Pearl Bodies as Ant Food: An Ecological Role for Some Leaf Emergences of Tropical Plants-猫眼课题宝

珍珠体作为蚂蚁食物：热带植物某些叶子出现的生态作用

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1982

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D. O'dowd

珍珠体，单叶或多叶，其特征是有光泽的珍珠状外观；球形或棒状，具底部收缩；容易从植物上脱离；储存相对大量的脂类；有有限的大小范围，通常为0.5至3.0毫米长。它们广泛分布于双子叶植物亚科中，已报道有19科50属，均具有热带或亚热带的亲缘关系。有几条证据表明这些结构与植物-蚂蚁的关系有关；这些证据是：在属的水平上，珍珠体的存在通常与植物的其他与蚂蚁有关的特征有关(48%)，如具角蜜腺和花外蜜腺；当存在洞口或花外蜜腺时，珍珠体的产生在时间和空间上是平行的；以及已知蚂蚁从叶子中收集珍珠体并将它们送回巢中。这些观察和关联表明，珍珠体在许多热带植物-蚂蚁互惠关系中发挥着重要作用。最近对植物-蚂蚁相互作用的田间研究将注意力重新集中在一个古老的争议上：花外花蜜(EFN)的作用。证据表明，EFN的产生决定了蚂蚁在植物上觅食的模式(Bentley 1977a，O‘Dowd 1979，Pickett和Clark 1979)，并可能导致蚂蚁与植物草食动物和种子捕食者发生干扰，以及维持有利的植物营养环境(Bentley 1977a，Inouye and Taylor 1979，Kleinfeldt 1978)。在植物-蚂蚁相互作用具有高度互惠专业性的地方，例如仙人掌-假蚂蚁；Cecropia-Azteca；和Macaranga-Firematogaster，强调不同的食物来源。在这些协会中，食物体(分别是贝尔体、穆勒体和贝卡体)由蚂蚁收获，并在蚂蚁群体的福祉和繁殖中发挥重要作用(Janzen 1966,1969；Rickso‘n 1980)。珍珠体在许多方面与这些蚂蚁食品体相似，在许多热带物种的叶子和枝条上产生。虽然相思的贝尔体实际上是改良的叶尖(Rickson 1969)，并显示出与整个珍珠体的发育差异，但Macaranga spp.蚂蚁食体的解剖和发育。和Cecropia Splp.与珍珠体的描述一致(Meyen 1837，Holmgren 1911)。事实上，早期的研究人员将这些蚂蚁食物体描述为“Perldrusen”(Meyen 1837)、“Perlules”(Rouppert 1926)和“parlharen”(Holmgren 1911)。作为Delpino(1886)提出的EFN分泌作用的推论，Penzig(1892)假设珍珠体作为蚂蚁的食物，是EFN的平行和互补的食物来源。正如他的同时代人，如霍姆格伦(1911)所恰当地指出的那样，不幸的是，彭齐格没有为他的有趣的假设提供证据。我在这里汇集了描述珍珠体的特征和分布的文献，并将其与上个世纪关于EFN和植物-蚂蚁相互作用的大量研究结合在一起。直接和间接的证据都支持彭齐格的观点，即这些叶子的出现具有营养导向和蚂蚁食物的功能。珍珠体的定义和术语Meyen(1837年)首次报道了“珍珠腺”，并将其描述为具有球状和光泽特征的单个或多个突起在枝叶上。根据他最初的观察，它们被不同地命名为珍珠膀胱(de Bary，1877)、珍珠毛发(Holmgren，1911)和珠体(Wheeler，1910)。由于没有证据表明这些叶子的突起是分泌的，我称它们为“珍珠体”，这与梅恩最初的描述一致，但没有被误导地描述为腺体。珍珠体来源不一，形态和发育也各不相同。它们的范围从单细胞到多细胞(图2)。La-f)，来源于表皮和真皮下组织(Penzig，1892)。然而，它们有许多共同之处，例如一个特色、有限尺寸范围1现有地址：南澳大利亚州弗林德斯大学生物科学系，南澳大利亚州贝德福德公园，南澳大利亚州5042。40 Biotropica 14(1)：40-49 1982年此内容于2016年4月23日星期六从157.55.39.201下载，UTC时间05：26：53所有用户均须遵守http://about.jstor.org/terms

