Literature - Pityogenes chalcographus
Compiled by John A. Byers

Annila, E. 1977. Seasonal flight patterns of spruce bark beetles. Ann.
Entomol. Fenn. 43 (1). 31-35. ab The flight activity of bark beetles
(Hylurgops palliatus, H. glabratus, Hylastes cunicularius, Xylechinus
pilosus, Cryphalus saltuarius, Polygraphus poligraphus, P. punctifrons,
Dryocetes, Pityogenes chalcographus, Ips typographus and Xyleborus
dispar) breeding in spruce was studied in southern and northern Finland
in 1968-1973 by collecting beetles with window flight traps. The seasonal
flight patterns are presented. The catches of a number of insects
associated with bark beetles are given. LG EN.█FINN FOR RES INST,

Baader, E.J. 1989. Comparative studies on semiochemicals in
pityogenes-spp Coleoptera: Scolytidae and their potential usage in pest
management systems. J. Appl. Entomol. 107 (1). 1-31. ab The analysis of
volatiles released by six species of genus Pityogenes upon attack of host
material revealed pheromone components which comprise grandisol,
cis-verbenol, .gamma.-isogeraniol, trans-cyclohexlidenethanol, ipsdienol
and 2-methyl-3-buten-2-ol. Chalcogran was found in P. chalcographus and
P. quadridens, E, Z-2,4-methyldecadienoate only in P. chalcographus.
Strong synergism between chalcogran, the obvious attractive principle,
and E, Z-2,4-methyl-decadienoate, the landing signal, suggests these
compounds as obligatory components of the aggregation pheromone.
(S)-cis-verbenol and oleoresin of the host trees somewhat increase
attraction. The Ö produced 1-hexanol reduces response. Baits combining
synthetic chalcogran and E, Z-2,4-methyldecadienoate induce mass
aggregation of P. chalcographs on traps. Formulating the synthestics for
use in polyethylene dispensers, 2-methyl-3-butyn-2-ol appears superior
compared to other solvents presently used in commercial products; the
concentrations of chalcogran and E, Z-2,4-methyldecadienoate used in
baits should consider two problems: the strong response of the ostomid
predator Nemosoma elongatum to chalcogran and the spill over particularly
in younger stands of the aggregating pest population to trees near traps.
Conceptual models indicate mass trapping using pheromone traps superior
to the conventional "Fangreisig" method, economically as well as
17, D-7800 FREIBURG/BR

Benz, G., Bovey, P., & Junod, P. 1986. On the specific attraction of the
males of the six-toothed spruce bark beetle Pityogenes chalcographus to
a mixture of synthetic pheromones of the eight-toothed spruce bark beetle
Ips typographus Coleoptera: Scolytidae. Experientia. Basel. 42 (3).
325-326. ab Traps containing a mixture of attractants for Ips typographus
also caught other species of bark beetles. The numbers of Pityogenes
chalcographus, Pityogenes conjunctus, and Trypodendron lineatum were too
high to be interpreted as accidental captures. The males of the
polygamous P. chaleographus were specifically attracted, indicating that
the I. typographus pheromone or one of its components acts as a kairomone
and primary attractant for the pioneering males of P. chalcographus. LG

Birgersson, G., Byers, J.A., Bergstrom, G., & Lofqvist, J. 1990.
Production of pheromone components chalcogran and methyl e z-2 4
decadienoate in the spruce engraver Pityogenes chalcographus. J. Insect.
Physiol. 36 (6). 391-396. ab Capillary gas chromatography and mass
spectrometry were used to quantify the amounts of E- and Z- chalcogran
amd methyl (E, Z)-2,4-decadienoate (E, Z-MD), pheromone synergists of the
bark beetle Pitygenes chalcographus (Coleoptera: Scolytidae). Males were
exposed or not to vapours of host (Norway spruce, Picea abies) oleoresin
or allowed to feed in host logs prior to extraction of body parts and
hindguts for pheromone synergists. E, Z-MD and chalcogran were produced
sex-specifically in males, and only after feeding on host-plant tissue.
The pheromone synergists were not produced during exposure to oleoresin
vapours. Several oxygenated monoterpenes (including trans-verbenol,
myrtenol, and trans-myrtanol) were found in feeding males. The amounts
of the pheromone synergists in unmated feeding males remained relatively
constant over a 3.5 day period. In contrast to many other pheromone
components of bark beetles, including chalcogran, E, Z-MD was found
primarily in the male's body (head and thorax) with less in the hindgut
(abdomen). The probable acetogenic origins of both pheromone components
indicate that the species has evolved control over production and is thus
not dependent on host precursors as expected in many other bark beetles.

Bombosch, S., & Dedek, W. 1994. Integrated pest control against Ips
typographus (L.): Combined use of pheromones and the systemic insecticide
methamidophos (IPIDEX). Zeitschrift. Fuer. Pflanzenkrankheiten. Und.
Pflanzenschutz. 101 (5). 508-518. Issn 0340-8159. ab IPIDEX, a paste
containing the systemic insecticide methamidophos, is spread as a strip
20 cm wide and about 2 mm thickness to the pretreated bark of spruce and
protected with foil. The treatment is performed between the end of March
and the middle of April, according to the height a.s.l. ; some weeks
later, immediately before the attack of beetles, trees are baited with
a pheromone dispenser (Pheroprax for Ips typographus and Chalcoprax for
Pityogenes chalcographus). Spruce inside a circle of about 10 m around
the pheromon tree are created with IPIDEX only. The insecticide is
translocated in the ascending sap in upward direction only and has
reached during this time interval the top of the tree, killing beetles
boring in until the middle of June (halflife in spruce 40-45 d, 15 degree
C). The insecticide is present only inside the tree and no outwashing or
evaporation can be observed. The rapid degradation in the bark beetle
(halflife about 2 h) and the smell of a repellent at the site of
application as well prevent unwanted ecological side effects. Extensive
experiments during the increase of bark beetle gradation 1992/93 in
southern Germany yielded the decrease of freshly attacked trees inside
the near environment of the experimental locations in 1992 to 4-14 and
in 1993 to 18-28%, relative to the number of attacked trees in the year
before, although IPIDEX was applied once only in spring against the first
but not later against the second generation of Ips typographus. LG
GE.█Inst Forstzoologie, Univ, Buesgenweg 3, D-37077 Goettingen, Germany

Byers, J.A. 1983. Electronic fraction collector used for insect sampling
in the photoperiod induced diel emergence of bark beetles. Physiol.
Entomol. 8 (2). 133-138. ab An electron timer and fraction collector
consisting of CMOS integrated circuits is described. It converts 50- or
60-Hz AC to real-time pulses in programmable whole-number increments (1,
10 or 60 s) from 3 to 16,659, producing timing periods from 3 s to > 11
days. The fraction collector contains a leaf-switch feedback circuit that
automatically adjusts to various gear motor speeds and sample tube
spacings so that proper positioning results. Hourly collections by the
device of the bark beetles Ips typographus L. and Pityogenes
chalcographus L. (Scolytidae) emerging from logs of Norway spruce, Picea
abies, indicated that both species emerged with a diel periodicity. A
unimodal emergence peak for both sexes of both species occurred at midday
in LD (L = light, D = dark) 20:4 at a constant and 80% r.h.

