全 文 ：第 26卷第 6期
2006年 6月
生 态 学 报
ACTA ECOLOGICA SINICA
Vo1．26．No．6
Jun．，2006
食虫沟瘤蛛的饥饿耐受性
肖永红 ，贺一原 ’，杨海明
(1．湖南师范大学生命科学学院，长沙 410081；2．井冈山学院生命科学学院，吉安 343009
3．中南林学院资源与环境学院，长沙 410004)
摘要：对我国多数稻区共享的优势种蜘蛛食虫沟瘤蛛进行饥饿耐受性测定，结果显示食虫沟瘤蛛的耐饥时间以及组内个体耐饥
力的变异幅度与温度高低呈负相关。在35、25~C和 15~C 3种试验温度下 ，食虫沟瘤蛛的耐饥时间及变异幅度均与龄期呈正相
关；在 5~C时，耐饥时间极大延长，组内变异系幅度亦急剧增大 ，但此时耐饥时间和变异幅度均与龄期呈负相关。在高温 35~C和
低温 5℃下，3个龄期组的耐饥时间差异不显著(P>0．05)；但在适宜温度 25~C和 15~C时，3个龄期组之间的耐饥时间差异显著
(P<0．05)。同一龄期组的食虫沟瘤蛛在任意两种不同的温度条件下，其耐饥时间的差异性均达到极显著水平(P<0．O1)。4种
1
试验温度条件下，处于饥饿状态的食虫沟瘤蛛各龄期组的阶段死亡率均具有正态分布特点 ，可用正态分布模型 』If=— ·
√ 2 71"
f r一 、2
e一 ： 进行拟合；并可根据正态分布的特点得出食虫沟瘤蛛的饥饿半致死时间( )和致死时间( )。
关键词：食虫沟瘤蛛；饥饿耐受性；温度；龄期
文章编号 ：1000-0933(2006)06-1725—07 中图分类号：Q959．226 文献标识码 ：A
The starvation endurance of Ummeliata insecticeps
XIAO Yong．Hong 一，HE Yi．Yuan · ‘ ，YANG Hai．Ming (1．cbz ＆ ， MnⅣ0 f ，c^∞ 口^410081，
China；2．Colege of Sciences，Jinggangshan University，Ji’∞，343009，China；3．Colege ofLife~iertces and Technology，Central South Forestry University，
Changsha 410004，China)．ActaEcologica Sinica，20O6，26(6)：1725—1731．
Abstract：The paddy spider is one of the primary predators of rice posts——accounting for around 51．2％ to 89．5％ of the total
kinds of predators in paddy fields．The paddy ecosystem is complex and unstable because of climate variation and the activities of
living organisms and human beings．Th e paddy spiders must endure starvation periods because the populations of their prey aye
occasionally reduced in a large amount by the cold climate in winter or by farming activities like spraying chemicals，gathering，
plowing and so on．Studying the starvation endurance of the paddy spider can provide more detailed and precise parameters for
protecting and utilizing them．This paper focuses primarily on the starvation endurance of Ummeliata insecticeps(Boesenberg et
Strand)，which is common in most rice—planting areas in China．The results show that the survival time of U．insecticeps and the
variation among individuals are positively related with the spider s instar age at 35，25℃ and 1 5℃ 。but negatively related at 5℃ ．
In addition。both the survival time and the variation among individuals of al test groups increased greatly at 5℃ ．Whereas the
differences of survival time among the three instar—groups are insignifcant(P>0．05)at 35℃ and 5℃ ，the survival times of the
基金项目：国家自然科学基金重点资助项目(39830040)；湖南省教育厅青年科技基金资助项目(03B025)
收稿日期：2005—03—29：修订日期：2006一Ol—l5
作者简介：肖71*通讯作者 Coresponding author．E-mail：yiyuanH@fm365．com
Foundation item：The project was supported by Key Item of National Natural Science Foundation of China(39830040)and Youth s Fund Item of Technology and
Science of Education Department in Hu nan Province(03B025)
Received date：2005-03—29；A0cepted date：2006一Ol—l5
Biography：XIAO Yong-Hong，Master，mainly engaged in animal ecology．E—maih yonghongxiao01@126
． corn
Acknowledgements：We’d like to express our appreciation to Profesor Wang Hongquan(Colege of Life Sciences，Hu’nail Normal University for his instructive
sugestions on this study；We are also grateful to Alex Daue(Arts and Science Colege，University of Colorado。USA)for reviewing and signifcantly improving an
earlier version of the manuscript；The senior author thanks Wen Juhua and Liu Feng for colecting the spider specimens(Colege of Life Sciences，Hunan Normal
University)
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1726 生 态 学 报 26卷
three instar-groups are significantly diferent(P<0．05)at the favorable temperatures of25~C and 15℃ ．Temperature influences
the spider s starvation endurance greatly．The survival time of a certain instar-group that lived in one kind of temperature is greatly
diferent from that of the same instar-group at any other kind of temperature(PThere is a close relationship between the stage—mortality(M)and the survival time(T)of U．insecticeps in starvation．It
1 ( 一“)
can be fitted with a normal distribution model： M =— = · e一 ．rhe starvation endurance thresholds and lethal times of
