The floral morphology and ontogeny of some Chinese representatives of orchid subtribe Orchidinae

The flower structure and development of ten species in six genera of the orchid subtribe Orchidinae are described and illustrated by scanning electron micrographs. Particular attention is given to the structure of the gynostemium, which for most species is interpreted from ontogenetic data. All the species studied here share a series of features, e.g. the sequence of tepal and anther initiation, the shape and position of the anther, the presence of auricles and basal bulges, the three-lobed condition of the median carpel apex and the lateral lobes of the median carpel embracing the basal ends of the thecae. However, the form and structure of the three carpel apices are most varied in the later development stages or in the adult flower. The genus Hemipilia shows a series of peculiar characters that are quite different from those of the other genera in Orchidinae. The peculiar structure and development of the viscidia in both Amitostigma and Neottianthe indicate that both of them are different from other genera in Orchidinae. The adult floral morphology shows that the genera Galearis and Chusua are not congeneric with Orchis. The separation of the lateral lobes of the rostellum in most genera studied here as well as in the Brachycorythis group from South Africa suggests that this is the ancestral state in the subtribe Orchidinae. In contrast, the conjoining of lateral lobes in Dactylorhiza and Orchis is suggested as a derived character.     

Developmental studies in orchid flowers II: Orchidoid species

The floral development of 11 species of Orchidoideae (sensu Rasmussen 1985) was studied by means of SEM, paying special attention to the early development of the gynostemium and its appendages. In contrast to the staminodes found in epidendroid and vandoid orchids, the 'auricles' of the tribe Orchideae are developed on the dorsal side of the fertile anther and therefore are not interpreted as staminodes. Presumed vestiges of the staminodes corresponding to those of the Epidendroideae and Vandoideae are differentiated in early developmental stages, but remain inconspicuous structures later on. The three-lobed rostellum originates entirely from the median carpel. The outstanding systematic position of the tribe Orchideae is briefly discussed.     

寒兰花器官发生及花发育过程的研究

为了揭示寒兰的成花机理,利用石蜡切片和花芽实体解剖记录了濒危植物寒兰花芽分化和发育的过程,并着重观察唇瓣和合蕊柱早期及中期的发育(在合蕊柱伸长之前)。结果表明:寒兰花芽分化沿着花序轴从下往上可分为4个阶段:花序原基分化,花原基分化,花被片分化和合蕊柱形成。唇瓣分化分为3个阶段:褶片分化,侧裂片分化和色块形成。唇瓣侧裂片和褶片产生较晚,与退化雄蕊可能没有关系。在合蕊柱形成过程中,首先分化出花药,随后分化产生中心皮顶部,侧心皮顶部,并形成花柱道,最终分化出蕊喙和黏盘。     

The gynostemium of Hemipiliopsis purpureopunctata and Senghasiella glaucifolia, two taxonomically disputed species of Habenariinae (Orchidaceae)

The gynostemium structure and ontogeny of two taxonomically disputed orchids, Hemipiliopsis (= Habenaria ) purpureopunctata and Senghasiella (= Habenaria ) glaucifolia , are described and illustrated by scanning electron micrographs. The early gynostemium ontogeny of Hemipiliopsis purpureopunctata is shown to be fundamentally similar to that of the species of the tribe Orchideae that have been previously studied. This includes the initiation sequence of sepals, petals and lip, form and orientation of anthers, three-lobed condition of median carpel apex, and presence of auricles and basal bulges. During the later developmental stages some differences occur. The stigma processes of Senghasiella glaucifolia are united into a tongue-shaped organ, and the lateral rostellum lobes of Hemipiliopsis purpureopunctata protrude forwards with their viscidia positioned above the spur-mouth. Based on gynostemium characters, the generic rank of Hemipiliopsis was confirmed, but that of Senghasiella was not supported.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 191–196.     

