New Zealand Entomologist 24: 79–85 (December 2001)

Observations of the flight activity of Hylastes ater and Hylurgus ligniperda

(Curculionidae: Scolytinae) in Pinus radiata forests in the central North Island,
New Zealand
Stephen D. Reay1,3 and Patrick J. Walsh2,4
1 School of Forestry, University of Canterbury, Private Bag 4800, Christchurch, N.Z.
2 FRASS, PO Box 1041, Rotorua, N.Z. 79
3 Current address and address for correspondence: Department of Biological Sciences,
The University of Waikato, Private Bag 3105, Hamilton, New Zealand. sreay@waikato.ac.nz
4 Current address: Forest Management Group, School of Engineering, Galway Mayo Institute of Technology,
Dublin Road, Galway, Ireland.

Abstract both observed three overlapping generations on H.

Hylastes ater and Hylurgus ligniperda adults were
trapped using Lindgren type funnel traps in Pinus
radiata forests in the central North Island, New
Zealand, from October 1998 through April 2000,
to observe the annual flight activity of both species.
The results of the trapping programme indicated
that H. ligniperda might be bivoltine and H. ater uni-
voltine. Since the introduction of H. ligniperda into
New Zealand, the peak period of H. ater flight activ-
ity has shifted from mid-summer to late autumn,
while H. ligniperda is the dominant species during
the spring and summer months. Therefore, most of
the habitat created by the harvesting of P. radiata is
likely to be colonised by H. ligniperda. In the past H.
ater was the dominant bark beetle colonising this
material. These changes in seasonality may have
resulted in a reduction in damage to P. radiata
seedlings in some areas.
Keywords: Hylastes ater, Hylurgus ligniperda, bark
beetles, colonisation activity
Introduction
The pine bark beetle, Hylastes ater (Paykull), of
European origin, was first discovered in New
Zealand in 1929 (Clark 1932). It feeds on the cam-
bium and inner bark of stumps and logs of Pinus
species. It is widely distributed throughout New
Zealand wherever pines are grown. A major cause
of economic damage by H. ater in New Zealand, is
when adults emerge from Pinus radiata D. Don
stumps and feed on seedlings. This can result in
widespread seedling mortality in certain areas
(Reay 2001). During these feeding attacks H. ater
can vector sapstain fungi to seedlings (Reay et al. in
press).
Clark (1932) and Crowhurst (1969) have given a
general account of H. ater in New Zealand. They
ater in the field. Crowhurst (1969) described the
development time of H. ater to be as short as six
weeks from egg to adult during the warmest sum-
mer months. Eggs laid at other times may take up to
ten months to develop to maturity, due to larval
diapause during winter (Crowhurst 1969).
Hylurgus ligniperda L., also of European origin,
was first discovered in New Zealand in 1974 (Bain
1977) and has since become widely established.
Like H. ater, H. ligniperda breeds in stumps, logs and
roots of harvested pine species. In New Zealand it is
mainly found in P. radiata (Bain 1977). The adult
beetles superficially resemble H. ater, but are larger
and are a reddish-brown colour. The rates of devel-
opment of H. ligniperda and H. ater are similar (Bain
1977, Reay 2001). Unlike H. ater, adult H. ligniper-
da does not attack seedlings following emergence
(Bain 1977, Reay 2001).
Differences in the recorded flight activity of
H. ater and H. ligniperda give an indication as to how
these two species interact with each other. Adult
flight represents the period during which new
adults emerge from brood material, search for and
colonise new breeding sites.The primary objectives
of this study were to monitor the flight activity of
these two species to determine whether there were
differences in flight activity. Secondly, the flight
activity data were examined to identify possible
environmental cues relating to activity.
Methods
In October 1998, 19 sites were selected in
Kaingaroa (Fletcher Challenge Forests) and Kinleith
(Carter Holt Harvey Forests) Forests. Sites were
selected to include a variety of site types (e.g.
aspect, slope, altitude, soil types etc.), and occurred
across the two forest estates. Sites were selected and
 New Zealand Entomologist 24: 79–85 (December 2001)
the trapping programme commenced within one
week of harvesting activities.
Flight activity, used as an index of colonisation at
each site, was assessed using five-funnel Lindgren
type traps (Lindgren 1983). A 100 ml glass test-
tube, containing 80 ml of raw turpentine was placed
80 in each plastic collection jar. Raw turpentine has
been described as an attractant for Hylastes species
(Gara & Vité 1962, Löyttyniemi et al. 1988, Phillips
1990). An assumption was made that the raw tur-
pentine would attract both H. ater and H. ligniperda.
