Reassessment of the Biology and Host Range of Acanthoscelides

macrophthalmus (Chrysomelidae: Bruchinae), a Seed-Feeding

Beetle Released for the Biological Control of Leucaena
leucocephala in South Africa
Author(s): Z. Shoba & T. Olckers
Source: African Entomology, 18(2):1-9.
Published By: Entomological Society of Southern Africa
https://doi.org/10.4001/003.018.0222
URL: http://www.bioone.org/doi/full/10.4001/003.018.0222

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Reassessment of the biology and host range of Acanthoscelides macrophthalmus
(Chrysomelidae: Bruchinae), a seed-feeding beetle released for the biological
control of Leucaena leucocephala in South Africa

Z. Shoba & T. Olckers*

School of Biological and Conservation Sciences, University of KwaZulu-Natal,
Private Bag X01, Scottsville, 3209 South Africa

Introduced for agroforestry, Leucaena leucocephala (Lam.) de Wit (Fabaceae) has become
naturalized in several countries worldwide and constitutes an ‘emerging weed’ in South
Africa. The seed beetle Acanthoscelides macrophthalmus (Schaeffer) (Chrysomelidae:
Bruchinae) was inadvertently introduced to many of these countries via contaminated
seeds, but was deliberately introduced into South Africa for biological control. Following an
isolated report of non-target feeding in the field in southeastern Asia, this study was aimed at
re-evaluating aspects of the beetle’s biology, in particular the likelihood of non-target attacks
on native mimosoid Fabaceae that are closely related to L. leucocephala. A preliminary field
assessment of the beetle’s susceptibility to native parasitoids was also undertaken. The
beetle’s life cycle includes several attributes (e.g. rapid reproductive maturation, high rates of
oviposition) that favour high population densities in the field. Although adults oviposited
on the seeds of all test plant species during no-choice tests, oviposition was significantly
higher on L. leucocephala. While several hatching larvae survived to adulthood on
L. leucocephala, single adult individuals were reared from the seeds of each of three non-target
species. Also, during multichoice tests, the beetles displayed strong oviposition preferences
for L. leucocephala. Two hymenopteran parasitoid species were reared from seeds containing
larvae which were exposed in the field, but overall levels of parasitism were low. These results
confirm that the host range of A. macrophthalmus is largely restricted to the genus Leucaena,
that native mimosoid Fabaceae are unlikely to be at risk and that the decision taken in 1999 to
release the beetle was justified.
Key words: biological weed control, host specificity, leucaena, non-target effects, parasitism,
seed beetles.

