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The Neotropical genus Averrhoidium (Sapindaceae) is classified in Doratoxyleae (Radlkofer 1934) and comprises four species with disjunct distribution: Averrhoidium dalyi is found in east Peru and northwest Brazil, Averrhoidium gardnerianum in east Brazil, Averrhoidium paraguaiense in Paraguay, and Averrhoidium spondioides in west Mexico. All specie...
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Sapindaceae Jussieu is a family of flowering plants in the Order Sapindales. Members of the family exist as trees and shrubs, and tendril-bearing vines with about 140-150 genera and 1400-2000 species worldwide. They are economically, medicinally and aesthetically useful. This research aimed at exploring the diversity of Sapindaceae in West and Cent...
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... At the centre of this short style a large canal is formed (Peck and Lersten 1991a). In Dodonaeoideae, Weckerle and Rutishauser (2003) showed in detail a mostly syncarpous gynoecium for four species of Averrhoidium (Fig. 1a, e, ii, iii; Table 2). At the level of the ovary, an outer median dorsal furrow is present on each carpel, but these do not represent zones of fusion but early developing loculicidal dehiscence lines. ...
... Averrhoidium (Dodonaeoideae) and Koelreuteria (Paullinieae) share similarities regarding the PTTT organization in the distal part of the stigma, comparable to Nitrariaceae (see above). Although not equally described in all studies, the PTTT in these Sapindaceae taxa represents a system of three external lateral papillate areas of the stigma continuous with the inner PTTT on the lateral and ventral sides of the three postgenitally united asymplicate carpels (Avalos et al. 2019;Ronse De Craene et al. 2000;Weckerle and Rutishauser 2003). So, this continuity between the outer papillate areas and inner PTTT can be considered a compitum that is partially external and internal at this level, as similarly found in Nitrariaceae. ...
... From the base of the stigma and style (symplicate zone) an internal compitum also develops in Averrhoidium and K. elegans, since the three stylar canals (PTTT) alternate with the locular cavities. All these canals and cavities are contiguous at the centre, forming a central and relatively wide canal full of secretion in the style (Avalos et al. 2019;Weckerle and Rutishauser 2003 ; Table 3). However, in K. paniculata the PTTT form three narrow canals that alternate with three locular furrows but do not form a wide canal. ...
... At the centre of this short style a large canal is formed (Peck and Lersten 1991a). In Dodonaeoideae, Weckerle and Rutishauser (2003) showed in detail a mostly syncarpous gynoecium for four species of Averrhoidium (Fig. 1a, e, ii, iii; Table 2). At the level of the ovary, an outer median dorsal furrow is present on each carpel, but these do not represent zones of fusion but early developing loculicidal dehiscence lines. ...
... Averrhoidium (Dodonaeoideae) and Koelreuteria (Paullinieae) share similarities regarding the PTTT organization in the distal part of the stigma, comparable to Nitrariaceae (see above). Although not equally described in all studies, the PTTT in these Sapindaceae taxa represents a system of three external lateral papillate areas of the stigma continuous with the inner PTTT on the lateral and ventral sides of the three postgenitally united asymplicate carpels (Avalos et al. 2019;Ronse De Craene et al. 2000;Weckerle and Rutishauser 2003). So, this continuity between the outer papillate areas and inner PTTT can be considered a compitum that is partially external and internal at this level, as similarly found in Nitrariaceae. ...
... From the base of the stigma and style (symplicate zone) an internal compitum also develops in Averrhoidium and K. elegans, since the three stylar canals (PTTT) alternate with the locular cavities. All these canals and cavities are contiguous at the centre, forming a central and relatively wide canal full of secretion in the style (Avalos et al. 2019;Weckerle and Rutishauser 2003 ; Table 3). However, in K. paniculata the PTTT form three narrow canals that alternate with three locular furrows but do not form a wide canal. ...
