蚂蚁与舞草互惠共生关系研究
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摘要
蚁运植物（myrmecochores）指种子由蚂蚁传播扩散的植物。
蚂蚁与蚁运植物的互惠共生关系是种群生态学研究中的一个重要方
面，在国际上已广泛开展，而中国没有这方面的研究报道。作者在云南西
双版纳等地发现了蚂蚁与植物的互惠共生现象；舞草Codariocalyx motorius
（Houtt.）Ohashi、圆叶舞草Codariocalyx gyroides（Roxb.ex Link）Hassk作为
蚁运植物在此前的国际国内文献中未见报道。本论文较系统地研究了蚂蚁
与蚁运植物互惠共生关系，并对其共生机理进行了有益探索，为蚂蚁与蚁
运植物互惠共生关系研究的深入开展奠定了重要基础。
一．新蚁运植物的生物生态学特性
本次研究发现的蚁运植物—舞草、圆叶舞草，主要生长在土壤营养贫
瘠、火烧频繁的生境。这与澳大利亚蚁运植物发生的主要环境比较一致。
但也有与之不同点：在澳大利亚，搬运种子的蚂蚁大多也是种子的捕食者，
而搬运舞草种子的蚂蚁与舞草仅为互惠共生关系，对种子并不取食。
舞草具有典型蚁运植物特征，即种子上附生有吸引蚂蚁并产生搬运行
为的油质体。
舞草的物候期因各地气温变化而异。在景谷，舞草开花期为8-10月，
结果期为9-11月；在西双版纳，舞草开花期为9-11月，结果期为10-12月。
由于种皮坚硬，自然条件下，种子萌发率低于20％，萌发的适宜温度
为20-30℃。通过物理、化学处理可显著提高种子萌发率，减少萌发所需时间。
二．扭运舞草种子的蚂蚁种类及其生物生态学特性
在西双版纳和思茅地区景谷县，共发现12种蚂蚁搬运舞草种于。对义
觅食及搬运行为观察表明，伊大头蚁是两个点搬运舞草种子的主要种类，
广泛分布于舞草生长的地域内。伊大头蚁主要在土壤中筑巢，蚁巢喜欢建
在阳光充足、温暖的生境。蚁巢为均匀分布，且巢群间相互排斥。
诱集试验结果表明：搬运舞草种子的蚂蚁对诱饵有一定的选择性，特
别趋于糖和鱼。
由于邻巨首蚁量较大，且对食物有很强的趋性，因此选择糖作诱饵会
影响其它种类的诱集。
伊大头蚁对糖、鱼和肉均有趋性，同时对种子也有一定的趋性。
甲醛仅对邻b首蚁觅食有影响，但该蚁对搬运舞草种子作用不大。
三．其它动物对舞草种于传播的影响
脊椎动物对舞草种于取食传播的试验表明，牛对种于的远距离传播有
一定的作用，但传括的概率很小。牛喜欢取食舞草。舞草种子经牛取食后，
大多被消化，只有极少量随粪便排出。经牛取食排泄的种子，种子上的油
质体还存在，蚂蚁可再次传播。所以认为，由于牛对舞草取食，因而总体
上对舞草种群的繁衍有抑制作用。
鸡可完全消化舞草种于。
经鸟取食的种于，油质体已被消化，但种子可随粪便排出，且萌发率
有所提高。
四．蚂蚁与舞草的互惠共生机理
蚂蚁通过＼舞草建立丘惠共生关系，叮从舞草种于油质体中获得＊。长
发育所需的氨基酸、无机Jh ％等营养。舞草通过与蚂蚁的互惠共生获得如
下益处：逃避呐齿类动物捕食。排除蚂蚁搬运种子后，啮齿类动物对舞草
种于的捕食率在景谷和励仑分别为28．33％和 16．5％。舞草种子释放时间
且且
‘Jw5蚁活动时问吻合，蚂蚁可先于啮齿类动物将趴牛种－厂运大。表明舞草
’J蚂蚁1忽共生更有益于减少啮齿类动物的批食u
研究结果表明，七壤表面的种于在刀耕火忡中将个部被烧死。但被蚂
蚁搬运到地「蚁巢处的种于，火烧后种厂汕为？街判提高。这是舞草通过
‘。业】蚁的I惠共生，得以避免刀耕火种对义种丁的3卜向。
蚂蚁取食油质体后丢弃的舞草种于与未经取食的种于萌发试验结果相
比，6U者萌发率有明显提高，萌发所需时间 ｝）di少。这种作用与人工去除油
质体促进了种于萌发相似。
蚂蚁搬运舞草种于的距离一般较短，似均超过种于自身的弹射距离。
儿种王要蚂蚁传播曲线显示，传插的距离各自都有－－高峰，平均传播的距
离在0．93＿－二．53ffi之问。传播曲线都有一个K K的尾部，即少量的蚂蚁传
播的距离相对较远。蚂蚁搬运还使种于到达营养较十富的蚁巢。伊大头蚁
蚁巢与附近七壤营养物质比较显示，蚁巢有机质及N、P、K含量均略高于
附近l：壤。
五．影响蚂蚁搬运舞草种于率的因子
蚂蚁搬运舞草种于率受多种因子影响。蚂蚁周年数量动态与搬运率有
关。此外，蚁运植物种类，种子离蚁巢远近，种于油质体等因素对搬运率
