Summer activity patterns of nocturnal Scarabaeoidea (Coleoptera) of the southern tablelands of New...
10
Australian Journal of Entomology (2007) 46, 7–16 © 2007 The Authors Journal compilation © 2007 Australian Entomological Society doi:10.1111/j.1440-6055.2007.00579.x Blackwell Publishing AsiaMelbourne, AustraliaAENAustralian Journal of Entomology1326-6756© 2006 The Authors; Journal compilation © 2006 Australian Entomological Society? 2007461716original articleScarabs of the southern tablelands of New South Wales M J Steinbauer and T A Weir *[email protected] †Present address: Australian Plague Locust Commission, Australian Government – Department of Agriculture, Fisheries and Forestry, GPO Box 858, Canberra, ACT 2601, Australia. Summer activity patterns of nocturnal Scarabaeoidea (Coleoptera) of the southern tablelands of New South Wales Martin J Steinbauer 1 *† and Tom A Weir 2 1 University of Tasmania and CRC for Sustainable Production Forestry, c/– CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia 2 CSIRO Entomology, Australian National Insect Collection, GPO Box 1700, Canberra, ACT 2601, Australia Abstract Australia has a rich diversity of Scarabaeoidea; however, little is known about the majority of them. Because adults of Anoplognathus, Automolius, Heteronychus, Heteronyx and Liparetrus in particular are reliant upon eucalypts, a number of bluegum plantation companies supported the commencement of research into the biology and ecology of scarabs of economic significance to them. Consequently, it was decided that the occurrences of species endemic to this area would be studied and it was assumed that information on the nocturnal species in the aforementioned genera would be obtained. From late November 2003 until late February 2004, the abundances of Scarabaeoidea caught in two light traps that partition insects caught on a given night into seven time periods each of 1.75 h duration were recorded. A total of 48 263 scarabs representing 21 genera were caught. Within the 14 species caught most often, six types of summer activity pattern were apparent: late spring to early summer (Austral- obolbus gayndahensis), early to mid-summer (Scitala sericans), mid-summer only (Sericesthis ignota), mid- to late summer (Acrossidius tasmaniae, Aphodius lividus, Heteronyx chlorotica, Het. praecox and Antitrogus morbillosus), late summer only (Ataenius picinus) and all summer (Anoplognathus pallidicollis, Phyllotocus macleayi, Sericesthis geminata, Ser. micans and Ser. nigrolineata). Abundances of nine species peaked between 21:30 and 23:15 h (Aph. lividus, Phy. macleayi, Het. chlorotica, Sci. sericans, Ser. geminata, Ser. micans, Ser. nigrolineata and possi- bly also Ant. morbillosus), three were most abundant between 19:45 and 21:30 h (Ano. pallidicollis, Ser. ignota and possibly also Ata. picinus), another two were most abundant from 19:45 to 23:15 h (Acr. tasmaniae and Het. praecox) and Aus. gayndahensis was most abundant between 23:15 and 01:00 h. Of course, it is not just a knowledge of the identity of the species and the timing of their occurrence that are important when making insect management decisions, but also the size of popu- lation needed to inflict economically significant loss. It is now beholden upon bluegum plantation companies to support further research to determine the relationships between light trap catches of eucalypt-feeding scarabs, tree age and/or size and level of defoliation in order to improve their confidence in this method of monitoring over ground surveys. Key words Bolboceratidae, Christmas beetles, cockchafers, pasture scarabs, Scarabaeidae, Trogidae. INTRODUCTION Scarabaeoidea are a very characteristic and often abundant element of the insect fauna of Australian summers (e.g. Carne et al. 1974, 1981; Roberts et al. 1982a). Nevertheless, detailed information relating to the seasonal phenologies of these insects is rather scarce. Of the published information, Allsopp and Logan (1999) recognised four patterns of seasonal activity in the 14 species of scarab they monitored, i.e. brief spring activity, brief summer activity, prolonged summer activity and prolonged spring to autumn activity. Only the nocturnal activ- ity patterns of some Australian dung beetles have been rela- tively well documented. Within species of Australian and exotic dung beetle there is an array of activity patterns that encompass the spectrum from entirely diurnal to entirely nocturnal (Houston & McIntyre 1985; Howden et al. 1991; Caveney et al. 1995; Davis 1999). Seasonal and daily activity patterns of any organism are so fundamental to understanding any organism that they are usu- ally the first aspects of a species’ biology that needs to be studied if not already known. The expansion of eucalypt plan- tations in parts of temperate Australia has increased the need to improve the level of knowledge of the basic biology of many eucalypt-feeding insects. The often sudden need to gain an understanding of the biology of insects of concern to human activities is not new; for example, the study of species of
Transcript of Summer activity patterns of nocturnal Scarabaeoidea (Coleoptera) of the southern tablelands of New...
Australian Journal of Entomology (2007) 46, 7–16
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society doi:10.1111/j.1440-6055.2007.00579.x
Blackwell Publishing AsiaMelbourne, AustraliaAENAustralian Journal of Entomology1326-6756© 2006 The Authors; Journal compilation © 2006 Australian Entomological Society? 2007461716original articleScarabs of the southern tablelands of New South WalesM J Steinbauer and T A Weir
†Present address: Australian Plague Locust Commission, AustralianGovernment – Department of Agriculture, Fisheries and Forestry, GPOBox 858, Canberra, ACT 2601, Australia.
Summer activity patterns of nocturnal Scarabaeoidea (Coleoptera) of the southern tablelands of New South Wales
Martin J Steinbauer1*† and Tom A Weir2
1University of Tasmania and CRC for Sustainable Production Forestry, c/– CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia 2CSIRO Entomology, Australian National Insect Collection, GPO Box 1700, Canberra, ACT 2601, Australia
Abstract Australia has a rich diversity of Scarabaeoidea; however, little is known about the majority of them.Because adults of Anoplognathus, Automolius, Heteronychus, Heteronyx and Liparetrus in particularare reliant upon eucalypts, a number of bluegum plantation companies supported the commencementof research into the biology and ecology of scarabs of economic significance to them. Consequently,it was decided that the occurrences of species endemic to this area would be studied and it was assumedthat information on the nocturnal species in the aforementioned genera would be obtained. From lateNovember 2003 until late February 2004, the abundances of Scarabaeoidea caught in two light trapsthat partition insects caught on a given night into seven time periods each of 1.75 h duration wererecorded. A total of 48 263 scarabs representing 21 genera were caught. Within the 14 species caughtmost often, six types of summer activity pattern were apparent: late spring to early summer (Austral-obolbus gayndahensis), early to mid-summer (Scitala sericans), mid-summer only (Sericesthisignota), mid- to late summer (Acrossidius tasmaniae, Aphodius lividus, Heteronyx chlorotica,Het. praecox and Antitrogus morbillosus), late summer only (Ataenius picinus) and all summer(Anoplognathus pallidicollis, Phyllotocus macleayi, Sericesthis geminata, Ser. micans andSer. nigrolineata). Abundances of nine species peaked between 21:30 and 23:15 h (Aph. lividus,Phy. macleayi, Het. chlorotica, Sci. sericans, Ser. geminata, Ser. micans, Ser. nigrolineata and possi-bly also Ant. morbillosus), three were most abundant between 19:45 and 21:30 h (Ano. pallidicollis,Ser. ignota and possibly also Ata. picinus), another two were most abundant from 19:45 to 23:15 h(Acr. tasmaniae and Het. praecox) and Aus. gayndahensis was most abundant between 23:15 and01:00 h. Of course, it is not just a knowledge of the identity of the species and the timing of theiroccurrence that are important when making insect management decisions, but also the size of popu-lation needed to inflict economically significant loss. It is now beholden upon bluegum plantationcompanies to support further research to determine the relationships between light trap catches ofeucalypt-feeding scarabs, tree age and/or size and level of defoliation in order to improve theirconfidence in this method of monitoring over ground surveys.
