POPULATION ECOLOGY
Development of Immature Stages of Sesamia nonagrioides
(Lepidoptera : Noctuidae) Under Alternating and
Constant Temperatures
ARGYRO A. FANTINOU,1 DIONYSSIOS Ch. PERDIKIS,2
AND
COSTAS S. CHATZOGLOU1
Environ. Entomol. 32(6): 1337Ð1342 (2003)
ABSTRACT Laboratory studies on the temperature-dependent development of Sesamia nonagrioides (Lefebvre) were performed under Þve constant temperatures ranging from 20 to 30⬚C as well
as under the corresponding alternating temperatures of 25:10, 27.5:12.5, 30:15, 32.5:17.5, and 35:20⬚C,
at a photoperiod of 16:8 (L:D) h. Both linear and nonlinear (Lactin formula) models provided a reliable
Þt of developmental rates versus temperature for all immature stages either at constant or alternating
temperatures. Developmental thresholds that were estimated by a linear model for eggs, larvae, or
pupae were approximately the same and estimated to be 10.57, 8.89, and 10.75⬚C, respectively, at
constant temperatures. Lower values were estimated when immature stages were exposed to the
corresponding alternating temperatures (7.23, 6.20, and 6.50⬚C for eggs, larvae, and pupae). Therefore,
the evaluation of the development of immature stages under constant temperatures resulted in an
overestimation of the lower thresholds. The Lactin-2 model also was applied, and larval and egg
developmental thresholds did not differ from those that were estimated by the linear model. Differences, however, occurred at the pupal stage. Nevertheless, the differences in the estimated values of
the lower developmental thresholds of the immature stages under constant or alternating temperatures indicate that a model predicting phenology in the Þeld should be based on ßuctuating temperature.
KEY WORDS Sesamia nonagrioides, developmental rate, constant temperature, alternating temperatures
THE ROLE OF TEMPERATURE on insect developmental rate
has been well documented, and several approaches
have been used to relate its role to the seasonality of
insect life stages. Among them, the linear model permits estimation of the lower developmental temperature, i.e., the temperature at which the developmental rate approaches zero. A mean number of degreedays is required to complete development of a given
life stage of an insect (Lamb 1992). Nonlinear models
also have been developed that describe development
of insects under variable temperature (Stinner et al.
1975, Sharpe and DeMichele 1977, Hilbert and Logan
1983, Wanger et al. 1984).
Under natural conditions organisms are exposed to
daily ßuctuations of temperature. However, developmental thresholds usually are determined at constant
temperature regimes, and few studies regard the role
of alternating temperatures on insect developmental
rate. However, developmental rates can differ greatly
for insects reared at constant temperature or alternating temperatures (Behrens et al. 1983, Huffaker et
1 Laboratory of Ecology and Environmental Sciences, Agricultural
University of Athens, Iera Odos 75, 188 55 Athens, Greece.
2 Laboratory of Agricultural Zoology and Entomology, Agricultural
University of Athens, Iera Odos 75, 188 55 Athens, Greece.
al. 1984) and the larger the temperature ßuctuation,
the bigger the difference (Messenger 1964, Ratte
1985). This difference, which probably results from
the nonlinearity of development, was Þrst described
by Kaufmann (1932) and has been called the Kaufmann effect (Behrens et al. 1983, Worner 1992). Moreover, Beck (1983) argued that daily temperature
cycles might decrease mortality or increase developmental rate beyond predictions based on constant
temperature studies, when temperatures in the cycling regime exceed the upper or lower thresholds.
The corn borer Sesamia nonagrioides (Lefebvre) is
distributed throughout southern Europe, North Africa, and the Middle East, causing severe damage to
late corn (Melamed-Madjar and Tam 1980, Tsitsipis
1990, Gillyboeuf et al. 1994) and sugarcane (Baniabassi 1981). This insect is multivoltine with three to
four generations and overwinters as a mature larva
(Fantinou et al. 1995). In Greece, the Þrst adults of the
overwintered generation occur from early March to
early May, in mild and cold areas, respectively (Tsitsipis 1990). Many studies have investigated the biology, control, and natural enemies of this species (Tsitsipis 1990; Eizaguirre et al. 1994; Fantinou et al. 1995,
1998, 2003; Lopez et al. 2001). Recently, a phenology
model was developed estimating the developmental
0046-225X/03/1337Ð1342$04.00/0 䉷 2003 Entomological Society of America
1338
ENVIRONMENTAL ENTOMOLOGY
threshold and thermal constant (degree-days) of the
different stages, and predicting the appearance of the
spring generation (Lopez et al. 2001). However, the
developmental rate has not been studied under alternating temperatures, though such data are necessary
to explain the population ßuctuations under natural
conditions. In addition, this knowledge will reduce the
discrepancies between Þeld and laboratory observations, permitting the formulation of more reliable phenological models. The present investigation compares
effects of constant temperatures and corresponding
alternating temperatures on the developmental rates
of S. nonagrioides immature stages (egg, larval, and
pupal stage).