的影响程度较高：新旧种子对搬运率也有一定的影响。
六．蚂蚁与舞草在热带雨林中的分布
主要蚂蚁种类和蚁运植物分布调查发现，在热带雨林及片断化热带雨
林中，搬运舞草种于的主要蚂蚁和舞草己没有分hJ。这表明随着植被的恢
复，搬运舞草种于的主要蚂蚁和舞草均己退出植被的演替。舞草作为先锋
植物，其生长、建群与分hJ影响特殊生境植被的恢复，利用和保护蚂蚁与
蚁运植物的互惠共生关系，叮促进植被的恢复和演替。
The study on mutualism between ants and myrmecochores has been intensely carried out worldwide, especially in U S., Australia and South Africa. However, any study on this field in Chinese continent has not reported. Phenomenon ot?mutualism between some ants and several plants is not previously reported in Xishuangbanna, Yunnan province, China. The Codariocalyx moi~orius (Houtt.) Qhashi as well as Codarioculyx gyroides (Roxb.ex Link) Hassk is firstly identified to be myrmecochore in the world and their mutualism with some ants has been systematically studied together with discussion of its mechanism in the paper.
1. The biological and ecological characters of new myrmecochores
Codariocalyx mowrius and C. gyroides grow in infertile and frequently burned habits. The habits of the myrmecochores in Xishuangbanna, Yunnan, are nearly similar to main ones of the myrmecochores in Australia, and there are, however, some differences between two places that ants of dispersing myrmecochores?seeds are predators in Australia and the relationship between ants and the myrmecochores in Xishuangbanna is only mutualism.
Codariocalyx motorius is typically characteristic of myrmecochores; that is, theye being elaiosomes on seeds attracting ants for seeds?removal.
In Jing Gu region, C. moforius blossoms from Aug. to Oct. and fruits from
SePt. to Nov.; in Xishuangbanna, C moIorius blossoms frOm SePt. to Nov. and
fruits from Oct. to Dec.
Naturally, germinating rate of C motOrius's seed is below 20% and the
suitable temperature is 20--30C. Afler Physical and chemical treatment, the
germinating rate is significantly increased and the germinating time reduced.
