Bulletin of Entomological Research (2006) 96, 379–386 DOI: 10.

1079/BER2006442

Field infestation, life history and

demographic parameters of the fruit fly
Bactrocera invadens (Diptera: Tephritidae)
in Africa
S. Ekesi *, P.W. Nderitu and I. Rwomushana
International Centre of Insect Physiology and Ecology, PO Box 30772-00100
GPO, Nairobi, Kenya

Abstract

Field infestation rates of an invasive fruit fly species, Bactrocera invadens Drew
Tsuruta & White on mango was determined at different localities in Kenya. At
most of the locations and especially at low elevations, B. invadens frequently shared
the same fruit with the indigenous fruit fly species Ceratitis cosyra (Walker) but
often occurred at higher numbers than C. cosyra. The level of infestation varied
with location ranging from 3.0 to 97.2 flies per kg of fruit. There was a significant
inverse relationship between numbers of flies per kg of fruit and elevation at which
fruit was collected, suggesting that B. invadens is a predominantly lowland pest.
On an artificial diet, development of B. invadens immatures lasted 25 days; egg
incubation required 1.2 days, larval development 11.1 days and puparia-adult
development 12.4 days. About 55% of eggs developed to the adult stage. Life
expectancy at pupal eclosion was 75.1 days in females and 86.4 days in males.
Average net fecundity and net fertility were 794.6 and 608.1 eggs, respectively,
while average daily oviposition was 18.2 eggs. Daily population increase was 11%
and mean generation time was 31 days. Results are discussed in relation to the
biology and ecology of the insect and in the development of mass rearing and
control measures for B. invadens.

Keywords: Bactrocera invadens, Ceratitis cosyra, mango, field infestation, elevation,