6 0 2
diferent U．insecticeps instar-groups can be calculated according to the equation of the normal distribution mode1．
Key words~Ummeliata insecticeps；starvation endurance；temperature；instar
The natural predator is one of the important biotic factors to regulate pest populations in ecosystems 卜。 ．A predator
s capacity to adapt to unfavorable conditions is an important characteristic for evaluating it．McMurtry J A took “the
survival capacity during starvation periods’ as one of the general characteristics of efective predators[
． The paddy
ecosystem is complex and unstable，to some degree，because of climate variations and the activities of living organisms and
human beings．The paddy predators have to deal with starvation aperiodically because the populations of their prey in the
paddy field are occasionally reduced in a large amount by the cold climate in winter or by farming activities like spraying
chemicals，gathering，plowing and so on[ 吲
．
There were already some reports on starvation endurance of phytoseiid
mites，Chrysopa phyUochroma，Microvelia horvathi，Chilocorus rubidus，etc．_8_l
．
Spiders are essential predators against many insects in paddy fields because of their vigorous predation habits and
large population sizes in a variety of speciesll2
． It is importan t to investigate spiders’starvation endurance to strengthen
their role as insect regulators．There have been quite a few studies focusing on community structure and predation function
of spiders 引
． As far as spiders starvation endurance is concerned，however，there are just a few rough descriptions at
natura1．variable temperatures[
． In order to provide more detailed and precise parameters for protecting and utilizing
spiders，this paper aims to test the starvation endurance of U．insecticeps，the domi nant species of spiders found in most
rice planting areas in China，at four kinds of controlled temperatures．
1 Material and Method
1．1 Material
The spiders(U．insecticeps)were collected from a rice field in a western suburb of Changsha，Hu nan Province，
China．Only healthy，uninjured individuals of U．insecticeps were colected for the test．
1．2 Method
Each spider was weighed by electronic analytical balance(1／1 0000，Shanghai Balance Instrument Plant)，then put
into a 30ml cuvete with a litle wet cotton to provide suitable humi dity．Then the spiders were separated into three instar．
groups according to their instar age： 1 一 2 instar·group，3 一 4 instar-group and 5 instar—adult．group
． Four
temperatures were set in the test：35，25，15 oC and 5~C．At each temperature，three control groups and three test groups
were studied．The spiders were fed with sufficient drosophilae for three days，after which no drosophila were ven to the
test groups．The control groups continued to be fed with suficient drosophila every week．The number of dead spiders in
each group was recorded every 5 days．Th e experiment was repeated 2 times．
1．3 Data analysis
1·3．1 The stage mortality M =M 一M (M stands for mortality of the test groups，and M stands for mortality of the
control groups)
1 (r )2
1·3．2 The model of normal distribution M = ·e一 (M stands for mortality，T stands for survival time，
is the standard deviation， is the mean)
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6期 肖永红 等：食虫沟瘤蛛的饥饿耐受性 1727
1．3．3 The data accorded with the normal distribution．The variances of al the data groups were homogeneous according
to the method of Bartlett．Th e analysis of variance was taken by the fixed effect model，and the means were separated by
the method of Duncan s Multiple Range Test．Th e analysis of linear regression，correlation coefficient and variance
analysis were completed by EXCEL．Curve equations to fit the model of normal distribution，starvation endurance threshold
( )and lethal time(T95)were calculated by SPTOOL in software MATLAB5．3．
2 Result and Analysis