Developmental studies in orchid flowers IV: Cypripedioid species

The floral development of four species of Cypripediaceae (sensu Rasmussen 1985) was studied by means of scanning electron microscopy, with special attention to the early development of the organs that constitute the gynostemium. At the ventral base of the gynostemium a prominent structure was observed. It is most probably a vestige of the median adaxial stamen a₃ based on its early initiation and place of origin. In Cypripedium calceolus the median carpel primordium is, according to expectation, initiated slightly earlier than the lateral carpel primordia, and later develops into the largest stigma lobe. Interestingly, Cypripedium irapeanurn shows an opposite sequence in the initial phase of the carpel development in that the primordia of the lateral carpels are initiated before the primordium of the median carpel.     

Floral morphology of southern African Orchideae. I. Orchidinae

The floral morphology of the southern African genera of Orchidaceae-Orchideae-Orchidinae ( Brachycorythis, Schwartzkopffia, Neobolusia, Schizochilus, Holothrix and Bartholina ) is surveyed paying special attention to the gynostemium. Ontogenetic data are provided for a number of species that appear to be essential in formulating a proper interpretation of the gynostemium. The floral architecture is shown to be basically similar to that of the (much better known) European representatives of the subtribe. This, however, does not fully apply to the homology of the lateral gynostemium appendages ("auricles"): In Brachycorythis, Neobolusia and Schizochilus these develop like in Orchis and Dactylorhiza. Their prominent sculptured portions originate from dorsal stamen outgrowths and correspond to filament excrescences. Structures obviously homologous to lateral inner stamens can be recognized in the early ontogeny, but are in the mature flower incorporated in the 'arch' connecting the lip with the gynostemium. In contrast, in Holothrix and Bartholina the gynostemium appendages correspond entirely to staminodes, while the filament excrescences are missing. It is also shown that the 'concave' stigma said to be characteristic of the Orchidinae is in fact ± convex or even pad-like, but is generally positioned in a cavity under the rostellum. The 'erect' anther (the main diagnostic feature of the Orchideae) is reflexed up to 45° in some taxa. Affinities of the genera are briefly discussed. The generic separation of Schwartzkopffia and Neobolusia from Brachycorythis does not appear justified. Neobolusia virginea is obviously misplaced in the respective genus, and eventually merits generic status. The affinities of Schizochilus remain ± obscure at the moment. Bartholina appears to be merely a small group of specialized Holothrix species.     

Developmental studies in orchid flowers I: Epidendroid and vandoid species

The floral development of 47 epidendroid and vandoid orchids was studied by means of scanning electron microscopy, paying special attention to the early development of the gynostemium (column) and its appendages. The following main conclusions are drawn: the lateral appendages of the adult gynostemium are homologous with the two lateral stamens of the inner whorl; their primordia are present even in species which lack prominent appendages in the adult gynostemium (incorporation of the sta-minodial primordia into the gynostemium during development). Ventral appendages observed in some species are supposed to be vestiges of the adaxial stamens on account of their early initiation. It is confirmed that the rostellum is the upper part of the median stigma lobe and that the lip corresponds to the inner median tepal. The affinities of the epidendroid and vandoid orchids are briefly discussed.     

The gynostemium of Bulbophyllum ecornutum (J. J. Smith) J. J. Smith (Orchidaceae)

RASMUSSEN, F. N., 1985. The gynostemium of Bulbophyllum ecornutum (J. J. Smith) J.J. Smith (Orchidaceae) . Stages in the development of the gynostemium of Bulbophyllum ecornutum demonstrate that the pollinium stalk is a hamulus in this and in a closely related species, B. gibbolabium . A hamulus arises by apical growth and reflexion of the median carpel. Hamuli have recently been discovered in several orchid genera, and a transverse fold of the rostellar apex is already known from a large group of orchids. The closely related B. cornutum has a quite different gynostemium structure.     

POSTPOLLINATION PHENOMENA IN ORCHID FLOWERS. V. PARTICIPATION BY THE ROSTELLUM AND GYNOSTEMIUM TIP

Removal of the rostellum following pollination does not prevent stigmatic closure in Cymbidium flowers and has a minimal effect on straightening of the gynostemium (column). However, this treatment does depress anthocyanin levels in both gynostemia and labella. Excision of the rostellum 30 or 60 min after pollination has a more pronounced effect than removal after 150 min. Stigmatic closure is not inhibited by removal of the gynostemium tip, but column swelling is reduced. These findings are discussed relative to rostellar functions and theories regarding their origin.     

