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Biological Control of Leucaena leucocephala (Lam.) de Wit

(Fabaceae) in South Africa: A Tale of Opportunism, Seed Feeders
and Unanswered Questions

Article  in  African Entomology · August 2011

DOI: 10.4001/003.019.0219

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Biological control of Leucaena leucocephala (Lam.) de Wit (Fabaceae) in
South Africa: a tale of opportunism, seed feeders and unanswered questions

T. Olckers
School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville,
3209 South Africa. E-mail: olckerst@ukzn.ac.za

Leucaena leucocephala (Lam.) de Wit (Fabaceae) is a typical ‘conflict species’, possessing both
useful and damaging attributes, that has become naturalized in several countries world-
wide, following deliberate introductions for agroforestry. Considered one of the ‘World’s
100 Worst Invaders’, the plant in South Africa is an incipient or ‘emerging weed’, with
infestations mostly occurring in the eastern subtropical regions of the country. Deliberate
biological control efforts have been confined to South Africa, although these have been
opportunistic, unfunded and, given the plant’s propensity for prolific seed production, have
focused exclusively on the seed beetle, Acanthoscelides macrophthalmus (Schaeffer)
(Chrysomelidae: Bruchinae). Following confirmation of host specificity, and after several
low-intensity releases, the beetle has become widely established in KwaZulu-Natal, where
releases were mostly made, but also in Gauteng, Mpumalanga and the North West
provinces. A sap-sucking psyllid, Heteropsylla cubana Crawford (Psyllidae), was inadver-
tently introduced into South Africa, but has not had any impact on weed infestations. The
biological control programme against L. leucocephala in South Africa is reviewed, including:
(i) aspects of the target plant that are pertinent to its invasiveness and management;
(ii) details pertaining to the two natural enemies that have featured so far; (iii) progress that
has been achieved; and (iv) the direction of future research efforts.
Key words: Acanthoscelides macrophthalmus, emerging weeds, Heteropsylla cubana, leucaena,
seed beetles, weed biological control.