2. Xinds of ants di8persing Co-- -- seed8 and their hiological
--and -ccoIogical charactcrs
In Xishuangbanna and Jinggu County Shoao District, l2 kinds of ants
removing C moIorius j seed have been fotmd. The results from observing their
behavior of looking fOr fOod and removing Seeds shOw that Pheidole yeensis
Forel is a main remover of C. motoriusk seed in the studied areas. PheidOle
yeensis Forel spreads widely over the habits snitab1e to C motorius and builds its
nest mainly in soils where it is sunshine and warm. The Nests evenly scatter, hut
mutually exclude in detinite distance.
The ants removing C. motoriusk seed are selective to baits and prefer to
candy and fish.
Because of larger number, Phebotogton atheis has a strong selectivity to
food, and therefOre collecting it with candy as bail will reduce numbers of other
ants collected.
Phebole yeensis has the same selectivity to candy, fish and meat. Its
selectivity to seed is also observed.
3. Thc cffcct of othcr animals on C motorus' 8ced rcmoval
The test of vertebrates' eating C motorius k seed and remc ving it has been
carried out and the results are the following. (l) Ox has some influence on the
removing seed distance, but the possibility is much lower, only about 0.067% of
test seed being found in ox excrement and them containing elaiosomes and still
v
attracti11g ants fOr removal again. So, ox plays a retarda11l role in deve1opment of
C' nlolt)l.ius3 POpulation with ox pretbrring to eat ' l,loIorius. (2) No seed in
excrefllent of 'chicken is found and chicken digests alI seeds, which it eats. (3)
'I'he seed tbund in excrement of birds do 11ot have claiosome, but have higher
germinating rate.
4. Thc mcchanism of mutualism bctWccn ants and Codariocalyx motOrlks
By mutuaIism, ants can get some nutriments, for examples, arnino acid and
inorganic elements from elaiosomes of C molorius'seed, and C motOrius can
also gets some benefitS, such as escaping froIn prey of rodent animals. Afler
excIuding ants' removing seed, the rate of rodent aniInals predation on seed is
28.33% and l6.5% respectively in Jinggu and Menglun. The tests show that time
of releasing '). motorius seed misses time of rodent animals' movement and
coincides with time of ants movement, which implies that mutualism between
ants and C) motorius is more beneficial to t
蚂蚁与蚁运植物的互惠共生关系是种群生态学研究中的一个重要方
面，在国际上已广泛开展，而中国没有这方面的研究报道。作者在云南西
双版纳等地发现了蚂蚁与植物的互惠共生现象；舞草Codariocalyx motorius
