Introduction, Rearing, and Host Range of Aerenicopsis championi Bates

(Coleoptera: Cerambycidae) for the Biological Control of Lantana

camara L. in Australia
W. A. Palmer,* B. W. Willson,† and K. R. Pullen‡
*Alan Fletcher Research Station, Queensland Department of Natural Resources, 27 Magazine Street, Sherwood, Queensland,
4075, Australia; †33 Ben Street, Chermside, Queensland, 4032, Australia; and ‡Division of Entomology, CSIRO,
P.O. Box 1700, Canberra, ACT, 2601, Australia

Received March 10, 1999; accepted September 8, 1999

been not been achieved in Australia nor in many other

The biology and host range of the cerambycid beetle
Aerenicopsis championi Bates, a potential biological
control agent for the weed Lantana camara L., were
studied. A. championi is a univoltine species associ-
ated with L. camara, L. urticifolia, and L. hirsuta in
Mexico and Central America. In Mexico, adult emer-
gence occurred in May and June at the start of the
rainy season. Larvae fed within the stems over a 9- to
12-month period and caused damage to the plant. The
insect was imported into Australia, where a procedure
for rearing it in the laboratory was developed. Host-
range tests indicated that adults oviposited and larvae
commenced development in L. camara and L. monte-
vidensis but not in any of 57 other species tested. A
CLIMEX model indicated that most areas infested with
lantana in Australia would have a favorable climate for
A. championi. Permission to release this insect in
Australia was obtained and three small releases were
made in southern Queensland in February 1995. r 2000
Academic Press
Key Words: biological control; Lantana; Aerenicopsis
championi; host range; biology; introduction; CLIMEX.
countries.
The stem borer Aerenicopsis championi Bates (Coleop-
tera: Cerambycidae) was first found in L. camara in
Mexico in 1902 by Albert Koebele, who described it as a
‘‘horn-tailed larva’’ (Perkins and Swezey, 1924). It was
released in Hawaii in 1902 (perhaps the first biological
control agent for a weed to be released in the United
States) and again in 1955 but did not establish on
either occasion (Perkins and Swezey, 1924; Fullaway,
1956).
We decided to introduce this insect into Australia
because it had many characteristics of a good control
agent. This necessitated collecting the insect from
Mexico, understanding its biology and phenology, devel-
oping a suitable method for rearing it in the laboratory,
reviewing the literature and any host information on
museum specimens, conducting new host-range tests to
satisfy Australian authorities that it was safe to re-
lease, and releasing the insect. This paper describes
these procedures.

MATERIALS AND METHODS

INTRODUCTION
The woody shrub known as lantana, Lantana camara
L. (family Verbenaceae), infests millions of hectares of
grazing and cropping land in 47 countries (Holm et al.,
1977) and is regarded as one of the world’s 10 worst
weeds (Muniappan et al., 1992). L. camara has long
been a target for biological control, with the first project
started in 1902, followed by others at intervals through-
out the century. These investigations (Perkins and
Swezey, 1924; Mann, 1954; Winder and Harley, 1983;
Palmer and Pullen, 1995) have resulted in 36 insect
species being released in various countries (Julien and
Griffiths, 1998), a number easily surpassing that for
any other weed species. However, effective control has
Literature review. Information about the biology,
geographic distribution, and host range of A. cham-
pioni and its congeners was sought by literature review
and by examining specimens in major collections, includ-
ing the Smithsonian, Texas A&M University, and the
Universidad Nacional Autónoma de México.
Biology and phenology. Field observations on its
phenology, distribution, and biology in Mexico were
made whenever possible during a 4-year survey of the
insects associated with lantana. One area near Al-
varado, 50 km south of the city of Veracruz, Mexico, was
visited repeatedly over a 2-year period to observe the
insect’s phenology under natural conditions and to
collect material for shipment to Australia. As the insect

