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Apostasia odorata. Androecium and gynoecium development. SEMs and transverse section series of the same flowers (D-F). A, From abaxial. Stamens are differentiated into filaments (arrows) and anthers (A). B, From the side. Incipient papillate stigma marked by arrow. C, Young bud laterally opened. Below the androecium the gynoecium is formed in the concave floral center (arrow). Vascular bundles indicated with thin lines. D, Carpels primordial. Gynoecium still open (a-d). E, Gynoecium closed. Free carpel tips (a, b). Style fused with staminode (c). Portioning of the ovary begins with three ridges along the inside of the ovary wall. Placental ridges contiguous in the lower part of the ovary (d-g). F, Style elongating. Stylar part (a-f). Transition between style and ovary (g). Placental ridges contiguous over the whole length of the symplicate part of the ovary (h-j). Synascidiate zone (k). Perianth removed in A and B. Scale mm; abbreviations as in figs. 2, 11. bars p 100

Apostasia odorata. Androecium and gynoecium development. SEMs and transverse section series of the same flowers (D-F). A, From abaxial. Stamens are differentiated into filaments (arrows) and anthers (A). B, From the side. Incipient papillate stigma marked by arrow. C, Young bud laterally opened. Below the androecium the gynoecium is formed in the concave floral center (arrow). Vascular bundles indicated with thin lines. D, Carpels primordial. Gynoecium still open (a-d). E, Gynoecium closed. Free carpel tips (a, b). Style fused with staminode (c). Portioning of the ovary begins with three ridges along the inside of the ovary wall. Placental ridges contiguous in the lower part of the ovary (d-g). F, Style elongating. Stylar part (a-f). Transition between style and ovary (g). Placental ridges contiguous over the whole length of the symplicate part of the ovary (h-j). Synascidiate zone (k). Perianth removed in A and B. Scale mm; abbreviations as in figs. 2, 11. bars p 100

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Floral development, structure, and pollination biology of species of Apostasia and Neuwiedia constituting the basalmost orchid subfamily Apostasioideae were studied. The perianth organs arise from a ring primordium. The two adaxial sepals appear first, followed by the petal primordia. This contrasts with monandrous orchids, in which the median peta...

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... In contrast, orchids from Vanilloideae and Cypripedioideae species can set fruits through self-pollination, although their floral morphology promote cross-pollination (Suetsugu & Fukushima 2014, de Oliveira et al. 2022. Finally, Apostasioideae orchids might be pollinated by xenogamy (Kocyan-& Endress 2001, Yin et al. 2016. A comparison of pollination systems among different orchid subfamilies is shown in Table 1. ...
... Kocyan & Endress 2001, Yin et al. 2016 Vanilloideae ...
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... Apostasioideae comprises the genera Apostasia and Neuwiedia, and their position in the phylogenetic tree has been controversial [7]. Some previous studies suggested that there is a closer relationship between Apostasioideae and lower Asparagales, Liliales, or Haemadorales [7,56]. However, anatomical and DNA studies indicate an exclusive association of these two genera with the rest of the orchids [7,57]. ...
... However, anatomical and DNA studies indicate an exclusive association of these two genera with the rest of the orchids [7,57]. Our phylogenetic tree based on CDS sequences strongly supports the monophyly of Apostasioideae and their sister-group relationship to the remaining orchids, consistent with previously reported results [56,58]. N. zollingeri var. ...
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... As such, the millions of dust-like seeds can be easily uplifted and carried away by the wind. Nevertheless, there are a few notable exceptions [31,42,[45][46][47][48][49][50][51][52][53][54][55][56][57][58] (Table 1; Figure 1). Hydrochory, or the dispersal of seeds by water, is an extremely rare phenomenon that has been documented in only two species within the Epidendroideae subfamily, Disa uniflora P.J.Bergius [52,59] and Epipactis gigantea Dougl. ...
... Certain orchids-primarily found in early diverging clades-show fleshy fruits that bear hard, rounded, dark seeds. These orchids are known to occur in Apostasia Blume and Neuwiedia Blume, the two genera of the Apostasioideae subfamily, the genus Selenipedium Rchb.f. of the Cypripedioideae subfamily, and in the genera Cyrtosia Blume and Vanilla of the Vanilloideae subfamily [31,42,46,[49][50][51][52][53][54][55][56][57][58]. Fleshy fruits and sclerified seeds seem to be very rare in the Orchidoideae and Epidendroideae subfamilies. ...
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... Resupination is closely related to flower zygomorphy in several families. For instance, in the basal subfamily Apostasioideae of the Orchidaceae, resupination does not occur in the actinomorphic genus Apostasia but is present in the zygomorphic Neuwiedia (Kocyan & Endress 2001). In orchids, zygomorphy is closely linked with the position of the median petal, known as the labellum (Darwin 1877;Ames 1938;Van der Pijl & Dodson 1966; Mondrag on-Palomino & Theißen 2009). ...