2．1 Th e survival time of U．unsecticeps individuals in starvation．Th e first death occured eadier in the young instar．
group living at hish temperature than the first death in old instar—group living at low temperature ．Th e times of the first
death occurence and last death occurence of the test spider groups are listed in Table 1．
Spiders can survive longer in a cold environment than
in a warnl environment．Furthermore， spiders with high
instar survived longer than those with low instar at 35℃ ．
25℃ and 15 oC but died more quickly at 5 oC．Th e average
survival time of the test spider groups is shown in Table 2．
Th e results of variance analysis are as follows：Th e
differences of survival time among three instar-groups are
insignificant at the hi【gh temperature of 35℃ and low
temperature 5℃ (F5℃ = 0．34 < F35℃ = 2．90 <
F(2
．
80，0 05)= 3．11， P > 0．05)． At the favorable
Tab el Su time of U．insecticeps individuals ofdiferent groups
temperature of 25~C and 15％ ，however，the difference of average survival time between 1 - 2 instar．group and 3 - 4
instar-group is significant(p<0．05)．Furthermore，the differences between the 5th instar-adult．group and the other two
instar-groups are extremely remarkable(P<0．O1)．The differences between the test groups at any two different
temperatures are also extremely remarkable(P<0．01)．
Table 2 The average survival tim e of test spider groups
Data are mean± SD·Means of saⅡ iow or column folowed by diferent leters are statistically diferent(ANOVAfolowed by LSD P(0．05，Duncan sTest
P<0．05)
It can be concluded that the older spiders display great superiority in resisting starvation when the environmental
temperature is favorable but are nearly the same as the young ones at the extreme temperatures
． Because the spider s bodv
and instar are limited by the exuvial time，the higher the spider s instar is，the larger its body is
． Large spiders need m0re
food and are able to prey on more insects．Large spiders also maintain much more energy storing material
， like fat，than
small spiders．Moreover，with growth，spiders physiological function improves and their regulation abilitv strengthens
gradually．Therefore，large spiders can endure dificult conditions like prey scarcity more efficiently and suⅣive longer
than small spiders．
Th e environmental temperature makes a great impact of spiders starvation endurance
． In ect0thenns the metab0lic
rate is relatively slow at low temperatures and more rapid as the environment becomes waimer【2 ．Therefore．the
consumption of materials that store energy ， such as fat， etc
． increases at high temperatures．In additi0n．high
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1728 生 态 学 报 26卷
temperatures may hinder normal metabolism of the spiders and disorder their physiological function． U．insecticeps has
evolved to slow its metabolic rate by lessening activities to lengthen survival time in starvation when the temperature is
extremely cold．This adaptation has played an important role in the spider’s starvation endurance at cold conditions
．
Regression analysis of the relationship between the survival time and instar shows that they display direct corelation at
35，25，15 oC，but inverse corelation at 5 oC．The linear equations are as follows：
Y35℃ = 1．625x +7．477
Y25ac = 3．944x + 12．947
Yl5℃ = 6．500x +25．421
Y5℃ =一6．750x + 106．35
： 0．9951
： 0．9934
： 0．9951
： 0．9951
The survival time and temperature are inversly corelated as the linear equ ations show．
1,t一 2 instar-group： Y = 一28．624x + 111．00 r = 0．8982
3 一 4 instar．group： Y = 一26．450x + 106．71 r = O．9282
5 instar-aduh．group： Y = 一24．110x + 102．37 r = 0．9663
2．2 Variances of starvation endurance of the test·groups
e variation degree of data groups can be found by
comparing the coeficient of variability． According to the
formula CV ： (the denominator stands for the standard
deviation，the numerator the mean)，variance coe佑cients of