INTRODUCTION
Native to Mexico and Central America, Leucaena
leucocephala (Lam.) de Wit (Fabaceae) is a shrub or
small tree that has become naturalized in several
tropical, subtropical and warm temperate countries
worldwide, as a result of introductions for agro-
forestry purposes (ISSG 2006). Commonly known
as leucaena, the plant is considered to be one of the
“World’s 100 Worst Invaders” (ISSG 2006), but is
still an ‘emerging weed’ in South Africa (Olckers
2004) where it occurs in several provinces, notably
the KwaZulu-Natal coastal region (Henderson
2001). Self compatibility, allelopathy and prolific
seed production (Raghu et al. 2005; ISSG 2006;
Tuda 2007) promote invasions of forest margins,
river banks, roadsides and wasteland in South
Africa (Henderson 2001). However, the plant
constitutes a typical ‘conflict species’ because of
*To whom correspondence should be addressed.
E-mail: olckerst@ukzn.ac.za
several useful attributes that include nitrogen
fixation, pulping for paper, sand-binding, refores-
tation and use as a source of fuel wood, construc-
tion material and livestock fodder (Henderson
2001; ISSG 2006; Tuda 2007; López et al. 2008).
Biological control of L. leucocephala was initiated
in South Africa in 1999, with the release of the seed
beetle Acanthoscelides macrophthalmus (Schaeffer)
(Chrysomelidae: Bruchinae) (Olckers 2004). Seed
beetles (formerly the family Bruchidae) have been
released against other agroforestry species, notably
Prosopis species, in South Africa because of their
potential to reduce seed dispersal and hence,
invasion potential, without negatively affecting
utilization (Impson et al. 1999). Because the pro-
gramme against L. leucocephala was not officially
funded, the beetle was opportunistically intro-
duced from Mexico during surveys for agents of
other plants that are invasive in South Africa
African Entomology 18(2) Supplement No. 1: 1–9 (2010)
2 African Entomology Vol. 18, No. 2, Supplement No. 1, 2010
(Olckers 2004). Prior to release, the beetle’s host
range was evaluated via exposure of adults to the
seeds of several non-target plants in the sub-
families Mimosoideae, Caesalpinioideae and
Papilionoideae of the family Fabaceae (Neser &
Klein 1998), although many species were not
tested due to the large size of the family.
Females deposit eggs on the surfaces of ripe
pods and ripe loose seeds of L. leucocephala. In the
field, ripe leucaena pods split open and release
their seeds onto the ground; however, it is unclear
whether the beetles typically select dehisced seeds
on the soil surface, or whether there is a preference
for ovipositing onto the canopy held pods which
still contain seeds. Whatever the case, oviposition
onto loose seeds takes place easily under laboratory
conditions. Although A. macrophthalmus oviposited
on the seeds of several non-target species, survival
to adulthood was confined to Leucaena species
(Neser & Klein 1998) and the beetle was cleared
for release in 1999. However, the project has
remained a low priority and despite releases of
beetles at a few sites, both the extent of the beetle’s
establishment and its impact on the weed’s seed
dynamics remain unknown (Zimmerman et al.
2004).
Although South Africa is currently the only
country where A. macrophthalmus was deliberately
introduced, the beetle was accidentally introduced
into several regions worldwide, including West
Africa (Delobel & Johnson 1998), Australia (Raghu
et al. 2005) and southeastern Asia (Tuda 2007),
presumably via contaminated seeds. Although the
literature indicates that the beetle has remained
specific to Leucaena species in its native and intro-
duced range (see references in Tuda (2007)), a
recent field survey in the Far East and South Asian
tropics and subtropics revealed one case of host
range expansion on mimosoid Fabaceae (Tuda
et al. 2009). This raised concerns of similar host
range expansions on native South African mimosoid
Fabaceae, notably species of Acacia, Albizia and
Dichrostachys, that are closely related to Leucaena
species (Sulaiman et al. 2003) and that are known
to be host plants for seed beetles (Southgate 1979;
Kingsolver 2004). Although representatives of
these genera (but individual species not listed)
were included in the original host-range evalua-
tion of A. macrophthalmus (Neser & Klein 1998),
these trials involved multichoice tests and did not
include no-choice tests which provide a more
conservative estimate of insect host range.
The primary aim of this study was thus to
reassess the host range of A. macrophthalmus by
exposing the beetles to seeds of selected native
mimosoid Fabaceae under both no-choice and
multichoice situations. Due to a lack of published
information on the biology of A. macrophthalmus,
an initial study was carried out to determine the
pre-oviposition period, adult longevity and dura-
tion of the immature stages, all aspects that could
influence the manner in which the host-range
tests should be conducted and interpreted. Finally,
because parasitism of A. macrophthalmus has been
reported in countries where it was introduced
(Delobel & Johnson 1998; Raghu et al. 2005), a
preliminary field trial was conducted to determine
any impacts by native parasitoids.
MATERIAL AND METHODS
Biology of A. macrophthalmus
Observations of the beetle’s biology were made
in an insectary at the University of KwaZulu-Natal
(Pietermaritzburg), where the beetles were cultured
at 28 °C in 5-litre plastic jars containing loose seeds
of L. leucocephala, where water only was provided
with an atomiser. The laboratory culture was
initiated from infested seeds that were collected at
field sites around Durban and Pietermaritzburg in
2008. To determine female pre-oviposition period,
10 unsexed, newly emerged beetles (≤24 hours
old) were confined in 500 ml aerated plastic
containers with 20 ripe seeds of leucaena. The
seeds were monitored daily for newly laid eggs
and the trial was terminated when the first egg
was recorded on a seed. Pre-oviposition period
was calculated from the time when the beetles
were confined in the container to the time when
the first egg was observed. The trial was replicated
20 times.
To determine the duration of larval develop-
ment to adulthood, 125 seeds with single newly
laid eggs (≤24 hours old) were each placed in
isolated compartments in microtitre trays for
individual observation and the dates were recorded.
The infested seeds were monitored daily for adult
emergence. Because the immature stages complete
their development exclusively within the seed,
the time from oviposition to adult emergence was
recorded. Newly emerged beetles were removed
from the tray compartments and added to the
laboratory cultures.
To determine adult longevity, five unsexed,
 Shoba & Olckers: Biology and host range of Acanthoscelides macrophthalmus (Chrysomelidae) 3

Table 1. Test plant species in the subfamily Mimosoideae (Fabaceae) that were used to reassess the host range of
Acanthoscelides macrophthalmus.

