recognized by pronotum
devoid of punctation and other ornamentation. Head with stray sparse
punctation, sparse erect hairs, and distinct longitudinal suture between
antennae. Antennae extend insignificantly beyond apex of elytra (male)
or just reach it; 1st to 5th segments with dense hairs on lower side, 3rd
to 9th segments with projecting apical spinule.

Pronotum slightly wider (female) or not wider (male) in middle than
long, rounded or slightly angular laterally, near base with narrow trans-
verse groove or narrow flange (female), smooth fringe medially on anterior
margin; disk slightly convex, with small round paramedial impression,
smooth, lustrous, with sparse erect hairs. Scutellum slightly elongate,
highly impressed, smooth.

Elytra elongate, 3.0 times longer than width at base, posterior to
humeri slightly or notably compressed, obliquely rounded apically (with
narrowly rounded inner angle and broadly tapered outer), notably flatten-
ed, with large flat punctation at base, minute dense punctation else-
where, lustrous in anterior third, posteriorly matte. Prosternum with
punctation and transverse striation. Metasternum with sparse puncta-

184

tion on disk, very dense but minute punctation laterally. Abdominal
sternites with sparse punctation. Body ventrally with sparse thin brown-
ish hairs. Body black, sometimes brownish. Pronotum orangish-red,
on posterior margin with black border. Prosternum orange, prosternal
process darkened. Elytra dark brown, with faint violet tinge. Body length
12.5 to 13,0 mm.

Distribution'. Ussuri -Primor’e region, southern Sakhalin. Described
from collection of the Zoological Museum, Moscow State University.
Rare. We did not find it in southern Primor’e fauna even after many
years of study. Judging from the collection, beetles fly from July through
August.

5. Rhopalopus ruficollis Mats.

Matsumura, 1911, /. Coll. Agric. Sapporo, 4, 1, 138; Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 249-250.

Adult: Differs from other species in red pronotum with two impres-
sions and uneven punctation. Head with moderate punctation, sparse
erect hairs, longitudinal suture between antennae, and smooth between
upper ocular lobes, without punctation. Antennae slender, extend beyond
apex of elytra; 5th to 7th segments produced apically, spinelike.

Pronotum transverse, broadens angularly or unevenly laterally, with
sparse erect hairs there, broadly fringed on anterior margin and narrow-
ly on posterior, on disk with flat rugose punctation, medially with
smooth band with longitudinal impression along each side of band.
Scutellum comparatively elongate, longitudinal, smooth, with trough-
like longitudinal impression.

138 Elytra elongate, long, 3.0 times longer than width at base, slightly
compressed posterior to humeri, obliquely rounded apically, in anterior
third coarsely punctate, lustrous, from middle third with minute rugulose
punctation, matte. Prosternum with rugulose punctation, metasternum
on disk with sparse and laterally very dense punctation. Abdominal
sternites with sparse minute punctation. Body ventrally with sparse short
brownish hairs. Body black or blackish-brown, pronotum rusty-red, some-
times darkened on posterior margin and in middle. Pronotum in anteri-
or half reddish-rust. Elytra violet or purple. Body length 12 to 14 mm
(Plavil’shchikov, 1940).

Distribution: Ussuri-Primor’e region, southern Sakhalin. Rare. Only
stray specimens known. We did not find this species in our long-term
investigations.

3. Genus Pronocera Motsch.

Motschulsky, 1875, Bull. Soc. Nat. Moscow, 49, 1, 149; Plavil’shchi-
kov, Fauna SSSR,22, 2, 261-262; Gressit, 1951, Longicorn Beetles

185

of China, vol. 2, p. 219; Linsley, 1964, Cerambycidae of North America,
22, 5, 12-13.

Adult: Characterized by elongate body, slender antennae, and compa-
ratively broad notch between lower and upper ocular lobes. Pronotum
transverse (female) or almost square (male), with uneven punctation on
disk. Prosternal process long, slender, completely divides forecoxae.
Mesosternal process narrows posteriorly, rounded or almost emarginate
apically. Elytra elongate, with parallel sides, with rugulose and minute
punctation. Femora with insignificantly, gradually thickening clava.

Larva: Distinguished from larvae of other genera by well-developed
ocelli on anterior flat part of parietals behind antennal bases. Head
with parallel sides, glabrous mediolaterally, without transverse hairy
band, with only stray sparse hairs around brown border on anterior
margin. Pronotal scutum white, mth longitudinal striation, bound lateral-
ly by longitudinal grooves. Thoracic legs short, with sharp claw. Loco-
motory ampullae developed on abdominal segments I to VII, with very
fine sculpture, matte.

Pupa: Characterized by elongate body. Antennae arcuate in second
half. Pronotum lustrous, rounded laterally, transversely striate on disk,
basally with well-developed spinules forming transverse band. Abdomi-
nal tergites with minute spinules forming transverse indistinct row or
narrow transverse band interrupted medially.

The genus Pronocera Motsch. is very small, with just three species
included in it. Of these, P. angusta Kriechb. inhabits West Europe, P.
brevicollis (Gebl.) southern regions of northern Asia, and P. collaris
Kirby the boreal zone of North America. All three species are ecologi-
cally associated with coniferous vegetation — Picea, Abies, and Pinus.

Type species: Callidium brevicollis Gebler, 1833.

1. Pronocera brevicollis (Gebl.)

Gebler, 1833, Bull. Soc. Nat. Moscow, vol. 6, p. 302 {Callidium)',
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 263-265; Gressit, 1951,
Longicorn Beetles of China, vol. 2, p. 219; Cherepanov and Cherepa-
nova, 1973, Nov. i maloizv.vidy fauny Sibiri, 6th ed., pp. 68-69.

Adult (Figure 80): Head between antennae highly convex transversely,
with median longitudinal suture, dense punctation, and brown hairs.

139 Eyes broadly emarginate, septum between ocular lobes slightly narrower
or almost not narrower than width of upper lobe. Antennae slender, in
female extend beyond 0.75 length of elytra, in male beyond elytral apex;
2nd segment short, nodular; 3rd distinctly longer than 5th.

Pronotum wider (female) or almost not wider (male) than long, gently
rounded (male) or slightly angularly produced laterally, on disk with sparse
and laterally denser punctation, with light brown hairs, medially usually

186

140

with longitudinal smooth band. Scutellum small, flat, narrowly rounded
posteriorly.

Elytra elongate, with parallel sides, insignificantly convex, with uni-
formly rounded humeri (humeral tubercles do not project), individually
narrowly rounded apically, posterior to humeri sometimes slightly com-
pressed, with very dense punctation forming transverse wrinkles, with
minute light-colored or light brown hairs. Prosternal process pointed
apically, reaches posterior margin of forecoxae. Femora with thin clava.
First segment of hind tarsi slightly longer than two successive together.
Body ventrally light brown, with semiadherent hairs. Entire body black
or oily black (f. typica), often pronotum red with black border on ante-
rior and posterior margins, prosternum partly red (m. daurica Motsch.)

187

141

or pronotum black only on disk and red along margins (ab. divisa
Baeckm.). Body length 9.0 to 13.0 mm.

Egg: White, elongate, rounded at poles. Chorion smooth, without
perceptible sculpture. Length about 1.2 mm, width 0.4 mm.

Larva (Figure 81): Head with parallel sides. Epistoma insignificantly
convex, on anterior margin with broad, coarsely longitudinal, dark brown
striate border (streaks diverge, slope laterally somewhat), in posterior
half with just perceptible median longitudinal suture, fuses laterally with
parietals, frontal sutures not visible. Hypostoma broadly divided by
gula into two triangular sclerites, on anterior margin with narrow rusty
border, near inner angles with slight notch. Gula comparatively broad,
slightly convex, narrows insignificantly toward apex. Parietals on anterior
margin with rusty-red border encircling articulate antennal sockets and
ocelli, with short stray hairs behind border. Ocelli convex, hyaline,
lateral to base of antennae but somewhat ventral on anterior flat section.

Figure 81. Larva of Pronocera brevicollis (Gebl.).
a— head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla.

188

Antennae short, project slightly beyond anterior margin of cephalic
capsule, brownish. Clypeus short, trapezoid, barely perceptible. Labrum
convex, small, brownish, on anterior margin broadly rounded, with
short setae. Mandibles short and black, reddish-rust basally, on outer
side with transverse groove, convex, broadly rounded apically. Inner
masticatory lobes of maxillae thick, obtuse apically and with short rusty
hairs here. Maxillary palps not long, project somewhat beyond apex of
inner lobes.

Pronotum 2.0 times wider than long, slopes forward markedly, later-
ally and on disk with rusty hairs forming two transverse bands, between
which two glabrous, yellowish-rust, transversely extended spots occur,
laterally with faint longitudinal spot. Pronotal scutum white and modera-
tely convex, with longitudinal striation, basally with stray, barely visible
setae forming transverse row, bound laterally by deep longitudinal groov-
es. Prothoracic presternum laterally with very dense long hairs and on
140 disk sparse short rusty hairs; eusternum with minute furrows, divided
anteriorly by gap into two round glabrous plates. Thoracic legs short,
with spinelike claw, brownish-rust. Dorsal locomotory ampullae slightly
convex, with very fine sculpture, matte, separated by faintly perceptible
transverse groove on anterior margin and longitudinal grooves laterally
and medially receding from it. Ventral locomotory ampullae with fairly
continuous transverse groove or latter interrupted medially, sometimes
with lateral longitudinal grooves. Body length of mature larvae 15 to
28 mm, width of head up to 2.0 mm.

Pupa (Figure 82): Body elongate. Head narrows markedly anterior
to antennae, convex between antennae, broadly impressed posterior to
them between upper ocular lobes, uniformly rounded on occiput. Anten-
nae in second half arcuate, pressed to sides.

Pronotum almost square, somewhat rounded laterally, uniformly con-
vex on disk and with transverse striation there, near base with sharp
subulate spinules forming continuous or medially interrupted transverse
band. Mesonotum transverse, with faint angularly produced scutellum
posteriorly, in posterior half with very minute, slightly sclerotized spi-
nules forming tuft. Metanotum convex, broadly rounded posteriorly,
lustrous, posterior to middle with minute spinules forming common tuft.
Femora somewhat clavate.

Abdomen elongate, narrows gradually from anterior to posterior end.
Abdominal tergites uniformly convex, lustrous, laterally with flat longi-
tudinal impression, medially or immediately posterior to middle with
minute acute spinules forming transverse indistinct row or transverse
narrow band. Posterior margin of abdominal tergite VII broadly rounded,
posterior to middle with four, around posterior margin with two seta-
ceous spinules forming two transverse rows or an indistinct diffuse tuft.

189

Abdominal tergite VIII lustrous, medially with pair of transverse broadly
separated spinules, or without them. Tip of abdomen rounded. Valvifers
of female minute, hemispherical, highly contiguous. Body length 10 to
14 mm, width of abdomen 3.5 mm.

Material: Collected in Ob’ region, Altai, Salair, Tuva, Ussuri-Pri-
141 mor’e region, and Sakhalin. Adult insects 49, larvae 63, pupae — one
male and four females, larval and pupal exuviae with beetles from cells
seven.

Distribution: West from Altai and Central Ob’, east to Sakhalin in-
clusive, northern Mongolia, northeast China, and Korea. Distributed in
maximum numbers in southeastern regions of western Siberia and
southern parts of eastern Siberia.

Biology: Inhabits spruce, fir, maple, and pine. Found in strip pine
forests of Kulunda. Flight of beetles prolonged, commencing mid-June
and ending in July. Stray insects even found in first half of September.

Figure 82. Pupa of Pronocera brevicollis (Gebl.).

190

Beetles lead cryptic mode of life, not seen on flowers. During reproduc-
tion found on drying trees. After mating, female oviposits in bark cre-
vices. Inhabits branches and tops of mature trees, sometimes under-
growth. Diameter of colonized shoots 1.0 to 12.0 cm. We did not find
it on trunks with thick bark. Young larvae seen from July through
September.

Larvae live under bark, make longitudinal meandering galleries deeply
impressed in alburnum (up to 2.0 mm), and plug them with fine frass.
Galleries sometimes narrow, sometimes broader, sometimes like plat-
forms. Before second hibernation larva bores wood, usually at an angle
of 45°, leaves transversely elongate-oval opening (3.0 to 5.0 mm wide)
on surface, and plugs it with frass. Longitudinal gallery in wood at depth
of 0.4 to 3.5 cm. After second hibernation larva makes cell in this
gallery and pupates there with its head toward inlet. Length of gallery
under bark 18 to 20 cm, width from 0.2 cm initially to 1.5 cm at end.

142 Total area of gallery under bark up to 16 cm^. Length of gallery in v/ood
2.1 to 5.1 cm, width 0.4 to 1.0 cm.

Pupation occurs from May through June. Beetles emerge two weeks
after pupation. In the laboratory, at 21°C a beetle emerged from a pupa
in 13 days and another beetle in 16 days. Young beetles push frass away
from inlet, nibble flight opening (4.0 to 5.0 mm in diameter) on bark
surface, and abandon cell through it. Emergence of beetles from wood
commences in June and ends in July. Beetles emerge from wood with
underdeveloped gonads. Weight of larvae before pupation 30 to 76 mg,
pupae 25 to 68 mg, and young beetles before emerging from wood 14
to 52 mg. Weight reduction during metamorphosis is illustrated by these
three specimens. Larva before pupation weighed 162 mg (100%), pupa
142 (87.6%), and young beetle formed from pupa before emerging from
wood 109.8 mg (67.8%), i.e., in this period the total weight reduction
was 32.2%. Generation completed in two years (Table 12).

Pronocera brevicollis (Gebl.) damages mainly spruce, more rarely
other species. For example, during forest inspections we found 84 in-
sects in larval, pupal, and adult stages; of these, 67 came from spruce
(Picea obovata), five from Siberian maple {Pirns sibiricus), 1 1 from pine

Table 12. Periods of developmeiit of Pronocera brevicollis (Gebl.)

Year of

development

April

May

June

July

August

September

October

1st

LPA

PAEL

AEL

2nd

3rd

LPA

PAEL

AEL

191

F. sylvestris), and one from fir (Abies sibirica). Motor chus minor (L.)
and Pogonocherus fasciculatus Deg. sometimes colonize the same shoots
together with this species.

4. Genus Semanotus Muls.

Mulsant, 1839, Col. France Long., p. 154; PlaviFshchikov, 1940, Fauna
SSSR, 22, 2, 274-275; Gressit, 1951, Longicorn Beetles of China, vol. 2,
p. 221; Linsley, 1964, Cerambycidae of North America, 22, 5, 18-19;
Mamaev and Danilevskii, \91 5, Lichinki zhukov-drovosekov,^^. 196-198.

Adult: Characterized by dense punctation on head, comparatively
short antennae, extending in female beyond 0.50 length of elytra, in male
beyond apex of elytra or even not reaching it. Eyes deeply emarginate.
Pronotum rounded laterally, with long erect hairs, on disk with glabrous
elevated shields with dense deep punctation between them. Prosternal
process long, reaches posterior margin of forecoxae. Elytra with parallel
sides, insignificantly convex, with white sinuous or straight transverse
bands. Femora clavate. First segment of hind tarsi equal to two succes-
sive together.

Larva: Very similar to larvae of the genus Rhopalopus. Differs in
ocelli not visible near antennal bases. Parietals medially with hairs form-
ing dense transverse band. Pronotum in anterior half with vague yellow-
ish-rust spots, laterally and on disk with light rusty hairs. Pronotal
scutum bound laterally by long longitudinal grooves, with minute longi-
143 tudinal striation. Thoracic legs short, with small, slightly sclerotized
claw. Locomotory ampullae developed on abdominal segments I to VII,
with very minute reticulate-squamiform of shagreen sculpture, imparting
matte tone, separated dorsally on anterior margin by transverse arcuate
or bracketlike groove, ventrally with short lateral longitudinal grooves.

Pupa: Characterized by head slightly bent under, insignificantly con-
vex frons between antennae, and broadly rounded occiput. Antennae
pressed to sides, in second half ventrally arcuate. Pronotum uniformly
convex, rounded laterally, glabrous, without spinules. Abdominal tergi-
tes in posterior half with very delicate, barely visible or fully developed
(especially on tergites V to VII) spinules.

The genus Semanotus Muls. developed in the early phase of coni-
ferous forest formations and occurs in the Holarctic. In USSR fauna
four species are known, of which two inhabit northern Asia; Southeast
Asia is characterized by five species and North America by four. All
species of this genus are ecologically associated with coniferous woody
plants.

Type species: Cerambyx undatus Linnaeus, 1758.

192

KEY TO SPECIES
Adult Insects

1 (2). Bases of elytra black. Transverse white bands on elytra sinuous.

Palearctic 1. S. undatus (L.).

2 (1). Bases of elytra red or reddish-rust. Transverse white bands on

elytra not sinuous, with straight margins. East Asia

2. S. bifasciatus Motsch.

Larvae

1 (2). Abdominal sternite VII with two sclerotized lobes bent inward.

Found on trunks and branches of spruce .... 1. S. undatus (L.).

2 (1). Abdominal sternite VII without sclerotized lobes. Found on juni-

per j 2. S. bifasciatus Motsch.

Pupae

1 (2). Body narrowly elongate. Abdominal tergites V to VII with weak

spinules 1. S. undatus (L.).

2 (1). Body broad, less elongate. Abdominal tergites V to VII with large

distinct spinules 2. S. bifasciatus Motsch.

1 . Semanotus undatus (L.)

Linnaeus, 1758, Syst. Nat., 10thed.,p. 396 (Cerambyx); Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 217-2^0; Dufify, 1953, Monograph Immat.
Stages of British and Imported Timber Beetles, p. 214; Demelt, 1966,
Tierwelt Deutschlands, vol. 2. pp. 69-70; Cherepanov and Cherepanova,
1973, Nov. i maloizv. vidy fanny Sibiri, 6th ed., pp. 59-61.

Adult (Figure 83): Readily recognized by long hairs on pronotum
and yellow transverse sinuous bands on elytra. Head with very dense
punctation, brownish hairs; transversely convex between antennae, with
smooth median longitudinal suture, flat between upper ocular lobes.
Eyes finely faceted, broadly and deeply (almost up to posterior margin)
144 emarginate. Antennae slender, extend beyond apex (male) or barely be-
yond 0.50 (female) of elytra. First segment of antennae notably longer
than 5th, with very dense and much longer hairs; successive segments
with dense, very short, compactly adherent hairs.

Pronotum transverse (female) or not wider than long (male), broad-
ens in anterior half, rounded laterally, narrows less anteriorly, more
posteriorly, with dense punctation, long light brown, sometimes dense
hairs, with smooth convex shields on disk, two anterolateral to middle,

193

Figure 83. Semanotus undatus (L.).

two posterolateral, and one between them medially. Sometimes lateral
shields fuse to form two smooth longitudinal bands. Scutellum small,
broadly rounded posteriorly, with adherent hairs.

Elytra with parallel sides, convex, individually rounded apically, some-
what compressed on suture posterior to scutellum and posterior to mid-
dle, with coarse punctation, semiadherent brown hairs. Femora thicken
gradually almost from base to apex, with long clava. First segment of
hind tarsi longer than two successive together. Body ventrally with adhe-
rent and erect, sometimes dense hairs. Body black, femora dark brown,
tibiae and tarsi more rusty, antennae dark brown or reddish-rust. Elytra
dark brown or black, with two transverse, broad, sinuous, light yellow
bands, one in anterior half, the other posterior to middle, more often
shortened on suture (f. typica), sometimes extended from lateral margin
to suture (ab. transversefasciatus Plav.), more rarely reduced to stray

194

spots (ab. biinterruptus PJav., ab. semireductus Plav., and ab. qmdri-
lunatus Hell.). Body length 8.0 to 15.0 mm.

Egg: White, elongate, almost identically rounded at poles. Chorion
smooth, lustrous, transparent. Length 1.6 to 1.8 mm, width 0.5 to
0.6 mm.

Larva (Figure 84): Readily recognized by sclerotized lobes on abdo-
minal sternite VII. Head narrowly rounded anteriorly. Epistoma notably
convex, with broad, longitudinally striate, dark brown border on ante-
rior margin, distinctly projecting longitudinal brown suture in posterior
half, and fuses laterally with parietals. Frontal sutures not visible. Hy-
postoma short, slightly convex, almost flat, laterally with straight parallel
sutures, on anterior margin with very narrow brownish border. Parietals
with short setaceous hairs forming medial transverse band extending
from hypostoma to frontal plate (epistoma), on anterior margin with
broad brownish-rust border encircling antennal sockets. Antennae short,

145 project insignificantly beyond anterior margin of cephalic capsule. Cly-
peus trapezoid, light brown. Labrum slightly elongate, narrowly rounded
apically, barely narrows toward base, in anterior half with short setae.
Mandibles thick, broadly rounded apically, dark red basally, with trans-
verse groove on outer side at junction of red and black parts. Inner
masticatory lobes of maxillae thick, taper apically and with short setae
there. Maxillary palps somewhat longer than inner lobes.

Pronotum 2.0 times wider than long, transverse yellow band in an-
terior half on disk separated medially by white clearance into two even
transverse tetragonal spots, laterally and in zone of yellow spots with
dense rusty hairs, before scutum with sparse, much shorter hairs. Pro-
notal scutum white, slightly convex, with faint longitudinal striation
bound laterally by long straight longitudinal grooves. Prothoracic pre-
sternum laterally with denser hairs, on disk with moderately dense rusty

146 hairs; eusternum glabrous, rugulose, anteromedially divided by hairy
clearance into two plates. Legs small but well developed, brownish-yellow
or brown and lustrous; claw short.

Abdomen laterally with short rusty hairs. Dorsal locomotory am-
pullae slightly convex, with common median longitudinal groove, trans-
verse groove in front, recurved lateral groove, with fine sculpture impart-
ing matte silvery tone. Ventral locomotory ampullae laterally with longi-
tudinal grooves, medially with transverse groove joining them. Abdo-
minal sternite VII with two lobular sclerotized processes lateral to loco-
motory ampulla; lobes bend down and inward. Body length of mature
larva 15 to 20 mm, width of head 2.1 mm.

Pupa (Figure 85): Characterized by absence of spinules on pronotum,
presence of weak spinules on abdominal tergites. Head short and gla-
brous, without setae and spinules, transversely convex between antennae,

195

145

Figure 84. Larva of Semanotus mdatus (L.).
a— head and pronotum; b — abdominal tergite IV with
dorsal locomotory ampulla; c — abdominal
sternite VIT.

flat or flatly impressed between upper lobes of eyes, rounded and lustrous
on occiput. Antennae pressed to sides, arcuate in second half.

Pronotum laterally slightly rounded, with almost parallel sides, broad-
ly rounded on posterior margin, uniformly convex on disk, transversely
very finely and densely striate, glabrous, without setae or spinules.
Mesonotum convex, with angularly or conically produced scutellum pos-
teriorly, in posterior half with short setae visible under high magni-
fication, without spinules. Metanotum broad, uniformly convex, without
median longitudinal groove, lustrous, with or without weak setae.

196

145

Abdomen elongate, broadens slightly in region of segment IV, narrows
gradually posteriorly. Abdominal tergites uniformly convex, without
median longitudinal groove, in posterior half with weak spinules form-
ing transverse row. Tergite VII more elongate, narrowly rounded poste-
riorly, in posterior third with fine spinules forming common tuft. In
female tip of abdomen more produced, valvifers large, hemispherical,
highly contiguous. Body length 10 to 15 mm, width of abdomen 3.5 mm.

Material. Collected in western and eastern Siberia, in Ussuri-Primor’e
region. Adult insects 192 (including 119 raised in the laboratory from
larvae collected in nature), larvae 190, pupae — nine males and five fe-
males, larval exuviae with beetles from cells 14.

197

Distribution: Palearctic, from Atlantic to Pacific Ocean, northern
boundary of range of coniferous forests to central Europe, northern
Mongolia, northern China, and Korea. Found in large numbers in forests
of foothills and hills of southern Siberia.

Biology: Semanotus imdatus (L.) lives in coniferous forests and is
ecologically associated with spruce. Beetles fly from May through July.
Found on withering and physiologically weakened trees; not seen on
flowers. Female lays eggs singly in bark crevices. Weight of egg just laid
0.25 mg. Female can lay about 50 eggs in her lifetime. Beetles mate the
very day of emergence from wood and female begins to oviposit two
days later. Colonizes trunks of wind-fallen and standing drying trees.

Incubation period of eggs extends up to two weeks. In the laboratory
at 14.6 to 19.6°C (average 17.5®C) larvae hatched in one experiment 11
days after oviposition, and in two other experiments in 12 to 13 days.
Larvae rupture the chorion, bore bark, make longitudinal galleries, and
plug inlet with fine frass. Galleries under bark meandering, width 0.3
147 to 0,7 mm, deeply impressed in wood, with steep, sharp, or gentle walls,
sometimes fuse to form common area. One such platformlike gallery,
made by a single larva, was 6.5 cm long, 2.5 to 3.8 cm wide, with four
small branches 0.7 to 1.1 cm wide. In one of these branches the larva
had bored wood. Total area of gallery under bark 25 cm^.