survival time of diferent spider instar·groups are displayed in
Figure 1．
，I1Ile variance coeficient and temperature are inversely
correlated as the linear equations show．
喜0．9
暑 0．8
骚
三 0．4
g 0．3
U
Fig．1 Variance coefficients of starvation enduran ce of the test—groups
1“一 2 instar·group： Y = 0．2449 一 0．1855 r = 0．9629
3 ——4 instar-group： Y = 0．2265 一 0．1135 r = 0．9739
5 instar—adult·group： Y = 0．1983 一 0．0468 r = 0．9793
The relationships between variance coefficients and the spider s instar are positively corelated at 35．25 oC and 15 cI=
but negatively at 5 oC．They can be fitted by liner equations as follows：
Y35℃ = 0．0325 + 0．1027 r = 0
．
9994一
Y25 = 0．0353 + 0．2036 r = 0．9389
Yl5 = 0．0183 + 0．4548 r = 0．9355
Y5℃ =一0．0394 + 0．9184 r = 0．9994
It could be deduced that some constitutional diversities existed among the spider individuals．As individuals grow，
environmental factors exert more influences on their physiological functions．That’s why the variability of starvation
endurance increases with the spiders growth at normal temperatures．At the low temperature of 5 cI=，however，some
individuals in the 1 一 2 instar·group were especially able to resist low temperature and starvation and survived
．
Accordingly，the average survival time of the 1 s,一 2 instar·group lengthened greatly。and the range of variation increased
correspondingly．Young spiders are able to resist starvation and cold temperatures more efficiently than old 0nes
． The
reason could be that U． insecticeps larvae have strengthened their resilience in the evolutionary process because U
．
insecticeps survives the harsh winter as larvae or sub．aduh spiders．Another explanation could be that U
． insecticeps larvae
ale insensitive to cold so they can survive longer in starvation than older spiders in low temperatures
．
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6期 肖永红 等：食虫沟瘤蛛的饥饿耐受性 l729
2．3 Stage—mortality of U．insecticeps in starvation
Setting 5 test days as a stage when U．insecticeps lived at 35 oC，10 test days as a stage at 25 oC and 15 oC，and 30
test days as a stage at 5℃ ，the number of dead in each instar—group was recorded in each stage．The stage mortalities of
the three instar-groups were calculated according to M ：MI—M (in 1．3．1)．Taking survival time as abscisa and stage
mortality as ordinate，the relationship between stage mortality an d survival time in four kinds of temperatures are displayed
in fig．2．
— — l 一2nd instar 一一 一 3rd一4th instar ⋯ ⋯ 一 5th—adultinstar
Survival time(d)
Fig．2 Stage‘mortalities of U．insecticeps in starvation at diferent temperatures
It ea／i be observed from fig．2 that all curves take the form of a bell jar，which indicates that the curves possess
ch锄 cteristics。f n。肿 aJ distributi。n． Fiting the curves with n。m al distributi。n Ⅱl。de1 M = ——— == ． e一
． the
． 6 2 7I"
correlation coeficients and the significance levels are shown in Table 3
．
Obviously all the curves can be fairly wel fitted with
norm al distribution．It Van be concluded that at the four test
temperatures，35，25，15 oC and 5 oC，stage mortalities of
U． insecticeps instar-groups in starvation have norm al
distribution．
2．4 Starvation endurance threshold( )and lethal time
(T95)
Table 3 The imitation of stage mortality CHIVES of U ． insecticeps t0
normal distribution model equation
2·4·1 Starvation endurance threshold( )of U．insecticeps Starvation endurance threshold may be c0nsidered the
time when 50％ 。f the test spiders had died from starvati。n and 50％ of the individuals still survived
． Th e cunres。f stage
mortality are symmetrical ab。ut the line X u acc。rding to the property。f n。rm al distributi。n
． Because is the peak。f the
∞ ∞ 如 们 如 加 m 0 如 加 ：2 m 0
∞ 如 柏 加 m 0 柏 加 m 0
扫一H鲁 。目． 基∞
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1730 生 态 学 报 26卷
curve．whieh can be explained by the equation of normal distribution，the value corresponding to is the starvation