life history, demographic parameters

Introduction worldwide (Clarke et al., 2005). In March 2003, three

Asian fruit fly pests from the genus Bactrocera are
regarded as some of the most destructive insects of fruits
and vegetables worldwide (White & Elson-Harris, 1992).
Losses are due to direct feeding damage and loss of export
market opportunities through quarantine restrictions
imposed by importing countries to avoid entry and estab-
lishment of unwanted fruit flies. With adult traits that can
include high mobility and dispersive powers, high repro-
ductive rates, and extreme polyphagy, Bactrocera species are
well-documented invaders and rank high on quarantine lists
*Fax: +254 20 863 2001
E-mail: sekesi@icipe.org
specimens of such a fly were detected in Kenya (Lux et al.,
2003), and the insect was described as Bactrocera (B.) invadens
Drew Tsuruta & White (Drew et al., 2005).
Bactrocera invadens is believed to belong to the Bactrocera
dorsalis complex of tropical fruit flies (French, 2005). This
complex comprises more than 75 species largely endemic to
South-East Asia (Drew & Hancock, 1994) with undescribed
species remaining in collections (Lawson et al., 2003) and the
group is arguably one of the most important pest species
complexes in world agriculture (Clarke et al., 2005; Drew
et al., 2005). Bactrocera invadens is believed to have invaded
Africa from the Indian subcontinent and was discovered in
Sri Lanka after it was first reported from Africa, where it
has become a significant pest of quarantine and economic
importance (Mwatawala et al., 2004). The insect is rapidly
380 S. Ekesi et al.
spreading across tropical Africa and in addition to Kenya, it
is now reported from Benin, Cameroon, DR Congo, Ghana,
Guinea, Mali, Nigeria, Senegal, Sudan, Tanzania, Togo,
Uganda and Equatorial Guinea and the Comoros Island
(Drew et al., 2005; French, 2005; S. Ekesi, unpublished data).
Preliminary results from host range studies show that it
attacks both cultivated and wild hosts including mango
Mangifera indica L. (Anacardiaceae), lemon orange Citrus
limon (Burman f.) (Rutaceae), tomato Lycopersicon esculentum
Miller (Solanaceae), banana Musa sp., (Musaceae), guava
Psidium guajava L. (Myrtaceae), marula Sclerocarya birrea
H. (Anacardiaceae), custard apple, Annona muricata L.
(Annonaceae), avocado Persea americana M. (Lauraceae) and
Indian almond Terminalia catappa L. (Combretaceae) (I.
Rwomushana, unpublished data; Z. Seguni, unpublished
data). The pest has also been reared from some unidentified
fruits in indigenous coastal forests; hence sufficient repro-
ductive bases exist even during the off-season for cultivated
fruits. Among the cultivated fruits sampled so far, mango
appears to be the primary host. However, because of the
‘novelty status’ of B. invadens, nothing is known with regard
to the actual level of damage to mango fruits from different
geographical areas of Kenya.
Only a few species of Bactrocera have become established
outside their native oriental range, namely Bactrocera
cucurbitae (Coquillet) in eastern Africa and Hawaii, Bactrocera
latifrons (Hendel) in Hawaii, Bactrocera oleae (Gmelin) in
California and members of the B. dorsalis species complex in
both Hawaii and Surinam (White & Elson-Harris, 1992; Rice
et al., 2003; Clarke et al., 2005). The invasion of B. invadens
in Africa adds to this list and compounds the existing fruit
fly problem in Africa. If the presence of B. invadens is not
addressed quickly through efficient control measures, this
fly may dramatically limit the freedom of fruit trade and
market access.
Most fruit flies from the genus Bactrocera are considered
to be lowland residents. In studies at an upper-elevation
fruit-producing area on the slopes of Maui volcano in
Hawaii, Wong et al. (1985) observed that captures of the
Oriental fruit fly Bactrocera dorsalis (Hendel) were inversely
related to elevation, and were positively correlated with host
fruiting. Similarly, in Kauai, Harris et al. (1986) found that
trap captures of the melon fly B. cucurbitae were high in
coastal agricultural areas but declined sharply with increas-
ing elevation. In a recent survey using methyl eugenol as an
attractant, with sampling points across Kenya, Uganda and
Tanzania (S. Ekesi et al. unpublished data) high numbers of
Table 1. Sampling sites of mangoes in Kenya.
Province Location Latitude
Central Kiambu 00 320 58
Central Kamiti 00 490 58
Central Muranga 00 320 56
Coast Malindi 03 110 45
Coast Taveta 03 240 48
Coast Muhaka 04 190 17
Eastern Kitui 01 240 32
Eastern Embu 00 280 34
Eastern Meru 00 200 45
Rift Valley Nguruman 01 480 28
Western Kisumu 00 060 75
B. invadens were detected as high as 1600 m above sea level
(masl). However, it is still unknown if the fly infests
mangoes grown at such elevations.
Fruit infestation data and demographic population analy-
sis have diverse applications: developing control strategies,
analysing population stability and structure, estimating
extinction probabilities, predicting life history evolution and
outbreaks, understanding intra- and interspecific relation-
ships, examining the dynamics of colonizing or invading
species development and application of control methods
(Levins & Wilson, 1980; Liedo et al., 1992; McPeek & Kalisz,
1993; Aluja et al., 1996; Peck & McQuate, 2004). The reso-
lutions upon which pest management decisions are made
and applied are not attainable without an explicit knowledge
of damage and biology of the pest. Also, studies on field
infestation rates on mangoes are important in establishing
the pest status of B. invadens and for formulating control
strategies for the fly’s suppression. The main objectives of
this study were therefore to establish the level of infestation
by B. invadens in mango fruits from different localities in
Kenya, in order to determine pre-adult (egg to adult) life
history parameters when reared on an artificial diet. These
data can then be used to estimate adult demographic
(biological and population) parameters and to define
strictures on population increase.
Materials and methods
Collection of mango fruits and handling of insects
Ripe and, occasionally, unripe fruits were sampled from
the tree and the ground (as ‘windfalls’) between January
2004 and March 2005. A mixture of mango varieties were
sampled but the most common varieties encountered were
Apple, Kent, Ngowe and Boribo. For each site, GPS data on
longitude, latitude and elevation were recorded (table 1).
Fruits were transported to the laboratory in screened 3-l
plastic buckets, weighed and counted. Fruits from separate
collections were placed individually in 1-l rectangular,
plastic containers that had
0.5 and 2.5 cm ellipsoidal holes
cut at the bottom (Copeland et al., 2002). The container was
then nested inside a 2-l rectangular plastic container with
heat-sterilized moistened sand at the bottom for pupation.
The holes in the smaller container allowed mature pupariat-
ing larvae to fall into the larger container below to pupariate
after leaving the mango fruits. The formation of visible
structures that follows the last larval instar in Tephritidae is
Longitude Elevation (m)
S 37 080 21 E 1654
S 37 080 21 E 1401
S 37 240 46 E 1602
S 40 050 16 E 27
S 37 420 47 E 601
S 39 310 29 E 43
S 38 020 34 E 1010
S 37 310 03 E 1508
S 37 370 28 E 1552
S 36 030 28 E 700
S 34 450 38 E 1196
 Field infestation and life history of Bactrocera invadens 381