Byers, J.A. 1992. Grid cell contour mapping of point densities bark
beetle attacks fallen pine shoots and infested trees. Oikos 63 (2).
233-243. ab A method for calculating and displaying patterns of local
density of plants and animals is presented for use with personal
computers. The algorithm, coded in the BASIC programming language, uses
x, y spatial point coordinates of organisms to calculate and display a
colored or shaded map of local densities within grid cells. The radius
of the local areas about the grid points within which densities are
calculated, the density-class interval boundaries when coloring the
cells, and the grid resolution can all be varied to facilitate
exploratory investigations. Contour density maps resulting from the
method are shown for an attack distribution of the bark beetle Pityogenes
chalcographus (Coleoptera: Scolytidae) on Norway spruce bark, Picea abies
and for a random distribution. Minimum allowed distance analysis of the
attack distribution indicated a distance of behavioral avoidance of
attack sites of about 1.6 cm for the bark beetle. Density maps of
different local radii are also shown for Norway spruce trees killed by
the bark beetle Ips typographus near Trebon, Czechoslovakia. Density maps
using different local radii and density increments are presented for
aggregated patterns of fallen Scots pine shoots, Pinus sylvestris,
infested by Tomicus minor bark beetles in southern Sweden. A method is
presented for statistical comparison of randomly-generated spatial point
data with natural data by using Chi-square analysis of the histograms of
differently colored grid cells. LG EN.█DEP ECOL, ANIM ECOL, LUND UNIV,

Byers, J.A. 1993. Avoidance of competition by spruce bark beetles Ips
typographus and Pityogenes chalcographus. Experientia. Basel. 49 (3).
272-275. ab Bark beetles, Ips typographus and Pityogenes chalcographus,
attracted by synthetic or natural pheromone to Norway spruce logs, Picea
abies, preferred to colonize uninfested logs rather than logs occupied
by these beetles, probably as a means of avoiding intra- and
interspecific competition. The aggregation pheromone components of P.
chalcographus, chalcogran and methyl (E, Z)-2,4-decadienoate, inhibited
the attraction response of I. typographus to its pheromone components
(methyl butenol and cis-verbenol), while the converse was not true.
However, verbenone released from colonized bark inhibited pheromonal
response of P. chalcographus. LG EN.█DEP ECOLOGY, LUND UNIV, S-223 62

Byers, J.A. 1993. Orientation of bark beetles Pityogenes chalcographus
and Ips typographus to pheromone-baited puddle traps placed in grids: A
new trap for control of scolytids. Journal. Of. Chemical. Ecology. 19
(10). 2297-2316. Issn 0098-0331. ab A puddle trap was designed that is
simple to build and efficient in catching bark beetles (Coleoptera:
Scolytidae). The trap is insensitive to wind and should be much easier
to manufacture than the more complicated perforated pipe and barrier
traps commercially available. A 7 times 7 grid of 49 puddle traps baited
with aggregation pheromone components of Pityogenes chalcographus
(chalcogran and methyl decadienoate) was placed at either 1.5-, 3-, 6-,
or 12-m spacing between traps in the field for two or more replicates of
one day length (June 1989, Torsby, Sweden). The resulting catches showed
that beetles were trapped as they flew into the grid since the inner
square-ring of 24 traps caught less beetles per trap than the outer
square-ring trap average (36 traps) in most experiments. Ips typographus
also landed in puddle traps primarily on the periphery of the grid (6-m
spacing only) when traps were baited with its pheromone components,
(S)-cis-verbenol and methyl butenol. Computer simulation of flying bark
beetles in grids of traps of various spacings and catch radii estimated
that the experimental pheromone traps had an effective catch radius of
1.3 m or less for P. chalcographus, depending on the spacing between
traps. An effective catch radius of 2 m for L. typographus was found for
the 6-m grid spacing. P. chalcographus beetles were increasingly
disrupted in their orientation to pheromone at the closer trap spacings
since the effective catch radius declined linearly with closer trap
spacing. However, landing was still precise since unbaited puddle traps
within the grid did not catch any bark beetles. LG EN.█Pheromone Res
Group, Dep Ecology Animal Ecology, Lund Univ, S-223 62 Lund, SWE

Byers, J.A., Anderbrant, O., & Lofqvist, J. 1989. Effective attraction
radius a method for comparing species attractants and determining
densities of flying insects. J. Chem. Ecol. 15 (2). 749-766. ab The
catches of bark beetles (Coleoptera: Scolytidae) were compared between
attractive traps releasing semiochemicals and passive traps (cylindricol
sticky screens long at 10 heights of 0.7-11.5 m, on poles). A central
attractive-trap pole was surrounded by three passive-trap poles spaced
50 or 100 m away at the apices of an equilateral triangle. The catches
of Tomicus piniperda and other scolytid species on the attractive-trap
pole baited with host monoterpenes, or the catches of Ips typographus
attracted to synthetic pheromone, were compared to passive trap catches
in a Scots pine forest or in a Norway spruc clear-cut, respectively.
Information about flight height distributions of the above scolytid
species, and Hylurgops palliatus, Cryphalus abietis, Pityogenes
chalcographus, P. quadridens, P. bidentatus, and Trypodendron domesticum
were obtained on the passive and attractive trap poles. A new method is
presented for determining the densities of flying insects based on the
passive trap's dimensions and catch, duration of test, and speed of
insect. Also, a novel concept, the effective attraction radius (EAR), is
presented for comparing attractants speices, which is independent of
insect density, locality, or duaration of test. The EAR is obtained by
the ratio of attractive and passive trap catches and the dimensions of
the passive trap, and thus should correlate positively with the strength
of the attractant and the distance of attraction. EARs are determined
from catch data of T. piniperda and I. typographus as well as from the
data of previous investigations on the same or other bark beetles. LG

Byers, J.A., Birgersson, G., Lofqvist, J., & Bergstrom, G. 1988.
Synergistic pheromones and monoterpenes enable aggregation and host
recognition by a bark beetle. Naturwissenschaften 75 (3). 153-155. ab.
We report the strong synergism between two pheromone components in
causing the attraction of the six-spined spruce bark beetle or
"Kupferstecher", Piyogenes chalcographus L. (Coleoptera: Scolytidae), in
the field. Our field results also suggest that several monoterpenes of
Norway spruce, Picea abies (L.) Karst, when presented with the pheromone
components, play a role in host recognition and colonization by the
beetle because they caused relatively more beetles to enter holes in
artificial hosts. Furthermore, sexual differences in strategies of host
resource competition are indicated because of the lower proportions of
males than females that entered the holes as well as were attracted to
higher release rates of pheromone. █DEP ECOL, UNIV LUND, S-223 62 LUND

Byers, J.A., Birgersson, G., Lofqvist, J., Appelgren, M., & Bergstrom,
G. 1990. Isolation of pheromone synergists of bark beetle Pityogenes
chalcographus from complex insect-plant odors by fractionation and
subtractive-combination bioassay. J. Chem. Ecol. 16 (3). 861-876. ab
Capillary gas chromatography with columns of different polarity and
two-dimensional fractionation of effluents were used with novel
subtractive-combination bioassays to rigorously isolate host- and
insect-produced pheromone synergists of the bark beetle Pityogenes
chalcographus (Coleoptera: Scolytidae). Methyl (E, Z)-2,4-decadienoate
(E, Z-MD) and the previously identified chalcogran were found to be
synergistically attractive to both sexes. E, Z-MD was produced
sex-specifically in males, and only when they had fed on host-plant
tissue. A Norway spruce monoterpene fraction (including .alpha.-pinene,
.beta.-pinene, and camphene) increased the attractive response to the
pheromone components. Dose-response curves for E, Z-MD and chalcogran in
the laboratory bioassay indicated the two components are highly
synergistic. The isolation methods are important for further progress in
identifying certain semiochemical synergists found in trace amounts in
complex chemical mixtures, such as when insects must feed in host plants
in order to produce pheromone. LG EN.█DEP ECOL, UNIV LUND, 223 62 LUND,

Byers, J.A., Hogberg, H.E., Unelius, C.R., Birgersson, G., & Lofqvist,
J. 1989. Structure-activity studies on aggregation pheromone components
of Pityogenes chalcographus Coleoptera: Scolytidae all stereoisomers of
chalcogran and methyl-2 4-decadienoate. J. Chem. Ecol. 15 (2). 685-696.
ab Syntheses of all four stereoisomers (2S,5S; 2S,5R; 2R,5R; and 2R,5S)
of chalcogran, a major component of the aggregation pheromone of
Pityogenes chalcographus, and of all four isomers (2Z,4Z; 2Z,4E; 2E,4E;
and 2E,4Z) of methyl 2,4-decadienoate (MD), the second major pheromone
component, are briefly described. Attraction responses of walking beetles
of both sexes were tested to mixtures of the synergistic pheromone
components or analogs. These bioassays showed that the E, Z isomer of MD
is the most active when tested with chalcogran. When tested with (E,
Z)-MD, (2S,5R)-chalcogran was the most active stereoisomer, while 2R,5R
and 2R,5S isomers had intermediate activities, and the 2S,5S isomer was
inactive. There was no evidence that the relatively less active
stereoisomers of chalcogran inhibited or promoted attraction to
(2S,5R)-chalcogram with (E, Z)-MD. Male beetles only produce the active
E, Z isomer of MD (inactive alone) and their hindguts contain the most
active (2S,5R)- and least active (2S,5S)-chalcogran. A mixture of all MD
isomers with racemic chalcogran was not significantly different in
attractivity compared to (E, Z)-MD with racemic chalcogran, indicating
no synergistic or inhibitory effects of the inactive isomers of MD. LG