Mature larvae bore wood to a depth of up to 2.5 cm, make longi-
tudinal hollow gallery, and do not fill it with frass. Length of longitu-
dinal gallery in wood 3.0 to 10.0 cm, width 0,6 cm. Inlet into wood
plugged with frass, elongate along trunk, its width 0.4 to 0.6 cm. At end
of hollow gallery (blind alley) larva makes cell, seals it off from gallery
with plug of fine frass, then pupates with head toward plug. Length of
cell 1.4 to 2.3 cm, width 0.35 to 0.60 cm. Length of plug between cell
and hollow gallery up to 1.4 cm. Pupal cells in wood at depth of 0.2 to
4.0 cm or more.

According to our observations in Salair, larvae begin to pupate in
midsummer, mainly in first and second 10 days of July. Pupal develop-
ment at an average daily temperature of 19.7°C takes 20 to 28 days,
average 24.3 + 0.8 days. Beetles emerge from pupae at end of July and
in August, but remain in cells for hibernation. In spring, with the onset
of warm weather, beetles rupture plug sealing cell, push back frass,
penetrate bark through hollow gallery and larval inlet, nibble oval flight
opening (0.3 to 0.4 cm wide) in surface, and exit. Emergence of hiber-
nating beetles from wood commences in May and ends in June, Some
mature larvae remain in cell for second hibernation and pupate in May
or early June. Young beetles developing from these pupae emerge from
wood within a week and begin to reproduce almost immediately. Beetles
not seen on flowers and do not require supplementary feeding; they

198

emerge from wood with developed gonads. For example, ovaries of one
female dissected just after emergence from wood contained 45 eggs, of
which 38 were mature. Another female on emerging from wood laid 25
eggs in a garden, and another 14 eggs were found in her ovaries on
dissection.

Weight of mature larvae at time of entry into wood increases up to
178.0 to 295.5 mg but decreases considerably in prepupal stage. During
metamorphosis insects lose an average of up to 31.5% weight. For
example, 12 insects under observation in the laboratory weighed 842 mg
(100%) in the larval stage before pupation, pupae developed from them
749.9 (89%), and young beetles 578.3 mg (68.5%). Records of 37 in-
sects established: weight of larvae before pupation 23.0 to 107.5 mg
(68.9 + 3.4), pupae 21.5 to 103.0 mg (62.3 + 3.2), and young beetles
148 before emerging from cells 17.5 to 90.0 mg (49.1 + 2.6). One newly emer-
ged beetle (female) recovered from a cell weighed 116 mg.

Under laboratory conditions at room temperature life cycle from time
of oviposition to emergence of adult was completed in about 13 months
(refrigerated June-August). In nature life cycle completed in two years
(Table 13).

147 Table 13. Periods of development of Semanotus undatus (L.)

Year of
development

May

June

July

August

September

October

1st

AEL

2nd

LPA

3rd

AEL

Semanotus undatus (L.) inhabits drying and physiologically weakened
spruce (Piceo obovata). We did not find it on any other species. While
inspecting forests, 257 insects were collected from trees in larval, pupal,
and adult stages. From the larvae collected in nature, 119 beetles were
raised, all of them on spruce trunks. Population density was compara-
tively high. For example, from a spruce trunk 20 cm in diameter at chest
height and 10.8 m high, 178 mature larvae were recovered from wood.
Over 16 larvae were recovered from each meter length of the trunk.
Trunk riddled with an extensive network of larval galleries.

Pogonocherus fasciculatus Deg., Molorchus minor (L.), and Saperda
interrupta Gebl. are often recovered with this species from branches,
and Clytus arietoides Reitt,, Monochamus saltuarius Gebl., and others
simultaneously from the same trunks.

199

2. Semanotus bifasciatus Motsch.

Motschulsky^ 1875, Bull, Soc. Nat. Moscow, 49, 1, 149; = sinensis,
Gahan, 1888, Ann. Mag. Nat. Hist., 6, 2, 61; = ab. fasciatus, Plavilst-
shikov [Plavirshchikov], 1934, Best. Tabel. Entom. Coleopt., voL 112, p.
186; = ssp. sincauster, Gressit, 1951, Longicorn Beetles of China, vol. 2,
p. 222; PlaviFshchikov, 1940, Fauna SSSR, 22, 2, 276-277; Kojima and
Okabe, 1960, Food Plants of Japan. Cerambycidae, pp. 123-124; Ivliev
and Kononov, 1972, Trudy Biologo- Pochv. In-Ta DVNTs AN SSSR, 7,

no, 100.

Adult (Figure 86): Well distinguished from S. undatus (L.) by mark-
edly hairy pronotum and elytra with nonsinuous transverse white bands.
Head with dense deep punctation, spaces between punctures smaller
than punctures, with erect brownish hairs, with or without narrow me-
dian longitudinal suture. Eyes minute but sharply faceted, broadly emar-
ginate, with narrow septum between ocular lobes, and one or two rows
of facets. Antennae shorter than body; apices reach hind clivus of elytra
(male) or slightly beyond 0.50 elytral length (female); 1st segment with

Figure 86. Semanotus bifasciatus Motsch.

149

200

dense long semiadherent hairs, remaining segments with very short ad-
herent hairs; 5th to 10th segments angularly produced apically.

Pronotum equal in width and length, broadens roundly in anterior
half, gently toward posterior margin, narrows abruptly anteriorly, con-
vex on disk, with dense large deep punctation, laterally with dense long
erect hairs, with five smooth convex shields, two anterolateral, tv/o pos-
terolateral, and one usually longitudinally elongate posteromedial. An-
terolateral shields usually circular, posterolateral ones longitudinally
elongate; sometimes latter well developed and join anterolateral ones
or, contrarily, reduced, even absent in some specimens. Scutellum fiat,
broadly rounded anteriorly, with sparse minute hairs.

Elytra comparatively broad, with parallel sides, convex, individually
rounded apically, basally in front of anterior white band and also in
region of white bands with large flat punctation, elsewhere with fine
dense punctation, with short semiadherent hairs (on black background
black, on white background light colored), around suture with long erect

149 white hairs forming longitudinal distinct row. Femora clavate; hind
femora with long insignificantly thickened clava. First segment of hind
tarsi equal to two successive ones. Body ventrally with long semi-adhe-
rent light brown hairs, looks densely hairy. Body black, antennae dark
brown, apically dark rust. Legs black or dark brown , tarsi with rusty
tinge. Elytra in anterior third reddish-rust, anterior to middle with nar-
row white band, anterior to posterior third with broad transverse white
band slightly curved anteriorly, white apically; black or blackish-brown
between white transverse bands and also behind posterior bands. Im-
pression created of elytra with red base, white apex, two white narrow
and two black much broader transverse bands (f. typica). Other forms
(ab. fasciatus Plav., ab. latifasciotus Matsuch.) with minor differences.
Only ssp. sincauster Gress. differs from nominal form in very smooth,
not coarse punctation in region of medial black band on elytra. Latter in
anterior half or only posteriorly appears rugose. Length 8.0 to 14.0 mm.

Egg: White, elongate, broadly rounded at one pole, narrows mark-
edly toward the other and pointed. Chorion smooth, lustrous, trans-
parent. Length 2.1 mm, width 0.4 mm.

Larva (Figure 87): Similar to larva of S. undatus (L.). Differs in
markedly oblique, almost transverse striation on anterior margin of
epistoma and absence of sclerotized lobes on abdominal sternite VII.
Half of head retracted into prothorax; general outline of head broadly
rounded anteriorly. Epistoma insignificantly convex, whitish, with broad

150 dark brown border on anterior margin and almost transverse striation,
in posterior half with distinct median suture. Frontal sutures not visible.
Hypostoma laterally with straight sutures, in anterior half with trans-
verse striation, on anterior margin with narrow clavate border. Gula

201

basally insignificantly broadened, narrows moderately, anteriorly, notably
elongate. Parietals more yellowish, on anterior margin with broad brown-
ish-rust border encircling articulate antennal sockets, around base of
antennae without perceptible ocelli, medially with dense rusty hairs
forming transverse band dorsally up to epistoma. Antennae compara-
tively long, project beyond anterior margin of cephalic capsule. Clypeus
short, very broad, projects as narrow band from behind anterior margin
of epistoma. Labrum transversely oval, with broadly rounded anterior
margin, in anterior half with short setae, whitish, basally brownish.
Mandibles apically broadly rounded, black, on outer side with lac sheen,
reddish basally. Inner masticatory lobes of maxillae thicken slightly to-
ward apex, white and broadly rounded apically, with short light-colored
setae there. Maxillary palps basally thick, narrow conically apically,
slightly longer than inner masticatory lobes.

Pronotum almost 2.5 times wider than long, in anterior half with
two transverse, tetragonal, rusty-yellow spots, laterally and on disk with
light rusty hairs forming transverse band in zone of rusty spots. Anterior
to scutum hairs comparatively short and sparse. Pronotal scutum white.

202

moderately convex, bound laterally by longitudinal grooves that are in-
significantly outcurved, slightly produced forward on anterior margin me-
dially and laterally, with minute, barely perceptible longitudinal striation.
Prothoracic presternum laterally v/ith long hairs and on disk with short-
ened rusty hairs; eusternum in form of two glabrous, circular, rugulose
plates separated in front by comparatively broad hairy clearance. Thora-
cic legs minute, with acicular, slightly sclerotized claw.

Abdomen laterally with short, not very dense light-colored hairs.
Dorsal locomotory ampullae insignificantly convex, with faint common
median longitudinal groove, on anterior margin with transverse groove
that is laterally arcuate or posteriorly bracketlike, with minute shagreen
or reticulate-squamiform sculpture, matte silver. Ventral locomotory
ampullae laterally with short longitudinal grooves, on sternite VII with
deep transverse groove, laterally without sclerotized lobes. Body length
of mature larvae 16 to 20 mm, width of head 2.8 mm.

Pupa: Differs from pupa of S. undatus (L.) in very broad body. Head
short, slightly bent under, glabrous, without spinules, barely convex bet-
ween antennae, flat between upper ocular lobes, uniformly rounded on
occiput. Antennae pressed to sides, ventrally arcuate in second half.

Pronotum broadens in anterior half, narrows gradually posteriorly, in-
significantly convex on disk, lustrous, without spinules. Mesonotum almost
equal in length and width, with slightly produced scutellum posteriorly.
Metanotum uniformly convex, without median longitudinal groove.

Abdomen flat, broadens in middle, gradually narrows posteriorly.
Abdominal tergites on posterior margin with minute spinules forming
transverse row or transverse band. Tergite VII with large spinules form-
ing extensive tuft in posterior half. Body length up to 16 mm, width of
abdomen up to 4.0 mm.

151 Material: Collected in Ussuri-Primor’e region (Suvorovka River near
Zmeinaya hill). Adult insects 1 5 (raised from larvae collected in nature),
larvae 10 (of which five raised from eggs laid by beetles in the labora-
tory), pupa one (injured), larval and pupal exuviae from cells with beet-
les six.

Distribution: Ussuri-Primor’e region, .northeast China, Korea, and
Japan.

Biology: Inhabits coniferous forests and ecologically associated with
juniper. Beetles fly from June through August. Found on trunks of wind-
fallen and standing withering trees, mate there, and oviposit in bark
crevices. One female can lay 48 eggs in her lifetime. In the laboratory
beetles begin to reproduce soon after emerging from wood. Larvae hatch
from eggs 11 to 16 days (average 14 days) after oviposition. Nineteen
eggs were kept under observation. Atmospheric temperature during this
period 11.0 to 20.6°C (average 17.4°C).

Larvae live under bark, make meandering galleries that broaden gra-
dually or sharply, terminate in platform deeply imprinted in alburnum,
and plug them with fine frass. Width of gallery imprinted in sapwood
initially 0.2 cm, then 0.9 cm, and at end of platform up to 1.6 cm. Length
of gallery up to 11 cm or more. Mature larvae bore wood, leaving an
oval inlet on surface that is transversely elongate or oblique to axis of
trunk. Width of inlet 0.6 cm. Larva makes cell along trunk in wood to
a depth of 2.0 cm or more and pupates in it with head toward inlet
plugged with frass. Sometimes cell disposed in wood not along trunk
but radially. Length of cell up to 2.1 cm and width 0.9 cm. Length of
plug of fine frass sealing cell from inlet opening 0.9 cm. Pupae seen at
end of summer; beetles emerge by autumn and hibernate in pupal cells.

Young beetles emerge from wood in June, break plug around inlet,
push back frass, reach bark, nibble oval flight opening, and emerge from
wood. Dimensions of opening 3.0 mm X 5.5 mm. Weight of beetles
just emerging from wood 53.4 to 81 .7 mg, average 63.6 mg (five insects
weighed). Generation completed in two years.

Semanotus bifasciatus Motsch. inhabits trunks and thick branches of
juniper {Juniperus rigida). For example, on a cutting 64 cm long and
5.0 to 7.5 cm in diameter, 11 adults and one larva were found in wood.
On the same cutting, under bark, 1 5 insects (larvae, pupae, and adults)
of Atimia nadezhdae Tsher. were also recovered simultaneously.

In Japan, Korea, and China this species colonizes Juniperus, Chamae-
cyparius, Thuja, and Thujopsis.

5. Genus Oupyrrhidium Pic

Pic, 1900, Catal. Longic., p. 50; Plavil’shchikov, 1940, Fauna SSSR,
22,2, 285-286 {Upyrrhidium); Gressit, 1951, Longicorn Beetles of China,
vol. 2, p. 226.

Adult'. Differs from those of other genera of this tribe in long 3rd
antennal segment, fairly long prosternal process, and sharply dilated,
comparatively short clava of hind femora. Head with dense punctation
and adherent hairs. Eyes deeply emarginate. Antennae long, slender; 3rd
segment longer than 5th. Pronotum laterally rounded, with very dense
minute punctation, short adherent hairs. Elytra elongate, with parallel
sides, individually broadly rounded at apex, matte, with longitudinal
152 ribs. Prosternal process pointed apically, with apex almost reaching
posterior margin of forecoxae. Femora sharply clavate, markedly convex
apically. First segment of hind tarsi considerably longer than two suc-
cessive together.

Larva’. Legs poorly developed, very short. Epistoma longitudinally
weakly striate only medially on anterior margin, laterally smooth.

204

Parietals glabrous in middle, with stray hairs around brownish- rust border
on anterior margin. Locomotory ampullae developed on abdominal seg-
ments I to VII, with minute furrows imparting characteristic sculpture,
with silvery sheen.

Pupa: In arrangement of spinules very similar to pupae of Pronocera
Motsch. Well distinguished, however, in sharply clavate femora. Head
short. Second half of antennae bent ventrad, looplike. Pronotum longi-
tudinally oval, usually with biapical spinules mediobasally. Meso- and
metanota on posterior margin or behind middle with minute spinules
forming one small tuft each. Abdominal tergites in posterior half with
minute mono- or biapical spinules forming transverse row, transverse
band, or one small paramedial tuft each. Abdominal tergite VII transverse
(male) or elongate (female), with single tuft of minute spinules in posterior
half.

Oupyrrhidium Pic is a monotypic genus characteristic for eastern
regions of Asia, and probably evolved in broad-leaved forests at com-
mencement of Quaternary period.

Type species: Callidiiim cinnabarinum Blessig, 1872.

1. Oupyrrhidium cinnabarinum (Bless.)

Blessig, \%12, Horae. Soc. Entom. Ross., vol. 9, p. 179 (Callidium);
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 286-287 (Upyrrhidium); Gressit,
1951, Longicorn Beetles of China, vol. 2, p. 226.

Adult (Figure 88): Characterized by flat body and vermilion elytra.
Body elongate, flat, head transversely convex between antennae, with
narrow median longitudinal suture extending onto vertex, with minute
extremely dense punctation, dense adherent reddish hairs on vertex,
elsewhere sparse gray hairs. Eyes emarginate almost up to posterior
margin. Antennae longer than body; 9th (male) or 10th (female) segment
extends beyond elytral apex; 1st segment thick (female) or slightly elongate
(male), not longer than 3rd segment, with large punctation.

Pronotum somewhat longer than wide, unevenly rounded laterally,
narrows more posteriorly, less anteriorly, with very dense minute puncta-
tion, dense adherent red hairs directed forward throughout disk from
posterior to anterior margin, dark brown hairs laterally in anterior half
forming one large black spot on each side. Scutellum longitudinal, flat,
broadly rounded at apex, with minute punctation, and short dense
adherent reddish hairs.

Elytra elongate, flat, with parallel sides, individually broadly rounded
at apex, slightly impressed around humeri, with longitudinal ribs (two
or three ribs each), very dense minute punctation imparting matte tone,
and minute semiadherent, barely visible, reddish hairs. Legs long; femora
clavate, highly convex in second half; hind tibiae somewhat curved.

205

154 Figure 88. Oupyrrhidium cinnabarimm {^\qss.).

First segment of hind tarsi 1.5 times longer than two successive together.
Abdominal sternite V transverse, posteriorly truncate (male) or elongate,
153 narrowly rounded posteriorly (female). Body, antennae, and legs black;
tarsi rusty; pronotum red on disk, black laterally; scutellum and elytra
vermilion (f. typica); pronotum sometimes entirely black (ab. nigricollis
Plav.). Body length 7.0 to 17.0 mm.

Egg: White, elongate, somewhat thickened in one half, rounded at
poles. Chorion smooth, without perceptible sculpture. Length 1.4 to 1.8
mm, width 0.5 mm.

Larva (Figure 89): Head narrows somewhat anteriorly, with almost
parallel sides. Epistoma slightly convex, divided medially by distinct
longitudinal suture, laterally without distinct frontal sutures, on anterior
margin with narrow dark brown border, in middle with longitudinal

206

155

Figure 89. Larva of Oupyrrhidium cinnabarimm (Bless.),
a — head and pronotum; b~prosternum; c— abdominal
tergite IV.

striation, smooth around anterior angles^ with setaceous hairs behind
border forming transverse row. Hypostoma short, on anterior margin with
smooth narrow brownish border, in anterior half with faint transverse
furrows or without them. Parietals on anterior margin with narrow rusty-
brown border, behind which stray hairs in anterior half do not form
distinct tuft. Antennae slender, with apices barely projecting beyond
anterior margin of cephalic capsule. One unevenly pigmented ocellus

207

near each base of antennae ventrally. Clypeus short, projects from behind
epistoma as narrow band. Labrum small, whitish, rounded or narrowly
rounded anteriorly, with short setae. Mandibles on outer side with
transverse groove, black, basally reddish-rust. Inner masticatory lobes of
maxillae thick, lustrous, narrowly rounded apically, distinctly shorter
than maxillary palps.

Pronotum in anterior half with pair of large yellow or yellowish- rust
spots, short fine rusty hairs laterally and on disk forming two transverse
bands, one denser along anterior margin of yellow spots, the other less
dense anterior to scutum. Pronotal scutum with two emarginations on
anterior margin, medial one prominent, with anterior angles insignifi-
cantly produced forward, white, with minute longitudinal striation, fine
sculpture at base imparting matte silver tone, and laterally with deep
longitudinal grooves. Prothoracic presternum with short erect hairs on
disk, laterally with long rusty hairs bent down and sideways; euster-
num glabrous, rugulose, anteriorly with faint very short hairy septum.
Thoracic legs very short, with minute thin claw.

Abdomen laterally with sparse short light-colored hairs. Dorsal
locomotory ampullae moderately convex, with minute furrows imparting
characteristic sculpture, with silvery hue, common median longitudinal
groove, lateral longitudinal grooves, and faint short transverse grooves
on anterior margin receding inward from lateral longitudinal grooves.
Ventral locomotory ampullae similar in structure, laterally with short
grooves flexed angularly in form of small dent near posterior end.
Ampullae on abdominal sternite VII divided medially by continuous
transverse groove. Body length of mature larva up to 20 to 23 mm, width
of head 2.8 mm.

Pupa (Figure 90): In arrangement of spinules on pronotum very
similar to pupa of Pronocera brevicollis (Gebl.). Differs in markedly
clavate (apically dilated) hind femora. Head rounded, between antennae
transversely convex, flat on vertex, uniformly rounded on occiput, lus-
trous. Antennae long, pressed to sides, bent forward, looplike, in second
half ventrad.

Pronotum not wider or even narrower than long, broadly rounded
154 laterally, uniformly convex on disk, transversely striate medially in
posterior half, basally with biapical spinules forming small tuft. Meso-
notum longitudinally elongate, convex, lustrous, with barely extended
scutellum posteriorly bearing small tuft of small spinules. Metanotum
transverse, rounded broadly on posterior margin, in posterior half
medially with minute spinules forming tuft. In some specimens stray
spinules replace tuft. Hind femora long, sharply clavate, almost reach
posterior margin of tergite VI.

208

155

Figure 90. Pupa of Oupyrrhidium cinnabarinum (Bless.).

Abdomen widens in region of tergites III and IV, narrows slightly
anteriorly and markedly posteriorly, more produced at tip in female.
Abdominal tergites uniformly convex, laterally with longitudinal, streak-
like, slightly convex furrows, in posterior half with mono- or biapical
spinules forming one paramedial tuft each (two to five spinules per tuft).
On abdominal tergites I and II these spinules form transverse row or
transverse band. Tergite VII in male transverse, broadly rounded posterior-
ly minute spinules around posterior margin form sparse tuft, in female
elongate, longitudinal, narrowly rounded posteriorly, posterior to mid-
dle with minute dispersed spinules. Valvifers of female oval, transversely
slightly elongate, contiguous. Body length 9.0 to 18.0 mm, width of
abdomen 3.5 to 4.0 mm.

209

Material: Collected in Ussuri- Prim or’ e region (Komarovka, Man-
zovka, and Volkhovka Rivers). Adult insects 32, larvae 11, pupae — three
males and one female, larval and pupal exuviae with beetles from cells
three.

Distribution: Southeastern northern Asia, from Khabarov to coast of
Sea of Japan, from Sikhote-Alin’ to Lake Khanka and Khasan, north-
east China, and Korea. Often found in southern areas of Ussuri-Primor’e
region.

Biology: Inhabits broad-leaved, mainly oak-elm forests. Prefers
clearances, fringes of large forests, and sparse damaged cuttings. Beetles
fly from mid-June to end of July. Found only on recently dried and

155 drying shoots of elm and oak, often on seedlings, mate there and oviposit.
Female lays eggs under bark strips singly or spaced in batches. Colonizes
thin shoots 2.0 to 3.0 cm in diameter. One female can lay up to 48 eggs.
Larvae hatch from eggs in two weeks. Larvae hatched from eggs laid
in the laboratory in 14 to 16 days. Larvae began hatching on the 19th
of July from eggs laid in nature on July 6th.

Newly hatched larva ruptures egg chorion, emerges, and bores bark.
Larvae make meandering galleries under bark, deeply impressed in
alburnum, and plug them with fine frass. Larval galleries sometimes
widen and narrow alternately, and sometimes resemble platforms with
rather sharp edges. Length of gallery under bark up to 13 cm, width
0.4 to 1.2 cm. Mature larvae bore wood to a depth of 0.6 to 1.0 cm and
nibble pupal cell there along shoot. Width of inlet 5.0 to 6.0 mm. Length
of pupal cell 1.8 to 3.4 mm and width 0.5 to 0.6 mm.

156 Larvae pupate after second hibernation. Pupation commences in
May, ceases in June. Pupae lie in cells with head toward inlet. Young
beetles emerge from pupae in two to three weeks. Weight of larvae be-
fore pupation 29.6 to 139.0 mg, pupae 26.5 to 11 8.8 mg, and young beet-
les before emerging from cell 21.5 to 98.0 mg. Young beetles nibble oval
flight opening (3.0 mm x 2.0 mm to 5.0 mm x 3.5 mm) in bark sur-
face and emerge. Emergence of young adults from wood commences mid-
June and ceases early July. Adults begin to reproduce soon after emer-
gence from cells. Usually not seen on flowers. Generation completed in
two years (Table 14).

Table 14. Periods of de?elopmeiit of Oupyrrhidium cinnabarinum (Bless.)

Year of

development

April

May

June

July

August

September

1st

LPAE

PAEL

2nd

3rd

LPAE

PAEL

210

Oupyrrhidium cinnabarinum (Bless.) damages thin shoots mainly of
elm, more rarely oak, of viable as well as withering trees. Thus of the
49 insects (larvae, pupae, and adults) found by us on shoots, 46 were
from elm and three from oak. Population density is illustrated in these
examples. On an elm shoot 43 cm long and 2.3 cm in diameter three
adults and one pupa were found. On another shoot 40 cm long and 2.0
cm in diameter four larvae were found. Similar population density found
on other shoots. Exocentrus marginatus Tsher., more rarely Pterolophisa
ussuriensis Plav., colonize shoots of elm concomitant with this species.

6. Genus Callidium (L.)

Fabricius, 1775, Syst. Entom., p. 187; Mulsant, 1839, Hist. Nat. Col.
France Longic., p. 42; Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 287-
291; Gressit, 1951, Longicorn Beetles of China, vol. 2, pp. 223-224;
Linsely, 1964, Cerambycidae of North America, 22, 528-30; Kojima and
Hayashi, 1969, Insect Life in Japan, vol. 1, p. 72; Cherepanov and Chere-
panova, 1973, Nov. i maloizv. vidyfauny Sibiri, 6th ed., pp. 53-54.