often erroneously called pupation but it is actually puparia-
tion as true pupation takes place unseen within the
puparium (see White & Elson-Harris, 1992).
Sample containers were checked every 3–4 days for
puparia and adult flies. Puparia were sieved from the sand
and held for adult emergence in transparent Plexiglas cages
(25r25r25 cm). Adult flies were fed on diet consisting of 3
parts sugar and 1 part enzymatic yeast hydrolysate ultrapure
(USB Corporation, Cleveland, Ohio, USA), and water on
pumice granules for 5 days until body colour had fully
developed, and then they were allowed to die within the
cages or were killed by freezing. Fruit samples were
discarded after about 4 weeks.
Life history parameter
Egg stage duration
Eggs of B. invadens were collected using a mango dome
(that had the pulp and the seed removed), placed for 1 h into
stock fly colonies maintained in the laboratory on a yeast–
carrot-based artificial diet (Ekesi et al., 2005), hereafter
simply referred to as diet. The insects originated from rotten
mangoes collected at a local market in Nairobi, Kenya, and
have been maintained on the diet for about 30 generations.
Eggs were carefully removed with a fine camel’s hair brush
and placed on a 4-cm square of moist blotting paper. Egg
hatch was determined every 12 h for a period of 3 days. Eight
replications were maintained for a total of 100 eggs per
replicate.
Larval and pupal development
Fifty newly emerged larvae from the lots of eggs
above were gently introduced with a fine camel’s hair
brush onto the surface of 50 g of diet in 50 ml plastic cups.
An experiment consisted of 4 cups of diet containing
50 larvae, with each group of 4 cups replicated 3 times.
Mature larvae were allowed to leave the rearing cup
ad libitum into larger 150 ml plastic cups that contained a
layer of heat-sterilized sand on the bottom. Observations
were performed every 6 h by sifting the sand to record
developmental duration. Puparia were held individually in
30 ml plastic containers for emergence. Length of pupal
period and sex were recorded for each fly that emerged from
puparia.
Adult demographic parameter
At eclosion, 25 pairs of newly emerged adults were held
individually in transparent Plexiglas cages (15r15r15 cm)
with diet and water as above. A small mango dome was
placed in each cage daily to monitor egg laying. Eggs on the
dome were gently removed as previously described, counted
and placed on a moist blotting paper and held in Petri dishes
for determination of egg hatch. Hatching was assessed after
72 h. All life history and demographic parameters were
generated in a room maintained at 28+1 C, 50+8% RH with
a photoperiod of L12 : D12.
Statistical analysis
Fruit flies per kg of field-collected mango fruits were
regressed against elevation from the GPS readings to test for
a positive relationship between these two parameters (PROC
REG; SAS Institute, 1985). Population parameters, life and
fertility tables were estimated following the methods
described in Carey (1993).
Results and Discussion
Mango fruit infestation
Field infestation rate of mangoes collected from the
different locations in Kenya is shown in table 2. In most
locations, B. invadens frequently shared the same fruit with
Ceratitis cosyra (Walker); therefore the number of C. cosyra
that emerged from collected fruits is also presented.
Infestation rates ranged from 3.0 to 97.2 flies per kg of fruit.
Highest numbers of B. invadens were collected at low
elevations, while C. cosyra appeared to dominate at high
elevations. None of the fruit samples collected at Kiambu,
Meru and Embu had B. invadens and, in general, in the
highlands (elevation > 1500 masl) fruit infestation did not
exceed 5 flies per kg of mangoes. Regression analysis yielded
a significant inverse relationship between the numbers of
fruit flies per kg of fruits and elevation (fig. 1). The present
result shows that B. invadens is predominantly a lowland
species. Bess (1953) reported an inverse relationship between
numbers of the Oriental fruit fly B. dorsalis and elevation
from 240–1220 masl on the islands of Hawaii and Maui.
Similarly, Vargas et al. (1983) demonstrated that fruit
infestation by B. dorsalis in native and exotic forests on
Kauai Island was moderate at middle (579–800 masl) eleva-
tions and low at high (> 800 masl) elevations. Generally,

Table 2. Level of infestation of mango by Bactrocera invadens and Ceratitis cosyra at different locations in Kenya.