Sapindales is a monophyletic order within the malvid clade of rosids. It represents an interesting group to address questions on floral structure and evolution due to a wide variation in reproductive traits. This review covers a detailed overview of gynoecium features, as well as a new structural study based on Trichilia pallens (Meliaceae), to provide characters to support systematic relationships and to recognize patterns of variations in gynoecium features in Sapindales. Several unique and shared characteristics are identified. Anacrostylous and basistylous carpels may have evolved multiple times in Sapindales, while ventrally bulging carpels are found in pseudomonomerous Anacardiaceae. Different from previous studies, similar gynoecium features, including degree of syncarpy, ontogenetic patterns, and PTTT structure, favors a closer phylogenetic proximity between Rutaceae and Simaroubaceae, or Rutaceae and Meliaceae. An apomorphic tendency for the order is that the floral apex is integrated in the syncarpous or apocarpous gynoecium, but with different length and shape among families. Nitrariaceae shares similar stigmatic features and PTTT structure with many Sapindaceae. As the current position of both families in Sapindales is uncertain, floral features should be investigated more extensively in future studies. Two different types of gynophore were identified in the order: either derived from intercalary growth below the gynoecium as a floral internode, or by extension of the base of the ovary locules as part of the gySapindales is a monophyletic order within the malvid clade of rosids. It represents an interesting group to address questions on floral structure and evolution due to a wide variation in reproductive traits. This review covers a detailed overview of gynoecium features, as well as a new structural study based on Trichilia pallens (Meliaceae), to provide characters to support systematic relationships and to recognize patterns of variations in gynoecium features in Sapindales. Several unique and shared characteristics are identified. Anacrostylous and basistylous carpels may have evolved multiple times in Sapindales, while ventrally bulging carpels are found in pseudomonomerous Anacardiaceae. Different from previous studies, similar gynoecium features, including degree of syncarpy, ontogenetic patterns, and PTTT structure, favors a closer phylogenetic proximity between Rutaceae and Simaroubaceae, or Rutaceae and Meliaceae. An apomorphic tendency for the order is that the floral apex is integrated in the syncarpous or apocarpous gynoecium, but with different length and shape among families. Nitrariaceae shares similar stigmatic features and PTTT structure with many Sapindaceae. As the current position of both families in Sapindales is uncertain, floral features should be investigated more extensively in future studies. Two different types of gynophore were identified in the order: either derived from intercalary growth below the gynoecium as a floral internode, or by extension of the base of the ovary locules as part of the gynoecium. Sapindales share a combination of gynoecial characters but variation is mostly caused by different degrees of development of the synascidiate part relative to the symplicate part of carpels, or the latter part is absent. Postgenital fusion of the upper part of the styles leads to a common stigma, while stylar lobes may be separate. Due to a wide variation in these features, a new terminology regarding fusion is proposed to describe the gynoecium of the order.noecium. Sapindales share a combination of gynoecial characters but variation is mostly caused by different degrees of development of the synascidiate part relative to the symplicate part of carpels, or the latter part is absent. Postgenital fusion of the upper part of the styles leads to a common stigma, while stylar lobes may be separate. Due to a wide variation in these features, a new terminology regarding fusion is proposed to describe the gynoecium of the order.
... The micropyle can be endostomic (Dodonaea Mill., Delavaya Franch., Cardiospermum, Paullinia, Serjania, Urvillea, Allophylus L., and Magonia A. St. Hil., Karkare-Khushalani and Mulay, 1964;Weckerle and Rutishauser, 2005;Cao et al., 2014;González et al., 2014González et al., , 2017 or amphistomic (Filicium Thwaites, Gulati and Mathur, 1977). However, an amphistomic micropyle of zigzag shape, as was observed here, has also been reported in Averrhoidium and within many other lineages of the Sapindales (Weckerle and Rutishauser, 2003;reviewed in Endress and Matthews, 2006;Bachelier and Endress, 2009;Bachelier et al., 2011). The obturator prevails in Sapindaceae, and it also occurs in Acer and many Rutaceae (Weckerle and Rutishauser, 2005;Zhou et al., 2004;Zhou and Liu, 2012;Ramp, 1988, Table 1). ...
