J. Appl. Entomol.

ORIGINAL CONTRIBUTION

Demographic parameters of the two main fruit fly (Diptera:

Tephritidae) species attacking mango in Central Tanzania
J. K. Salum1*, M. W. Mwatawala1, P. M. Kusolwa1 & M. D. Meyer2
1 Department of Crop Sciences and Production, Sokoine University of Agriculture, Morogoro, Tanzania
2 Entomology Section, Royal Museum for Central Africa, Tervuren, Belgium

Keywords
Africa, development, invasive species
Correspondence
M. D. Meyer (corresponding author), Royal
Museum for Central Africa, Entomology
Section, Leuvensesteenweg 13, B3080
Tervuren, Belgium. E-mail:
marc.de.meyer@africamuseum.be
*Present address: District Agriculture and
Livestock Development Office, Manyoni
District, Singida, Tanzania.
Received: July 12, 2012; accepted: February 5,
2013.
doi: 10.1111/jen.12044
Abstract
Fruit flies (Diptera: Tephritidae) are among the most important pests
damaging fruit crops growing in Tanzania. Demographic parameters for
Ceratitis cosyra (Walker) and Bactrocera invadens (Drew, Tsuruta & White)
are reported and compared. These include pre-adult survival and develop-
ment rates, adult survival and fecundity and population parameters such
as the intrinsic rate of increase, and mean generation time. Population
and biological parameters of the C. cosyra and B. invadens were measured
in an environmental chamber at 25°C and 30°C with 75% RH and at
room temperature ranging 23–28°C. The results showed that B. invadens
had shorter embryonic, larval and pupal development times than C. cosyra
independent of temperature. Life expectancy of male B. invadens was
significantly greater than that of C. cosyra, while life expectancy of female
B. invadens was greater than that of C. cosyra but not significantly. Average
net fecundity was higher for B. invadens than for C. cosyra. Both species
attained their highest intrinsic rate of increase and net reproductive rate
at 30°C. B. invadens exhibited higher intrinsic rate of increase and net
reproductive rate than C. cosyra at all temperatures tested.

in July–September 2008. The most challenging task is

Introduction
Fruit and vegetable production is one of the fastest-
growing agricultural sectors in Africa (Ekesi and
Billah 2005) and is important as a source of nutrients
to supplement daily dietary requirements. Further-
more, fruit production provides a source of income
and employment to many growers and traders.
Tropical and subtropical regions are becoming
important areas for fruit production, and there is a
great potential for export of fruits from these regions
to international markets if export qualities can be
met. For example, Vayssi eres et al. (2008) reported
that export of mango from Benin to Europe has multi-
plied fivefold over the past fourteen years, increasing
from 42 000 tons in 1992 to over 210 000 tons in
2006. BOT (2008) recorded substantial increases in
non-traditional exports from Tanzania, including hor-
ticultural products, which went up to USD 4.6 mil-
lions from July–September 2007 to USD 8.3 millions
to export pest-free fruits. Fruit flies (Diptera: Tephriti-
dae) have been listed among the major pests of many
fruits (CABI 2007). They are reported to be present in
all fruit growing regions of the world, causing losses
as high as USD 910 millions (White and Elson-Harris
1994). The main losses that fruit flies cause are
through direct damage to fruits and fruit vegetables.
Vayssi eres et al. (2008) reported that, in tropical
regions, mangoes are attacked by fruit flies (Diptera
Tephritidae), which inflict great economic devastation
in both East Africa and West Africa. In Tanzania, a
number of fruit flies have been recorded (Mwatawala
et al. 2009a), and in the last decade, two species of
fruit flies were introduced into the country: Bactrocera
invadens Drew Tsuruta & White, which was detected
in 2003 (Mwatawala et al. 2004), and Bactrocera
latifrons (Hendel), which was detected in 2006
(Mwatawala et al. 2007). Such introductions give
raise to interspecific competition, resulting in