Province Location No. of No. of Fruit % of fruits No. of No. of flies emerged No. of flies
samples fruits weight (kg) infested puparia per kg fruit
B. invadens C. cosyra
Central Kiambu 2 65 21.7 3.1 3 0 3 4.5
Central Kamiti 4 113 37.8 7.1 8 1 5 3.0
Central Muranga 1 56 18.2 19.6 13 2 11 3.5
Coast Malindi 2 92 30.8 56.5 813 803 10 46.9
Coast Taveta 1 55 18.3 61.8 324 303 31 28.6
Coast Muhaka 2 51 17.5 74.5 1231 1056 175 97.2
Eastern Kitui 1 32 11.6 25.3 56 15 41 21.0
Eastern Embu 2 104 34.7 3.8 5 0 5 3.8
Eastern Meru 4 133 44.2 1.5 2 0 2 3.0
Rift Valley Nguruman 6 121 40.5 76.3 654 624 30 21.3
Western Kisumu 1 36 12.5 13.9 19 10 9 4.8
382
120.0
Fruit flies per kg mango
100.0
80.0 y = – 0.0416x + 64.307
R2 = 0.7608
60.0
40.0
20.0
0.0
0 500 1000 1500
Elevation (m)
Fig. 1. Linear regression of mean number of fruit flies per kg of
mango fruits vs. elevation at different localities in Kenya.
elevation by itself does not determine fruit fly distribution
but associated factors such as temperature, rainfall and host
plants at such elevations play a significant role (Nishida
et al., 1980).
With the exception of the interaction between B. dorsalis
and previously established Ceratitis capitata (Wiedemann) on
the Hawaiian Islands (Vargas et al., 1995), the impacts of
invasion of Bactrocera sp. on indigenous fruit fly species have
been little studied (Liquido et al., 1994; Clarke et al., 2005). In
Kenya, prior to the invasion of B. invadens, C. cosyra was the
primary pest of mango. For example, at Nguruman, damage
due to C. cosyra ranged from 60 to 70% in 1998 to 2001 (Lux
et al., 1999; Dimbi, 2003). At the same locality in the current
study, over 80% of mango fruits were infested with both fly
species, with B. invadens accounting for 91% of puparia
collected from the fruits. It will be of great ecological interest
to assess the ability of C. cosyra to coexist with the highly
aggressive B. invadens. In Hawaii, the disappearance of
C. capitata at low elevation was initially assumed to be as a
result of the displacement by B. dorsalis and eradication
strategies were developed based on this analysis (Vargas
et al., 1995). Recent studies have, however, attributed species
replacement to differences in life history strategy between
the two fly species, with C. capitata regarded as an r-selected
species, (smaller and capable of rapid colonization of newly
planted coffee) and B. dorsalis a K-selected species, (larger
with later onset of reproduction) (MacArthur & Wilson,
1967; Vargas et al., 2000). The late onset of reproduction re-
ported by these authors corresponds to the current informa-
tion about B. invadens. So far, these results suggest some
evidence of competitive displacement of C. cosyra by
B. invadens at Nguruman as indicated by the large difference
in pupal harvest from mangoes. However, using data
maintained in our database from 1998 to date, further study
is underway to quantify temporal and spatial population
changes, and the level of fruit infestation by C. cosyra at
Nguruman to clearly establish evidence of competitive
displacement. Additional studies are also investigating
interspecific competition between the two species and
mechanisms involved in such interactions.
The concept of pest free areas and low pest prevalence
zones is being advocated in various parts of the world to
allow for export of fruits from a particular area of a country,
or the entire country, with minimal or zero quarantine
restrictions (Simpson, 1993; Anon., 1999, 2000; FAO, 1999).
The absence of B. invadens on mango in Kiambu and Meru in
S. Ekesi et al.
Table 3. Pre-adult development time and survival of Bactrocera
invadens reared on artificial diet at 28+1 C.
Developmental Development (days) Survival
stage mean+SE (%)
Eggs 1.2+0.02 90.0
Larvae 11.1+3.12 75.6
Puparia 12.4+0.15 80.9
Egg to adult 24.6 55.0
2000
the current survey has an important bearing in the establish-
ment of pest free areas. In 2003 alone, fruit production from
high elevation areas of Kiambu and Meru accounted for

195,024 metric tons (HCDA, 2004). These locations are
ranked among the major fruit production areas of Kenya and
the main fruits grown, targeting both export and domestic
urban markets, are mangoes, citrus, bananas and guava, all
of which are already recognized as hosts to B. invadens
(I. Rwomushana, unpublished data). Although the fruit
samples were rather small, the preliminary data showing
lack of mango infestation by B. invadens and limited
infestation by C. cosyra suggest the possibility of exploiting
such areas as either pest free (for B. invadens) or low pest
prevalence (for C. cosyra) zones. This designation could
permit export of mangoes without strict quarantine restric-
tions if is confirmed that other species of fruit flies do not
occur in these localities. A large-scale delimiting survey for
B. invadens is underway in Kenya under an FAO Technical
Cooperation programme with the aim of exploring for pest
free and/or low pest prevalence areas in the country.
Development and survival of immature stages
The average duration of egg development was 1.2 days,
with a survival rate of 90% (table 3). Larval and pupal
development lasted 11 and 12 days, respectively. Larval
survival was 76% while that of puparia was 81%. Larval
development duration was highly variable, spreading
between 9 and 13 days to pupation, but eclosion from the
pupal stage was spread over just 2 days. Total development
lasted 25 days and only 55% of eggs laid reached the adult
stage (table 3). The duration of egg development observed
for B. invadens in this study falls within the range reported
for other Bactrocera species. In B. dorsalis and B. cucurbitae,
egg duration lasted 1.8 and 1.3 days, respectively while in
B. latifrons it was much longer reaching 3.2 days at the
same temperatures as in the present study (Vargas et al.,
1996). In general, larval and pupal durations were similar to
previous reports by other authors (Vargas et al., 1984, 1996;
Carey et al., 1985; Yang et al., 1994).
Adult life parameters
The average pre-oviposition period was 7.1 days and the
oviposition period lasted 73 days. At 28 C, Vargas et al.
(1997) reported pre-oviposition periods of 5.3–5.7 days for
B. cucurbitae, B. dorsalis and B. latifrons. Table 4 summarizes
biological parameters of adults. Life expectancy at pupal
eclosion was 75.1 days in females and 86.4 days in males.
In B. dorsalis, Yang et al. (1994) found no significant
difference in longevity among male and female flies. In
other fruit fly species, such as C. capitata, life expectancy in
adult males often exceeds that of females (Carey et al., 1995;
 Field infestation and life history of Bactrocera invadens 383

Table 4. Biological parameters of adult Bactrocera invadens reared on artificial diet at

28+1 C.