227 1049-9644/00 $35.00

Copyright r 2000 by Academic Press
All rights of reproduction in any form reserved.
228 PALMER, WILLSON, AND PULLEN
feeds within woody stems, much of its life cycle was not
easily amenable to close observation.
Development of laboratory method. A laboratory
method for rearing the insect was developed at the Alan
Fletcher Research Station in Brisbane to provide mate-
rial for host-range tests within the quarantine facility
and to meet the regulatory requirement that the insect
be reared through a full generation in quarantine
before being released.
Initially, late-instar larvae, pupae, and teneral adults
were shipped in May and June 1989–1991 from Al-
varado to the Alan Fletcher Research Station, where
emerging adults were released into cages of potted
lantana. This method was not satisfactory because it
was difficult to collect a sufficient quantity of insects, a
high proportion of those collected was parasitized, and
other mortality factors were also significant. It was
therefore decided to collect the much more abundant
early instars later in the year. These were shipped in
the stems in which they were found. Five shipments
were made between August 1992 and January 1993,
with approximately 200 larvae surviving shipment. On
arrival in Brisbane they were transferred into artifi-
cially prepared galleries bored into cut stems of lan-
tana. These cut stems were prepared by selecting
60-cm lengths of 4- to 5-mm-diameter stem of the
‘‘Helidon White’’ variety. The stems were stripped of
leaves and placed immediately into water. Within 24 h
the stem lengths were treated with a rooting hormone
gel containing 0.3% indole butyric acid and then stood
in a sand–peat mixture.
Emerging adults were placed onto potted ‘‘Helidon
White’’ lantana plants on which they oviposited. How-
ever, it was soon noted that excessive sap flow in these
potted plants caused high mortality, particularly of
early instars, and attempts to rear the larvae on
artificial diet (Harley and Willson, 1968) were also
unsuccessful. Larvae were therefore recovered from the
potted plants and reared by the cut-stem method
described above. At least two changes of stems were
required to complete development. When feeding ceased,
the stems were held until the adults emerged.
Host-range studies. A study of the insect’s host
range was conducted at the Alan Fletcher Research
Station to ensure that the insect was safe for release in
Australia. The plant species used for this test were
selected by the centrifugal phylogenetic method (Wap-
shere, 1975), whereby closely related taxa are repre-
sented by proportionately larger number of species in
the testing program. Of the 59 species included in the
tests (Table 1), 6 were members of the Verbenaceae,
while an additional 17 species were from the very
closely related Lamiaceae. The genus Clerodendrum
was particularly well represented because Chock and
Chong (Hawaiian Department of Agriculture and For-
estry, unpublished) observed oviposition in Cleroden-
drum.
Ten adults from the laboratory colony were placed in
a gauze cage (90 3 90 3 55 cm) containing eight test
plants and one plant of L. camara (‘‘Helidon White’’) as
a control. The plants were observed every 2 or 3 days
and any feeding by the adults was noted. After 14 days,
the plants were closely examined for oviposition and
adult feeding. Any plant on which oviposition was
observed was retained and the subsequent larval devel-
opment was noted. By this method, each species in
Table 1 was tested on two occasions, a process that
required 16 separate cage tests.
Climate matching for Australian release. The cli-
mate matching model CLIMEX (Skarratt et al., 1995)
was used to assess where the insect was likely to be
effective in Australia and as an aid to the selection of
release sites. This model uses an integrated climate
index to describe the climate suitability of an area for
the species. The climate profile of the insect was first
determined by recursively trying various sets of param-
eter values until the model’s distribution matched the
known distribution of the insect in North America. The
set of parameters was then used to predict the potential
distribution of the insect in Australia.
Release. Application was made to the Australian
Quarantine Inspection Service (AQIS) and Environ-
ment Australia to release this insect in Australia. At
the conclusion of the present study, three releases, each
of about 20 adults, were made in February 1995 in the
Brisbane area and its hinterland at Gatton (where
‘‘Helidon White’’ was predominant), Jimboomba, and
Greenbank. On each occasion a cage was placed over a
suitable lantana plant and the adults were released
within. At one of these sites (Greenbank, Brisbane) 40
larvae were also placed into plants by first drilling
selected stems to provide a suitable gallery for them.
RESULTS
Literature review. The neotropical genus Aerenicop-
sis Bates (Cerambycidae: Lamiinae: Aerenicini) con-
sists of nine species distributed from Mexico to South
America (Martins, 1984), with two species, A. cham-
pioni and A. costaricensis Breuning, known from Mexico
or Central America (Chemsak et al., 1992). A. cham-
pioni is endemic to Mexico and Central America. Lar-
vae and adults were first associated with lantana in
Mexico by Koebele (Perkins and Swezey, 1924). In 1954,
N. Krauss collected it from lantana in Mexico, Guate-
mala, Honduras, and El Salvador, where it was found
in well-drained situations at low elevations (,122 m),
so that it was considered to be a species of the immedi-
ate coastal strip (Mann, 1954). There is no record of A.
championi having hosts other than Lantana spp. and
 INTRODUCTION, REARING, AND HOST RANGE OF A. championi 229