© 2013 Blackwell Verlag, GmbH 1

Demography of Bactrocera invadens and Ceratitis cosyra
competitive displacement (Ekesi et al. 2009).
Mwatawala et al. (2009a) reported that B. invadens
dominates the native mango fruit fly (Ceratitis cosyra
(Walker)), a predominant pest of mangoes, in its
fundamental niche. The same pattern was observed in
the study Kenya by Ekesi et al. (2009), while
Rwomushana et al. (2009) showed significant effects
of temperature on competitive success of B. invadens
compared with C. cosyra. The information by
Mwatawala et al. (2009a) is based on trap catches’
data in the field as well as incidence and infestation
rates on fruits. The study did not study possible
changes in life-history strategies, such as development
rates of immature stages (i.e. embryonic developmen-
tal time, larval and pupal development), adult longev-
ity, fecundity, mortality, pre-oviposition period, net
reproductive rate, mean generation time and doubling
time. These parameters have been studied for several
major fruit fly pests in other parts of the world. Specif-
ically for Africa, Duyck and Quilici (2002) looked at
the survival and development of the Mediterranean
fruit fly, Ceratitis capitata (Wiedemann), the Natal fruit
fly, C. rosa Karsch and the Mascarenes fruit fly, C. cato-
irii Guerin-M eneville and compared them at five con-
stant temperatures spanning from 15 to 35°C. Ekesi
et al. (2006) presented the comparative developmen-
tal biology of B. invadens maintained at one fixed
temperature (28
1°C), while Rwomushana et al.
(2008) compared development and survival at differ-
ent temperatures for this particular species. A compar-
ative study, conducted at different temperatures, for
the predominant mango pests in Africa is lacking, and
the current study aims to complement the data by
Mwatawala et al. (2009b) and the other studies
mentioned above. The main objective of this study
therefore is to determine the life-history strategies
related to competition among the two main fruit fly
species (Diptera: Tephritidae) attacking mango in
Central Tanzania: the invasive B. invadens and the
indigenous C. cosyra.
Material and Methods
Establishment of fruit fly colonies
The studies were conducted under laboratory condi-
tions at the Sokoine Unversity of Agriculture (SUA),
situated in Morogoro (Central Tanzania) at about
520 m altitude. Colonies of adult fruit flies were
established from infested Mangifera indica L., Annona
muricata L., Psidium guajava L. and Terminalia catappa
L., collected from the Horticulture Unit at SUA (see
Mwatawala et al. 2006 for detailed description of the
2 © 2013 Blackwell Verlag, GmbH
J. K. Salum et al.
unit) and Morogoro town centre. Adults were fed
simultaneously with enzymatic yeast hydrolysate
(ICN Biomedical, Irvine, CA) and sucrose in a ratio of
1 : 3 and water on pumice granules in different plastic
containers for each species. Fruit fly rearing protocols
established by Vayssieres et al. (2008) were followed.
The colonies of B. invadens were maintained on artifi-
cial diet for ten generations while those of C. cosyra
were maintained for eight generations.
Embryonic developmental time
Eggs of B. invadens and C. cosyra were collected from
mangoes. The duration of the egg stages was
established by, for each experiment, dissecting 100
eggs out of fruits, removing them with a camel hair
brush and placing them on a moist filter paper in a
Petri dish at 25 and 30°C with 75% relative humidity
maintained in an environmental growth chamber.
Eggs were placed in the environmental growth cham-
ber for 20 h, and then egg hatch was determined
every 12 h using a microscope. A separate experiment
(also using 100 eggs) was set up in an open rearing
room of the horticulture unit, at room temperature
ranging from 23 to 28°C. This range corresponds to
the average temperature in the region at this altitude
during mango fruit maturation period (Geurts et al.
2012), and the purpose was to allow comparison
with natural conditions. Three replications were
maintained.
Larval and pupal development
Fifty newly emerged larvae from the above described
eggs were gently introduced with a camel hair brush
onto the 50 g of specific artificial larval diet, as
described by Ekesi et al. (2007) and adopted by Dimbi
et al. (2009), in a Petri dish. Changes in larval stages
were observed twice per day. The time needed for
50% of individuals to develop to a particular stage
was noted at the 25 and at 30°C with 75% relative
humidity and at room temperature ranging from 23
to 28°C. Larval instars were differentiated by their
size, body surface sculpturing, mouth-hook morphol-
ogy and colour according to White and Elson-Harris
(1994), but results indicate the duration for the whole
larval development.
By the end of the third larval instars, mature
larvae were allowed to leave the Petri dish to pupate
in the sand in a plastic container. Pupae were col-
lected by sifting the sand, counted and carefully
cleaned. They were transferred to a plastic container
covered with fine mesh and containing a small moist
J. K. Salum et al.
sponge. At the end of the pupal stage, the number
of newly emerged adults was recorded twice a day.
Effect of temperature (subfactor) and species (main
factor) on larval and pupal stage duration (days)
were tested using the two-way analysis of variance
(ANOVA). A cohort of a life stage of each fruit fly
species formed a replicate. The least significance
difference test was used to compare differences
between means. All analyses were performed using
the GenStat statistical software (VSN International,
Hemel Hempstead, UK).
Adult demographic parameters
Adult life-history traits were assessed using 15 pairs
of newly emerged adults of each species isolated in
individual plastic container covered with fine mesh
and placed at 25, 30 and 23–28°C for each fly spe-
cies. Adults were fed with enzymatic yeast hydroly-
sate (ICN Biomedicals) and sugar in a ratio of 1 : 3.
A single ball of wrapped artificial larval diet was
placed in a plastic container for B. invadens and
mango (single fruit) for C. cosyra daily to monitor
egg laying (larval diet is based on protocols and
recommendations of the African Fruit Fly Initiative,
as developed by International Centre for Insect
Physiology and Ecology, S. Lux personal communi-
cation, see Ekesi et al. 2007). The wrapping of the
diet and skin of the fruit were punctured using an
entomological pin to facilitate oviposition. Eggs on
the wrapping of the diet and skin of the fruit were
gently removed as previously described and
counted by using binocular microscope, and daily
mortality of flies was recorded. Each treatment was
2.5
2
Days ± SE
1.5
0.5
0
B. invadens
25
Fig. 1 Mean duration of egg development for Bactrocera invadens and Ceratitis cosyra at three temperature regimes.
© 2013 Blackwell Verlag, GmbH
Demography of Bactrocera invadens and Ceratitis cosyra
replicated three times. The same analytical methods
as for larval and pupal development were used.
Standard life table parameters among the different
species and treatments were calculated from daily
records of mortality of immature stages (through
observation under microscope), fecundity and fertil-
ity of pairs of adults. Parameters symbols, formulae
and definition follow Carey (1993).
Results
Embryonic developmental time
The egg stage duration varied significantly between
species (F = 22.40; df = 1; P = 0.001) and among
temperature regime (F = 33.76; df = 2; P = 0.001)
(fig. 1), being shorter for both species at 30°C. Bactro-
cera invadens exhibited an egg development period
shorter than that of C. cosyra for all temperature
regimes tested.
Larval and pupal development
The larval duration (fig. 2a) varied significantly
between species (F = 31.18; df = 1; P = 0.001) and
among temperature regime (F = 30.62; df = 2; P =
0.001). The pupal duration (fig. 2b) also varied signifi-
cantly with species (F = 21.51; df = 1; P = 0.001) and
temperature (F = 69.87; df = 2; P = 0.001). The same
tendencies as in egg stage duration were observed with
the larval and pupal development time of B. invadens
being shorter than those of C. cosyra for all temperature
tested and the average developmental time for both
species being shortest at 30°C.
C. cosyra B. invadens C. cosyra B. invadens C. cosyra
25 30 30 23-28 23-28
Temperature °C
3
Demography of Bactrocera invadens and Ceratitis cosyra J. K. Salum et al.