General trait/parameter Formula Value

Adult life expectancy e0 (days) P
w
Female lx 75.1
x=a
Pw
Male lx 86.4
x=a
Fecundity
b
P
Gross fecundity (eggs) mx 1056.8
x=a
Pb
Net fecundity (eggs) lxmx 794.6
x=a
Pb P
w
Eggs per female per day lxmx= lx 18.2
x=a x=a
Pb Pb
Mean age fecundity schedule xmx= mx 21.6
x=a x=a

Egg hatch (hx) 0.76

Fertility b
P
Gross fertility mxhx 678.7
x=a
Pb
Net fertility lxmxhx 608.1
x=a
Pb P
w
Fertile eggs per female per day lxmxhx= lx 13.9
x=a x=a
Pb b
P
Mean age fertility schedule xmxhx= mxhx 14.9
x=a x=a

a = Age at start of reproduction; b = age at end of reproduction; x = age interval in days;

w = maximum age; lx = proportion of individuals surviving to age x; total eggs laid by
average female at age x; hx = proportion of eggs hatching for those eggs laid at age x.

Papadopoulos et al., 2002). These variations between sexes
have largely been attributed to high reproductive cost in
females compared to males, and to hormonal as well as other
behavioural and physiological differences (Vargas & Carey,
1989; Carey et al., 1995). Life time gross fecundity was 1056.8
eggs whereas net fecundity was 794.6 eggs, with an average
daily egg production of 18.2 eggs. However, due to a com-
bination of adult mortality and an egg viability of only 76%,
net fertility was 608.1. In B. dorsalis and B. cucurbitae gross
fecundity has been estimated to range from 380 to 1428 eggs
at temperatures within the range tested here (Vargas et al.,
1984, 1997; Carey et al., 1985; Yang et al., 1994).
A survivorship curve for both sexes is presented in fig. 2.
Survival for both sexes was high (> 80%) until day 16 and
gradually declined between days 25 to 60, but at a much
higher rate for females beyond day 60. More males survived
beyond day 16 compared to females. Daily egg production
peaked between days 8–25 and thereafter decreased asymp-
totically (fig. 3). Percentage egg hatch was high (> 70%)
between days 8–22 and then decreased, fluctuating widely,
from 30–66% between days 25–53 (fig. 3).
Life table statistics
Life table parameters of B. invadens are presented in
table 5. The intrinsic rate of increase was 0.113 with a net
reproductive rate of 273, indicating a daily increase of 11%
and 273-fold increase from one generation to another. The
mean generation time was 31 days and the population was

1
100
Eggs per female per day

0.9
0.8 90
80
Survival (lx)

0.7
% egg hatch

0.6
0.5 60 60
0.4
0.3 40
30
0.2
0.1 20
0 0
0
0
10
20
30
40
50
60
70
80
90
0
0
13 0
0
0
0
0
0
0

0 10 20 30 40 50 65 75
10
11
12

14
15
16
17
18

Age (days) Age (days)

Fig. 2. Age-specific adult ( , female; —, male) survival of Fig. 3. Age-specific oviposition rate ( ) and egg hatch (—) of
Bactrocera invadens reared at 28+1 C. Bactrocera invadens reared at 28+1 C.
384 S. Ekesi et al.