（Houtt.）Ohashi、圆叶舞草Codariocalyx gyroides（Roxb.ex Link）Hassk作为
蚁运植物在此前的国际国内文献中未见报道。本论文较系统地研究了蚂蚁
与蚁运植物互惠共生关系，并对其共生机理进行了有益探索，为蚂蚁与蚁
运植物互惠共生关系研究的深入开展奠定了重要基础。
一．新蚁运植物的生物生态学特性
本次研究发现的蚁运植物—舞草、圆叶舞草，主要生长在土壤营养贫
瘠、火烧频繁的生境。这与澳大利亚蚁运植物发生的主要环境比较一致。
但也有与之不同点：在澳大利亚，搬运种子的蚂蚁大多也是种子的捕食者，
而搬运舞草种子的蚂蚁与舞草仅为互惠共生关系，对种子并不取食。
舞草具有典型蚁运植物特征，即种子上附生有吸引蚂蚁并产生搬运行
为的油质体。
舞草的物候期因各地气温变化而异。在景谷，舞草开花期为8-10月，
结果期为9-11月；在西双版纳，舞草开花期为9-11月，结果期为10-12月。
由于种皮坚硬，自然条件下，种子萌发率低于20％，萌发的适宜温度
为20-30℃。通过物理、化学处理可显著提高种子萌发率，减少萌发所需时间。
二．扭运舞草种子的蚂蚁种类及其生物生态学特性
在西双版纳和思茅地区景谷县，共发现12种蚂蚁搬运舞草种于。对义
觅食及搬运行为观察表明，伊大头蚁是两个点搬运舞草种子的主要种类，
广泛分布于舞草生长的地域内。伊大头蚁主要在土壤中筑巢，蚁巢喜欢建
在阳光充足、温暖的生境。蚁巢为均匀分布，且巢群间相互排斥。
诱集试验结果表明：搬运舞草种子的蚂蚁对诱饵有一定的选择性，特
别趋于糖和鱼。
由于邻巨首蚁量较大，且对食物有很强的趋性，因此选择糖作诱饵会
影响其它种类的诱集。
伊大头蚁对糖、鱼和肉均有趋性，同时对种子也有一定的趋性。
甲醛仅对邻b首蚁觅食有影响，但该蚁对搬运舞草种子作用不大。
三．其它动物对舞草种于传播的影响
脊椎动物对舞草种于取食传播的试验表明，牛对种于的远距离传播有
一定的作用，但传括的概率很小。牛喜欢取食舞草。舞草种子经牛取食后，
大多被消化，只有极少量随粪便排出。经牛取食排泄的种子，种子上的油
质体还存在，蚂蚁可再次传播。所以认为，由于牛对舞草取食，因而总体
上对舞草种群的繁衍有抑制作用。
鸡可完全消化舞草种于。
经鸟取食的种于，油质体已被消化，但种子可随粪便排出，且萌发率
有所提高。
四．蚂蚁与舞草的互惠共生机理
蚂蚁通过＼舞草建立丘惠共生关系，叮从舞草种于油质体中获得＊。长
发育所需的氨基酸、无机Jh ％等营养。舞草通过与蚂蚁的互惠共生获得如
下益处：逃避呐齿类动物捕食。排除蚂蚁搬运种子后，啮齿类动物对舞草
种于的捕食率在景谷和励仑分别为28．33％和 16．5％。舞草种子释放时间
且且
‘Jw5蚁活动时问吻合，蚂蚁可先于啮齿类动物将趴牛种－厂运大。表明舞草
’J蚂蚁1忽共生更有益于减少啮齿类动物的批食u
研究结果表明，七壤表面的种于在刀耕火忡中将个部被烧死。但被蚂
蚁搬运到地「蚁巢处的种于，火烧后种厂汕为？街判提高。这是舞草通过
‘。业】蚁的I惠共生，得以避免刀耕火种对义种丁的3卜向。
蚂蚁取食油质体后丢弃的舞草种于与未经取食的种于萌发试验结果相
比，6U者萌发率有明显提高，萌发所需时间 ｝）di少。这种作用与人工去除油
质体促进了种于萌发相似。
蚂蚁搬运舞草种于的距离一般较短，似均超过种于自身的弹射距离。
儿种王要蚂蚁传播曲线显示，传插的距离各自都有－－高峰，平均传播的距
离在0．93＿－二．53ffi之问。传播曲线都有一个K K的尾部，即少量的蚂蚁传
播的距离相对较远。蚂蚁搬运还使种于到达营养较十富的蚁巢。伊大头蚁
蚁巢与附近七壤营养物质比较显示，蚁巢有机质及N、P、K含量均略高于
附近l：壤。
五．影响蚂蚁搬运舞草种于率的因子
蚂蚁搬运舞草种于率受多种因子影响。蚂蚁周年数量动态与搬运率有
关。此外，蚁运植物种类，种子离蚁巢远近，种于油质体等因素对搬运率
的影响程度较高：新旧种子对搬运率也有一定的影响。
六．蚂蚁与舞草在热带雨林中的分布
主要蚂蚁种类和蚁运植物分布调查发现，在热带雨林及片断化热带雨
林中，搬运舞草种于的主要蚂蚁和舞草己没有分hJ。这表明随着植被的恢
复，搬运舞草种于的主要蚂蚁和舞草均己退出植被的演替。舞草作为先锋
植物，其生长、建群与分hJ影响特殊生境植被的恢复，利用和保护蚂蚁与
蚁运植物的互惠共生关系，叮促进植被的恢复和演替。
The study on mutualism between ants and myrmecochores has been intensely carried out worldwide, especially in U S., Australia and South Africa. However, any study on this field in Chinese continent has not reported. Phenomenon ot?mutualism between some ants and several plants is not previously reported in Xishuangbanna, Yunnan province, China. The Codariocalyx moi~orius (Houtt.) Qhashi as well as Codarioculyx gyroides (Roxb.ex Link) Hassk is firstly identified to be myrmecochore in the world and their mutualism with some ants has been systematically studied together with discussion of its mechanism in the paper.
