The host range of aconophora compressa walker (homoptera: membracidae): a potential biological control agent for lantana Camara L. (verbenaceae)

8 Pages·1996·5.7 MB·

by  W A Palmer

PROC. ENTOMOL. SOC. WASH.
98(4), 1996, pp. 617-624
THE HOST RANGE OF ACONOPHORA COMPRESSA WALKER
(HOMOPTERA: MEMBRACIDAE): A POTENTIAL BIOLOGICAL CONTROL
AGENT FOR LANTANA CAMARA L. (VERBENACEAE)
W. A. PALMER, B. W. WILLSON, AND K. R. PULLEN
(WAP) Tropical Weeds Research Centre, Queensland Department of Natural Resources,
PO. Box 187, Charters Towers, Queensland, Australia 4820; (BWW) Alan Fletcher Re-
search Station, Queensland Department of Natural Resources, 27 Magazine Street, Sher-
wood, Queensland, Australia 4075; (KRP) 112 Irvine Street, Watson, ACT, Australia
2602.
Abstract.-The membracid Aconophora compressa Walker is endemic to the Neotropics
where it is found on various species of Lantana. Host range tests were conducted to
ascertain whether it would be safe to introduce and release this insect in Australia for the
biological control of Lantana camara, a serious weed in many tropical and subtropical
countries. These tests, conducted in both Mexico and Australia, indicated that A. com-
pressa was narrowly stenophagous. In all tests A. compressa oviposited and immatures
developed on L. camara. In one of three tests oviposition and nymphal development
occurred on Duranta repens and in two tests oviposition but not nymphal development
occurred on Jacaranda spp. Oviposition did not occur on 55 other plant species tested.
Permission to release this treehopper was obtained, and it was released in Queensland in
1995.
Key Words: Lantana, biological control, treehopper, Aconophora
The woody shrub Lantana camara L. bicide programs have not been particularly
(family Verbenaceae; subfamily Verbeno- effective. Over a period of nearly 100 years
ideae) is one of the world(cid:8217)s ten worst nox- there has been a number of investigations
ious weeds (Muniappan et al. 1992) and in- (Perkins and Swezey 1924, Winder and
fests millions of hectares of grazing and Harley 1983) resulting in 32 insect species
cropping land in 47 countries (Holm et al. being released in various countries (Julien
1977). Its highly aggressive nature leads to 1992); a number easily surpassing that for
its taking over rangeland (Perkins and any other weed species. However, effective
Swezey 1924, Kleinschmidt and Johnson control has been achieved only in parts of
1977, Cilliers and Neser 1991), particularly Hawaii (Perkins and Swezey 1924, Harley
gullies and hillsides with rich soils. Some 1974). The biological control achieved in
varieties are also poisonous to livestock Australia is considered only partially effec-
(Everist 1974). tive (Winder and Harley 1983) and more
Lantana camara has long been a target biological control agents are desired (Taylor
for biological control because it is an intro- 1989).
duced plant, it infests rangelands where it A survey of the insect fauna on L. ca-
may not be economic to treat with herbi- mara and three closely related congeners in
cides or clear physically, and because her- North America was recently conducted
618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Known host plants of eight species of Aconophora most according to Dietrich and Dietz (1991).
Hosts on which immatures have also been found are denoted with an asterisk.
Species Hosts
. mexicana Stal Anacardiaceae, Fabaceae, Guanabanus sp.*, Piper auritum
. elongatiformis Dietrich Baccharis sp.*, Fabaceae, Piperaceae*
. elongata Dietrich Fabaceae
. compressa Lantana camara,* L. urticifolia*, L. hispida, Lippia myrocephala
. laminata Fairmaire Theobroma cacao, Lupinus sp., Baccharis conferta, Tetracera sp.*
. albipennis Dietrich Juniperus sp., Solanum tuberosum
. robusta Dietrich Alchornea sp.*, Croton gossipifolius*
SbPS.S mBar eginatsa Wa lker Gladiolus
(Palmer and Pullen 1995) to find further bi- over the period of deposition. The female
ological control agents. One species target- remains with this egg mass and its resultant
ed for further study in that investigation progeny for the rest of her life. The eggs,
was the membracid Aconophora compressa each approximately 1.2 < 0.4 mm, hatch
Walker, which was found to be damaging 14-20 days after oviposition. Nymphal de-
plants around Cuernavaca, Mexico. This velopment takes 4(cid:8212)6 weeks and usually in-
paper reports the studies into the treehop- volves 5 instars. The nymphs remain in an
per(cid:8217)s biology and host range that were con- aggregation close to the empty egg mass
ducted before it could be released in Aus- and their mother.
tralia.