TABLE 1—Continued
TABLE 1
Plant Species against Which the Oviposition and Adult No. of
Feeding Behavior of Aerenicopsis championi Were Tested in leaves No. of
No. of attacked eggs
Brisbane to Satisfy Requirements for its Release in Australia
Plant species replications by adults laid
No. of
Mimosaceae:
leaves No. of
Acacia podalyriifolia A. Cunn. ex
No. of attacked eggs
G. Don 2 0 0
Plant species replications by adults laid
Myoporaceae:
Myoporum parvifolium R. Br. 2 0 0
Annonaceae:
Myrtaceae:
Annona 3 squamosa L. 2 0 0
Eucalyptus microtheca F. J. Muell. 2 0 0
Apiaceae:
Oleaceae:
Daucus carota L. 2 0 0
Ligustrum sinense Lour. 2 0 0
Asteraceae:
Passifloraceae:
Carthamus tinctorius L. 2 0 0
Passiflora edulis Sims. 2 0 0
Helianthus annuus L. 2 0 0
Pedaliaceae:
Lactuca sativa L. 2 0 0
Sesamum indicum L. 2 0 0
Avicenniaceae:
Poaceae:
Avicennia marina (Forssk.) Vierh. 2 0 0
Cenchrus ciliaris L. 2 0 0
Bignoniaceae:
Sorghum bicolor (L.) Moench 2 0 0
Jacaranda acutifolia Humb. &
Triticum aestivum L. 2 0 0
Bonpl. 2 0 0
Zea mays L. 2 0 0
Pandorea pandorana (Andr.)
Rosaceae:
Steenis. 2 0 0
Prunus persica (L.) Batsch. 2 0 0
Boraginaceae:
Rosa sp. 2 0 0
Cordia dichromata G. Forster 2 0 0
Scrophulariaceae:
Ehretia acuminata R. Br. 2 0 0
Duboisia myoporoides R. Br. 2 0 0
Brassicaceae:
Solanaceae:
Brassica oleracea L. 2 0 0
Lycopersicon lycopersicum (L.)
Chenopodiaceae:
Karst. ex Farw. 2 0 0
Beta vulgaris L. 2 0 0
Solanum tuberosum L. 2 0 0
Chloanthaceae:
S. nemophilum F. Muell. 2 0 0
Chloanthes parviflora Walp. 2 0 0
Theaceae:
Spartothamnela juncea (Cunn. ex
Camellia sinensis (L.) O. Kuntze. 2 0 0
Walp.) Briq. 2 0 0
Verbenaceae:
Fabaceae:
Duranta erecta L. 2 0 0
Arachis hypogaea L. 2 0 0
Lantana camara L. 16 85.3 48.6
Glycine max (L.) Merrill 2 0 0
L. montevidensis (K. Spreng.) Briq. 2 3.0 1
Macroptilium atropurpureum
Phyla nodiflora (L.) Greene 2 0 0
(D.C.) Urban 2 0 0
Stachytarpheta cayennensis (Rich.)
Medicago sativa L. 2 0 0
Vahl. 2 0 0
Phaseolus vulgaris L. 2 0 0
Verbena bonariensis L. 2 0 0
Lamiaceae:
Ajuga australis R. Br. 2 0 0
Calicarpa pedunculata R. Br. 2 0 0
Clerodendrum cunninghamii
Benth. 2 0 0 nothing is known of the biology or host range of any of
C. speciosissimum Van Geert 2 0 0 its congeners.
C. floribundum R. Br. 2 0 0 In an unpublished Hawaiian study in 1955, Q. C.
C. inerme (L.) Gaertn. 2 0 0 Chock and M. Chong conducted tests on the adult
C. tomentosum (Venten.) R. Br. 2 0 0
Faradaya splendida F. Muell. 2 0 0
feeding and oviposition range of the insect. Their plant
Glossocarya hemiderma (F. Muell. test list comprised 25 species, most of which were
ex Benth.) Benth. & J. D. Hook. 2 0 0 economically important horticultural plants that are
Gmelina leichardtii (F. Muell.) distantly related to Lantana. Also included were the
Benth. 2 0 0 more closely related Clerodendrum sp., Duranta sp.,
Mentha 3 peperita L. 2 0 0
Plectranthus argentatus S. T.
and Vitex sp. They confined 10 adults (collected in
Blake 2 0 0 Mexico) on a cutting of each plant species after first
Premna lignum-vitae (Cunn. ex confirming successful oviposition on a lantana cutting.
Shauer) Pieper 2 0 0 After 48 h any oviposition or adult feeding was recorded
Prostanthera ovalifolia R. Br. 2 0 0 before the insects were returned to lantana to again
Salvia coccinea Juss. ex J. Murray 2 0 0
Teucrium argutum R. Br. 2 0 0 confirm ability to oviposit. A. championi oviposited
Westringia fruticosa (Willd.) Druce 2 0 0 successfully only on Clerodendrum sp. Some nibbling