Adult: Characterized by moderately elongate, comparatively flat body.
Head short, frons between antennae, last segment of maxillary palps
large, markedly broadened apically. Pronotum transverse, rounded later-
ally, narrows insigniflcantly anteriorly, more posteriorly, on disk usually
with large and laterally minute punctation. Elytra with parallel sides, with
dense minute punctation, without furrows [C. violaceum (L.) and others]
or broadened from base posteriorly, with large transverse furrows, flatter
or flattened posteriorly (C. aeneum Deg.). Prosternal process short, does
not reach middle of forecoxae, or long, extending far beyond middle of
forecoxae. Femora clavate. Hind femora of female do not reach apex of
elytra, of male extend almost beyond it.

Egg: White, moderately (C. aeneum Deg.) or markedly [C. chlori-
zans (Sols.)] elongate.

Larva: Body moderately elongate. Head small, more or less retracted
into prothorax. Epistoma on anterior margin with brownish-rust border
on which striation transverse [C. violaceum (L.)] or longitudinal and
157 oblique (C. aeneum Deg., C. coriaceum Payk.), or absent [C. chlorizans
(Sols.)]. Hypostoma with spinules on anterior margin (C. aeneum Deg.)
or without them [C. violaceum (L.) and others]. Pronotal disk with uni-
formly rusty hairs forming two transverse flelds. In some species basally
with sclerotized ringlet (C. coriaceum Payk., C. aeneum Deg.), in others
without ringlet [C. violaceum (L.), C. chlorizans (Sols.)]. Pronotal scutum
with fine dense longitudinal striation, basally with minute shagreen
sculpture forming transverse matte silver border. Thoracic legs short,
with sharp acicular claw.

211

Pupa: Characterized by slightly convex frons between antennae. Head
lustrous, without setae or spinules, broadly rounded on occiput. Anten-
nae pressed to sides, with apices ventrad (female) or arcuate, directed
forward (male). Pronotum transverse, rounded laterally, slightly covex or
somewhat flattened on disk. Upper side of body with spinules [C. viola-
ceum (L.)] or without them (C. aeneum Deg.). Tip of abdomen in male
narrowly rounded, in female more extended, without spinules.

This genus is distributed in the Holarctic. No less than 12 species are
found in the Palearctic and 17 species in North America; of these, one
[C. violaceum (L.)] is widespread in Eurasia as well as in North America.
Four species are known in northern Asia. All of them are ecologically
associated with conifers and mainly colonize drying and withered trees.
Type species: Cerambyx violaceum Linnaeus, 1758.

KEY TO SPECIES

Adult Insects

1 (4). Prosternal process short, does not reach middle of forecoxae (sub -

genus Callidium s. str.).

2 (3). Elytra with parallel sides, uniform on disk, not rugose, without

longitudinal carinae, with violet tinge. Eurasia and North America.
1. C. violaceum (L.).

3 (2). Elytra broaden notably from base posteriorly, in posterior half as

though flattened, on disk posterior to base coarsely, transversely
rugose, with distinct longitudinal carinae, with greenish-bronze
hue. Eurasia 2. C. aeneum Deg.

4 (1). Prosternal process long, elongate, extends far beyond middle of

forecoxae (subgenus Palaeocallidium Plav.).

5 (6). Elytra rusty-brown, with bronze or bluish-bronze iridescence.

Eurasia 3. C. coriaceum Payk.

6 (5). Elytra dark green, with bronze or purplish-bronze metallic irides-

cence. Northern Asia from Altai to Pacific Ocean coast

4. C. chlorizans (Sols.).

Larvae

1 (4). Dorsal locomotory ampullae with single transverse groove.

2 (3). Hypostoma on anterior margin, lateral to gula, without spinules.

1. C. violaceum (L.).

3 (2). Hypostoma on anterior margin, lateral to gula, with pair of large

spinules 2. C. aeneum Deg.

212

4 (1). Dorsal locomotory ampullae with two transverse grooves, of which

anterior one continuous, posterior one widely interrupted medi-
ally.

5 (6). Epistoma on anterior margin with dense thin striation. Hairs on

pronotal disk basally with sclerotized ringlets

3. C. coriaceum Payk.

158 6 (5). Epistoma on anterior margin without striation. Hairs on pronotal
disk without basal sclerotized ringlets. . . . 4. C. chlorizans(Sols.).

Pupae

1 (2). Pro-, meso-, and metanota, and abdominal tergites with spinu-

les 1. C. violaceum (L.).

2 (1). Pro-, meso-, and metanota invariably without spinules, abdomi-

nal tergites with faint spinules or without them.

3 (4). Head broadly concave on vertex. Abdominal tergites without

spinules 2. C. aeneum Deg.

4 (3). Head not concave on vertex, forming there common plane with

frons between antennae. Abdominal tergites with distinct or faint
spinules 4. C. chlorizans (Sols).

1. Callidium violaceum (L).

Linnaeus, 1758, Syst. Nat., 10th ed., p. 395 {Cerambyx); Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 297-298; Duffy, 1953, Monograph Beetles,
pp. 214-216; Linsley, 1964, Cerambycidae of North America, 22, 5, 30-
32, Starzyk,1968, Przeglad Zoologizni, 12, 4, 401-404; Kojima and
Hayashi, 1969, Insect Life in Japan, vol. 1, p. 72; Cherepanov and
Cherepanova, 1973, Nov. i maloizv. vidy fauny Sibiri, 6th ed. pp. 56-59.

Adult (Figure 91): Characterized by double punctation on pronotum
in male and more uniform large punctation in female, and violet elytra.
Head with large punctation, markedly retracted into prothorax, trans-
versely convex between antennae, with narrow median longitudinal
suture. Eyes finely faceted, straightly truncate posteriorly, deeply emar-
ginate anteriorly on inner side. Antennae barely extend beyond 0.50
(female) or 0.66 length of elytra (male), lustrous in anterior half and
matte in posterior half.

Pronotum laterally rounded anterior to middle, sometimes broadens
angularly, narrows markedly posteriorly, on disk in posterior half with
large punctation, laterally and in anterior half with dense minute puncta-
tion (male) or dense large punctation throughout entire surface (female),
with sparse erect rusty hairs. Scutellum short, rounded posteriorly, medial-
ly impressed, troughlike. Elytra with parallel sides, insignificantly convex,
uniform, without longitudinal carinae, apically with narrowly rounded

213

Figure 91 . Callidium violaceum (L.).

159 inner angle and hollow tapered outer angle, with dense coarse punctation,
and semiadherent short thin hairs. Femora with long flattened clava. Hind
ferriora extend almost beyond apex of elytra (male) or do not reach it
(female). Body ventrally with long light-colored semiadherent hairs. Head,
pronotum, prosternum, and elytra violet or bluish-violet with metallic
iridescence, meso- and metanota and abdomen brownish-rust or chestnut
with rusty tinge. Antennae and legs dark brown (f. typica), rarely elytra
green (ab. virescens Stierl.) or violet with purple iridescence, and legs
rusty-red (ab. salessei Pic). Body length 9.0 to 15.0 mm.

Egg: White, elongate, rounded at poles. Chorion smooth, without
perceptible cellular sculpture. Length 1.8 mm, width 0.6 mm.

Larva (Figure 92): Head short, retracted into prothorax up to apex of
median suture. Epistoma slightly convex, almost flat, uniform on anterior

214

Figure 92. Larva of Callidium violaceum (L.).
a — head and pronotum; b — abdominal tergite with dorsal loco-
motory ampulla; c—ventral view of head (maxillae, labium,
hypostoma, and gula)

' margin, with rusty-red border, laterally on clypeus with dark brown sp©t,
with transverse thin curved streaks visible under high magnification,
fuses laterally with parietals, frontal sutures not visible, median longi-
160 tudinal suture visible only at apex. Hypostoma lustrous, transversely
finely striate together with gula in anterior half, without spinules on
anterior margin. Parietals in anterior half with short thin hairs. One
convex gray ocellus near base of each antenna. Antennae project forward
insignificantly; 1st segment thick, not shorter than remainder together.

215

Clypeus white, trapezoid, lustrous. Labrum small, white, narrows toward
apex, narrowly rounded there, broadens angularly in posterior half, con-
vex on disk, with short light-colored setae along margins. Mandibles
massive, broadly rounded apically, dark red basally, convex and smooth
on outer side. Inner masticatory lobes of maxillae apically with short
light-colored setae, somewhat shorter than maxillary palps.

Pronotum transversely oval, slightly narrower anteriorly, with one
longitudinally elongate spot laterally, two transverse tetragonal yellow
spots on disk in anterior half, before scutum, and also in anterior half in
region of yellow spots and laterally with rusty hairs forming two transverse
fields, between which longitudinal hairy clearance retained medially; clear-
ance separates two glabrous plates posterior to yellow spots. Pronotal
scutum insignificantly convex, white, with barely perceptible longitudinal
striation, bound laterally by short longitudinal grooves, glabrous, only
basally with eight to ten short setae forming transverse row. Prothoracic
presternum with short hairs on disk, laterally with dense long rusty hairs;
eusternum in form of pair of round glabrous lustrous lateral plates,
divided anteriorly by hairy field, sometimes yellowish. Base (supporting
part or sternellum) of pro-, meso-, and metasterna glabrous in middle,
with rusty hairs laterally. Thoracic legs small, rusty-brown; claw sharp.

Abdomen laterally with thin light-colored hairs. Abdominal tergites
in anterior half with lateral constriction. Dorsal locomotory ampullae
moderately elongate, divided by common median longitudinal groove,
anteriorly with transverse groove curved forward, joining laterally with
lateral longitudinal grooves. Ventral locomotory ampullae divided by
common longitudinal groove, laterally near anterior angles with short
bracketlike, sometimes triradial grooves. Body length of mature larvae 18
to 21 mm, width of head 2.1 mm.

Pupa (Figure 93): Body comparatively elongate (male) or widens in
region of abdomen (female). Head between antennae transversely convex,
widely impressed between upper ocular lobes, glabrous, without setae.
Antennae pressed to sides, in second half bent ventrad, arcuate.

Pronotum laterally rounded, flat on disk or slightly convex, narrows
uniformly anteriorly and posteriorly, near base with sparse, dispersed,
broad short spinules which, in female, sometimes form compact row or
sometimes medially interrupted transversely. Mesonotum convex, with
insignificantly produced elevated scutellum with tuft of short widened
spinules in posterior part. Metanotum transverse, slightly convex, on
disk posterior to middle with short spinules forming extensive tuft sepa-
rated partly or fully in front by longitudinal groove.

Abdomen elongate, almost with parallel sides (male) or broadens
(female), narrows markedly anteriorly and especially posteriorly. Abdo-
minal tergites more (male) or less (female) convex, with common median

216

159 Figure 93. Pupa of Callidium violaceum (L.), female.

longitudinal groove, in posterior half laterally with minute spinules form-
161 ing transverse band (eight to ten paramedial spinules in male and 12 to
20 in female). Tergite VII narrows posteriorly, in second half with
minute spinules forming dispersed tuft. Tergite VIII in female elongate,
with almost parallel sides, triangular in male, not elongate, narrowly
rounded posteriorly. Valvifers of female hemispherical, contiguous, com-
paratively large. Body length 18 mm, width of abdomen 5.0 mm.

Material: Collected in western and eastern Siberia. Adult insects 187,
larvae 125, pupae six.

Distribution: Zone of growth of coniferous forests. Europe from the
Atlantic to the Urals, from Sweden and Finland to the Mediterranean;
Asia from the Urals to the Pacific Ocean, from forest tundra to northern
Mongolia, northern China, Korea, and Japan; North America.

217

Biology: Inhabits deciduous, pine, cedar, and mixed forests with
viable conifers. Large numbers occur in forests in plains and foothills,
found in hills to a height of 1,000 m above sea level. Beetles fly from
May to July. Maximum numbers seen in second half of June. During
systematic inspections of forests, mainly in the southern regions of
Siberia, 142 beetles were collected in one season; of these, 2.1% were
found in May, 89.4% in June, and 8.5% in July. Beetles usually not
seen on flowers during reproduction, but on stumps, trunks of irretriev-
ably withered, just dried, and felled coniferous trees. Inhabit trunks of
varying thickness and branches up to 3.0 cm or more in diameter.
Female lays eggs in bark crevices singly or in batches of five each.
According to observations made in Tuva, incubation period of eggs in
nature at 18.9 + 0.7°C varies from 17 to 19 days. Hatching of larvae
commences end of June and ceases in August.

Larvae live under bark, make meandering (sometimes platformlike)
longitudinal or transverse, quite often intersecting broad galleries,
impressed in alburnum, and plug them with fine frass. Mature larva bores
wood to a depth of up to 1.0 cm, makes short longitudinal gallery with
cell at end, plugs inlet with frass, turns head toward inlet, and pupates.
Length of gallery under bark up to 15 cm, width 1.0 to 2.5 cm. Width
of inlet into wood 0.4 cm. Length of longitudinal gallery in wood 3.0 to
7.0 cm, length of pupal cell 1.6 to 2.0 cm and width 0.5 cm.

Pupation of larvae commences early May, ceases in first half of June.
Young beetles nibble oval openings (3.0 mm X 5.0 mm to 4.0 mm X 8.0
mm) in bark surface and abandon cell through them. Emergence of
beetles from wood commences in last 10 days of May and ceases in early
July. Beetles begin to reproduce soon after emerging from wood. Records
of nine insects revealed: weight of larvae before pupation 101.0 to 214.1
mg, pupae 85.0 to 194.1 mg, and young beetles before emerging from
cells 68 to 1 54 mg. Weight of some larvae under bark before penetrating
wood 233 mg.

More often colonizes larch, rarely other conifers. For example, of
152 insects (larvae, pupae, and adults) collected by us in nature, 1 1 6
came from larch, 26 Siberian maple, nine pine, and one spruce. Not
found on fir. Technically described as a pest. Evidently does not attack
growing viable trees, not even weakened ones.

2. Callidium aeneum Deg.

Degeer, 1775, Mem. Ins., vol. 5, p. 89; Plavil’shchikov, 1940, Fauna
SSSR, 22, 2, 299-301; Gressit, 1951, Longicorn Beetles of China, vol. 2,
p. 226; Demelt, 1966, Tierwelt Deutschlands, vol. 2, p. 70; Kojima and
Hayashi, 1969, Insect Life in Japan, vol. 1, p. 72.

162 Adult (Figure 94): Differs from other species in broad transverse coarse
wrinkles on elytra. Head posterior to eyes with almost parallel sides,

218

with dense fused punctation, longitudinal suture medially between anten-
nae. Eyes very finely faceted, broadlye marginate. Antennae slender,
apices extend beyond middle or reach hind clivus of elytra.

Pronotum transverse, rounded laterally, narrows less anteriorly and
notably posteriorly, slightly convex on disk, with minute fused rugulose
punctation, laterally with two longitudinal grooved dents (male) or with
very distinct, separated punctures, with shagreen spaces between them,
laterally with slight, bandlike dents or without them (female). Scutellum
smooth, comparatively short, broadly rounded apically (male) or more
elongate, narrows notably toward apex, narrowly rounded posteriorly
(male) [s/c]. Elytra flat, broaden from base to apex, individually rounded
apically, with faint longitudinal carinae, fine punctation at base, elsewhere
with transversely folded coarse furrows. Femora broaden almost from base
to apex, quite flattened. Body dark brown, ventrally more rusty. Legs dark
rust or chestnut. Antennae rusty or dark brown. Elytra with green,
greenish-bronze, or bronze metallic iridescence. Body length 10 to 14 mm.

Figure 94. Callidium aeneum Deg.

219

Egg: White, matte, moderately elongate, broad at one pole, notably
more narrowly rounded at the other. Chorion with fine sculpture visible
under high magnification. Length 1.6 to 1.8 mm, width 0.6 mm.

Larva (Figure 95): Well distinguished from all species of this genus
by spinules on anterior margin of hypostomal sclerites. Head almost
with parallel sides, transverse. Epistoma flat, laterally without perceptible
frontal sutures, medially in posterior half with distinctly visible brown
longitudinal suture, on anterior margin with broad brownish-rust border,
obliquely (directed laterally) striate; fine streaks extend from middle of
posterior margin of border forward toward base of mandibles [these
streaks in Callidium violaceum (L.) transverse but, contrarily, on clypeus
narrowly bent backward]. Hypostoma slightly convex, lustrous, laterally
with straight sutures, on anterior margin with rusty-brown border, two
large dark brown spinules on each sclerite, of which outer one some-
what larger than inner and slightly posterior relative to it. Gula narrows
163 somewhat anteriorly in anterior half, flat, does not project beyond

Figure 95. Larva of Callidium deneum Deg.
a — hypostoma; b — ^head and pronotum; c — abdominal tergite with dorsal
locomotory ampulla.

120

hypostoma. Parietals in anterior half with hairs forming extensive tuft.
Antennae rusty, project insignificantly from antennal sockets. One uneven-
ly pigmented ocellus located laterally near base of each antenna. Clypeus
short, widely flattened at base, hyaline. Labrum white, basally with slight
brownish tone, narrowly rounded apically, with short light- colored setae.
Mandibles thick, short, smooth on outer side, medially in posterior half
with longitudinal, not very deep groove.

Pronotal disk with dense, comparatively long rusty hairs anterior to
scutum and laterally, two transverse, comparatively narrow, yellowish-
rust spots in anterior half, with one longitudinally elongate yellowish-
rust spot laterally. Hairs on anterior margin of yellow spots basally with
sclerotized ringlet and form compact transverse band here. Pronotal
scutum insignificantly convex, with two emarginations on anterior mar-
gin, medially extends markedly forward, bound laterally by longitudinal
groove, matte at base. Prothoracic presternum on disk with dense short,
laterally dense very long rusty hairs. Thoracic legs short but fully devel-
oped, with small poorly sclerotized claw, and diffusey ellowish colora-
tion.

Abdomen moderately elongate, narrows insignificantly from thorax
posteriorly, with sparse rusty hairs laterally. Dorsal locomotory ampul-
lae white, matte, divided by common median longitudinal groove, in
front by transverse groove joining longitudinal lateral grooves. Ventral
164 locomotory ampullae laterally with raylike dent. Abdominal sternite VI
with pair of hyaline ampullar-spherical processes on locomotory ampulla.
Body length of mature larvae 18 to 20 mm, width of head 2.2 to 2.8
mm.

Pupa (Figure 96): Differs from pupa of Callidium violaceum (L.) in
absence of spinules on dorsal side of body. Head short, moderately bent
under, transversely slightly convex between antennae, broadly curved on
vertex. Antennae pressed to sides, arcuate, bent ventrad posterior to mid-
femora.

Pronotum transverse, rounded laterally, narrows anteriorly and poste-
riorly, slightly convex on disk, almost flattened, without spinules. Meso-
notum short, medially longitudinally convex, impressed pitlike in region
of posterior angles around elytra, with extended elevated scutellum
posteriorly, without spinules, lustrous. Metanotum transverse, slightly
convex, medially with longitudinal groove, broadly rounded posteriorly,
glabrous, without spinules. Femora clavate; apices of hind femora extend
beyond abdominal tergite IV.

Abdomen elongate, with almost parallel sides (male). Abdominal
tergites uniformly convex, with slight median longitudinal groove, gla-
brous, without spinules. Body length 10 mm, width of abdomen 3.5 mm
(male).

221

163

Material. Collected in Trans-Baikal, Yakutia (Zhigansk region), Tuva,
Altai, and Ob’ region. Adult insects 43, larvae 177, pupa one (male).

Distribution: Europe, from boundaries of coniferous forests in the
north to the Mediterranean Sea, Asia from Salekhard, Zhigansk in the
north to Altai, Sayan, and Amur in the south. Common in Trans-Baikal
(Shilka River basin).

Biology: Mainly occurs in fir forests, more rarely in other forests.
Found in hills up to 2,000 m above sea level. Beetles sighted in first half
of June up to end of July. Lead cryptic mode of life. Not seen on flowers.
Fly to drying trees, mate there, and female later oviposits in bark cre-
vices. Mainly colonizes branches from lower to upper level, including
crowns. Quite often colonizes freshly felled and wind-fallen trees. Ferti-
lity of female comparatively high. Ovaries of one female just emerging
from cell contained 48 mature eggs.

Larvae live under bark and make meandering, usually longitudinal,
more rarely transverse galleries, impressed in alburnum. Mature larva
bores v/ood and makes cell there along trunk. Length of gallery in wood

222

4.0 cm, length of pupal cell 1.5 to 1.6 cm and width up to 0.5 cm. Cell
sealed from inlet with plug of frass. Larva pupates with head toward
inlet.

Pupation commences mid-May and ends in June. Emergence of young
beetles from pupal cells completed in June. Beetles emerge with develop-
ed gonads and do not need supplementary feeding. Reproduction com-
mences soon after emergence from wood. Weight of larvae before pupa-
tion (data of 11 specimens) 53 to 120 mg, pupae 45 to 105 mg, beetles
after emerging from cells 37 to 84 mg. Generation completed in two
years.

Mainly damages fir. For example, while collecting these insects in
nature, 140 were found on fir shoots, 20 on spruce, 20 larch, five pine,
and two maple; total, 187 insects. Population density sometimes compara-
tively high. On one fir shoot 2.5 cm thick and 18 cm long four insects
were recovered from wood, including two larvae and one beetle. On the
same shoot three larvae of Pogonocherus fasciculatus Deg. were found.
Not found on deciduous species.

165 3. Callidfum conaceum Payk.

Paykull, 1800, Fauna Suecica, vol. 3, p. 91; Plavil’shchikov, 1940,
Fauna SSSR, 22, 2, 291-294; Gressit, 1951, Longicorn Beetles of China,
vol. 2, pp. 224-225; Cherepanov and Cherepanova, 1973, Nov, i maloizv.
vidy fauny Sibiri, 6th ed., p. 56.

Adult (Figure 97): Head comparatively small, with uneven rugose
punctation, broad longitudinal suture between antennae, quite often here
less punctate, more lustrous. Eyes finely faceted, broadly but not very
deeply emarginate, with broad septum between lobes barely narrower than
upper lobes. Antennae slender, apices extend considerably beyond 0.50
(female) or 0.66 length of elytra (male); 3rd segment longer than 4th,
equal to 5th.

Pronotum broadens in posterior half, narrows less anteriorly, more
posteriorly, slightly convex on disk or even flat and here medially with
large, sometimes uneven punctation throughout length, laterally with dense
minute punctation, quite often with lustrous smooth band anterior to
scutellum. Scutellum minute, broadly rounded apically, smooth or with
sparse punctation, flat or longitudinally somewhat impressed. Pro-, meso-,
and metasterna with dense minute punctation, prosternal process long,
cuneiform and pointed apically. Elytra with parallel sides, slightly con-
vex, with insignificantly projecting rounded humeri, individually rounded
and with densely rugose punctation apically, in anterior half with large
and posterior half less large punctation, with semiadherent minute light-
colored hairs. Body ventrally with dense semiadherent light-colored hairs.
Hind femora extend beyond apex of elytra (male) or considerably short

223

Figure 97. Callidium coriaceum Payk.

of reaching it (female). First segment of hind tarsi longer than two suc-
cessive together.

Body black, blackish-brown; abdomen generally with rusty or chest-
nut hue, legs dark brown; elytra rusty-brown, with bronze or bluish-
bronze iridescence, lighter in color toward apex; antennae dark brown or
166 dark rust (f. typica). Sometimes elytra greenish, legs light rust (ab. aenei-
penne Muls.). Body length 8.0 to 14.0 mm.

Egg: White, elongate, broadly and obtusely rounded at one pole,
pointed at the other. Chorion with minute noncellular sculpture. Length
1.9 mm, width 0.6 mm.

Larva (Figure 98): Characterized by fine longitudinal striation of pro-
notal scutum, structure of locomotory ampullae characteristic of the sub-
genus Palaeocallidium Plav., and other features. Head markedly retract-
ed into prothorax. Epistoma with slight median longitudinal suture,

224

fuses laterally with parietals (frontal sutures not visible), on anterior
margin with dense, thin, longitudinally directed striation and also rusty-
brown border. Hypostoma smooth on anterior margin, without spinules,
with narrow rusty-brown border. Hypostomal suture straight, very dis-
tinct. Gula narrows markedly anteriorly, with narrow anterior margin,
two transverse narrow grooves at base. Hairs on anterior half of parie-
tals sparse, form dispersed tuft. Antennae slender; 1st segment insigni-
ficantly thicker than 2nd. Ocelli (one on each side) hyaline, somewhat
dorsoventrally elongate. Clypeus very short, white, looks like transverse
band. Labruni almost triangular, narrows anteriorly from base, not
pointed apically, with short setae along margins, white, brownish at base.
Mandibles on outer side of base with faint longitudinal groove.

