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Human Anatomy & Physiology (11th Edition)
11th Edition
ISBN:9780134580999
Author:Elaine N. Marieb, Katja N. Hoehn
Publisher:Elaine N. Marieb, Katja N. Hoehn
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Density effects on pupal and adult mass
The weights of the pupae (F= 91; df = 2, 1319; P = 0.0001)
and adults (F= 13; df = 2, 1319; P = 0.0001) were influenced
by larval density. Pupae and adults were significantly heavier
when reared from uncrowded larvae than those from crowded
or moderately crowded larvae. There were no significant pupal
or adult weight differences between those reared from
crowded and moderately crowded larvae (P = 0.05, Tukey's test).
Fecundity and longevity
The fecundity (total number of eggs laid per living female)
and longevity of A.diaperinus adults at 25°C varied during the
9-month period of observations. The mean (±SEM) numbers of
eggs per day laid by a female adult beetle were 3.57 ± 0.09,
4.76 ± 0.28, 7.33±0.15 and 5.48 ± 0.23 when the beetle was 0.5,
3, 6 and 9 months old, respectively. The mean proportion sur-
vival of the female and male adults at 0.5 month of age was 0.99;
at 9 months it was 0.73 and 0.77, respectively. Preiss & Davidson
(1971) reported that t mean life-span of A.diaperinus is greater
than 400 days. They observed a female beetle deposit 2684 eggs
in 703 days (3.81 eggs/female/day) at room temperature. Wilson
& Miner (1969) reported the effect of temperature on the fecun-
dity of A.diaperinus during a 25-day oviposition period. Although
the age of the adult females used in their study was unknown,
they reported the mean numbers of eggs laid per female per
day were 2.0. 2.7. 5.5 and 3.7 at 21.1, 26.7, 32.2 and 37.8°C,
respectively. In our study at 25°C, each female adult laid from
3.6 to 7.3 eggs per day. Dass et al. (1984) reported that the mean
fecundity per female at 27°C was 4.6 eggs per day throughout
the oviposition period that averaged 324 days. Barke & Davis
(1969) noted that each female laid 7-8 eggs per day at 21°C.
Data from the literature on development times for A.diaperinus
are often not detailed, based on very limited numbers of speci-
mens, and difficult to use for comparisons. In our study, mean
total development times for the various life stages were shorter
than those reported by Ichinose et al. (1980). For example, the
mean days for development from egg to adult emergence at 20,
25, 30 and 35 C in our study were 164, 60, 36 and 29, respec-
tively, whereas Ichinose et al. (1980) reported 238, 76, 49 and
38, respectively. Most of the differences were in the develop-
ment times of the larvae. The mean for development of the lar-
vae at 20. 25. 30 and 35°C in our study were 134, 46, 26 and 22,
respectively, whereas Ichinose et al. (1980) reported 212, 63, 40
and 31, respectively. Data on much fewer larvae (40-92 per tem-
perature) were obtained by Ichinose et al. (1980) than in our
study (358-815 per temperature). Swatonek (1970) reported de-
velopment times at 25°C for only 15 specimens and the days
from egg to adult emergence was 48 compared to 60 in our study.
At 38°C, our data on total development times from egg hatch to
adult emergence and egg incubation times were 14.5 and 0.5
days shorter, respectively, when compared with those of Wilson
& Miner (1969). Ichinose et al. (1980) reported no development
for eggs, larvae and pupae of A.diaperinus at 15.7°C, 17.9°C
and 14.7°C. respectively. Wilson & Miner (1969), however, ob-
served egg and larval development at 15.6°C, whereas Swatonek
(1970) noted egg. larval and pupal development at 15°C. These
Alphitobius diaperinus development 85
differences could be attributed to different strains used in the
studies as well as differences in handling procedures and
temperature control.
Temperature is the major influence on the development of
immature A.diaperinus, as indicated in our study and other pre-
vious reports. Although there were density effects on larval de-
velopment rates that were statistically significant, the differences
were relatively small. It is reasonable to expect that under severe
conditions (e.g. high temperature, overcrowding, low food sup-
ply, poor food quality), some individual beetles may exhibit
different developmental times compared to individuals at opti-
mum habitat conditions. For example, Wilson & Miner (1969)
observed that larvae had a variable number of instars depending
upon temperatures; at high temperatures (32-38°C), there were
eight larval instars whereas there were eleven instars at
15.6°C.