Table 5. Population parameters of Bactrocera invadens reared on
artificial diet at 28+1 C.
General trait/parameter Formula Value
Rates b
P the true situation in the field. Furthermore, high rates of
Net reproductive rate (Ro) lxmx 273.0
x=a
Intrinsic rate of increase (r) ln(Ro)/T 0.113
Finite rate of increase (l) er 1.120
P
w the data presented here should provide some baseline in-
Intrinsic rate of birth (b) 1= Lx exrx 0.117
x=0
Intrinsic rate of death (d) bxr 0.230
Times
Doubling time (DT) ln(2)/r 6.16
b b
P P
Mean generation time xlxmx= lxmx 30.7
x=a x=a
a = Age at start of reproduction; b = age at end of reproduction;
w = maximum age; Lx = fraction per capita lived between age x
and x+1; mx = hxmx/2, number of female progeny produced by
an average female at age x.
estimated to double in 6 days. These values are close to
those reported by Vargas et al. (1984) for B. dorsalis and
B. cucurbitae but considerably lower than those reported by
Yang et al. (1994) for a Chinese strain of B. dorsalis. The
intrinsic rate of increase has been suggested as an important
index for identification of tephritid fruit fly populations with
good biological attributes for mass rearing and to develop
population growth models (Rössler, 1975; Ichinose &
Nakasone, 1979; Vargas et al., 1984). Values calculated here
for B. invadens may be useful both for mass rearing purposes
and predicting population trends.
Conclusions
The data presented here provide new information
regarding the level of damage by B. invadens to mangoes
and detail the fly’s development on artificial diet. The high
level of infestation recorded at low elevations is an
indication that B. invadens may be well adapted to a hot
climate and thus represents a real threat to mango crops
grown in the warmer low elevation regions of Kenya. Most
studies have demonstrated that fruit fly species from the
genus Bactrocera are adapted to low elevations (Bess, 1953;
Haramoto & Bess, 1970; Vargas et al., 1983; Wong et al., 1985).
However, B. invadens is currently in the process of spreading
and the limit of its altitudinal distribution is probably not yet
known. Studies on the distribution and abundance of this
pest as it relates to different host plants and temperature is
therefore warranted.
Whilst the present results add to our understanding of
life history parameters of the new pest and may be useful in
the development of population models and formulation of
management strategies, certain caveats exists that require
attention. In most fruit fly species, reproductive and life
history traits of population parameters are known to vary
among different geographic areas (Vargas & Carey, 1989).
Additional studies on the demographic parameters of
B. invadens from different geographic areas within the
country are therefore warranted. It must also be recognized
that the culture of B. invadens used in this study was
laboratory-reared, and certain aspects of the biology of
laboratory-adapted flies will obviously differ from those of
flies in the wild. A high protein level in the larval diet
obviously results in shorter developmental time and larger
sizes of individuals (Kaspi et al., 2002) and may not reflect
population increase and shorter mean generation time have
been reported for laboratory-adapted compared to wild
populations of tephritids (Vargas & Carey, 1989). Such
differences probably exist in B. invadens. Overall, however,
formation for development of control measures and perhaps
mass rearing procedures for this pest, which has recently
been rated by the Inter-African Phytosanitary Council as
‘a devastating quarantine pest’ (French, 2005). Technically,
eradication of fruit flies that respond to methyl eugenol such
as B. invadens is feasible using male annihilation techniques
and this has been achieved in other tropical regions of the
world (Tan, 2000). Such actions, however, require rigorous
operational standards and huge investments, even if under-
taken within hours after pest introduction and detection. The
scale of the already known distribution of B. invadens in
Africa, the fragmented structure of farms, poor quarantine
services, vast reservoir and lack of state or donor resources
makes eradication a difficult if not an impossible task.
However, the areas with low pest prevalence identified in
this study might benefit from a simple community-based
quarantine system such as the restriction of uncontrolled
movements of fruits.
Although procedures for mass rearing of B. invadens will
be reported elsewhere, generally the rearing diet used in this
study was adapted from Hooper (1987) by substituting
bran with carrot. This diet can be easily utilized in any mass
rearing facility worldwide, and thus large numbers of this
insect can be produced on a routine basis as with other
Bactrocera species such as B. dorsalis, B. cucurbitae and
B. latifrons (Vargas et al., 1993). In on-going studies we are
evaluating the host range of B. invadens, seasonal distribu-
tion, its temperature tolerance and response to food
attractants as some of the major steps towards developing
effective management tactics. Field explorations are also