Pronotum on disk and laterally with dense rusty hairs forming two
transverse bands, one in anterior half, the other anterior to scutum. Hairy
band with glabrous clearance between them. Hairs basally with fairly
distinct sclerotized ringlet. Yellowish-rust spots in anterior third of pro-
notum distinct, on disk broad, tetragonal, transversely elongate, laterally
more oval, longitudinally elongate. Pronotal scutum insignificantly convex,
only slightly produced anteromedially on anterior margin, tapers straight-
ly from there to sides, with dense thin longitudinal striation, at base
shagreen, matte, with very minute nonstriate sculpture.

Prothoracic presternum glabrous as in other species of this genus, on
disk with short hairs and laterally with long dense hairs; eusternum

165

Figure 98. Larva of Callidium coriaceum Payk.
a — head and pronotum; b — abdominal tergite
with dorsal locomotory ampulla.

225

basally in form of glabrous plates divided anteriorly by hairy clearance.
Thoracic legs small, with thin claw, dark brown on anterior and poste-
rior sides and more sclerotized there.

Abdomen narrows posteriorly from thorax, in region of segments III
to VI with parallel sides, segment VII usually somewhat broadened,
laterally with thin rusty, sometimes rather dense hairs. Dorsal locomo-
tory ampullae matte, divided by two transverse grooves joined laterally
with lateral longitudinal grooves, anterior groove invariably continuous,
apically outcurved, posterior groove often medially interrupted and repli-
cate. Ventral locomotory ampullae moderately convex, divided by single
transverse groove, broadly interrupted medially, and laterally joins short
lateral longitudinal groove. Sometimes these grooves are shortened,
resemble depressions with three rays, or curve backward, bracketlike.
Length of mature larvae 20 to 22 mm, width of head 2.8 to 3.0 mm.

Pupa: Not known.

Material: Collected in the eastern Urals, Ob’ region, Altai, Tuva,
north of Yakutia (Zhigansk and Aldan), Primor’e, and Ussuri- Primor’e
167 region. Adult insects 28, larvae 40.

Distribution: Viable zone of coniferous forests. Europe from Sweden,
Finland, and Karelia almost up to the Mediterranean Sea (mainly in hills
there), northern Asia from the Urals to the Pacific Ocean coast, including
Siberia, northern Mongolia, northern China, and Korea. Sporadic every-
where, comparatively few in numbers.

Biology: Ecologically associated with coniferous forests. Inhabits
plains as well as hilly regions. We found it around Lake Telets at a height
of 2,000 m above sea level. Beetles fly in June and July. At this time
found on trunks of drying and freshly felled trees where mating takes
place. Female oviposits in bark crevices.

Newly hatched larvae bore bark, live under it, and make longitudinal
meandering, at places platformlike galleries. They penetrate wood before
pupation and make cell in upper layer along trunk. We detected one
gallery and cell. Gallery extended top downward. Length of gallery under
bark 23 cm, width 0.6 to 1.6 cm. Length of cell 2.5 cm, width 0.9 cm.
Beetle (female) lay in cell with head toward inlet. Ovaries of this female
contained 48 fully mature eggs. Generation completed in not less than
two years. Colonizes spruce, larch, and Siberian maple, in that order.
While inspecting forests we found 45 insects (larvae and adults) in wood;
of these, 23 were recovered from Siberian spruce, 12 from larch, nine
from Siberian maple, and one from fir.

4. Callidium chlorizans (Sols.)

Solsky, 1870, Horae Soc. Entom. Ross., vol. 7, p. 3S4 (Semanotus);
Plavil’shchikov, 1940, Fauna SSSR, 22, 2, 294-295; Gressit, \95\, Longi-
corn Beetles of China, vol. 2, p. 224.

226

Adult (Figure 99): Similar to Callidium coriaceum Payk. Differs in
greenish-bronze upper part of body, which is very long. Head with dense
large punctation, spaces between punctures narrow, smaller than punc-
tures, and broad longitudinal groove between antennae. Eyes distinctly
and finely faceted, with broad shallow notch; septum between lobes slight-
ly narrower than upper lobe. Antennae apically reach only 0.50 (female)
or slightly more than 0.66 length of elytra (male), with minute gray
adherent hairs, basally with much longer hairs, apically shorter ones.

168

Figure 99. Callidium chlorizans (Sols.).

227

Pronotum broadens anterior to middle, broadly rounded laterally in
anterior half, narrows basally (but gently) and less apically (steeply),
straightly truncate on anterior margin or more often broadly and insig-
nificantly emarginate and notably curved medially here, fairly convex,
with deep large punctation, fine light-colored erect hairs; spaces between
punctures on disk sometimes fuse to form one smooth (usually median)
or two or three facets that are usually longitudinally elongate. Scutellum
minute, smooth, broadly rounded posteriorly, glabrous, without hairs.
Elytra long, elongate, with distinctly parallel sides, individually rounded
apically, basally on disk more convex with rounded or slightly projecting
humeri, longitudinally impressed in second third along suture, with large
dense punctation forming transverse wrinkles on disk, and light-colored
semiadherent hairs. Prosternum with dense, almost fused punctation.
Prosternal process long, elongate, pointed or more rarely rounded apically.
Metasternum with dense notchlike punctation, abdomen with sparser
punctation.

168 Body ventrally with dense light-colored adherent and erect hairs.
Abdominal sternites laterally with deep circular, comparatively large pits.
Sternite V in female more elongate, rounded apically or sometimes narrow-
ly emarginate, transverse in male and gently rounded apically. Hind
femora extend slightly beyond apex of elytra or fall considerably short
of it. Top of head, pronotum, and elytra dark green with bronze or pur-
plish-bronze metallic iridescence. Legs dark brown, with reddish-rust or
chestnut tone. Body ventrally rusty-brown, chestnut. Antennae dark brown
or almost black, sometimes lighter in color, rusty. Body length 9.0 to
17.0 mm.

Egg: White, markedly elongate, slender, narrows gradually more
toward one pole and pointed, narrowly rounded at the other. Chorion
transparent, smooth, hyaline, without perceptible sculpture. Length 2. 1
mm, width 0.5 mm.

Larva (Figure 100): Well distinguished from larvae of proximate spe-
cies Callidium coriaceum Payk. by absence of striation on anterior margin
of epistoma and sclerotized ringlets at base of hairs on pronotum. Head

169 small, narrows somewhat anteriorly. Epistoma on anterior margin with
broad, sharp, dark brown, smooth border altogether devoid of striation,
in posterior half with dark brown median longitudinal suture, fuses later-
ally with parietals. Frontal sutures lacking. Hypostoma narrows some-
what anteriorly, on anterior margin with narrow dark rust border, deep
notch near inner angles of sclerites bearing articulate spinule of maxillae.
Gula gradually or angularly narrows steeply anteriorly, slightly convex,
basally without perceptible transverse grooves. Parietals in anterior half
with thin, sometimes fairly dense hairs. Antennae slender, comparatively
long; 1st segment white, subsequent ones yellowish-brown. One convex

228

hyaline ocellus near base of each ant enna. Clypeus short, projects some-
what from behind epistoma, white, trapezoid, comparatively broad. Lab-
rum minute, broadly rounded apically, with sparse short setae, whitish,
basally brownish. Mandibles massive, thick, smooth and convex on
outer side in anterior half, flattened in posterior half, medially without
perceptible longitudinal groove, broadly rounded apically. Maxillary
palps short, not longer than inner masticatory lobes. Latter broadly
rounded apically, with short light-colored setae.

Pronotum laterally rounded, narrows notably more anteriorly, slopes
toward head, with two transverse tetragonal yellowish (sometimes faint)
spots in anterior^ half, longitudinally elongate spot laterally, thin light
170 rust hairs on disk forming two compact transverse bands. Hairs basally
without sclerotized ringlets. Laterally, light rust hairs form extensive field.
Pronotal scutum insignificantly convex, with usually two emarginations
on anterior margin, medially more elongate, anterior angles less elongate,
with dense thin longitudinal striation, basally matte and with minute
shagreen sculpture, with short setae forming transverse row separating
striate section of scutum from nonstriate, matte, bound laterally by short

169 Figure 1 00. Larva of Callidium chlorizans (Sols.).

a — head and pronotum; b — abdominal tergite with
dorsal locomotory ampulla.

229

longitudinal grooves. Alar lobes glabrous on inner side. Prothoracic pre-
sternum generally with short hairs on disk, laterally with very long and
very dense rusty hairs; glabrous plates of eusternum divided medially by
compact hairy band, with coarse punctation or minute furrows. Thoracic
legs dark brown on outer side, light rust on inner side, with sharp aci-
cular claw.

Abdomen laterally with dense long rusty hairs. Dorsal locomotory
ampullae fairly convex, matte, with common median longitudinal groove,
with two transverse grooves, of which anterior one continuous, posterior
one broadly interrupted medially; transverse grooves merge laterally with
short longitudinal grooves. Sometimes transverse grooves merge among
themselves at an acute angle. Ventral locomotory ampullae also matte,
divided by common broad median longitudinal groove; short transverse
groove on disk broadly interrupted medially and laterally merges with
short longitudinal grooves. Body length of mature larvae 20 to 22 mm,
width of head 2.2 to 2.5 mm.

Pupa (Figure 101): Characterized by poorly developed spinules on
abdominal tergites, convex vertex in same plane as frontal gap between
antennae. Head short, narrows anteriorly from antennae. Frons between
antennae broad, almost flat, in same plane as vertex. Occiput broadly
rounded, with sparse large punctation forming clusters. Antennae com-
paratively short, pressed to sides, with apices bent ventrad posterior to
midfemora (female) or ventrad and forward (male).

Pronotum laterally rounded, narrows less toward apex, more so to-
ward base, slightly convex on disk, lustrous, smooth, without spinules,
medial width not more or only slightly more than length. Mesonotum
lustrous, longitudinally convex, slightly slopes laterally, with produced
scutellum on posterior margin, and devoid of spinules. Metanotum insig-
nificantly convex, with faint median longitudinal groove, broadly round-
ed posteriorly, without spinules. Femora clavate, thicken gradually toward
apex.

Abdomen elongate (male) or broadens somewhat in region of segment
IV (female), sometimes narrows gradually from base to apex. Abdominal
tergites uniformly convex, with common median longitudinal groove, in
posterior half with minute sharp spinules forming transverse, usually
indistinct row. Spinules more distinct in female, barely or not at all per-
ceptible in male. Tergite VII narrowly (male) or broadly rounded at
apex, convex and smooth on disk, without spinules. Tergite VIII in
female elongate, with parallel sides, in male short, narrows toward apex.
Valvifers of female (ventral view) large, hemispherical, matte at apex,
with fine sculpture. Body length 10 to 15 mm, width of abdomen 3.0 to
4.0 mm.

230

169 Figure 101. Pupa of Callidium chlorizans (Sols.).

Material: Collected in Altai, Tuva, Baikal region, and Ussuri-Primor’e
region. Adult insects 25, larvae 58, pupae 10 (seven males and three
females).

171 Distribution: Northern Asia, including Altai, Tuva, eastern Siberia,
northern Mongolia, northern China, Korea, northern Japan. Sporadic.
We sighted large numbers in northern offshoots of Tannu-01 range in
Tuva.

Biology: Inhabits coniferous forests and is ecologically associated with
larch. Beetles fly in first half of June up to end of July, and sighted at
this time on trunks of trees. Mainly colonizes lower section of larch
trunks in zone of thick bark.

Larvae live in bark and make meandering galleries in bark layer with-
out affecting bast. On removing bark, thin layer of beast covering gallery
filled with fine frass visible on inner side. Width of gallery 0.6 to 1.0 cm.
Gallery ends in bark with pupal cell. Beetle nibbles short exit to surface

231

from cell, leaving fine layer of bark outside. Cell along trunk axis or obli-
que. Length of pupal cell 1.5 to 2.0 cm, width 0.6 to 0.9 cm.

Pupation of larvae commences end of May and ends in June. Dura-
tion of pupal stage about three weeks. Beetles develop from pupae in
June, but emerge from cells in June and first half of July. In Tuva beet-
les and Il-instar larvae were seen simultaneously on trees on July 10,
indicating that the same tree is colonized repeatedly. Beetles emerge from
cells with developed gonads. Ovaries of two females recovered from cells
contained 16 and 30 mature eggs respectively.

Weight of insects varies considerably. Weight of larvae 37.0 to 114.2
mg, pupae 34 to 100 mg, and beetles in cells 22.5 to 96.0 mg. The follow-
ing example demonstrates weight variation during metamorphosis. Three
specimens in the larval stage before pupation weighed 263.7 mg (100%),
pupal stage 230.2 mg (87,2%), and adults 185 mg (70.1%). Generation
completed in two years (Table 15).

Table 15. Periods of development of CaUidium chlorizans (Sols.)

Year of
development

April

May

June

July

August

September

1st

LPAE

PAE

2nd

3rd

LPAE

PAE

Note: Periods of development of other species of this genus, described above,
are almost identical.

Found only on larch. In Tuva in 1976 often found on drying trees
priorly damaged by the Siberian silkworm. Tetropium gracilicorne Reitt.
and Clytus arietoides Reitt. colonize together with this species.

7. Genus Phy mat odes Muls.

Mulsant, 1839, Col. France Longic., p. 47; Plavil’shchikov, 1940,
Fauna SSSR, 22, 2, 303-307; Gressit, 1951, Longicorn Beetles of China,
vol. 2, pp. 226-227; Linsley, 1964, Cerambycidae of North America, 22,
5, 44-45; Kojima and Hayashi, 1969, Insect Life in Japan, voX. l,pp.
12-1 y, Mamaev and Danilevskii, 1975, Lichinki zhukov-drovosekov, pp.
201-202.

Adult: Characterized by comparatively small body (especially in the
subgenus Phymatodellus Reitt.). Pronotum laterally rounded, with
smooth shields on disk [P. testaceous (L.), P. abietinus Plav. and Lur.]
172 or without them {P. zemlinae Plav. and Anufr., P. mediofasciatus Pic).

232

Apex of femora thickened, clavate, markedly dilated [P. maaki (Kr.)]
or thickens gradually (P. abietinus Plav. and Lur.), Elytra slightly convex
or notably flattened. First segment of hind tarsi 1.5 times longer than
successive two together {Phymatodes s. str.) or not longer (Phymatodellus
Reitt.).

Larva: Differs from larvae of other genera in pronotum with yellow-
ish-rust transverse band interrupted three times by white band into
four equal tetragonal spots covered with dense hairs. Head markedly
retracted into prothorax; one ocellus near base of each antenna. Epi-
stoma fuses with parietals, frontal sutures not visible, median longitu-
dinal suture projects more distinctly only in posterior half. Pronotum
usually narrowly rounded anteriorly, on disk in region of proscutellum
and laterally with dense hairs. Pronotal scutum uniformly convex, rugu-
lose or bulges more basally and here with minute reticulate- squamiform
sculpture, matte, notably flattened anteriorly and lustrous, longitudi-
nally striate. Thoracic legs poorly developed, wartlike, sometimes almost
imperceptible, especially in young larvae. I-instar larvae (P. ermolenkoi
Tsher.) laterally on abdominal segments VI and VII with one acute,
slightly sclerotized spinule on each side, which disappears after molt.

Pupa: Body moderately elongate. Head short, with widely separated
antennae, wide frons between them. Antennae pressed to sides, posterior
or midfemora bent ventrad, with apices directed forward. Femora clavate.
Abdominal tergites II to VII with pair of widely separated spinules (P.
ussuricus Plav., P. vandykei Gress.) or without them (P. abietinus Plav.
and Lur.) or with four to twelve spinules [P. mediofasciatus Pic, P. maaki
(Kr.)] forming transverse rows. Tip of abdomen rounded, without spi-
nules.

No less than 16 species of this genus are known in the USSR; of
these, nine are found in northern Asia including one [P. alni (L.)] rang-
ing west to the Urals inclusive, one (P. abietinus Plav. and Lur.) cover-
ing southern regions of western Siberia, five east in Ussuri-Primor’e
region, and one [P. testaceus (L.)] in Europe, southern Urals, Japan, and
North America. Not known on the continent between the Urals and
Japan. In Southeast Asia 10, Japan five, and North America 24 species
of this genus are known.

All species of the genus Phymatodes Muls. inhabiting northern Asia
are ecologically associated with forests. Among them, P. maaki (Kr.), P.
mediofasciatus Pic, P. ussuricus Plav., and P. vandykei Gress. develop
on grapevine, P. testaceus (L.) and P. alni (L.) develop on oak and other
deciduous species. Phymatodes abietinus Plav. and Lur. develops on Sibe-
rian fir and P. ermolenkoi Tsher. on oak.

Type species: Cerambyx testaceus Linnaeus, 1758.

233

KEY TO SPECIES
Adult Insects

1 (10). Elytra without transverse white bands.

2 ( 3). First segment of hind tarsi 1.5 times longer than two successive

together (subgenus Phymatodes s. str.). Europe up to southern

Urals inclusive 1. P. testaceus (L.).

3(2). First segment of hind tarsi not longer, or only slightly longer
than two successive together (subgenus Phymatodellus Reitt.).
173 4 ( 5). Body red or reddish-rust; elytra bluish. Ussuri-Primor’e region.

2. P« zemlinae Plav. and Anufr.

5 ( 4). Body black or blackish-brown; elytra without bluish iridescence.

6 ( 9). Elytra comparatively short, not more than 3.0 times longer than

pronotum. Femora sharply thickened toward apex, clava mar-
kedly dilated.

7 ( 8). Humeral tubercles of elytra do not project, gently rounded, in-

ner side with faintly visible impression. Ussuri-Primor’e region.
3. P. ussuricus Plav.

8 ( 7). Humeral tubercles of elytra distinctly project forward, inner side

with very distinct longitudinal impression. Kunashir and Japan.

4. P. vandykei Gress.

9(6). Elytra very elongate, long, not less than 4.0 times longer than
pronotum. Femora thicken gradually toward apex, clava not

markedly dilated. Western Siberia

5. P. abietinus Plav. and Lur.

10 ( 1). Elytra with transverse white bands.

11 (12). Elytra with single white band, on suture without erect hairs

posterior to scutellum (subgenus Paraphymatodes Plav.) Ussuri-
Primor’e region 6. P. mediofasclatus Pic.

12 (11). Elytra with two white bands, on suture with tuft of upright

black hairs (subgenus Poecilium Fairm.).

13 (16). Pronotum with dense, long, erect brownish hairs forming es-

pecially dense brush laterally. Hairs not shorter than 3rd an-
tennal segment. Ussuri-Primor’e region.

14 (15). First to 6th antennal segments with stray hairs, 2nd segment

almost not longer than wide 7. P. maaki (Kr.).

15 (14). First to 6th antennal segments with dense setaceous hairs, 2nd

segment almost 2.0 times longer than wide

9. P. erraolenkoi Tsher.

16 (13). Pronotum with short erect hairs, not forming dense brush. Hairs

shorter than 3rd antennal segment. Europe to the Urals in-
clusive 8. P. alni (L.).

234

174

Larvae

1(2). Pronotum anterior to scutum with sparse thick setaceous hairs.
Pronotal scutum lustrous, with deep longitudinal striation.

Found on trunks of oak and other deciduous species

1. P. testaceus (L.).

2(1). Pronotum anterior to scutum with dense hairs; if with sparse
hairs, then hairs thin and nonsetaceous. Pronotal scutum with
dense thin, fairly distinct longitudinal striation or without it.

3 (10). Pronotal scutum more convex at base, with fine reticulate-

squamiform sculpture imparting matte tone, lustrous anteriorly,
longitudinally striate.

4 ( 5). Transverse yellowish-rust band in anterior third of pronotum

continuous, without white clearances, only with three small

flanges. Found on grapevine

2. P. zemlinae Plav. and Anufr.

5 ( 4). Transverse yellowish-rust band in anterior third of pronotum

with three longitudinal interruptions separating it into four
transvere spots.

6 ( 9). Pronotal scutum in anterior half with sparse longitudinal streaks;

distance between streaks several times more than their width.

7 ( 8). Dorsal locomotory ampullae with coarse longitudinal striation.

Inner masticatory lobes of maxillae narrowly rounded at apex.
Pronotal scutum at base with narrow, convex, matte transverse
band covering not more than 0.25 of scutum. Found on grape-
vine 3. P. ussuricus Plav.

8(7). Dorsal locomotory ampullae without perceptible longitudinal
striation. Inner masticatory lobes of maxillae broadly rounded
at apex, as though obtuse. Pronotal scutum at base with much
wider matte transverse band covering not less than 0.33 of
scutum. Found on grapevine. ........ 4. P. vandykei Gress.

9 ( 6). Pronotal scutum in anterior half with very dense longitudinal
streaks; distance between streaks not more than their width.
Found on fir 5. P. abietinus Plav. andLur.

10 ( 3). Pronotal scutum uniformly convex, lustrous, at base with faint

reticulate- squamiform sculpture or without it, rugulose, with
longitudinal streaks.

11 (12). Hairs on yellowish-rust spots of pronotum simple, basally with-

out sclerotized ringlets. Found on grapevine

6. P. mediofasciatus Pic.

12 (11). Hairs on yellowish-rust spots of pronotum basally with distinct

sclerotized or barely projecting ringlet.

235

13 (14). Hairs on yellowish-rust spots of pronotum basally with highly

sclerotized ringlet. Found on grapevine

7. P. maaki (Kr.).

14 (13). Hairs on yellowish-rust spots of pronotum basally with slightly

sclerotized ringlet. Found on other species.

15 (16). Yellowish-rust spots in anterior third of pronotum on disk

transverse, slightly elongate, almost square, slightly longer in

cross section than lengthwise. Found on oak

8. P. alni (L.).

16 (15). Yellowish-rust spots in anterior third of pronotum on disk

transverse, highly elongate; in cross section (relative to body

axis) almost 2.0 times length. Found on oak

9. P. ermolenkoi Tsher.

Pupae

1 ( 2) Abdominal tergites II to VI with six to nine paramedial spinules
in posterior third forming transverse distinct or indistinct row.

Found mainly on oak 1. P. testaceus (L.).

2(1). Abdominal tergites II to VI with one to four well-developed
or faint paramedial spinules in posterior third forming one or
tv/o indistinct transverse rows. If with large number of spinules,
found on grapevine.

3 (12). Clava of hind legs thickens gradually from base to apex. Spi-

nules on abdominal tergites minute, poorly developed.

4 (11). Spinules on abdominal tergites II to VI form single transverse

row.

5(6). Abdominal tergites II to VI with four to six paramedial spi-
nules forming indistinct transverse row

2. P. zemlinae Plav. and Anufr.

6 ( 5). Abdominal tergites II to VI with or without one or two para-

medial spinules.

7 (10). Pronotum laterally in anterior third without flange. Abdominal

tergites II to VI with perceptible spinules.

8 ( 9). Spinules (two) on abdominal tergite VII erect or bent down

and backv/ard 3. P. ussuricus Plav.

175 9 ( 8). Spinules (tv/o) on abdominal tergite VII large, bent down and
forward 4. P. vandykei Gress.

10 ( 7). Pronotum laterally in anterior third with fairly distinct flange.

Abdominal tergites II to VI without visible spinules .

5. P. abietinus Plav. and Lur.

11 ( 4). Spinules on abdominal tergites II to VI form two transverse

rows 6. P. mediofasciatus Pic.

236

12 ( 3). Clava of hind legs in second half sharply thicken, dilated. Spi-

nules on abdominal tergites well developed; if poorly develop-
ed [P. alni (L.)], insects develop mainly on oak.

13 (16). Spinules on abdominal tergites (two to four on each side of

longitudinal groove) large, acute, directed backward.

14 (15). Abdominal tergites II to VI with three or four spinules on each

side of longitudinal groove. Found on grapevine

7. P. maaki (Kr.).

15 (14). Abdominal tergites II to VI with two or three spinules on each

side of longitudinal groove. Found on oak

9. P. ermolenkoi Tsher.

16 (13). Spinules on abdominal tergites minute, barely developed. Found

mainly on oak 8. P. alni (L.).

1. Phymatodes testaceus (L.)

Linnaeus, 1758, Syst. Nat., 10th ed., p. 396 {Cerambyx); Plavil’shchi-
kov, 1940, Fauna SSSR, 22, 2, 307-312; Duffy, 1953, Monograph Beetles
pp. 217-219; Romadina, 1954, Trudy Zool. In-ta, vol. 16, p. 219; Naka-
mura, 1958, Miscell. Report of Hiwa Museum for Nat. History, vol. 2,
pp. 14-15; Nakamura, 1960, ibid., vol. 3, pp. 1-2; Linsley, 1964, Ceram-
bycidae of North America, 22, 5, 63-64; Demelt, 1966, Tierwelt Deutsch-
lands, 52, 2, 71-72.

Adult {VigutQ 102): Differs from other species of this genus in mar-
kedly elongate, with parallel sides, yellow or metallic blue elytra. Head
between antennae transversely convex, with median longitudinal suture,
smoothened punctation, transversely impressed between upper ocular
lobes. Eyes sharply faceted, deeply emarginate. Antennae with short
adherent hairs; 10th segment extends beyond apex of elytra (male) or
considerably short of it (female).