Key words Bolboceratidae, Christmas beetles, cockchafers, pasture scarabs, Scarabaeidae, Trogidae.
INTRODUCTION
Scarabaeoidea are a very characteristic and often abundantelement of the insect fauna of Australian summers (e.g. Carneet al. 1974, 1981; Roberts et al. 1982a). Nevertheless, detailedinformation relating to the seasonal phenologies of theseinsects is rather scarce. Of the published information, Allsoppand Logan (1999) recognised four patterns of seasonal activityin the 14 species of scarab they monitored, i.e. brief springactivity, brief summer activity, prolonged summer activity and
prolonged spring to autumn activity. Only the nocturnal activ-ity patterns of some Australian dung beetles have been rela-tively well documented. Within species of Australian andexotic dung beetle there is an array of activity patterns thatencompass the spectrum from entirely diurnal to entirelynocturnal (Houston & McIntyre 1985; Howden et al. 1991;Caveney et al. 1995; Davis 1999).
Seasonal and daily activity patterns of any organism are sofundamental to understanding any organism that they are usu-ally the first aspects of a species’ biology that needs to bestudied if not already known. The expansion of eucalypt plan-tations in parts of temperate Australia has increased the needto improve the level of knowledge of the basic biology of manyeucalypt-feeding insects. The often sudden need to gain anunderstanding of the biology of insects of concern to humanactivities is not new; for example, the study of species of
8 M J Steinbauer and T A Weir
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Anoplognathus was initiated when rural dieback of woodlandeucalypts was of wider public concern (Landsberg et al. 1990).In parts of temperate Australia, plantations of Eucalyptus glob-ulus ssp. globulus can suffer considerable defoliation as a resultof the activities of a range of scarab species, especially speciesof Anoplognathus, Automolius, Heteronychus, Heteronyx andLiparetrus (Bashford 1993; Neumann 1993; Stone 1993; Loch& Floyd 2001; Bulinski & Matthiessen 2002; Floyd et al. 2002;Hurley & Patel 2003; Lawson & King 2003). Heteronychusarator, species of Automolius, Heteronyx and Liparetrus canmass-attack and kill seedling E. globulus, sometimes requiringthat they be re-planted at considerable economic cost (Bulinski& Matthiessen 2002; Lawson & King 2003).
Our study was initiated to provide basic information on theactivity patterns of Heteronyx adults in particular (note:because we used light traps, we were unlikely to catch eitherAutomolius or Liparetrus species because members of thesegenera are diurnal). Rather than only record the numbers ofHeteronyx species caught, we decided to record the numbersof individuals belonging to the entire superfamily.
MATERIALS AND METHODS
Site
This study was undertaken in a field trial of 500 (numberoriginally planted) rain-fed eucalypts that were planted inOctober 1998 as 6-month-old seedlings. The site is located onland that formerly belonged to the CSIRO Ginninderra Exper-iment Station (35°09′55.7″S and 149°02′49.9″E; altitude615 m above sea level), which is on the southern tablelands ofNew South Wales.
Light trapping and recording temperature and rainfall
The two 8 W ultraviolet light traps used in this study aredescribed in Steinbauer (2003). The traps were programmedto operate for 12 h from 18:00 to 06:00 (Australian easternstandard time) and could partition catches into seven timeperiods, i.e. 18:00–19:45 h, 19:45–21:30 h, 21:30–23:15 h,23:15–01:00 h, 01:00–02:45 h, 02:45–04:30 h and 04:30–06:00 h.
We trapped for 48 nights, beginning on the evening of 27November 2003 (week 48) and continuing until the morningof 26 February 2004 (week 9). Traps did not operate on Fridayand Saturday nights. Both traps were turned off from theevening of 19 December (week 52) to the morning of 4January 2004 (week 1).
A Starlog (UNIDATA Australia, Perth) portable data-logger (model 6003A), with version 3.09 software, was placedapproximately mid-way between the two light traps. The data-logger was fitted with a model 6501 ambient temperaturesensor and a model 6506 tipping bucket rainfall gauge. Thedata-logger was programmed to record daily maxima and min-ima for ambient temperature and daily rainfall. These data aresummarised in Figure 1.
Insect identification and grouping
Species identifications were made by comparison withspecimens in the Australian National Insect Collection, inconjunction with the following taxonomic papers: Aphodiinae– Stebnicka and Howden (1995, 1997); Dynastinae – Carne(1957a); Melolonthinae – Britton (1957, 1980, 1987, 1988,2000) and Allsopp (2003); Rutelinae – Carne (1957b, 1958);Bolboceratidae – Howden (1992); Trogidae – Scholtz (1986).
Visual inspection of our trap catch data, as well as referenceto the categorisations used by Allsopp and Logan (1999), ledus to identify six types of summer activity pattern. These were:all summer, late spring to early summer, early to mid-summer,mid-summer only, mid- to late summer and late summer only.
To interpret nocturnal activity patterns, we grouped theinsects that we caught frequently according to aspects of theirbiology, i.e. dung-feeding (as adults, not necessarily as larvae),diurnal, eucalypt-feeding, purportedly eucalypt-feeding andnon-feeding and unknown biology. We refer to some of theseinsects as ‘purportedly eucalypt-feeding’ because we did nottrap them in 20 funnel traps hung beneath the canopies of fiveeucalypts in the field trial nor did Sericesthis micans eat anyof the leaves of the five species of eucalypt offered to them(MJ Steinbauer unpubl. data 2004). Sericesthis ignota was notincluded in the lists of ‘tree-feeding scarabs’ given in Robertset al. (1982a,b) but is grouped with the purportedly eucalypt-feeding species by virtue of the genus to which it belongs.
Data analysis
Trends in weekly and nightly light trapping data were statis-tically analysed only for those species of which we caught atleast 40 individuals during the whole period using both lighttraps. Week number was used to facilitate both the presenta-tion of the data and the statistical analyses. Catches of scarabs(not including zeroes) according to week and time period were
Fig. 1. Maximum and minimum ambient temperature (data aremeans ± standard errors) and rainfall (vertical bars) at CSIROGinninderra Experiment Station for 27 November 2003 (week 48)to 26 February 2004 (week 9).