underway in Sri Lanka for natural enemies that might be
useful in suppressing the pest.
Acknowledgements
Thanks are due to Drs F. Schulthess and N.K. Maniania
(International Centre of Insect Physiology and Ecology) for
valuable comments on earlier draft of the manuscript. The
authors appreciate the assistance of E. Mlato, J. Kiilu and
P. Agola in collection of data. Financial support for the
research came from the International Fund for Agricultural
Development and the International Atomic Energy Agency.
References
Aluja, M., Celedonio-Hurtado, H., Liedo, P., Cabrera, M.,
Castillo, F., Guillen, J. & Rios, E. (1996) Seasonal popu-
lation fluctuations and ecological implications for manage-
ment of Anastrepha fruit flies (Diptera: Tephritidae) in
commercial mango orchards in southern Mexico. Journal of
Economic Entomology 89, 654–667.
Anon. (1999) Areas in Mexico free from fruit flies (ALMF, 8/96).
30 pp. Bilingual Document, English–Spanish, Support
 Field infestation and life history of Bactrocera invadens
document to the Quarantine Bilateral Agreement between
MAF New Zealand and SAGAR, Mexico.
Anon. (2000) A submission supporting area freedom from Queensland
fruit fly and Mediterranean fruit fly for the Riverland, Sunraysia
and Riverrina pest free areas of mainland Australia. 32 pp.
Department of Agriculture, Fisheries and Forestry, Austra-
lia.
Bess, H.A. (1953) Status of Ceratitis capitata in Hawaii following
the introduction of Dacus dorsalis and its parasites. Proceed-
ings of the Hawaiian Entomological Society 15, 221–234.
Carey, J.R. (1993) Applied demography for biologists. 206 pp. New
York, Oxford University Press.
Carey, J.R., Harris, E.J. & McInnis, D.O. (1985) Demography of
a native strain of the melon fly, Dacus cucurbitae, from
Hawaii. Entomologia Experimentalis et Applicata 38, 195–199.
Carey, J.R., Liedo, P., Orozco, D., Tatar, M. & Vaupel, J.W.
(1995) A male–female longevity paradox in medfly cohorts.
Journal of Animal Ecology 64, 107–116.
Clarke, A.R., Armstrong, K.F., Carmichael, A.E., Milne, J.R.,
Raghu, S., Roderick, G.K. & Yeates, D.K. (2005) Invasive
phytophagous pests arising through a recent evolutionary
radiation: the Bactrocera dorsalis complex of fruit flies.
Annual Review of Entomology 50, 293–319.
Copeland, R.S., Wharton, R.A., Luke, Q. & de Meyer, M. (2002)
Indigenous hosts of Ceratitis capitata (Diptera: Tephritidae)
in Kenya. Annals of the Entomological Society of America 95,
672–694.
Dimbi, S. (2003) Evaluation of the potential of Hyphomycetes
fungi for the management of the African tephritid fruit flies
Ceratitis capitata (Weidemann), Ceratitis cosyra (Walker) and
Ceratitis fasciventris (Bezzi) in Kenya. 146 pp. PhD thesis,
Kenyatta University, Nairobi.
Drew, R.A.I. & Hancock, D.L. (1994) The Bactrocera dorsalis
complex of fruit flies (Diptera; Tephritidae: Dacinae) in
Asia. Bulletin of Entomological Research Supplement Series, 2,
1–68.
Drew, R.A.I., Tsuruta, K. & White, I.M. (2005) A new species of
pest fruit fly (Diptera: Tephritidae: Dacinae) from Sri Lanka
and Africa. African Entomology 13, 149–154.
Ekesi, S., Maniania, N.K., Mohamed, S.A. & Lux, S.A. (2005)
Effect of soil application of different formulations of
Metarhizium anisopliae on African tephritid fruit flies and
their associated endoparasitoids. Biological Control 35, 83–
91.
FAO (1999) Requirements for the establishment of pest free places of
production and pest free production sites. 34 pp. ISPM no. 10,
Food and Agriculture Organization, Rome.
French, C. (2005) The new invasive Bactrocera species. pp. 19–20
in Insect Pest Control Newsletter, No. 65. International Atomic
Energy Agency, Vienna, Austria.
Haramoto, F.H. & Bess, H.A. (1970) Recent studies on the
abundance of the oriental and Mediterranean fruit flies and
the status of their parasites. Proceedings of the Hawaiian
Entomological Society 20, 551–556.
Harris, E.J., Takara, J.M. & Nishida, T. (1986) Distribution of the
melon fly Dacus cucurbitae (Diptera: Tephritidae) and host
plants on Kauai, Hawaiian Islands. Environmental Entomol-
ogy 15, 488–493.
HCDA (2004) Progress report on good agricultural practices,
MOA/HCDA/JICA-training team. Horticultural Crops
Development Authority Horticultural News 30, 6–9.
Hooper, G.H.S. (1987) Application of quality control procedures
for large scale rearing of the Mediterranean fruit fly.
Entomologia Experimentalis et Applicata 44, 161–167.
385
Ichinose, F. & Nakasone, S. (1979) Intrinsic rate of increase
of laboratory populations of Dacus cucurbitae Coquillett
and D. dorsalis Hendel (Diptera: Tephritidae). Japanese
Journal of Applied Entomology and Zoology 23, 42–43.
Kaspi, R., Mossinson, S., Drezner, T., Kamensy, B. & Yuval, B.
(2002) Effects of larval diet on developmental rates and
reproductive maturation of male and female Mediterranean
fruit flies. Physiological Entomology 27, 29–38.
Lawson, A.E., McGuire, D.J., Yeates, D.K., Drew, R.A.I. &
Clarke, A.R. (2003) Dorsalis: an interactive identification tool
to fruit flies of the Bactrocera dorsalis complex. CD-ROM
Publication, Griffith University, Brisbane, Australia.
Levins, R. & Wilson, M. (1980) Ecological theory and pest
