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Ansari, M.A., Casteels, H., Tirry, L. & Moens, M. (2006). Biology

of Hoplia philanthus (Coleoptera: Scarabaeidae), a new and
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 POPULATION ECOLOGY

Biology of Hoplia philanthus (Col., Scarabaeidae, Melolonthinae):

A New and Severe Pest in Belgian Turf
MINSHAD ALI ANSARI,1,2 HANS CASTEELS,1 LUC TIRRY,3 AND MAURICE MOENS1,3

Environ. Entomol. 35(6): 1500Ð1507 (2006)

ABSTRACT We studied the adult behavior, oviposition, and larval and pupal development of Hoplia
philanthus Füessly (Scarabaeidae: Melolonthinae) in the Þeld and laboratory in Belgium. Adult
emergence was observed in the Þrst week of June 2000, 2001, and 2002, peaked ⬇2 wk later, and
continued until the last week of the month. The average sex ratio of emerging adults was 1.3:1 (male
and females) during the collection period. Adults were observed feeding mainly on leaves of Betula
utilis variety jacquemontii (B. u. Doorenbos) and Carpinus betulus L. During the day, H. philanthus
adults were most active at 1400 hours (GMT ⫹ 1.00). Oviposition started in the last week of June and
lasted until the end of July. Each female deposited ⬇25Ð 40 white eggs at a depth of 10 Ð15 cm in soil.
Eggs hatched 28 ⫾ 5 d after being laid at an average monthly soil temperature of 18.1⬚C. Three larval
instars could be discerned by head capsule width; all larval instars fed on grass roots. The Þrst-instar
larvae were found in the last week of July 2000; second instars appeared mostly in the second week
of September 2000 and could be found until May 2001. Third instars were found from the second week
of June 2001 until April 2002. Pupae could be found from the Þrst week of May until the end of the
month. The duration of the pupal stage was 28 ⫾ 5 d at an average monthly soil temperature of 16.5⬚C.
According to these observations, H. philanthus has a 2-yr life cycle.

KEY WORDS Hoplia philanthus, Scarabaeidae, turfgrass, white grub

A number of scarabaeid beetles, such as Hoplia philan- various plants, including turf, herbaceous ornamen-
thus Füessly (Scarabaeidae: Melolonthinae), the Eu- tals, nursery trees, and shrubs (Ansari 2004). Heavily
ropean cockchafer, Melolontha melolontha L., the gar- infested turf can often be lifted from the soil or rolled
den chafer, Phyllopertha horticola L., the summer back like a carpet because the bigger part of the root
chafer, Amphimallon solstitialis L., and the brown cha- system has been consumed. Secondary damage from
fer, Serica brunnea L., are pests in Belgium. The four foraging insectivorous birds and mammals often
latter species are present at low population densities causes further disruption to the turf surface. Adults
and rarely cause economically important damage. H. feed on the foliage of various plants but generally
philanthus, sometimes called the Welsh chafer, how- cause little damage.
ever, has become an economically important pest of Little information is available on the biology and
turf and pastures (Ansari et al. 2003b). Its presence distribution of H. philanthus. To develop a control or
was Þrst noted during a survey in the surroundings of management strategy for H. philanthus, more infor-
Ghent (Casteels and De Clercq 1998). Since then, it mation is needed on its life cycle and ecology. There-
has spread continuously and has reached pest status in fore, we observed the species in the Þeld and in the
a large part of Flanders. Elsewhere in Europe, prob- laboratory.
lems of H. philanthus are reported from Germany
(Ehlers 2000), Poland (Bunalski 1995), Spain (Baraud
Materials and Methods
1992, Micó et al. 2003), The Netherlands (Vlug 2001),
and the United Kingdom (Gratwick 1992). Observation Site. The Þeld studies were conducted
At the moment, millions of dollars are spent each in the period 2000 Ð2002 in a heavily infested lawn
year for the control of the white grubs and for renew- (⬇0.5 ha) at the Research Station for Ornamental
ing or replacing damaged turf (Klein et al. 2000). The Plants (PCS) at Destelbergen, province of East
larvae of H. philanthus feed on underground parts of Flanders. The lawn was composed of a mixture of
perennial ryegrass (Lolium perenne L.) and bentgrass
1 Institute for Agricultural and Fisheries Research, Burg. Van Gan- (Agrostis tenuis Sibth.) on a sandy soil (89.9% sand,
sberghelaan 96, B-9820 Merelbeke, Belgium. 6.6% silt, 3.5% clay; 3.02% organic matter; pH 4.52).
2 Corresponding author: Department of Biological Science, Uni-
Ornamental trees [Carpinus betulus L., Betula utilis
versity of Wales, Swansea SA2 8PP, UK (e-mail: m.a.ansari@
swan.ac.uk).
variety jacquemontii (B. u. Doorenbos) and Liqui-
3 Laboratory of Agrozoology, Department of Crop Protection, Gh- dambar styraciflua L.], shrubs [Thuja plicata Donn ex
ent University, Coupure Links 653, B-9000 Gent, Belgium. D. Don, Picea sp., Cupressocyparis leylandii (Jacks.