INTRODUCTION
Leucaena leucocephala (Lam.) de Wit (Fabaceae:
Mimosoideae), commonly known as leucaena, is a
shrub or small tree that has become naturalized in
more than 120 tropical, subtropical and warm
temperate countries worldwide, following delib-
erate introductions for agroforestry and other
purposes (Hughes 2006). The plant is regarded as
one of the ‘World’s 100 Worst Invaders’ (Hughes
2006), but is still considered to be an incipient or
‘emerging weed’ in South Africa (Olckers 2004;
Henderson 2007). Several features, notably self
compatibility and prolific seed production, promote
invasions of forest margins, river banks, drainage
lines, roadsides and other disturbed areas in
several countries worldwide (Henderson 2001;
Raghu et al. 2005; Hughes 2006; Tuda 2007). The
plant constitutes a typical ‘conflict species’ because,
although it is highly invasive, it has a multitude of
useful attributes that include nitrogen fixation for
crops, livestock fodder, fuel wood, pulping for
paper, sand-binding and reforestation (Henderson
2001; Hughes 2006; Orwa et al. 2009). In South
Africa, leucaena is cultivated mainly as a source of
fuel wood and livestock fodder (Neser & Klein
1998).
In many situations in South Africa, the plant is
unable to realize its full invasive potential because
of suboptimal environmental conditions and
extensive browsing by livestock and wild animals
(Neser & Klein 1998). However, when conditions
are suitable and browsing is limited, leucaena
populations display rapid growth rates and high
seed production, and trees taller than 2–3 m
escape browsing. In such situations, mechanical
control is largely ineffective because the plants
coppice readily unless the cut stumps are treated
with diesel or herbicides (Hughes 2006) and may
form denser thickets that are difficult to utilize.
Although herbicides like picloram and meta-
sulfuron-methyl have been used elsewhere in the
world (Hughes 2006), no herbicides are currently
registered for use against the plant in South Africa.
Despite its incipient weed status in South Africa,
there were concerns that leucaena would cause
problems similar to those caused by some of South
Africa’s worst invaders, notably Australian wattles
African Entomology 19(2): 356–365 (2011)
 Olckers: Biological control of Leucaena leucocephala (Fabaceae)
and Prosopis species, which share the same attrib-
utes and were imported for very similar reasons
(Neser & Klein 1998). Biological control was thus
considered as an attractive option to restrict the
plant’s spread at an early stage (Olckers 2004)
without negating its benefits, tactics that, in hind-
sight, should have been employed for other such
‘conflict species’.
Although biological control provides an alterna-
tive to conventional control methods, South Africa
is currently the only country to have actively pur-
sued this option. However, efforts in South Africa
have not been officially funded and have thus
relied on opportunism, notably the collection of
agents during exploration for agents of other
higher-priority invasive species (Olckers 2004). In
addition, the considerable benefits provided by
the plant’s vegetative organs, coupled with its
extensive seed production, necessitated a focus on
seed-feeding agents and culminated in the release
of a seed-feeding beetle in 1999. However, biological
control efforts in South Africa have remained at
a low level and besides information that has
appeared in unpublished reports and popular
articles, only one scientific paper has so far been
published on the topic (Shoba & Olckers 2010).
The aim of this review is thus to consolidate all
available information on the biological control
programme against L. leucocephala in South Africa,
in particular: (i) aspects of the target plant that are
pertinent to its invasiveness and management; (ii)
details of the natural enemies that have featured
so far; (iii) progress that has been achieved; and
(iv) the direction of future research efforts.
THE TARGET PLANT
Description
Several published and electronic sources
(Henderson 2001; Hughes 2006; Orwa et al. 2009)
provide a description of the plant, which comprises
an unarmed, evergreen shrub to medium-sized
tree that varies from 3–15 m in height. Plants in the
genus Leucaena Benth., comprising some 23 species
(USDA 2010), differ from all other mimosoid
Fabaceae by their hairy anthers, while L. leucocephala