management. Annual Review of Entomology 25, 287–308.
Liedo, P., Carey, J.R., Celedonio, H. & Guillen, J. (1992)
Size specific demography of three species of Anastrepha
fruit flies. Entomologia Experimentalis et Applicata 63,
135–142.
Liquido, N.J., Harris, E.J. & Dekker, L.A. (1994) Ecology of
Bactrocera latifrons (Diptera, Tephritidae) populations: host
plants, natural enemies, distribution, and abundance.
Annals of the Entomological Society of America 87, 71–84.
Lux, S.A., Zenz, N. & Kimani, S. (1999) Economic role and
distribution of fruit flies. pp. 78–79. ICIPE Annual Scientific
Report, 1998/199.
Lux, S.A., Copeland, R.S., White, I.M., Manrakhan, A. &
Billah, M.K. (2003) A new invasive fruit fly species from
the Bactrocera dorsalis (Hendel) group detected in East
Africa. Insect Science and Its Application 23, 355–360.
MacArthur, R.H. & Wilson, E.O. (1967) The theory of island
biogeography. 215 pp. Princeton, Princeton University Press.
McPeek, M.A. & Kalisz, S. (1993) Population sampling and
bootstrapping in complex designs: demographic analysis,
pp. 273–289 in Scheiner, S.M. & Gurevtch, J. (Eds) Design
and analysis of ecological experiments. New York, Chapman
& Hall.
Mwatawala, M.W., White, I.M., Maerere, A.P., Senkondo, F.J.
& de Meyer, M. (2004) A new invasive Bactrocera species
(Diptera: Tephritidae) in Tanzania. African Entomology 12,
154–156.
Nishida, T.E., Harris, E.J. & Vargas, R.I. (1980) Life system
components in relation to fruit fly eradication strategies in
Hawaii. pp. 133–142 in Proceedings of the Symposium on Fruit
Fly Problems. Kyoto and Naha, August 9–12, 1980.
Papadopoulos, N.T., Katsoyannos, B.I. & Carey, J.R. (2002)
Demographic parameters of the Mediterranean fruit
fly (Diptera: Tephritidae) reared in apples. Annals of the
Entomological Society of America 95, 564–569.
Peck, S.L. & McQuate, G.T. (2004) Ecological aspects of
Bactrocera latifrons (Diptera: Tephritidae) on Maui, Hawaii:
movement and host preference. Environmental Entomology
33, 1722–1731.
Rice, R.E., Phillips, P.A., Stewart-Leslie, J. & Sibbett, G.S.
(2003) Olive fruit fly populations measured in central and
southern California. California Agriculture 57, 122–127.
Rössler, Y. (1975) Reproductive differences between laboratory-
reared and field-collected populations of the Mediterranean
fruit fly Ceratitis capitata. Annals of the Entomological Society
of America 68, 987–991.
SAS Institute (1985) SAS User’s Guide: statistics, version 5. SAS
Institute, Carey, North Carolina.
Simpson, S.E. (1993) Caribbean fruit fly-free zone certification
protocol in Florida (Diptera: Tephritidae). Florida Entomol-
ogist 76, 228–232.
386 S. Ekesi et al.
Tan, K.-H. (2000) Area-wide control of fruit flies and other insect
pests. 782 pp. Penerbit Universiti Sains Malaysia, Pulau
Pinang.
Vargas, R.I. & Carey, J.R. (1989) Comparison of demographic
parameters for wild and laboratory reared tephritids
(Diptera: Tephritidae). Annals of the Entomological Society
of America 82, 55–59.
Vargas, R.I., Nishida, T. & Beardsley, J.W. (1983) Distribution
and abundance of Dacus dorsalis (Diptera: Tephritidae)
in native and exotic forest areas of Kauai. Environmental
Entomology 12, 1185–1189.
Vargas, R.I., Miyashi, D. & Nishida, T. (1984) Life history and
demographic parameters of three laboratory reared tephri-
tids (Diptera: Tephritidae). Annals of the Entomological
Society of America 77, 651–656.
Vargas, R.I., Mitchell, S., Hsu, C.-L. & Walsh, W.A. (1993)
Evaluation of mass-rearing procedures for Bactrocera
latifrons (Diptera: Tephritidae). Journal of Economic Entomol-
ogy 86, 1157–1161.
Vargas, R.I., Walsh, W.A. & Nishida, T. (1995) Colonization of
newly planted coffee fields: dominance of Mediterranean
fruit fly over oriental fruit fly (Diptera: Tephritidae). Journal
of Economic Entomology 88, 620–627.
Vargas, R.I., Walsh, W.A., Jang, E.B., Armstrong, J.W. &
Kanehisa, D.T. (1996) Survival and development of
immature stages of four Hawaiian fruit flies (Diptera:
Tephritidae) at five constant temperatures. Annals of the
Entomological Society of America 89, 64–69.
Vargas, R.I., Walsh, W.A., Kanehisa, D.T., Jang, E.B. &
Armstrong, J.W. (1997) Demography of four Hawaiian
fruit flies (Diptera: Tephritidae) reared at five constant
temperatures. Annals of the Entomological Society of America
90, 162–168.
Vargas, R.I., Walsh, W.A., Kenehisa, D., Stark, J.D. & Nishida,
T. (2000) Comparative demography of three Hawaiian fruit
flies (Diptera: Tephritidae) at alternating temperatures.
Annals of the Entomological Society of America 93, 75–81.
White, I.M. & Elson-Harris, M.M. (1992) Fruit flies of economic
importance: their identification and bionomics. 601 pp. Wall-
ingford, Oxon, CAB International.
Wong, T.M.Y., McInnis, D.O. & Mochizuki, N. (1985) Seasonal
distribution and abundance of adult male oriental fruit flies
(Diptera: Tephritidae) in Kula, Maui, Hawaii. Journal of
Economic Entomology 78, 1267–1271.
Yang, P., Carey, J.R. & Dowell, R.V. (1994) Temperature
influences on the development and demography of
Bactrocera dorsalis (Diptera: Tephritidae) in China. Environ-
mental Entomology 23, 971–974.
(Accepted 3 May 2006)
Ó CAB International, 2006