Histological and Micro-CT Evidence of Stigmatic Rostellum Receptivity Promoting Auto-Pollination in the Madagascan Orchid Bulbophyllum bicoloratum

Background

The rostellum, a projecting part of the gynostemium in orchid flowers, separates the anther(s) from the stigma and thus commonly prevents auto-pollination. Nonetheless, as a modified (usually distal) portion of the median stigma lobe, the rostellum has been frequently invoked of having re-gained a stigmatic function in rare cases of orchid auto-pollination. Here it is shown that a newly discovered selfing variant of Madagascan Bulbophyllum bicoloratum has evolved a modified rostellum allowing the penetration of pollen tubes from in situ pollinia.

Methods

Gynostemium micro-morphology and anatomy of selfing and outcrossing variants of B . bicoloratum was studied by using light and scanning electron microscopy and histological sections. Pollen tube growth in the selfing variant was further observed via X-ray computed microtomography (micro-CT), providing 3D reconstructions of floral tissues at a micron scale.

Findings

Selfing variants possess a suberect (‘displaced’) rostellum rather than the conventional, erect type. Very early in anthesis, the pollinia of selfers are released from the anther and slide down onto the suberect rostellum, where pollen tube growth preferentially occurs through the non-vascularized, i.e. rear (adaxial) and (semi-) lateral parts. This penetrated tissue is comprised of a thin layer of elongate and loosely arranged cells, embedded in stigmatic exudates, as also observed in the stigmatic cavity of both selfing and outcrossing variants.

Conclusions

Our results provide the first solid evidence of a stigmatic function for the rostellum in orchid flowers, thereby demonstrating for the first time the feasibility of the micro-CT technique for accurately visualizing pollen tube growth in flowering plants. Rostellum receptivity in B . bicoloratum probably uniquely evolved as an adaptation for reproductive assurance from an outcrossing ancestor possessing an erect (non-receptive) rostellum. These findings open up new avenues in the investigation of an organ that apparently re-gained its ‘primordial function’ of being penetrated by pollen tubes.     

A molecular phylogeny for the large African orchid genus Disa

Phylogenetic relationships were inferred for the African subtribe Disinae (Orchidoideae, Orchidaceae), which include the large genus Disa and the small genus Schizodium. One nuclear (ITS) gene region and two plastid (trnLF and matK) gene regions were sequenced for 136 ingroup, representing 70% of all known Disinae species, as well as for 7 outgroup taxa. The combined data matrix contained 4094 characters and was analysed using parsimony and Bayesian inference. Our results show that the generic status of Schizodium can no longer be supported, as it is deeply embedded within the genus Disa. Furthermore, the currently recognised subgenera do not reflect the phylogenetic relationships and should be rejected. Several of the currently recognised sections are monophyletic, others contain misplaced elements, while some are polyphyletic. Morphological divergence, rather than convergence, has hampered previous attempts at a phylogenetic classification of the Disinae. On the basis of our molecular phylogenetic hypothesis, we propose a monotypic subtribe Disinae and a subdivision of the genus Disa into 18 sections.     

马兜铃属(马兜铃科)花的形态发生研究

对马兜铃属植物北马兜铃(Aristolochia contorta Bge.)花形态发生的扫描电镜观察表明:其花萼在发生时与苞片相似,6枚雄蕊呈4枚先发生、2枚后发生的方式,心皮的发生以6个胎座突出到子房室中为特征,由侵入的侧膜胎座合生为中轴胎座。在胎座发生发育过程中,在花药的腹面各产生一个突起,此突起后来与胎座上端相连,最终发育为合蕊柱裂片。结合文献资料,我们认为北马兜铃的花被与苞片是同源的,其合蕊柱裂片来自于雄蕊,中轴胎座是次生的。     

Developmental studies in orchid flowers III: Neottioid species

The floral development of 19 species of Neottioideae (sensu Rasmussen 1985) was studied by means of scanning electron microscopy, paying special attention to the early differentiation of the organs that constitute the gynostemium. The gynostemium development of the Epipactieae proved to be similar to that of the Epidendroi deae and Vandoideae, in particular in that massive primordia corresponding to inner lateral staminodes are differentiated in early stages and later constitute the lateral appendages of the gynostemium. In the Neottieae a progressive reduction and delayed initiation of these staminodes was observed: the lateral teeth of the gynostemium originate from large staminode primordia in one species ( Corymborkis veratrifolia ), in the remaining species they are initiated in later stages or are missing.     