0046-225X/06/1500Ð1507$04.00/0 䉷 2006 Entomological Society of America

December 2006 ALI ANSARI ET AL.: BIOLOGY OF H. philanthus 1501

Fig. 1. Environmental conditions at observation site (lawn consisted of perennial ryegrass and bentgrass). The monthly
mean maximum, minimum, and average air temperature and the average soil temperature at 10, 20, and 50 cm depth were
recorded over the observation period or obtained from the local weather station. Daily rainfall totals (cm).

Dallimori) Dallimori, Viburnum plicatum Thunb., and Field Population. To estimate the initial population
V. opulus L.], and bushes (Rhododendron sp. and Erica density of larvae at the observation site, for each ob-
carnea L.) bordered the lawn. Temperature at the site servation date (5, 15, and 26 June 2000), 15 plugs (30
varied greatly within a day. The monthly mean max- by 30 by 20 cm) were randomly taken by cutting
imum, minimum, and average air temperature and the through the turf and thatch on three sides. Grubs were
average soil temperature at 10, 20, and 50 cm depth carefully collected from the upper 5Ð20 cm of soil,
were recorded over the observation period at the below the thatch layer, identiÞed to the genus level by
study site or obtained from the local weather station means of the raster pattern on the ventral side of the
(Fig. 1). pygidium (in the genus Hoplia, spines are irregu-
1502 ENVIRONMENTAL ENTOMOLOGY Vol. 35, no. 6