Materials and Methods
A colony of S. nonagrioides, derived from larvae
collected in Kopais (latitude 38⬚14⬘, central Greece) in
1999, which has been maintained for 12 successive
generations in the laboratory, was the source of insects
used in this investigation. Maintenance and handling
of insects have been described in a previous article
(Fantinou et al. 1995). The larvae were kept at 25 ⫾
1⬚C and 55 ⫾ 5% RH under a photoperiod of 16:8 (L:D)
h and were reared on artiÞcial diet (Tsitsipis 1984)
that was changed twice a week. All experiments were
carried out in small incubators provided with 7-d programmers for illumination, temperature, and humidity. Light intensity in the incubators was measured as
22.5 ␮Ein m⫺2 s⫺1 (400 Ð700 nm) by a quantum sensor
(Model Li 188 B, LI-COR, Lincoln, NE). Humidity
was ⬇60 Ð 65%. The reported temperatures were accurate to within ⫾1⬚C, whereas the transition from
one temperature to another was essentially complete
within ⬇30 min after switching.
The developmental period of the immature stages of
S. nonagrioides was measured by exposing the different stages either to constant temperatures of 20, 22.5,
25, 27.5, and 30⬚C, or to corresponding alternating
temperatures with thermocryophase (T:C) temperatures of 25:10, 27.5: 12.5, 30:15, 32.5: 17.5, and 35:20⬚C,
under a constant photoperiod of 16:8 (L:D) h. This
photoperiod was selected because shorter photoperiods such as 12:12, or 14:10 (L:D) h were reported to
induce a portion of the population to diapause
(Fantinou et al. 1995), whereas under constant darkness, thermoperiods also could induce diapause
(Fantinou et al. 2002). The mean temperatures of the
given alternating temperature regimes were calculated on the basis of the arithmetic averages of hourly
temperatures through the 24-h cycle. Thus, each regime had a mean temperature corresponding to that of
the constant selected, and the high temperature was
synchronized with the photophase. Because our purpose was the comparison of development of immature
stages under constant and alternating temperatures,
temperatures ⬍20⬚C or ⬎35⬚C were not tested, because the cryophase or the thermophase probably
would be under or over the lethal values.
Hatchability and larval and pupal growth were measured under the temperature treatments noted above.
Vol. 32, no. 6
Newly collected eggs (⬍24 h old) were acquired from
the colony. The eggs were placed in 4-cm plastic petri
dishes on a Þlter paper moistened with propionic acid
(1:1000 dilution), and placed in incubators at the designated temperatures. Four replicates of 50 eggs each
were tested under each experimental regime. Hatchability was measured daily. Larval development was
investigated by placing newly hatched larvae (⬍24 h
old) on artiÞcial diet, in four replicates of 25 larvae
each, and rearing them through to pupation in plastic
boxes. Larvae were checked daily for pupation and
mortality. Four replicates of 15 newly formed pupae
each were placed in plastic boxes and exposed to the
experimental regimes. The number of emergent adults
was recorded daily. Observations were taken about
noon, which was between the 10th and 12th h of the
photophase.
The time (days) needed for 50% of the population
to complete each immature life stage at each temperature regime (either constant or alternating) was
noted. The developmental rate of each life stage of S.
nonagrioides was derived according to the temperature summation model and was calculated using the
reciprocal of the average days (1/d) of its duration.
The relationship between developmental rate (1/d)
and temperature (T) was Þrst estimated by the linear
function 1/d ⫽ ⫺ t/k ⫹ (1/k) T, where t and k are the
lower developmental threshold and the thermal constant (cumulative degree-days), respectively. Therefore, 1/d ⫽ a ⫹ bT, where t ⫽ ⫺a/b and k ⫽ 1/b. In
addition the improved nonlinear model of Lactin et al.