Pronotum laterally rounded or angularly slightly produced, narrows
less anteriorly, notably posteriorly, on posterior margin with narrow
transverse groove, convex on disk, with moderate punctation, short dark
brown or light-colored hairs, with three smooth lustrous shields, of which
two anterolateral to middle and one median in posterior half. Scutellum
fiat, broadly rounded posteriorly, with minute punctation, and adherent
light-colored hairs. Elytra markedly elongate, with parallel sides, bulge
slightly on disk, apically individually or almost jointly rounded broadly,
with minute uniform, sometimes smoothened punctation, rugulose sculp-
ture in clearances between punctures, with short semiadherent light-
colored hairs. Prosternum with large deep (male) or minute faint (female)
punctation. First segment of hind tarsi longer than two successive
together.

Body yellowish-rust, with reddish tinge (especially on pronotum),
metathorax black, abdomen for most part (male) or only at base (female)

237

black or dark brown, elytra light colored, rusty or rusty-brown, legs
and antennae reddish-rust (f. typica); quite often elytra light rust, dark
brown or blue at apex or most of posterior half (ab. praeustus F.), more
often elytra entirely blue (ab. variabilis L.). Color highly variable. Large
176 number of aberrant forms known (Plavil’shchikov, 1940). Body length
7.0 to 16.0 mm.

Egg: White, moderately elongate, narrows more toward one pole,
broadly rounded or notably obtuse at one end, narrowly rounded at the

238

other. Chorion smooth, hyaline. Length 1.2 mm, width 0.5 mm.

Larva (Figure 103): Head short, markedly retracted into prothorax,
Epistoma faintly convex, broadly emarginate on anterior margin, with
smooth narrow dark brown border, behind which stray short setae form
transverse row, with faint or almost imperceptible longitudinal suture in
posterior half. Frontal sutures not visible, Epistoma laterally fuses with
parietal s. Hypostoma slightly convex, smooth, with narrow brownish
border on anterior margin; hypostomal sclerites with extended sharp in-
ner angles, narrow markedly on posterior inner margin. Gula basally
very broad, narrows anteriorly, without brownish border on anterior
margin. Parietals on anterior margin ventral and dorsal to antennae with
177 broad rusty-brown border, posterior to antennae with extensive, trans-
versely oval, elongate, pigmented ocellus, with not very long hairs form-
ing transverse band. Antennae slender, apices project forward beyond
anterior margin of cephalic capsule; 2nd and 3rd antennal segments brown-
ish, whitish ringlet at apex. Clypeus lustrous, semitransparent or whitish,
narrows toward apex, broadens markedly at base, flattened, strip-
like. Labrum semitransparent, broadens in posterior half, narrows an-
teriorly, rounded apically, with short sparse setae in anterior half. Labial
submentum transverse, slightly convex; mentum slightly longitudinal,
narrows slightly apically, with long setae laterally. Inner masticatory lobes
of maxillae comparatively thick, rounded apically and with short sparse

Figure 103. Larva of Phymatodes testaceus (L.),
a— head and pronotum; b — abdominal tergite IV
with dorsal locomotory ampulla.

239

setae there. Maxillary palps thin, barely longer than inner lobes. Mandi-
bles black, with contrasting red border basally, and median longitudinal
groove on outer side of base.

Pronotum slopes distinctly toward head, with yellowish or rusty spots
in anterior third forming transverse band (two of these spots on disk
transverse, two on sides longitudinally elongate), in anterior half in front
of scutum and laterally with rusty, not very long hairs, without distinct
ringlet at base. Pronotal scutum convex, white, with narrow median longi-
tudinal groove, bound laterally by short longitudinal folds, with two
emarginations on anterior margin, medially produced anteriorly, with
coarse longitudinal striation, matte silver basally and here with minute
reticulate- squamiform sculpture. Prothoracic presternum uniformly con-
vex, with dense rusty hairs; eusternum not bound by groove, merges
with presternum, glabrous at base, lustrous, coriaceous, with rounded
hairy clearance only on anterior margin medially. Thoracic legs not visi-
ble in I-instar larvae, fully developed in Il-instar and mature larvae.

178 Abdomen comparatively thick, narrows insignificantly posteriorly,
with sparse short hairs laterally. Dorsal locomotory ampullae convex,
transversely elongate, with barely visible common median longitudinal
groove, without grooves laterally and on disk, only sometimes with pro-
minent longitudinal, barely visible narrow dents, or with transverse,
barely visible narrow groove (on tergites I, II, and VII), with minute
reticulate-squamiform sculpture that imparts matte silver tone. Ventral
locomotory ampullae bulge, transversely elongate, cover much of ster-
nites, with minute squamiform sculpture. Pattern on dorsal ampullae
given by Duffy (1953) and Mamaev and Danilevskii (1975) evidently not
correct; either described from a damaged larva of Xylotrechus antilope
(Schonh.) or some other species. Body length of mature larva 10 to 18
mm, width of head 2.1 mm.

Pupa (Figure 104): Differs from other species of this genus in very
large body and innumerable minute spinules on abdominal tergites. Head
narrower than prothorax, on occiput convex, smooth, on vertex between
upper ocular lobes widely impressed, transversely striate. Frons between
antennae insignificantly convex, with median longitudinal suture, bet-
ween lower ocular lobes with coarse transverse furrows. Antennae com-
paratively long, slender, markedly bent ventrad in second half.

Pronotum transverse, angularly produced mediolaterally, narrows
identically anteriorly and posteriorly; disk slightly convex, with thin
transverse striation, glabrous, without setae and spinules, with small
medial tubercle near anterior margin. Mesonotum glabrous, in anterior
half slightly convex, on posterior margin with angularly produced scutel-
lum, transversely impressed slightly in middle. Metanotum glabrous,
without setae, broad, with median broad longitudinal cavity, transverse

240

177

Figure 104. Pupa of Phymatodes testaceus (L.), female.

striation on disk, broadly rounded on posterior margin.

Abdomen elongate, with almost parallel sides, narrows slightly an-
teriorly and markedly posteriorly from segment VI. Abdominal tergites
convex, laterally with thin longitudinal outcurved furrows, in posterior
third with minute spinules forming transverse row on tergite I; tergites
II to VI with transverse band interrupted medially (six to nine parame-
dial spinules in band). Tergite VII narrows posteriorly, broadly rounded
on posterior margin, bulges slightly on disk, with very minute, specklike
spinules forming transverse row near posterior margin directed backward.
Valvifers of female comparatively large, hemispherical, notably narrow
toward base on inner side. Hind femora gently clavate and apices barely
reach posterior margin of tergite IV. Body length 9.0 mm, width of
abdomen 2.8 mm.

Material: Collected in broad-leaved forests of the southern Urals.
Adult insects 14, larvae six, pupa-— one female, exuviae with beetles and
pupae from cells three. One pupa and three adults raised from larvae
hatched from eggs laid in the laboratory.

241

Distribution: Extends from Atlantic Ocean coast to the southern Urals
inclusive, north from Sweden and Norway, south to northern Africa and
Syria; also occurs in Japan and North America. Difficult to consider
this species imported from Europe. It probably covered the entire Holarc-
tic, then disappeared from Siberian territory but was preserved within
the present range.

Biology: Inhabits various forests; confined more to oak-broad-leaved
forests. Beetles fly from May through June, sometimes up to mid-July.

179 Flight occurs in second half of day, with insects active in warm weather
before sundown. Often confined to drying oak and mates there. Female
lays eggs singly (some distance apart) in bark crevices of trunk. Fertility
comparatively high. For example, ovaries of one female dissected before
oviposition contained 42 eggs, of another female 124. Embryonic develop-
ment takes more than two weeks. Larvae began hatching on July 9 to
10 from eggs laid by beetles in a forest under a gently sloping tree on
June 22 to 24. Total duration of egg development 15 to 17 days. Atmo-
spheric temperature during this period 15 to 27°C (22.1 + 0.5°C).

Larvae live in and under bark, make meandering longitudinal galleries,
leaving no impression on alburnum, and plug them with fine frass of
bark particles. Galleries made by larvae sometimes narrow, sometimes
broader, sometimes resemble platforms. Mature larva makes cell on
thick-barked tree under bark or even in upper layer of wood parallel or
oblique to trunk axis. Length of gallery made by larva 13.5 cm, width

9.0 to 15.0 mm. Length of pupal cell 14 to 21 mm, width 5.0 to 8.0 mm.
Exit from cell to surface up to 6.0 mm long. Thin bark layer remains
between exit and surface. Larvae pupate early spring. Pupa lies in cell
with head upward, develops for 15 to 17 days. In the laboratory at 18.3
to 21.2°C one beetle emerged on the 15th, another on the 17th day after
appearance of pupa. Young beetles nibble oval flight opening (1.5 mm x

3.0 mm to 3.0 mm X 5.0 mm) on bark surface and emerge from cell
through it. Emergence of beetles commences in May and ceases by mid-
June, Life cycle from egg to mature adult completed in one to two years.
It is significant that development of eggs of the same batch under labora-
tory conditions at the same temperature (16 to 25®C) was completed in
nine months by some and over 1 0 months by others. Weight of larvae
before pupation (records of six insects) 44 to 84 mg, pupae 28.5 mg,
adults 22 and 69 mg.

Mainly colonizes oak. According to Plavil’shchikov (1940), Duffy
(1953), and Demelt (1966), it also attacks chestnut, elm, beech, alder,
willow, and other deciduous species. We found it in the southern Urals
in the basal section of trunks and on thick branches 1 8 to 30 cm in
diameter on drying and just desiccated oak trees.

242

2. Phymatodes zemlinae Plav. and Anufr.

Plavilstshikov [Plavil’shchikov] and Anufriev, 1964, Zool Zhurn., 43,
10, 1565-1569; Cherepanov and Cherepanova, 1974, Usachi vinograda
amurskogo, pp. 34-36.

Adult (Figure 105): Characterized by absence of smooth shields on
pronotum, red body color, and bluish metallic iridescence of elytra.
Head with dense deep punctation, between antennae with broad longi-
tudinal suture. Eyes markedly convex, coarsely faceted, with broad deep
notch. Antennae markedly shorter than body, apices barely reach poste-
rior margin of middle third of elytra (male) or extend slightly beyond
middle (female), with adherent rusty hairs; 1st to 8th segments with long
erect hairs, 5th segment shorter than 3rd, equal to 4th.

Pronotum convex, uniformly rounded laterally, near base with narrow
transverse groove, with uniform dense deep punctation, short rusty non-
adherent hairs, without smooth shields. Scutellum broad, insignificantly

180

Figure 105. Phymatodes zemlinae Plav. and Anufr.

243

180 longer than width at base, narrowly or broadly rounded posteriorly, with
dense deep punctation. Elytra with parallel sides, rather convex, rounded
jointly at apex, on inner side around humeri with short longitudinal
groove, with dense punctation forming rather distinct transverse wrinkles,
and short adherent hairs. Hind femora with elongate clava. First seg-
ment of hind tarsi slightly longer than 2nd and 3rd segments together.
Head, pronotum, underside of body, and scutellum rusty-red, eyes dark
brown, antennae chestnut, elytra bluish with metallic iridescence, femora
yellowish- red, tibiae and tarsi brownish. Body length 5.0 to 8.0 mm.

Larva: Body moderately elongate. More than half of head retracted
into prothorax. Epistoma with flat or medially faintly notched anterior
margin, with lustrous brownish border there, behind which setae form
transverse band; divided by median longitudinal suture, bound laterally
by faint frontal sutures. Hypostoma with triangular sclerites widely se-
parated by gula, slightly convex, with three striae diverging from poste-
rior angle. Clypeus trapezoid. Labrum small, rounded, in anterior half
with short setae. Mandibles massive, broadly rounded apically, hollow
inside, smooth outside, posterior to middle with transverse narrow groove,
black, reddish-rust at base.

Pronotum transverse, rounded laterally, narrows more anteriorly,
on anterior margin with rusty transverse continuous band extending

181 laterally, and anterior to scutum and laterally with rusty hairs. Pronotal
scutum convex, longitudinally rugose, with two emarginations on an-
terior margin, bound laterally by longitudinal folds, divided by median
longitudinal groove. Thoracic legs short, minute, wartlike.

Abdomen laterally with thin light-colored hairs. Locomotory am-
pullae developed on abdominal segments I to VII, divided by median
longitudinal groove, laterally with radial folds, faint on sternite VII.
Body length of mature larvae up to 10 mm, width of head 1.1 mm.

Pupa (Figure 106): Body moderately elongate, somewhat flattened.
Head broad, bulges between antennae, with median longitudinal suture,
without setae, uniformly rounded on occiput. Antennae pressed to sides,
bent ventrad posterior to midfemora, with apices directed forward.

Pronotum insignificantly convex on disk, rounded laterally, narrows
anteriorly and posteriorly, with setae on disk, without spinules. Meso-
notum short, transverse. Metanotum longitudinal, with median longi-
tudinal groove, and very gently rounded on posterior margin. Apices of
hind femora extend slightly beyond abdominal tergite IV.

Abdomen in region of segments IV and V broadens insignificantly,
narrows gradually posteriorly, with median longitudinal groove on dorsal
side. Abdominal tergites II to VI with minute sharp spinules in posterior
half forming indistinct transverse row (eight to twelve spinules in row).
Tergite VII apically broadly rounded, in posterior half close to posterior

244

180

Figure 106. Pupa of Phymatodes zemlinae
Plav. and Anufr.

margin with spinules forming transverse row, of which two bent down
toward center. Sometimes spinules wanting. Body length 5.0 to 8.0 mm,
width of abdomen 1.8 to 2.1 mm.

Material: Collected in Ussuri-Primor’e region. Adult insects (raised
in laboratory) three, pupae— -two males and one female, larvae 12, larval
exuviae with beetles from cells three.

Distribution: Ussuri-Primor’e region.

Biology: Colonizes broad-leaved forest biotopes. Ecologically associat-
ed with grapevine. Beetles fly in May and June, lead cryptic mode of
life. Not seen on flowers. After mating, female oviposits under intact
bark strip. Colonizes thin moist viable shoots of Amur grapevine (Vitis
amurensis) 3.0 to 7.0 mm in diameter.

Newly hatched larvae bore wood, make galleries along pith, and
plug them with fine frass. Upper layer of wood adhering to bark not
damaged. Length of gallery 8.0 to 15.0 cm, width up to 3.0 mm. Mature

245

larva hibernates. The following spring makes cell and pupates in it.
Length of cell 21 mm, width 3.0 mm.

Pupae found in May. Young beetles emerge from them in second half
of May and early June. Beetles nibble oval openings on shoot surface and
abandon cell. Size of openings 1.5 mm X 2.5 mm. Generation complet-
ed in one to two years.

Brachyclytus singularis Kr. colonizes thicker portions of the same
shoots, with diameter of 5.0 mm or more.

3. Phymatodes ussuricus Plav.

PlaviPshchikov, 1940, Fauna SSSR, 22, 2, 318; Cherepanov and
Cherepanova, 1974, Usachi vinograda amurskogo, pp. 26-30.

182 Adult (Figure 107): Characterized by small body, hind femora mar-
kedly dilated at apex, and other features. Head minute, retracted up to
eyes into prothorax, lustrous, with minute deep punctation, sparse long
hairs, near base of antennae elevated tubercularly, with median longitudi-
nal suture. Eyes distinctly faceted, markedly emarginate, upper lobes of
eyes half size of lower ones, septum between them narrow, with two rows of
facets. Antennae slightly thickened apically, last segment extends beyond
0.50 (female) or almost beyond 0.66 (male) length of elytra, with minute
adherent hairs; 1 st to 5th segments with long hairs.

Pronotum narrows less anteriorly, more posteriorly, rounded later-
ally, markedly produced near base and poorly near apex, flat on disk,
with minute deep punctation, light-colored erect hairs. Scutellum longer
than width at base, broadly rounded posteriorly, with uneven puncta-
tion. Elytra with parallel sides, 3.0 times longer than general width, with
projecting humeral tubercle, longitudinally impressed inward from it, flat
on disk, with narrowly rounded inner apical angle and gently tapered
outer, in anterior half with distinct deep punctation, in posterior half
with fine smoothened punctation, with rusty semiadherent short hairs.
Legs with long setae. Second half of femora markedly dilated, smooth,
with fine smoothened punctation. Hind tarsi 0.50 length of tibiae, 1st
segment equal to two successive together.

Abdomen convex ventrally, with sparse minute punctation, and long
semiadherent hairs. Body dark brown chestnut, elytra in anterior half
reddish-rust, gradually darken toward apex, quite often entirely dark
brown or chestnut. Body length 4.0 to 5.0 mm.

Egg: White, semitransparent, narrows more toward one pole. Chorion
smooth, hyaline. Length 1.0 mm, width 0.4 mm.

(Figure 108): Body small, elongate. Head narrowly rounded
anteriorly, markedly retracted into prothorax. Epistoma insignificantly
emarginate on anterior margin, with dark brown lustrous border; in
posterior half divided by entirely distinct median longitudinal suture;

246

poorly demarcated laterally; frontal sutures not visible. Hypostoma divid-
ed into two convex minute lustrous triangles, with tapered or rounded
outer anterior angle and sharply produced inner angle, on anterior mar-
gin with rusty diffuse border. Parietals in anterior half with stray hairs.
Clypeus minute, trapezoid, rusty-brown. Labrum narrowly rounded
183 apically, with sparse setae. Mandibles black, rusty at base, rounded at
apex, with cultrate edge; dorsoventral width not less than length. Inner
lobes of maxillae convex, thin, narrowly rounded apically. Maxillary palps
thin, do not project anteriorly or project slightly beyond apex of inner
masticatory lobes.

Prothorax narrowly rounded anteriorly, at base not narrower or some-
what wider than mesothorax. Pronotum transverse, convex on disk, slopes
slightly toward head, with four yellowish-rust spots on anterior margin
forming transverse band interrupted three times, in anterior half before
scutum and laterally with long rusty hairs. Pronotal scutum white, glab-
rous, without hairs, bound laterally by barely perceptible short longitudinal

247

Figure 108. Larva of Phymatodes ussuricus Plav., head and pronotum.

grooves, more convex at base, with minute reticulate-squamiform sculp-
ture, matte, lustrous for most part in front, with long longitudinal stria-
tion, with spaces between striae several times wider than striae. Alar
lobes with rusty hairs. Prosternum in anterior half and laterally with
long rusty hairs, on disk posterior to middle with pair of widely separat-
ed, lustrous, glabrous, sometimes faint spots. Thoracic legs lacking, or
barely perceptible in mature larvae (under high magnification) as wart-
like processes; claw brownish.

Abdomen narrows slightly posteriorly, laterally with sparse short
light-colored hairs. Locomotory ampullae better developed on abdomi-
nal segments I to VI and less so on segment VII, matte, with minute
reticulate-squamiform sculpture, medially divided by common longitudi-
nal groove, laterally with coarse longitudinal rugose streaks. Body length
of mature larva 5.0 to 8.0 mm.

Pupa (Figure 109): Characterized by widely separated, minute paired
spinules on abdominal tergites. Body comparatively flat. Head short,
transversely convex between antennae, with median longitudinal suture,
without setae, hemispherically rounded on occiput. Antennae pressed to
sides, bent ventrad posterior to midfemora.

184 Pronotum uniformly convex, transverse, rounded laterally, narrows
gradually posteriorly, more steeply anteriorly, glabrous, without setae.
Mesonotum convex, with broadly rounded scutellum posteriorly. Meta-
notum lustrous, not shorter than wide, with median longitudinal groove,
glabrous, without spinules. Hind femora distinctly clavate, markedly
thickened in second half, with apices extending almost beyond abdominal
tergite IV, closely pressed to sides.

248

183

Figure 109. Pupa of Phymatodes ussuricus Plav.

Abdomen moderately elongate, widens somewhat in region of seg-
ment IV, narrows significantly posteriorly. Abdominal tergites uniformly
convex, with median longitudinal groove, with one barely visible para-
medial spinule, flat laterally, not produced. Tergite VII in posterior half
with pair of widely separated, more distinct, much larger spinules. Tergite
VIII elongate, rounded posteriorly. Valvifers of female hemispherical,
somewhat separated. Body length 5.0 mm, width of adomen 1.1 mm.

Material: Collected in Ussuri-Primor’e region (Komarovka, Suvo-
rovka, Artemovka Rivers, environs of Vladivostok, and so forth). Adult
insects 13, larvae 41, pupa — one female, larval exuviae with beetles from
cells four.

Distribution: Ussuri-Primor’e region. We found it in the region of
Komarovka and Suvorovka Rivers, along Kaban’ spring near Ussuriisk,
and in “Kedrovaya Pad” sanctuary.

Biology: Found in broad-leaved forests on Amur grapevine {Vitis
amurensis). Beetles fly from last few days of June up to mid-August. En
masse flight in middle 10 days of July. Beetles lead cryptic mode of life.

249

not seen on flowers. Found on grapevine creepers, mate there. Female
usually lays eggs singly under intact bark strips. Colonizes only thin
shoots 4.0 to 8.0 mm in diameter. Larvae hatch from eggs 18 to 20 days
after oviposition. For example, larvae began hatching from eggs laid July
3 through 6 from the 24th of that month. In another instance larvae
began hatching from eggs laid on a creeper on July 7 to 9 from July 29th
and immediately commenced boring wood.

Larvae live in wood, make galleries along shoots initially in upper
layer directly under bark, later along pith, and plug them with fine frass.
They then hibernate in wood. In spring of the following year they con-
tinue to make galleries and in June make a cell along the shoot in upper
layer of wood or in pith. Thin layer of wood left between cell and bark.
Length of cell 6.0 to 8.0 mm, width 1.8 to 2.0 mm. Larva passes through
preparatory phase in cell before pupation. It evacuates intestinal con-
tents and loses considerable weight. For example, in the laboratory four
larvae lost 23.6 to 28.8% of their weight during this period.

Pupation of larvae is generally completed in June. Pupae found up
to early July. Young beetles emerge from pupae in second half of June
and early July. They nibble an oval opening (1.5 mm X 2.0 mm to 2.0
mm X 2.5 mm) on shoot surface and emerge from pupal cell through
it. Complete life cycle from egg to adult requires one year.

Population density on grapevine creeper ^comparatively high. Thus
in a creeper section 52 cm long and 5.0 to 7.0 mm in diameter eight
insects were found; in another section 90 cm long and 4.5 to 7.0 mm in
diameter 32 insects (seven adults, 23 larvae, and two pupae); in a third
185 section 1.0 m long and 4.5 to 5.0 mm in diameter 10 insects, i.e., over
20 insects of this species were found on the average for every meter of
creeper. Teratoclytus plavilstshikovi Zaitz. sometimes colonizes together
with this species but usually lives on much thicker sections of the
creeper.

4. Phymatodes vandykei Gress.

Gressit, 1960, Kontyu, 9, 4, 172-173; Kojima and Hayashi, 1969,
Insect Life in Japan, vol. 1, p. 73.

Adult (Figure 110): Very similar to Phymatodes ussuricus Plav. and
almost indistinguishable from it. Only very minor differences in structure
of eyes and other features evident. Head retracted into prothorax up to
antennae, flat between them, without distinct tubercular elevation, with
narrow streaklike median longitudinal suture, with sparse minute puncta-
tion, sparse light-colored hairs. Eyes broadly emarginate, upper lobes
0.25 size of lower one, covered for most part by pronotum, septum be-
tween ocular lobes narrov/, with one row of facets. Antennae thicken
toward apex, extend beyond middle of elytra; 11th segment elongate, coni-

250

cally (female) or more acutely produced (male) at apex, with sparse hairs
at base, and denser adherent hairs toward apex, with long erect hairs on
first five segments.

Pronotum convex, markedly narrower at base than at apex, rounded
laterally, with broad shallow constriction in anterior third, more distinct-
ly sloped constriction near posterior margin, convex (not flat) on disk;
with uniform, fairly dense punctation and rusty hairs. Scutellum flat,
narrows posteriorly, narrowly or broadly rounded apically. Elytra with
parallel sides, posterior to scutellum along suture somewhat impressed,
with moderate punctation (in anterior half much larger, in posterior
comparatively small but entirely distinct), with short semiadherent hairs.

Body dark brown with rusty tinge, pronotum much darker. Elytra
in anterior half rusty-brown or chestnut, much darker in posterior half.
Body length 4.5 to 5.0 mm.

Egg: White, elongate, broadly rounded at one pole, narrows gradually
toward the other, appears pointed. Chorion smooth, hyaline, semitrans-
parent. Length 0.8 mm, width 0.35 mm.

186

Figure 110. Phymatodes vandykei Gress.

25l

Larva (Figure 111): Similar to larva of Phymatodes ussuricus Plav.
Differs in absence of longitudinal groove on dorsal locomotory ampullae,
and very thick inner maxillary lobes broadly rounded apically. Head
short, half of it retracted into prothorax. Epistoma smooth, uniform on
anterior margin, 'with lustrous brownish-rust border contrasting sharply,
indistinctly bound laterally (frontal sutures not visible), with faint median
longitudinal suture. Hypostomal sclerites smooth, lustrous, somewhat pro-
duced forward on inner angles, rounded on outer anterior angles. Gula
flat, markedly widens basally and slightly apically, laterally in anterior
half appears emraginate. Parietals in anterior half with stray long seta-
ceous hairs. Antennae long, with long thick 1st segment. Clypeus elongate,
trapezoid, semitransparent, Labrum oval, gently rounded on anterior
margin, with sparse short setae, and together with clypeus covers only
half of mandibular joint. Mandibles thick, broadly rounded and black
apically, reddish-rust at base. Inner lobes of maxillae thick, broadly
rounded apically, with sparse light-colored setae. Maxillary palps conical,
comparatively short, not longer than inner lobes.