larly implanted) (Niklas 1974). The grubs were put To determine the oviposition period and egg hatch,
back in the soil plug, which was returned to its Þve soil samples (5Ð10 cm depth and 10 cm diameter)
original place. were taken around entrance holes and searched for
Adult Emergence. Adult beetles were identiÞed to eggs every 3Ð 4 d. Eggs were collected from the soil and
species level using identiÞcation keys by Freude et al. counted. Simultaneously, the oviposition was ob-
(1969). Counting of emergence holes was evaluated as served at the observation site by taking sod of grass as
a method to determine adult emergence. Although described in the section on the Þeld population. The
holes made by earthworms were also present in the soil was inspected for eggs at 4-d intervals in June 2000,
observation site, the emergence holes of H. philanthus 2001, and 2002.
could be distinguished from the previous one by the Eggs. Egg hatch was monitored in the cage exper-
fact that the adults scattered soil on one side of the iment. Groups of 10 eggs were collected from soil with
emergence hole, whereas earthworms piled up their a camel hair brush, transferred to petri dishes (9 cm
casting all around holes. Emergence holes could also diameter), covered with moist soil, closed with a lid,
be distinguished from entrance holes made by the and kept in the dark at 18 ⫾ 2⬚C and 90% RH. Care was
female beetles for oviposition, by observing how soil taken to minimize mortality caused by manipulation.
was scattered around the hole. Emergence holes were The number of hatched or collapsed eggs was re-
always clean on one side of the hole with soil scattered corded every day. The egg sizes were measured upon
around the other side, whereas entrance holes were collection. Observations started on 24 June 2002, i.e.,
clogged with soil and had some soil scattered all when the Þrst adults laid eggs under the sod of grass.
around the hole. Larval Development. To study the larval develop-
Twenty-Þve (1.0 by 1.0 m) observation plots were ment, 10 plots (2.0 by 2.0 m) separated by 0.5-m
randomly established on the lawn. On each observa- buffers were established at the observation site in June
tion date, Þve plots were randomly selected, and the 2000. One soil plug (15 by 15 by 10 Ð50 cm range of
total number of emergence holes in these plots was depths) was taken from each of the 10 plots with a
counted. Observations were done on 2, 6, 12, 20, and shovel at bimonthly intervals from June 2000 until July
30 June 2000; 1, 8, 15, 22, and 29 June 2001, and 4, 11, 2002. In each of the plots, larvae from different size
18, 25, and 30 June 2002. were present, indicating the presence of overlapping
generations. On each sampling date, 25 individuals
Adult Activity. Trees and shrubs located at the edge
were collected from each larval size that was encoun-
of the study site (25 by 15-m area) were observed for
tered and brought to the laboratory. Head capsule
the presence of H. philanthus adults on 14, 15, and 16
width was measured from widest point to widest point
June 2000, 2001, and 2002. Observations were made
under dissecting microscope to differentiate the larval
from 1000 to 2000 hours (GMT ⫹ 1). All observed
instars. The number of larvae sampled varied accord-
adults were collected by hand net and stored per time
ing to the period and the instars present. The location
period of 2 h in separate petri dishes (9 cm diameter) where each larva was found (depth below the thatch
containing 70% ethanol, which were wrapped in layer), larval migration in the soil, overwintering, and
ParaÞlm. In the laboratory, adults were separated by natural larval mortality were recorded; soil tempera-
sex to get an estimate of the sex ratio. The number of ture at 10, 20, and 50 cm depth was measured (Fig. 1).
males and females per square meter was recorded per Statistical Analysis. The number of H. philanthus
time period. adults collected at different times from the observa-
For a more exact determination of the sex ratio, 150 tion site was compared by analysis of variance
pupae of H. philanthus were collected from the ob- (ANOVA). Means were separated with TukeyÕs test
servation site in the Þrst week of May 2005 and trans- (SPSS 2003). To compare adult emergence between
ferred to plastic containers (11 cm diameter by 8 cm years, the number of emergence holes observed in
height) Þlled with 350 g sandy soil (14% moisture). different plots on the observation site was averaged to
Ten pupae were placed in each container and were determine mean adult emergence/m2.
kept at 22 ⫾ 2⬚C. The development of pupa into adults
was observed at 3-d intervals.
Oviposition. To determine the oviposition period,
7Ð10 d after Þrst adult emergence, adults were col- Results and Discussion
lected from the turf and shrub border at the obser- Field Population. The density of H. philanthus lar-
vation site (on 7 June 2000, 11 June 2001, and 10 June vae in the Þeld varied from 75 to 450/m2 at different
2002). Adults (n ⫽ 20) were placed in each of three months of the year. At the start of the observations
cages (1.5 by 1.5 by 1.5 m), which were open at the (early June 2000), different larval instars and adults
bottom, placed on the ground, and Þlled with an ad- were found; eggs were detected only at the end of
ditional layer of 20 cm of soil. The cages were closed June. IdentiÞcation of the grubs to the genus level
at the top and the four sides by an iron mesh to prevent showed that only larvae of the genus Hoplia were
adult escaping. Commercially produced sod of rye- present. All adult beetles were identiÞed as H. philan-
grass (L. perenne), 3.0 Ð 4.0 cm thick, were placed on thus.
the soil surface and watered. In each cage, a 1.0-m- Adult Emergence. Overall, we did not Þnd signiÞ-
high, pot grown C. betulus was administered as food cant differences in total number of emergence holes
source for the adults. among the 3 yr (F ⫽ 0.657; df ⫽ 2,42; P ⬎ 0.524).
December 2006 ALI ANSARI ET AL.: BIOLOGY OF H. philanthus 1503

Fig. 2. Emergence of H. philanthus adults from a lawn at different days in June 2000, 2001, and 2002. Bars with different
letters represent means that are signiÞcantly different based on Tukey test (P ⬍ 0.05); error bars represent SEM.