(1995) was used. This model, which is based on a
version of Logan et al. (1976), eliminates a redundant
parameter and introduces an intercept to allow estimation of a low developmental threshold. In addition,
it satisfactory describes the relationship between developmental rate and temperature above optimum,
and provides an estimation of the upper temperature
threshold. The model is composed of four parameters
and has the following form:
r共T兲 ⫽ e ␳T ⫺ e关 ␳Tmax⫺共Tmax⫺T兲/⌬兴 ⫹ ␭ ,
where r(T) is the developmental rate at temperature
T, and ␳(rho), T(max), ⌬, and ␭ are Þtted parameters.
The curves were Þtted with nonlinear regression using
the Marquardt algorithm in SPSS 8 (SPSS 1997).
Data were analyzed using the procedures of Statistica (StatSoft, Inc. 1995). Before statistical analysis, the
data were checked for analysis of variance (ANOVA)
assumptions and transformed, if needed, either to arcsine 公(x) (hatchability and emergence percentage),
or according to the Box-Cox method (Box and Cox
1964). Means were separated according to the least
signiÞcant difference (LSD) test (a ⫽ 0.05).
Results
S. nonagrioides achieved complete development
from egg to adult emergence at temperatures that
ranged from 20 to 30⬚C, when exposed to constant
temperatures or at the corresponding alternating temperatures. An exception was at (T:C) 35:20⬚C where
December 2003
FANTINOU ET AL.: TEMPERATURE AND DEVELOPMENT OF S. nonagrioides
1339
Table 1. Egg hatch, pupation, and adult emergence (%) ⴞ SE of S. nonagrioides, when exposed to different constant or corresponding
alternating temperatures
Eggs
Temperature
(T:C)⬚C
20:20
22.5:22.5
25:25
27.5:27.5
30:30
25:10
27.5:12.5
30:15
32.5:17.5
35:20
ANOVA
Temp.
Level
Temp. level
Larva
Pupa
Mean (%)
⫾SE
Mean (%)
⫾SE
Mean (%)
⫾SE
65.00ab
64.50ab
60.50ab
42.00ac
40.50
75.50b
69.00b
27.50c
27.00c
0
7.68
7.09
7.31
2.94
12.95
8.18
7.42
1.71
1.29
0
88.00a
86.00ab
87.00a
90.00a
66.00cde
81.00ac
73.00bcd
64.00cde
61.00de
50.00e
4.32
2.58
7.51
4.16
3.46
3.41
3.42
3.26
4.43
2.58
90.83a
90.18a
93.82a
93.30a
75.00b
79.02b
87.92a
82.55b
81.35b
76.36b
6.03
4.28
2.17
3.71
3.95
3.30
3.89
6.14
3.78
5.63
df
F
P
df
F
P
df
F
P
1,24
3,24
3,24
2.42
10.76
3.48
1326
.0001
.0315
1,30
4,30
4,30
30.34
6.19
1.45
⬍.0001
⬍.0009
⬍.239
1,30
4,30
4,30
17.68
3.02
2.89
⬍.0001
.0233
.0282
Data (%) were transformed using the arcsine square-root transformation before the ANOVAs were conducted. Values followed by same
letter in the same column are not signiÞcantly different (P ⫽ 0.05), LSD test. Means separation of egg hatchability was contacted at the range
of 20 Ð27.5⬚C for constant and 25:10 to 32.5:17.5⬚C for alternating temperatures.
no eggs hatched (Table 1), indicating that 35⬚C exceeded the upper threshold. Egg hatch rate was
greater under the alternating temperatures (T:C) 25:
10⬚C and (T:C) 27.5:12.5⬚C than under the corresponding constant. However, signiÞcantly lower hatch
occurred at (T:C) 30:15 or 32.5:17.5⬚C than under the
corresponding constant temperatures. Pupation and
adult emergence (percentage) under constant temperatures did not differ in the range of 20 Ð27.5⬚C but
was signiÞcantly lowered at 30⬚C (Table 1). Conversely, pupation decreased with increasing temperature under the alternating temperature regimes.
Slightly lower, but signiÞcant, percentages of adults
emerged when pupae were exposed to alternating
temperatures.
