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Seeds of Theaceae. A. Gordonia lasianthus (Sheridan & Telford 1116). B. Gordonia brandegeei (Schultes & Reko 798). C. Franklinia alatamaha (Atha 518). D. Schima wallichii (Soepadmo & Suhaimi s76). E. Schima argentea (Fang 5685). F. Gordonia (=Polyspora) axillaris, a: embryo, b: wing (Rock 7299). G. Camellia sinensis (Brach & Palomino 1754). H. Tutcheria greeniae (Steward & Cheo 1045). I. Tutcheria shinkoensis (Boufford and Bartholemew 25109). J. Stewartia pteropetiolata (Tsang 24021). K. Stewartia malacodendron (Mackenzie 1698). L. Stewartia monadelpha (Ahles 35399). M. Stewartia ovata (Kearney 548). See Table I for voucher information. Scale bar = 5 mm. 

Seeds of Theaceae. A. Gordonia lasianthus (Sheridan & Telford 1116). B. Gordonia brandegeei (Schultes & Reko 798). C. Franklinia alatamaha (Atha 518). D. Schima wallichii (Soepadmo & Suhaimi s76). E. Schima argentea (Fang 5685). F. Gordonia (=Polyspora) axillaris, a: embryo, b: wing (Rock 7299). G. Camellia sinensis (Brach & Palomino 1754). H. Tutcheria greeniae (Steward & Cheo 1045). I. Tutcheria shinkoensis (Boufford and Bartholemew 25109). J. Stewartia pteropetiolata (Tsang 24021). K. Stewartia malacodendron (Mackenzie 1698). L. Stewartia monadelpha (Ahles 35399). M. Stewartia ovata (Kearney 548). See Table I for voucher information. Scale bar = 5 mm. 

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Species of Gordonia s.l. are characterized by having seeds with prominent flattened apical wings. However, recent molecular phylogenetic studies show that this concept of Gordonia is not monophyletic, with species occurring in two tribes of Theaceae. We examine seed coat micromophology of 14 species of Gordonia s.l., including representatives from...

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... of the 24 species of Theaceae examined vary from globose to compressed, ovoid, oblong, or reniform in profile (Fig. 1). Seeds of several specimens have thin, flattened regions that may be described as wings, although the size, shape, and location of these wings vary across taxa. All specimens examined of Gordonia s.l. have prom- inent apical wings. These wings are longer than the length of the embryo in all examined seeds of Gordonia s.l. in Theeae and in half of Gordonia s.l. of Gordonieae. Such well-developed wings were not observed in seeds from any other taxon of ...
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... images of seeds from the five species sampled from Gordonieae are shown in Fig. 2. These seeds include the laterally flattened and winged seeds of Gordonia s.s., includ- ing G. lasianthus and G. brandegeei, and angular, reniform seeds of Franklinia W. Bartram ex Marshall and Schima Reinw. ex Blume (Fig. ...
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... images of seeds from the four species sampled from Stewartieae are shown in Fig. 5. Seeds of species examined were ovoid to oblong, angular or compressed, some with a small wing surrounding the entire embryo (e.g., Stewartia monadelpha Siebold & Zucc.; Fig. 1L). In all seeds observed, testa cells appear consistently small and isodiametric (Fig. 5). In two of the four species (S. pteropetiolata W.C. Cheng (=Hartia sinensis Dunn) and S. ovata (Cav.) Weath.; Fig. 5A, B), testa cells show sculpting on the periclinal surface giving them a plicate appearance. A third species, S. monadelpha, has less prominent sculpting of even bumps on the periclinal surface. In contrast, testa cells of S. malacodendron L. appear exceptionally ...
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... there is wide variation in seed morphol- ogy across Theaceae, seeds of Gordonia s.l. are all flattened and oblong with a conspicuous apical wing (Fig. 1). This morphology appears unique within the family. Indeed, similarity in fruit and seeds was cited by Sealy (1958) and Keng (1980) to justify combining older generic concepts of Laplacea and Polyspora with Gordonia, resulting in the more recent concept of Gordonia s.l. Other taxa of Theaceae have seeds described as winged but these wings are not apical or as prominent as in Gordonia s.l. (e.g., Wang et al., 2006) and may have different ontogenies (Tsou, ...
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... were observed with a light microscope then mounted and left uncoated for observation under SEM with a Jeol JSM 5600LV in a low vacuum mode (20-100 Pa) at 30KV, with a back- scatter detector. A seed of Gordonia s.l. can be divided into two sections: the part that contains the embryo, and the flattened membranous exten- sion of the seed coat that forms an apical wing ( Fig. 1). We obtained SEM images at three loca- tions on the seed: at the proximal end (over the embryo), midway (at the embryo -wing inter- face), and at the distal end (over the wing). Images were obtained at X50, X100 and ...