However, there were signiÞcant differences between females (F ⫽ 0.62 df ⫽ 2,51; P ⬍ 0.542) was not
the number of emergence holes at different dates in signiÞcantly different among the years in 2000, 2001,
June 2000 (F ⫽ 45.9; df ⫽ 4,10; P ⬍ 0.001), June 2001 and 2002. The timing at which males (F ⫽ 6.0; df ⫽
(F ⫽ 22.2; df ⫽ 4,10; P ⬍ 0.001), and June 2002 (F ⫽ 5,12; P ⬍ 0.005) and females (F ⫽ 3.2; df ⫽ 5,12; P ⬍
29.8; df ⫽ 4,10; P ⬍ 0.001). In 2000, 2001, and 2002, the 0.046) were collected was signiÞcantly different in
Þrst H. philanthus adult emergence hole was observed 2000. However, males (F ⫽ 2.80; df ⫽ 5,12; P ⬎ 0.067)
on 2 June, 1 June, and 4 June, respectively. Each year, and females (F ⫽ 0.81; df ⫽ 5,12; P ⬎ 0.565) were not
adult emergence increased until 12, 15, and 18 June signiÞcantly different in 2001. In 2002, the numbers of
and decreased until completion by the end of the collected males were signiÞcantly different at differ-
month (Fig. 2).
ent hours of the day (F ⫽ 4.4; df ⫽ 2,12; P ⬍ 0.017), but
Because scarab beetles often tend to reinfest their
the number of females was not signiÞcant (F ⫽ 1.7;
site of origin because of favorable conditions (envi-
ronmental and biological) for oviposition, the obser- df ⫽ 2,12; P ⬍ 0.20; Fig. 3).
vation of adult emergence sites and patterns might be The number of adults collected in the period 14 Ð16
useful to predict where turf damage by the larvae of June in 2000, 2001, and 2002 was plotted as the mean
the next generation is most likely going to increase. A number of adults collected every 2 h, with the repli-
similar conclusion with respect to the oriental beetle, cations being the 3 d in the month (Fig. 3). The
Exomala orientalis (Waterhouse), in golf courses was greatest number of adults was collected at 1400 hours.
drawn by Choo et al. (2002). In general, more males than females were collected.
Adult Activity. Overall, the number of males caught Of 298 adults collected, 67.1% were males and 32.9%
per square meter (F ⫽ 0.34; df ⫽ 2,51; P ⬍ 0.710) and were females.

Fig. 3. Average number of H. philanthus adults collected on shrubs and trees surrounding a lawn. Adults were collected
every 2 h during the period 14Ð16 June 2000, 2001, and 2002.
1504 ENVIRONMENTAL ENTOMOLOGY Vol. 35, no. 6