Duelli, P., Studer, M., & Naef, W. 1986. The flight of bark beetles
outside of forest areas. J. Appl. Entomol. 102 (2). 139-148. ab Flight
phenologies and vertical distribution of bark beetle flight were
investigated in an agricultural area at least 420 m way from potential
breeding places. Pheromone traps (Pheroprax, Linoprax) and unscented
sticky traps (square grids of 1 m2) were fixed on a meterological mast
at 9 different heights from 1.7 m up to 150 m. Of the 12 scolytid species
recorded in 1984, Ips typographus L. (N = 287) and Pityogenes
chalcographus L. (N = 319) were the two most abundant species. The
vertical flight distribution of I. typographus shows a marked peak at 5
m. Less than 5% flew higher than 10 m, with 1 individual at 100 m. More
females than males were caught in the pheromone traps. Maximum catches
of P. chalcographus were at the lowest level, at 1.7 m. Decrease in
numbers with height is slower than in I. typographus; 14% flew higher
than 10 m. In the pheromone traps, 82% were males. In the sticky traps,
the sex ratios of both species were more balanced. The phenologies of
both I. typographus and P. chalcographus show two peak flight periods.
Flight in May/June respresents the overwintered generation, July/August
presumably the summer generation. A comparison with reference traps
within forest area 700 m north and south of the meteo mast reveals that
the proportion of Ips typographus flying out of forest areas varies
greatly between the two generations while less than 10% of the
overwintered beetles were trapped outside of the forest, more than a
third of the summer catches were trapped far away form any potential
breeding places. It is suggested that habitat changes (innerforest
movements) are mainly performed by the summer generation, while the
overwintered beetles are less mobile. From March to September,
Trypodendron lineatum was abundant in forest areas, but virtually absent
in our traps outside the forest, suggesting a far less dispersive flight

Eidmann, H.H. 1987. Insects attacking logs of exotic conifers in sweden.
J. Appl. Entomol. 103 (3). 278-283. ab Insect attack on logs from
25-year-old trees of 22 species in the genera Picea, Abies, Larix, and
Pseudotsuga was studied in the province of Ostergotland in Sweden.
Attacks by the bark beetle Pityogenes chalcographus predominated, more
rarely occurred Ips typographus. All other frequently found insects were
typically secondary species with broad host ranges, i.e. Hylurgops
palliatus, Dryocoetes autographus, Trypodendron lineatum, and Rhagium
inquisitor. The intensity of insect attack varied between tree species
and genera. The Abies species were least attacked. Abies, Larix, and
Pseudotsuga were less frequently attacked by P. chalcographus and were
less suitable as breeding substrate for this bark beetle than was Picea.

Eidmann, H.H. 1992. Impact of bark beetles on forests and forestry in
sweden. J. Appl. Entomol. 114 (2). 193-200. ab The impacts of bark
beetles in Sweden and the role and management of bark beetle breeding
substrate are discussed. Of the six important species damaging living
trees (Tomicus piniperda, Tomicus minor, Hylastes cunicularius,
Polygraphus poligraphus, Pityogenes chalcographus, Ips typographus),
Tomicus piniperda and Ips typographus have the greatest impact. The
damage caused by Tomicus adults feeding in pine shoots is directly
related to beetle numbers, which in turn depend on the availability of
non-resistant breeding substrate. Intensity of shoot pruning, tree size,
and geographic region are important factors determining the extent of
losses, which can amouunt to 45% of the annual volume growth. The extent
of tree mortality caused by Ips typographus attack depends on
interactions between host tree vigour, densities of colonizing
populations, and the availability of other, non-resistant breeding
substrate. During the latest outbreak, the trees killed represented about
6 million cubic meters of wood. The heaviest losses were reduction of
timber quality, unsalvaged timber, and the cost of control measures. The
economic impact of reduced growth depends on the shortening of rotation
by insect attack and on interest rates. During the 1960s and 1970s bark
beetle attacks and their impact increased owing to natural causes and
forestry practices. Hereafter, integrated forest protection efforts,
based on research, organisation, information, and legislation, reduced
bark beetle attacks to a low level. LG EN.█SWEDISH UNIV AGRICULTURAL
7044, S-750 07 UPPSALA, SWEDEN

Francke, W., Heemann, V., Gerken, B., Renwick, J.A., A., & Vite, J.P.
1977. 2 ethyl-1 6-dioxa spiro-4 4 nonane principal aggregation pheromone
of Pityogenes chalcographus. Naturwissenschaften 64 (11). 590-591. no
abstract available, however, this is the paper where chalcogran (the
chemical above) was identified as one of the aggregation pheromone
components of P. chalcographus.█

Fuehrer, E. 1977. Studies on intraspecific incompatibility in Pityogenes
chalcographus Coleoptera: Scolytidae. Z. Angew. Entomol. 83 (3). 286-297.
ab The fertility of 13 North and Central European strains of P.
chalcographus was comparatively studied in the laboratory and crossing
experiments between some of these strains were performed. Differences in
fertility and reproductive incompatibility between certain strains were
recognized. Incompatibility comes from increased egg mortality reduced
egg production. There is much variation in the degree of incompatibility,

Fuehrer, E. 1978. Race differentiation in Pityogenes chalcographus
Coleoptera Scolytidae part 1 morphological characteristics. Z. Angew.
Entomol. 86 (4). 392-402. ab Morphological variation in P. chalcographus
revealed significant differences between north-east European and central
European populations and between populations within each region. The
pattern of geographical distribution of the different types corresponds
to the natural distribution of spruce in Europe. LG GE.█INST FORSTZOOL,

Fuehrer, E. 1980. Applicability of fungicides as an experimental tool for
investigations of bark beetles. Anz. Schaedlingskd. Pflanzenschutz.
Umweltschutz. 53 (3). 36-40. ab The fungicides Euparen and Orthocid 50
were efficient in suppressing growth of saprophagous fungi on spruce
phloem chips, which are used as a food of Pityogenes chalcographus adults
in laboratory experiments. Beauveria bassiana infection of the beetles
is also checked, when fungicides are applied to the food. LG GE.█INST

Fuehrer, E. 1981. Seasonal quality fluctuations of norway spruce phloem
picea-excelsa as a brood material of the bark beetle Pityogenes
chalcographus Coleoptera: Scolytidae. Z. Angew. Entomol. 91 (1). 74-83.
ab Food quality of Norway spruce phloem for breeding P. chalcographus
females and for their progeny is indicated by the density of egg niches,
number of progeny produced per time unit, number of adult progeny, the
time needed for its development and by adult body weight, respectively.
Food quality changes during the year, particularly in Sept.-Oct., when
breeding is at a minimum. LG GE.█INST FORSTZOOL, UNIV GOTTINGEN

Fuehrer, E., Hausmann, B., & Wiener, L. 1991. Bark beetle Coleoptera
Scolytidae colonization and terpene patterns in the bark of norway spruce
picea-abies karst. trap trees. J. Appl. Entomol. 112 (2). 113-123. ab
Trap trees of the Norway spruce were used to investigate correlations
between colonization density of bark beetles and terpene patterns of the
trunk phloem. Ips typographus and Pityogenes chalcographus were the
dominant species, which demonstrated the usual differentiated patterns
of attack concentration along the trunks. There were, however also
exceptions made by P. chalcographus, extending its mass colonization down
to more basical parts of the trunk. Phloem samples were collected from
7 trap trees, when bark beetle flight was just starting in middle of May
1985. Three samples per trap tree were each taken from the center of the
bottom, middle and top third of the trunk, respectively, and analyzed by
a GC/MS device. The average density of enterance holes of I. typographus
and P. chalcographus as calculated for each third of an individual trunk,
was now compared with the varying mono- and sesquiterpene patterns of the
respective trunk sections. A strong positive correlation of P.
chalcographus attack density was found to exist to myrcene, and to the
.beta.-pinene: .alpha.-pinene relation, and to .beta.-phellandrene
.times. limonene as well. Correlation was a little higher, when these
three terpene values were combined by addition. With I. typographus it
was more difficult to find correlations with terpene contents of the
bark. A r = 0.679 (p < 0.01) was calculated for the term (.alpha.-pinene
.times. limonene):(myrcene .times. camphene); neglecting some outliers
correlation could be increased to r = 0.912. Considering the total number
of values, correlation is improved by addition of .beta.-phellandrene:
(.DELTA.3-carene + 1) and tricylene .times. (terpinolene + 1) (r = 0.739,
p < 0.001). It is suggested that the differing preference of I.
typographus and P. chalcographus for completely different terpene
patterns of the spruce phloem may play an important role in host tree
selection as well as in selection of the colonization site within a
trunk. Further experiments should demonstrate, whether these terpene
fractions are involved in the "primary attraction principle" of Norway
spruce to the investigated bark beetle species. LG GE.█INST FUER