(a) 18
16
14
12

Days ± SE
10
8
6
4
2
0
B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra

25 25 30 30 23-28 23-28
Temperature °C

(b) 18

16

14

12
Days ± SE

10

0
B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra

25 25 30 30 23-28 23-28
Temperature °C

Fig. 2 Mean developmental time (days) of (a) larval and (b) pupal development for Bactrocera invadens and Ceratitis cosyra at three temperature
regimes.

for B. invadens and 316.4 for C. cosyra, while at 23–28°

Pre-oviposition period
Pre-oviposition period varied significantly between
temperatures (F = 26.61; df = 2; P = 0.001) (fig. 3),
while no significant differences were observed for
pre-oviposition period (in days) between the two fruit
fly species (P > 0.05) in all the experimental set-ups.
Gross and net fecundity
The average gross fecundity (theoretical natality rate
during lifetime of female) at 25°C was 577.6 eggs for
B. invadens and 499.1 eggs C. cosyra. At 30°C, it was,
respectively, 969.6 and 768.4 eggs, while at 23–28°C
it was 745.2 and 670.0 eggs. The average net fecun-
dity rate of B. invadens at 25°C was 371.8 eggs, while
it was 294.5 eggs for C. cosyra. At 30°C, it was 544.5
C, it was, respectively, 516.4 and 457.3.
Daily egg production per female per day
Figure 4 presents eggs production per female per day
which varied significantly among species (F = 27.37;
df = 1; P = 0.001) and temperature (F = 48.46;
df = 2; P = 0.001). Age-specific reproduction data for
the two species at the temperature regimes of 25 and
30°C are given in Appendices S1–S4.
Longevity
Life expectancy at adult emergence varied
significantly between species for males (F = 51.20;
df = 1; P = 0.001) but not significantly between