Of 150 pupae collected from the observation site, and 22.7⬚C in June and July, the mean minimum tem-
143 adult beetles emerged from rearing containers, perature was 12.8, 13.8, and 13.6⬚C, and the average
and 58% were males and 42% were females, which is mean temperature was 17.1, 17.8, and 18.1⬚C for 2000,
a sex ratio of ⬃1.3:1. The difference with the sex ratio 2001, and 2002, respectively. During the different
observed in the Þeld might be caused by a different years, the data on oviposition obtained at the obser-
migratory behavior between the sexes. vation site were similar to those observed in the cages.
Observation of adult activity throughout the study Females were sexually mature after 7Ð10 d of feeding
showed that emergence of adults mainly occurred and ßew to deposit eggs in soil.
during the day. Temperature conditions inßuenced Eggs. Eggs are glossy, milky-white, and oval. The
foraging activity, with adult beetles becoming more average length of the eggs was 1.9 ⫾ 0.15 mm (n ⫽ 50),
active at 1400 hours. We could conÞrm some of the and the average width was 1.6 ⫾ 0.10 mm (n ⫽ 50).
observations made by Facundo et al. (1999) on E. Before hatching, larvae are visible through the chorion
orientalis. In Belgium, little beetle activity was ob- of mature eggs. Preliminary observations conducted in
served when air temperature was ⬍14⬚C and when the laboratory in June and July 2000 on eggs kept in
days were overcast. petri dishes showed that high moisture was necessary
There is no information available on mating activity for eclosion (data not shown); this is similar to what
of H. philanthus beetles. However, during the obser- they need in their natural habitats. The average egg
vation period, mating was observed on surrounding development time was 28 ⫾ 5.0 (n ⫽ 50) days. More
shrubs and trees from 900 to 1600 hours. It lasted ⬇2 than 95% of the eggs hatched under these conditions.
min and was repeated several times. We also noticed Larval Development. On 26 June 2000, the Þrst eggs
that males of H. philanthus often ride on the dorsum were found in the soil at the observation site. By 24
of the females and tried copulating repeatedly. Fe- July of that year, only a few eggs remained, while the
males that were not receptive to copulation tried to population was mainly composed of Þrst instars (L1).
escape when males mounted them. In 2000, 2001, and Their average head capsule width was 1.1 ⫾ 0.5 mm
2002, the Þrst mating was observed on 7, 9, and 11 June, (n ⫽ 125). In the second week of September 2000, L1s
respectively. Additional studies on the sexual behavior had molted to L2s. The average head capsule width
of H. philanthus are necessary for understanding the was 2.2 ⫾ 0.20 mm (n ⫽ 450). With decreasing soil
life cycle of H. philanthus. temperature, L2 larvae descended into the soil where
No study has been conducted on the feeding be- they spent the winter (December and January) below
havior of H. philanthus adults. During our observation, the frost line (below 5Ð10 cm deep). In March
we frequently observed H. philanthus beetles feeding (spring) 2001, as soil temperature increased, L2 larvae
on leaves of B. utilis variety jacquemontii, and C. began their upward migration. By late March to early
betulus, causing moderate damage to plants. However, April, the overwintering L2 larvae had moved close to
other scarab species such as Adoretus tenuimaculatus the grass roots and resumed feeding. L2 larvae
(Waterhouse) (Lee et al. 1998), Ectinohoplia rufipes moulted to third instar (L3) by June 2001; their av-
(Motschulsky) (Choo et al. 1999), and Popillia ja- erage head capsule width was 3.0 ⫾ 0.25 (n ⫽ 550).
ponica (Newman) (Fleming 1972) feed heavily on the This is the phase in the grubsÕ life cycle during which
leaves and ßowers of trees and shrubs. The signiÞ- it feeds most actively on grass roots and causes the
cance of the damage caused by the adult beetle of H. most important damage. In late October and early
philanthus remains unknown and warrants further November 2001, L3 larvae migrated deeper into the
study. soil. In late March 2002 overwintering L3 larvae be-
Oviposition. On their release in the cages, the bee- came active and moved toward the soil surface for a
tles stayed a few hours on the cage net or on the C. brief feeding period. At this time, minor turf damage
betulus leaves, but eventually buried deep into the soil. occurred as the mature larvae soon stopped feeding to
The following days, the beetles moved out of the soil, initiate pupation in May 2002. The life span of the
spent a few hours on the plant leaves, and Þnally third-instar larvae was ⬎330 d. From these studies and
returned to the soil. Oviposition started during the last observations, it could be concluded that in Belgium H.
week of June 2000 depending on the age of the beetle philanthus has one generation per 2 yr with three larval
and was completed the last week of July 2000. Ovi- stages (Fig. 4).
position started around 1000 hours and was most com- Through continuous soil sampling during the larval
mon in the afternoon at 1400 hours. Gravid females development in plots, we observed larval migration to
generally oviposited alone, although sometimes two different levels in the soil. The depth at which the
females were found on a single entrance hole. The larvae could be found depended on the soil temper-
female deposited a cluster of 25Ð 40 eggs in the same ature in late autumn, winter, and early spring and
hole at a depth of 10 Ð15 cm below the thatch layer. mainly on soil moisture during the rest of the year.
The depth of oviposition seemed to depend on the Second- and third-instar larvae were found as deep as
moisture content of the soil; if the water content of the 50 cm in the soil during the winter (Fig. 5). However,
soil was near to its Þeld capacity, ⬎90% of the eggs during the grub sampling, we observed that ⬎50% of
were laid in the upper 10 cm. In the last week of July the larvae were present at a depth of 10 cm (Fig. 5).
in each of the 3 yr, the females stopped laying eggs and We did not estimate the larval population below 20-
reduced feeding and general activity. During ovipo- and 50-cm soil depth. Further studies on larval distri-
sition, the mean maximum temperature was 22.1, 23, bution and percentage of soil moisture at different
December 2006 ALI ANSARI ET AL.: BIOLOGY OF H. philanthus 1505