186

Figure 111. Larva of Phymatodes vandykei Gress.
a — head and pronotum; b— -abdominal tergite IV with dorsal locomotory

ampulla.

252

186 Pronotum laterally rounded, narrows more anteriorly, on disk with
short and laterally much denser longer hairs, in anterior half with trans-
verse yellowish band divided by three white longitudinal interruptions
into four spots; of these, middle ones more distinct than lateral. Pronotal
scutum short, convex, white, lustrous, on anterior margin slightly pro-
duced medially, with barely visible thin longitudinal striation, bound
laterally by well- developed longitudinal grooves, basally more convex,
matte (this section covering not less than 0.33 of scutum). Prothoracic
presternum laterally with dense long, on disk sparse short light-colored
hairs; eusternum in form of pair of widely separated, lustrous, glabrous
round spots. Thoracic legs lacking; only in mature larvae present as slight
warts. Base (supporting part or sternellum) of pro-, meso-, and meta-
sterna glabrous, matte.

Abdomen elongate, narrows from thorax to middle, in second half
with parallel sides, quite often obtuse at posterior end. Abdominal seg-
ments laterally with sparse light-colored hairs, with insignificant flange
in anterior half or without it. Locomotory ampullae on abdominal seg-
ments I to VII slightly convex, transversely elongate, with minute reti-
culate-squamiform sculpture, matte, less developed on sternite VII. Body
length of mature larva 7.0 to 8.0 mm, width of head 1.0 mm.

Pupa (Figure 112): Differs from pupa of Phymatodes ussuricus Plav.
in more developed spinules on abdominal tergite VII. Body elongate.
Head short, narrows slightly in front of antennae. Frons broad, flat,
around antennal bases on inner side transversely slightly convex, with
median longitudinal suture. Vertex posterior to antennae slightly im-
pressed. Occiput hemispherically rounded, glabrous, lustrous. Antennae
short, pressed to sides, with apices insignificantly bent ventrad.

Pronotum not longer than width in middle, almost transverse, round-
ed laterally, convex on disk, with median longitudinal, transversely
striate band in anterior half. Mesonotum slightly convex, broadly round-
ed posteriorly, not produced in region of scutellum. Metanotum broad,
distinctly transverse, insignificantly convex, with slight median groove.
Femora in second half highly thickened, dilated, clavate. Hind femora
just reach posterior margin of abdominal tergite IV.

187 Abdomen elongate, with almost parallel sides in anterior half, broad-
ens very slightly in region of segments IV and V, narrows gradually poste-
riorly. Abdominal tergites highly convex, even, with one barely visible
paramedial spinule each directed backward. On some (especially anterior)
tergites, these spinules are altogether imperceptible. Tergite VII broad,
transverse, broadly rounded posteriorly, in posterior half with pair of
transversely separated, much larger, entirely visible spinules directed for-
ward. Tip of abdomen rounded, glabrous, without setae or spinules.
Body length 5,1 mm, width of abdomen l.I mm.

253

Figure 112. Pupa of Phymatodes vandykei Gress.

Material: Collected on Kunashir Island. Adult insects five, larvae
six, pupa—one male, exuviae with beetles from cells four.

Distribution: Islands of Japan and southern Kuril’ (Kunashir Island),
within viable zone of grapevine (Vitis).

Biology: Inhabits broaddeaved and mixed forests. Beetles fly from
second half of July. They do not require supplementary feeding and mate
immediately on emergence from pupal cells. Female oviposits under
intact bark strips of grapevine shoots. Colonizes mainly thin shoots up
to 1 .0 cm in diameter. Eggs generally laid singly.

In hatching larvae rupture chorion, emerge, and bore shoot. Their
longitudinal galleries under bark are deeply impressed in wood and
densely plugged with fine frass of bark and wood. In June mature larva
makes cell under bark in upper layer of tree along shoot and pupates
in it. Length of galleries 9.0 to 14.5 cm, width around cell up to 3.5
188 mm. Length of pupal cell 6.0 to 7.0 mm, width 2.5 to 3.0 mm. Some-
times larva bores wood before pupation, makes cell in pith of shoot,

254

turns head tovi^ard inlet, and then pupates. Pupation commences end of
June. Pupae found up to mid-July.

Emergence of young beetles from cells was observed in last 10 days
of July near Alekhino village on Kunashir Island. Beetles nibble oval
opening (1.0 mm x 1.2 mm) on shoot surface and emerge. By this time
their gonads are fully developed. Ovaries of a female just emerging from
cell contained 22 mature eggs. Weight of beetles immediately after emer-
gence from shoots 3,0 to 4.5 mg (male) to 6.0 mg (female).

A given shoot is colonized repeatedly. Three beetles were found in
cells on a shoot 19 cm long and 0.7 cm in diameter. Phymatodes maaki
(Kr.) inhabits grapevine shoots together with this species.

5. Phymatodes abietinus Plav. and Lur.

Plavilstshikov [Plavil’shchikov] and Lurie, 1960, ByuU. Mask. 0-va
Ispyt. Prirody, Otd. Biol, 65, 4, 124-125.

Adult (Figure 113): Characterized by short slender antennae, poorly
developed smooth shields on pronotum, and markedly elongate elytra
with parallel sides. Body minute, comparatively narrow. Head insigni-
ficantly retracted into prothorax, with uneven punctation, short smooth
median suture between antennae, and sparse erect hairs. Eyes indistinctly
faceted, on inner side slightly and angularly emarginate. Antennae slender,
comparatively short, with apices extending beyond 0.66 (male) or 0.50
(female) length of elytra. Third antennal segment only 2.0 times longer
than 2nd, notably lognger than 5th; latter equal to 4th.

Pronotum not longer than width in middle, rounded laterally, with
distinct groove near base and poorly developed transverse groove near
apex, moderately convex on disk, with uneven punctation, sparse light
brown hairs not forming compact cover, on disk with smooth distinct
or barely visible shields, two anterolateral to middle and one, rarely well
developed, on hind clivus. Scutellum longitudinal, rounded posteriorly,
with adherent hairs.

Elytra with parallel sides, markedly elongate, 3,0 times longer than
wide at humeri, with faint impression around humeri at base, individually
rounded apically, with dense punctation forming transverse wrinkles, and
189 short light-colored semiadherent hairs. Femoral clava comparatively long,
moderately thickened. Hind tibiae straight in male and female. First
segment of hind tarsi shorter than two successive together. Body ventrally
with sparse hairs and lustrous sheen.

Entire body, antennae, and elytra dark brown, legs much lighter
colored, rusty, sometimes antennae rusty. Body length 4.5 to 6.0 mm.

Egg: White, oval, obtusely rounded at poles. Chorion smooth, lus-
trous, hyaline. Length 0.8 mm, width 0.3 mm.

255

Larva (Figure 114): Characterized by uniform distribution of hairs
laterally and on disk of pronotum, faint yellowish spots on pronotum,
and very dense longitudinal striation on pronotal scutum. Head short,
markedly retracted into prothorax, narrowly rounded anteriorly. Epi-
stoma on anterior margin with broad lustrous brownish border. Median
longitudinal suture and frontal sutures not visible. Hypo stoma narrows
anteriorly, laterally with straight sutures and pointed inner angles of
sclerites widely separated by gula. Parietals in anterior half with stray
long piliform setae. Antennae with three segments, comparatively thick
at base. Clypeus small, basally flattened and white. Labrum oval, convex,
hyaline, in anterior half with sparse short setae. Mandibles black, basally
190 thick, broadly rounded apically, with angularly produced upper margin,
smooth outwardly, inner side broadly emarginate as though hollow.
Inner lobes of maxillae broadly rounded apically, lustrous, with sparse
short setae. Maxillary palps thick, long, with four segments.

Pronotum 2.0 times wider than long, somewhat narrowly rounded
anteriorly, with faint wide yellowish spots in anterior third (almost not
visible in some specimens), on disk with sparse short, and laterally much
longer fine hairs. Pronotal scutum insignificantly convex, white, with fine
dense longitudinal striation (distance between streaks less than their

256

189

Figure 114. Larva of Phymatodes abietinus Plav. andLur.
a — head and pronotum; b— abdominal tergite IV with dorsal locomotory

ampulla.

width), bound laterally by short longitudinal grooves, at base with
minute reticulate-squamiform sculpture. Alar lobes outwardly lustrous
lateral to grooves, with sparse hairs. Thoracic legs poorly developed,
barely perceptible, wartlike.

Abdomen moderately elongate, narrows somewhat posteriorly. Ab-
dominal segments I and VII laterally in anterior third with narrow deep
flange, behind which long fine hairs occur. Locomotory ampullae deve-
loped on abdominal segments I to VII, with fine reticulate-squamiform
sculpture, matte, without longitudinal streaks. Body length of mature
larvae 6.5 to 7.0 mm, width of head 1.1 mm.

Pupa (Figure 115): Differs from other species of this genus in absence
of spinules on dorsal side of body. Head narrows anteriorly from eyes,
notably impressed between antennae, on occiput spherically rounded,
hyaline, glabrous, without setae. Antennae slender, bent ventrad, loop-
like in second half.

Pronotum slightly elongate, in posterior third broadens angularly,
narrows gently anteriorly, flat on disk (female) or notably convex (male).

257

189

Figure 115. Pupa of Phymatodes abietims Plav. and Lur.

hyaline, with lustrous sheen. Mesonotum convex, with broadly rounded
(female) or slightly produced (male) scutellum posteriorly. Metanotum
transverse gently rounded on angles, slightly convex, with median longi-
tudinal groove in anterior half. Hind femora in second half moderately
thickened, with spices extending beyond base of abdominal tergite IV.

Abdomen elongate, narrows insignificantly anteriorly, not obtuse at
posterior end, rather somewhat pointed. Abdominal tergites convex,
rounded laterally, without setae or spinules. Valvifers of female large,
adjacent, broadly and uniformly rounded apically. Body length 5.5 to
6.0 mm, width of abdomen 1.3 mm.

Material: Collected in Salair fir forests. Adult insects eight, larvae
12, pupae six, larval exuviae with beetles from cells six.

Distribution: Southern regions of western Siberia. Found in forests
of Novosibirsk and Kemerov regions.

258

Biology: Inhabits fir forests of different densities. Beetles fly from
first half of July. Female oviposits on thin drying fir shoots {Abies sibi-
rica) 0.5 to 1.7 cm in diameter in lower tier of crown. Larvae hatch two
weeks later from eggs. Under laboratory conditions, at 20°C, larvae
began emerging from eggs 12 to 15 days after oviposition. Larvae begin
to hatch from early August.

In hatching larvae rupture chorion, bore bark, make gallery under
bark along shoot in upper layer of wood, and plug it with fine frass.
Length of gallery under bark up to 12 cm and width 2.0 mm. Before
second hibernation larvae bore wood, leaving an inlet 2.5 cm in width
on surface. The following spring mature larva makes pupal cell along
shoot and pupates in it. Length of pupal cell 7.0 to 1 1.0 mm, width 2.5
to 3.0 mm.

191 Pupation commences end of May, early June, and ceases in last 10
days of June. Young beetles appear mainly in second half of June. Found
in wood up to middle 10 days of July. Young beetles nibble oval flight
opening (1.0 mm X 1.5 mm to 1.4 mm X 2.5 mm) on shoot surface and
abandon cell. Emergence of beetles from wood completed in first half of
July. Young beetles do not require additional feeding, mate immediately
after emergence from wood, and female later oviposits. Female notably
larger than male. For example, in a series of weighings males (eight) in
larval stage before pupation weighed 3.4 to 7.4 mg, in pupal stage. 2.9
to 6.5 mg, and adults 1 .8 to 4.2 mg; females (nine) correspondingly weigh-
ed: 5.8 to 11.8, 5.3 to 10.1, and 4.3 to 8.2 mg.

Population density sometimes comparatively high. On a shoot 21.5
cm long and 1.7 cm in diameter, 11 specimens were found including
seven pupae and four larvae.

6. Phymatodes mediofasciatus Pic

Pic, 1933, Mel. Exot., vol. 62, p. 29; Plavil’shchikov, 1940, Fauna
SSSR, 22, 2, 324-325; Cherepanov and Cherepanova, 1974, Usachi vino-
grada amurskogo, pp. 30-33.

Adult (Figure 116): Readily recognized by a single white transverse
band on black elytra. Head markedly retracted into prothorax, with
faint anteromedian longitudinal suture, slightly impressed between upper
ocular lobes, with dense minute punctation, and light-colored adherent
hairs. Eyes finely and distinctly faceted, with individual facets in septum
between lobes forming narrow band. Antennae barely reach, more often
do not reach elytral apex (male) or 11th segment extends beyond 0.75
length of elytra (female). Third antennal segment equal to 5th, insigni-
ficantly longer than 4th.

Pronotum convex, roundly enlarged in anterior half, narrows more
toward base, near posterior margin with narrow transverse groove, with

192

Figure 116. Phymatodes mediofasciatus Vic.

deep uniform punctation, distance between punctures smaller than punc-
tures, with thin compactly adherent gray hairs combed forward and
from sides toward middle, laterally with long erect setaceous hairs.
Scutellum broad, broadly, more rarely narrowly rounded posteriorly,
with dense punctation, and adherent hairs. Elytra moderately elongate
with parallel sides, at base from inner side of humeral tubercle uniformly
convex, individually or almost jointly rounded at apex, basally with
large, elsewhere with very small dense punctation, medially with brown-
ish hairs, on transverse band with white short hairs, at base and on
hind clivus with very long whitish hairs forming two compact light-
colored fields — one at base, the other at apex. Hind tibiae straight (fe-
male) or notably curved (male). First segment of hind tarsi distinctly
longer than two successive together.

Entire body, antennae, and legs black only tarsi sometimes rusty-
brown. Elytra black, medially with curved white transverse band. Body
length 4.0 to 7.0 mm.

260

Egg\ Initially transparent, hyaline, later turns whitish and elongates.
Chorion smooth. Length 1.2 mm, width 0.4 mm.

Larva (Figure 117): Differs from larvae of the subgenus Phyma-
todellus in large body, entirely visible thoracic legs, elongate longitudinal
spots laterally on pronotum, and structure of dorsal locomotory am-
pullae. Head narrowly rounded anteriorly, more than half of it retracted
into prothorax. Epistoma laterally almost not demarcated, fuses with
parietals, medially divided by longitudinal suture, well distinguished
192 in posterior half, with broad rusty- white border on anterior margin.
Frontal sutures not visible. Hypostomal sclerites barely convex, smooth,
with sharply manifest dark rusty border on anterior margin, sharply
produced inner and straight outer anterior angles. Parietals in anterior
third rusty-brown, with setaceous hairs forming indistinct transvere row.
Antennae comparatively long; 1st antennal segment very thick, elongate,
almost conical. Clypeus small, trapezoid, semitransparent. Labrum small,
elongate, narrowly rounded apically, with short setae. Mandibles massive,
broadly rounded apically, with cultrate margin, on outside basally with
transverse groove. Inner lobes of maxillae thin, elongate, narrowly round-
ed apically.

Pronotum transverse, roundly enlarged in posterior half, narrows ante-
riorly, slopes moderately toward head in anterior half, with four yellow

193 Figure 117. Larva of Phymatodes mediofasciatus Pic.

a— head and pronotum; b — abdominal tergite with
dorsal locomotory ampulla.

261

spots on anterior margin; of these spots, middle ones transversely elon-
gate, tetragonal, lateral ones longitudinal, extend almost up to base,
narrowly cuneiform posteriorly, with hairs only on anterior margin,
elsewhere glabrous, lustrous (in species of Phymatodellus lateral spots
with long uniform hairs do not extend toward base). Pronotal scutum
white, lustrous, uniformly convex, without matte transverse band at
base, with two emarginations on anterior margin, anteromedially elon-
gate, bound laterally by short longitudinal grooves. Proscutellar zone
with short hairs; pronotum laterally with dense long rusty hairs. Trans-
verse yellow spots in anterior half with long hairs, without sclerotized
ringlets at base. Prothoracic presternum on disk with shortened, later-
ally with long rusty hairs; eusternum glabrous. Thoracic legs minute,
poorly developed but entirely perceptible.

Abdomen narrows slightly posteriorly, with dense light-colored hairs
laterally. Abdominal segments in anterior third laterally with deep con-
striction. Dorsal locomotory ampullae markedly convex, developed on
abdominal tergites I to VII, coriaceous, without squamiform sculpture,
with minute furrows, divided by common median longitudinal groove,
laterally with longitudinal short groove, on anterior margin with trans-
193 verse groove, mediolaterally with faint oblique groove that extends
posteromedially. Ventral locomotory ampullae less convex, weakly deve-
loped on sternite VII. Length of mature larvae 11 to 12 mm, width of
head 1.3 mm.

Pupa (Figure 118): Body moderately elongate, not flat. Head narrows
steeply from antennae anteriorly, transversely convex between antennae,
uniformly rounded on occiput. Antennae in male comparatively long,
bent ventrad round midfemora, with apices turned forward; in female
notably shorter, extend beyond hind femora, with apices bent ventrad.

Pronotum rounded, bulges uniformly on disk, not longer than width
in middle, smooth, lustrous. Mesonotum medially convex, longitudinally
impressed laterally, not produced on posterior margin, angularly rounded.
Metanotum with broad median longitudinal groove, laterally with distinct
oblique dents, broadly rounded on posterior margin, almost semi-
circular.

Abdomen moderately elongate, broadens notably in region of seg-
ments III and IV, narrows gradually posteriorly. Abdominal tergites
convex, with common, narrow, median longitudinal groove, with stray
fairly prominent spinules that medially and in posterior half form two
transverse rows. Tergite VII broadly rounded posteriorly, in posterior
half with minute spinules directed dorsally and forming diffuse tuft. Tip
of abdomen rounded. Valvifers of female conically produced laterally,
adjacent. Body length 8.5 mm, width of abdomen 2.3 mm.

Material: Collected in Ussuri-Primor’e region. Adult insects 25,

262

Figure 118. Pupa of Phymatodes mediofasciatus Pic.

larvae 16, pupae — two males and females, larval exuviae with beetles
from cells 10.

194 Distribution: Ussuri-Primor’e region, northeast China. We found it
around Komarovka and Suvorovka Rivers, near Vladivostok and Ochin-
nikovo village.

Biology: Inhabits broad-leaved forests of various densities, but spora-
dically distributed in them. Vitally associated with grapevine {Vitis amu-
rensis). Beetles fly from end of May to July inclusive; they require no
supplementary feeding. After emerging they mate, and later female ovi-
posits. One female can lay up to 24 eggs in her lifetime. Eggs are spaced
singly under intact (tightly adhering) bark strip. Thin shoots of grape-
vine 4.0 to 8.0 mm in diameter are usually colonized. At 21.6 + 0.6°C,
larvae hatch from eggs in eight to 14 days, average 11.5 days, after
oviposition. Hatching of larvae commences in second half of June and
ends in August.

263

Newly hatched larvae bore wood and during this process chorion of
eggs falling under bark is filled with frass. Larvae make longitudinal
galleries in wood, usually along pith, and plug them with fine frass. Some-
times they damage almost the entire plant; only bark filled with frass
remains. Mature larva after hibernation, in spring with the onset of warm
weather, makes cell 10 to 15 mm long and 4.5 to 5.0 mm wide in wood,
quite often in pith of shoot. Thin layer of wood usually remains between
cell and bark.

Pupation of larvae observed in May and first half of June. Young
beetles appear in second half of May and in June. One week after emer-
gence they nibble an oval opening (2.7 mm X 3.5 mm) on surface of
grapevine shoots and abandon pupal cell through it. Weight of mature
larvae before pupation 16.9 to 35.3 mg, pupae up to 18.5 mg or slightly
more, and beetles before emerging from wood 7.5 to 29,0 mg. Genera-
tion completed in one year.

7. Phymatodes maaki (Kr.)

Kraatz, 1889, Deutsch. Entom. Z., vol. 22, p. 105 (C^z/Z/J/ww); PlaviP-
shchikov, 1940, Fauna SSSR. 22, 2, 329-331; Nakamura, 1958, Miscell.
Report of Hiwa Museum Nat. History, vol. 2, pp. 15-16; Cherepanov
and Cherepanova, 1974, Usachi vinograda amurskogo, pp. 22-26.

Adult (Figure 119): Differs from other species of this genus in very
thick clava of hind femora, characteristic color of elytra, and compara-
tively large body. Head retracted into prothorax almost up to eyes, faintly
convex between antennae, flat between upper ocular lobes, with dense
deep punctation, and erect hairs. Eyes distinctly faceted, broadly emar-
ginate, with narrow septum between lobes. Antennal apices reach beyond
posterior band (male) or anterior band (female) of elytra. Third antennal
segment longer than 4th, equal to 5th or somewhat longer.

Pronotum laterally rounded, uniformly convex on disk, basally with
narrow flange, width in middle not less than pronotal length, with uni-
form deep punctation, long erect brownish hairs, sometimes thin median
longitudinal carinae. Scutellum triangular, narrowly rounded posteriorly,
with barely visible smoothened punctation.

Elytra slightly elongate, with parallel sides, individually rounded api-
cally, insignificantly convex on disk, in anterior half (before anterior
band) with large deep punctation and black erect hairs forming very
dense tuft on posterior margin of scutellum, in posterior half with minute
smoothened, poorly distinguishable punctation and tender brownish (on
bands white) compactly adherent hairs. Clava of hind femora markedly
thickened, hind tibiae notably curved. First segment of hind tarsi not
195 longer than two successive together. Head and prothorax black. Meso-
and metathorax, abdomen, and scutellum red or reddish^rust, anterior

264

196

Figure 119. Phymatodes maaki (Kr.).

half of elytra rea, posterior half black, with two white transverse bands,
one (anterior) narrow and anterior to middle, the other (posterior) broad
and anterior to hind clivus. Antennae rusty-brown, darken more toward
apex. Base of femora, tibiae, and tarsi rusty, femoral clava black (f.
typica). Quite often anterior white band anteriorly bound by black band
(ab. anticemarginatus Plav.), sometimes abdom.en black (ab. obscuri-
ventris Plav.). Body length 6.0 to 10,0 mm.

Egg: White, semitransparent, narrow s more toward one pole, rounded
at ends. Chorion smooth, hyaline, without perceptible sculpture. Length
1.1 to 1.2 mm, width 0.5 to 0.6 mm.

Larva (Figure 120): Characterized by comparatively large body and
sclerotized ringlets at base of hairs along anterior margin of yellowash-
rust spots on pronotum. Head narrowly rounded anteriorly, markedly
retracted into prothorax. Epistoma in general color merges laterally with
parietals, its boundaries not perceptible (frontal sutures not distinct),
divided by median longitudinal suture, more distinct in posterior half,

265

anterior margin with brownish border, distinct median emargination,
and lateral to notch with translucent black, almost square spot. Hypo-
stoma slightly convex, with narrow brownish border on anterior margin,
laterally with straight sutures, divided into two triangular sclerites on
which inner angles broadly rounded and anterior outer ones straight or
even somewhat pointed. Gula laterally notched, rusty-brown in anterior
half, lighter colored posteriorly. Parietals in anterior half with stray
coarse hairs. Antennae long, project far beyond base of mandibles.
Clypeus minute, lustrous, semitransparent, almost rectangular. Labrum
rounded, convex, not narrower or slightly narrower than clypeus, in
second half with minute setae, narrows markedly toward base. Mandibles
massive, black, basally project somewhat and with rusty tinge, broadly
rounded apically. Inner masticatory lobes of maxillae narrowly rounded
apically, shorter than maxillary palps.

Pronotum transverse, narrowly rounded anteriorly, broadens in poste-
rior half, with four sharp light red-rust spots in anterior third forming
transverse band interrupted three times by white clearance, laterally and
on disk anterior to scutum with dense light rusty hairs. Rusty spots with
long hairs and dark red sclerotized ringlet at base of hairs. Pronotal
scutum white, glabrous, uniformly convex, without longitudinal striation,
with fine, barely visible furrows, bound laterally by short longitudinal
grooves. Prothoracic presternum laterally and on disk with dense rusty
hairs; eusternum in form of two round glabrous plates divided anteriorly
by hairy clearance. Base of pro-, meso-, and metasterna (basisternum s.
sternellum) glabrous, matte, laterally with sparse hairs. Thoracic legs
poorly developed, wartlike. Better developed in mature larvae before
pupation, not seen in young larvae.

197

Figure 120. Larva of Phymatode^ maaki (Kr.),
head and pronotum.