Fuehrer, E., Wiener, L., & Hausmann, B. 1992. Dynamics of terpene
patterns and bark beetle colonization in spruce trap trees showing
different crown depletion Coleoptera: Scolytidae. Entomol. Gen. 17 (3).
207-218. ab Dynamics of cortical terpene patterns, which are considered
to be relevant for orientation of Ips typographus (Linnaeus 1758) and
Pityogenes chalcographus (Von Linne 1761), respectively; have been
studied from logging until insect colonization in several trap trees
(Picea abies Karsten). During this period, significant changes of the
respective terpene patterns did not occur. It is, therefore, assumed that
the terpene equipment of the safe tree may represent its potential
attractivity, while its actual attractivity may then be determined by
structures and water condition of phloem and by physical factors, which
influence the emission of single compounds. The potential attractivity,
however, appears determined by the stress condition of the tree, which
was judged by the degree of depletion and discoloration of the crown. LG

Gries, G. 1986. As to the importance of maturation-feeding for the
dispersal flight of Pityogenes chalcographus l. Coleoptera: Scolytidae.
Z. Angew. Zool. 73 (3). 267-280. ab The emergence and successive feeding
of adults in vitro were studied under standardized conditions during 16
days. During that time, the adults showed changes in colour and became
sclerotized. These processes were influenced by nutrition. When the
feeding was purposely interrupted, these processes came into a
stand-still. Without feeding the beetles were not able to fly. During the
maturation-feeding their flight-ability developed, also more or less
parallel their flight-disposition. The ability to starve (and therefore
the later-flight-capacity) improved. That is caused by a growing amount
of fat content. The strengthening structure of the cuticula may have a
supporting effect. As the physiological abilities develop parallel with
colouring the colour of the young beetles become indicator of their
physiological state and ability. During the maturation-feeding pheromones
were produced already. Therefore the production of pheromones did not
depend on the degree of maturity of the beetles but only on the freshness

Harding, S., Lapis, E.B., & Bejer, B. 1986. Observations on the activity
and development of Pityogenes chalcographus Coleoptera: Scolytidae in
stands of norway spruce in denmark. J. Appl. Entomol. 102 (3). 237-244.
ab Exploitation of thinnings in young stands of Norway spruce for the
production of chips necessitates an period of seasoning of the felled
trees in the stand. This infers a potential risk of bark beetle attack
on standing trees. The seasonal activity and reproduction of Pityogenes
chalcographus L. were studied in shade and half-shade.sbd.conditions
representative of thinning by rows. In half-shade chipping is required
before August in order to avoid emergence of the offspring from trees
infested during the early spring flight in May. From trees attacked later
on or lying in shade emergence is delayed and chipping can be carried out

Hilker, M. 1984. Attractivity of various white paints with different
rates of uv and visible light reflectance for Pityogenes chalcographus
and Ips typographus Coleoptera: Scolytidae. Z. Angew. Entomol. 98 (5).
463-473. ab Laboratory tests on the attractivity of various white paints
with different rates of reflectance for UV and visible light were
performed on the bark beetles P. chalcographus L. and I. typographus L.
In binary choices traps with different white paints were tested against
a standardized black trap. The criterion of the attractivity of the white
paint was the quantity of specimens caught in the respective trap. When
illumination with UV rich light of low luminous intensity was applied on
P. chalcographus, white painted traps, which showed with respect to the
band of wavelength .lambda. = 390-470 nm rates of reflectance .gtoreq.
74%, were more attractive than the black trap (i.e., traps with Pb
white-, Pb white plus varnish-, magnesium oxide- and barium sulfate
paint); .ltoreq. 70%, were equally (or less) attractive as (than) the
black trap (i.e., traps with Zn white- and latex-paint, covered with an
Al strip, white cotton cloth and white cotten cloth plus varnish).
White-paints with rates of reflectance at 70-74% in the band .lambda. =
390-470 nm may represent a threshold of attractivity for P.
chalcographus. For I. typographus distinct differentiation between traps
with UV light reflectance of various degrees was found as well; the Pb
white-trap (with high reflectance for UV and visible light) was much more
attractive than the Zn white-trap, which has lower rates of reflectance
in respect to the UV and blue-violet bands of the spectrum than the Pb
white-trap. Alterations of the spectral radiation flux from the source
of light and variations of the luminous intensity at the trap resulted
in changes of the attractivity of white-paints for P. chalcographus; when
compared to illumination with UV rich light of low luminous intensity,
illumination with light of low UV content, but high luminous intensity,
resulted in decreasing captures in traps with rates of reflectance
.gtoreq. 74% for .lambda. = 390-470 nm (i.e., Pb white-, Pb white plus
varnish-, magnesium oxide- and barium sulfate-paint). Furthermore, these
traps were more attractive than the black trap. The efficiency of capture
for traps with low reflectance within .lambda. = 390-470 nm increased
(i.e., Zn white- and latex-paint); however, it did not overtop that of

Jahn, E. 1982. Studies on the attack on spruce trees by bark beetles in
connection with moon phases. Anz. Schaedlingskd. Pflanzenschutz.
Umweltschutz. 55 (10). 145-149. ab The interaction of moon phase, felling
of spruce, weather conditions and severity of scolytid attack was studied
in Austraia. Scolytids, Ips typographus, I. amitinus and Pityogenes
chalcographus attacks were most severe at the full moon. This pattern was
maintained through out the year although scolytid pheromone release
during the later phases of swarming did complicate the pattern.
Production by the tree of chemical attractants coincident with lunar
phase is the suggested explanation of the pattern observed. LG

Johansson, L., Andersen, J., & Nilssen, A.C. 1994. Distribution of bark
insects in "island" plantations of spruce (Picea abies (L.) Karst.) in
subarctic Norway. Polar. Biology. 14 (2). 107-116. Issn 0722-4060. ab
Eleven isolated plantations of Norway spruce Picea abies (L.) Karst. in
Troms county, northern Norway, were examined for bark insects in summers
1990 and 1991. Field-infested bolts from controlled fellings were reared
in laboratory to complement field-investigations. Results are compared
to records made from the same plantations 1974-78. Eight scolytid
species, two curculionids and two cerambycids were found inhabiting
spruce in 1990-91. Dryocoetes autographus was found in ten, and Tetropium
castaneum in 9 plantations. Hylastes cunicularius has apparently expanded
its distribution since 1978. One plantation had no bark insects confined
to spruce, except old galleries of Pityogenes chalcographus. No new
immigrants to the region were found, but one curculionid species may have
become extinct. Reproduction of Dryocoetes autographus and Tetropium
castaneum was studied in the laboratory D. autographus bred consecutive
generations from the same bolt, whereas T. castaneum displayed varying
generation time, partly within the same bolt. This enhances their
viability in plantations where felling activity is low. Several scolytid
species widely distributed elsewhere in naturally spruce-forested areas
in Scandinavia north of the Arctic circle are still absent in Troms,
including all the smallest species. It is suggested that the success of
colonizers in unpredictable climates depends on both dispersal
capabilities (larger species have a higher dispersal power than the
smallest species), life cycle flexibility and versatility in habitat
requirements. It is concluded that only species with certain properties
have established in the plantations, i.e. the immigration and
colonization is not stochastic. LG EN.█Grunnvag, 9372 Gibostad, NOR

Klipstein, E.L. 1986. Cytology and sperm polyploidy in Pityogenes
chalcographus Coleoptera: Scolytidae. J. Appl. Entomol. 102 (3). 285-295.
ab These experiments are based on the biological relationship that a
partial reproductive incompatibility does exist between certain
populations of Pityogenes chalcographus. Besides this and the existence
of unusual sperm polyploidy in spermatogonial development four regionally
distinct strains of this bark beetle were cytologically studied.
Karyotypic determinations were made on orcein squashes and MEREDITH
preparations of testes taken from callow adults. The karyotypic formula
during the spermatogonial metaphase is 18 A + Xy and during the first
meiotic metaphase 9 AA + Xyp. No diagnostic chromosome polymorphism
within the different populations was detected. In first meiotic metaphase
the first bivalent is ring-shaped in almost every nuclei. This ringform
is explained as the result of obligate chiasma. In general, the
spermatogonial development is normal, as is known for insects, but in
some cases very large spermatids have been observed, which are the result
of spermatogonial polyploidy. This genom multiplication takes place
before meiosis. The parental strains differ slightly in frequency of
sperm polyploidy. Examinations of six F1 interpopular crosses provide a
somewhat increased occurrence of sperm polyploidy. The origin of these
remarkable cytological features and the conclusions pertaining to them
are discussed. LG GE.█MOERIKESTR 16, D-4790 PADERBORN