is characterized by intermediate leaflets and large
pods that occur in clusters of 5–20 per inflores-
cence. The leaves are bipinnate with an elliptic
convex extra-floral nectary on the petiole, 4–9
pairs of pinnae and 13–21 pairs of leaflets on the
pinnae (Fig. 1). The creamy-white flowers occur in
357
globose heads while the pods occur in crowded
clusters and are pendulous, flattened and papery
(Fig. 1). The specific name ‘leucocephala’ refers to
the flowers and is derived from ‘leu’, meaning
white, and ‘cephala’, meaning head. The pods
are passively dehiscent, splitting into two non-
recurving halves (Henderson 2001), with 8–18
seeds per pod. Combinations of flowers, immature
and mature pods are often present on plants at the
same time.
Origin and taxonomy
The plant is native to Mexico and Central
America and possibly parts of the West Indies,
northern South America and the southern U.S.A.,
although the precise extent of its native range has
been confounded by natural variation and exten-
sive cultivation in many of these countries
(Hughes 2006). Indeed, the exact origin of L. leuco-
cephala remains unknown and neither centres of
morphological diversity nor clearly natural popu-
lations have been confirmed (Orwa et al. 2009).
Synonyms for L. leucocephala include Acacia glauca
Willd., Acacia leucocephala (Lam.) Link, Leucaena
glabrata Rose, Leucaena glauca (L.) Benth. and
Mimosa leucocephala Lam. (Hughes 2006; USDA
2010).
Three subspecies are currently recognized,
namely leucocephala, glabrata (Rose) Zárate and
ixtahuacana C.E. Hughes (Orwa et al. 2009; USDA
2010). The natural distribution of subspecies
glabrata is unclear, since it is widely cultivated in
Mexico and Central America and difficult to differ-
entiate from natural populations (Orwa et al.
2009). By contrast, subspecies ixtahuacana is
restricted to a small area of northern Guatemala
and neighbouring southern Mexico and has not
been introduced outside this range (Orwa et al.
2009). Subspecies leucocephala is particularly wide-
spread and abundant on Mexico’s Yucatán Penin-
sula, leading to speculation that this is the centre of
origin (Orwa et al. 2009).
The subspecies glabrata and leucocephala have
been spread pantropically (Hughes 2006), with
leucocephala believed to have been introduced out-
side of Mexico during the 16th Century and
glabrata only in the last few decades (Orwa et al.
2009). These subspecies comprise arborescent
(glabrata) and shrubby (leucocephala) forms and are
commonly referred to as the ‘giant or Salvador
type’ and the ‘common or Hawaiian type’, respec-
tively (Hughes 2006). By the late 19th Century,
358 African Entomology Vol. 19, No. 2, 2011
X 1/2
100 mm
Fig. 1. Leucaena leucocephala. (Drawn by G. Condy; first published in Henderson (1995), ARC-Plant Protection
Research Institute, Pretoria.)
L. leucocephala leucocephala had spread or been
introduced throughout Asia and Africa and is now
considered to be a widespread tropical weed in
more than 20 countries (Hughes 2006; Orwa et al.
2009). The plants that have become invasive in
South Africa conform to the leucocephala form
(Neser & Klein 1998) which is considerably more
aggressive than the glabrata form.
Introduction and distribution in South Africa
Although the exact date of introduction is
unknown, L. leucocephala has been present in
South Africa since early in the 20th Century (Tribe
1995). Environmental constraints, notably climatic
limitations, were perceived as barriers to invasion
and only low altitude (<300 m) or high-rainfall
(>800 mm of annual rainfall) regions were
deemed to be at risk, these comprising the eastern
escarpment and coastal regions of South Africa
(see references in Tribe 1995). These predictions
have largely been accurate and although infesta-
tions have been reported in several South African
provinces, notably Limpopo, Mpumalanga,
KwaZulu-Natal and the Eastern Cape, these are
mostly in the KwaZulu-Natal coastal region
(Fig. 2). The highest densities occur in the
Durban-Tongaat area (Neser & Klein 1998), while
scattered populations occur southwards into the
Eastern Cape and northwards to the Mozambique
border. Leucaena leucocephala is not regarded as a
‘prominent invader’, according to the Southern
African Plant Invaders Atlas (SAPIA): most records
of invasion have arisen from the Savanna Biome
(115 records), followed by watercourses and
 Olckers: Biological control of Leucaena leucocephala (Fabaceae) 359