Scarabs of the southern tablelands of New South Wales 9
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
first analysed using Kruskal–Wallis ranking (adjusted forties) tests, followed by pair-wise comparisons using Mann–Whitney tests if initial analyses were statistically significant(and providing sufficient data). Only pair-wise comparisonsfor consecutive weeks (ignoring the period between weeks 52and 1) were conducted. Analyses were conducted usingMINITAB Release 13.32 for Windows. Only weekly catchesof species that exhibit statistically significant variations inabundance are graphically illustrated. Statistical results forcatches by time period are presented in the Results section.
RESULTS
Species composition
During our light trapping we caught 48 263 insects represent-ing 21 genera and 46 species (Table 1). Four of the speciescaught represent accidental introductions into Australia.Adults of most of the species trapped are nocturnal (but seeDiscussion). Of the 21 genera, Heteronyx was the most spe-cies-rich, with 13 species. The next most species-rich generawere Anoplognathus (seven species) and Sericesthis (six spe-cies). We caught 40 or more individuals of 14 species. Of thesespecies, Acrossidius tasmaniae was the most abundant; it com-prised 87.7% of all the insects caught (Table 1). The next mostabundant species was Ser. micans (6.2%), followed by Aphod-ius lividus (1.2%), followed by Phyllotocus macleayi (1.1%).
Dung-feeding scarabs
Of the three species of dung-feeding scarabs, Acr. tasmaniaeand Aph. lividus were active from mid- to late summer andAtaenius picinus was active in late summer only (Table 2). Theweekly fluctuations in the catches of Acr. tasmaniae were sta-tistically significant (H = 21.13, P = 0.004). Pair-wise testsrevealed that catches in week 7 were significantly lower thanthose in week 8 (W = 662.0, P = 0.003; see Fig. 2); no otherpair-wise tests were significant. There were no statisticallysignificant differences in weekly catches of either Aph. lividusor Ata. picinus (H = 7.40, P = 0.284 and H = 1.95, P = 0.377,respectively).
Catches by time period for both Acr. tasmaniae andAph. lividus exhibited statistically significant variation (seeTable 2). In the case of Acr. tasmaniae, catches between 21:30and 23:15 h were significantly larger than those between 23:15and 01:00 h (W = 1882.0, P = 0.026). For Aph. lividus, nopair-wise tests of catches for the three consecutive periodsspanning 19:45–02:45 h were statistically significant and thenumber of catches in periods from 02:45 to 06:00 were toofew for analysis. No catches by time period in the case ofAta. picinus were statistically significant (Table 2).
Diurnal scarabs
Both Anoplognathus pallidicollis and Phy. macleayi wereactive all summer. Catches of Ano. pallidicollis did not exhibitstatistically significant weekly (H = 8.78, P = 0.361) or timeperiod fluctuations (Table 2). In contrast, catches of
Phy. macleayi exhibited statistically significant weekly varia-tion (H = 20.30, P = 0.016; Fig. 3). Pair-wise tests revealedthat catches in week 51 were significantly less than those inweek 2, catches in week 2 were larger than those in week 3as were catches in week 7 compared with week 8 (W = 34.0,P = 0.019; W = 83.0, P = 0.014 and W = 131.0, P = 0.014,respectively). It seems likely that the abundance of this speciesrose markedly during the 2 weeks (52 of 2003 and 1 of 2004)when we were not light trapping. Catches of Phy. macleayi bytime period did not, however, exhibit statistically significantvariation (see Table 2).
The anomaly concerning captures of these two supposedlydiurnal species at night is considered in the Discussion.
Eucalypt-feeding scarabs
Adult Heteronyx chlorotica and Het. praecox were caught inmodest numbers during mid-summer only. The light trapcatches of neither species exhibited statistically significant
Fig. 2. Weekly light trap catches of dung-feeding adults ofAcrossidius tasmaniae (data are means ± standard errors).
Fig. 3. Weekly light trap catches of the diurnal scarab Phyllo-tocus macleayi (data are means ± standard errors).
10 M J Steinbauer and T A Weir
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Tabl
e 1
The
spe
cies
of
Scar
abae
oide
a, t
heir
tax
onom
ic a
ffilia
tions
and
bio
logi
es, c
augh
t on
Gin
nind
erra
Exp
erim
ent
Stat
ion
Spec
ies
nam
eFa
mily
Subf
amily
Com
mon
nam
eL
ife
cycl
e du
ratio
n in
yea
rs
Adu
lt fo
odR
efer
ence
(s)
Tota
l nu
mbe
rca
ught
Acr
ossi
dius
tas
man
iae
(Hop
e)Sc
arab
aeid
aeA
phod
iinae
Bla
ckhe
aded
pas
ture
co
ckch
afer
1D
ung
Car
ne (
1957
c)42
447
Ano
plog
nath
us c
hlor
opyr
us(D
rapi
ez)
Scar
abae
idae
Rut
elin
aeC
hris
tmas
bee
tle2
(can
be
1)E
ucal
yptu
sC
arne
et a
l. (1
974)
38
Ano
plog
nath
us h
irsu
tus
Bur
mei
ster
Scar
abae
idae
Rut
elin
aeC
hris
tmas
bee
tle2
Euc
alyp
tus
Rob
erts
et a
l. (1
982a
); G
oody
er (
1985
)26
Ano
plog
nath
us m
onta
nus
Mac
leay
Scar
abae
idae
Rut
elin
aeC
hris
tmas
bee
tle2
Euc
alyp
tus
Goo
dyer
(19
85)
4A
nopl
ogna
thus
pal
lidi
coll
isB
lanc
hard
Scar
abae
idae
Rut
elin
aeC
hris
tmas
bee
tle2
Euc
alyp
tus
Goo
dyer
(19
85)
70
Ano
plog
nath
us p
inda
rus
Car
neSc
arab
aeid
aeR
utel
inae
Chr
istm
as b
eetle
2E
ucal
yptu
sG
oody
er (
1985
)1
Ano
plog
nath
us s
utur
alis
Boi
sduv
alSc
arab
aeid
aeR
utel
inae
Chr
istm
as b
eetle
2E
ucal
yptu
sG
oody
er (
1985
)32
Ano
plog
nath
us v
elut
inus
Boi
sduv
alSc
arab
aeid
aeR
utel
inae
Chr
istm
as b
eetle
2E
ucal
yptu
sG
oody
er (
1985
)1
Ant
itro
gus
mor
bill
osus
(B
lack
burn
)Sc
arab
aeid
aeM
elol
onth
inae
Tabl
elan
d pa
stur
e sc
arab
2N
on-f
eedi
ngR
ober
ts e
t al.