Fig. 4. Life cycle of H. philanthus in lawn at the Research Station for Ornamental Plants (PCS), Destelbergen, under
natural conditions showing the presence of multiple generations in 3 consecutive sampling yr. Plain block: adults emerged
in June 2000. Horizontal lines: generation started from eggs laid by adults in June 2000. Vertical lines: generation started from
eggs laid by adults in June 1999. Multi-blocks: generation started from eggs laid by adults which emerged in June 2001. Cross
lines: generation started from eggs laid by adults which emerged in June 2002.

levels in the soil are planned. Both second- and third- Pupae. Pupae were present in the soil from the Þrst
instar larvae of H. philanthus overwintered. High num- week of May 2002 to the end of that month; no pupae
bers of larvae survived in winter because of their vertical were found later than May. Initially, pupae were pale
migration (the soil does not usually freeze at ⬍50 cm; Fig. yellow; but turn brownish-yellow with aging. They
5). A preliminary observation indicated that larvae could were ⬇14.0 ⫾ 2.0 mm long and 4.0 ⫾ 1.0 mm wide (n ⫽
resist soil temperatures below 0⬚C for several days. When 80). In May 2002, at an average soil temperature of
observing 150 larvae left to overwinter in 2000, 2001, and 16.5⬚C, the development from pupa to adult took 28 ⫾
2002 at 7Ð10 cm depth under the thatch layer for 250 h 5 d under Þeld conditions. Approximately 80% of the
in frozen soil in the Þeld, an average larval survival rate pupae were found in the most superÞcial layer (0 Ð10
of ⬎90% was recorded for L2 and 80% for L3 larvae. An cm) and 20% in the layer (10 Ð20 cm).
average survival of 98% was observed for larvae kept at Mortality Factors. No parasitized eggs were found
50 cm depth. All the L2 and L3 resumed feeding within at the observation site. During our study, we collected
24 h when moved in DecemberÐFebruary from the open several naturally infected larvae of H. philanthus from
Þeld to the laboratory at 20⬚C. observation sites (lawns, sport turf, and pastures) and
1506 ENVIRONMENTAL ENTOMOLOGY Vol. 35, no. 6

Fig. 5. Presence of H. philanthus larvae in the soil over a 2-yr period. Clear bars indicate the number of larvae in the Þrst
10 cm of an 0.1-m2 area; dark bars indicate average maximal depth (in cm) at which larvae were found data average of at
least two samplings per month.