Klipstein, E.L. 1986. Intraspecific differentiation in gradology and
ecology of Pityogenes chalcographus Coleoptera: Scolytidae in view to
interpopular crossing. Anz. Schaedlingskd. Pflanzenschutz. Umweltschutz.
59 (7). 131-135. ab Four regionally different strains and six F1
interpopular crosses of Pityogenes chalcographus were used for
infestation and starvation experiments. In the infestation experiment
beetles were placed for three days on vital Norway spruce and the amount
of nuptial chambers were recorded. During the viability experiment the
insects were kept under a daily varying temperature for 16 h at and for 8 h at The rel. humidity was set at 93%,
77% and 57%. The infestation potential differs in the susceptibility of
host trees and within the populations. Especially the F1 crosses provide
an enhanced attack and potential of injury which are to be considered a
case of heterosis. Because of the irregular host-tree preference in some
populations it is assumed that there is a highly specific interaction in
feeding stimulation. The effect of heterosis is also present in
starvation experiments, but it seems to be only restricted to an optimal
survival range. It is emphasized that genetic factors play a remarkable
role in the interactions between bark beetles and host trees as well as
in population dynamics. LG GE.█MOERIKESTR 16, D-4790 PADERBORN

Kohnle, U. 1985. Investigations of chemical communication systems in
secondary bark beetles Coleoptera: Scolytidae. Z. Angew. Entomol. 100
(2). 197-218. ab The chemical communication system of several secondary
European bark beetles was investigated. The ash bark beetle, Leperisinus
varius, responded in the field to ethanol-baited traps in significant
numbers. Male-released exo-brevicomin increased trap catches of L. varius
while another male specific compound, 7-methyl-1,6-dioxaspiro (4.5)
decane, proved inhibitory. Hylurgops palliatus, a species colonizing
Norway spruce was attracted to traps containing ethanol as a bait; spruce
resin acted synergistically. For Dryocoetes autographus, another spruce
bark beetle, females were shown to be the host selecting sex, suggesting
this species to be monogynous. However, the males release endo- and
exo-brevicomin which attracted predominantly female D. autographus in the
field. With endo-brevicomin, only the (+)-enantiomer elicited response
of D. autographus while the antipode did not affect beetle response. In
polygynous bark beetle species males are the host selecting sex and as
a rule release the aggregating pheromone(s) which attract both males and
females. Dispersal losses of hostsearching males are compensated through
male polygyny. In contrast, some monogynous bark beetles such as D.
autographus seem to balance the sex ratio during host colonization
through sexual cross-attractiveness of sex-specific pheromones: Male and
female beetles release different pheromone components attracting
predominantly beetles of the opposite sex; a phenomenon characteristic
for sex pheromones of Lepidoptera. Within associations of bark beetles
colonizing Norway spruce investigated in this study, kairomonal effects
of pheromones were common. E.g., the aggressive species Ips typographus
and Pityogenes chalcographus were attracted to some extent to endo- or
exo-brevicomin, pheromone components of secondary D. autographus. In
contrast, response of D. autograpuys to baited traps was inhibited when
pheromone components of the aggressive species were added. LG

Kubisz, D. 1992. Occurrence of predators from the genus rhizophagus
herbst col. rhizophagidae in pheromone traps. J. Appl. Entomol. 113 (5).
525-531. ab The observations were carried out from 1984 to 1989 in
Krynica (Beskid Sadecki, Western Carpathians (Poland)) during the
experiments concerning the utilization of pheromone traps against
Xyloterus spp. and Pityogenes chalcographus under the mountain
conditions. Drainpipe and screen traps baited with LINOPRAX, CHALCOPRAX,
and mixture of lineatin and ethanol were used in these experiments. There
were 7 species from the genus Rhizophagus found in traps (R. depressus,
R. nitidulus, R. dispar, R. cribratus, R. parvulus, R. perforatus, R.
bipustulatus). Their numbers, sex ratio, time of appearance, maximum of
occurrence, and the correlation between their appearance and flight of
Xyloterus spp. and P. chalcographus are discussed. The percentage of
Rhizophagidae in the total number of trapped insects, other than
Xyloterus spp. and P. chalcographus, was calculated and the selectivity
of both types of traps used was evaluated. The numbers of Rhizophagidae
were directly proportional to the distance between the traps. LG EN.█DEP

Lobinger, G. 1994. Air temperature as a limiting factor for flight
activity of two species of pine beetles, Ips typographus L. and
Pityogenes chalcographus L. (Col., Coleoptera: Scolytidae). Anzeiger.
Fuer. Schaedlingskunde. Pflanzenschutz. Umweltschutz. 67 (1). 14-17. Issn
0340-7330. ab Single catches of bark beetles in pheromone baited traps
were registered by the aid of a new instrument. It consists of an
electronical weather station in connection with a pheromone trap with
infrared sensors as additional equipment. So it was possible also to
register weather data at the moment of every catching event. It could be
shown that the flight behaviour of both species of bark beetles was
influenced by air temperature. I. typographus did not fly beneath a
minimum temperature of 16.5 degree C. There was also an upper limit of
30 degree C for flight activity of this species. P. chalcographus has a
threshold of temperatures about 16.8-17 degree C for activity. No upper
limit of temperature could be observed up to 35 degree C. Both of them,
I. typ. and P. chalc., reacted very sensitive and spontaneous to these
temperature thresholds. LG GE.█Landesanstalt Wald Forstwirtschaft,
Hohenbachernstr 20, D-85354 Freising, GER

Michalski, J., Kaczmarek, S., & Ratajczak, E. 1991. Mites acari
mesostigmata in the bark-beetle qualleries Coleoptera: Scolytidae of the
gorczanski national park. Pol. Pismo. Entomol. 61 (3-4). (1992). 137-142.
ab Twenty six species of Mesostigmata of 7 families were found to occur
in the galleries of Ips typographus (L.), Polygraphus poligraphus (L.)
and Pityogenes chalcographus (L.) in the Gorczanski National Park. The
most abundant and diversified mite fauna was found in the galleries of
Ips typographus, and populations of Dendrolaelaps quadrisetus (BERLESE)
constituted c. 40% of the association, The mite species listed in the
paper are new for the fauna of the Gorczanski National Park. LG

Moor, H., & Neffeler, M. 1983-. A bark beetle-eating spider
troxochrus-nasutus araneae erigonidae. Faun. Oekol. Mitt. 5 (9-10). 1984
ETH-ZENTRUM, CH-8092 ZUERICH Moor, H., & Nyffeler, M. 1983. A note on
spiders killing bark beetles. Mitt. Schweiz. Entomol. Ges. 56 (1-2).

Mortiz, B., & Fuehrer, E. 1988. Phloem compounds of picea-abies analyses
on bark beetle trap-trees. J. Appl. Entomol. 105 (5). 502-509. ab In the
phloem of Picea abies, the amounts of starch and soluble carbohydrates
of phenols, resin and nitrogen were measured. During storage of five
weeks, the amounts of starch decreased rapidly, while that of the other
investigated substances did not change significantly. The amounts of
starch and phenols are significantly higher in dominant trees than in
overtopped individuals. Brood experiments showed that dominant trees were
more resistant to the attack of the sixth-toothed bark beetle (Pityogenes

Naumann, & Etienne, K. 1978. The influence of forest structure and the
availability of trees equipped with traps on the flying behavior of some
species of beetles in storm damaged conifer woods. Z. Angew. Zool. 65
(4). 397-414. ab In a spruce stand near Celle, Germany, some problems
concerning spatial distribution during flight of common bark- and
wood-inhabiting Coleoptera were investigated by means of 40 window traps
and a trap tree barrier. Most of the 38 spp. seem to have micro- and
mesoclimate preferences during flight. Numerous species ignored trap
trees situated away from their climatic requirements. Pathogenic spruce
bark beetles Ips typographus and Pityogenes chalcographus behave
similarly, except for a pronounced restriction of I. typographus to the
stratum of tree tops. Both species fly mainly against the wind and in
spring they only swarm on warm days with southerly winds. These facts may
counteract the dispersal of populations during spring flight.
Interspecific competition between I. typographus and P. chalcographus for
suitable trees was observed, but seems to have little influence on their