(198
2a,b
);
Goo
dyer
(19
85)
186
# A
phod
ius
livi
dus
(Oliv
ier)
Scar
abae
idae
Aph
odiin
ae–
?D
ung
Steb
nick
a an
d H
owde
n (1
995)
586
# A
taen
ius
pici
nus
Har
old
Scar
abae
idae
Aph
odiin
ae–
?D
ung,
dec
ayin
g O
rgan
ic m
atte
rSt
ebni
cka
and
How
den
(199
7)42
Aus
tral
obol
bus
gayn
dahe
nsis
(Mac
leay
)B
olbo
cera
tidae
––
?U
nkno
wn
–56
Aut
omol
ius
vulg
aris
Bri
tton
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1?
Euc
alyp
tus
Law
son
and
Kin
g (2
003)
1D
asyg
nath
us t
ritu
bera
latu
sB
lack
burn
Scar
abae
idae
Dyn
astin
ae–
1U
nkno
wn
(not
lea
ves)
Car
ne (
1957
a)1
Ele
phas
tom
us m
eral
dus
Car
neB
olbo
cera
tidae
––
2 pe
r ye
arD
ung
Car
ne (
1965
)2
# H
eter
onyc
hus
arat
or (
Fabr
iciu
s)Sc
arab
aeid
aeD
ynas
tinae
Afr
ican
bla
ck b
eetle
1W
ide
vari
ety
of p
lant
sW
alla
ce (
1946
); C
arne
(19
57a)
4
Het
eron
yx s
p. n
ov.
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1
or m
ore
Euc
alyp
tus
Rob
erts
et a
l. (1
982b
); B
ritto
n (2
000)
1H
eter
onyx
sp.
nov
. [K
ey 1
4]Sc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)3
Het
eron
yx s
p.Sc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)11
Het
eron
yx a
equa
lis
Bla
ckbu
rnSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)8
Het
eron
yx a
gric
ola
Bla
ckbu
rnSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)5
Het
eron
yx a
ustr
alis
Gué
rin-
Mén
evill
eSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)4
Het
eron
yx c
ervi
na B
oisd
uval
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1
or m
ore
Euc
alyp
tus
Rob
erts
et a
l. (1
982b
); B
ritto
n (2
000)
1H
eter
onyx
chl
orot
ica
(Gyl
lenh
al)
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1
or m
ore
Euc
alyp
tus
Rob
erts
et a
l. (1
982b
); B
ritto
n (2
000)
80
Nam
es o
f sp
ecie
s pr
efac
ed b
y #
are
intr
oduc
ed s
peci
es. A
NIC
, Aus
tral
ian
Nat
iona
l In
sect
Col
lect
ion;
–, n
o da
ta.
Scarabs of the southern tablelands of New South Wales 11
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Het
eron
yx e
xcis
us B
lack
burn
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1
or m
ore
Euc
alyp
tus
Rob
erts
et a
l. (1
982b
); B
ritto
n (2
000)
18H
eter
onyx
gra
nula
tus?
Bla
ckbu
rnSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)4
Het
eron
yx g
ranu
m B
urm
eist
erSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)25
Het
eron
yx p
raec
ox (
Eri
chso
n)Sc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)28
5H
eter
onyx
sim
ius
Bla
ckbu
rnSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
mor
eE
ucal
yptu
sR
ober
ts e
t al.
(198
2b);
Bri
tton
(200
0)1
Mae
chid
ius
sp.
Scar
abae
idae
Mel
olon
thin
ae–
?U
nkno
wn
–1
Om
orgu
s co
stat
us (
Wie
dem
ann)
Tro
gida
e–
Car
cass
bee
tle?
Dry
car
cass
es,
bat
guan
oSc
holtz
(19
86);
Mat
thew
s (1
984)
20
Ont
hoph
agus
sp.
nov
. nr.
pent
acan
thus
Har
old
Scar
abae
idae
Scar
abae
inae
Dun
g be
etle
?D
ung
AN
IC d
ata
1
Oph
ropy
x hi
spid
a (B
lack
burn
)Sc
arab
aeid
aeM
elol
onth
inae
Cha
fer
?E
ucal
yptu
sB
ritto
n (1
987)
; AN
IC d
ata
9P
hyll
otoc
us b
imac
ulat
us E
rich
son
Scar
abae
idae
Mel
olon
thin
aeN
ecta
r sc
arab
?N
ecta
r &
pol
len
Don
aghe
y (1
998)
8P
hyll
otoc
us m
acle
ayi
Fisc
her
Scar
abae
idae
Mel
olon
thin
aeN
ecta
r sc
arab
?N
ecta
r &
pol
len
Don
aghe
y (1
998)
516
Phy
llot
ocus
rufi
penn
is (
Boi
sduv
al)
Scar
abae
idae
Mel
olon
thin
aeN
ecta
r sc
arab
?N
ecta
r &
pol
len
Don
aghe
y (1
998)
2Sc
ital
a se
rica
ns E
rich
son
Scar
abae
idae
Mel
olon
thin
aeSh
iny
past
ure
scar
ab1
Euc
alyp
tus?
Har
dy (
1977
); R
ober
ts e
t al.
(198
2b)
155
Sem
anop
teru
s su
bcos
tatu
s(C
aste
lnau
)Sc
arab
aeid
aeD
ynas
tinae
–?
Non
-fee
ding
Law
renc
e an
d B
ritto
n (1
991)
1
Seri
cest
his
gem
inat
a B
oisd
uval
Scar
abae
idae
Mel
olon
thin
aePr
uino
se s
cara
b1
Euc
alyp
tus
Wen
sler
(19
71);
R
ober
ts e
t al.
(198
2a,b
)51
Seri
cest
his
igno
ta B
ritto
nSc
arab
aeid
aeM
elol
onth
inae
Cha
fer
1 or
2?
Euc
alyp
tus
Bri
tton
(198
7)17
8Se
rice
sthi
s m
ican
s B
lack
burn
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r2
Euc
alyp
tus
Rob
erts
et a
l. (1
982a
,b);
B
ritto
n (1
987)
2 98
7
Seri
cest
his
nigr
olin
eata
(Boi
sduv
al)
Scar
abae
idae
Mel
olon
thin
aeD
usky
pas
ture
sca
rab
2E
ucal
yptu
sC
arne
and
Chi
nnic
k (1
957)
; R
ober
ts e
t al.
(198
2a,b
);
Bri
tton
(198
7)
338
Seri
cest
his
para
llel
a B
lack
burn
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r1
or 2
?E
ucal
yptu
sB
ritto
n (1
987)
2Se
rice
sthi
s su
tura
lis
(Mac
leay
)Sc
arab
aeid
aeM
elol
onth
inae
Cha
fer
2E
ucal
yptu
sR
ober
ts e
t al.
(198
2a,b
);
Bri
tton
(198
7)22
# Tr
ox s
cabe
r (L
inna
eus)
Tro
gida
e–
Car
cass
bee
tle?
Dry
car
cass
esSc
holtz
(19
86)
2W
ebbe
lla
sp.
Scar
abae
idae
Mel
olon
thin
aeC
hafe
r?
