ECOLOGICAL AND BEHAVIORAL BASIS FOR REPRODUCTIVE ISOLATION IN THE SYMPATRIC ENCHENOPA BINOTATA COMPLEX (HOMOPTERA. MEMBRACIDAE)
ECOLOGICAL AND BEHAVIORAL BASIS FOR REPRODUCTIVE ISOLATION IN THE SYMPATRIC ENCHENOPA BINOTATA COMPLEX (HOMOPTERA. MEMBRACIDAE)
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同翅双翅目双翅目复合体（同翅目、膜科）生殖隔离的生态和行为基础
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发表时间：
1982
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通讯作者：
S. Guttman
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作者：
T. Wood;S. Guttman
进化生物学中的一个普遍问题是，是否可以通过开发新的寄主植物或改变生活史来促进共域物种形成。最近的研究(Maynard Smith，1966；Alexander，1968；Huettel和Bush，1972；Knerer和Atwood，1973；Bush，1969,1975；Tauber和Tauber，1977a，1977b；Edmunds和Alstad，1978)已经使共生物种形成比过去更容易被接受(Mayr，1970)，但它仍然受到争论(Futuyma和Mayer，1980)。我们过去与树蝉Enchenopa binotata Say的工作(Wood，1980；Guttman等人，1981)表明，这种昆虫沿着寄主植物线分裂成一个共域繁殖隔离物种的复合体。Enchenopa在北美东部的落叶乔木和灌木上很常见，远至巴拿马南部。来自北美7种寄主植物的Enchenopa在分类上都被列为同一种(Metcalf和Wade，1965)，尽管它们在若虫着色、产卵地点、卵泡成分、产卵时间和若虫取食地点上有所不同。特定寄主物种的雌性在有选择的情况下选择该寄主产卵，当被限制在一个小笼子里时，根据植物种类选择交配(Wood，1980)。来自每一寄主物种的昆虫在电泳上是可区分的，尽管从不同同种树收集的样本中可以发现“寄主小种”内的差异。Enchenopa在黑核桃、Ptela trifoliata、刺槐和Cercis canamsis上的电泳图谱彼此不同，也不同于两个彼此相似的Viburnum和Celstrus jobens上的Enchenopa(Guttman等人，1981)。因此，双齿棘豆是繁殖隔离的物种的复合体。Enchenopa的北美寄主植物代表了进化上的多样性植物，这些植物在广泛和局部上是同域的。当几个寄主物种被发现在一起时，Enchenopa可能发生在每个寄主上。雌性将卵插入寄主植物的枝条，并用一种称为卵泡的分泌物覆盖它们，这种分泌物保护越冬的卵子(Wood和Patton，1971)，并含有一种产卵引诱剂，负责在单一枝条上聚集卵块。由此产生的大量若虫聚集吸引了互惠互利的蚂蚁，这显然提高了若虫的存活率(Wood和Seilkop，Unpubl)。Wood(1980)在Bush(1975)工作的基础上提出了以下假设，以解释Enchenopa binotata复合体的分歧。根据目前的地理分布(Metcalf和Wade，1965)、哥斯达黎加的寄主植物和季节记录(Ballou，1936)以及我们中的一人(T.K.W.)的观察，推测原始种群为热带、多食性和多变性。当Enchenopa的祖先在北温带气候中定居时，他们遇到了落叶寄主，这促进了树蝉生活史协调的选择
A pervasive question in evolutionary biology is whether sympatric speciation can be promoted by the exploitation of new host plants or changes in life history. Recent studies (Maynard Smith, 1966; Alexander, 1968; Huettel and Bush, 1972; Knerer and Atwood, 1973; Bush, 1969, 1975; Tauber and Tauber, 1977a, 1977b; Edmunds and Alstad, 1978) have made sympatric speciation more readily accepted than in the past (Mayr, 1970) but it is still subject to debate (Futuyma and Mayer, 1980). Our past work (Wood, 1980; Guttman et al., 1981) with the treehopper Enchenopa binotata Say suggests that this insect diverged along host plant lines into a complex of sympatric reproductively isolated species. Enchenopa is common in eastern North America on deciduous trees and shrubs as far south as Panama. Enchenopa from each of seven North American host plants are all listed taxonomically as the same species (Metcalf and Wade, 1965), although they differ in nymphal coloration, oviposition site, egg froth composition, time of day they deposit eggs, and nymphal feeding sites. Females from a given host species select that host for oviposition when given a choice, and when confined to a single small cage select mates assortatively by plant species (Wood, 1980). Insects from each host species are electrophoretically distinguishable although differences within "host races" can be found among samples collected from different conspecific trees. Enchenopa on Juglans nigra, Ptelea trifoliata, Robinia pseudoacacia, and Cercis canadensis are electrophoretically different from each other and from those on two species of Viburnum and Celastrus scandens, which are similar to each other (Guttman et al., 1981). Thus E. binotata is a complex of reproductively isolated species. The North American host plants of Enchenopa represent evolutionarily diverse plants which are broadly and locally sympatric. When several host species are found together Enchenopa may occur on each host. Females insert eggs into the branch of the host plant and cover them with a secretion known as egg froth, which protects overwintering eggs (Wood and Patton, 1971), and contains an ovipositional attractant which is responsible for the clumping of egg masses on single branches. The large aggregations of nymphs that result attract mutualistic ants, which apparently increase nymphal survival (Wood and Seilkop, unpubl.). Wood (1980) suggested the following hypothesis based on the work of Bush (1975) to explain the divergence of the Enchenopa binotata complex. Based on present geographical distributions (Metcalf and Wade, 1965), host plant and seasonal records in Costa Rica (Ballou, 1936) and observations by one of us (T.K.W.), the original stock was postulated to be tropical, polyphagous and multivoltine. As Enchenopa progenitors colonized north temperate climates they encountered deciduous hosts which promoted selection for coordination of treehopper life history