Tribe Test plant species (common name) Overlap with

leucaena

Mimoseae Leucaena leucocephala (Lam.) de Wit (leucaena)* –

Ingeae Albizia adianthifolia (Schumach.) W.F. Wight (flat-crown)* Extensive
Acacieae Acacia erioloba E. Mey. (camel thorn) Very little
Acacia hebeclada DC subsp. hebeclada (candle thorn)* Very little
Acacia karroo Hayne (sweet thorn)* Extensive
Acacia nilotica (L.) Willd. Ex Del. subsp. kraussiana (Benth.) Brenan (scented Extensive
thorn)
Acacia robusta Burch. subsp. robusta (ankle thorn)* Extensive
Acacia sieberiana DC var. woodii (Burt Davy) Keay & Brenan (paperbark thorn) Extensive
Acacia swazica Burt Davy (Swazi thorn)* Limited
Acacia tortilis (Forsk.) Hayne subsp. heteracantha (Burch.) Brenan (umbrella Extensive
thorn)
Adenantherae Dichrostachys cinerea (L.) Wight & Arn subsp. africana Brenan & Brummet Extensive
(sickle bush)

*Pods split on the tree to release seeds.

newly emerged beetles, which had emerged on
the same day, were confined in similar aerated
plastic containers with 20 seeds of leucaena and
were provided with water only. The beetles were
monitored daily and the time from emergence to
death was recorded for each beetle. This arrange-
ment was replicated 16 times, providing a sample
size of 80 beetles.
Host range of A. macrophthalmus
All trials were conducted in the insectary under
the same conditions described previously. Ovi-
position behaviour was first tested by exposing the
beetles to loose seeds of 10 native species from
three tribes in the subfamily Mimosoideae, as well
as L. leucocephala, in both no-choice and multi-
choice situations (Table 1). The test species, from
the genera Acacia, Albizia and Dichrostachys, were
selected because of their close relatedness to
L. leucocephala (Sulaiman et al. 2003) and because
several occur naturally in areas that are typically
invaded by leucaena.
In the no-choice tests, 15 unsexed newly
emerged adults were presented with 20 seeds of
a single test plant species in a 500 ml plastic
container. All tests were replicated four times,
using different beetles and different seeds on
each occasion, and the different replicates were
staggered over several weeks. Cotton wool placed
in the containers was kept moist for the duration
of the trials to provide the beetles with water. The
trials were terminated after 14 days, surviving
adults were removed and the number of eggs on
each seed was recorded. The seeds which had been
oviposited on were then kept in the containers for
several weeks to allow development of the imma-
ture stages and were monitored daily to record
adult emergence. Emerging beetles were removed
from the containers to avoid reinfestation.
Following oviposition on seeds of several
non-target species (in particular Ac. sieberiana)
during the above trials, an additional set of
no-choice tests was conducted using seeds of
Ac. sieberiana and separately, seeds of leucaena to
quantify egg-laying patterns and determine
whether the beetles readily oviposit on non-target
species or whether they delay oviposition and
eventually ‘dump’ their eggs in the absence of the
natural host. These trials were conducted as before
but the seeds were glued to the base of the con-
tainer to limit oviposition to the upper surfaces
and facilitate egg counts which were recorded
daily. Eggs laid on the sides of the container were
also recorded in these trials, which were replicated
four times and terminated after 15 days.
Multichoice tests were conducted to compare
the egg-laying decisions of females when presented
with a choice. Twenty seeds from each of the
11 test plant species were grouped according to
species in Petri dishes, which were arranged
4 African Entomology Vol. 18, No. 2, Supplement No. 1, 2010
randomly inside a 60 × 30 × 30 cm glass tank that
was covered with gauze. Fifty newly emerged
beetles were confined in the glass tank with the
different arrangements of seeds, and water was
provided as before. This design allowed the
beetles to orientate freely and oviposit on seeds of
their choice. These trials were replicated five times
and were also terminated after 14 days. Eggs laid
on the seeds were counted and each Petri dish
was covered with a lid to prevent escape of the
emerging beetles. The seeds were similarly moni-