Unk
now
n–
27
Spec
ies
nam
eFa
mily
Subf
amily
Com
mon
nam
eL
ife
cycl
e du
ratio
n in
yea
rs
Adu
lt fo
odR
efer
ence
(s)
Tota
l nu
mbe
rca
ught
Nam
es o
f sp
ecie
s pr
efac
ed b
y #
are
intr
oduc
ed s
peci
es. A
NIC
, Aus
tral
ian
Nat
iona
l In
sect
Col
lect
ion;
–, n
o da
ta.
12 M J Steinbauer and T A Weir
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
weekly (H = 4.26, P = 0.513 and H = 9.25, P = 0.160, respec-tively) or time period fluctuations (see Table 2).
Purportedly eucalypt-feeding scarabs
Three of the species of purportedly eucalypt-feeding scarabswere caught all summer (Sericesthis geminata, Ser. micans andSer. nigrolineata), one (Sci. sericans) was caught from earlyto mid-summer and another (Ser. ignota) was caught in mid-summer only (Table 2). Only the weekly catches ofSci. sericans, Ser. micans and Ser. nigrolineata exhibited sta-tistically significant fluctuations (H = 10.25, P = 0.006;H = 30.48, P < 0.001 and H = 20.38, P = 0.005, respectively).Because the number of times Sci. sericans was caught in weeks2 and 3 was so few, no pair-wise tests were able to be performed.Individuals of Sci. sericans may also have been active fromweek 48 onwards; however, this is not indicated in Figure 4abecause of the loss of a group of specimens prior to havingtheir identities confirmed. Catches of Ser. micans during week2 were larger than those during week 51 and catches duringweek 7 were larger than those during week 6 (W = 132.0,P < 0.001 and W = 207.0, P = 0.047, respectively; Fig. 4b).Catches of Ser. nigrolineata during week 51 were significantlylarger than those during week 2 (W = 351.0, P = 0.040); how-ever, no other pair-wise tests of catches from consecutive weekswere statistically significant (Fig. 4c). Catches of all three ofthese species would likely have been modest to large duringthe 2 weeks when no light trapping was conducted.
Of Sci. sericans, Ser. geminata, Ser. ignota, Ser. micansand Ser. nigrolineata, only the catches of Ser. micans exhibitedstatistically significant fluctuations according to time period(see Table 2). No pair-wise tests of catches from consecutiveweeks were statistically significant and catches in week 7 weretoo few to allow comparison with catches from week 6.
Non-feeding and unknown biology scarabs
Adults of Antitrogus morbillosus (non-feeding) and Austral-obolbus gayndahensis (unknown adult biology) were caughtduring mid- to late summer and late spring to early summer,respectively (Table 2). Only the weekly catches of Ant. mor-billosus exhibited statistically significant weekly fluctuations(H = 13.88, P = 0.008; Fig. 5). Only two pair-wise tests ofcatches from different weeks could be conducted. Of thesetests, catches in week 4 were significantly larger than those inweek 5 (W = 158.5, P = 0.027).
The catches of neither Ant. morbillosus nor Aus. gaynda-hensis exhibited statistically significant variation according totime period (Table 2).
Weekly catches and catches by time period of the other 32species for which ≤40 individuals were caught are summarisedin Table 3. The data are presented in order of the earliestspecies to be caught to the latest species to be caught.
DISCUSSION
This study was primarily conducted to obtain informationabout the summer and nocturnal activity patterns of species ofTa
ble
2A
bund
ance
s of
spe
cies
of
whi
ch ≥
40 in
divi
dual
s w
ere
caug
ht a
ccor
ding
to w
eek
duri
ng s
umm
er w
hen
caug
ht a
nd ti
me
of n
ight
whe
n ac
tive
(not
e: n
o sc
arab
s w
ere
caug
ht d
urin
gth
e pe
riod
18:
00–1
9:45
h)
Spec
ies
Wee
k(s)
cau
ght
19:4
5–21
:30
21:3
0–23
:15
23:1
5–01
:00
01:0
0–02
:45
02:4
5–04
:30
04:3
0–06
:00
HP
Phy
llot
ocus
mac
leay
i49
, 50,
51,
2, 3
, 4, 5
, 6, 7
, 816
831
326
22
56.
230.
284
Aus
tral
obol
bus
gayn
dahe
nsis
50, 5
11
1728
73
4.52
0.34
0A
nopl
ogna
thus
pal
lidi
coll
is50
, 51,
2, 3
, 4, 5
, 6, 7
, 834
278
31
14.
940.
424
Scit
ala
seri
cans
†51
, 2, 3
3769
2816
32
6.89
0.22
9Se
rice
sthi
s ni
grol
inea
ta51
, 2, 3
, 4, 5
, 6, 7
, 839
145
7849
234
6.19
0.28
8Se
rice
sthi
s m
ican
s51
, 2, 3
, 4, 5
, 6, 7
, 8, 9
791
1 66
535
812
445
412
.59
0.02
8Se
rice
sthi
s ge
min
ata
†51
, 2, 3
, 4, 8
1129
73
16.
730.
151
Seri
cest
his
igno
ta2,
3, 4
, 616
91
81.
700.
428
Acr
ossi
dius
tas
man
iae
2, 3
, 4, 5
, 6, 7
, 8, 9
14 7
1117
443
8672
1009
235
377
35.9
0<0
.001
Het
eron
yx c
hlor
otic
a2,
4, 5
, 6, 7
, 827
512
4.31
0.11
6A
ntit
rogu
s m
orbi
llos
us2,
4, 5
, 8, 9
1471
4931
174
4.43
0.48
9A
phod
ius
livi
dus
3, 4
, 5, 6
, 7, 8
, 919
729
874
91
713
.54
0.01
9H
eter
onyx
pra
ecox
3, 4
, 5, 6
, 7, 8
, 913
912
513
41
37.
830.
166
Ata
eniu
s pi
cinu
s6,
7, 8
2417
11.
840.
399
Stat
istic
al r
esul
ts a
re f
or K
rusk
al–W
allis
ran
king
(ad
just
ed f
or ti
es)
test
s of
sca
rab
catc
hes
acco
rdin
g to
tim
e pe
riod
onl
y (a
naly
ses
cond
ucte
d us
ing
all t
rap
catc
h da
ta n
ot to
tals
pre
sent
ed h
erei
n).
†An
addi
tiona
l86
ind
ivid
uals
rep
rese
ntat
ive
of e
ither
of
thes
e tw
o sp
ecie
s w
ere
caug
ht i
n w
eeks
48
(tw
o be
etle
s), 4
9 (s
even
bee
tles)
and
50
(77
beet
les)
but
wer
e lo
st p
rior
to
confi
rmat
ion
of t
he i
nitia
l id
entifi
catio
ns.
Scarabs of the southern tablelands of New South Wales 13
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Heteronyx because members of the genus are pests of newlyplanted bluegums. The two species of concern in Tasmania areHeteronyx crinitus and Het. dimidiata that we did not catch atour study site (Hurley & Patel 2003). We did, however, catchvarying numbers of another 13 species of Heteronyx. BothHet. chlorotica and Het. praecox were most abundant frommid- to late summer. Of the other species of Heteronyx, thereappeared to be a trend towards early to mid-summer activity.