Our data on A.diaperinus development as influenced by
temperature and the estimates of the parameters for the Sharpe &
DeMichele model, can be used in the construction of computer
simulation models to predict development and population
dynamics of this species under variable temperature conditions.
Although temperature is the major factor affecting the rates of
development and survival, the effects of additional factors (e.g.
food quality and availability, moisture, adult longevity and
fecundity, and effects of microorganisms in the poultry litter)
must be included in those simulation models.
Acknowledgments
The authors thank B. Cassel and G. Burke for laboratory
assistance and R. E. Stinner for statistical and modeling advice.
The research in this publication was funded in part by the North
Carolina Agricultural Research Service.
References
Axtell, R.C. & Arends, J.J. (1990) Ecology and management of arthro-
pod pests of poultry. Annual Review of Entomology, 35, 101-126..
Berke, H.E. & Davis, R. (1967) Sexual dimorphism in the lesser
mealworm, Alphitobius diaperinus (Panzer) (Coleoptera:
Tenebrionidae). Journal of the Georgia Entomological Society, 4, 119-
121.
Barke, H.E. & Davis, R. (1969) Notes on the biology of the mealworm,
Alphitobius diaperinus (Coleoptera: Tenebrionidae). Journal of the
Georgia Entomological Society, 4, 46-50.
Calnek, B.W., Barnes, H.J., Beard, C.W., Reid, W.M. & Yoder, H.W., Jr
(1991) Diseases of Poultry, 9th edn. Iowa State University Press, Ames,
Iowa.
Dass, R., Paul, A.V.N. & Agarwal, R.A. (1984) Feeding potential and
biology of the lesser mealworm, Alphitobius diaperinus (Panz.) (Col.,
Tenebrionidae), preying on Corcyra cephalonica St. (Lep., Pyralidae).
Zeitschrift für Angewandte Entomologie, 98, 444-447.
Despins, J.L. & Axtell, R.C. (1994) Feeding behavior and growth of
turkey poults fed larvae of the darkling beetle, Alphitobius diaperinus.
Poultry Science, 73, 1526-1533.
:
Transcribed Image Text:Density effects on pupal and adult mass The weights of the pupae (F= 91; df = 2, 1319; P = 0.0001) and adults (F= 13; df = 2, 1319; P = 0.0001) were influenced by larval density. Pupae and adults were significantly heavier when reared from uncrowded larvae than those from crowded or moderately crowded larvae. There were no significant pupal or adult weight differences between those reared from crowded and moderately crowded larvae (P = 0.05, Tukey's test). Fecundity and longevity The fecundity (total number of eggs laid per living female) and longevity of A.diaperinus adults at 25°C varied during the 9-month period of observations. The mean (±SEM) numbers of eggs per day laid by a female adult beetle were 3.57 ± 0.09, 4.76 ± 0.28, 7.33±0.15 and 5.48 ± 0.23 when the beetle was 0.5, 3, 6 and 9 months old, respectively. The mean proportion sur- vival of the female and male adults at 0.5 month of age was 0.99; at 9 months it was 0.73 and 0.77, respectively. Preiss & Davidson (1971) reported that t mean life-span of A.diaperinus is greater than 400 days. They observed a female beetle deposit 2684 eggs in 703 days (3.81 eggs/female/day) at room temperature. Wilson & Miner (1969) reported the effect of temperature on the fecun- dity of A.diaperinus during a 25-day oviposition period. Although the age of the adult females used in their study was unknown, they reported the mean numbers of eggs laid per female per day were 2.0. 2.7. 5.5 and 3.7 at 21.1, 26.7, 32.2 and 37.8°C, respectively. In our study at 25°C, each female adult laid from 3.6 to 7.3 eggs per day. Dass et al. (1984) reported that the mean fecundity per female at 27°C was 4.6 eggs per day throughout the oviposition period that averaged 324 days. Barke & Davis (1969) noted that each female laid 7-8 eggs per day at 21°C. Data from the literature on development times for A.diaperinus are often not detailed, based on very limited numbers of speci- mens, and difficult to use for comparisons. In our study, mean total development times for the various life stages were shorter than those reported by Ichinose et al. (1980). For example, the mean days for development from egg to adult emergence at 20, 25, 30 and 35 C in our study were 164, 60, 36 and 29, respec- tively, whereas Ichinose et al. (1980) reported 238, 76, 49 and 38, respectively. Most of the differences were in the develop- ment times of the larvae. The mean for development of the lar- vae at 20. 