Recurrent polymorphic mating type variation in Madagascan Bulbophyllum species (Orchidaceae) exemplifies a high incidence of auto‐pollination in tropical orchids

The transition from outcrossing to self‐fertilization is one of the most common evolutionary changes in angiosperms. The orchid family exemplifies this evolutionary trend but, because of a general lack of large‐scale surveys on auto‐pollination in orchid taxa, the incidence and modes of auto‐pollination among (sub)tropical orchids remain poorly known. In the present study, we assessed the frequency and mode of auto‐pollination within and among species of a largely monophyletic group of Madagascan Bulbophyllum. The capacity for autonomous fruit set was investigated by bagging experiments in the greenhouse and the field, complemented with detailed floral micromorphological studies of the gynostemium. Our survey comprises 393 accessions, representing at least 78 species, and thus approximately 37% of the species diversity of the genus in the Madagascan region. Our studies revealed that mating type is directly related to gynostemium structure, most often involving the presence or absence of a physical barrier termed ‘rostellum’. As a novel and unexpected finding, we identified eight species of a single lineage of Madagascan Bulbophyllum (termed ‘clade C’), in which auto‐pollinating morphs (selfers), either lacking a rostellum or (rarely) possessing a stigmatic rostellum, co‐exist with their pollinator‐dependent conspecifics (outcrossers). We hypothesize that auto‐pollination via rostellum abortion has a simple genetic basis, and probably evolved rapidly and recurrently by subtle changes in the timing of rostellum development (heterochrony). Thus, species of clade C may have an intrinsic genetic and developmental lability toward auto‐pollination, allowing rapid evolutionary response under environmental, perhaps human‐disturbed conditions favouring reproductive assurance. Overall, these findings should stimulate further research on the incidence, evolution, and maintenance of mating type variation in tropical orchids, as well as how they adapt(ed) to changing environmental conditions. © 2014 The Authors. Botanical Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 242–258.     

Trends in evolution of floral ontogeny in Cassia sensu stricto,Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); a study in convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off-median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).     

ORIGIN AND DEVELOPMENT OF THE TERMINAL CARPEL IN PSEUDOWINTERA TRAVERSII

Flowers of Pseudowintera traversii (Buchan.) Dandy possess a terminal unicarpellate gynoecium. The present study of carpel morphogenesis was initiated for the purposes of (1) providing additional developmental documentation of the occurrence of terminal carpels in the Winteraceae and (2) comparing the mode of initiation and development of the ascidiate terminal carpel of P. traversii with the essentially conduplicate terminal carpel of Drimys lanceolata. Following its axillary origin, the floral apex of P. traversii initiates 2–3 connate sepals, 5–6 petals, 4–15 stamens, and usually a single terminal carpel, in acropetal succession. Bicarpellate gynoecia may occur with a frequency of up to 15 % on a given plant. The floral apex is zonate and shows increased expression of its zonation during later stages of floral development. The terminal carpel is ascidiate from inception and originates as a cylindrical growth around the entire circumference of the floral apex; transformation of the floral meristem into a carpel primordium terminates apical growth of the floral axis. Carpel growth continues to be cylindrical and is mediated by a ring of marginal and submarginal initials at its summit. Earlier and more extensive division of initials and their derivatives on the dorsal rim causes the primordium to become canted adaxially, shifting the apical cleft to a subterminal adaxial position. Continued marginal meristematic activity results in closure of the cleft as well as elevation and elaboration of the stigmatic crests. Five to seven bitegmic ovules are initiated at the same time as crest elaboration and arise in two rows from the adaxial (laminar) position. Carpel maturation is signified by tannin deposition and oil cell differentiation, beginning at the base and proceeding acropetally; carpel margins bordering the cleft are the last to differentiate. Carpel procambialization is continuous and acropetal from inception, with the dorsal median bundle differentiating before the ventral strands. The significance of occasional bicarpellate flowers is discussed.     