Niemeyer, H. 1992. Monitoring Ips typographus and Pityogenes
chalcographus col. Scolytidae in lower saxony and schleswig-holstein. J.
Appl. Entomol. 114 (1). 98-102. ab Due to the intensive management of
German forests, the main purpose of bark beetle monitoring is not to
detect stands of high infestation risk. Much more, monitoring shall give
a rough idea of regional population trends. Some years experience show
that single monitor traps more than 50 m apart from each other use to
cause pheromone induced infestations in nearby standing spruce. Thus, in
1988 a system of monitoring has been established in Lower Saxony and
Schleswig-Holstein which consists of mass trapping lines along margins
as usual (distances of 30-50 m between traps and 10-15 cm between traps
and next spruce) but each of them with one fortnightly checked trap: the
monitor trap. This system helps reducing infections and provides records
of semimonthly data from April to September. The analysis of 3 years of
flight activity patterns and data tables did not reveal a method to use
the data for a long term prediction. However, a short term prediction
based on monthly warning messages reffering to regions seems to be
feasible, provided that corresponding short term thresholds of
infestation risks will be elaborated. LG EN.█NIEDERSAECHSISCHE FORSTLICHE

Nilssen, A.C. 1979. Development of a bark fauna in plantations of spruce
picea-abies in north norway. Astarte 11 (2). 151-170. ab The spruce bark
fauna (phytophagous Coleoptera: Scolytidae, Curculionidae, Cerambycidae)
was investigated in 141 spruce plantations in northernmost Norway beyond
the natural distribution of spruce. In addition to the species from the
indigenous pine (Pinus sylvestris (L.)), several host specific spruce
beetles were found. Dryocoetes autographus Ratz., Hylastes cunicularius
Er., Tetropium castaneum L and D. hectographus Reitt. were most common,
detected in 73, 35, 23 and 17 plantations, respectively. Hylurgops
glabratus Zett., Ips typographus L., Pityogenes chalcographus L. and
Pissodes gyllenhali Gyll. were more frequent. Evidence of long-distance
aerial dispersal from the natural spruce forests in north Sweden and
Finland was shown by finds of spruce beetles in gut contents of fish from
mountain lakes. On 1 occasion I. typographus, P. chalcographus and
Tetropium castaneum were carried at least 180 km by aerial currents.
Occurrence of spruce beetles on some islands may be explained by
anthropochorous dispersal. LG EN.█ZOOL DEP, TROMSO MUS, UNIV TROMSO,

Nilssen, A.C. 1984. Long range aerial dispersal of bark beetles and bark
weevils Coleoptera: Scolytidae and curculionidae in northern finland.
Ann. Entomol. Fenn. 50 (2). 37-42. ab Trap logs of spruce (Picea abies)
were placed at different distances north of the spruce forests along the
Muonio-Kilpisjarvi road in northwestern Finland. At the greatest
distance, 171 km (Kilpisjarvi), 3 scolytids were found: Dryocoetes
autographus, Hylastes cunicularius and H. brunneus, and the curculionid
Hylobius abietis. These were evidently dispersed anemochorously from the
Finnish/Swedish forests. At the other sites spruce bark beetles were
found at the following maximum distances from the spruce forests:
Pityogenes chalcographus and H. cunicularius: 86 km, D. autographus: 52
km, Ips typographus: 43 km (trap log baited with synthetic pheromones),
Hylurgops glabratus: 19 km, and D. hectographus: 10 km. Some of the
species may have originated from small populations living on nearby
non-host trees Pinus sylvestris, but most of them were probably blown by
winds or actively flew from the spruce forests. LG EN.█ZOOLOGY DEP,

Pettersen, H. 1976. Chalcid-flies hymenoptera chalcidoidea reared from
Ips typographus and Pityogenes chalcographus at some norwegian
localities. Norw. J. Entomol. 23 (1). 47-50. ab A total of 12 spp. of
chalcid-flies were found as parasites or hyperparasites on Ips
typographus and Pityogenes chalcographus from Norwegian localities. Of
these, 7 are not previously reported from Norway (not indicated). The
species Eurytoma blastophagi Hedqv., Roptrocerus xylophagorum Ratz., R.
brevicornis Thoms., Dinotiscus eupterus (Walk.) and Tomicobia seitneri
(Ruschka) were predominant in the material from the localities examined.
The species Eurytoma arctica Thoms. and Heydenia pretiosa Forst. were not
frequently found. Eurytoma arctica Thoms. and especially E. morio Boh.
and Rhopalicus tutela (Walk.) are species sporadically found on these
spruce bark beetles, but known to be more frequent on pine bark beetles.
(The remaining species found were Rhopalicus brevicornis Thomson,
Karpinskiella pityophtori Boucek and Mesopolobus typographi (Ruschka).

Ritzengruber, O. 1990. Isoenzyme analysis of different populations in
Pityogenes chalcographus l. Coleoptera: Scolytidae ii. population
structure differentiation of populations. J. Appl. Entomol. 109 (1).
55-63. ab Six populations of the spruce bark beetle Pityogenes
chalcographus from Scandinavia (Finland, Norway) Eastern and Central
Europe (Poland, Federal Republic of Germany, Austria) were analysed for
their genetic structure of seven enzymes by polyacrylamide
gel-electrophoresis. Significant differences between the populations were
observed in GOT (glutamate oxaloacetate transaminase), EST (esterase) ,
APH (acid phosphatase) and AMY (amylase). CAT (catalase) and LAP (leucine
aminopeptidase) were also polymorphic but differed very little. LIP
(lipase) was monomorphic in all populations. By calculating the genetic
identities and after carrying out an UPGMA-cluster analysis two
population groups were observed. Within the first are RA from Finland and
WA from Poland. The second group contains the remaining populations VI
from Finland, NO from Norway and the populations from Central Europe JA
and LI. The differences respectively the similarities between the
populations are interpreted by the history of distribution and

Ritzengruber, O., & Fuehrer, E. 1986. Methods for analyzing isoenzymes
of different populations in Pityogenes chalcographus Coleoptera:
Scolytidae i. adaptation of methods enzyme polymorphism. J. Appl.
Entomol. 101 (2). 187-194. ab Methods for the distinct separation of the
isozymes of peroxidase, glutamate-oxaloacetate-transaminase, esterase,
acid phosphatase, amylase and leucine aminopeptidase of the bark beetle
Pityogenes chalcographus (Col., Scolytidae) by polyacrylamide gel
electrophoresis are described. All six enzymes are found polymorphic. The
electrophoretic patterns are typical for each population from six
different regions of northern and central Europe. LG GE.█INST FUER

Sanders, W., & Horn, M. 1982. Studies on the preference for white of the
bark beetle Pityogenes chalcographus. Z. Angew. Zool. 69 (4). 495-508.
ab During the flightperiod males of the bark beetle P. chalcographus
preferred in binary choices white trunkmodels made from pasteboard
(Fotokarton). Gray, black and black-white colored models were neglected.
The beetles showed the same behavior when a white model was compared with
a spruce log of the same size. White models were more attractive in a
white than in a black colored environment. The same effect was given,
when a bright environment was compared with a shady one. A white model
in a bright and white colored environment was highly attractive. Not the
contrast between model and environment but the light intensity around the
model was important. Spruce logs in a bright area were much more
attractive than spruce logs in a shady region. The boring activity on the
surface of the spruce logs was higher in a bright than in a shady region.
P. calcographus showed 2 characteristic types of flight: the spiral
flight released by optical stimuli and the round dance flight released