Fig. 2. Distribution of Leucaena leucocephala and the seed-feeding beetle Acanthoscelides macrophthalmus (o) in
South Africa. (Drawn by L. Henderson; data source: SAPIA database, ARC-Plant Protection Research Institute,
Pretoria.)

wetlands (43), forest habitats (11) and the Grass-
land Biome (8) (Henderson 2007).
Biology and ecology
The plant’s biology and ecology (summarized
from Hughes 2006; Orwa et al. 2009) predisposes it
to invasion. The flowers are self-fertile with most
seeds arising from self-pollination, and isolated
trees are thus capable of seed production. How-
ever, the plants are also capable of out-crossing
and are pollinated by a wide range of generalist
insects, notably bees. With sufficient moisture,
flowering and seed-pod production occurs
throughout the year and seed set is extensive, with
individual plants capable of producing in excess of
30 000 seeds (Raghu et al. 2005). The pods ripen in
10–15 weeks and release their seeds in response to
moisture but will retain them on the plant for
longer in times of drought (Raghu et al. 2005). The
seeds have hard waxy coats that enable them to
remain viable for at least 10–20 years in the soil
seed-banks, permitting germination for pro-
longed periods after dispersal. In addition, seed-
lings are able to produce seed within one year.
Given its economic importance, there is a consid-
erable body of information (see Hughes 2006;
Orwa et al. 2009) on the plant’s requirements and
limitations. Leucaena is a short-lived (20–40 years)
tropical plant that requires warm temperatures
and high moisture regimes for optimal growth
and thus displays reduced growth during cold
winter months. Optimal growth conditions are
limited to 15–25 degrees north or south of the
equator, meaning that most of South Africa is
climatically suboptimal. The plant is also frost
sensitive, shedding its leaves after light frosts.
Heavy frosts kill the above-ground growth, but
the plants re-sprout during summer. Sub-humid
360 African Entomology Vol. 19, No. 2, 2011
or humid climates with moderate dry seasons are
preferred and the plant thrives under high rainfall
(>1200 mm/year) conditions, whether natural or
irrigated. Leucaena is also tolerant of fast fires but
can re-grow if damaged by slower fires. In general,
the plant is considered to be limited by altitude
(0–1500 m), mean annual temperature (25–30 °C),
mean annual rainfall (650–3000 mm) and soil con-
ditions (neutral to alkaline). Although leucaena is
drought tolerant, it is intolerant of soils with low
pH, low phosphorus and calcium, high salinity
and water logging. The plant’s deep root system
ensures that it persists on a wide range of soils but
grows best on deep clay soils which are fertile,
moist and alkaline.
The plant aggressively colonizes disturbed sites,
even in its native range, because of several inva-
sive properties that include self-fertility, year-
round flowering and fruiting, abundant seed
production, extensive seed banks, and its ability to
coppice after frost, fires or clearing. Areas that are
invaded include open (often coastal) habitats,
semi-natural, disturbed and degraded habitats
and even agricultural land following deliberate
plantings. There is, however, little evidence of
invasion of undisturbed natural vegetation, at
least in South Africa (Neser & Klein 1998).
BIOLOGICAL CONTROL
The biological control programme against
L. leucocephala commenced during the late 1980s
when surveys in the Americas, for agents of other
higher-priority weeds, also permitted importations
of agents for leucaena. Given that this programme
was low-priority and unfunded, intensive surveys
for candidate agents were never carried out and
collections of agents were sporadic. Also, given
the plant’s agricultural and economic potential,
biological control options were restricted to
seed-feeding agents, a strategy often used in
South Africa where conflicts of interest have arisen
(Impson et al. 1999). In particular, leucaena’s status
as an incipient weed suggested that it was consid-
erably more vulnerable to biological control than
highly invasive related weeds (e.g. Australian
wattles) that had been naturalized in South Africa
for far longer.
Agroforestry databases list only one insect species,
the leucaena psyllid Heteropsylla cubana Crawford
(= Heteropsylla incisa (Sulc)), as capable of causing
extensive damage to leucaena plantations (Orwa
et al. 2009). Since this insect was inadvertently
introduced to Africa, including South Africa, in the
1990s, its potential contribution to biological
control will be considered. Only one agent, the
seed-feeding beetle Acanthoscelides macrophthalmus
(Schaeffer), has otherwise been considered and
will thus be the main focus of this review. Besides
its deliberate introduction into South Africa, 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), presum-
ably via contaminated seeds.
Heteropsylla cubana (Psyllidae)
Native to Central and South America, the
leucaena psyllid achieved international notoriety
in the early 1980s when outbreaks and devastating
defoliation of leucaena plantings were first detected
in Florida (U.S.A.) and then Hawaii, from where
it spread rapidly throughout the Asian-Pacific
region, including Australia, during the late 1980s
and early 1990s (Napompeth 1995). First reported
in Africa (Kenya) in 1992, the insect spread into the
moist tropical and subtropical regions of several
neighbouring East African countries (Napompeth
1995), but was first recorded in South Africa, near
Hazyview (25.02.48S 31.07.46E) in Mpumalanga
Province, as late as 1998 (Morris 2000; I. Millar,
pers. comm.). One year later, it was recorded
several hundred kilometres southwards on experi-
mental plantations of leucaena in Pietermaritz-
burg (29.40.26S 30.24.51E), KwaZulu-Natal, where
it caused an immediate reduction in leaf growth
and forage yield (Morris 2000). The manner in