Mean developmental time of immature stages was
longer at the low constant temperature of 20⬚C than at
the corresponding alternating temperature (T:C) 25:
10⬚C, whereas the opposite was observed at 30⬚C (Ta-
ble 2). From 20 to 27.5⬚C, when insects were exposed
either to constant or alternating temperatures, mean
developmental time of each life stage signiÞcantly
decreased as temperature increased. Above this range,
this relationship no longer consistently held for the
egg or pupal stage. More time was needed for egg
maturation at 30⬚C than at 27.5⬚C. Pupal stage duration
was similar between 27.5 and 30⬚C but took longer at
(T:C) 35:20⬚C than at (T: C) 32.5:17.5⬚C (Table 2).
Based on these results, the lower threshold and the
degree-days required for the development of each life
stage of S. nonagrioides were estimated by the linear
model (Table 3). Regressions were conducted in the
ranges of temperatures in which this relationship was
linear. For the egg and pupal stage this range was from
20 to 27.5⬚C, whereas for larvae it was from 20 to 30⬚C
for both constant and alternating temperatures.
Higher coefÞcients of determination (R2) were obtained for all immature stages at constant than alter-
Table 2. Mean developmental time (days) ⴞ SE of egg, larval, and pupal stage of S. nonagrioides, when exposed to different constant
or corresponding alternating temperatures
Temperature
(T:C)⬚C
Egg
Temp.
Level
Temp. level
Pupa
⫾SE
Mean (d)
⫾SE
Mean (d)
⫾SE
11.92a
9.78b
8.28c
6.63d
7.70
10.67c
9.45b
8.45c
6.58d
0.08
0.08
0.08
0.10
0.10
0.08
0.08
0.13
0.19
51.38a
37.81bc
33.98de
30.97e
25.56f
48.69a
40.58b
35.87cd
34.75cd
27.08f
2.66
0.78
1.53
0.81
0.36
0.40
0.58
1.05
0.14
0.14
18.98a
14.06b
11.42c
10.38d
10.27d
18.17e
13.40f
12.88f
11.12c
12.17g
0.12
0.12
0.14
0.12
0.14
0.13
0.13
0.13
0.13
0.14
20:20
22.5:22.5
25:25
27.5:27.5
30:30
25:10
27.5:12.5
30:15
32.5:17.5
35:20
ANOVA
Larva
Mean (d)
Estimated total
period (d)
82.28
61.65
53.66
47.98
43.53
77.53
63.43
57.20
52.45
df
F
P
df
F
P
df
F
P
1,24
3,24
3,24
23.34
692.91
25.73
⬍.0001
⬍.0001
⬍.0001
1,30
4,30
4,30
40.09
422.59
5.53
⬍.0001
⬍.0001
.0002
1,30
4,30
4,30
74.44
1072.51
52.26
⬍.0001
⬍.0001
⬍.0001
Values followed by same letter in the same column are not signiÞcantly different (P ⫽ 0.05), LSD test. Means separation of egg duration
was contacted at the range of 20 Ð27.5⬚C for constant and 25:10 to 32.5:17.5⬚C for alternating temperatures.
1340
ENVIRONMENTAL ENTOMOLOGY
Vol. 32, no. 6
Table 3. Parameters of linear regression model and R2 values for temperature-dependent developmental rates of immature stages
of S. nonagrioides
Temperature
Stage
Constant
Alternating
Constant
a
Egg
Larva
Pupa
0.0087
0.0018
0.0059
Alternating
Constant
0.0071
0.0015
0.0043
0.0921
0.0016
0.0634
Alternating
Constant Alternating
R2
b
0.0513
0.0093
0.0283
0.9602
0.9222
0.9663
Constant
t
0.9186
0.9107
0.9014
10.57
8.89
10.75
Alternating
k
7.23
6.20
6.58
114.94
555.56
169.49
140.85
666.67
232.56
a, intercept; b, slope; t, lower developmental threshold in ⬚C; k, cumulative degree-days required for stage development.
Linear parameters were estimated in the range of 20 Ð27.5⬚C for the egg and pupal stage and from 20 to 30⬚C for larvae under both constant
and alternating temperatures.
nating temperatures (Table 3). The estimated developmental thresholds of the various stages ranged from
8.9 to 10.8⬚C (constant) and from 6.2 to 7.2⬚C (alternating). Lower developmental thresholds were estimated when the insects were exposed to alternating
temperatures than when they were exposed to the
constant temperatures.