endum ce threshold(T5o)of the spider(the average survival time in table 1)．It can be seen that Tso of U．insecticeps
instar．grouDs increases as the temperature decreases．When the test temperatures ale 35，25，1 5~C and 5℃ ，the Tso was
10．73，20．84，38．42 days and 92．85 days，respectively．
2．4．2 StaⅣ 0n lethal time(T95)of U．insecticeps When T obeys normal distribution(the mean is ，the standard
deviati0n is )，X = _二_ 0beys standard n0mal distribution N(O，1)．It can be known that P(X≤1．65)=0．95 by
0
checking the standard normal distribution table．From this the starvation lethal time(T95)of different spider instar。groups
can be calculated．The T95 values of the spider instar-groups at the four test temperatures are listed in Table 4．
Table 4 shows that when the environmental tempe-
ratures are 35，25，15cc and 5cC，respectively，95％ U．
insecticeps individuals，which were provided with water and
space but no food，would die in 18．50，32．54，54．02 days
and 98．30 days，on average．
3 Conclusion and discussion
Table 4 The lethal times of U．insecticeps instar-groups in starvation
Starvation le山al time(d) ．
TempeIature —1t- 2~a instar 3d- 4t—h instal"5th instar-aduh M k
3．1 Th e spider is a predat0r that can endure starvation for a long time⋯ 2。
． Liu F X et al measured the starvation
endurance of 11 spider species(except for U．insecticeps)at four controled temperatures，20，25，30~C and 35~C．
Their studv showed that the spiders in starvation survive longer and longer as they grow and develop，as well as with
decreasing temperature[
．
Th is is consistent with the result of starvation endurance from 15℃ to 35℃ in this paper．By
testing an extremely low temperature，this paper reveals further that the starvation endurance of U． insecticeps increases
greatly at 5℃ and the young survive even longer than the old．It can be concluded：
First．10w temperature plays a crucial role in starvation endurance of U． insecticeps． U． insecticeps has evolved to
endure the challenges 0f the winter environment，such as cold and food scarcity．It survives cold conditions by reducing the
amount and intensity of activities and by slowing down metabolism．
Second，U．insecticeps survive winter mainly as larvae．A population with more larvae is a growing population，and
growing populations can control pests more eficiently．Th e U．insecticeps larvae have evolved to be superior to adults at
enduring starvation and cold temperatures．For this reason，they are wel adapted to rebuild the spider population each
spring，making them ideal predators for controlling pe st populations．
3．2 Human activities have a very signifcant influence on the development and decline of paddy ecosystems．One of the
distinct characteristics of paddy ecosystems is that their communities decline and rebuild periodically along with fanning
activities．Th e starvation endurance of U．insecticeps can help them endure food scarcity caused by cold climate in winter
or by fanning activities such as spraying pesticide and plowing the paddy field．When the prey population increases，U．
insecticeps can prey on them quickly and control the pest s population size effectively．Th erefore，U．insecticeps is one of
the most efective predator species in paddy fields．
3．3 Th e use of natural predators as a form of pesticide is not wel developed in China．Th e challenge of preserving the
commercial predators is one of the problems that are holding up production of predator stocks ．Strong starvation
endurance of predators at low temperatures brings forward the theoretical foundation for a preservation technique for them．
Further research on more spider species at lower temperatures is needed so that the best suitable temperature for preserving
spiders can be obtained．
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维普资讯 http://www.cqvip.com
6期 肖永红 等：食虫沟瘤蛛的饥饿耐受性 1731
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