Schlyter, F. 1992. Sampling range attraction range and effective
attraction radius estimates of trap efficiency and communication distance
in Coleopteran pheromone and host attractant systems. J. Appl. Entomol.
114 (5). 439-454. ab There is a need for information on communication
distances in natural systems and the ranges of action of synthetic
pheromone traps. The differences between sampling range (RS, time
dependent ecological measure) and attraction range (RA, behavioral
measure) are stressed and compared to the simple but artificial effective
attraction radius concept (REA, mainly trap and bait dependent). To
estimate such ranges defined theoretical concepts and proper analysis of
data are needed. Quantitative estimates are rare in the literature but
maximum RA for Lepidoptera (moths) may reach 200-400 m and the
corresponding Rs functions may be in the range 200-500 m. RS estimated
from regression of recapture of marked Cyclas formicarius (Curculionidae)
after 16 h of sampling was 390 m for a high dose (10 .mu.g) and 94 m for
a low dose (0.01 .mu.g). RA for bark beetles (Scolytidae) estimated from
regression analysis of trap interactions are in the range of 19-97 m, but
have large confidence intervals due to the few data points available.
Passive traps at 1-35 m distance from pheromone traps yield RA values
based on log-log regression values of between 17 and 34 m depending on
pheromone release(10-5 to 10-2 g/h) for Ips typographus. Effective
attraction radii's were estimated, both as simple point-estimates and as
means with confidence intervals. Minima and maximum estimates obtained
for bark beetles were 0.2-1 m for Tomicus piniperda, 0.6 m for T. minor,
0.3-3 m for IPs typographus, 3-10 m for Pityogenes chalcographus and
0.2-0.4 m for Scolytus (scolytus) triarmatus. It is concluded that RS and
RA are of great basic and applied value but both dififcult to estiamte.
REA lacks a direct relation to natural behaviour but is a useful index
of trapping power and specificity. LG EN.█PHEROMONE GROUP, DEP ECOL ANIM

Schroeder, L.M., & Eidmann, H.H. 1993. Attacks of bark and wood-boring
Coleoptera on snow-broken conifers over a two-year period. Scand. J. For.
Res. 8 (2). 257-265. ab The attacks of bark- and wood-boring Coleoptera
on broken conifer stems after severe snow-breakage in early 1988 were
studied in autumn 1988 and 1989. The study included twelve stands in the
county of Varmland in central Sweden differing in age and edaphic
conditions. The tops on the ground as well as the remaining rooted stem
stumps of 94 Picea abies and 61 Pinus sylvestris were inspected. The
presence of living branches on the stumps strongly influenced the
incidence of insect attack. All stumps without branches were attacked
during the two-year period, whereas only a few spruce stumps with more
than ten branches and no pine stumps with more than five branches were
attacked. On spruce stumps, the most frequently encountered Coleoptera
were all scolytids, i.e. the species Pityogenes chalcographus, Hylurgops
palliatus, and the genera Polygraphus, Dryocoetes, and Trypodendron. On
pine stumps, Tomicus piniperda was the most common species. Most of the
spruce tops and virtually all pine tops were attacked during the two-year
period, and most of these attacks occurred during the first summer. P.
chalcographus, Dryocoetes, and weevils of the genus Pissodes were the
most frequent Coleoptera on spruce tops. On pine tops, the dominant
insects belonged to the genera Pissodes and Pityogenes. LG EN.█SWED UNIV

Skatulla, U., & Feicht, E. 1992. Studies on the flight behaviour of
Pityogenes chalcographus l. col. Scolytidae and of some other insect
species in connection with pheromone traps by aid of a new electronic
instrument. Anz. Schaedlingskd. Pflanzenschutz. Umweltschutz. 65 (1).
4-7. ab The flight behavior of Pityogenes chalcographus, Nemosoma
elongatum, Epuraea muehli and Karpinskiella pityophthori (a bark beetle
parasite) on pheromone traps depending on temperature and day time was
studied by aid of a new electronic instrument. It could be shown that
each of these species has a characteristical threshold of temperature for
the beginning of its flight activity. Also the day time had a great
influence on the swarm activity of the investigated species. P.
chalcographus even showed significant differences between males and
females in regard of their daily swarm time. LG GE.█BAYER FORSTL

Tunset, K., Nilssen, A.C., & Andersen, J. 1993. Primary attraction in
host recognition of coniferous bark beetles and bark weevils col.
Scolytidae and curculionidae. J. Appl. Entomol. 115 (2). 155-169. ab The
hypothesis of primary attraction in Solytidae and Curculionidae was
tested in the field from late May throughout June 1979 and 1990 with 20
and 14 flight traps, respectively. The traps, designed especially to
exclude the possiblity of secondary attraction, were baited with billets
of Scots pine (Pinus sylvestris) or Norway spruce (Picea abies), and
empty controls, Tomicus piniperda, Hylastes brunneus, Hylastes
cunicularius, Pityogenes bidentatus, Pityogenes chalcographus, Hylobius
abietis and Pissodes pini were significantly more frequently caught in
traps baited with pine or spruce than to empty controls, and the random
flight hypothesis was rejected. The same species, except P.
chalcographus, were also able to discriminate significantly between pine
and spruce before landing. H. cunicularius was exclusively attracted to
traps baited with Norway spruce, whereas the rest of the species
preferred the traps baited with Scots pine. The mechanism used by the
beetles in their host selection is olfactory sensing of chemical
compounds. Host acceptance and nonhost rejecting starts before landing,
implying the ability the perceive plant characterisatics at a distance
(primary attraction). LG EN.█ZOOL DEP, TROMSO MUSEUM, UNIV TROMSO, N-9000

Wegensteiner, R., & Fuehrer, E. 1991. Flight activity of some conifer
bark beetles Coleoptera: Scolytidae in relation to altitude. Anz.
Schaedlingskd. Pflanzenschutz. Umweltschutz. 64 (2). 25-33. ab Results
are provided for insects caught by LINOPRAX-, CHALCOPRAX- or
PHEROPRAX-baited Theysohn traps from March to August 1988 at three
altitudes (380 m, 50 m and 660 m) in the model forest Roslia (Lower
Austria). Different altitudes altered flight activity by one or two
weeks. Trypodendron lineatum appeared first, with Pityogenes
chalcographus appearing at the same time or later than Ips typographus.
Flight began in spring at weekly average temperature-minima/-maxima of
2.5/ for T. lineatum, 3.5/ for P. chalcographus
and 1.7/ for I. typographus. PHEROPRAX had the higher
pheromone-specific allurement for other scolytids. CHALCOPRAX was very
attractive to insects of specific orders (whereby Nemosoma elongatum was
most dominant). The time of bark beetle appearance in spring is discussed
in relation to former examinations in respect to trap trees and pheromone

Wegensteiner, R., Lorbeer, E., & Fuehrer, E. 1989. Additional attraction
of Pityogenes chalcographus l. Coleoptera: Scolytidae to
chalcoprax-baited traps with spruce branches. J. Appl. Entomol. 108 (3).
250-259. ab The beetle catch-quota of Chalcoprax-baited traps, with or
without healthy or damanged spruce bundles were registered over a
timespan of two years. Traps including brushwood proved to be more
effective than those baited with pheromone dispensers only. In addition
to this, the change in the terpene composition of healthy and damaged
brushwood and its effect on the beetle catch-quota were studied during
1987. It was found that the synergistic effect on emitted terpenes on P.
chalcographus differed from spring to summer. LG EN.█INST FUER

Wigger, H. 1994. Response of adult feeding capacity of the bark beetle
predator Nemosoma elongatum L. (Col., Coleoptera Ostomidae) to different
prey quantities in artificial galleries. Anzeiger. Fuer.
Schaedlingskunde. Pflanzenschutz. Umweltschutz. 67 (1). 8-13. Issn
0340-7330. ab Feeding capacity of adult N. elongatum was tested at
different prey quantities of the bark beetle Pityogenes chalcographus L.
(Scolytidae) in artificial galleries. Maximum feeding capacity of 2 P.
chalcographus/day was observed when 3 prey items/day were offered. There
was no increase in prey consumption with further increase of prey
quantities, i. e. there was no functional response. Possible reasons for
restricted feeding capacity are discussed. It is suggested that N.
elongatum adults exhibit a "threshold strategy" which tends to reach an
optimum feeding level instead of a maximum food intake. The consequences
of restricted feeding capacity for limiting bark beetle populations are
discussed. LG GE.█Inst Forstzool, Univ Goettingen, Buesgenweg 3, 37077
Goettingen, GER

Winter, K. 1980. Date of thinning and attack by engraver beetle
Pityogenes chalcographus in spruce stands of the harz mountains west
germany. Z. Pflanzenkr. Pflanzenschutz. 87 (9). 523-532. ab In 5
differently exposed young spruce stands in the Harz Mountains (Lower
Saxony), the correlation between the time of thinning and attack as well
as fly-out of the engraver beetle P. chalcographus was investigated. All
investigated stems were attacked by P. chalcographus. Of the bark area
of all plots 87% were colonized by P. chalcographus, 13% by other
species, i.e., Hylurgops palliatus, Dryocoetes autographus, Polygraphus
polygraphus, Cryphalus abietis, Ips typographus, Xyloterus lineatus,
Pissodes harzyniae and Tetropium sp. I. typographus infested 4% of the
controlled trees, all of them with a diameter of more than 12 cm. Less
than 1% of the controlled trees were attacked by X. lineatus. On the
average, 70% of the bark area of the controlled samples were attacked by
P. chalcographus. The plots of all areas cleared in July and Aug. showed
the lowest attack, i.e, 47% (July) and 49% (Aug.). The values of fly-out
also were lowest in the July and Aug. plots. There is no correlation
between stem diameter and attack, and between attack of the inferior and
the superior part of stem. The development of P. chalcographus depends
on the exposition of an individual stem more than on the exposition of
the area. On southern slopes more bark beetles emerged from shaded stems
than from sunned, on the northern slopes more emerged from sunned than