Anther and gynoecium structure and embryological information in Koelreuteria and Sapindaceae as a whole remain understudied, as well as the evolution of imperfect flowers in the latter. The aims of this study were to analys in K. elegans subsp. formosana the anther and gynoecium structure and the development of male and female gametophytes in the two floral morphs of putatively imperfect flowers. Standard techniques were applied for LM and SEM. Compared to the normal anther development in staminate flowers, a delayed programmed cell death of tapetum, septum and middle layers on the onset of microspore stage result in indehiscent anthers in the functionally pistillate flowers. Orbicules are reported for the first time in Sapindaceae. Gynoecium development in functionally pistillate flowers is normal, whereas in functionally staminate ones a pistillode with degenerated ovules at the tetrad stage is formed. The pollen tube transmitting tract consists of one layer of epithelial cells with a small lumen in the style and ovary. The anatomy of the latter revealed an axile placentation and complete septum. Reproductive characters of Koelreuteria shared with other Sapindaceae taxa are a secretory binucleate tapetum, simultaneous cytokinesis, unicellular stigmatic papillae, Polygonum type megagametophyte with ephemeral antipodals, and ovules campylotropous, with funicular obturator and amphistomic micropyle. Imperfect flowers in the family are triggered by defective sporophytic tissues impacting on the normal development of microspores and megaspores, disrupting embryological events in staminate and pistillate organs at different developmental stages. Reconstructing ancestral character state under parsimony revealed that two ovules per locule, absence of obturator, presence of hypostase, and binucleate tapetum may be regarded as plesiomorphic for Sapindaceae. These analyses have provided information on the evolution of embryological characters in the family and its phylogenetic significance.
... h Detail of nonsecretory tissue in the hermaphrodite flower, showing a glandular trichome (gt). Scale bars: a, b = 1 mm; c-e = 250 μm; f, g = 500 μm; and h = 100 μm morphology (Weckerle and Rutishauser 2003;present study). The genus Dodonaea s.l. ...
... In K. elegans, the fleshy crest at the base of the petal blade and the stamen filament trichomes offset the absence of the appendages (Avalos et al. 2016). However, exposed nectaries are mostly observed in genera lacking petal appendages, such as Diplokeleba and Magonia of the subfamily Dodonaeoideae, and were also noted in the apetalous flowers of Averrhoidium, Diplopeltis, Llagunoa (Dodonaeoideae) (Weckerle and Rutishauser 2003;present study), and Allophylastrum Acev-Rodr. (Paullinioideae) . ...
We investigated the morphology and structure of the floral nectary in 11 Neotropical genera belonging to the subfamilies Dodonaeoideae and Paullinioideae (Sapindaceae) from southern South America representing three tribes (Dodonaeaeae, Paullinieae, and Melicocceae), in relation to other floral traits in species with contrasting morphological flower characteristics. Nectary organization was analyzed under light, stereoscopic, and scanning electron microscopes; Diplokeleba floribunda N.E. Br. was also observed using transmission electron microscopy. Our comparative data may contribute to the understanding of floral nectary evolution and systematic value in this family. The nectaries were studied in both staminate and pistillate flowers. All the floral nectaries are typical of Sapindaceae: extrastaminal, receptacular, structured, and persistent. The anatomical analysis revealed a differentiated secretory parenchyma and an inner non-secretory parenchyma; the nectary is supplied by phloem traces and, less frequently, by phloem and xylem traces. Nectar is secreted through nectarostomata of anomocytic type. The anatomical analysis showed the absence of nectary in the three morphs of Dodonaea viscosa flowers. Nectary ultrastructure is described in D. floribunda. In this species, the change in nectary color is related to progressive accumulation of anthocyanins during the functional phase. We found relatively small variation in the nectary structural characteristics compared with large variation in nectary morphology. The latter aspect agreed with the main infrafamilial groupings revealed by recent phylogenetic studies, so it is of current valuable systematic importance for Sapindaceae. In representatives of Paullinieae, the reduction of the floral nectary to 4–2 posterior lobes should be interpreted as a derived character state.
... Both sepal and petal whorls arise in a spiral sequence (which is considerably shortened in the petals). A similar transition between sepals and petals is observed in Averrhoidium Baillon (1874: 244) where the petal number fluctuates between one and four (Weckerle & Rutishauser 2003). The flower symmetry of Eurycorymbus cavaleriei is oblique monosymmetric at mid-developmental stages and there is a shift in the symmetry towards polysymmetry during later development. ...