The nonlinear model Lactin-2, when Þtted to values
of mean developmental rate, gave a good Þt to the data
sets for the range of temperatures used (Fig. 1aÐ c;
Table 4). Although the range of temperature did not
extend to lower values (e.g., 5, 10, or 15⬚C), we tested
this model for estimating optimal and lethal thresholds
for the immature stages. The resulting coefÞcients of
determination were between 0.91 and 0.98 and had
higher values at constant than alternating temperatures (Table 4). The low developmental thresholds for
the egg and larval stages were similar to those estimated by the linear model. However, higher values of
the low developmental threshold for the pupal stage
were estimated by Lactin-2 model than that by the
linear, either for constant or alternating temperatures.
The optimal developmental temperatures estimated
for egg, larval, and pupal stages were 29.5, 41.25, and
29.25 under constant and 38.00, 44.25, and 27.75⬚C
under alternating temperatures, respectively (Table
4). In the range of 20 Ð27.5⬚C, the curves obtained
were approximately linear, whereas at the high temperatures the lethal points did not differ from the
observed values for egg and pupal stages (30.25 and
37.25⬚C under constant and 39.75 and 35.50⬚C under
alternating temperatures, respectively). Above the
optimal temperature, the developmental rate decreased and for eggs and pupae the lethal thresholds
values were between 30 and 35⬚C, whereas for larvae
the drop from optimal development to lethality
showed a very wide range and was placed at 40 Ð 45⬚C.
1). Hence, developmental time of egg maturation began to increase at high temperature indicating that
very warm as well as cold temperatures can retard. In
Discussion
Egg hatch, pupation, adult emergence (%) and developmental time at each life stage of S. nonagrioides
were strongly inßuenced by temperature (Tables 1,2).
Whatever the life stage, increasing temperature resulted in predictable increases in rates of development, with the exceptions of the egg stage at the
constant temperature of 30⬚C and the pupal stage
under the alternating regime of (T: C) 35:20⬚C (Table
Fig. 1. Developmental rate (day⫺1) of (a) eggs, (b)
larvae, and (c) pupae of S. nonagrioides as a function of
constant (F) and alternating (E) temperatures (⬚C). Fitted
curves according to Lactin et al. (1995).
December 2003
FANTINOU ET AL.: TEMPERATURE AND DEVELOPMENT OF S. nonagrioides
1341
Table 4. Parameters (ⴞSE) of Lactin et al. (1995) model and R2 values for temperature-dependent developmental rates of immature
stages of S. nonagrioides
Stage
Temperature
Egg
constant
alternating
constant
alternating
constant
alternating
Larva
Pupa
Parameter estimates
␳
Tmax
⌬
␭
R2
t
0.0073 ⫾ 0.0003
0.0061 ⫾ 0.0004
0.0017 ⫾ 0.0001
0.0014 ⫾ 0.0001
0.0091 ⫾ 0.0040
0.0077 ⫾ 0.0051
30.3896 ⫾ 5.5785
40.8473 ⫾ 0.0000
44.2713 ⫾ 0.0000
48.4214 ⫾ 0.0000
48.1542 ⫾ 9.1664
45.9451 ⫾ 10.9093
0.1144 ⫾ 0.0001
0.5375 ⫾ 0.0000
0.4165 ⫾ 0.0000
0.6937 ⫾ 0.0000
6.3328 ⫾ 5.2501
5.5673 ⫾ 6.0261
⫺1.0757 ⫾ 0.0096
⫺1.0403 ⫾ 0.0125
⫺1.0151 ⫾ 0.0033
⫺1.0083 ⫾ 0.0028
⫺1.1307 ⫾ 0.0513
⫺1.0983 ⫾ 0.0762
0.957
0.921
0.922
0.909
0.985
0.916
10.00
6.45
8.60
5.85
14.08
12.50
␳, rate of increase to optimum temperature; Tmax, lethal temperature; ⌬, difference between the lethal temperature and the optimum
temperature of development; ␭, parameter that makes the curve intercept the x-axis; t, lower developmental threshold in ⬚C.
addition, egg hatchability was reduced at these very
high temperatures. The data on partial development of
eggs under high but alternating temperatures indicated that this stage could achieve a substantial
amount of development at unfavorable temperatures.