Host RANGE AND PHENOLOGY
BIOLOGY Aconophora compressa occurs from Co-
The tribe Aconophorini (Homoptera: lombia in South America, through Central
Membracidae) consists of three genera: America, to Mexico. Three specimens were
Aconophora Fairmaire, Calloconophora collected in Arizona, USA in 1930 (Die-
Dietrich and Guayaquila Goding (Die- trich and Deitz 1991). As this is the only
trich and Deitz 1991). All of the Acono- record of an Aconophorini occurring north
phorini are gregarious and subsocial, that of Mexico it may be regarded with some
is females guard their egg-masses and doubt. In all probability the northern end of
usually remain with nymphal aggrega- its range is Sinaloa, Mexico, some 400 k
tions throughout their development (Die- south of the of the United States border.
trich and Deitz 1991). Females guarding Aconophora compressa is therefore essen-
eggs sometimes aggressively confront tially Neotropical. Within Mexico it is more
predators with buzzing wings and physi- common in central and southern areas. In
cal contact. When disturbed, aggregations the recent survey (Palmer and Pullen 1995)
of adults usually disperse explosively but it was found at Veracruz and Oaxaca but
usually reaggregate, often at the original was most abundant around Cuernavaca in
site. Ant mutualism is associated with the the state of Morelos.
genus Guayaquila but has been observed Aconophora compressa is a multivoltine
with no species of Aconophora (Dietrich species found throughout the year. Popula-
and Deitz 1991). tions increase throughout the growing sea-
Female A. compressa oviposit up to 65 son (June(cid:8212)December) and are greatest at the
eggs to form a single cluster on stems. end of that time and in the ensuing three
These eggs are oviposited over a number of months. Damaging numbers may occur
days in a white froth, which hardens when from November to February.
dry so that the egg mass increases in size The genus Aconophora includes 13 spe-
VOLUME 98, NUMBER 4 619
cies (Dietrich and Deitz 1991). Hosts for 8 made between the 16th and 37th days were
of these species are known (Table 1). Most used to estimate the mean number of egg
congeners are fairly polyphagous. Older masses on each plant species. Counts made
host record data must be treated with cau- between the 30th and 37th days were used
tion because the genus was substantially re- to estimate the mean population of nymphs
vised (Dietrich and Deitz 1991). The con- on each plant species.
cept of A. compressa in particular has been Experiment 2: Two plants each of 12
in confusion since Goding (1928) trans- species were selected in March 1991 for a
ferred it to the genus Guayaquila and syn- large cage experiment. The selection in-
onomized it with A. caliginosa Walker and cluded 6 species of Verbenaceae, with the
G. maxima Goding without examining the remainder being from closely related fami-
type species (Dietrich and Deitz 1991). lies. All the plants were examined prior to
Aconophora compressa has recently (Palm- their selection to ensure that they were in
er and Pullen 1995) been collected from L. good condition and were free of Aconopho-
camara, L. urticifolia, L. hirsuta and also ra. They were then watered before being
from Lippia myrocephala. It has also been placed inside a 2 X 3 X 7 m walk-in cage
collected from an unidentified shrub with situated on the grounds of the University of
densely pubescent bark (Dietrich and Deitz Morelos.
1991). The plants were placed inside the cage in
two rows; each row containing one plant of
Host SPECIFICITY
each of the 12 species. Within these rows
Materials and methods.(cid:8212)Three indepen- the plants were randomly assigned a posi-
dent experiments were conducted to ascer- tion which were equidistant so that no plant
tain the host range of this insect. The first was shadowed by its neighbours.
two were conducted at the University of Adult Aconophora compressa were col-
Morelos, Cuernavaca, Mexico prior to the lected from one large bush of L. urticifolia
insect being imported into Australia. The growing on the university grounds. The in-
third experiment was conducted within sects were collected either by hand or as-
quarantine facilities at the Alan Fletcher pirator and placed in plastic vials; fifteen to
Research Station, Brisbane. each vial. One vial was placed at the base
Experiment 1: The host specificity was of each plant in the experiment so that a
assessed using standard, gauze covered cag- total of 360 insects were placed in the cage.