1. The biological and ecological characters of new myrmecochores
Codariocalyx mowrius and C. gyroides grow in infertile and frequently burned habits. The habits of the myrmecochores in Xishuangbanna, Yunnan, are nearly similar to main ones of the myrmecochores in Australia, and there are, however, some differences between two places that ants of dispersing myrmecochores?seeds are predators in Australia and the relationship between ants and the myrmecochores in Xishuangbanna is only mutualism.
Codariocalyx motorius is typically characteristic of myrmecochores; that is, theye being elaiosomes on seeds attracting ants for seeds?removal.
In Jing Gu region, C. moforius blossoms from Aug. to Oct. and fruits from
SePt. to Nov.; in Xishuangbanna, C moIorius blossoms frOm SePt. to Nov. and
fruits from Oct. to Dec.
Naturally, germinating rate of C motOrius's seed is below 20% and the
suitable temperature is 20--30C. Afler Physical and chemical treatment, the
germinating rate is significantly increased and the germinating time reduced.
2. Xinds of ants di8persing Co-- -- seed8 and their hiological
--and -ccoIogical charactcrs
In Xishuangbanna and Jinggu County Shoao District, l2 kinds of ants
removing C moIorius j seed have been fotmd. The results from observing their
behavior of looking fOr fOod and removing Seeds shOw that Pheidole yeensis
Forel is a main remover of C. motoriusk seed in the studied areas. PheidOle
yeensis Forel spreads widely over the habits snitab1e to C motorius and builds its
nest mainly in soils where it is sunshine and warm. The Nests evenly scatter, hut
mutually exclude in detinite distance.
The ants removing C. motoriusk seed are selective to baits and prefer to
candy and fish.
Because of larger number, Phebotogton atheis has a strong selectivity to
food, and therefOre collecting it with candy as bail will reduce numbers of other
ants collected.
Phebole yeensis has the same selectivity to candy, fish and meat. Its
selectivity to seed is also observed.
3. Thc cffcct of othcr animals on C motorus' 8ced rcmoval
The test of vertebrates' eating C motorius k seed and remc ving it has been
carried out and the results are the following. (l) Ox has some influence on the
removing seed distance, but the possibility is much lower, only about 0.067% of
test seed being found in ox excrement and them containing elaiosomes and still
v
attracti11g ants fOr removal again. So, ox plays a retarda11l role in deve1opment of
C' nlolt)l.ius3 POpulation with ox pretbrring to eat ' l,loIorius. (2) No seed in
excrefllent of 'chicken is found and chicken digests alI seeds, which it eats. (3)
'I'he seed tbund in excrement of birds do 11ot have claiosome, but have higher
germinating rate.
4. Thc mcchanism of mutualism bctWccn ants and Codariocalyx motOrlks
By mutuaIism, ants can get some nutriments, for examples, arnino acid and
inorganic elements from elaiosomes of C molorius'seed, and C motOrius can
also gets some benefitS, such as escaping froIn prey of rodent animals. Afler
excIuding ants' removing seed, the rate of rodent aniInals predation on seed is
28.33% and l6.5% respectively in Jinggu and Menglun. The tests show that time
of releasing '). motorius seed misses time of rodent animals' movement and
coincides with time of ants movement, which implies that mutualism between
ants and C) motorius is more beneficial to t
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