Eurycorymbus is an unusual monotypic genus of Sapindaceae endemic to China. The floral organogenesis of E. cavaleriei has been investigated with the light and scanning electron microscope to clarify its floral characters and its uncertain systematic position. Eurycorymbus cavaleriei possesses several unique characteristics in Sapindaceae. Flowers are unisexual. Sepals initiate in a spiral sequence, followed by five alternating petals which resemble sepals at maturity. The sequential initiation of eight stamens partly overlaps with that of the petals. Three carpellary lobes are simultaneously demarcated and merge gradually to form a tricarpellate ovary. Mature stamens of male flowers have long and smooth filaments, which are folded in buds. In female flowers only one or two ovules per ovary develop into seeds. The extrastaminal nectary forms doubly scalloped lobes. Symmetry changes from oblique monosymmetry at a mid-developmental stage to polysymmetry at maturity. It is suggested that monosymmetry might precede polysymmetry in the evolution of Sapindaceae.
... Only rarely do inner perianth parts also have long plastochrons, and then the transition from spiral to whorled phyllotaxis is delayed. Such delay occurs sporadically in the core eudicots such as in Paeoniaceae ( Paeonia , Hiepko, 1965a ), Dilleniaceae ( Dillenia , Endress, 1997 ), Sapindaceae ( Averrhoidium , Weckerle and Rutishauser, 2003 ), Cactaceae ( Pereskia , Leins and Schwitalla, 1985 ), Phytolaccaceae ( Phytolacca , Ronse Decraene et al., 1997 , clearly seen in their fi g. 3B , although they interpret the pattern differently), Theaceae ( Stewartia , Erbar, 1986 ; Hartia , Tsou, 1998 ). ...
Angiosperms and their flowers have greatly diversified into an overwhelming array of forms in the past 135 million years. Diversification was shaped by changes in climate and the biological environment (vegetation, interaction with other organisms) and by internal structural constraints and potentials. This review focuses on the development and structural diversity of flowers and structural constraints. It traces floral diversification in the different organs and organ complexes (perianth, androecium, gynoecium) through the major clades of extant angiosperms. The continuously improved results of molecular phylogenetics provide the framework for this endeavor, which is necessary for the understanding of the biology of the angiosperms and their flowers. Diversification appears to work with innovations and modifications of form. Many structural innovations originated in several clades and in special cases could become key innovations, which likely were hot spots of diversification. Synorganization between organs was an important process to reach new structural levels, from which new diversifications originated. Complexity of synorganization reached peaks in Orchidaceae and Apocynaceae with the independent evolution of pollinaria. Such a review throughout the major clades of angiosperms also shows how superficial and fragmentary our knowledge on floral structure in many clades is. Fresh studies and a multidisciplinary approach are needed.
... Weckerle and Rutishauser (2005) studied the gynoecium, fruit and seed structures and ontogeny in selected species of Paullinieae. Further studies on other tribes of the family are those of Ronse Decraene et al. (2000) and Weckerle and Rutishauser (2003). ...
... The nectaries here studied belong to the axial persistent type (Smets & Cresens 1988, Smets 1989). Weckerle and Rutishauser (2003) mention the presence of a persistent extrastaminal flower nectary disk in Averrhoidium (Sapindaceae, Doratoxyleae). From an evolutionary point of view the persistent nectaries are concentrated in the medium-and higher-evolved groups of the dicots (Endress 1994). ...
The morphology, anatomy and ontogeny the floral nectaries of two species of Sapindaceae (Paullinieae), viz. Cardiospermum grandiflorum and Urvillea chacoensis, were studied using light and scanning electron microscopy. Both species are monoecious with oblique monosymmetric flowers. The floral nectaries are located at the base of the androgynophore. Vascular tissue supplying the nectaries consists of phloem. Cardiospermum grandiflorum has a posterior nectary with two lobes without any specially differentiated tissue, and with nectarostomata. In Urvillea chacoensis the nectary has four lobes; two posterior ones with distinctive nectariferous tissue with nectarostomata on the abaxial side, and two minor anterior ones that are histologically similar but lack nectarostomata. The nectaries in both species have a common base with the petals. Since the nectaries are macroscopically differentiated they are considered as structural nectaries.
... In other Sapindales, flowers are also largely isomerous and polysymmetric (Meliaceae, Rutaceae and Simaroubaceae). Largely monosymmetric flowers with oblique monosymmetry are reported in Sapindaceae (Weckerle & Rutishauser, 2003, 2005Endress & Matthews, 2006b;Ronse de Craene & Haston, 2006). Figure 41. ...