and gnawing by the adults was observed on Gmelina
230 PALMER, WILLSON, AND PULLEN
sp., Persea americana Mill., Leucaena leucocephala
(Lam.) de Wit, and Pimenta sp., but this was considered
inconsequential. These scientists did not rear the insect
through a complete generation in the laboratory. The
insect was released in Hawaii on their recommenda-
tion, using further material collected as adults in
Mexico, but, again, it did not establish (Fullaway,
1956).
Biology and phenology. The egg is pale yellow,
approximately 0.8 3 1.8 mm, and tapers at both ends.
The female inserts eggs singly into the petioles of
terminal leaves and occasionally into tender stems,
flower peduncles, or the midrib of larger leaves. Eclo-
sion occurs from 6 days onward.
Immediately after eclosion, the larva, which has a
characteristic, highly chitinized ‘‘horn tail,’’ feeds down
through the petiole into the terminal shoot that withers
and dies after approximately 2 weeks of larval feeding.
This period in the terminal shoot is a critical time in the
life cycle of the insect and over 90% mortality has been
observed in the laboratory when sap flow was exces-
sive. The larva then makes its way to a woody main
stem of the plant. As it continues feeding, it makes
small holes in the side of the stem, through which it
pushes frass. Larvae can feed along 1–2 m of the stem
and cause it to die progressively and break off. Feeding
can continue for up to 6 months before the larva seals
off its tunnel with chewed fiber and enters a bright
yellow, diapausing prepupal stage.
Pupation occurs within the chamber 9–12 months
after eclosion of the egg. The pupa is naked and yellow,
FIG. 1. Locations (q) at which Aerenicopsis championi was collected in Mexico in relation to four major regional cities (
up to 16 mm in length and 2–3 mm in diameter. The
pupal stage lasts 10–14 days before adult eclosion
occurs. The teneral adult remains in the chamber for a
further 2 weeks before eating its way out. The adult
beetles, brown with diagonal bands of grey, live for up
to 3 months and feed on the midrib and veins of leaves,
petioles, and terminal shoots.
At the site at Alvarado, larvae, pupae, and teneral
adults were found within stems in May prior to the
commencement of the rainy season in June. By June,
mainly adults were found in the stems, while in the
following month the characteristic terminal damage of
the early instars was observed. Larvae developed
throughout the July–December period, with one ten-
eral adult being found as early as December.
Collections of late-instar larvae and pupae in May
and June at Alvarado indicated that A. championi was
heavily parasitized. One ichneumonid parasite Ag-
onocryptus chichimecus (Cresson) is known from this
species.
During the course of these studies, larvae were
collected at various locations in Mexico (Fig. 1) from three
very closely related species: L. camara, L. urticifolia Mill.,
and L. hirsuta Mart. & Gal. The insect was found near
Lake Chapala (1500 m) in the state of Jalisco, and at
Jalapa and Orizaba (both nearly 1300 m) in the state of
Veracruz. These locations are at considerably higher alti-
tudes than where it had been previously found.
Development of laboratory method. The cut-stem
method proved satisfactory, and sufficient adults were
reared over a period of 2 years to complete the host-
).
 INTRODUCTION, REARING, AND HOST RANGE OF A. championi 231