which H. cubana achieved its rapid global distribu-
tion is unclear since this does not appear to have
been human-assisted, particularly because
leucaena is typically transported via seeds and not
vegetative material (Napompeth 1995).
Although the native host range of H. cubana is
considered to be restricted to Leucaena and other
closely related legume genera (Napompeth 1995),
some Leucaena species, and even some varieties of
L. leucocephala, are resistant to the insect, suggesting
some degree of host specificity and possibly some
potential as a biological control agent. Although
older leaves are generally avoided, sap-sucking of
the developing leaflets at the shoot tips, by the
adults and immature stages, may inflict substantial
damage to leucaena cultivations and wild popula-
tions. In particular, their rapid life cycles (10–20
 Olckers: Biological control of Leucaena leucocephala (Fabaceae)
days from egg to adult) and population doubling-
times of 2.52 days facilitate massive population
increases in tropical regions (Napompeth 1995).
Studies in tropical Asia revealed that population
increases are negatively correlated with tempera-
ture, and that the populations peak in cooler areas
and during the cooler parts of the year (Napompeth
1995; Geiger & Gutierrez 2000). The insect also has
a distinct upper thermal limit and numbers often
decline substantially during warmer periods
(Napompeth 1995).
Original reports from several countries (Napom-
peth 1995) described extensive defoliation that
resulted in stunted growth, economic losses and
sometimes widespread mortality of trees. In eastern
Africa, initial damage was so severe that farmers’
confidence in the plant’s economic viability declined
substantially (Morris 2000). South African popula-
tions of leucaena were deemed to be equally
vulnerable to the psyllid because of the narrow
genetic base of the ‘Hawaiian-type’ stocks that
comprised most stands in the country (Morris
2000). However, despite major concerns at the
time, damage in South Africa did not reach the
levels described in other parts of the continent.
The 1999 infestation that was reported in Pieter-
maritzburg did not proliferate and appeared to
have disappeared by the subsequent growing
season (Morris 2000). This appeared to be unre-
lated to specific site conditions (e.g. leaf abscission
during cold dry winters) because H. cubana has
also not proliferated in the warmer coastal areas.
Indeed, the insect appears to be intolerant of high
temperatures and recent observations in Kwa-
Zulu-Natal have failed to provide any evidence of
flourishing populations (T. Olckers, pers. obs.).
Currently, H. cubana does not appear to pose any
significant threats to L. leucocephala populations in
South Africa or elsewhere in the world. Although
cyclical defoliations have been reported in several
countries, the psyllid appears to have been
brought under control by the cultivation of resis-
tant varieties of leucaena and by several generalist
predators and parasitoids, including native species
and exotic species introduced for biological
control (Hughes 2006). In South Africa it is possible
that generalist predators were acquired from
presumably native Cicadellidae, identified in 1990
as ‘nr. Accacididia sp.’ (M. Webb, Natural History
Museum, London; AcSN 1575), that were often
found in considerable numbers on young, stunted
growth of the plant, at least around Pretoria
361
(25.43.39S 28.14.13E) (S. Neser, pers. comm.).
Consequently, H. cubana is not expected to play
any role in the biological control of leucaena in
South Africa.
Acanthoscelides macrophthalmus
(Chrysomelidae: Bruchinae)
A small colony of A. macrophthalmus was origi-
nally collected from infested leucaena seed in an
area of Texas (U.S.A.) bordering Mexico, on a
general collecting trip in 1989 and imported into
South Africa. Unfortunately, details and coordi-
nates for the locality(ies), for this and for subse-
quent collections, are no longer on record. The
biology and host range of the beetle was first
investigated in quarantine (Neser & Klein 1998)
and later re-assessed following release (Shoba &
Olckers 2010) and details from these studies are
summarized below. The insect remained in quar-
antine for some 10 years because the conflicts of
interest surrounding leucaena were only resolved
in the late 1990s. Following its decline in quaran-
tine, the original culture was later augmented by a
collection from Florida (U.S.A.), in July 1999, while
in September 2005 a fresh laboratory culture was
established following collections in Sonora,
Mexico (S. Neser, pers. comm.).
Under laboratory conditions (28 °C), adult
beetles lived for around 12 days when maintained
on water alone and females laid eggs some three
days after emergence. Since adult Bruchinae often
feed on pollen and nectar in the field (Southgate
1979), adult longevity may be considerably longer
with food supplementation. In the laboratory,
eggs are typically deposited on the surfaces of the
ripe pods and ripe, loose seeds of L. leucocephala. In
the field, ripe leucaena pods split open and release
their seeds and it is unclear whether the beetles
typically oviposit on dehisced seeds on the soil
surface or whether they prefer canopy-held pods
that still contain seeds. Hatching larvae burrow
into the seed and consume much of the seed con-
tents during their development. Before pupating,
the final instars excavate a circular ‘emergence
hatch’ on the seed surface which is forced open by
the emerging adult. The duration of the immature
stages (egg to adult) is around 36 days at 28 °C.
The beetle’s host range was originally assessed
by multi-choice tests, in which adults were exposed
to the seeds of several non-target plants in the
subfamilies Mimosoideae, Caesalpinioideae and
Papilionoideae of the family Fabaceae (Neser &
362 African Entomology Vol. 19, No. 2, 2011