Concerning the nocturnal activity patterns of Heteronyx, thecatches of Het. chlorotica peaked between 19:45 and 21:30 hand those of Het. praecox peaked between 21:30 and 23:15 h.Heteronyx chlorotica was not caught after 01:00 h whereasHet. praecox was caught at all times between sunset and sun-rise, although in lesser numbers than earlier in the night. Theother less abundant species of Heteronyx were caught between19:45 and 23:15 h with most caught between 21:30 and23:15 h.
Because Allsopp and Logan (1999) monitored a suiteof scarab species endemic to southern Queensland, only oneof the species we commonly caught was also caught bythese authors. Allsopp and Logan (1999) reported thatAno. pallidicollis exhibited prolonged summer activity, whichagrees with our findings. Similarly, they too did not find anycorrelation between catches of Ano. pallidicollis and rainfall(analyses not shown). In contrast, Carne et al. (1981) reportedthat the abundance of Ano. montanus was correlated withrainfall. Allsopp and Logan (1999) did, however, catch otherspecies of genera also caught by us, i.e. other species ofAnoplognathus and Antitrogus. According to them, Antitrogusconsanguineus exhibited brief spring activity, Antitrogus par-vulus exhibited brief summer activity and Anoplognathusboisduvalii and Ano. porosus both also exhibited prolongedsummer activity, i.e. as did Ano. pallidicollis. The species ofAntitrogus we caught, Ant. morbillosus, was active betweenmid- and late summer. The suite of species monitored byRoberts et al. (1982a) is typical of the northern tablelands ofNew South Wales but nevertheless has a number of species incommon with our study. However, Roberts et al. (1982a)present year-to-year fluctuations in abundance, thereby pre-venting comparisons of summer activities of species in com-mon with our study.
Both Anoplognathus and Phyllotocus are widely regardedas diurnal; however, modest numbers of Ano. pallidicollis andlarge numbers of Phy. macleayi were attracted to our traps
Fig. 4. Weekly light trap catches of three purportedly eucalypt-feeding scarabs, namely (a) Scitala sericans, (b) Sericesthismicans and (c) Sericesthis nigrolineata (all data aremeans ± standard errors). Note different scales on y-axes.
(a)
(b)
(c)
Fig. 5. Weekly light trap catches of the non-feeding adults ofAntitrogus morbillosus (data are means ± standard errors).
14 M J Steinbauer and T A Weir
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Tabl
e 3
Abu
ndan
ces
of s
peci
es o
f w
hich
≤40
ind
ivid
uals
wer
e ca
ught
acc
ordi
ng t
o w
eek(
s) w
hen
caug
ht a
nd t
ime
of n
ight
whe
n ac
tive
(not
e: n
o sc
arab
s w
ere
caug
ht d
urin
g th
e pe
riod
18:0
0–19
:45
h)
Spec
ies
Wee
k(s)
cau
ght
19:4
5–21
:30
21:3
0–23
:15
23:1
5–01
:00
01:0
0–02
:45
02:4
5–04
:30
04:3
0–06
:00
Phy
llot
ocus
rufi
penn
is48
2E
leph
asto
mus
mer
aldu
s48
, 49
11
Ont
hoph
agus
sp.
nov
. nr.
pent
acan
thus
491
Het
eron
yx a
rato
r49
, 51,
84
Het
eron
yx e
xcis
us49
, 2, 3
, 6, 7
, 87
91
1H
eter
onyx
aus
tral
is50
, 51
31
Het
eron
yx a
equa
lis
50, 5
1, 2
, 73
41
Om
orgu
s co
stat
us50
, 51,
2, 3
, 5, 6
, 7, 8
610
13
Het
eron
yx s
p.50
, 51,
3, 4
, 53
8A
nopl
ogna
thus
chl
orop
yrus
50, 5
1, 3
, 4, 6
, 7, 8
1023
21
2H
eter
onyx
agr
icol
a50
, 4, 6
14
Ano
plog
nath
us v
elut
inus
511
Het
eron
yx s
imiu
s51
1Se
man
opte
rus
subc
osta
tus
511
Ano
plog
nath
us h
irsu
tus
†51
, 2, 3
138
1A
nopl
ogna
thus
mon
tanu
s51
, 2, 4
22
Seri
cest
his
sutu
rali
s51
, 2, 3
, 4, 5
, 63
710
11
Ano
plog
nath
us s
utur
alis
†51
, 2, 4
, 6, 7
1010
111
Oph
ropy
x hi
spid
a51
, 2, 5
, 7, 8
23
31
Trox
sca
ber
51, 6
11
Ano
plog
nath
us p
inda
rus
21
Seri
cest
his
para
llel
a2
2H
eter
onyx
gra
num
2, 3
, 5, 7
, 84
154
11
Het
eron
yx c
ervi
na3
1W
ebbe
lla
sp.
4, 5
, 7, 8
716
31
Aut
omol
ius
vulg
aris
51
Das
ygna
thus
tri
tube
rala
tus
51
Het
eron
yx s
p. n
ov.
61
Het
eron
yx s
p. n
ov. [
Key
14]
7, 8
3H
eter
onyx
gra
nula
tus?
84
Mae
chid
ius
sp.
81
Phy
llot
ocus
bim
acul
atus
86
11
†An
addi
tiona
l 45
ind
ivid
uals
rep
rese
ntat
ive
of e
ither
of
thes
e tw
o sp
ecie
s w
ere
caug
ht d
urin
g w
eeks
49
(tw
o be
etle
s) a
nd 5
0 (4
3 be
etle
s) b
ut w
ere
lost
pri
or t
o co
nfirm
atio
n of
the
ini
tial
iden
tifica
tions
.
Scarabs of the southern tablelands of New South Wales 15
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
which seems at odds with this view. It is possible that individ-uals of both species that were residing on the nearby treesmight have become aroused by our trap’s light and beencaught by them. However, only small to modest numbers(i.e. 1–12 Anoplognathus per tree in December 1999 (17 outof 500 trees) and January 2000 (6 out of 500 trees)) ofAno. pallidicollis and almost no Phy. macleayi were observedon the nearby eucalypts during the entire trapping period (MJSteinbauer pers. obs. 2004). This would appear to suggest thatthese species were attracted to our traps from areas outside thefield trial. Large numbers of adults of a species of Anoplog-nathus have been observed flying at night to conspecificbeetles crushed by traffic beneath a large Eucalyptuspolyanthemos in Cook, Australian Capital Territory (ACT)(MJ Steinbauer pers. obs. 2001). Based on the roles of plantand insect volatile compounds in the attraction of males andfemales of the forest cockchafer and the European cockchaferto aggregations of feeding and mating conspecifics (e.g.Ruther et al. 2000, 2002; Reinecke et al. 2002), it is possiblethat these Anoplognathus adults were attracted by the largequantities of volatiles released by their crushed counterpartsthat they could have been using for indicating the location ofa large mating aggregation. That some species of Anplog-nathus are attracted to eucalypts where conspecifics are feed-ing and mating is supported by their highly variable pattern ofabundance, e.g. of 249 E. globulus surveyed in a nearby fieldtrial on 1 February 2001, adult Anoplognathus were presenton 23 trees and their abundance on these trees varied between1 and 30 beetles (19 trees) while three trees had 50 beetles andone tree had 200+ beetles (combined mean and standard devi-ation 27.1 ± 40.7). If mating aggregations are shown to format night, it would indicate that at least some species of Anop-lognathus are not exclusively diurnal. Whether Phy. macleayiflies at night to find mates or for some other reason cannot bespeculated upon at this time.