range testing and to make three releases. Further develop- TABLE 2

ment of the method was hampered by the relatively small The Parameter Values for the CLIMEX Model Derived for
numbers of insects available and by the long life cycle. Aerenicopsis championi by Recursive Iteration
Host-range studies. In the host-range tests the
adults fed extensively on L. camara, with the number of Index Parameter Value
leaves attacked ranging from 60 to 131 (mean 5 85) per
Temperature DV0 15
plant. Considerably less feeding occurred on L. monte- DV1 22
vidensis, with 3 leaves being attacked in both replica- DV2 28
tions. Feeding was not observed on any other plant DV3 36
listed in Table 1. Moisture SM0 0.2
SM1 0.4
The females oviposited extensively into L. camara
SM2 1.75
with a mean of 48 eggs (range 27 to 66) recorded per SM3 2.5
plant. Two eggs were also oviposited in L. monteviden- Heat stress TTHS 31
sis. Both hatched and the larvae fed for 3 months along THHS 0.003
25 and 30 cm of stem. They were then prevented from Cold stress TTCS 4
THCS 0.02
further feeding by the small diameter of the stems, DTCS 18
which were of normal thickness for this plant. At that DHCS 0.0012
time the larvae were about half the size of those feeding Wet stress SMWS 2.5
on L. camara. Oviposition did not occur on any other HWS 0.002
plant listed in Table 1. Dry stress SMDS 0.2
HDS 0.015
These tests clearly showed that A. championi has a
very narrow host range confined to some species of
Lantana. The host range exhibited by A. championi,
Lamiaceae (Cantino et al., 1992). As a result, the
both in our tests and in those of Chock and Chong, was
Verbenaceae is now represented in Australia by only 10
sufficiently narrow to recommend this insect’s release
naturalized genera and there is no species of this family
in Australia.
that is native to Australia (L. Jessup, Queensland
Climate matching for Australian release. The pa- Herbarium, personal communication). A high propor-
rameter set for the CLIMEX model (Table 2) was tion of the approximately 30 naturalized species of
arrived at by iteration using the known presence of the Verbenaceae are also minor weeds (Kleinschmidt and
insect at Veracruz, Jalapa, Orizaba, Tampico, Guadala- Johnson, 1977). Thus, even if A. championi had a host
jara, and Chilpancingo. The final parameter set indi- range spanning the Verbenaceae, it would present little
cated that this insect would be adversely affected by risk to the Australian flora or to agriculture. This
cold conditions. This parameter set was then used to dearth of close relatives perhaps explains the high
predict favorable areas in Australia. It indicated that number of agents which have been introduced for this
the climate along the Queensland and northern New weed.
South Wales coast from Grafton north to Cape York The investigations into the host range indicated that
should be favorable for the establishment of A. cham- A. championi was very host specific. The formal tests
pioni (Fig. 2). Areas to the south of Grafton would be too indicated that adult feeding and oviposition were con-
cool, while inland areas to the west of this coastal plain fined to L. camara and L. montevidensis. The oviposi-
would be either too hot and dry (in the north) or too cool tion on Clerodendrum sp., reported by Chock and
(in the south). Chong, was not reproduced, even though five Cleroden-
Release. Approval for release in Australia was drum spp. were included in the host test list. As A.
granted by the Australian Quarantine Inspection Ser- championi was found in three Lantana spp. in Mexico,