Table 1. Chronological record of releases of Acanthoscelides macrophthalmus against Leucaena leucocephala in

KwaZulu-Natal, South Africa.

Site Town: (map reference) Site Vegetation First/last No. of Total Outcome
No. (disturbance) release dates releases released1

1 Umhlanga Rocks: Outskirts Transformed 17 Apr 00 1 140a Established

(29.41.93S 31.04.55E) (roadside)
2 Hilton: Cedara Farm Transformed 02 May 00 13 660a Uncertain
(29.32.30S 30.16.33E) (planted stand) 21 Nov 00 (site cleared)
3 Pietermaritzburg: UKZN Campus Transformed 14 June 04 1 70a Established
(29.37.46S 30.24.07E) (planted stand)
4 Pietermaritzburg: UKZN Farm Transformed 03 Feb 05 6 4010a,s Established
(29.40.26S 30.24.51E) (pasture) 01 July 05
5 Westville: UKZN Conservancy Forest along 27 Jan 06 2 3400a,s Established
(29.49.24S 30.57.05E) watercourse 23 Feb 06
6 Verulam: Sewage treatment Transformed 13 Apr 06 1 2310a,s Established2
(29.40.46S 31.02.08E) works (wasteland)
7 Durban: Nr UKZN Campus Transformed 18 Dec 07 1 650a,s Established2
29.51.26S 30.57.09E) (roadside)
8 Durban: Old quarry in Transformed 09 Apr 09 1 600a,s Established2
(29.48.49S 31.00.45E) Umgeni Road (wasteland)
9 Durban: Along Umgeni Transformed 09 Apr 09 1 1000a, s Established2
(29.48.25S 30.58.48E) Road (wasteland)
10 Karridene: Nr Protea Hotel Transformed 24 June 09 1 200a,s Established2
(30.07.26S 30.50.12E) (wasteland)
11 Amanzimtoti: Outskirts Transformed 24 June 09 1 400a,s Established2
(30.02.00S 30.53.28E) (roadside)
12 Mtubatuba: Nr Primary School Transformed 30 June 09 1 400a,s Established2
(28.25.38S 32.10.56E) (roadside)
13 Mtubatuba: Outskirts Transformed 30 June 09 1 400a,s Established2
(28.25.04S 32.10.43E) (roadside)
1
Stages released included adultsa or seeds infested with larvaes.
2
Populations were already established at the time of release.