4 © 2013 Blackwell Verlag, GmbH

J. K. Salum et al. Demography of Bactrocera invadens and Ceratitis cosyra

14

12

10

Days ± SE
8

0
B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra
25 25 30 30 23-28 23-28
Temperature °C

Fig. 3 Mean pre-oviposition period (in days) for Bactrocera invadens and Ceratitis cosyra reared at three temperature regimes.

14

12
Mean number of eggs ± SE

10

0
B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra
25 25 30 30 23-28 23-28
Temperature °C

Fig. 4 The daily egg production of Bactrocera invadens and Ceratitis cosyra at three temperature regimes.

Table 1 Survival rate (%) of Bactrocera invadens and Ceratitis cosyra reared on artificial diet at three temperature regimes

25°C 30°C 23–30°C

Developmental
stage B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra

Eggs 82.7 85.0 90.7 92.3 88.7 84.7

Larvae 77.3 77.3 81.3 75.3 76.7 76.0
Puparia 50.6 43.2 64.3 72.6 61.2 52.5
Egg to adult 38.7 33.3 52 54.7 46.7 40.0

temperatures (F = 15.77; df = 2; P = 0.001). In
females, life expectancy at adult emergence was
higher for B. invadens than for C. cosyra but was not
significantly different. Bactrocera invadens also had a
slightly higher survivorship from egg to adult com-
pared with C. cosyra at some temperature regimes
(Table 1). This is reflected in net reproductive rate
being higher and expectation of life being greater for
B. invadens at all temperatures tested. Age-specific
mortality rates for the two species at the temperature
regimes of 25 and 30°C are given in Appendices
S5–S8.

© 2013 Blackwell Verlag, GmbH 5

Demography of Bactrocera invadens and Ceratitis cosyra J. K. Salum et al.

Table 2 Population parameters of Bactrocera invadens and Ceratitis cosyra reared on artificial diet at three temperature regimes

25°C 30°C 23–28°C

General trait/parameters
Rates Formula B. invadens C. cosyra B. invadens C. cosyra B. invadens C. cosyra

P
b
Net reproductive rate (R0) lxmx 30.5 96.6 196.1 107.5 199.1 164.9
x¼a
Intrinsic rate of increase (r) Ln(Ro)/T 0.128 0.109 0.174 0.154 0.159 0.159
Finite rate of increase (k) er 1.135 1.116 1.19 1.16 1.17 1.15
Times
Doubling time (DT) Ln(2)/r 6.3 5.4 3.98 4.51 4.37 4.97
Pb P
b
Mean generation time (T) xlxmx= lxmx 38.08 41.77 30.34 30.4 33.3 36.6
x¼a x¼a

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 eggs laid by average female at age x.

