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The Embryology and Systematic Relationships of Prionium Serratum (Juncaceae: Juncales) Author(S): Sioban L

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The Embryology and Systematic Relationships of Prionium Serratum (Juncaceae: Juncales) Author(S): Sioban L The Embryology and Systematic Relationships of Prionium serratum (Juncaceae: Juncales) Author(s): Sioban L. Munro and H. Peter Linder Source: American Journal of Botany , Jun., 1997, Vol. 84, No. 6 (Jun., 1997), pp. 850- 860 Published by: Wiley Stable URL: https://www.jstor.org/stable/2445821 JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at https://about.jstor.org/terms Wiley is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:55:19 UTC All use subject to https://about.jstor.org/terms American Journal of Botany 84(6): 850-860. 1997. THE EMBRYOLOGY AND SYSTEMATIC RELATIONSHIPS OF PRIONIUM SERRATUM (JUNCACEAE: JUNCALES)1 SIOBAN L. MUNRO2 AND H. PETER LINDER Botany Department, University of Cape Town, Rondebosch 7700, Cape Town, South Africa Although Prionium is included in Juncaceae, rbcL sequence data indicate that Juncaceae is paraphyletic, with most genera closer to Cyperaceae than to Prionium. Cyperaceae and Juncales have embryological synapomorphies: thus embryology is used to test the monophyly of Juncaceae. The embryology of Prionium is described and its systematic position discussed. Material was prepared using standard methods of paraffin embedding. Additional embryological data were extracted from the literature. The anther in Prionium is tetrasporangiate, and the wall has an epidermis, an endothecium and middle layer, and an irregularly bilayered, glandular secretory tapetum. Microsporogenesis is probably simultaneous; pollen is ulcerate with a granular exine, in tetrahedral and cross tetrads, and trinucleate at release. The trilocular ovary contains many cras- sinucellate ovules probably having a Polygonum-type embryo sac. Endosperm is helobial and the embryo is of the Onagrad type, Juncus variation. The seed is testal-tegmic and germination is epigeal. The embryology of Prionium is most like that of Juncaceae, which shares several synapomorphies with Cyperaceae. Some of the characters in Cyperaceae may be inter- preted as specialized forms of those found in Juncaceae. Embryology supports the monophyly of Cyperaceae and Juncales, but not Juncaceae; thus the position of Prionium remains unresolved. Key words: Cyperaceae; embryology; Eriocaulaceae; Flagellariaceae; Juncaceae; Prionium serratum; Typhaceae. Prionium, a monotypic genus, is restricted to South in Juncaceae are closer to Cyperaceae than to Prionium, Africa where it occurs on oligotrophic soils along streams thus suggesting that Juncaceae sensu Dahlgren, Clifford, in the South Western Cape, extending along the coast into and Yeo (1985) are paraphyletic. This is corroborated by southern Kwazulu/Natal. Prionium has a woody, decum- the peculiar leaf anatomy of Prionium (Cutler, 1969) and bent rhizome, which is clothed in old sheathing leaf bases the presence of flavone C-glycosides found only in and is topped by a crown of serrate, tristichously inserted Prionium among members of Juncaceae (Williams and leaves. In addition, the plant often grows to several me- Harborne, 1975). tres in height. This growth form is bizarre in comparison Embryological data provide numerous synapomor- to other members of Juncaceae, which are usually small phies for both Cyperaceae and the Juncales (Linder and and herbaceous. Kellogg, 1995) and embryology therefore has the poten- Prionium forms dense monospecific stands, which usu- tial to test the monophyly of Juncaceae sensu lato. As ally grow in the beds of streams and rivers. The mass of Prionium is embryologically unknown, this study de- woody rhizomes act to bind the soil, thereby building up scribes the embryology of Prionium serratum (L.f.) Dre- river beds, ameliorating flooding events, and filtering wa- ge ex E. Mey and discusses its implications for the sys- ter. Prionium produces numerous seeds, which germinate tematic relationships of Prionium. in open spaces along river banks, but is also able to form new plants by budding along the woody rhizome. MATERIALS AND METHODS There is little morphological information available for Inflorescences of P. serratum at different developmental stages were Prionium except with regard to anatomy, which has been collected from three localities in the South Western Cape, South Africa well documented by Cutler (1969) and vascular construc- (Table 1) and fixed in formalin-acetic acid-ethanol (FAA) (17 ethanol: tion and development, which is covered by Zimmerman 2 formalin: 1 acetic acid vlv). Flowers and seeds were prepared ac- and Tomlinson (1968). The leaf and stem flavonoid cording to standard methods of paraffin embedding and sectioning. Se- chemistry have been surveyed by Williams and Harborne rial sections of 10 p.m were stained in a safranin-fast green combination (1975) for Prionium as well as for