tored to record adult emergence and emerging
beetles were removed from the Petri dishes to
avoid reinfestation of the seeds.
Parasitism of A. macrophthalmus
Parasitoid traps were prepared in which 20
larvae-infested seeds were glued onto plastic
sheets (30 × 12 cm) to expose the immature stages
to native parasitoids in the field. Each seed con-
tained 1–3 eggs and the total number of eggs
(larvae) per trap ranged from 34–41. Three study
sites containing populations of Ac. sieberiana and
leucaena were selected around Pietermaritzburg
and the traps were then hung from selected trees,
with one trap per tree of Ac. sieberiana and
leucaena. Five pod-bearing trees of each species
were randomly selected at the first site (University
arboretum), while three trees of each species were
selected at the other two sites (University research
farm) due to fewer numbers of eggs available at
the time. In total, 400 eggs (larvae) were exposed at
the first site, compared with 230 and 237 at the
other two sites, providing an indication of the
number of beetles expected to emerge at each site.
The traps bearing the infested seeds were exposed
in the field for five weeks, during September and
October 2008, and the seeds were then removed
and confined in Petri dishes in the laboratory to
record beetle and parasitoid emergence. Emerging
individuals were removed on a daily basis to avoid
reinfestation and all parasitoids were preserved
for identification.
Statistical analysis
In the no-choice and multichoice tests, means of
the numbers of eggs deposited and numbers of
emerging adults were compared among the test
plants using Kruskal-Wallis tests and where these
were significant (P < 0.05), pairwise comparisons
between L. leucocephala and each non-target
species were carried out using Mann-Whitney
U tests. Regression analyses, using oviposition
data on L. leucocephala from the laboratory and
field trials (except where affected by parasitism),
were used to examine the relationship between
the survival of the immature stages to adulthood
and egg-laying intensity.
RESULTS
Biology of A. macrophthalmus
The longevity of newly emerged adults varied
from 2–20 days, with a mean (±S.D.) of 11.9 ±
3.5 days, when maintained on water alone. Conse-
quently, beetles were exposed to the test seeds for
14 days during the subsequent host-range tests.
The pre-oviposition period lasted 2–4 days and
females laid eggs some 2.7 ± 0.8 days (mean ±
S.D.) after emergence. Observations on the breed-
ing colony revealed that eggs were normally laid
singly on the seeds, although multiple oviposition
(‘superparasitism’) occurred when oviposition
sites were limited. However, only one or two
adults normally emerged from ‘superparasitized’
seeds, although up to three were recorded.
Hatching larvae burrowed into the seed leaving
tiny entry holes visible and consumed much of the
seed contents during larval development. Prior to
pupation, the final instar excavated a perfectly
circular ‘emergence hatch’ on the seed surface
which was forced open by the emerging adult. Of
the 125 eggs (infested seeds) that were monitored,
117 (93.6 %) produced adults and the duration of
the immature stages (egg to adult emergence)
lasted 27–59 days, with a mean of 35.3 ± 5.7 days at
28 °C.
Host range of A. macrophthalmus
Oviposition was recorded on the seeds of all test
plants under no-choice conditions (Table 2). With
the exception of the trials involving leucaena
seeds, the beetles also oviposited extensively on
the sides of the plastic containers and this was
particularly evident in trials where few eggs
were laid on the actual seeds (e.g. Ac. karroo and
Ac. tortilis). Of the non-target seeds, those of
Ac. sieberiana appeared to be the most favoured for
oviposition (Table 2). However, the beetles laid
significantly more eggs on the seeds of leucaena
(P < 0.05) than on those of any of the non-target
species (Table 2). Consequently, significantly more
adults emerged from leuacena seeds (P < 0.05).
While a total of 87 adults emerged from eggs
 Shoba & Olckers: Biology and host range of Acanthoscelides macrophthalmus (Chrysomelidae) 5

Table 2. Mean (±S.D.) oviposition and adult emergence by Acanthoscelides macrophthalmus on seeds of different
test plants during no-choice tests in 500 ml plastic containers.