25. 30 and 35°C in our study were 134, 46, 26 and 22, respectively, whereas Ichinose et al. (1980) reported 212, 63, 40 and 31, respectively. Data on much fewer larvae (40-92 per tem- perature) were obtained by Ichinose et al. (1980) than in our study (358-815 per temperature). Swatonek (1970) reported de- velopment times at 25°C for only 15 specimens and the days from egg to adult emergence was 48 compared to 60 in our study. At 38°C, our data on total development times from egg hatch to adult emergence and egg incubation times were 14.5 and 0.5 days shorter, respectively, when compared with those of Wilson & Miner (1969). Ichinose et al. (1980) reported no development for eggs, larvae and pupae of A.diaperinus at 15.7°C, 17.9°C and 14.7°C. respectively. Wilson & Miner (1969), however, ob- served egg and larval development at 15.6°C, whereas Swatonek (1970) noted egg. larval and pupal development at 15°C. These Alphitobius diaperinus development 85 differences could be attributed to different strains used in the studies as well as differences in handling procedures and temperature control. Temperature is the major influence on the development of immature A.diaperinus, as indicated in our study and other pre- vious reports. Although there were density effects on larval de- velopment rates that were statistically significant, the differences were relatively small. It is reasonable to expect that under severe conditions (e.g. high temperature, overcrowding, low food sup- ply, poor food quality), some individual beetles may exhibit different developmental times compared to individuals at opti- mum habitat conditions. For example, Wilson & Miner (1969) observed that larvae had a variable number of instars depending upon temperatures; at high temperatures (32-38°C), there were eight larval instars whereas there were eleven instars at 15.6°C. Our data on A.diaperinus development as influenced by temperature and the estimates of the parameters for the Sharpe & DeMichele model, can be used in the construction of computer simulation models to predict development and population dynamics of this species under variable temperature conditions. Although temperature is the major factor affecting the rates of development and survival, the effects of additional factors (e.g. food quality and availability, moisture, adult longevity and fecundity, and effects of microorganisms in the poultry litter) must be included in those simulation models. Acknowledgments The authors thank B. Cassel and G. Burke for laboratory assistance and R. E. Stinner for statistical and modeling advice. The research in this publication was funded in part by the North Carolina Agricultural Research Service. References Axtell, R.C. & Arends, J.J. (1990) Ecology and management of arthro- pod pests of poultry. Annual Review of Entomology, 35, 101-126.. Berke, H.E. & Davis, R. (1967) Sexual dimorphism in the lesser mealworm, Alphitobius diaperinus (Panzer) (Coleoptera: Tenebrionidae). Journal of the Georgia Entomological Society, 4, 119- 121. Barke, H.E. & Davis, R. (1969) Notes on the biology of the mealworm, Alphitobius diaperinus (Coleoptera: Tenebrionidae). Journal of the Georgia Entomological Society, 4, 46-50. Calnek, B.W., Barnes, H.J., Beard, C.W., Reid, W.M. & Yoder, H.W., Jr (1991) Diseases of Poultry, 9th edn. Iowa State University Press, Ames, Iowa. Dass, R., Paul, A.V.N. & Agarwal, R.A. (1984) Feeding potential and biology of the lesser mealworm, Alphitobius diaperinus (Panz.) (Col., Tenebrionidae), preying on Corcyra cephalonica St. (Lep., Pyralidae). Zeitschrift für Angewandte Entomologie, 98, 444-447. Despins, J.L. & Axtell, R.C. (1994) Feeding behavior and growth of turkey poults fed larvae of the darkling beetle, Alphitobius diaperinus. Poultry Science, 73, 1526-1533. :
84 L. M. Rueda and R. C. Axtell
Table 3. Mean (SE) proportion survival of eggs, larvae. pupae and
total immatures of Alphitobius diaperinus at five constant temperatures.