... The role of the stigmatic head in the formation of an external compitum has only been studied in some Rutaceae among Sapindales (Ramp, 1988). A papillate stigmatic surface is common in Sapindales and also present in all other families, including Meliaceae (Gouvêa et al., 2008a, b), Rutaceae and Simaroubaceae (Ramp, 1988), Sapindaceae (Weckerle & Rutishauser, 2003, 2005 and Nitrariaceae (Ronse de Craene & Smets, 1991), but in most cases it is unknown whether the papillae are uni-or multicellular. Stigmas with uniseriate, multicellular papillae as in Burseraceae and Anacardiaceae are rare in angiosperms and occur elsewhere in Sapindales only in Kirkia (Kirkiaceae) (Bachelier & Endress, 2008). ...
... Kirkiaceae, Meliaceae, Simaroubaceae or Rutaceae) than of the syntropous ovules of Anacardiaceae. Two axile collateral placentae bearing two ovules as in Burseraceae are not so common in other Sapindales, whereas two axile placentae that are (almost) collateral with the two ovules becoming superposed, as in Beiselia, are known in Meliaceae (Wiger, 1935), Sapindaceae (Weckerle & Rutishauser, 2003) and Kirkiaceae (Bachelier & Endress, 2008). Bitegmic, antitropous and slightly campylotropous ovules with a longer and (sometimes) thicker inner integument and (long) S-shaped or zig-zag micropyle as in Burseraceae are also found in Kirkiaceae (Bachelier & Endress, 2008), Meliaceae, Simaroubaceae and Rutaceae (Wiger, 1935;Narayana, 1957Narayana, , 1958Narayana, 1960b;Nair & Sukumaran, 1960;Prakash, Lim & Manurung, 1977). ...
Anacardiaceae and Burseraceae are traditionally distinguished by the number of ovules (1 vs. 2) per locule and the direction of ovule curvature (syntropous vs. antitropous). Recent molecular phylogenetic studies have shown that these families are sister groups in Sapindales after having been separated in different orders for a long time. We present a comparative morphological study of the flower structure in both families. The major clades, usually supported in molecular phylogenetic analyses, are well supported by floral structure. In Anacardiaceae, there is a tendency to gynoecium reduction to a single fertile carpel (particularly in Anacardioideae). The single ovule has a long and unusually differentiated funicle, which connects with the stylar pollen tube transmitting tract in all representatives studied. In Anacardiaceae–Spondiadoideae, there is a tendency to form an extensive synascidiate zone, with a massive remnant of the floral apex in the centre; these features are also present in Beiselia (Burseraceae) and Kirkiaceae (sister to Anacardiaceae plus Burseraceae) and may represent a synapomorphy or apomorphic tendency for the three families. In core Burseraceae, gynoecium structure is much less diverse than in Anacardiaceae and has probably retained more plesiomorphies. Differences in proportions of parts of the ovules in Anacardiaceae and Burseraceae are linked with the different direction of ovule curvature. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159, 499–571.
... In other Sapindales, flowers are also largely isomerous and polysymmetric (Meliaceae, Rutaceae and Simaroubaceae). Largely monosymmetric flowers with oblique monosymmetry are reported in Sapindaceae (Weckerle & Rutishauser, 2003, 2005Endress & Matthews, 2006b;Ronse de Craene & Haston, 2006). Figure 41. ...
... The role of the stigmatic head in the formation of an external compitum has only been studied in some Rutaceae among Sapindales (Ramp, 1988). A papillate stigmatic surface is common in Sapindales and also present in all other families, including Meliaceae (Gouvêa et al., 2008a, b), Rutaceae and Simaroubaceae (Ramp, 1988), Sapindaceae (Weckerle & Rutishauser, 2003, 2005 and Nitrariaceae (Ronse de Craene & Smets, 1991), but in most cases it is unknown whether the papillae are uni-or multicellular. Stigmas with uniseriate, multicellular papillae as in Burseraceae and Anacardiaceae are rare in angiosperms and occur elsewhere in Sapindales only in Kirkia (Kirkiaceae) (Bachelier & Endress, 2008). ...