maximum temperature (i.e. 35°C) for which no

Population parameters
Population parameters of B. invadens and C. cosyra are
summarized in Table 2. The intrinsic rate of increase
varied significantly with temperature (F = 97.99;
df = 2; P = 0.001) and species (F = 59.42; df = 1;
P = 0.001). Both species exhibited their highest
intrinsic rate of increase at 30°C and their lower at 25°
C with B. invadens showing the highest rate at all
temperatures.
The net reproductive rate (R0) represents the mean
number of female offspring produced by a single
female from a cohort during the course of her life
span (Papadopoulos et al. 2002). In the current study,
the net reproductive rate varied significantly with spe-
cies (F = 22.01; df = 1; P = 0.001) and temperature
(F = 14.94; df = 2; P = 0.001). At 25°C, the average
net reproductive rate was lower for both species than
at 30°C. Also, the doubling time (DT) varied signifi-
cantly with temperature (F = 109.18; df = 2;
P = 0.001) and species (F = 58.86; df = 1; P = 0.001)
with lowest average at 30°C. The mean generation
times varied significantly with temperatures, but were
not significantly different between the two species.
Discussion
It is known that temperature exerts strong effects on
reproductive parameters (Vargas et al. 1997). In our
study, the duration of all development stages (embry-
onic, larval and pupal development) is temperature
dependent for both species, as was observed in other
fruit fly studies (see for example Vargas et al. 2000;
Duyck and Quilici 2002; Vayssi eres et al. 2008; Liu
and Ye 2009). Our study confirms the pattern that an
increase in temperature results in a faster develop-
ment. Duyck and Quilici (2002), however, also
indicated that mortality can increase and at certain
development was observed for Ceratitis capitata, C. cato-
irii and C. rosa. This maximum threshold was not
studied here because our main concern was to com-
pare the developmental rate between B. invadens and
C. cosyra. The experiments have shown that at all
stages, B. invadens develops faster than C. cosyra, for
all temperature regimes tested. Pre-oviposition period
was also lowest at 30°C, but no significant difference
could be found between the two species studied. The
developmental times at 25 and 30°C for B. invadens
are in general longer than those observed by Rwo-
mushana et al. (2008). The developmental duration
of C. cosyra was never studied. Our findings show a
remarkably longer developmental time than for other
Ceratitis species studied by Duyck and Quilici (2002).
Results regarding fecundity and longevity are some-
what lower than those observed by Ekesi et al. (2006)
for B. invadens. The cause of this is not clear, but
experimental set-up (differences in egging device,
rearing diet, etc.) cannot be excluded. In general, the
results are, however, comparable. The gross fecundity
for C. cosyra (499.1 eggs) is lower, but in a similar
range, than that observed for the main fruit fly pest C.
capitata (Vargas et al. 2000: 562.4 eggs) at a compara-
ble temperature (25°C). All other population parame-
ters confirm the above tendencies with higher
performance at higher temperature regime and
B. invadens scoring significantly higher than C. cosyra
for the majority of the parameters.
Our results therefore show that B. invadens has a
demographic advantage over the main indigenous
mango pest, C. cosyra. The immature growth stages for
the former are shorter, and it scores higher on all
adult parameters tested. Bactrocera invadens also
appears to have a broader host range (Mwatawala
et al. 2009b) and hence more alternative hosts to
build up populations throughout the year than

6 © 2013 Blackwell Verlag, GmbH

J. K. Salum et al.
C. cosyra does. Both aspects, in addition to factors such
as behaviour (see Duyck et al. 2006b) which have not
been studied, may explain the fact that B. invadens has
become the predominant infester of mangoes in Cen-
tral Tanzania and has led to outcompeting C. cosyra.
This is in congruence with the findings of Ekesi et al.
(2009) where it was shown that B. invadens is displac-
ing C. cosyra in certain parts of Kenya, with tempera-
ture being one of the mechanisms contributing to the
competitive success of B. invadens (Rwomushana
et al. 2009). Duyck et al. (2006a) demonstrated that
often climatic niche partitioning is observed after
invasion of exotic fruit flies, but it is not clear whether
that is the case here. Geurts et al. (2012) recently
looked at spatial and temporal partitioning of
indigenous and exotic fruit flies in Central Tanzania.
Specifically for B. invadens and C. cosyra, both are
predominant in the same agro-ecological zone, but a
temporal shift between both species could be observed
which could be the result of this competition,
although further studies are needed to establish the
exact interspecific interactions.
From a practical point, demographic parameters are
useful in predicting phenological events for ecological
and pest management purposes, and in the construc-
tion of basic fruit fly population growth and survival
models (Fletcher 1989). These in turn can be used for
predictive distribution models such as recently
conducted for the indigenous C. rosa (De Villiers et al.
2013). Such phenological and distributional informa-
tion is essential for the development of an effective
control program. Although development of such a
program was beyond the scope of this study, it is
anticipated that the data presented here will contrib-
ute to the development of such tools.
Acknowledgements
This study was partially financially supported by the
Belgian Development Cooperation (project S1_
TNZ_IPM). We also acknowledge the useful com-
ments provided by Serge Quilici and an anonymous
referee on an earlier version of this article, which
greatly improved the manuscript.
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Supporting Information
Additional Supporting Information may be found in
the online version of this article:
Appendix S1. Population parameters Bactrocera
invadens at 25°C.
Appendix S2. Population parameters of Ceratitis
cosyra at 25°C.
Appendix S3. Population parameters of Ceratitis
cosyra at 30°C.
Appendix S4. Population parameters of Bactrocera
invadens at 30°C.
Appendix S5. Life history parameters of Ceratitis
cosyra at 25°C.
Appendix S6. Life history parameters of Bactrocera
invadens at 25°C.
Appendix S7. Life history parameters of Ceratitis
cosyra at 30°C.
Appendix S8. Life history parameters of Bactrocera
invadens at 30°C.