Acrossidius tasmaniae was by far the most commonlycaught species. Carne (1957c; as Aphodius howitti Hope) sug-gested that large numbers of Acr. tasmaniae occur when thereis a combination of below-average yearly rainfall duringwhich rainfall does not exceed 100 mm per month. Much of2003 fulfilled both these criteria which may have been thereason for the large numbers of this beetle that were caughtduring the first 9 weeks of 2004 (also evident from the numer-ous enquiries from the public to CSIRO Entomology andmedia articles such as that in The Canberra Times, Saturday,24 January 2004). Similar conditions in 2005 resulted in largenumbers of this species occurring again in February 2006 inthe ACT and surrounding areas (TA Weir pers. obs. 2006).
Commercial plantation companies often plant E. globulusseedlings in late winter and early spring when soil moisture isless likely to be limiting to their establishment; the trees aremost vulnerable to the damaging effects of scarab attack in thefirst 12 months after planting (Hurley & Patel 2003). Accord-ing to Matthiessen (unpubl. data 2006), the ground-dwellinglarvae of Heteronyx elongatus have damaged seedling euca-lypts in Western Australia by chewing their roots while com-plexes of Heteronyx species adults are causing ever more
damage to bluegums of all ages, particularly around Albany.MJS and colleagues have planted over 2000 eucalypts (com-prising mostly E. globulus and E. nitens) in three field trialsin the location where this study was conducted but no evidenceof scarab defoliation was seen during the first 12 months posttheir being planting. Some trees were attacked at least onceby various species of Anoplognathus in their second and latergrowing seasons; for example, on 20 January 2000, 11% ofthe eucalypts in the field trial used for this study had hadanywhere between 10% and 100% of all leaves removed byChristmas beetles (MJ Steinbauer unpubl. data 2000). Smallnumbers of Automolius vulgaris, Liparetrus asper andLip. fulvohirtus were seen and caught in the funnel traps weused during the period of the study but they did not causesignificant defoliation (MJ Steinbauer unpubl. data 2004). Ourlight trap catches and observations suggest that were planta-tion companies to monitor scarab beetle populations usinglight traps of the same specifications as those that we used,they would likely need to be catching greater numbers ofeucalypt-feeding species than we caught before becomingconcerned about the level of defoliation the trees might beexperiencing. For example, they may need to be catchingsimilar numbers of eucalypt-feeding species as we caught ofSer. micans. If Acr. tasmaniae was a eucalypt-feeding species,it seems likely that in the numbers they were caught theywould have inflicted substantial defoliation on the trees in ourfield trial. The relationships between light trap catches of euca-lypt-feeding scarabs, tree age and/or size and level of defoli-ation needs focused investigation before time-consumingground surveys could be replaced with routine operation andinspection of light traps in plantations of seedling E. globulus.Light traps offer some considerable advantages over groundsurveys, which makes this a worthy objective. Specifically,light traps can largely negate staff variations in estimatinginsect abundances, they can simultaneously sample of numberof species (e.g. Heteronyx species, autumn gum moth andspecies of nocturnal wasp parasitoids) and the insects caughtare able to be retained for forwarding to relevant experts foridentification should they be unfamiliar to plantation staff (afar more reliable means of obtaining an identification thanfrom verbal descriptions or even a photograph).
ACKNOWLEDGEMENTS
We thank CSIRO Plant Industry, Ben Boyd, Brett Brewer,John Dowse, Bob Edmonds, Mark Hilhorst, Rex Sutherland,Craig Szabadics and Colin Tann for their advice and assis-tance. Rolf Oberprieler, Kim Pullen and Keith Wardhaughreviewed an earlier draft of the manuscript.
REFERENCES
Allsopp PG. 2003. Synopsis of Antitrogus Burmeister (Coleoptera: Scar-abaeidae: Melolonthini). Australian Journal of Entomology 42, 159–178.
16 M J Steinbauer and T A Weir
© 2007 The AuthorsJournal compilation © 2007 Australian Entomological Society
Allsopp PG & Logan DP. 1999. Seasonal flight activity of scarab beetles(Coleoptera: Scarabaeidae) associated with sugarcane in southernQueensland. Australian Journal of Entomology 38, 219–226.
Bashford R. 1993. Insect pest problems of eucalypt plantations in Aus-tralia. 4. Tasmania. Australian Forestry 56, 375–377.
Britton EB. 1957. A Revision of the Australian Chafers (Coleoptera:Scarabaeidae: Melolonthinae), Vol. 1. British Museum (NaturalHistory), London, UK.
Britton EB. 1980. A revision of the Australian chafers (Coleoptera: Scar-abaeidae: Melolonthinae), 3. Tribe Liparetrini: genus Liparetrus. Aus-tralian Journal of Zoology, Supplementary Series 76, 1–209.
Britton EB. 1987. A revision of the Australian chafers (Coleoptera: Scar-abaeidae: Melolonthinae), Vol. 5. Tribes Scitalini and Comophorinini.Invertebrate Taxonomy 1, 685–799.
Britton EB. 1988. Synopsis of the genera of Australian Heteronychini(Coleoptera; Scarabaeidae: Melolonthinae). Journal of the AustralianEntomological Society 27, 27–36.
Britton EB. 2000. A review of Heteronyx Guérin-Méneville (Coleoptera:Scarabaeidae: Melolonthinae). Invertebrate Taxonomy 14, 465–589.
Bulinski J & Matthiessen JN. 2002. Poor efficacy of the insecticide chlo-rpyrifos for the control of African black beetle (Heteronychus arator)in eucalypt plantations. Crop Protection 21, 621–627.
Carne PB. 1957a. A Systematic Revision of the Australian Dynastinae(Coleoptera: Scarabaeidae). CSIRO Publishing, Melbourne,Australia.
Carne PB. 1957b. A revision of the ruteline genus Anoplognathus Leach(Coleoptera: Scarabaeidae). Australian Journal of Zoology 5, 88–143.
Carne PB. 1957c. An ecological study of the pasture scarab Aphodiushowitti Hope. Australian Journal of Zoology 4, 259–314.
Carne PB. 1958. A review of the Australian Rutelinae (Coleoptera: Scar-abaeidae). Australian Journal of Zoology 6, 162–240.