Test plant species Number of eggs Eggs on container Number of emerging % Adult emergence
on seeds1 adults1

L. leucocephala 194.8 ± 30.9 None 21.8 ± 6.9 11.7 ± 4.8

Ac. erioloba 56.5 ± 19.3 Some 0 0
Ac. hebeclada 39.5 ± 23.2 Many 0 0
Ac. karroo 1.8 ± 2.1 Many 0 0
Ac. nilotica 54.0 ± 102.1 Many 0.3 ± 0.5 0.1 ± 0.2
Ac. robusta 53.5 ± 17.8 Some 0 0
Ac. sieberiana 100.8 ± 29.4 Many 0 0
Ac. swazica 41.0 ± 11.5 None 0 0
Ac. tortilis 8.8 ± 8.8 Many 0.3 ± 0.5 1.2 ± 2.4
Al. adianthifolia 34.8 ± 24.3 Many 0 0
D. cinerea 39.3 ± 21.6 Many 0.3 ± 0.5 0.8 ± 1.6
1
Means compared between test plants by Mann-Whitney U tests; all were significantly different from Leucaena leucocephala (P < 0.05).

deposited on seeds of leucaena, only one adult
emerged from eggs laid on the seeds of each of Ac.
nilotica, Ac. tortilis and D. cinerea. None of the eggs
laid on any of the seeds of the remaining seven
non-target species produced adults (Table 2), even
though the eggs had hatched and penetration of
the seeds had occurred. Of the high numbers of
eggs laid on leucaena (around 10 per seed) only
12 % produced adults, probably as a result of
excessive larval competition.
The evaluation of egg-laying patterns revealed
that beetles confined with seeds of Ac. sieberiana
delayed their oviposition by two days, relative to
those confined with seeds of L. leucocephala (Fig. 1).
Cumulative oviposition was lower in the Ac.
sieberiana trial by days 4–12 but was similar to that
in the leucaena trial by days 13–15, so that overall
levels of oviposition were similar in both trials
(Fig. 1). However, in contrast to the oviposition
trials using leucaena seeds, where around 120 eggs
were laid onto the seeds (Fig. 1), beetles confined
with Ac. sieberiana seeds oviposited almost exclu-
sively on the sides of the container and only two
eggs were recorded on the actual seeds (on
day 13). This contrasts the results of the earlier
no-choice tests where Ac. sieberiana was the most
‘preferred’ of the non-target species and where
eggs were typically laid on the seeds (Table 2).

Fig. 1. Accumulated mean number of eggs laid by Acanthoscelides macrophthalmus when exposed to seeds of
Leucaena leucocephala and Acacia sieberiana for 15 days during no-choice tests. Means for Ac. sieberiana include
eggs laid on the seeds and on the sides of the containers.
6 African Entomology Vol. 18, No. 2, Supplement No. 1, 2010

Table 3. Mean (± S.D.) oviposition and adult emergence by Acanthoscelides macrophthalmus on seeds of different
test plants during multichoice tests in 54 litre glass tanks.

Test plant species Number of eggs on seeds1 Number of emerging adults % Adult emergence

L. leucocephala 243.0 ± 72.2 23.0 ± 3.9 10.3 ± 3.9

Ac. erioloba 2.8 ± 3.7 0 0
Ac. hebeclada 1.0 ± 1.2 0 0
Ac. karroo 0.2 ± 0.4 0 0
Ac. nilotica 0.2 ± 0.5 0 0
Ac. robusta 5.0 ± 4.8 0 0
Ac. sieberiana 0.6 ± 0.9 0 0
Ac. swazica 0.2 ± 0.5 0 0
Ac. tortilis 0.2 ± 0.5 0 0
Al. adianthifolia 0.4 ± 0.6 0 0
D. cinerea 3.8 ± 4.9 0 0
1
Means compared between test plants by Mann-Whitney U tests; all were significantly different from Leucaena leucocephala (P < 0.01).