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• In many families, plants undergo floral resupination by twisting through approximately 180º during floral development so that the flower is effectively positioned upside down. In most orchids, resupination results in the median petal (i.e. the labellum) becoming lowermost, which plays a crucial role in pollination by serving as a landing platform or as a trapping device or both. Incomplete resupination is predicted to lead to reduced pollination, although tests of this assumption are still lacking. • We investigated the effect of resupination using Phragmipedium vittatum, a rare lady's slipper orchid whose specialized labellum forms a trapping device. First, we surveyed the natural occurrence of incomplete resupination. Then, we manipulated flowers into non- (≈ 0º), half- (≈ 90º), and fully resupinate (≈ 180º) positions to test the effect of orientation on pollen smear removal and deposition by pollinators (female hoverflies). • We found that ca. 10 % of flowers in the natural population were not fully resupinate, being either non- (upward, 0º - 60º) or half-resupinate (sideward, 60º - 120º). The change in orientation prevented the effectiveness of pollination by hoverflies since no pollen smear removal or deposition were found in flowers from non- and half-resupinate treatments. Although these flowers still attracted hoverflies, they were not trapped effectively. • As this orchid is incapable of autonomous self-pollination, flowers that do not resupinate fail to set fruits. These results highlight the importance of correct floral orientation provided by resupination to ensure pollination in orchids and other resupinate flowers.
... In contrast with all other orchids, Apostasia Blume (1825: 423; Orchidaceae: Apostasioideae) is characterized by their nearly actinomorphic flowers with recurved perianth, a simple gynostemium and three united stamens (the lateral stamens are fully developed, whereas the median is a staminode; Kocyan & Endress 2001, Rao 1974). This subfamily includes fewer than ten species mainly distributed in Southeast Asia from the Himalayas and Japan to northern Australia (de Vogel 1969). ...
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After examining morphological, molecular and genome-size evidence, we here describe a new orchid species, Apostasia fujianica, from Fujian, China. Morphological comparisons indicated that A. fujianica is similar to A. shenzhenica and A. nipponica, whereas the former displayed distinct differences in habit, roots, leaves, inflorescences and fruit shape and size. Apostasia fujianica (341 Mb) has a smaller genome size than A. shenzhenica (471 Mb), A. ramifera (366 Mb) and A. fogangica (931 Mb). Molecular analyses from combined nuclear (ITS, Xdh, naD1) and plastid (matK, rbcL, psbA-trnH, trnL-trnF and trnS-trnG) datasets indicated that A. fujianica is sister to A. shenzhenica. These results support the status of A. fujianica as a new species, distinguished in many aspects from A. shenzhenica, A. ramifera and A. nipponica.
... A distinctive feature is the fusion of the stamens and pistil into a single structure referred to as the column (van der Cingel 2001; Roberts & Dixon 2008). Reduction in stamen number has led to orchid groups with three, two or, for the vast majority (99%) of species, one stamen (van der Pijl & Dodson 1966;Kocyan & Endress 2001). Below the column is a petal that has been modified into a labellum, or lip, which acts as a landing area for pollinators and directs them to the (presumed) nectar source within the flower (van der Cingel 2001;Brown 2005). ...
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There are approximately 30,000 species in the family Orchidaceae, with some species growing terrestrially and others growing as epiphytes. Though the pollination biology of many of these species is not well known, there has been a diversity of taxa recorded as orchid pollinators. Insects, birds, and even a record of a mammal species have been documented as successful pollinators, while some orchids are able to reproduce without the use of a pollen vector. The goal of this review is to provide an overview of orchid pollination tactics, with references to more specific studies of each, and to analyze a large subset of publications to determine differences in pollinator taxa and specificity between epiphytic and terrestrial orchid genera. This review examines pollination data from over 400 orchid species, including 74 epiphytic and 83 terrestrial orchid genera. Two pollinator taxa, Coleoptera and Hymenoptera (Class: Insecta), were found to pollinate significantly more terrestrial than epiphytic orchid genera, while other taxa showed no significant differences. Hymenoptera were the dominant taxa of pollinator in regards to the overall number of species recorded; however, based on species interaction webs that were built, the Lepidoptera (Class: Insecta) have stronger interactions with the orchid species they pollinate, suggesting a more specific relationship between the two.
... However, these differentiating characters are very superficial, as is evident from Figure 2 in Lu et al. (2019), in which true A. shenzhenica is shown with open flowers in which the staminode is clearly shorter than the style and wings are clearly present toward the base of the staminode. It is also evident from a study on floral development in Apostasia by Kocyan and Endress (2001) that a shorter style is simply an effect of early developmental stage. We conclude that it is likely that incompletely developed floral parts had been observed at the time of the original publication of A. shenzhenica, and hence the dimensions mentioned in the protologue are not an accurate reflection of the size of these structures as compared with those provided in the literature for other species. ...
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Apostasia nipponica and Crepidium cordilabium are newly added to the orchid flora of Hong Kong and their taxonomy is reviewed. A detailed description and line drawing are provided for both species, as are notes on flowering phenology and conservation status. Following careful examination of type material and other authentic specimens from throughout their range, as well as of photographs of plants in the wild, A. nipponica is removed from the synonymy of A. wallichii and reinstated as a valid species, under which A. ramifera and A. shenzhenica are newly synonymized. Additionally, Crepidium cordilabium, which was originally described from Taiwan as a natural hybrid between C. matsudae and Dienia ophrydis, is treated here as a distinct species belonging to the section Hololobus and sub‐section Maximowiczianae.