Juncaceae. (Johansen, 1940). For observations of microspore nuclei, pollen grains Prionium is usually included in Juncaceae (Cutler, at different stages in development were cleared using Herr's (1971) 1969; Dahlgren and Clifford, 1982; Dahlgren, Clifford, clearing agent [2 lactic acid: 2 chloral hydrate: 2 phenol: 2 clove oil: 1 and Yeo, 1985; Simpson, 1995), a small family of eight xylene (by mass)] for 12-18 h. Pollen and seeds were also sputter- to ten genera, which is restricted to the southern hemi- coated with gold-palladium for scanning electron microscopy (SEM). sphere, except for the cosmopolitan genera Juncus and Embryological data on Juncaceae and related families viz. Cyperaceae Luzula. On the other hand, rbcL sequence data (Chase et (Juncales), Thurniaceae (Juncales), Flagellariaceae (Poales), Eriocaula- al., 1993; Plunkett et al., 1995) indicate that most genera ceae (Commelinales), and Typhaceae (Typhales) were extracted from the available literature for comparison. I Manuscript received 17 June 1996; revision accepted 8 November 1996. RESULTS The authors wish to thank Andrew Spinks and Tony Verboom for Anther and microspores-The anther is tetrasporan- reading through an earlier draft version and also, Paula Rudall and Dennis Stevenson for critically reviewing this manuscript. This research giate (Fig. 1), the wall comprising a single-layered epi- was supported by the Foundation for Research and Development (FRD). dermis, endothecium, and middle layer and a bilayered, 2 Author for correspondence. uninucleate tapetum of the glandular secretory type (Fig. 850 This content downloaded from 86.59.13.237 on Tue, 06 Jul 2021 08:55:19 UTC All use subject to https://about.jstor.org/terms June 1997] MUNRO AND LINDER-EMBRYOLOGY OF PRIONIUM 851 TABLE 1. Voucher list of specimens collected for embryology. (Fig. 23). The endosperm in the micropylar chamber is initially acellular (Fig. 22), becoming cellular later, at Collector Collector's Locality no. which time the chalazal chamber consists of a few crushed cells and several disorganized nuclei (Fig. 23). Linder 5 770 Palmiet River bridge, S. W. Cape Linder 5 771 Palmiet River, S. W. Cape Linder 5773 Silvermine Nature Reserve, Cape Peninsula Embryogeny-The zygote (Fig. 24) formed through Munro 1 Algeria Forest Station, Clanwilliam, S. W. Cape syngamy divides to form a two-tiered proembryo con- sisting of a terminal cell, CA, and a basal cell, CB (Fig. 25). This develops into a three-celled, two-tiered proem- 2). Anther wall formation was not observed, but the num- bryo when CA divides longitudinally (Figs. 26, 27) and ber of layers present suggests the monocotyledonous further into a four-celled, three-tiered proembryo when type. The middle layer is ephemeral, degenerating shortly CB divides transversely producing two daughter cells, M before the microspores develop into pollen grains. Mi- and CI (Fig. 28). The latter division is the only contri- crosporogenesis is probably simultaneous (Figs. 3-5) bution of the basal cell to embryo development and this with pollen released in tetrahedral (Fig. 6) as well as is thus of the Onagrad type. The two cells of CA divide tetragonal (Fig. 7) tetrads, each grain being ulcerate with longitudinally to form a quadrant, tier Q. This is followed a slightly granular exine (Fig. 6) and three-celled at the by a longitudinal division of M, the middle cell. Tier CI shedding stage (Fig. 8). divides transversely to form two superposed cells N and N', which results in a four-tiered (Q, M, N, and N'), four- Ovary, nucellus, megagametophyte, and integu- celled proembryo (Figs. 29-31). At this stage, the cells ments-The trilocular ovary (Fig. 9) contains numerous of the epidermal initials (tier Q) are cut off without the anatropous ovules borne in two rows on each axile pla- formation of octants (Figs. 32, 33). This pattern of de- centa (Fig. 10). The style is much reduced, such that only velopment is the Juncus variation. Following this, tier Q the three stigmata remain at the apex of the ovary (Fig. undergoes several divisions to form two parts L and L' 11). The surface of each stigma is covered in sessile pa- causing the basal region of the embryo to become pear pillae. shaped (Fig. 34). Tier N' divides transversely forming The ovule is crassinucellate, with an archesporial cell two superposed cells 0 and P, which form the suspensor cutting off a parietal cell (Fig. 12), which undergoes fur- (Fig. 34). At this stage tiers M, L', and L undergo several ther divisions to form the nucellar tissue (Fig. 13). This divisions and increase in size (Fig. 34). Following this, results in the megaspore mother cell being deep seated tiers P and 0 are reduced to a single cell (N'), while tiers within the ovule (Fig. 13). N and M proliferate (Figs. 35, 36). Later in the devel- Although the tetrad stage of megasporogensis was not opment, the suspensor (tiers N' and N) is reduced leaving observed it is likely that the development of the embryo tier M in this region (Figs.
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