isolated entomopathogenic nematodes and Hypho- ample, use of traps might help to reduce beetle pop-
mycetes fungi. An entomopathogenic fungus of the ulation and plant damage, although this has to be
genus Metarhizium infected a total of 10% of the larvae conÞrmed in Þeld studies Þrst. Cultural control meth-
collected and was identiÞed as M. anisopliae variety ods that could be deployed by home gardeners (e.g., trap
anisopliae (Metschnikoff) Sorokin (Ansari et al. cropping, companion planting, repellent mulches) war-
2004a). A second fungus species was identiÞed as rant further study. Also, more detailed studies of the
Beauveria bassiana (Balsamo) Vuillemin (Humber biology (adult fertility and longevity and host range) of
1997). In addition to fungi, entomopathogenic nem- this species are needed. Recently, biological control
atodes of the genus Heterorhabditis (Nematoda: Het- has been examined as control tool for H. philanthus.
erorhabditidae) were also isolated from infected lar- Experiments with entomopathogenic nematodes (An-
vae of both H. philanthus and M. melolontha; the sari et al. 2003b), fungi (Ansari et al. 2004a), and their
nematodes were identiÞed as H. bacteriophora (Poi- combined use (Ansari et al. 2004b) have been carried
nar) (Ansari et al. 2003a). Recently, we isolated Stein- out in laboratory, greenhouse and under Þeld condi-
ernema glaseri (Steiner) (Ansari et al. 2005) from tion (Ansari et al. 2006) against third-instar H. philan-
naturally infected third-instar H. philanthus in football thus. Results from combined application had an addi-
Þeld in Eeklo, province of East Flanders. tive to synergistic effect on larval mortality. This new
In conclusion, the studies and observations con- approach may offer a powerful and reliable tool for
ducted show that in Belgium one H. philanthus gen- biological control of H. philanthus.
eration takes 2 yr; the life cycle is spread over 3 cal-
endar yr. The life cycle of scarabs varies considerably
with climate and species. Species belonging to the Acknowledgments
subfamily Melolonthinae, tribe Sericini usually have a We thank H. Van De Sype and J. Witters for very valuable
1-yr life cycle, but some species in more temperate help with the Þeldwork. We also thank the Research Station
climates have a 2-yr life cycle (Ritcher 1957). The for Ornamental Plant Production (PCS), Destelbergen, Bel-
European cockchafer or May beetle, M. melolontha, gium, for providing access to a Þeld heavily infected with H.
has a 4-yr life cycle (Keller et al. 1999), especially in philantus.
temperate regions. In North America, many species of
Phyllophaga have a 2- or 3-yr life cycle (Ritcher 1957). References Cited
To our knowledge, there has been no extensive study
of the life cycle of H. philanthus. However, Freude et Ansari, M. A. 2004. Biological control of Hoplia philanthus
al. (1969) have mentioned a 2-yr development. (Coleoptera: Scarabaeidae) with entomopathogenic
The larvae of H. philanthus are a serious pest of nematodes and fungi. PhD dissertation, Ghent University,
Ghent, Belgium.
turfgrass. Soil insecticides are still the primary means Ansari, M. A., P. K. Long, and M. Moens. 2003a. Heterorh-
for turfgrass managers, growers, and millions of home- abditis bacteriophora (Heterorhabditidae: Rhabditida),
owners for white grubs control (Vlug 1996). However, parasitic in natural populations of white grubs (Co-
because in Belgium, this type of control is not allowed leoptera: Scarabaeidae) in Belgium. Russ. J. Nematol. 11:
in turf, alternative products will be needed. For ex- 57Ð59.
December 2006 ALI ANSARI ET AL.: BIOLOGY OF H. philanthus 1507

Ansari, M. A., L. Tirry, and M. Moens. 2003b. Entomopatho- Fleming, W. E. 1972. Biology of the Japanese beetle. U.S.
genic nematodes and their symbiotic bacteria for the Dept. Agric. Res. Stat. Tech. Bull. 1449: 1Ð129.
biological control of Hoplia philanthus (Coleoptera: Scar- Freude, H., K. W. Harde, and G. A. Lohse (eds). 1969. Die
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