Life stage
Egg
Larva
Pupa
Immature*
Life stage
Pupae
Temp.
(°C)
Female
Male
Both sexes
20
25
30
35
38
Adults
20
25
30
35
38
Female
Male
Both sexes
20
25
30
35
38
20
25
30
35
38
Mean (SE)
survival'
0.61 ± 0.03a
0.72 ± 0.05ab
0.86 ± 0.02c
*Egg hatch to adult emergence.
Means followed by the same letter within each life stage category
are not significantly different (P = 0.05, Tukey's test; SAS Institute, 1988).
20
0.81 ± 0.02bc
0.79± 0.04bc
(Fig. 3). Overall, uncrowded larvae had significantly higher
rates of survival compared with crowded larvae but not with
moderately crowded larvae. There were no significant survival
differences between moderately crowded and crowded larvae
(P = 0.05; Tukey's test). When density-survival data were ana-
lysed for each temperature, there were no significant differences
in larval survival among the three densities at 20, 25, 30 and
38°C. At 35°C, survival of the larvae was significantly greater
at the uncrowded than at the other densities but there was no
significant difference between the two highest densities
(P=0.05, Tukey's test).
20.63±0.39a
16.16±0.30a
18.39 ± 0.32ab
0.32 ±0.03a
0.60 ± 0.03b
0.73 +0.03b
0.70 ± 0.03b
0.68 ± 0.04b
0.85 ± 0.04a
0.94 0.01b
0.93±0.01a
0.95 +0.01b
0.91±0.01a
0.28 0.03a
0.56 ± 0.03b
0.68 ± 0.03b
0.66 ± 0.03b
0.62± 0.04b
25
MEAN PROPORTION SURVIVAL
20.39±0.21a
16.57 ± 0.20a
18.48± 0.18a
1.0
0.9-
0.8-
0.7
0.6-
0.5-
0.4
0.3-
0.2-
0.1-
0.0
Table 4. Mean (SE) weights (mg) of pupae and adults of Alphitobius diaperinus reared as larvae at five constant temperatures.
Temperature (°C)*
20
25
30
TEMPERATURE C
30
Fig. 3. Mean proportion survival of Alphitobius diaperinus larvae reared
at three densities (1, 3 and 6 larvae per gram of medium) at each of five
constant temperatures (°C).
21.08± 0.22a
17.48±0.16b
19.28±0.17b
Temperature effects on pupal and adult mass
Pupal weight was significantly affected by the temperature at
which the larvae were reared (F= 9.86, df = 4, 1319; P = 0.0001)
(Table 4). Pupae produced from larvae reared at 35°C had the
lowest weights. Perhaps the smaller pupae reflected a reduced
number of larval instars and the adults produced from those pu-
pae would have reduced fecundity but these possibilities were
not investigated.
1 LARVA PER G
3 LARVAE PER G
6 LARVAE PER G
Adult weight also was influenced by the temperature at which
the larvae were reared (F= 280.83; df = 4, 1319; P = 0.0001).
Adults from larvae reared at 38°C had the lowest weight. When
separately analysed according to sex, female adults (16.2 mg)
were significantly heavier than male adults (13.2 mg). Similarly,
female pupae (20.3 mg) weighed significantly more than male
pupae (16.8 mg).
35
35
19.07 0.24b
16.41±0.20a
17.74 +0.18c
16.76 ± 0.25ac
14.58 +0.25b
15.67 ± 0.19b
38
38
16.77 ± 0.27a
14.08±0.21ab
15.43±0.21ab
16.49 ± 0.25a
13.11±0.23a
14.80 ± 0.20a
19.81 ±0.21b
16.05 ± 0.14c
17.93±0.17c
Means followed by the same letter within a row are not significantly different (P = 0.05, Tukey's test; SAS Institute, 1988).
20.45 0.18a
17.01±0.21ab
18.73±0.17ab
11.56 ± 0.14d
8.84 0.15d.