However when tested with a number of other
potential attractants, the turpentine used in this
study was found not to be attractive to either H. ater
or H. ligniperda (Reay 2001). Therefore, any H. ater
or H. ligniperda individuals caught during this study
were due to accidental catches, or visual attraction
to the traps.While more individuals may have been
caught if a more effective attractant had been used,
the number of H. ater and H. ligniperda individuals
caught is assumed to reflect the relative activity of
each species.
Two traps, approximately 80m apart, were erect-
ed in each site. Traps were located 30m into each
site to avoid any potential edge effects. Traps were
serviced weekly from 1 October 1998 through 5
May 1999.The turpentine was replaced during each
Fig. 1. Mean number of H. ater and H. ligniperda individuals caught per trap during the period
1 October 1998 through 5 May 1999, showing the potential influence of atmospheric pressure
on the mean weekly catch.
trap service. Traps were left in a few sites to moni-
tor flight activity during winter months.These traps
were serviced once each month during winter. In addi-
tion to the 19 sites first selected, two further groups of
sites were added to the 1998/99 trapping programme
to cover different harvesting times. Four sites were
added in January 1999 and nine sites were added in
March 1999 giving a total of 32 trapping sites.
In October 1999 traps were shifted to 17 new
sites. As with the previous summer, three additional
sites were added later in the summer period (March
2000). Unlike the previous summer, all traps were
serviced at fortnightly intervals.While there was an
associated loss in the sensitivity of the data collect-
ed, the large effort required for weekly servicing
over such a long period was not possible during the
1999/2000 trapping period.
The National Institute of Water and Atmosphere
supplied climatic data for the study area during the
period of study. These data were recorded at the
Rotorua Airport climate station (B86133). While
there was likely to be variation in climate between
the Rotorua Airport and the sites in this study, it was
not feasible to collect climate data within the forest
estates. The data were used to investigate the influ-
ence of relative changes in climate on flight activity
over time.

Fig. 2. Mean number of H. ater and H. ligniperda individuals caught per trap during the period
4 October 1999 through 20 April 2000.
Results
The flight activity of H. ater and H. ligniperda fluc-
tuated over the first seven-month trapping period
from 1 October 1998 through 5 May 1999 (Fig. 1).
Flight activity during spring and early summer
(7 October to mid-November) was dominated by
H. ligniperda. A second period of increased activity
by H. ligniperda began during January 1999 and con-
tinued for approximately three months.This period
of activity by H. ligniperda did not peak as sharply as
that observed during spring. However, it did contin-
ue for a longer time. H. ligniperda flight activity
declined at the end of summer.
During most of the spring and summer months,
the flight activity of H. ater was very low in compar-
ison to H. ligniperda (Fig. 1). There were slight
increases in activity, which coincided with H. lig-
niperda flight. At the end of the 1998/99 summer
period the activity of H. ater increased substantially
(during March) before sharply peaking in the latter
part of April.This sharp peak in flight activity lasted
for a short time (2-3 weeks). Following this, num-
bers of both species caught fell dramatically.The activ-
ity of both species during the 1999 winter period was
minimal (few or no individuals were caught).
Flight activity during the second trapping period
4 October 1999 through 20 April 2000 (Fig. 2) was
Reay & Walsh: Scolytine flight activity in pine forest
81
consistent with that observed the previous summer
(Fig. 1).The activity of H. ligniperda peaked in spring
as with the previous year. There appeared to be a
greater amount of H. ater activity at this time com-
pared with the previous year. However, spring activ-
ity was still dominated by H. ligniperda. Following
this, the numbers of both species fell to relatively
low levels.At the end of January there was a marked
increase in H. ligniperda abundance, which peaked at
the end of February before falling away during
April.The flight activity of H. ater increased slightly
at the end of January 2000, before increasing
sharply during April (Fig. 2). This sharp increase in
activity was consistent with that recorded for the
previous summer.
The potential effects of temperature and atmos-
pheric pressure were investigated using the flight
activity data from the 1998/99 period.The data col-
lected at fortnightly intervals during the
1999/2000 period were not sensitive enough to
determine meaningful relationships. H. ligniperda
activity appears to have been influenced by peaks in
atmospheric pressure (Fig. 1). The three largest
peaks in H. ligniperda flight activity corresponded
with the three most significant peaks of mean week-
ly atmospheric pressure. Mean weekly temperature
appears not to be related to H.ligniperda flight (Fig. 3).
 New Zealand Entomologist 24: 79–85 (December 2001)
82
Fig. 3. Mean number of H. ater and H. ligniperda individuals caught per trap during the period
1 October 1998 through 5 May 1999, showing the potential influence of temperature on the
mean weekly catch.