On Diplandrorchis,a very primitive and phylogenetically significant new genus of Orchidaceae

Diplandrorchis is a very curious new orchid genus, containing only one species D. sinica S. C. Chen, found in Huan-ren County of Liaoning Province in northeastern China. It, as the genus name implies, has two fertile stamens, borne on the upper part of the column near the terminal stigma. One of them is opposite to the dorsal sepal, while the other to the median petal (lip). Thus, they represent two median stamens of both inner and outer whorls. This is quite unique in the whole family, including the diandrous group, in which the two stamens are opposite to the lateral petals and thus represent the lateral ones of the inner whorl. The flower is erect, with its pale greenish or whitish perianth nearly regular. Two lateral sepals are more or less oblique, showing some difference from the dorsal one, but the three petals, which are thinner and narrower, are very similar to each other. Neither rostellum nor any other appendages are found in its column, but a terminal stigma and two erect stamens. The pollinia are naked and granular, which, in almost all flowers examined, have naturally fallen out of their cells on to the stigma. Apparently, it is self-fertilised, as found in Tangtsinia and some other primitive orchids. Besides, this interesting orchid is a dwarf saprophyte, with its habit very similar to that of Neottia, another saprophytic genus assigned to the subtribe Neottiinae. The fact that the saprophytic orchids are largely found in the primitive subtribes, such as the Neottiinae, Limodorinae, Vanillinae and Pogoniinae, is worthy of special attention. Theoretically, the saprophytic plants must have developed from green-leaved plants. In the Orchidaceae, however, what ancestor are the most primitive saprophytes derived from ? This is indeed an interesting question closely related to the origin of the Orchidaceae. From the facts mentioned above, the present genus is a very primitive or relic one and of great phylogenetic interest. It is placed here as the most primitive member in the subtribe Neottiinae, although it is sharply distinct from the remaining generaof this subtribe. It may deserve a separate subtribe, but further study is needed.     

Comparative vegetative anatomy and classification of Diseae (Orchidaceae)

The leaf, stem, root, tuber and dropper anatomy of the orchid tribe Diseae (including the subtribes Satyriinae, Disinae, Brownlecinac, Huttonaeinae and Coryciinae) is reviewed. The study is largely based on investigations of 123 species, and data from several previous publications have also been incorporated. Two characters were identified as being taxonomically valuable: (1) the presence of sclerenchyma caps associated with leaf vascular bundles, and (2) the degree of dissection of the siphonostele of the tuber ('polystelic' or 'monostelic'). The phylogenetic analysis shows that anatomical characters do not change the basic structure of a cladogram that is based on morphological characters. The taxa of Diseae are discussed on the basis of anatomical data. Subtribes Satyriinae (excluding the anatomically unusual genus Pachites), Brownleeinae, Huttonaeinae, and Coryciinae are uniform in. critical anatomical characters. However, subtribe Disinae is rather diverse in vegetative anatomy. Disa sect. Micranthae differs from the rest of the genus in its leaf anatomy. The occurrence of foliar sclerenchyma bundle caps and 'polystelic' tubers supports the incorporation of Herschelianthe in Disa sect. Stenocarpa.     

The column types of Neottia and Archineottia of the family Orchidaceae and their taxonomic and phylogenetic significance