es approximately | X 1 X 1 m in dimen- The insects then made their own way out
sion. Two replications were conducted; one of the vial and onto the plant.
in January 1991 and the other in February The test was assessed twice a week for
1991. In each replication one potted plant approximately seven weeks, a time suffi-
of eight species was selected and placed in cient for the insect to develop through a
the cage. Adult A. compressa from a labo- generation on the test plants. Each plant
ratory colony (20(cid:8212)40) were then introduced was examined and the number of colonies,
into the cage and scattered about so that adults, egg masses, and presence of nymphs
there was every chance that they would be noted. Six counts taken between the 23rd
exposed to all the plants in the cage. and 40th day after the experiment com-
The plants were examined approximately menced, when all stages of the insect were
twice a week for at least five weeks. Each present on the plants, were used to calculate
plant was examined and the number of the mean populations of adults, egg masses
adults, egg masses, and nymphs noted. and nymphs reported here.
Counts made between the 6th and 37th days Experiment 3: Adult choice tests were
were used to estimate the mean population carried out using gauze covered cages ap-
of adults on each plant species. Counts proximately 90 <X 90 X 90 cm. Three rep-
620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. A summary of the abundance of adults, egg masses and nymphs found on experimental plants in
three experiments conducted to determine the host range of A. compressa.
Mean No. of Mean No. of Mean No. of
Plant Species Expt. No. Adults Egg Masses Nymphs
Verbenaceae
Verbenoideae
Lantana camara L. 1 7.8 3.3 33.7
L. camara D 3.9 Del 24.9
L. camara 3 19.4 8.4
Lantana montevidensis (K. Spreng.) Briq. 1 0.3 0.0 0.0
L. montevidensis 2 0.0 0.0 0.0
L. montevidensis 3 0.03 0.0 0.0
Verbena carolina L. 2 0.0 0.0 0.0
V. bonariensis L. 3 0.0 0.0
Duranta repens L. 1 0.4 0.0 0.0
D. repens 2 10.6 2.4 4.5
D. repens var. lorentzii 3 0.0 0.0
Phyla nodiflora (L.) Greene 3 0.03 0.0
Stachytarpheta urticifolia (Salsb.) Sims 3 0.14 0.0
Viticoidae
Callicarpa pedunculata R. Br. 3 0.0 0.0
Clerodendrum sp. 2 0.0 0.0 0.0
C. cunninghamii Benth. 3 0.0 0.0
C. tomentosum (Venten.) 3 0.0 0.0
Petraea volubilis L. 2 0.0 0.0 0.0
Faradaya splendida F. Muell. 3 0.0 0.0
Gmelina leichhardtii (F. Muell.) Benth. 3 0.0 0.0
Premna lignum-vitae (Cunn. ex Shauer) Pieper 3 0.0 0.0
Vitex trifolia L. 3 0.0 0.0
Caryopteridoideae
Glossocarya hemiderma (F. Muell. ex Benth.) 3 0.0 0.0
Annonaceae
Annona X squamosa L. 3 0.0 0.0
Asteraceae
Baccharis halimifolia L. 3 0.33 0.0
Avicenniaceae
Avicennia marina (Forssk.) var australasica 3 0.0 0.0
Bignoniaceae
Jacaranda sp. 1 0.3 0.0 0.0
Jacaranda sp. 2 1.1 0.3 0.0
Jacaranda acutifolia Humb. & Bonpl. 3 3.1 0.3
Pandorea pandorana (Andr.) Steenis 3 0.0 0.0
Boraginaceae
Cordia dichromata G. Forster 3 0.0 0.0
Ehretia acuminata R. Br. 3 0.03 0.0
Chloanthaceae
Chloanthes parviflora Walp. 3 0.0
Spartothamnela juncea (Cunn. ex Walp.) Briq. 3 0.0 0.0
Convolvulaceae
Evolvulus alsinoides (L.) L. 3 0.0 0.0
Ipomoea batatas (L.) Lam. 3 0.0 0.0
VOLUME 98, NUMBER 4
Table 2. Continued.
Mean No. of Mean No. of Mean No. of
Plant Species Expt. No. Adults Egg Masses Nymphs
I. plebeia R. Br. 0.03 0.0
Polymeria calycina R. Br. 0.0 0.0
Dilleniaceae
Hibbertia scandens (Willd.) Dryand. 0.0 0.0
Euphorbiaceae
Croton insularis Baillon
Ericaceae
Azalea sp.