A relationship between H. ater flight activity and
atmospheric pressure was not apparent during the
1998/99 summer period (Fig. 1). This may be a
function of the low levels of H. ater activity during
this period.The peak of H. ater flight activity during
the latter part of April 1999 follows a sharp decline
in temperature and atmospheric pressure, indicating
a potential relationship.
Discussion
In general, the activity of each species was rea-
sonably consistent between the two years, although
the periods of peak activity of H. ater and H. lig-
niperda were at different times.The flight activity of
H. ater was characterised by one dominant period of
flight just before winter. This indicates that, despite
earlier observation (Crowhurst 1969), H. ater is
now essentially univoltine in New Zealand. This
period of H. ater activity was more intense com-
pared with periods of H. ligniperda activity. Outside
of this period of activity, populations of active adult
H. ater were low. The flight activity of H. ligniperda
during the summer periods was characterised by
two periods of peak activity.Therefore, H. ligniperda
may be bivoltine in New Zealand. The dominant
periods of adult flight activity of the two species
were separated in time, although they often inhabit-
ed the same stumps (Reay 2001).
It is possible to suggest hypotheses regarding the
adult flight activity and the relationship of this activ-
ity to the lifecycle of the two species. The flight
activity of the two species represents a search for
new breeding (and food) material. It is during these
periods of flight that new sites are colonised and
new generations are established. The H. ligniperda
adults trapped in spring were most likely to have
overwintered as late instar larvae or as adults. The
second period flight activity was most likely of
adults resulting from oviposition during spring
colonisation. Larvae resulting from H. ligniperda
colonisation and breeding during the mid-summer
period may develop through to adult beetles that
may emerge and colonise new habitats before the
end of summer, or perhaps overwinter as adults.The
offspring produced by colonising adults during the
later part of the second activity period of H. lig-
niperda would have been unable to develop fully
before winter. These individuals would have over-
wintered as late instar larvae and completed their
development prior to the spring emergence.
In contrast, a period of emergence and the coloni-
sation of new habitats during autumn dominated the

breeding and development activity of the H. ater
populations.The offspring produced by these adults
would have overwintered as eggs and/or early instar
larvae. These larvae presumably continued their
development during the following summer and
their emergence coincided with the period of peak
H. ater activity at the end of that summer.
Both species co-existed in the same breeding
material (Reay 2001), yet had discrete and identifi-
able periods of peak activity. A similar situation
occurs in South Africa, although activity occurs dur-
ing different periods to New Zealand, with Hylastes
angustatus (Herbst) and H. ligniperda (Tribe 1990,
1991a,b). The activity of H. angustatus is charac-
terised by one sharp peak of activity per year, dur-
ing a period of spring emergence in September and
October (Tribe 1990). This period of activity
accounted for 95% of the annual catch of H. angus-
tatus. In South Africa, H. ligniperda has a peak of
activity during April and May, but this accounts for
only 37% of the annual catch (Tribe 1991a). The
flight activity of this species is variable throughout
the year (Tribe 1991b).
These observations of H. ater in New Zealand are
similar to those made by Tribe (1991b) of H. angus-
tatus in South Africa with respect to an annual peak
in activity. However, where the activity of H. angus-
tatus peaks in spring in South Africa (Tribe 1990),
the peak activity of H. ater in New Zealand was in
autumn. Tribe (1990) suggests that the flight of H.
angustatus be triggered by a combination of temper-
ature and rainfall thresholds. Most flight occurs
when there is adequate moisture and the tempera-
ture is between 12-16ºC. In New Zealand, the peak
of H. ater flight occurred when the mean tempera-
ture was approximatly 13ºC (following a sharp
decline from approximately 16-18ºC) and following
a decrease in atmospheric pressure.
The peak period of H. ligniperda in South Africa is
during autumn, during periods of high moisture and
low temperatures (a mean temperature of approxi-
mately 15ºC) (Tribe 1991a). In contrast, H. ligniper-
da was most active in spring and summer in New
Zealand, although mean temperatures during these
periods are similar to the optimum temperatures
described for South Africa during autumn (Tribe
1991a,b). The variability of H. ligniperda flight in
New Zealand is consistent with that observed in
South Africa (Tribe 1991b). In France, H. ligniperda
activity is more representative of the New Zealand
situation, with the dominant period of activity in
Reay & Walsh: Scolytine flight activity in pine forest
spring and a smaller peak in autumn (Fabre & Carle
1975).The activity peaks of each species in a differ-
ent season indicates that different species may be
responding to different environmental cues (Tribe
1991b). Further research is clearly necessary to
accurately determine any role that climatic variables
might play in the life cycle of both species in New 83
Zealand.