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... In particular, Gordonia s.l. may have species on both sides of the Pacific. The relationships within this genus have been contested by several authors, and even its status as a monophyletic clade is in question (Prince and Parks 2001;Yang et al. 2004;Gunathilake et al. 2015). The Neotropical distribution of Laplacea in Theeae, on the other hand, has been argued to result from a single long-distance dispersal event from Asia to South America , though the position of key taxa involved in the disjunct distribution is uncertain. ...
... All the Asian species of Gordonia were placed in the genus 12 SYSTEMATIC BIOLOGY Polyspora of tribe Theeae, putatively placed as the sister group to Camellia, corroborating some previous studies (Yang et al. 2004;Li et al. 2011b;Yu et al. 2017b). These findings based on molecular evidence are also supported by recent morphological and cytogenetics studies (Gunathilake et al. 2015;Hembree et al. 2019). ...
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... species is a prominent flattened apical wing, micromorphological study of the seed coat was used to corroborate the above placement of Gordonia s.l. species (Gunathilake et al. 2015). ...
... The isodiametric pattern of the seed coat above the wing (Fig. 1d) is in agreement with that observed in Gordonia s.l. species of Theeae by Gunathilake et al. (2015). The seed coat of species in Gordonieae (G. ...
... The seed coat of species in Gordonieae (G. lasianthus (L.) J.Ellis and G. brandegeei H.Keng) shows protrusions though it is not consequently present on the wing (Gunathilake et al. 2015). Our seeds show an intermediate size between the size ranges of Gordonia s.l. ...
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... In addition to horticultural and historical inquiries, a modest number of biological traits of Franklinia have been studied, including cytology (Santamour, 1963), ovule and anther development (Tsou, 1997), early floral development (Tsou, 1998), external seed morphology and anatomy (Gunathilake, Prince & Whitlock, 2014) and wood anatomy (Liang & Baas, 1991). Among the most striking traits of Franklinia, however, is its reproductive phenology, specifically its prolonged duration of fruit maturation, which has never been studied. ...
... (Prince & Parks, 2001;Li et al., 2013). Gordonieae also includes Gordonia J.Ellis, which used to encompass more species (now shown to belong in Theeae), but is currently understood to comprise only two extant species (Prince & Parks, 2001;Yang et al., 2004;Gunathilake et al., 2014). These are the neotropical G. brandegeei H.Keng and G. lasianthus (L.) J.Ellis, which, like Franklinia, is native to the south-eastern United States. ...
... Aside from Franklinia, the longest intervals between flowering and fruit maturation in Theaceae occur in the entirely evergreen Theeae. In Camellia L. specifically, this process requires many months, but also results in the largest seeds in the family (for seed size comparisons see Gunathilake et al., 2014). Climate and genotype may affect duration of fruit growth, as illustrated by the variation in time needed for fruit development in commercial tea that is grown in widely varying climates. ...
Article
In Franklinia alatamaha (Theaceae) an extended period of dormancy, associated with winter, separates pollination in the late summer and autumn from observable fruit growth, which occurs during the subsequent summer season. Here, ovule and early seed development were examined to decipher the timing of reproductive events that underlie this unusual phenological pattern. Female gametophytes were found to be mature before pollination. Evidence for double fertilization was observed soon after pollination. Early endosperm development progresses for up to 3 months after fertilization but comes to a standstill at the onset of winter. The zygote becomes dormant shortly after fertilization and does not divide during the autumn or winter. At the start of the following growing season, endosperm development is reinitiated and the first events associated with the formation of an embryo occur. Seed development is completed roughly at the same time as new flowers are opening, a full year after pollination and fertilization. Dehiscence of fruits does not occur until later in autumn. The prolonged zygotic dormancy in Franklinia is exceedingly rare among angiosperms and differs markedly from patterns of delayed fertilization that typically underlie extended periods between pollination and seed maturation in other temperate, perennial species. http://onlinelibrary.wiley.com/doi/10.1111/boj.12409/abstract
... Indeed, these characters have been used to differentiate species within many genera, including Allium (Alliaceae; Fritsch et al. 2006), Ephedra (Ephedraceae; Ickert-Bond and Rydin 2011), Frailea (Cactaceae; Metzing and Thiede 2001), Lathyrus (Fabaceae; Güneş and Ali 2011), and Mentzelia (Loasaceae; Thompson and Prigge 1986;Holmgren and Holmgren 2002;Schenk and Hufford 2010). Tes-tal microsculpturing traits have also been used to infer deeper evolutionary relationships within families but with mixed success (e.g., Barthlott 1981;Shetler and Morin 1986;Johnson et al. 2004;Zhang et al. 2005;Cupido et al. 2011;Kasem et al. 2011;Sousa-Baena and Menezes 2014;Gunathilake et al. 2015). Despite the utility of these characters for species delimitations, botanists have yet to identify the ecological or evolutionary reasons why closely related species differ in testal microsculpture traits. ...
... cells. Thus, the seeds of section Bartonia are similar to other species where seed size is a function of cell size (Alonso-Blanco et al. 1999;Ohto et al. 2009;Gunathilake et al. 2015). ...
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... Polyspora Sweet (1825: 205, cited by Sweet, 1826 of the Theaceae has been traditionally placed within the genus Gordonia J. Ellis (1770: 520) but recent studies using molecular markers (Prince & Parks, 2001;Yang et al., 2004;Prince, 2007& Zhang et al., 2014 and seed coat micromorphology (Gunathilake et al., 2014) support its delimitation from the latter. Polyspora species are characterized within the family by having sepals indistinct from the petals, stamens basally united with the corolla, oblong-ellipsoid woody capsules loculicidally dehiscent by five valves, and apically winged seeds (Sealy, 1958;Orel et al., 2013). ...
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