Lamiuaceae
Ajuga australis R. Br.
A. cf. reptans L. 0.0
Ocimum basilicum L. 0.0
Lavendula dentata L.
Plectranthus argentatus S. T. Blake
Prosthantera ovalifolia R. Br.
Mentha sp.
M. spicata L.
Salvia splendens F. Sellow ex Roem. & Scult.
S. coccinea Juss. ex J. Murr. 0.03
Scutellaria formosana N. E. Br.
Teucrium argutum R. Br.
Westringia fruticosa (Willd.) Druce WWWWWHeW e e WW0.23 WN
Mimosaceae
Acacia podalyriifolia G. Don 0.18
Acacia concurrens Pedley
Malvaceae
Hibiscus rosa-sinensis L. 0.0
Myoporaceae 0.0
Myoporum parvifolium R. Br. 0.04
Myrtaceae
Eucalyptus crebra F. Muell.
Eucalyptus curtisii Blakely & C. White
Eucalyptus sp.
Pedaliaceae
Sesamum indicum L.
Piperaceae
Piper novae-hollandiae Miq.
Scrophulariaceae
Duboisia myoporoides R. Br.
Mazus pumilio R. Br.
Solanaceae
Cestrum nocturnum L.
Lycopersicon lycopersicum L.
Solanum nemophilum F. Muell.
S. pseudocapsicum L.
S. tuberosum L. WWe WN
622 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A
i
Fig. 1. Predicted distribution of A. compressa in Australia based on the climate matching model CLIMEX.
lications were conducted concurrently. In ments. The egg masses on D. repens pro-
each replication, 9 plant species, always in- duced nymphs that then developed normal-
cluding L. camara, were arranged in a cage. ly on that host. Nymphs were never found
Thirty adult A. compressa were then scat- associated with the egg masses on Jacaran-
tered into the cage. Counts of the adults on da indicating that this plant is an unsuitable
each plant were made between the 6th and host.
21st day after this introduction. Egg masses Altogether this insect was tested against
were counted on the 21st day. 62 plant species of which 17 were of the
Results.(cid:8212)The results of all three exper- Verbenaceae and 13 and 5 species were of
iments are given in Table 2. Lantana ca- the closely related Lamiaceae and Solana-
mara was the only plant species on which ceae, respectively. The insect was in fact
A. compressa consistently oviposited. Ovi- thoroughly tested along the centrifugal de-
position also occurred on both plants of Du- sign advocated by Wapshere (1975). All
ranta repens L. in one of three experiments evidence suggests that the host range of
and on Jacaranda spp. in all three experi- this insect is limited to Lantana and close-
VOLUME 98, NUMBER 4 623
ly related genera such as Duranta and Lip- leased against Mimosa invisa L., and the
pia. aphid Aphis chloris Koch, released against
Hypericum perforatum L., are other exam-
RELEASE IN AUSTRALIA ples of Sternorrhyncha listed by Julien
Permission to release this insect in Aus- (1992). The only instance, to our knowl-
tralia was sought even though it could breed edge, of the Auchenorrhyncha being so
on Mexican varieties of D. repens (subfam- used is that of Stobaera concinna (Stal) re-
ily Verbenoidae). From an Australian per- leased for Ambrosia artemisiifolia (L.) and
spective D. repens is an introduced orna- Parthenium hysterophorus L. (Julian 1992).
mental suspected of being poisonous to The reasons why the Homoptera other than
children and stock (Everist 1974). The Ver- Coccoidea have been so eschewed include
benoidae is represented in Australia by one the perception that most are polyphagous,
native species, Verbena macrostachya, that many are vectors of plant diseases, that
which is found only in inland Queensland they may be a greater than normal risk in
far from the habitat of L. camara. quarantine facilities, that they are difficult
Approval to release A. compressa was to ship, and that their effects on the plant
obtained in early 1995 and a mass rearing are slight or difficult to define.