Clark (1932b) and Crowhurst (1969) indicated
that there were three overlapping broods of H. ater
per year in New Zealand. However, the establish-
ment of H. ligniperda in New Zealand has resulted
in the dominance of this species during the spring
and summer months. Ciesla (1988) reported that
H. ligniperda is dominant over H. ater in Chile.
Observations by Tribe (1990, 1991a,b) indicate that
H. ligniperda is the most dominant species in South
Africa. One effect of the introduction of H. ligniper-
da has been to influence the activity of H. ater dur-
ing the summer period. For example, the reduction
of H. ater populations during this period may be due
to competition by H. ligniperda. Population dynam-
ics of H. ater were difficult to observe when the
species was present at low levels.
Crowhurst (1969) reports that overwintering H.
ater larvae enter a true diapause, which is broken in
early January. Larvae from eggs laid by H. ater dur-
ing early spring were also observed to pupate in
January. Only larvae derived from eggs laid in
March and April enter diapause (Crowhurst 1969).
The larval resting stage prior to pupation is up to
two weeks for larvae from eggs oviposited at all
other times of the year (Crowhurst 1969).
Crowhurst (1969) suggests that the function of this
larval diapause mean that overwintering larvae and
those larvae that develop during the summer reach
maturity at the same time, during January. Thus
developing larvae would reach maturity during the
warmest period of the year, with larval offspring
produced by these adults developing when condi-
tions are optimal (Crowhurst 1969). Observations
by Crowhurst (1969) indicate that the peak of H.
ater flight activity was likely to have occurred during
January prior to the introduction of H. ligniperda.
Following Crowhurst’s (1969) observations of H.
ater, H. ligniperda was accidentally introduced into
New Zealand. A short research note was published
on H. ater in New Zealand four years following the
arrival of H. ligniperda (Milligan 1978). Milligan
(1978) reports the swarming of H. ater in
September and October, and again in April and May.
 New Zealand Entomologist 24: 79–85 (December 2001)
It is conceivable that the trapping programme in this
study missed an earlier H. ater spring flight. There
may be evidence of this in a small peak of H. ater
activity in mid-October (Figure 2). However, traps
left in a few sites during the winter period caught
minimal numbers of both species. As H. ligniperda
84 flight activity peaks during the mid-summer period,
it is likely that the delayed emergence by H. ater is in
response to competition from H. ligniperda. While
both species are able to occupy the same habitat
(Reay 2001), a competitive advantage by H. ligniper-
da over H. ater is most apparent during the summer
months when populations of H. ligniperda dominate.
Presumably, H. ater would have colonised all habitat
created during the summer months prior to the
introduction of H. ligniperda into New Zealand.
Breeding habitat resulting from harvesting during
spring and summer is now dominated by H.ligniperda.
In the natural forest, access of both H. ater and H.
ligniperda to breeding material is likely to be limited
during endemic conditions (i.e. an occasional tree
fall) (Raffa & Berryman 1983, Tribe 1991b,
Kirkendall et al. 1997). Secondary or non-aggres-
sive bark beetles are very successful at exploiting
such habitat (Tribe 1991b, Wilson & Day 1995,
Örlander et al. 1997, Rieske & Raffa 1999). Large-
scale disturbances, such as wind-throw, would result
in an increase of breeding habitat.These bark beetles
would quickly exploit this increase in available sub-
strate and populations would reach epidemic pro-
portions, until the increased supply of breeding
material was exhausted (Rudinsky 1962, Berryman
1972, Christiansen et al. 1987). In New Zealand P.
radiata forests, large-scale harvesting activities are
continuous year round.A consequence of this is that
enough breeding habitat is created for populations
levels to constantly reach epidemic proportions.
Observations made during this study indicate that
the vast majority of stumps created during harvest-
ing activities were colonised by either H. ater or H.
ligniperda, or both species. Historically, all breeding
material would have been occupied by H. ater in
New Zealand. The introduction of H. ligniperda has
probably had some influence on the pest status of H.
ater. H. ligniperda is the dominant species and
colonises the majority of habitat created during har-
vesting activities. Consequently, the abundance of
H.ater may have decreased.Therefore, it may be rea-
sonable to suggest that the impacts of H. ater on the
forest industry will also have reduced.
Acknowledgements
We would like to thank Carter Holt Harvey
Forests, Fletcher Challenge Forests, Scott Downs,
Mark Forward and the staff at the School of Forestry
for their assistance during this project, and to Nod
Kay for his useful comments during this study.
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