Carne PB. 1965. A revision of the genus Elephastomus Macleay(Coleoptera: Geotrupidae). Journal of the Entomological Society ofQueensland 4, 3–13.
Carne PB & Chinnick LJ. 1957. The pruinose scarab (Sericesthis pruinoseDalman) and its control in turf. Australian Journal of AgriculturalResearch 8, 604–616.
Carne PB, Greaves RTG & McInnes RS. 1974. Insect damage to planta-tion-grown eucalypts in north coastal New South Wales, with partic-ular reference to Christmas beetles (Coleoptera: Scarabaeidae).Journal of the Australian Entomological Society 13, 189–206.
Carne PB, McInnes RS & Green JP. 1981. Seasonal fluctuations in theabundance of two leaf-eating insects. In: Eucalypt Dieback in Forestsand Woodlands (eds KM Old, GA Kile & CP Ohmart), pp. 121–126.CSIRO Publishing, Melbourne, Australia.
Caveney S, Scholtz CH & McIntyre P. 1995. Patterns of daily flightactivity in onitine dung beetles (Scarabaeinae: Onitini). Oecologia103, 444–452.
Davis AJ. 1999. Species packing in tropical forests: diel flight activity ofrainforest dung-feeding beetles (Coleoptera: Aphodiidae, Scara-baeidae, Hybosoridae) in Borneo. Raffles Bulletin of Zoology 47,473–486.
Donaghey RH. 1998. Associations among plants, birds and insects. Aus-tralian Plants on Line. [Cited 19 May 2004.] Available from URL:http://farrer.riv.csu.edu.au/ASGAP/APOL10/jun98-4.html
Floyd RB, Farrow RA & Matsuki M. 2002. Variation in insect damageand growth in Eucalyptus globulus. Agricultural and Forest Entomol-ogy 4, 109–115.
Goodyer GJ. 1985. Scarab grubs in northern tableland pastures. Agfact 1,1–11.
Hardy RJ. 1977. The biology and behaviour of the pasture beetle Scitalasericans Erichson (Scarabaeidae: Melolonthinae). Journal of the Aus-tralian Entomological Society 15, 433–440.
Houston WWK & McIntyre P. 1985. The daily onset of flight in thecrepuscular dung beetle Onitis alexis. Entomologia Experimentalis etApplicata 39, 223–232.
Howden HF. 1992. A revision of the Australian beetle genera EucanthusWestwood, Bolbobaineus Howden & Cooper, Australobolbus
Howden & Cooper and Gilletinus Boucomont (Scarabaeidae:Geotrupinae). Invertebrate Taxonomy 6, 605–717.
Howden HF, Howden AT & Storey RI. 1991. Nocturnal perching ofScarabaeine dung beetles (Coleoptera, Scarabaeidae) in an Australiantropical rain forest. Biotropica 23, 51–57.
Hurley M & Patel V. 2003. Effect of imidacloprid on the growth ofEucalyptus nitens seedlings. Australian Forestry 66, 100–101.
Landsberg J, Morse J & Khana P. 1990. Tree dieback and insect dynamicsin remnants of native woodlands on farms. Proceedings of the Eco-logical Society of Australia 16, 149–165.
Lawrence JF & Britton EB. 1991. Coleoptera (Beetles). In: The Insectsof Australia. A Textbook for Students and Research Workers, pp. 543–683. CSIRO Publishing, Melbourne, Australia.
Lawson SA & King J. 2003. Swarming scarab beetles. DPI Note. [Cited19 May 2004.] Available from URL: http://www.dpi.qld.gov.au/hard-woodsqld/9353.html
Loch AD & Floyd RB. 2001. Insect pests of Tasmanian blue gum,Eucalyptus globulus globulus, in south-western Australia: history,current perspectives and future prospects. Austral Ecology 26, 458–466.
Matthews EG. 1984. A Guide to the Genera of Beetles of South AustraliaPart 3 Polyphaga: Eucinetoidea, Dascilloidea and Scarabaeoidea.South Australian Museum, Adelaide, Australia.
Neumann FG. 1993. Insect pest problems of eucalypt plantations in Aus-tralia. 3. Victoria. Australian Forestry 56, 370–374.
Reinecke A, Ruther J & Hilker M. 2002. The scent of food and defence:Green leaf volatiles and toluquinone as sex attractant mediate matefinding in the European cockchafer, Melolontha melolontha. EcologyLetters 5, 257–263.
Roberts RJ, Ridsdill-Smith TJ, Porter MR & Sawtell NL. 1982a. Fluctu-ations in the abundance of pasture scarabs over an 18-year period oflight trapping. In: Proceedings of the 3rd Australasian Conference onGrassland Invertebrate Ecology (ed. KE Lee), pp. 75–79. SouthAustralian Government Printer, Adelaide, Australia.
Roberts RJ, Campbell AJ, Porter MR & Sawtell NL. 1982b. The distri-bution and abundance of pasture scarabs in relation to Eucalyptustrees. In: Proceedings of the 3rd Australasian Conference on Grass-land Invertebrate Ecology (ed. KE Lee), pp. 207–214. South Austra-lian Government Printer, Adelaide, Australia.
Ruther J, Reinecke A, Thiemann K, Tolasch T & Hilker M. 2000. Matefinding in the forest cockchafer, Melolontha hippocastani, mediatedby volatiles from plants and females. Physiological Entomology 25,172–179.
Ruther J, Reinecke A & Hilker M. 2002. Plant volatiles in the sexualcommunication of Melolontha hippocastani: response towards timedependent bouquets and novel function of (Z)-3-hexen-1-ol as asexual kairomone. Ecological Entomology 27, 76–83.
Scholtz CH. 1986. Revision of the genus Trox Fabricius (Coleoptera:Trogidae) of the Australasian region. Australian Journal of Zoology,Supplementary Series 125, 1–99.
Stebnicka ZT & Howden HF. 1995. Revision of Australian genera in thetribes Aphodiini, Aegialiini and Proctophanini (Coleoptera: Scara-baeidae: Aphodiinae). Invertebrate Taxonomy 9, 709–766.
Stebnicka ZT & Howden HF. 1997. Revision of the Australian species ofAtaenius Harold (Coleoptera: Scarabaeidae: Aphodiinae: Eupariini).Invertebrate Taxonomy 11, 735–821.
Steinbauer MJ. 2003. Using ultra-violet light traps to monitor autumngum moth, Mnesampela privata (Lepidoptera: Geometridae), insouth-east Australia. Australian Forestry 66, 279–286.
Stone C. 1993. Insect pest problems of eucalypt plantations in Australia.2. New South Wales. Australian Forestry 56, 363–369.
Wallace CR. 1946. The black beetle as it affects coastal dairy farmers.Agricultural Gazette of New South Wales 23, 121–124.
Wensler RJ. 1971. Biology and rearing of the scarabaeid, Sericesthisgeminata (Coleoptera). Journal of the Australian Entomological Soci-ety 10, 285–289.
Accepted for publication 21 May 2006.