... Most orchids possess dehiscent, capsular fruits, while indehiscent, fleshy fruits have been observed only in a few genera across different subfamilies, such as Neuwiedia (subfamily Apostasioideae), Selenipedium (subfamily Cypripedioideae), Cyrtosia (subfamily Vanilloideae), and Yoania (subfamily Epidendroideae) (Dressler 1989;Clements and Molvray 1999;Kocyan and Endress 2001;Suetsugu 2018a,b). So far no fleshy fruit is reported in any genus of subfamily Orchidoideae. ...
... The subfamily Apostasioideae (containing two genera-Apostasia and Neuwiedia) has been considered as the earliest-diverging lineage of Orchidaceae, which possesses several unique characters, including actinomorphic flower, three stamens and powdery pollen grains (Kocyan and Endress 2001). In addition, the presence of indehiscent fruits and the thickened seed coat in some Apostasia and Neuwiedia species (Nishimura and Tamura 1993;Clements 1999) are suggested to be plesiomorphic characters. ...
... In the subfamily Apostasioideae, only two Neuwiedia species, i.e. N. griffithii and N. veratrifolia are known to possess capsular fruits (Kocyan and Endress 2001). Such an observation would suggest that fleshy fruit might be a plesiomorphic character in Orchidaceae, and the production of fleshy fruits arose a few times in other orchid subfamilies. ...
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Background Seed dispersal allows plants to colonize new habitats that has an significant influence on plant distribution and population dynamics. Orchids produce numerous tiny seeds without endosperm, which are considered to be mainly wind-dispersed. Here, we report avian seed dispersal for an early diverging orchid species, Neuwiedia singapureana , which produces fleshy fruits with hard seed coats in the understory of tropical forests. Results Neuwiedia singapureana produced fleshy fruits that turned red in autumn, and birds were confirmed to be the primary seed dispersers. As compared to its sister species, N. veratrifolia with dehiscent capsular fruits, embryos of N. singapureana were larger and enclosed by thickened and lignified seed coats. After passing through the digestive tracts of birds, the seeds still stayed alive, and the walls of seed coat contained several cracks. The germination percentage increased significantly for digested seeds as compared with seeds from intact fruits. Conclusion The thickened and lignified seed coat may protect seeds as they passed through the digestive tracts of birds. Taken together with a recent report of insect-mediated seed dispersal system in the subfamily Apostasioideae, the animal-mediated seed dispersal may be an adaptive mechanism promoting the success of colonization in dark understory habitats.
... To achieve clear evidence of the status of ovule development in the Apostasioideae is difficult because of the reduced number of species and their rarity. According to the literature (Kocyan and Endress, 2001) on two species of Neuwiedia, one of the two genera of the Apostasioideae, ovule development is almost complete at anthesis. This finding suggests that members of Apostasioideae have the same ovule developmental stage at anthesis as some members of Asparagales such as Iridaceae and Hypoxidaceae (i.e., Vos, 1948;Rudall, 1994). ...
Article
Premise: Unlike most flowering plants, orchid flowers have under-developed ovules that complete development only after pollination. Classical studies reported variation in the stage in which ovule development is arrested but the extent of this variation and its evolutionary and ecological significance are unclear. Methods: Here, we performed light microscopy observations and surveyed the literature gaining information on 94 orchid species including tropical and temperate members of all subfamilies as well as species with contrasting pollination strategies (rewarding versus deceptive) and life forms (epiphytic versus terrestrial). We analysed the data using statistical comparisons and a Phylogenetic Generalized Least Square (PGLS) analysis. Key results: Apostasioideae, the sister to the rest of the orchids, have mature ovules similar to other Asparagales, while under-differentiated ovules are present in the other subfamilies. Ovule developmental stages showed high variation even among closely related groups. Ovules were more developed in terrestrial than in epiphytic, in temperate than in tropical, and in rewarding than in deceptive pollination orchid species. This latter comparison was also significant in the PGLS analysis. Conclusions: These results suggest that ovule developmental stage in orchids can be shaped by ecological factors, such as seasonality and pollination strategy, and can be selected for optimizing female reproductive investment. This article is protected by copyright. All rights reserved.
... In Orchidaceae, floral bud differentiation has been investigated in several taxa, including Cattleya, Bletilla striata, Cypripedium, Phragmipedium, Oncidium, Apostasioideae, Chloraea, Telipogon, Phalaenopsis and Dendrobium (Rotor and MacDaniels 1951;Sano et al. 1961;Tanaka et al. 1986;Kurzweil 1993;Kocyan and Endress 2001;Weng et al. 2002;Pabón-Mora and González 2008;Wei et al. 2010;Steinfort et al. 2012;Wang et al. 2019;Zhang et al. 2019). In general, the shoot apical meristem begins to differentiate, initiating the development of floral organs after flowering induction is completed. ...
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