Klein 1998). Although A. macrophthalmus oviposited
on the seeds of several of these species, survival to
adulthood was confined to Leucaena species and
the beetle was cleared for release in 1999. The
insect’s host range was later re-assessed following
a report of a single host-range expansion on a
native species of Mimosoideae in Taiwan (Tuda
et al. 2009) which raised concerns of similar attacks
on native South African Mimosoideae (e.g. Acacia
species) that are closely related to the genus
Leucaena. These trials, which involved both
no-choice and multi-choice tests, confirmed that
native South African Mimosoideae are not at risk
and thus that the decision to release the beetle was
justified (Shoba & Olckers 2010).
Following clearance for release in 1999, A.
macrophthalmus was released around Pretoria
(25.43.39S 28.14.13E) in Gauteng Province where it
became established. Most releases have taken
place in KwaZulu-Natal, the first of these in 2000
and the remainder on several occasions during
2004–2009 (Table 1). Evidence of establishment in
KwaZulu-Natal became apparent from 2005
onwards, with even small releases (<200 beetles;
Table 1) proving successful and there were signs
that the beetle was already present at several sites
when initial releases were undertaken (Table 1).
The beetle has now become widely established in
the province (Fig. 2), even at localities distant from
the original release sites. Around 2005, the beetle
was released and established near Buffelspoort
(25.46.54S 27.32.23E) in the North West Province
 Olckers: Biological control of Leucaena leucocephala (Fabaceae)
(S. Neser, pers. comm.). During 2008, the beetle
was also established at two release sites in the low-
altitude region of Mpumalanga, near Boulders
(25.53.94S 31.32.33E) and Buffelspruit (25.49.81S
31.51.46E) (Fig 2), and has since become wide-
spread in the southeastern region bordering
Swaziland (D. Strydom, pers. comm.). All releases
in KwaZulu-Natal, and elsewhere in the country
prior to 2005, were carried out with stocks originat-
ing from Florida/Texas, while releases in the
northern regions of the country after 2005 were
carried out using the Mexican stocks.
Although A. macrophthalmus has become wide-
spread in the KwaZulu-Natal coastal region,
where leucaena infestations are most abundant,
its impact on the weed’s seed dynamics has not
been quantified. By contrast, levels of seed dam-
age in southeastern Queensland (Australia) varied
from 11–54 %, depending on the length of time
(1–4 months) that the pods remained on the trees
(Raghu et al. 2005), while 67 % seed damage was
reported in West Africa (Delobel & Johnson 1998).
In particular, the plant’s ability to produce seeds
throughout the year, coupled with the longevity of
seeds in the seed bank, suggest that the phenology
of the beetle in relation to seed availability will be
crucial in determining whether sufficient seed
damage is inflicted consistently throughout the
year. A study to examine the rates of damage on
canopy-held seeds over time is currently in prog-
ress in the KwaZulu-Natal coastal region. Another
important aspect is whether oviposition in the
field is restricted to canopy-held seeds and thus
whether dehisced seeds escape damage. The
study in Australia (Raghu et al. 2005) which indi-
cated that seed damage by A. macrophthalmus was
limited by pod retention times, suggested that, de-
spite their acceptance of loose seeds in the labora-
tory, the beetles failed to locate dehisced seeds in
the field. If this is true, then their impact on
leucaena seed survival may be limited even fur-
ther. Experiments to test the beetle’s selection of
oviposition sites under natural conditions are also
currently in progress.
Another factor that could influence rates of seed
damage by A. macrophthalmus is the recruitment of
parasitoids, particularly those on native Bruchinae
that feed on indigenous South African mimosoid
Fabacaeae (Impson et al. 1999). Parasitism of the
immature stages of A. macrophthalmus has been
reported in countries where it was inadvertently
introduced (Delobel & Johnson 1998; Raghu et al.
363
2005) and also in South Africa (Shoba & Olckers
2010; S. Neser, pers. comm.). Preliminary surveys
in KwaZulu-Natal have already recorded at least
five species of hymenopteran parasitoids (all
Chalcidoidea) of the larvae/pupae and have
suggested that mortality caused by parasitism