Pearl bodies, single or multicelled leaf emergences, are characterized by a lustrous, pearl-like appearance; a spherical or club-like shape with a basal constriction; ease of detachment from the plant; storage of relatively large quantities of lipid; axd a limited size range, usually from 0.5 to 3.0 mm in length. They are broadly distributed in the Dicotyledonae and have been reported in 19 families and 50 genera, all of tropical or subtropical affinity. Several lines of evidence implicate these structures in plant-ant association; these are: at the generic level, the presence of pearl bodies is often associated (48%) with other ant-related features of the plant such as specializel domatia and extrafloral nectaries; when domatia or extrafloral nectaries are present, pearl-body production is parallel in time and space; and ants are known to collect pearl bodies from leaves and return them to their nests. These observations and correlations suggest that pearl bodies play an important role in many tropical plant-ant mutualisms. RECENT FIELD STUDIES OF PLANT-ANT INTERACTIONS have refocused attention on an old controversy: the role of extrafloral nectar (EFN). Evidence indicates that EFN production defines the pattern of ant foraging on the plant (Bentley 1977a, O'Dowd 1979, Pickett and Clark 1979) and can result in interference by attending ants with plant herbivores and seed predators as well as the maintenance of a favorable plant-nutrient environment (Bentley 1977a, Inouye and Taylor 1979, Kleinfeldt 1978). Where there is a high degree of reciprocal specialization in plant-ant interaction, e.g., A cacia-Pseudomyrmex; Cecropia-Azteca; and Macaranga-Crematogaster, a different food resource has been emphasized. In these associations, food bodies (Beltian, Mullerian and Beccarian bodies, respectively) are harvested by the ants and play an essential role in the well-being and reproduction of the ant colonies (Janzen 1966, 1969; Rickso'n 1980). Pearl bodies, similar in many respects to these ant-food bodies, are produced on the leaves and shoots of many tropical species. While the Beltian bodies of Acacia cornigera are actually modified leaf tips (Rickson 1969) and show developmental differences from pearl bodies as a whole, the anatomy and development of the ant-food bodies of Macaranga spp. and Cecropia splp. are consistent with descriptions of pearl bodies (Meyen 1837, Holmgren 1911). In fact, earlier researchers describe these ant-food bodies as "Perldrusen" (Meyen 1837), "perlules" (Rouppert 1926), and "parlharen" (Holmgren 1911). As a corollary to the role of EFN secretion suggested by Delpino (1886), Penzig (1892) hypothesized that pearl bodies serve as ant-food and represent a parallel and co'mplementary food resource to EFN. As was aptly pointed out by his contemporaries, e.g., Holmgren (1911), Penzig unfortunately provided no evidence for his interesting hypothesis. I bring together here the literature describing the characteristics and distribution of pearl bodies and couple it to the substantial research accrued over the last century on EFN and plant-ant interactions. Evidence, both direct and circumstantial, supports Penzig's contention that these leaf emergences are nutritionally directed and function as ant food. DEFINITION AND TERMINOLOGY OF PEARL BODIES Meyen (1837) first reported "pearl glands" and described them as single or multicelled emergences on shoots and leaves with a characteristic spherical shape and lustrous aspect. Following his initial observations they have been variously termed pearl bladders (De Bary 1877), pearl hairs (Holmgren 1911), and bead bodies (Wheeler 1910). Since there is no evidence that these leaf emergences are secretory, I term them "pearl bodies" consistent with Meyen's initial description but without the misleading description as glands. Pearl bodies are of heterogeneous origin and differ in morphology and development. They range from single to multicelled (fig. la-f) and are derived from both epidermal and subepidermal tissues (Penzig 1892). Nevertheless, they share many common features such as a characteristic, limited size range 1Present address: Department of Biological Sciences, The Flinders University of South Australia, Bedford Park, South Australia 5042, Australia. 40 BIOTROPICA 14(1): 40-49 1982 This content downloaded from 157.55.39.201 on Sat, 23 Apr 2016 05:26:53 UTC All use subject to http://about.jstor.org/terms

Pearl Bodies as Ant Food: An Ecological Role for Some Leaf Emergences of Tropical Plants

珍珠体作为蚂蚁食物：热带植物某些叶子出现的生态作用

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