However, high temperatures were not detrimental to
pupal or larval development under the alternating
temperatures of (T: C) 35:20⬚C. Thus, the detrimental
effects of high temperatures (⬎35⬚C) during the day
on larval development in nature are probably not
pronounced, because they alternate with lower temperatures during the night. Temperatures of 35 or even
40⬚C are common in Greece during July and early
August, when larvae are present in the Þelds.
The linear model provided a good Þt to the data for
developmental rate of immature stages of S. nonagrioides, both at constant and alternating temperatures.
Regardless of the temperatures tested, there were not
great differences in the developmental thresholds estimated for the various stages. However, lower values
were obtained when the insects were exposed to alternating temperature cycles (Table 3). Similarly, Behrens et al. (1983) found that alternating temperatures
with an amplitude of 12⬚C lowered the estimated temperature threshold for embryonic development of
Gryllus bimaculatus De Geer from 16.6⬚C at constant
temperatures to 11.8⬚C. Yamashiro et al. (1998) found
that alternating temperatures reduced the thermal
requirements for oviposition in Arrachya menetriessi
Feldermann.
Developmental thresholds obtained via the linear
model are approximately equal to those reported by
Lopez et al. (2001) for the egg and pupal stages, but
they differed for the larval stage. The differences may
be due to the different origin of the insect populations
or to the different temperature ranges used in these
two studies; Lopez et al. (2001) tested larval development at a range of 15Ð27.5⬚C. Conversely, Thanopoulos and Tsitsipis (1987) reported a lower developmental threshold for larval S. nonagrioides of
10.85⬚C for a population from Greece.
The high egg hatchability of S. nonagrioides we
observed under alternating temperatures of (T:C) 25:
10⬚C indicates that 10⬚C probably was not detrimental,
although it was close to the lower developmental
threshold. This suggests that some insect development
occurs at temperatures lower than those at which
development can be completed at constant tempera-
tures. There may not be a distinct temperature developmental threshold with alternating temperatures,
but developmental rates may become asymptotically
lower as temperature decreases. Therefore, developmental zero, as estimated by linear extrapolation of the
middle portion of the developmental curve, does not
correspond to the temperature level below which development ceases. However, it probably corresponds
to the temperature level at which biological materials
such as eggs or pupae often can survive for long periods with little or no development or hatching (as at
higher temperature levels) (Karandinos and Axtell
1967). Therefore, there is an indication that constant
temperatures outside a fairly narrow intermediate
range yield abnormally low developmental rate and
survival values. For example, under the alternating
temperatures of (T:C) 35:20⬚C, 76.36% of adult
emerged, whereas no pupal survivorship occurred at
a constant temperature of 36⬚C (Lopez et al. 2001).
Our data also showed that a temperature of 30⬚C
caused a decrease in the developmental rate of eggs
and pupae, and a consequent departure from the linear trend. Therefore, we applied the Lactin model to
investigate the adverse effect of extreme temperatures
on developmental rate. That model satisfactorily described the relationship of developmental rate and
temperature for immature stages of S. nonagrioides.
The model Þt the data well and predicted values for
the lower developmental thresholds for the egg and
larval stages similar to those estimated by the linear
model. However, estimates for the pupal stage differed
between the two models. The lack of data at lower
temperatures may have led to an underestimation of
the threshold by the nonlinear model. Also, the values
for the lower developmental thresholds estimated by
the Lactin model were lower under the alternating
than constant temperatures. The estimated lethal
thresholds seemed to be close to the high temperatures tested at which high mortality rates were observed. Although high mortality at high temperatures
makes the study of developmental rate difÞcult, more
data are needed at extreme temperatures to accurately
estimate the upper temperature threshold. Nevertheless, the high larval survival and the linear trend of
larval developmental rate within the temperatures
tested in this study suggest that this species can withstand relatively high temperatures due to its tropical
1342
ENVIRONMENTAL ENTOMOLOGY
origin (Commonwealth Institute of Entomology,
1979).
The data presented here show that this insect can
undergo a signiÞcant amount of development at temperatures outside the optimal range, especially at
lower temperatures. This suggests that in many localities it may be necessary to consider development
under alternating temperatures when predicting population events in the Þeld. The survival estimates and
developmental model generated in this study will require further testing and validation in the Þeld.
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Merkel-Wallner. 1983. Effects of diurnal thermoperiods
and quickly oscillating temperatures on the development
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Received for publication 29 April 2003; accepted 5 September
2003.