program outside the quarantine facilities Apart from A. compressa being the first
was initiated. The climate matching model use of a membracid for biological control,
CLIMEX (Sutherst and Maywald 1985, its progress will be of interest because it
Skarratt et al. 1995) was utilized in the se- feeds on stems externally. Endophagous
lection of release sites. The model indicated species such as stem borers and gall for-
(Fig. 1) that A. compressa should adapt to mers are more commonly used but there are
conditions along the eastern seaboard of few examples of ectophagous stem utiliz-
Australia where L. camara is a problem. CIs:
Conditions in southern Queensland and
ACKNOWLEDGMENTS
northern New South Wales should be par-
ticularly favorable. We thank C. H. Dietrich who provided
First releases were made near Brisbane identifications and other valuable advice to
in May 1995 and at Charters Towers in Oc- us. The experimental work in Mexico was
tober 1995. carried out by K. R. Pullen. We also wish
to thank the University of Morelos for mak-
DISCUSSION ing facilities available of this work. CLI-
We anticipate that A. compressa will be MEX simulations were undertaken by M.
a useful biocontrol agent if it establishes A. Hannan-Jones. The project was support-
successfully. In Mexico it is quite damaging ed financially by the Queensland Rural
especially when a plant is attacked by mul- Lands Protection Board.
tiple colonies. One reason for optimism is
LITERATURE CITED
that its action will be largely independent
of leaf status. Lantana has the ability to Cilliers, C. J. and S. Neser. 1991. Biological control
drop its leaves when stressed and uses this of Lantana camara (Verbenaceae) in South Afri-
ca. Agriculture, Ecosystems and Environment 37:
mechanism to survive periods of drought,
57-75.
cold, insect attack and possibly herbicide
Dietrich, C. H. and L. L. Deitz. 1991. Revision of the
application. This reduces the effect of in- neotropical treehopper tribe Aconophorini (Ho-
sects that attack foliage. moptera: Membracidae). North Carolina Agricul-
With the notable exception of the Coc- tural Research Service. Technical Bulletin 293.
134 pp.
coidea, the Homoptera have not been great-
Everist, S. L. 1974. Poisonous Plants of Australia.
ly used in biological control of weeds (Ju-
Angus and Robertson, Sydney, Australia. 966 pp.
lien 1992). The psyllid Heteropsylla sp., re- Goding, EFW . 1928. Membracidae of South America
624 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and Antilles, III. Subfamily Membracinae. Journal agous arthropods associated with Lantana ca-
of the New York Entomological Society 36: 201(cid:8212) mara, L. hirsuta, L. urticifolia and L. urticoides
234. (Verbenaceae) in North America. Biological Con-
Harley, K. L. S. 1974. Biological control of Lantana trol 5: 54-72.
in Australia. Proceedings of the III International Perkins, R. C. L. and O. H. Swezey. 1924. The intro-
Symposium on Biological Control of Weeds, duction into Hawaii of insects that attack Lantana,
Montpellier, France. (ed.), Commonwealth Insti- Bulletin of the Experimental Station of the Ha-
tute of Biological Control. Farnham Royal, UK. walian Sugar Planter(cid:8217)s Association 16: 1-83.
23-29 pp. Skarratt, D. B., R. W. Sutherst, and G. EF Maywald.
Holm, L. G., D. L. Plucknett, J. V. Pancho, and J. P. 1995. CLIMEX for Windows Version 1.0. CSIRO
Herberger. 1977. The World(cid:8217)s Worst Weeds. Uni- and CRC for Tropical Pest Management. Bris-
versity Press of Hawaii, Honolulu, USA. 609 pp. bane.
Julien, M. H. 1992. Biological Control of Weeds. A Sutherst, R. W. and G. E. Maywald. 1985. A com-
World Catalogue of Agents and their Target puterized system for matching climates in ecolo-
Weeds. Third Edition. CAB International, Wal- gy. Agriculture, Ecosystems and Environment 13:
lingford, UK. 186 pp. 281-299.
Kleinschmidt, H. E. and R. W. Johnson. 1977. Weeds Taylor, E. E. 1989. A history of biological control of
of Queensland. Queensland Department of Pri- Lantana camara in New South Wales. Plant Pro-
mary Industries, Brisbane. 469 pp. tection Quarterly 4: 61-65.
Muniappan, R., M. Marutani, J. McConnell, P. Singh, Wapshere, A. J. 1975. A protocol for programmes for
N. Wilson, and T. S. Lali. 1992. Bibliography of biological control of weeds. PANS 21: 295-303.
Lantana camara. Agricultural Experiment Station, Winder, J. A. and K. L. S. Harley. 1983. The phy-
University of Guam. Technical Report No. 94. 31 tophagous insects on Lantana in Brazil and their
Pp- potential for biological control in Australia. Trop-
Palmer, W. A. and K. R. Pullen. 1995. The phytoph- ical Pest Management 29: 346-362.