The column is the most characteristic part of an orchid flower. It is considered to be formed by the union of stamens with a central style and stigma. In the Apostasieae, for example, the column is rather primitive in the stamens and style only partially united, whereas in the majority of higher orchids it becomes more advanced through a eomplete union of them into a single organ. Within the family, indeed, the column structure is greatly diversified and of great taxonomic significance. It is interesting to note that a great range of diversity of column structure is bund in Neottia (sensu lato), a small but widespread genus consisting of 14 species, about two thirds of which, however, are of local occurance and seem to be little known to many botanists. In some speeies of this genus we find a very primitive column structure which is quite unique in the family, while in the others it is much more complicated. In all, five types of their column structure can be distinguished as fol- lows: (1) column rather longer; anther erect with a short filament attached to the back of the column near the apex; stigma terminal; neither clinandrium nor rostellure; (f. 2, 4) (2) as the preceding, 'except for the stigma more or less curved foreward and filament longer; (f. 6, 8) (3) column rather longer with a clinandrium at its summit, upon which a sessile and incumbent anther sits; rostellum large, horizontally projecting out over the concave stigma situated in the front of the column; (f. 10, 13, 15, 17) (4) as the preceding, except, for the anther and rostellum almost erect, and the stigma more or less bilabiate; (f. 19,21) (5) column very short; anther and rostellum erect; stigma lamellate, erect; reflexed and almost clasping the rostellum. (f.,2g) In these .five types, with the exception of the first one in which the labellum (the median petal) is very similar to the lateral: petals, they all possess zygomorphic perianth with labellum bilobed or entire which is quite different from the two lateral petals. Here, we see a great change in the column structure from one form with stamen and style not fully united to another form in which they have been well fused. Speaking strictly, these are two sorts of entirely different column structure. The former one, represented by (1) and (2) as stated above, is, in fact, an incomplete or s very primitive column in having a terminal stigma and an erect stamen with its free filament attached to the back of the column; and the absence of clinandrium and rostellum. Furthermore, there exists on the back of the column a thick ridge with its upper end joined to the filament, with which it is of the same texture and appearance. In Neottia pantlingii (=Arohineottia pantlingii) the free filament is even rather longer than the ridge, (f. 6) while in the other three species (f. 2, 4, 8) they are shorter. It is in my opinion the lower part of the filament adnate to the compound style or column. This is another fact of interest perhaps not occuring in any other living orchids. On the other hand, the latter one, represented by (3), (4) and (5), is a more advanced column structure, in which a higher level of specialisation with well-developed clinandrium and rostellum is reached. The stigma becomes shallow depressed on the anterior side of the column, or sometimes in the form of somewhat a bilabiate lip projecting out before or under the long rostellum. This is apparently a complete column both in structure and function quite different from the former and, contrarily, much like that of Listera. Basing upon the facts just mentioned, we may subdivided Neottia (sensu lato) into two distinct genera, with two and three sections respectively. They are as follows: 1. Archineottia S. C. Chen, gen. nov. (1) Sect. Archineottia 1) A. gaudissartii (Hand.-Mzt.) S. C. Chen, comb. nov. (China) 2) A. microglottis (Duthie) S. C. Chen, comb. nov. (India) (2) Sect. Furciila S. C. Chen, sect. nov. 3) A. pantlingii (W. W. Smith) S. C. Chen, comb. nov. (Sikkim) 4) A. smithiana (Schltr.) S. C. Chen, comb. nov. (China) 2. Neottia Guett. (1) Sect. Listeroides S.C. Chen, sect. nov. 1) N. listeroides (L.) Rchb. f. (China, Sikkim, Kashmir) 2) N. camtschatea (L.) Rchb. f, (China, Soviet Union) 3) N. megalochila S. C. Chen, nom. nov. (China) 4) N. inayatii (I)uthie) Schltr. (Pakistan, Kashmir) 5) N. tenii Schltr; (China) (2) Sect. Neottia 6) N. papilligera Schltr. (Chinas: Japan, Korea, Soviet Union, Sikkim) 7) N. nidus-avis (L.) L. C. Rich. (Europe, Iran, Western Siberia) 8) N. brevilabris Tang et Wang: (China) (3) Sect. Hologlossa S. C. Chen, sect. nov. 9) N. acuminata Schltr. (China, Japan, Korea, Soviet Union, Sikkim) Inperfeetly known species: 10) N. ussuriensis (Kom. et Nevski) S6o (Soviet Union) Thus, the subtribe Neottiinae are composed of four genera, namely, Diplandrorchis, Archineottia, Neottia and Listera. The new genus Archineottia, as one of the most primitive genera in the family, is of great interest from a phylogenetic point of view. It shows dose similarity to Diplandrorchis and Neottia in habit, but sharply distinct from them in column structure. These genera, as indicated By some authors, also show affinity in some respects with the subtribe Limodorinae, especially to Tangtsinia and Sinorchis, the other two quite primitive genera in the family. There is, indeed, a great need of further study of these interesting or relic genera and this, I think, would go a long way towards solving the problems concerning the origin ofthe Orchidaceae.