... Kirkiaceae, Meliaceae, Simaroubaceae or Rutaceae) than of the syntropous ovules of Anacardiaceae. Two axile collateral placentae bearing two ovules as in Burseraceae are not so common in other Sapindales, whereas two axile placentae that are (almost) collateral with the two ovules becoming superposed, as in Beiselia, are known in Meliaceae (Wiger, 1935), Sapindaceae (Weckerle & Rutishauser, 2003) and Kirkiaceae (Bachelier & Endress, 2008). Bitegmic, antitropous and slightly campylotropous ovules with a longer and (sometimes) thicker inner integument and (long) S-shaped or zig-zag micropyle as in Burseraceae are also found in Kirkiaceae (Bachelier & Endress, 2008), Meliaceae, Simaroubaceae and Rutaceae (Wiger, 1935;Narayana, 1957Narayana, , 1958Narayana, 1960b;Nair & Sukumaran, 1960;Prakash, Lim & Manurung, 1977). ...
Anacardiaceae and Burseraceae are traditionally distinguished by the number of ovules (1 vs. 2) per locule and the direction of ovule curvature (syntropous vs. antitropous). Recent molecular phylogenetic studies have shown that these families are sister groups in Sapindales after having been separated in different orders for a long time. We present a comparative morphological study of the flower structure in both families. The major clades, usually supported in molecular phylogenetic analyses, are well supported by floral structure. In Anacardiaceae, there is a tendency to gynoecium reduction to a single fertile carpel (particularly in Anacardioideae). The single ovule has a long and unusually differentiated funicle, which connects with the stylar pollen tube transmitting tract in all representatives studied. In Anacardiaceae–Spondiadoideae, there is a tendency to form an extensive synascidiate zone, with a massive remnant of the floral apex in the centre; these features are also present in Beiselia (Burseraceae) and Kirkiaceae (sister to Anacardiaceae plus Burseraceae) and may represent a synapomorphy or apomorphic tendency for the three families. In core Burseraceae, gynoecium structure is much less diverse than in Anacardiaceae and has probably retained more plesiomorphies. Differences in proportions of parts of the ovules in Anacardiaceae and Burseraceae are linked with the different direction of ovule curvature.
... In other Sapindales, flowers are also largely isomerous and polysymmetric (Meliaceae, Rutaceae and Simaroubaceae). Largely monosymmetric flowers with oblique monosymmetry are reported in Sapindaceae (Weckerle & Rutishauser, 2003, 2005Endress & Matthews, 2006b;Ronse de Craene & Haston, 2006). Figure 41. ...
... The role of the stigmatic head in the formation of an external compitum has only been studied in some Rutaceae among Sapindales (Ramp, 1988). A papillate stigmatic surface is common in Sapindales and also present in all other families, including Meliaceae (Gouvêa et al., 2008a, b), Rutaceae and Simaroubaceae (Ramp, 1988), Sapindaceae (Weckerle & Rutishauser, 2003, 2005 and Nitrariaceae (Ronse de Craene & Smets, 1991), but in most cases it is unknown whether the papillae are uni-or multicellular. Stigmas with uniseriate, multicellular papillae as in Burseraceae and Anacardiaceae are rare in angiosperms and occur elsewhere in Sapindales only in Kirkia (Kirkiaceae) (Bachelier & Endress, 2008). ...
... Kirkiaceae, Meliaceae, Simaroubaceae or Rutaceae) than of the syntropous ovules of Anacardiaceae. Two axile collateral placentae bearing two ovules as in Burseraceae are not so common in other Sapindales, whereas two axile placentae that are (almost) collateral with the two ovules becoming superposed, as in Beiselia, are known in Meliaceae (Wiger, 1935), Sapindaceae (Weckerle & Rutishauser, 2003) and Kirkiaceae (Bachelier & Endress, 2008). Bitegmic, antitropous and slightly campylotropous ovules with a longer and (sometimes) thicker inner integument and (long) S-shaped or zig-zag micropyle as in Burseraceae are also found in Kirkiaceae (Bachelier & Endress, 2008), Meliaceae, Simaroubaceae and Rutaceae (Wiger, 1935;Narayana, 1957Narayana, , 1958Narayana, 1960b;Nair & Sukumaran, 1960;Prakash, Lim & Manurung, 1977). ...