vice and Environment Australia. The released adults it appears that its host range is confined to Lantana
oviposited successfully and larval galleries and frass spp. (most probably to species in the section camara)
were seen at all three sites during the months following and as such, it presents no risk if released in Australia.
release. Larvae determined to be the second generation A. championi is the second stem-boring cerambycid
were also recovered at the Gatton site in 1997. However, at to be released on lantana in Australia. In 1969, the
later times further evidence of the insect’s presence could lamiine Plagiohammus spinipennis (Thomson) was re-
not be found and it is now considered that these few leased in Queensland and northern New South Wales.
small releases did not lead to establishment. This species established at only one site in New South
Wales, where it may contribute to a level of control
DISCUSSION (Taylor, 1989). A. championi could clearly contribute to
the utilization of this almost vacant feeding niche of the
Recently, the Verbenaceae was redescribed with ma- interior of the stem.
jor elements of the old family being moved into the The CLIMEX model indicated that a very substantial
232 PALMER, WILLSON, AND PULLEN
FIG. 2. Meteorological sites in Australia predicted by the climate matching model CLIMEX to be climatically favorable (d) or climatically
unsuitable (x) for the establishment of Aerenicopsis championi.
portion of the range of lantana in Australia would have
a climate favorable for the permanent establishment of
A. championi. Areas around Brisbane, Bundaberg,
Gladstone, Mackay, Innisfail, Cairns, and Cooktown,
all of which have Ecoclimatic Indices greater than 50,
would be very suitable for release sites. Elsewhere in
the world, the climates of lantana-infested areas of
South Africa, Kenya, and (to a lesser extent) India
would also be suitable for this insect.
One obstacle to the successful establishment of A.
championi may be its inability to utilize some of the
various weedy varieties of lantana found in Australia.
Unfortunately for biocontrol research, there are some
29 varieties of L. camara present in Australia, and it is
known that some of the introduced insects are not
equally effective on all varieties (Harley, 1971). The
relative susceptibility of the varieties to A. championi
is not known and should be addressed. The chances of
successful establishment will be improved by releasing
on the most susceptible varieties in areas climatically
most suitable.
Although this insect is recognized as a promising
biocontrol agent, a full-scale mass-rearing program has
not yet been initiated in Queensland because of other
priorities. The univoltine nature of the life cycle, the
heavy requirement for labor and other resources, and
difficulties in collecting sufficient material in Mexico
have led to other insects being released first. However,
it is anticipated that this insect will be mass-reared in
the future and released along the Queensland coast.
ACKNOWLEDGMENTS
We acknowledge the contributions of J. Cronin and H. Nahrung,
who assisted with the experiments at the Alan Fletcher Research
Station, and M. Day for his advice. The identity of the species was
confirmed by J. Chemsak of the University of California, Berkeley. M.
Hannan-Jones, W. Senaratne, and R. Sutherst (CSIRO, Division of
Entomology) fitted parameters to the CLIMEX model. The study was
supported by funds from the Queensland Rural Lands Protection
Board.
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