These results indicate that ‘egg dumping’ is occur-
ring when the beetles are deprived of seeds of
their natural host.
Similarly in the multichoice tests, substantially
higher numbers of eggs (around 12 per seed) were
laid on leucaena seeds (P < 0.01) than on those of
the 10 non-target plants (Table 3). Very low numbers
were laid on Ac. erioloba, Ac. robusta and D. cinerea,
which were the most ‘preferred’ of the non-target
plants during this trial (Table 3). Adults emerged
only from eggs laid on L. leucocephala seeds, with
10 % of the eggs producing adults.
Parasitism of A. macrophthalmus
The field exposure trials confirmed that the
endophagous immature stages of A. macroph-
thalmus are susceptible to native parasitoids. Two
wasp species (Chalcidoidea) that were both provi-
sionally identified as Pteromalidae (Prinsloo 1980)
were recovered from the traps, with 23 specimens
of a single species and one specimen of a different
species. The emergence holes of the adult wasps
were easily differentiated from those of the adult
beetles by virtue of their smaller size and jagged
edges. Parasitism was recorded at only one of the
three sites (Table 4), where wasps were recovered
from six of the 10 traps exposed in stands of both L.
leucocephala (all five traps) and Ac. sieberiana (one
trap). More wasps were recovered from the traps
exposed on the L. leucocephala trees (19) than on the
Ac. sieberiana trees (5). It is unclear why parasitism
was recorded only at this site and not at the other
two, since the native Acacia species (presumably
the source of the parasitoids) present at the three
sites were not obviously different. At this site,
fewer adults of A. macrophthalmus (9 %) emerged
from the seeds exposed in stands of L. leucocephala,
than on those exposed in stands of Ac. sieberiana
(45 % adult emergence). Adult emergence was
somewhat higher at the remaining two study sites
(46–62 %) where no parasitism was recorded
(Table 4).

Table 4. Numbers of beetles and parasitoids that emerged from larvae-infested seeds of Leucaena leucocephala
exposed in stands of L. leucocephala and Acacia sieberiana at three field sites.

Study site Tree stand No. exposed No. beetles No. parasitoids
larvae1 emerged2 emerged2

University arboretum L. leucocephala 199 18 (9.0 %) 19 (9.5 %)