266

Abdomen narrows slightly posteriorly, laterally with thin, light-color-
ed, not very dense hairs. Dorsal locomotory ampullae convex, rugulose,
without squamiform sculpture, without perceptible longitudinal streaks,
divided by common median longitudinal groove, laterally with short
longitudinal folds, on anterior segments oblique deep folds. Ventral loco-
196 motory ampullae similar in structure. Body length of mature larva 10 to
15 mm, width of head 1.4 mm.

Pupa (Figure 121): Differs from pupae of other genera in well-develop-
ed spinules on abdominal tergites and produced scutellum on posterior
margin of mesonotum. Head short, somewhat bent under, flat in region
of frons, between antennae with slight transverse elevation smoothened
somewhat in female, on vertex and occiput rounded hemispherically.
Antennae pressed to sides, bent ventrad around midfemora, with apices
directed forward.

Pronotum convex, not longer than width in middle, rounded later-
ally, narrows identically anteriorly and posteriorly (female), sometimes
gradually anteriorly and more steeply posteriorly (male), smooth on

197

Figure 121. Pupa of Phymatodes maaki (Kr.).

267

disk, glabrous, without spinules or setae. Mesonotum lustrous, convex,
on posterior margin with produced and somewhat elevated scutellum.
Metanotum transverse, broadly rounded posteriorly, with median longi-
tudinal groove. Femora clavate; clava of hind femora in male markedly
dilated, thickens gradually in female.

Abdomen elongate, narrows slightly toward base, gradually but more
so toward tip. Abdominal tergites uniformly convex, with common medi-
an longitudinal groove, on posterior margin with short spinules forming
transverse row, and three or four paramedial spinules each. Tergite VII
broadly rounded posteriorly, in posterior half with spinules forming one
or two tufts or transverse row. Tip of abdomen rounded, glabrous. Body
length 8.5 to 13.0 mm, width of abdomen 2.0 to 2.8 mm.

Material: Collected in Ussuri-Primor’e region (Komarovka River,
environs of Partizansk, Vladivostok, Nakhodka, and Ovchinnikovo
village) and Kunashir Islands (Alekhino, Sernovodsk). Adult insects 60,
larvae 109, pupae—five males and 11 females, exuviae of larvae with
beetles from cells 28. Large series of larvae raised from eggs laid by beet-
les in the laboratory.

Distribution: Ussuri-Primor’e region, Sakhalin, Kunashir Island, north-
east China, Korea, and Japan within viable zone of grapevine (Vitis).

Biology: Inhabits broad-leaved forests. Ecologically associated with
197 grapevine {Vitis). Found more often in thinned forests. Beetles fly end
of May, early June, with maximum numbers occurring from middle 10
days of June through first 10 days of July. Lead a cryptic mode of life;
not seen on flowers. During flight period seen more often on drying
grapevine trunks where they mate. Female lays eggs singly there under
intact bark strips. One female can lay up to 24 eggs. Colonizes shoots
of medium thickness, 1.2 to 5.0 cm in diameter. Colonizing commences
from apex and extends to base. Thick shoots rarely colonized.

Hatching of larvae from eggs commences two to three weeks after
oviposition. For example, from eggs laid by beetles on June 10 to 18
first larvae appeared on July 3, and from eggs laid after June 28 larvae
hatched July 16 and 17. Period of larval hatching in nature extends
from last 10 days of June to end of July. Newly hatched larvae bore
grapevine shoot, make longitudinal gallery under bark, deeply impressed
in wood, and plug it densely with fine frass. Length of gallery under
bark up to 15 cm, width 3.0 to 4.0 mm. Mature larva penetrates deeper
into wood, advances there along shoot, makes cell, later nibbles falcate
exit from it toward surface, leaving layer of wood usually not more
than 1.0 mm, more rarely up to 5.0 mm, and pupates in it. Length of
cell 12 to 15 mm, width 4.0 to 5.0 mm.

Pupation of larvae observed in May. Beetles emerge from pupae
three weeks later. Young beetles nibble oval-elongate openings

268

(1.5 mm X 2.0 mm to 2.0 mm X 3.0 mm) on bark surface along shoot,
and abandon pupal cell through them. Emergence of beetles from wood
completed mid-June. For example, on Kovarovka River in 1971 almost
90% of the population had emerged from wood by June 8. Beetles
emerge with developed gonads and hence are capable of reproduction
without supplementary feeding. Weight of mature larvae before pupation
28.6 to 79.8 mg, pupae 16.0 to 55.6 mg, and young beetles in wood be-
fore emerging from cells 9.6 to 42.5 mg.

Teratoclytus plavilstshikovi Zaitz. SindBrachyclytus singularis Kr. quite
often inhabit the same shoots together with this species. For example, we
collected from one Amur grapevine shoot 40 cm long and 4.0 to 5.0 cm
in diameter, Phymatodes maaki (Kr.) (21), T. plavilstshikovi Zaitz. (1),
and B. singularis Kr. (2). This species also develops on grapevine {Vitis
cognetiae) in Japan (Nakamura, 1958).

198 8. Phymatodes alni (L.)

Linnaeus, 1767, Fauna Suecica, p. 639 (Leptura); Plavil’shchikov,
1940, Fauna SSSR, 22, 2, 325-326; Duffy, 1953, Monograph Immat.
Stages of British Import. Tinib. Beetles,^^.l\9-220(Poecilium)\T>QmQ\i,
1966, Tierwelt Deutschlands, 52, 2, 73-74.

Adult (Figure 122): Similar to Phymatodes maaki (Kr.). Differs in
broad anterior transverse band on elytra, small size, and other features.
Body short, comparatively broad. Flead retracted into prothorax almost
up to antennae, with minute punctation and minute, barely visible hairs,
somewhat convex around base of antennae, with median longitudinal
suture. Eyes distinctly faceted, broadly and deeply emarginate. Antennae
somewhat thickened toward apex, reach hind clivus of elytra (male) or
extend beyond 0.50 length of elytra (female); 3rd antennal segment
equal to 5th, notably longer than 4th; 11th segment thick, narrows
steeply to pointed apex.

Pronotum laterally rounded, quite convex on disk, with narrow flange
at base, dense minute punctation, without smooth shields, with short
clavate erect hairs. Scutellum longitudinal, with almost parallel sides,
gently rounded posteriorly. Elytra 2.7 times longer than total width at
humeri, posterior to humeri somewhat compressed laterally, with pro-
jecting humeral tubercles and inward to them slight impression, indivi-
dually rounded at apex, with rather sparse punctation, more distinct at
base, with short, clavate, faint erect hairs, on disk posterior to scutellum

199 sometimes with long black hairs forming tuft. Hind femora thicken
gradually, clavate at apex; hind tibiae straight, not curved.

Body black, antennae reddish-rust, legs brownish-rust, elytra basally
light rust, elsewhere blackish-brown, with two white transverse bands,
one anterior to middle directed anterolaterally from suture, second

269

198

Figure 122. Phymatodes aini (L.).

anterior to hind clivus curves posterolaterally from suture (f. typica).
Sometimes elytra reddish-rust at base and apex, black between white
bands (ab. apicale Pic) or entirely black, only with two white bands
(ab. infuscatus Chevr.). Body length 3.5 to 7.0 mm.

Egg\ White, hyaline, somewhat matte, rounded at poles, broadens
anteriorly, narrows posteriorly. Chorion with minute faint sculpture.
Length 0.8 mm, width 0.4 mm.

Larva (Figure 123): Very similar to larva of P. ermoJenkoi Tsher.
Differs in denser hairy cover on pronotal disk, absence of hairy septum
on prothoracic eusternum, and other minor features. Head narrowly
rounded anteriorly, markedly retracted into prothorax. Epistoma con-
vex, with broad dark brown border on anterior margin, broadly emargi-
nate there, around anterior angles with narrow secondary notch, longi-
tudinal (median) suture and frontal sutures not visible. Hypostoma flat

270

Figure 123. Larva of Phymatodes alni (L.).
a— head and pronotum; b— abdominal tergite IV
with dorsal locomotory ampulla.

or slightly convex, on anterior margin with narrow rusty-brown border;
hypostomal sclerites with sharply produced anterior inner angles and
steeply rounded posterior inner margin. Gula between sclerites of hypo-
stoma whitish, without rusty border on anterior margin, 2.0 times longer
than width at apex. Parietals medially with stray or several setaceous hairs,
on anterior margin lateral to antennal sockets with broad rusty-brown
200 border. Antennae long, apices extend beyond anterior margin of cepha-
lic capsule. Posterior to antennae transversely elongate, unevenly pig-
mented, sometimes barely visible ocellus present. Clypeus short, flattened,
almost striplike, semitransparent, hyaline. Labrum semitransparent, mi-
nute, oval, with sparse short light-colored setae. Mandibles with trans-
verse groove on outer side around base, gently rounded apically and
there with sharp cultrate edge, black, reddish-brown basally. Labial sub-
mentum somewhat longitudinal, slightly convex in middle. Inner lobes
of maxillae whitish, lustrous, rounded apically, with stray short setae.
Maxillary palps narrow markedly apically, brownish, notably longer
than inner lobes of maxillae.

Pronotum slopes toward head, rounded on anterior margin, broadly
rounded in posterior third, in anterior half closer to anterior margin

271

with four sharp, broad, rusty, tetragonal spots forming transverse band,
separated medially and laterally by longitudinal whitish clearances; ante-
rior to scutum and in zone of rusty spots with dense, comparatively long,
light rust hairs basally with small, slightly projecting, sclerotized reddish-
brown ringlets (in P. ermolenkoi Tsher. these hairs are sparse and basal
ringlet almost imperceptible). Pronotal scutum white and convex, on
anterior margin angularly produced medially, slopes toward anterior
angles, with minute but sharp longitudinal striation, laterally with short
shallow longitudinal grooves, medially with long narrow longitudinal
groove. Prothoracic presternum convex, with moderately dense but not
very long hairs; eusternum glabrous at base, lustrous, not demarcated
by groove from presternum, anterior margin medially without hairy
clearance or barely perceptible one. Thoracic legs lacking or seen as
faint wartlike projections.

Abdomen laterally with fairly dense light-colored hairs. Dorsal loco-
motory ampullae insignificantly convex, transversely oval, with minute
silvery squamiform sculpture, laterally with faint longitudinal grooved
dents or without them. Ventral locomotory ampullae similar in struc-
ture. Abdominal tergite IX very small, short, considerably shorter and
narrower than tergite VIII, apically with sparse long setaceous hairs.
Body length up to 8.0 mm, width of head 1.1 mm.

Pupa (Figure 124): Well distinguished from that of P. ermolenkoi
Tsher. by very minute, barely perceptible spinules on abdominal tergites.
Body small, moderately elongate. Head short, on occiput hemispherically
rounded, lustrous, between antennae broad, slightly convex. Antennae
slender, pressed to sides, apices bent ventrad somewhat.

Pronotum broadens posterior to middle, rounded laterally, narrows
gradually anteriorly, steeply toward base, slightly convex on disk, almost
flattened. Mesonotum slightly convex, on posterior margin with triangu-
larly produced scutellum. Metanotum broad, insignificantly convex,
broadly rounded on posterior margin, without setae or spinules.

Abdomen broadens in region of segments III and IV, narrows very
slightly anteriorly but markedly posteriorly. Abdominal tergites convex,
with faint median longitudinal groove, with one or two specklike, barely
perceptible paramedial spinules or without them. Abdominal tergite
VII triangular, rounded posteriorly, bulges on disk, with four minute
spinules in posterior third forming transverse row. Apices of hind femora
barely extend beyond abdominal tergite IV. Body length 5.2 mm, width
of abdomen 1.5 mm.

201 Material. Collected in broad-leaved forests in the southern Urals.
Adult insect one, larvae three, pupa— one male, larval exuviae from
cells two. Collection of adult insects in Zoological Museum of Moscow
State University also studied.

199

Figure 124. Pupa of Phymatodes alni (L.).

Distribution: Almost all of Europe, east up to the Urals, north up
to Scandinavia, and south up to the Mediterranean Sea.

Biology: Inhabits deciduous forests and ecologically associated mainly
with oak. Beetles fly from May through June. Female oviposits on thin
drying shoots of viable as well as withering trees. Diameter of colonized
shoots 1.5 to 7.5 cm.

Newly hatched larva bores shoot, makes longitudinal straight or
meandering gallery under bark, impressed deeply in wood, and plugs
it densely with fine frass. Sometimes larva makes longitudinal gallery
under bark initially in one direction, then in another; gap between
galleries narrow, resembles septum. Mature larva bores deeper into wood
and makes cell there along shoot or sloping toward shoot surface. Some-
times cell slitlike. Length of cell 9.0 to 11.0 mm, width up to 3.0 mm.
Larva pupates in cell with head toward inlet.

Pupation observed in early spring. Pupa develops for about two
weeks, possibly more. In the laboratory, at 17.2 to 20.0°C (average
19.1 + 0.3°C) pupa developed in 12 days. Beetle developed from it

273

made an oval opening (1.5 mm X 2.5 mm) after one week on shoot
surface and emerged from wood through it. Weight of larvae before
pupation (eight insects) 8.0 to 16.0 mg, pupae 5.0 to 11.0 mg, and beet-
les 4.5 mg or more. According to our observations, this species coloniz-
es oak. According to data available in literature (Plavil’shchikov, 1940;
Demelt, 1966), also attacks elm, chestnut, ash, alder, maple, and dog
rose. However, found more often on oak. Phymatodes ermolenkoi
Tsher. found together with this species in oak forests in Ussuri-Primor’e
region.

9. Phymatodes ermolenkoi Tsher.

Cherepanov, 1980, Nov. i maloizv. vidy fanny Sibiri, pp. 88-89.

Adult (Figure 125): Similar to P. maaki (Kr.) in tuft of long black
setaceous hairs posterior to scutellum and shape of white transverse
bands on elytra. Differs in black antennae, elytra lustrous lac black on
hind clivus, and other features. Head short, between antennae with
faint longitudinal groove, with dense deep punctation, and dark brown
erect hairs. Eyes sharply and finely faceted, markedly emarginate, with
more convex lower lobes. Antennal apices extend beyond middle of
elytra, commencing from 5th segment more (male) or less (female) serrate,
with dentate produced outer angle, notably thicken toward apex, with
minute faint adherent hairs, and long (denser on 1st to 6th segments)
setae. First antennal segment elongate, apically thickened, slightly longer
than 3rd; 5th segment longer than 4th, equal to 3rd; 11th segment
slightly longer than 10th, apically pointed or narrowly rounded.

Pronotum slightly transverse, laterally rounded, with narrow flange
at base, posterior margin curved, dense large deep punctation, and erect
setaceous dark brown hairs; hind clivus sometimes with narrow smooth
longitudinal band. Scutellum flat, narrowly or broadly rounded posterior-
ly, with uneven punctation. Elytra with parallel sides, moderately con-
202 vex, with straight humeri, insignificantly projecting rounded humeral
tubercle, tapered outer angle at apex, and narrowly rounded inner
angle; in anterior third with large coarse punctation, middle third with
minute faint punctation, posterior third on hind clivus with barely per-
ceptible, smoothened punctation, with lac sheen; disk with two trans-
verse white bands (anterior one shortened, anterior to middle of elytra,
and curves forward; posterior one broader, not shortened or slightly so,
anterior to hind clivus), in anterior half with very long hairs, posterior
half with short hairs, on suture posterior to scutellum with dense long
black hairs forming distinct tuft. Body ventrally with long light brown
hairs. Abdominal sternites convex and lustrous, with sparse minute
punctation. Legs with dense erect setaceous hairs. Femora sharply cla-
vate, with short, markedly dilated, lustrous, finely and sparsely punc-

274

Figure 125. Phymatodes ermolenkoi Tsher,

tate clava, with very long, coarsely punctate shaft. Hind tibiae notably
shorter than femora, with dense coarse punctation. First segment of
hind tarsi longer than two successive together. Body and antennae black.
Legs black, shaft of femora red. Elytra black, in anterior third red, dark
red or dark brownish-red, more rarely entirely black. Body length 5.0 to
7.0 mm.

Differs from proximate species P. quadrimaculatusGxt^^,, dhinhuitd
203 in Japan, in very short clava of hind legs (clava shorter than shaft),
markedly smoothened punctation on hind clivus of elytra, and much
larger body size.

Egg: White, silvery, tapers gently toward one pole, steeply toward

275

the other, acute or narrowly rounded at ends. Chorion transparent,
lustrous. Length 1.0 mm, width 0.5 mm.

Larva (Figure 126): In shape of rust-colored spots on pronotum re-
sembles larva of P. maaki (Kr.). Differs in absence of sharply sclero-
tized ringlet at base of hairs covering rusty spots in anterior third of
pronotum. Head narrowly rounded anteriorly, markedly retracted into
prothorax. Epistoma slightly convex, lustrous, on anterior margin with
broad dark brown border, behind which stray setaceous hairs form
transverse row, anterior angles produced somewhat forward, with faint
light-colored median longitudinal groove, and laterally fused with parie-
tals. Frontal sutures not visible. Hypostoma slightly convex, on anterior
margin with indistinct, sometimes narrow rusty-brown border. Gula

Figure 126. Larva of Phymaiodes ermolenkoi Tsher.
a — head and pronotum; b — abdominal tergite with
dorsal locomotory ampulla; c— abdominal segments
VI to IX of I-instar larva.

276

between sclerites of hypostoma longer than apical width, broadens
basally, whitish, without brownish border on anterior margin. Parietal s on
anterior margin dorsal and ventral to antennae with broad rusty-brown
border that does not cover antennal sockets from behind; posteriorly
sockets with one transversely oval, unevenly pigmented, spotlike ocellus
204 each; medially parietals with stray long hairs. Antennae long; 1st seg-
ment almost as long as two successive segments together. Clypeus lus-
trous, semitransparent, broadly flattened basally. Labrum oval, whitish,
convex, with long light-colored setae. Mandibles black, dark red basally,
broadly rounded apically. Labial mentum somewhat transverse, convex,
with long setae laterally. Inner lobes of maxillae digitate, with sparse
short setae apically. Maxillary palps comparatively thin, shorter than
inner lobes.

Pronotum transverse, narrows anteriorly, broadly rounded on ante-
rior margin, in anterior third with four sharply manifested rusty or
rusty-yellow transverse spots forming transverse band interrupted medial-
ly and laterally by longitudinal white clearances, anterior to scutum with
rather dense hairs, in zone of rusty spots with sparse minute light-
colored hairs encircled basally by faint ringlet or without it. Pronotal
scutum silvery shagreen, with narrow median longitudinal groove and
deep straight lateral longitudinal grooves. Prothoracic presternum with
dense hairs, laterally sometimes long, on disk short; eusternum not
demarcated, merges with general surface of presternum, basally with
pair of lustrous glabrous plates separated anteriorly by broad hairy
clearance. Thoracic legs lacking, or not perceptible.

Abdomen thick, laterally with short, not very dense, light-colored
hairs. Dorsal locomotory ampullae slightly convex, transversely oval,
with silvery squamiform sculpture covering much of tergites. Ventral
locomotory ampullae broad, transversely elongate, with minute silvery
reticulate sculpture. In I-instar larvae segments VI and VII laterally
with one large sharp spinule each directed posterolaterally. After molt
these spinules disappear. Body length of mature larva up to 6.0 mm,
width of head 1.0 mm.

Pupa (Figure 127): Characterized by small body, well-developed
spinules on abdominal tergites, glabrous convex pronotum, and other
features. Head short, transversely convex between antennae, with narrow
faint median longitudinal suture, glabrous, without setae, on occiput
hemispherically rounded, lustrous. Antennae pressed to sides, with apices
turned dorsad (female) or slightly bent ventrad (male).

Pronotum roundly (female) or angularly (male) broadens medially,
and here notably wider than long, flattened on disk, medially with fine
transverse striation, narrows markedly anteriorly and posteriorly, with
rounded angles. Mesonotum medially with transverse striation, laterally

277

203 Figure 127. Pupa of Phymatodes ermolenkoi Tsher., female.

matte silver, posteriorly with produced, sometimes slightly elevated scu-
tellum. Metanotum broad, sometimes rounded, with median longitudi-
nal, barely visible, troughlike, transversely finely striate groove. Pro-,
meso-, and metanota glabrous, without setae.

Abdomen narrows insignificantly anteriorly and markedly poste-
riorly. Abdominal tergites convex, with common median longitudinal
groove, on posterior third of tergites with one (on I) or more often
three (on II to VI), more rarely two or four paramedial spinules form-
ing transverse row. Spinules in female much larger, in male slender.
Abdominal tergite VII narrows posteriorly and there narrowly or broad-
ly rounded, on posterior margin with four, more rarely one or two or
five comparatively large spinules that bend forward and form distinct
transverse or curved uneven row, with two small spinules each anterior
to this row forming additional transverse row. Tergite VIII moderately
205 elongate, trapezoid, narrows somewhat posteriorly and glabrous here,

278

rarely with one or two faint spinules. Valvifers of female hemi-
spherical, proximate, with apices slightly produced laterally. Apices of
femora markedly clavate, hind femora closely pressed to sides, with
apices extending beyond abdominal tergites IV and V. Body length 5.5
to 9.0 mm, width of abdomen up to 3.0 mm.

Material: Collected in Ussuri-Primor’e region, near village Khasan.
Adult insects (raised in laboratory) 21, larvae 30, pupae— five males and
five females, series of larvae raised from eggs laid by beetles in the
laboratory.

Distribution: Southeastern zone of Ussuri-Primor’e region.

Biology: Inhabits broad-leaved forests and ecologically associated
with oak. Beetles fly in first half of summer. Colonizes thin shoots of
undergrowth and physiologically weakened trees. Female lays eggs singly
in bark crevices. One female can lay up to 20 eggs in her lifetime.
Ovaries of one female dissected seven days after emergence contained
16 eggs. In the laboratory beetles lived for 31 to 38 days. Egg develop-
ment from moment of laying to hatching of larvae at 17.7°C extended
for 18 to 24 days, average 19 days (12 eggs under observation).

Newly hatched larvae bore bark, make longitudinal galleries, im-
pressed in wood and inside bark, and plug them with fine frass of wood
and bark particles. Galleries usually slitlike, radially elongate cross-
wise; larvae lie in them with one side toward bark, the other toward
pith. Mature larva makes slitlike pupal cell along shoot in which it
hibernates. Length of larval gallery under bark 9.0 cm or more. Length
of pupal cell 8.0 to 11.0 mm, width 3.5 to 4.0 mm. Cell impressed in
wood and inside bark.

Pupation of larvae commences early spring. In the laboratory, at
room temperature, pupae developed for 22 to 25 days. Newly formed
beetles nibble ellipsoid flight opening (2.5 mm X 1.5 mm to 3.0 mm X
1.8 mm) on bark surface and emerge through it from pupal cell. Records
of 33 insects showed: weight of larvae before pupation 8.0 to 27.0 mg,
pupae 7.5 to 23.5 mg, adults 7.0 to 23.0 mg. Some larvae before pre-
paration for pupation weighed about 37 mg. Females considerably
larger than males.

Colonizes oak shoots 1.8 to 3.0 cm in diameter. Population density
quite high. For example, on shoots with a total length of 89 cm and
diameter 1.8 to 2.4 cm, 29 insects were found: nine larvae, 12 pupae,
and eight adults.

SUPPLEMENT

206 A description of the morphology and biology of some species is given
below supplementing information in Usachi Severnoi Azii {Prionime —
Aseminae [Cerambycidae of Northern Asia (Prioninae — Aseminae)] releas-
ed in 1979.

III. Subfamily Leptiirinae
6. TRIBE STENOCORINI
2. Genus Rhamnusiwn Latr.

I . Rhamnusium gracilicorne Thery

Thery, 1894, Bull. Soc. Entom., p. 265; Cherepanov, 1979, Usachi
Severnoi Azii, pp. 85-87.

Adult (Figure 128): Two males raised from larvae collected in nature
belong to var. rufotestaceum Pic. Head, pronotum, and abdomen red;
eyes black, antennae red, matte only at apex from 5th segment; meso- and
metasterna and scutellum black; elytra red. Identical to nominal form in
rest of features.

Larva: See Usachi Severnoi Azii (Cerambycidae of Northern Asia)
(Cherepanov, 1979).

Pupa (Figure 129): Differs from pupae of all other genera of the tribe
Stenocorini in absence of spinules and long setae on abdominal tergites.
Rusty or light-colored setae faintly visible only under high magnification
against background of chitinous cover. Body of pupa moderately elon-
gate. Head short and bent under, around clypeus and at base of antennae
with very minute setae, frons between antennae transversely insignificant-
ly convex, vertex between upper lobes of eyes slightly impressed, occiput
broadly rounded, antennae short, their base recessed toward mandibles,
pressed to sides of body, with apices bent ventrad.

Pronotum narrows anteriorly, rounded on posterior angles, on ante-
rior and posterior margin slopes uniformly and there without transverse
groove and without flange, convex on disk, with stray thin rusty setae
seen only under high magnification. Mesonotum convex, on posterior
margin with thickly produced scutellum, laterally with very fine short
rusty setae forming elongate tuft on each side of scutellum. Metanotum

279

280

transverse, convex, in posterior half with minute rusty setae forming two
paramedial tufts (one tuft on each side).