10.20 0.13d
1996 Blackwell Science Ltd, Medical and Veterinary Entomology 10: 80-86
Transcribed Image Text:84 L. M. Rueda and R. C. Axtell Table 3. Mean (SE) proportion survival of eggs, larvae. pupae and total immatures of Alphitobius diaperinus at five constant temperatures. Life stage Egg Larva Pupa Immature* Life stage Pupae Temp. (°C) Female Male Both sexes 20 25 30 35 38 Adults 20 25 30 35 38 Female Male Both sexes 20 25 30 35 38 20 25 30 35 38 Mean (SE) survival' 0.61 ± 0.03a 0.72 ± 0.05ab 0.86 ± 0.02c *Egg hatch to adult emergence. Means followed by the same letter within each life stage category are not significantly different (P = 0.05, Tukey's test; SAS Institute, 1988). 20 0.81 ± 0.02bc 0.79± 0.04bc (Fig. 3). Overall, uncrowded larvae had significantly higher rates of survival compared with crowded larvae but not with moderately crowded larvae. There were no significant survival differences between moderately crowded and crowded larvae (P = 0.05; Tukey's test). When density-survival data were ana- lysed for each temperature, there were no significant differences in larval survival among the three densities at 20, 25, 30 and 38°C. At 35°C, survival of the larvae was significantly greater at the uncrowded than at the other densities but there was no significant difference between the two highest densities (P=0.05, Tukey's test). 20.63±0.39a 16.16±0.30a 18.39 ± 0.32ab 0.32 ±0.03a 0.60 ± 0.03b 0.73 +0.03b 0.70 ± 0.03b 0.68 ± 0.04b 0.85 ± 0.04a 0.94 0.01b 0.93±0.01a 0.95 +0.01b 0.91±0.01a 0.28 0.03a 0.56 ± 0.03b 0.68 ± 0.03b 0.66 ± 0.03b 0.62± 0.04b 25 MEAN PROPORTION SURVIVAL 20.39±0.21a 16.57 ± 0.20a 18.48± 0.18a 1.0 0.9- 0.8- 0.7 0.6- 0.5- 0.4 0.3- 0.2- 0.1- 0.0 Table 4. Mean (SE) weights (mg) of pupae and adults of Alphitobius diaperinus reared as larvae at five constant temperatures. Temperature (°C)* 20 25 30 TEMPERATURE C 30 Fig. 3. Mean proportion survival of Alphitobius diaperinus larvae reared at three densities (1, 3 and 6 larvae per gram of medium) at each of five constant temperatures (°C). 21.08± 0.22a 17.48±0.16b 19.28±0.17b Temperature effects on pupal and adult mass Pupal weight was significantly affected by the temperature at which the larvae were reared (F= 9.86, df = 4, 1319; P = 0.0001) (Table 4). Pupae produced from larvae reared at 35°C had the lowest weights. Perhaps the smaller pupae reflected a reduced number of larval instars and the adults produced from those pu- pae would have reduced fecundity but these possibilities were not investigated. 1 LARVA PER G 3 LARVAE PER G 6 LARVAE PER G Adult weight also was influenced by the temperature at which the larvae were reared (F= 280.83; df = 4, 1319; P = 0.0001). Adults from larvae reared at 38°C had the lowest weight. When separately analysed according to sex, female adults (16.2 mg) were significantly heavier than male adults (13.2 mg). Similarly, female pupae (20.3 mg) weighed significantly more than male pupae (16.8 mg). 35 35 19.07 0.24b 16.41±0.20a 17.74 +0.18c 16.76 ± 0.25ac 14.58 +0.25b 15.67 ± 0.19b 38 38 16.77 ± 0.27a 14.08±0.21ab 15.43±0.21ab 16.49 ± 0.25a 13.11±0.23a 14.80 ± 0.20a 19.81 ±0.21b 16.05 ± 0.14c 17.93±0.17c Means followed by the same letter within a row are not significantly different (P = 0.05, Tukey's test; SAS Institute, 1988). 20.45 0.18a 17.01±0.21ab 18.73±0.17ab 11.56 ± 0.14d 8.84 0.15d. 10.20 0.13d 1996 Blackwell Science Ltd, Medical and Veterinary Entomology 10: 80-86
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