Wulf, A. 1983. Investigations on the entomo pathogen fungus
beauveria-bassiana parasitizing the spruce wood engraver Pityogenes
chalcographus Coleoptera Scolytidae. Z. Angew. Entomol. 95 (1). 34-46.
ab The fungus penetrates through the cuticle of the bark beetle and
infests muscles and fat tissue. The whole body is completely infested
with saprophytic hyphase subsequent to the death of the beetle and then
the mycelium emerges and sporulates. Larvae and pupae of P. chalcographus
are more sensitive to B. bassiana then the adults. Temperature and the
amount of inoculum have a marked influence on fungal development. The
humidity of the bark is the main factor which effects the development of

Zuber, M., & Benz, G. 1992. Investigations on the temporary sequence of
the flight activities of Ips typographus l. and Pityogenes chalcographus
l. col. Scolytidae by means of the commercial pheromone preparations
pheroprax and chalcoprax. J. Appl. Entomol. 113 (5). 430-436. ab Traps
baited with the commercial population attractants Pheroprax and
Chalcoprax, developed to attract the eight-toothed spruce bark beetle,
Ips typographus (L.), and the six-toothed spruce bark beetle, Pityogenes
chalcographus (L.), respectively were used in 1988-1990 to study the
flight activity of these insects and their attraction by the
preparations. During the three years of investigations, the dates of
emergence and the peaks of the first and the second flight remained
almost unchanged. Whereas the Chalcoprax-baited traps captured almost
only individuals of P. chalcographus, the traps baited with Pheroprax
attracted mainly I. typographus but also a fair number of P.
chalcographus. On the average, 60-75% of the P. chalcographus specimens
caught by the traps baited with Chalcoprax were females, whereas their
percentage in the Pheroprax-baited traps was 20-30% only. Both of these
polygamous bark beetles when captured in the traps baited with the
corresponding specific attractants exhibited a remarkable predominance
of males (50-65%) at the beginning of the first flight period, but this
percentage dropped to 25-35% after 2-3 weeks. However, the predominance
of the males of P. chalcographus captured in Pheroprax-traps either
remained high (65-90%) or did not develop below 60% (except in July
1990). It was shown that the distance of Chalcoprax-baited traps from the
Pheroprax-baited traps could influence the number and the sex-ratio of
P. chalcographus in the Pheroprax-baited traps, the number being
increased in the presence of Chalcoprax. The potential components
responsible for attraction of the males in the traps baited with
Chalcoprax or Pheroprax are suggested. LG GE.█ENTOMOL INST, ETH-ZENTRUM,

Zumr, V. 1982. The data for the prognosis of spring swarming of main
species of bark beetles Coleoptera: Scolytidae on the spruce
picea-excelsa. Z. Angew. Entomol. 93 (3). 305-320. ab The release and
spring swarming of Ips typographus (L.), I. amitinus Eichh. and
Pityogenes chalcographus (L.) was observed in 3 variants of spruce stands
(a clearing, sparse and dense spruce stand). The release and spring
swarming of the bark beetles was observed using traps and special trap
devices placed into spruce logs. Spring swarming was observed beginning
with the insects leaving overwintering sites. The effect of maximum day
temperature (tmax) of the soil, of the phloem and of air on the release
and swarming was demonstrated. On the basis of measured temperature data,
release and spring swarming was determined by the sum of effective
temperature (.SIGMA.Et). When leaving overwintering sites, the bark
beetles appeared 1st from suspended logs in the margin of the stand and
from the soil litter in photoelectors in the clearing. Beetles left the
phloem of the suspended logs in the margin of the stand 14-16 days soon
in comparison to the photoelectors in the clearing and 11-13 days sooner
than inside the stand in comparison with photoelectors in the sparse
spruce stand. I. typographus spring attack started with the trap logs and
landings in the clearing, 1-2 days later in the sparse stand and 3-5 days
later in the dense spruce stand in comparison with the clearing. I.
amitinus and P. chalcographus spring attack occurred 0-1 day later in the
sparse forest and 0-2 days later in the dense spruce forest. The swarming
conditions for each of the 3 bark beetle species are discussed. LG

Zumr, V. 1988. The effectiveness of the aggregation pheromone chalcoprax
in the control of spruce wood engraver Pityogenes chalcographus l.
Coleoptera Scolytidae. Lesnictvi. Prague. 34 (6). 489-498. ab The
effectiveness of the aggregation pheromone Chalcoprax was studied as far
as for the control of spruce wood engraver Pityogenes chalcographus (L,)
in four variants of spruce stands located in Southern Bohemia
(Czechoslovakia). Using Chalcoprax, the following features were specified
in greater detail: the beginning of swarming and flitting of spruce weed
engraver beetles, sex ratio and attractivity for the natural enemies of
the given pest. The beginning of spruce wood engraver swarming and
flitting is influenced in a decisive manner by the temperature. The
swarming begins at the avreage daily air temperature (tmax) of Intensive flitting begins at tmax =,
with the maximum at tmax = Out of the total number of
spruce wood engravers that had been trapped, 39.9% of male beetles were
caught in the pheromone traps; the total of 212 980 spruce wood engraver
individuals were caught in twenty pheromone traps in all variants of
spruce stands. The highest numbers of spruce wood engravres were caught
at a clearing (variant C) -110 300 individuals and the lowest numbers
along the border of spruce pole stand -9040 individuals (variant D). As
to the predators, Nemosoma elongatum (L.) was repersented in the highest
numbers-60.1%; the following percent were 18.8 in Nudobius lentus Grav.,
12.3 in Thanasimus formicarius (L.) and 8.8 in Paromalus parallelopipedus
Hrbst. Using the aggregation pheromone, not only the occurrence of spruce
wood engraver can be well monitored in the forest stand but also of the
natural enemies of this beetle, and its ecological demands can be
invesigated in greater detail. LG CZ.█ENTOMOL USTAV CSAV, BRANISOVSKA 31,

Zumr, V. 1992. Attractiveness of introduced conifers to xylophagous
beetles and their acceptance. J. Appl. Entomol. 113 (3). 233-238. ab The
incidence of subcorticolous and xylophagous beetles (Coleoptera) on 11
conifers (including introduced species) was tested in a model area in
southern Bohemia by means of tree and window traps. The most infested
conifers were spruces and pines. The highest dominance among the bark
beetles was found in Pityogenes chalcographus (L.) averaging 48% in all
the conifers, and Xyloterus lineatus (Ol.) averaging 18%. Ips typographus
(L.) was dominant on the spruces (Picea spp.) (average dominance 41%),
Pityokteines spinidens Reitt. on firs (Abies spp.) (average dominance
47%) and Myelophilus piniperda (L.) on pines (Pinus spp.) (average

Zumr, V., & Soldan, T. 1981. Reproductive cycles of Ips typographus Ips
amitinus and Pityogenes chalcographus Coleoptera: Scolytidae. Acta.
Entomol. Bohemoslov. 78 (5). 280-289. ab Histological changes of ovaries
and testes and fecundity of the bark beetles I. typographus, I. amitinus
and P. chalcographus were investigated under laboratory conditions.
Although development of the gonads is completed in adults before feeding
starts, the terminal phases of gametogenesis (spermateliosis,
vitellogenesis) occur during maturation feeding. Forming of vitellarium
and egg production is repeated during regeneration feeding in females.
There are 3 reproductive cycles in I. typographus, which inhabits warm
localities at lower altitudes, living on the lower (tree) stem portion
below the crown. Two cycles occur in I. amitinus inhabiting relatively
cold mountain localities and the same host-plant niche as I. typographus,
and a single reproduction in P. chalcographus which inhabits both lowland
and mountain localities but colonizes a different niche (spruce
branches). Fecundity of these species is 30-50 eggs /reproductive cycle.
Different types of reproductive strategies are discussed. LG