29°37’S 30°24’E Ac. sieberiana 201 90 (44.8 %) 5 (2.5 %)
University farm (1) L. leucocephala 118 72 (61.0 %) 0
29°40’S 30°24’E Ac. sieberiana 112 69 (61.6 %) 0
University farm (2) L. leucocephala 123 60 (48.8 %) 0
29°40’S 30°24’E Ac. sieberiana 114 52 (45.6 %) 0
1
Assuming 100 % hatching success of eggs deposited on the seeds.
2
Percentage adult emergence and parasitism expressed in relation to the number of exposed larvae.
 Shoba & Olckers: Biology and host range of Acanthoscelides macrophthalmus (Chrysomelidae)
DISCUSSION
The results of the life-history features investi-
gated during this study were predominantly to
help with the design and interpretation of
the host-range studies. Under laboratory condi-
tions, when sustained on water alone, adults of
A. macrophthalmus reached reproductive maturity
soon after emergence from infested seeds (3 days)
but were short-lived (12 days) in relation to the
endophagous immature stages (36 days). How-
ever, adult Bruchinae are known to feed on pollen
and nectar in the field (Southgate 1979) and
longevity may thus be substantially higher under
such circumstances. In addition, all observations
and trials were carried out on loose seeds which
had been removed from pods. Bruchinae are
renowned for their good dispersal abilities in
relation to finding suitable oviposition sites, but it
is unclear whether oviposition by A. macrophthalmus
in the field is restricted to seeds that are retained in
their pods on the trees and thus whether dehisced
seeds escape predation. A study by Raghu et al.
(2005) in Australia indicated that seed predation
by A. macrophthalmus was restricted by pod reten-
tion times, suggesting that, despite their accep-
tance of loose seeds in the laboratory, the beetles
fail to locate dehisced seeds in the field. If this is
true, then their impact on leucaena populations in
the field may well be limited.
The reported host specificity of A. macroph-
thalmus (Neser & Klein 1998) was confirmed in
that, despite oviposition on non-target seeds and
the recovery of single adults from the seeds of
three non-target species in the no-choice trials,
development to adulthood was largely confined
to L. leucocephala. Deprivation of leucaena seeds
resulted in females ‘dumping’ their eggs on both
non-target seeds and the sides of the containers.
In contrast, clear oviposition preferences were
demonstrated in the multichoice tests when
L. leucocephala seeds were available. Seeds of
mimosoid legumes such as Acacia species have
unique defensive compounds, notably non-protein
amino acids, that limit the emergence of adult
Bruchinae by disrupting larval development
(Ward & Or 2004), which could explain the unsuit-
ability of the non-target seeds for species like
A. macrophthalmus which have previously not been
exposed to them. Our findings are thus consistent
with those of Neser & Klein (1998) and other
reports stating that the host ranges of Bruchinae
7
are largely limited to restricted plant taxa, mostly
at the level of tribes and subtribes (see references
in Tuda (2007)).
During the host-range trials, adult emergence
from seeds of L. leucocephala was low (10–12 %) in
relation to the high numbers of eggs laid (10–12
eggs per seed). In contrast, adult emergence was
considerably higher from seeds used in the para-
sitism study (46–62 % in unparasitized situations),
where egg densities were much lower (1–3 eggs
per seed). When these data were combined, there
was a strong negative relationship (y = 68.78e–0.16x,
R2 = 0.90, P < 0.001) between the percentage of
eggs that produced adults and the mean number
of eggs per seed (Fig. 2). Female Bruchinae are
known to display oviposition preferences for
seeds of higher nutritional quality (e.g. larger size)
to ensure larval survival and to resort to ‘super-
parasitism’ when resources are limited (Siemens
et al. 1991; Cope & Fox 2003). However, leucaena
seeds generally seem to have enough resources for
one or two larvae only, leading to increased larval
competition and high mortality inside ‘super-
parasitized’ seeds. Although ‘superparasitism’
seems unlikely to occur in the field due to the pro-
lific seed production of leucaena populations, it
may affect beetle population densities during
times of seed shortages, particularly if the beetles
avoid dehisced seeds (Raghu et al. 2005).
Parasitism of the immature stages of A. macro-
phthalmus by native hymenopteran parasitoids was
confirmed, with several individuals of a species of
Pteromalidae collected. Native Pteromalidae have
been reported to parasitize A. macrophthalmus in
the field in Senegal, West Africa where the beetle
was inadvertently introduced (Delobel & Johnson
1998). However, it is likely that a more intensive
investigation of parasitism will reveal more
parasitoid species, as several wasp families are
known to parasitize the immature stages of
Bruchinae (Hoffmann et al. 1993; Kingsolver 2004).
Although the rates of parasitism were low in this
preliminary study, more extensive sampling is
required to determine the extent to which field
populations of A. macrophthalmus are affected by
parasitism.
Although the biology of A. macrophthalmus dis-
plays several features typical of a good biological
control agent (e.g. high oviposition rates, a short
life cycle with the potential for several generations
per annum, good dispersal abilities), aspects such
as the selection of oviposition sites under natural
8 African Entomology Vol. 18, No. 2, Supplement No. 1, 2010
Fig. 2. Relationship between the survival to adulthood of the immature stages of Acanthoscelides macrophthalmus
and the intensity of oviposition on seeds of Leucaena leucocephala.
conditions need clarification. Given the single
instance of non-target feeding in the field in
southern Asia (Tuda et al. 2009), the justification
for extending the original host-specificity studies
was well founded, even though these data serve to
confirm the results of the original tests. However,
development to adulthood (albeit single individuals
only) recorded in this study on the seeds of three
non-target species, suggests that field monitoring
of seeds of native Acacia species, in areas invaded
by leucaena, should be undertaken in South Africa
to confirm that no host range expansions have
occurred.
Populations of A. macrophthalmus have recently
been confirmed at several sites in the KwaZulu-
Natal coastal region and the extent of seed damage
thus needs to be quantified. Similarly, the extent of
parasitism needs to be quantified at a range of field
sites, which are in close proximity to different
native Acacia species hosting different complexes
of native Bruchinae and their parasitoids. This
study has thus raised several possibilities for
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Accepted 22 April 2010