(10–14 %) is significant. Eggs on pods in the field in
Gauteng are often parasitized by unidentified
species of Mymaridae and Trichogrammatidae,
but the levels of egg parasitism have not been
determined (S. Neser, pers. comm.). Several wasp
families are known to parasitize the immature
stages of Bruchinae (e.g. Hoffmann et al. 1993;
Impson et al. 1999; Kingsolver 2004); often with
negative consequences in weed biological control
programmes (Van Klinken & Flack 2008). It is thus
possible that a more intensive assessment, currently
in progress, will reveal more parasitoid species
and higher rates of parasitism.
DISCUSSION
Although A. macrophthalmus is widely distributed
in the KwaZulu-Natal coastal region, the extent
of its distribution elsewhere in the country is
unknown. Surveys in other leucaena-infested
regions should thus be undertaken to resolve this
uncertainty and to determine whether additional
releases by implementing agencies are required.
The question of whether A. macrophthalmus has
made, or will make, a significant contribution to
the management of leucaena populations needs to
be addressed by assessing: (i) the population
dynamics of the beetle, and levels of seed destruc-
tion, in relation to seed availability over time; (ii)
the beetle’s oviposition preferences in relation to
canopy-held versus dehisced seeds and; (iii) levels
of predation and parasitism on the immature
stages of the beetle.
It is highly unlikely that A. macrophthalmus on its
own will be sufficient to regulate populations of
L. leucocephala as this will depend on very high
levels of seed destruction being achieved, during
periods of both seed abundance and seed limita-
tion. Seed feeders are often unable to regulate
plant populations because of their inability to
track temporal fluctuations in seed availability,
which may be aggravated by excessive mortality
of the immature stages (Van Klinken & Flack 2008).
The Australian study (Raghu et al. 2005) concluded
that the beetle populations were unable to reach
sufficiently high numbers to exploit available
364 African Entomology Vol. 19, No. 2, 2011
seeds and that, despite seed damage, soil
seed-banks doubled over the 4-month study
period. Several studies have suggested that in the
case of legumes with high levels of seed produc-
tion, seed feeders need to destroy at least >95 % of
seeds produced (see references in Raghu et al.
2005), and probably >99 % (Hoffmann & Moran
1998), in order to ensure population regulation.
Although preliminary observations have suggested
that such high levels will not be realized in South
Africa, this should be viewed in the context of
the weed’s overall management. Intensive and
sustained clearing of adult plants, but also of
recruited seedlings, by the Working for Water
Programme along watercourses in South Africa
could ensure that even moderate levels of pre-
dispersal seed destruction could be significant in
reducing the costs of long-term clearing operations
(Moran et al. 2004). In addition, seedlings and plants
coppicing after clearing are browsed extensively
by both domestic and wild animals and should
thus take considerably longer to reach a size where
seeds are produced. In any event, the release of
seed-feeding or seed-reducing agents is often
viewed as an important ‘first line of defence’ in
weed biological control and the release of addi-
tional agents that attack other parts of the plant are
crucial in achieving acceptable levels of control
(Hoffmann & Moran 1998; Moran et al. 2004).
Whether or not the release of additional agents is
required, or possible, given the weed’s benefits,
will depend on the outcomes of the proposed
studies.
Another aspect worthy of investigation is the
extent to which the consumption of dehisced
seeds by rodents is likely to affect seed bank densi-
ties and hence seedling recruitment. If ‘seed rain’
is regularly and significantly depleted by rodents,
then failure by A. macrophthalmus to exploit
dehisced seeds may not pose the problem that has
been suggested in Australia. The contention that
seed damage by A. macrophthalmus does not neces-
sarily imply seed mortality and that small propor-
tions of damaged seeds are still capable of
germination (Raghu et al. 2005) should also be veri-
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I thank M.P. Hill (Rhodes University), S. Neser,
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