POLLEN MORPHOLOGY AND THE RELATIONSHIPS OF AËTOXYLON, AMYXA, AND GONYSTYLUS TO THE THYMELAEACEAE

POLLEN MORPHOLOGY AND THE RELATIONSHIPS OF AËTOXYLON, AMYXA, AND GONYSTYLUS TO THE THYMELAEACEAE
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花粉形态以及 AËtoxylon、Amyxa 和 Gonystylus 与瑞香科的关系

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发表时间:
1985
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通讯作者:
J. Skvarla
J. Skvarla
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作者:
J. W. Nowicke;V. Patel;J. Skvarla

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在 SEM 和 TEM 中将三个密切相关的属 Aetoxylon、Amyxa 和 Gonystylus 的花粉与 Thymelaeaceae, s. 的花粉进行了比较。 s。瑞香科植物具有球形、全孔的颗粒,具有巴豆状顶盖,其中基本亚基形状为三角形并形成连续的三角形阵列。薄片(TEM)和断裂(SEM)显示这些亚基附着在水平杆的环状网络上。在瑞香科中,三角形亚基的细分数量和融合程度各不相同,并形成形态连续体。 Aetoxylon、Amyxa 和 Gonystylus 也有球形、全孔花粉,但其顶盖中几乎所有亚基的区别似乎都已丧失。然而,Aetoxylon、Amyxa 和 Gonystylus 的外壁结构迄今为止在被子植物中是独一无二的。薄切片显示出厚厚的顶盖,上面有一层短的甚至颗粒状的小柱,然后是一层薄的、不连续的层,较大的小柱似乎悬挂在上面。即使在孔区域也没有内壁的证据。独特的外壁结构支持将 Aetoxylon、Amyxa 和 Gonystylus 作为独立的科(Gonystylaceae)进行处理,与瑞香科(Thymelaeaceae)有联系。 AETOXYLON (Airy Shaw) Airy Shaw 和 Amyxa Van Tiegh. 在婆罗洲各有一个种,而 Gonystylus 有 25 种,分布在马来西亚、所罗门群岛和斐济的大部分地区,已被 Hutchinson (1959) 视为一个单独的科,Gonystylaceae,或被 Wagenitz 的 Domke (1934) 纳入 Thymelaeaceae (1964)、Airy Shaw (Willis, 1966) 和 Cronquist (1981),他们都承认这三个属的区别,将它们划分为 Gonystyloideae 亚科。 Aetoxylon、Amyxa 和 Gonystylus 可以通过以下特征的组合与其余瑞香科植物区分开来:蒴果果实、花瓣众多、花柱细长和透明的点状叶子。这些属也缺乏内部韧皮部,而这种内部韧皮部是大量 t 的特征。 1984 年 8 月 6 日收到出版;修订于 1985 年 2 月 5 日接受。我们感谢 J. L. Bittner、S. G. Braden 和 W. Chissoe 提供的出色技术援助。我们还感谢已故 Jan Muller 博士寄来的花粉样本。这项研究得到了 S. Dillon Ripley 流体研究中心的资助以及国家科学基金会资助 BSR-8002585 和 BSR-8216349 的部分支持。大多数瑞香科,以及大多数香桃木目。 Archangelsky (1971) 描述了瑞香科及其近缘科的花粉形态 (1971),将其主要的巴豆状图案称为星状。他认为大多数瑞香科植物都具有典型的星状雕刻,但将膝柱科描述为具有隐星状雕刻,并引用“其外壁的绝对特殊类型作为将其视为独立科的进一步标准”。更有趣的是,Archangelsky 认为星状雕刻是简单网状雕刻的复杂变量之一。作为桃金娘目(Patel、Skvarla 和 Raven,1983、1984)和大戟目(sensu Cronquist,1981;Nowicke,1984;Nowicke 和 Skvarla,1984)花粉形态广泛研究的一部分,对瑞香科 39 属 70 多个物种的花粉进行了检查。包括 Aetoxylon sympetalum、Amyxa pluriconis 和四种 Gonystylus(见表 1)。
The pollen of three closely related genera, Aetoxylon, Amyxa, and Gonystylus is compared in SEM and TEM with that of Thymelaeaceae, s. s. The Thymelaeaceae have spherical, pantoporate grains with a crotonoid tectum in which the basic subunit is tnangular in shape and forms a continuous triangular array. Thin section (TEM) and fractures (SEM) revealed that these subunits are attached to a ringlike network of horizontal rods. Within the Thymelaeaceae, the triangular subunits vary in the number of subdivisions and degree of fusion and form a morphological continuum. Aetoxylon, Amyxa, and Gonystylus also have spherical, pantoporate pollen but with a tectum in which almost all of the distinction of the subunits appears to have been lost. The structure of the exine in Aetoxylon, Amyxa, and Gonystylus, however, is unique thus far within the angiosperms. Thin section revealed a thick tectum with a layer of short or even granular columellae, then a thin, discontinuous layer from which larger columellae appear to hang. There is no evidence of an endexine even in the region of the apertures. The distinctive exine structure would support the treatment of Aetoxylon, Amyxa, and Gonystylus as a separate family, Gonystylaceae, allied to the Thymelaeaceae. AETOXYLON (Airy Shaw) Airy Shaw, and Amyxa Van Tiegh., each with a single species in Borneo, and Gonystylus with 25 species distributed through most of Malaysia, the Solomon Islands, and Fiji, have been treated as a separate family, Gonystylaceae, by Hutchinson (1959), or included in the Thymelaeaceae by Domke (1934), Wagenitz (1964), Airy Shaw (Willis, 1966), and Cronquist (1981), all of whom acknowledge the distinction of these three genera by segregating them as a subfamily, Gonystyloideae. Aetoxylon, Amyxa, and Gonystylus can be distinguished from the remaining Thymelaeaceae by a combination of characters: capsule fruit, numerous petals, elongate style, and pellucid-punctate leaves. These genera also lack the internal phloem that characterizes the vast t Received for publication 6 August 1984; revision accepted 5 February 1985. We thank J. L. Bittner, S. G. Braden, and W. Chissoe for their fine technical assistance. We are also grateful for the pollen samples sent by the late Dr. Jan Muller. This study was supported in part by a grant from Fluid Research from S. Dillon Ripley, and National Science Foundation Grants BSR-8002585 and BSR-8216349. majority ofthe Thymelaeaceae, as well as most of the order Myrtales. Archangelsky (1971 ), who described the pollen morphology of the Thymelaeaceae and allied families (1971), referred to their predominant crotonoid pattern as stellate. He considered the majority of the Thymelaeaceae to have typical stellate sculpturing, but described the Gonystylaceae as having cryptostellate sculpturing, and cited the "absolutely special type of their exines as a further criterion" for their treatment as an independent family. Of further interest, Archangelsky suggested that stellate sculpturing was one of the complex variables of simple reticulate sculpturing. As part of an extensive study of pollen morphology in the orders Myrtales (Patel, Skvarla and Raven, 1983, 1984) and Euphorbiales (sensu Cronquist, 1981; Nowicke, 1984; Nowicke and Skvarla, 1984), the pollen of more than 70 species representing 39 genera in the Thymelaeaceae have been examined. Aetoxylon sympetalum, Amyxa pluriconis, and four species of Gonystylus were included (see Table 1).