Abdomen elongate, narrows slightly posteriorly. Abdominal tergites
I to VI convex, with median transverse groove, behind which very minute
207 setae, barely visible under high magnification, form broad transverse
band. Tergite VII broadly rounded posteriorly, not longer than width at
base, convex on disk, with minute rusty hairs, more numerous in pos-
terior half. Tergite VIII transverse, 2.0 times wider than long, broadly
rounded posteriorly, convex, with minute, on posterior margin much
longer rusty setae. Tip of abdomen rounded, obtuse (ventral view), later-
ally with broad, horseshoe-shaped, lustrous carina bearing stray light-
colored, very minute setae. Valvifers of female small, proximate, apically

Figure 128. Rhamnusium gracilicorne Thery.

281

Figure 129. Pupa of Rhamnusium gracilicorne Thery.

with laterally extended tubercle, with minute furrows, rust colored. Body
length 15 to 22 mm, width of abdomen 5.0 mm.

Material: Collected in the southern Urals (Chesnokovka). Adult
insects— “two males (raised from larvae in the laboratory), pupae — six
females.

Biology: Found in broad-leaved forests in the southern Urals. Larvae
found in wood at base of thick dead branches on thick- trunked viable
elm trees (Ulmus). Mature larva makes cell along shoot, nibbles exit to
shoot surface, plugs it with fibrous frass, and pupates with head toward
exit. Length of cell 25 to 35 mm, width 9.0 to 15.0 mm. In the labora-
tory, at 11.0 to 20.2°C (average 16.4 + 0. 5°C) pupae developed in 23
208 days. Young beetles push back frass, nibble flight opening (8.0 mm x 6.0

282

mm to 10.0 mm x 6.0 mm) on shoot surface and emerge from cell.
Weight of two pupae (male) 190.5 and 198.0 mg; weight of two beetles
after emerging from cell 120 to 138 mg.

4. Genus Stenocorus F.

2. Stenocorus meridianus (L.)

Linnaeus, 1758, Syst» Nat., 10th ed., 398 (Leptura); Cherepanov,
1979, Usachi Severnoi Azii, pp. 95-97.

Pupa (Figure 130): Differs from proximate species S/ewocort/j
sis Kr. in long setae on head around base of antennae laterally and
posterior to upper ocular lobes. Body stocky, slightly concave. Head
markedly bent under, impressed on vertex between upper ocular lobes,
transversely convex between antennae, with median longitudinal suture,
in region of frons and vertex often with transverse striation (with uni-

Figure 130. Pupa of Stenocorus meridianus (L.).

283

form transverse furrows), with long rusty setae forming one tuft each
around base of antennae on inner side and in postorbital zone, posterior
to upper ocular lobes (in Stenocorus amurensis Kr. tufts of setae absent
on head, at most stray short setae, not forming tufts, present). Antennae
pressed to sides, arcuate in second half, bent ventrad, with apices usually
adjoining 1 st segment of midtarsi.

Pronotum convex on disk, transversely rugose, with narrow median
longitudinal groove, narrows markedly anteriorly, with broad sharp flange
anteriorly, with two steep emarginations on posterior margin and elevat-
ed there in form of carina, on anterior elevated margin with not very
dense long setae, basally with dense rusty setae forming compact band
replicated angularly in middle, elsewhere on surface with stray scattered
setae. Mesonotum small, insignificantly convex, on posterior margin
with produced scutellum, laterally with dense rusty setae forming two
distinct tufts. Metanotum glabrous in anterior half, transversely finely
striate, with pair of tubercular, densely setaceous processes in posterior
half. Femora markedly bent dorsad, apically with setae forming collar or
spread out.

Abdomen narrows from anterior to posterior end. Abdominal tergites
I to IV with sharp tubercular paramedial elevations covered with dense
rusty setae. Tergite V with two faint elevations covered with setae; ter-
gite VI without tubercular elevation, in posterior half with rusty setae
forming small paramedial tuft. Abdominal tergite VII convex, narrows
slightly posteriorly, broadly rounded at posterior margin, in posterior
209 half with sparse setae. Tergite VIII almost semicircular, in posterior half
with long rusty setae forming two dense lateral tufts. Posterior margin
of tergite IX elongate, with short, slightly bifurcate, coriaceous urogom-
phus, in posterior half with long rusty hairs. Tip of abdomen (ventral
view) obtuse, bound laterally by carina covered with sparse long rusty
hairs. Valvifers of female almost triangular, angularly produced inward,
with lateral conical tubercle apically. Body length 1 8 to 25 mm, width of
abdomen up to 6.0 mm.

Material: From broad-leaved forests of the southern Urals. Raised
from larvae collected in nature (additionally); adult insects six, pupae —
two males and one female.

Biology: Larvae live under root bark of elm, oak, and other deciduous
woody species, make galleries impressed in wood. Mature larva nibbles
opening on root surface and emerges through it into soil. Usually makes
pupal cell alongside root, polishing inner walls by rotary movement, and
pupates. Length of cell 21 to 25 mm, width 12 to 15 mm. Pupation of
larvae May-June. Pupae develop for up to three weeks. At low tempera-
tures pupal development is retarded. For example, in the laboratory
larvae pupated by August 18, pupae formed in cells placed in refrigera-

284

tor at 1.0®C, then kept at 12.0 to 13.8°C (average 13.1+0.8°C). Beetles
emerged from these pupae on December 22 to 27, i.e., 126 to 131 days
after pupation of larvae, or 1 7 to 22 days after refrigeration at optimum
temperature (four pupae under observation). Weight records of five spe-
cimens (male) showed: larvae before pupation 300 to 424 mg (average
381.2), pupae 235 to 314 mg (average 291.6), and beetles before emerg-
ing from soil 187 to 244 mg (average 223.4), i.e., during metamorphosis
weight loss was 44.1%. In another instance (one female specimen) larval
weight before pupation 719 mg (100%), pupa developed from it 536 mg
(74.5%), and beetle emerging from latter 435 mg (60%). Beetles remain
in cell for one week, emerge on soil surface with developed gonads, and
commence reproduction.

IV. Subfamily Aseminae
10. Tribe ASEMINI
2. Genus Arhopalus Serv.

3. Arhopalus tristis (F.)

Fabricius, 1787, Mant. Ins., vol. 1, p. 154 (Callidium); Cherepanov,
1979, Usachi Severnoi Azii, pp. 425-427.

Larva (Figure 131): Characterized as follows. Parietals along frontal
suture with white glabrous plate, more often without it. Thoracic legs
glabrous on outer side or with stray setae, inner side at apex of femora
with long setae forming transverse row or collar [thoracic legs in Arho-
palus rusticus (L.) with dense setae on both inner and outer side of
femora forming broad band encircling femur in distal half]. Posterior
margin of abdominal tergite IX with pair of widely separated spinules
set on thick extended base. Distance between these spinules 2.0 times
diameter of their base [in Arhopalus rusticus (L.) spinules at apex of
210 abdominal tergite IX more proximate; distance between them not more
than diameter of their base].

I-instar larvae characterized by more protruding pleural tubercles.
Posterior margin of abdominal tergite IX with pair of faint tubercular
formations devoid of sclerotized spinules. Latter seen in Il-instar.

Pupa (Figures 132 and 133): Differs from pupa of proximate species
Arhopalus rusticus (L ) in arrangement of spinules on abdominal tergites
and other features. Head slightly narrower than pronotum, faintly convex
between antennae, with fine but faint transverse striation, rounded on
occiput, posterior to upper ocular lobes with short setae that form nar-
row transverse band there, on inner side of which and around clypeus

285

Figure 131. Larva of Arhopalus tristis (F.).
a — ^head and pronotum; b — abdominal tergite IX;
c— outer and inner side of leg.

with stray, barely perceptible setae. Antennae short, pressed to sides,
with apices bent ventrad; outer side of 1 st segment with one to three
minute spinules, remaining segments without spinules [antennae in Arho-
palus rusticus (L.) with innumerable large spinules on outer side of 1st
segment, remaining segments with minute spinules],

Pronotum slightly transverse, rounded laterally, slightly convex on
disk, with sharp acicular spinules laterally and in anterior third set on
211 thin protruding coriaceous base, medially with minute setae forming lon-
gitudinal band. Mesonotum transversely impressed, with produced scu-
tellum on posterior margin, transverse furrows, laterally with numerous

286

210

Figure 132. Pupa of Arhopalus tristis (F.), female.

minute or sparse stray setae. Metanotum broadly rounded posteriorly,
with faint median longitudinal groove, with transverse furrows or streaks;
minute setae laterally in anterior half form one fairly distinct tuft each.

Abdomen elongate, broadens insignificantly in region of segments II
to IV, narrows gradually posteriorly. Abdominal tergites in posterior
third convex and there with twin (large and minute) spinules forming
transverse band narrowly interrupted in middle, laterally in anterior half
with distinct dent, in anterior third with sparse, fine, barely perceptible
setae [abdominal tergites in Arhopalus rusticus (L.) with dense spinules
in entire posterior half, forming broad continuous, almost uninterrupted
band]. Posterior margin of tergite VII rounded, not longer than width at
base, convex on disk, glabrous in anterior half, with sparse, rather small

287

Figure 133. Antennae in pupae of (a) Arhopalus
tristis (F.) and (b) A. rusticus (L.).

spinules in posterior half. Tergite VIII transverse, convex, with stray
setae barely visible under high magnification. Tip of abdomen with pair
of thin urogomphi ending in sharp sclerotized spinule curved inward.
Abdominal sternites laterally with tubercular process covered with minute
spinules. Anterior third of sternites with minute sparse setae visible under
high magnification. Valvifers of female small, hemispherical, apically with
slightly projecting tubercle, separated by rather small gap. Body length
18 to 24 mm, width of abdomen 5.0 to 5.5 mm.

Material: Collected in pine forest near Klyuchi (Kulunda) village.
Larvae 16, pupae — two males and three females (two females raised from
eggs laid by beetles in the laboratory), larval exuviae from cells with
pupae three.

Biology: Larvae live in upper layer of wood, make longitudinal galle-
ries, and plug them densely with fine frass. Larva makes cell at end of
gallery and pupates in it. Pupation in June. One week after emergence
beetles nibble oval flight opening (6.0 mm X 4.0 mm) on bark surface
and abandon cell. Length of cell 3.0 to 5.5 cm, width 10 to 12 mm.
Generation completed in two years. Hibernation in larval stage. Under
laboratory conditions simulating natural environment, beetles emerged
in 1979 from eggs laid in 1977. However, development was delayed up
to three years in dry wood.

Weight indices of specimens differ little from those of Arhopalus rus-
ticus (L.). One larva before pupation weighed 303 mg (100%) and pupa
developed from it 279 mg (92%); corresponding values in another speci-
men (female): 468 mg (100%) and 363.5 mg (77.7%). Specimens develop-
ing in dry wood small. Arhopalus tristis (F.) inhabits pine {Pinus sylves-
tris). We did not find it on any other species. Lives in root zone of thick-
trunked trees.

INDEX OF LATIN NAMES OF
CERAMBYCID BEETLES*'

abietinus, Phymatodes 13, 118, 119,
120, 171, 172, 173, 174, 175, 188,
189

Acanthoderes 36
Acmaeops 65

acutivittis, Rhaphuna 11, 116
adspersus, Xylotrechus 13
aeneipenneab., Callidium 166
aeneum, Callidium 156, 157, 158,
161, 162,163
Aglaophis 1 1

alni, Phymatodes 12, 13, 172, 173,
174, 175, 198, 199
alpina, Rosalia 112
alpinus, Cerambyx 112
altaicus, Xylotrechus 10, 11, 12, 13,
14

Amarysius 3, 10
amurensis, Stenocorus 208
angusta, Pronocera 138
antilope, Xylotrechus 12, 129, 178
anticemterruptus, ab. Polyzonus 108
anticemarginatus ab. Phymatodes
195

apicale ab., Phymatodes 199
apicalis ab., Molorchus 55
arcuatus, Plagionotus 12, 13
argentatus, Cerambyx 97
Arhopalus 209, 210. 211, 212
arietis, Clytus 3, 11
arietoides, Clytus 3, 11, 13, 148, 171
arnoldi, Xylotrechus 10
Aromia 9, 11, 13, 87, 88, 89, 90, 91,
92, 93

arvicola, Xylotrechus 12
Aseminac 3, 200, 209

Asemini 209
Asemum 21 1
Asias 11
Atimia 151

auranticollis, Rhopalopus 124, 137

bajulus, Cerambyx 120
bajulus, Hylotrupes 120, 121
batesi, Rosalia 112
bifasciatus, Rhopaloscelis 38, 42
bifasciatus, Semanotus 143, 148, 149,
151

biinterruptus ab., Polyzonus 108
biinterruptus ab., Semanotus 144
binotatum ab., Stenygrinum 27
bivittatum, Callichroma 93
Brachyclytus 197
Brachyta 3

brevicollis, Callidium 138
brevicollis, Pronocera 13, 118, 138,
140, 141, 142, 153
brevicorne, Obrium 32, 36, 38, 39,
40, 42

brevipennis, Nathrius 3, 47, 48, 49,
50

brunneipennis ab., Molorchus 66

Caenoptera 51
Callichroma 93

Callichromini 7, 8, 9, 10, 11, 12, 15,
16, 18. 87. 107, 111, 112
Callidiini 7. 8, 10, 11, 12, 15, 16, 17,
18,111.116,120,213
Callidiopini 7, 8, 12, 14, 25
Callidium 28, 82, 116, 117, 118, 119,
120, 124, 125, 138, 152, 156, 157,

*Reproduced from the Russian original. Russian page numbers appear in
the left-hand margin of the text — General Editor.

^All taxa higher than genus are printed in bold face and synonyms in
italics. Page numbers in bold face denote description of taxa.

288

2S9

158, 159, 161, 162, 163, 164, 165,

167, 168. 169, 171, 194, 209
Callimellum 51

campestris, Trichoferus 19, 20, 22
camharinum, Cerambyx 32
cantharinum, Obrium 31,32, 34, 35,
36. 37

capricornis, Xylotrechus 3
castanipennis ab., Molorchus 66
Cerambyddae 3. 72, 97, 101, 102
Cerambydnae 7, 8, 11, 12, 13, 29
Cerambydni 7, 12, 14, 15, 22
Cerambyx 19, 22, 32, 89. 97, 107,
112, 120, 143, 157.158, 172, 175
Chelidonium 10, 11, 13, 87, 88, 89,
97. 98. 99. 101, 106
chinensis, Xylotrechus 3
Chloridolum 11, 13. 87. 88,89,93.
94, 95. 97

chlorizans, Callidium 156, 157, 158,
167,168, 169, 171

Chlorophorus 3. 10. 11. 12,13, 16,
87

christophi, Plagionotus 12, 13
cinereus, Cerambyx 19
cinnabarinum, Callidium 152
cinnabarinum, Oupyrrhidium 152,
154, 155, 156

clavipes, Acanthoderes 36
clavipes, Rhopalopus 9, 13, 118, 123,
124, 125, 126, 127, 129, 132
Clytini 7, 8. 9, 10, 11. 12, 15, 17. 18
Clytus 3, 10, 11. 16, 148, 171
coelestis, Rosalia 11, 112, 113, 116
collaris, Acmaeops 65
collaris, Pronocera 138
colobotheoides, Aglaophis 11
conjunctum ab., Stenygrinum 27
coriaceum, Callidium 157, 165, 167
cuneipennis, Xylotrechus 12, 13

daurica m., Pronocera 139
decempunctata, Rosalia 112
Deilus 11, 82, 83, 86. 87
detritus, Plagionotus 12, 13
Dilus 82

Dilusini 7, 8. 9. 12. 15, 16. 17. 81,
213

Disteniinae 3
divaricata, Sydonia 46
divisa ab., Pronocera 139

Epania 51

ebenina, Necydalis 133
Epiclytus 16

ermolenkoi, Phymatodes 9, 12, 13,
172, 173, 174, 175, 199, 200. 201,
202, 203
Evodinus 3
Exocentrus 78, 156

fasciatus, Cerambyx 107
fasciatus, Polyzonus 8, 10, 106, 107,
111

fasciatus ab., Semanotus 148, 149
fasciculatus, Pogonocherus, 58, 142,
148. 164

fenestratus, Stenhomalus 43
figuratus, Chlorophorus 3
floralis, Plagionotus 8, 10
fugax, Callidium 82
fugax, Deilus 11, 82, 83, 86, 87
funebris, Rosalia 112

gilvipes, Tetrops 65
gracile, Obrium 10, 31, 32, 36, 38,
39, 40. 41, 42

Gracilia 14, 15. 27. 28. 29. 30
gracilipes, Chlorophorus 11
gracilicorne, Rhamnusium 206, 207
gracilicorne, Tetropium 171
Gradliini 7, 12, 14, 15, 17, 27, 81,
213

grallator, Amarysius 3

heptapotamicus, Molorchus 52, 53,
54, 66. 69. 70. 71
herbsti, Chlorophorus 12
Hesperophanes 19

Hesperophanini 7, 9, 12, 14, 15, 17,
18,213

Hylolrupes 117, 118. 119, 120, 121
hypocrita, Clytus 3

incarinatus ab., Molorchus 55
incognitus, Molorchus 52, 53, 54, 75,
76, 77

infuscatus ab., Phymatodes 199
insubricum, Callidium 124
interrupta, Saperda 58, 148

japonicus, Mallambyx 22
junebris, Rosalia 112

290

kaehleri, Purpuricenus 13
kiesenvetteri, Molorchus 52, 53, 54,
64, 65, 66, 68

kobotokensis, Molorchus 11, 52, 53,
54, 58, 72, 73

kunashiricus ssp., Molorchus 72

lameeri, Rosalia 112
latefasciatus ab., Polyzonus 108
lateritia, Rosalia 112
latifasciatus ab., Semanotus 149
Leontium 9, 10, 11, 87, 88. 89. 101,
102, 103, 105. 106
Leptidea 46, 47
Leptideella 47
Leptura 198, 208
Lepturinae 3, 7, 11, 206
Linomius 51, 52

maaki, Phymatodes 118, 172, 173,
174, 175, 188, 194,196,197,198,
201, 203

Mallambyx 7, 14. 22. 23. 24
marginatus, Exocentrus 156
Mausaridaeus 111

mediofasciatus, Phymatodes 118,
172, 173, 174, 175, 191, 192. 193
meridianus, Stenocorus 208
Mesosa 129

minor, Molorchus 11, 52, 53, 54. 55,
57. 142, 148
minor, Necydalis 52
minuta, Callidium 28
minuta, Gracilia 3. 14, 28, 29, 30
mixtus, Xylotrechus 3
Molorchini 7, 8. 10, 12, 13, 14, 16,
17, 46. 51.81
Molorchinus 51

Molorchus 3. 10. 11. 51. 53. 54. 55.
57. 58, 59, 61, 62.63.64.65,66,
68, 69, 70. 71, 72. 73. 75. 76, 77,
78, 142, 148
Monochamus 148
morio, Necydalis 116, 133
moschata, Aromia 9, 11, 13, 89, 90,
91. 92, 93

moschatus, Cerambyx 89
myops, Mesosa 129

Nadezhdiana 9, 51, 78, 79
nadezhdae, Atimia 151

Nathriini 7. 8, 9. 12, 14. 16, 17. 46
Nathrius 3, 46. 47, 48, 49, 50
Neocerambyx 22
Necydalis 52, 53. 116, 133
nigricollis ab., Oupyrrhidium 153
nigritulus, Clytus 3

Obriini 7. 8, 9, 10, 11, 12, 13. 14. 16.
17,30, 48,81

Obrium 10. 30. 31. 32. 34,35.36,
37. 38, 39. 40,41, 42, 43
Obscuriventris ab., Phymatodes 195
orientalis ssp., Aromia 90, 92
Oupyrrhidium 117, 118, 119, 120,
151, 152, 154, 155, 156

Palaeocallidium 157, 166
pantherinus, Xylotrechus 13
Paraphymatodes 173
pavlovskii, Xylotrechus 3
perforata, Saperda 36
petasifer, Purpuricenus 3, 13
Phymatodellus 171, 172, 191, 192
Phymatodes 9, 12, 13, 16, 117, 118,

119, 120, 129. 171, 172. 173, 174,
175.176,177, 179, 180, 181, 182,
183, 185, 186, 187, 188, 189, 191,
192, 193, 194, 196, 197. 198, 199,
200, 201, 202, 203

Plagionotus 8, 10, 12, 13
plavilstshikovi, Teratoclytus 13, 185,
197

Plocaederus 22

Polyzonus 8, 10, 87, 88, 89, 106, 107,
111

Pogonocherus 58, 142, 148, 164
Poecilium 173, 198
porteri, Nathrius 47
posticeinterruptus ab., Polyzonus
108

praeusta, Tetrops 65
praeustus ab., Phymatodes 175
Prioninae 206

Pronocera 13, 116, 117, 118, 119,

120, 138, 139, 140, 141, 142. 152,
153

Pronocerodes 124
Prorrhopalopus 123, 124
Psebiini 14, 16, 46, 81
Pseudocallidium 111
Pterolophia 156

291

puellus ab., Hylo trapes 121
pulcher, Plagionotus 12, 13
Purpuricenus 3, 10, 13
pygmaeum, CalUdium 28

quadrilunatus ab., Semanotus 144
quadrimaculatus, Phymatodes 203
quadrinotatum, Stenygrinum 3, 25,
26

raddei, Mallambyx 14, 22, 23, 24
raddei, Neocerambyx 22
Rhamnusium 206, 207
Rhaphuma 7, 11, 116
Rhopalopus 8, 9, 10, 13, 16, 117,
118, 119, 123, 124, 125, 126, 127,

129, 130, 131, 132, 133, 134, 135,
136, 137, 142

Rhopaloscelis 38, 42
Rosalia 11. Ill, 112, 113, 116
Rosaliini 7, 8, 9, 12. 15, 16. 18, 111
rufescens ab., Molorchus 55
ruficollis, Rhopalopus 124, 137
rufilius, Xylotrechus 3
rufithorax ab., Rhopalopus 124, 130
rufotestaceum var., Rhamnusium
206

rusticus, Arhopalus 209, 210, 211
rusticus, Xylotrechus 10, 13

salessei ab., Callidium 159
saltuarius, Monochamus 148
Saperda 28, 36, 58, 148
sartor, Chlorophorus 3
scalaris, Saperda 36
scutifer ab., Hylotrupes 122
Semanotus 11, 13, 14, 117, 118, 119,
120, 142, 143, 144, 145, 146, 147,
148, 149, 150, 151, 167
semireductus ab., Semanotus 144
sexmaculatus, Chlorophorus 12, 13
sibiricus, Cerambyx 107
sieversi, Chloridolum 11, 13, 88, 94,
95, 97

signaticollis, Rhopalopus 124, 125,

130, 131, 133, 134, 136
sinensis, Semanotus 148
singularis, Brachydytus 13, 197
sincauster ssp., Semanotus 148, 149
speciosus, Rhopalopus 124, 125, 133,

134, 135

Stenaspini 7, 8, 10, 11, 12, 15, 17, 18
Stenhomalus 10, 30, 31, 32, 42,43.

44, 45, 46
Stenocorini 206
Stenocorus 208
Stenopterus 51
Stenygrinum 3, 25, 26
stierlini, Exocentrus 78
striatum, Asemum211
Sydonia 46

Teratoclytus 8, 13, 185, 197
testaceus, Cerambyx 172
testaceus, Phymatodes 118, 129, 171,

172, 173. 174, 175, 176, 177
Tetropium 171

Tetrops 65

theresae ab., Hylotrupes 121
Tomentaromia 89

transversefasciatus ab., Semanotus
144

Trichoferus 18, 19, 20, 22
tristis, Arhopalus 209, 210, 211, 212
tsherepanovae, Purpuricenus 13
tuvensis, Asias 11

ubsanurensis, Chlorophorus 3
umbellatarum, Molorchus 3, 52, 53,
54, 58, 59, 61, 62, 63
undatus, Cerambyx 143
undatus, Semanotus 11, 13, 14, 118,
143, 144, 145, 146, 147, 148, 149,
150

Upyrrhidium 151, 152

ussuricus, Phymatodes 13, 118, 172,

173, 174, 181, 185, 186
ussuriensis, Molorchus 52, 53, 54,

58. 59, 61,62, 64
ussuriensis, Pterolophia 156

vandykei, Phymatodes 172, 173,

174, 175, 185

variabilis ab., Phymatodes 175
varius, Chlorophorus 12, 87
villioni, Xylotrechus 3
villosa, Nadezhdiana 78, 79
violaceum, Callidium 156, 157, 158,
159, 162, 164
violaceum, Cerambyx 15
virescens ab., Callidium 159

292

viride, Leontium 9, 11, 87, 102, 103,
105, 106

vulcanus, Stenhomalus 10, 43, 44,
45, 46

Xyloclytus 3

Xylotrechus 7, 9, 10, 11, 12, 13, 14,
16, 129, 178

zaitzevi, Chelidonium 11, 13, 97, 98,
99, 101, 106

zemlinae, Phymatodes 172, 173, 174,
179, 180

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