墨脱毛兰，一个可能的“异常整齐花”型，兼论兰科中“异常整齐花”现象

摘要/Abstract

摘要：

新种墨脱毛兰可能是足茎毛兰的“异常整齐花”型。 “异常整齐花”现象在兰科中时

有所见，在不同分类群中情况各异，至今尚未见系统的研究。本文认为辐射对称花被是兰科的

正常特征之一，主要见于原始的种类和少数进化的种类，也不同程度地存在于“异常整齐花”实

体中。具辐射对称花被的兰科植物大多数不是“异常整齐花”型，判断时要参酌其他特点，持慎

重态度。兰科“异常整齐花”有两种情况，一是异常型，亦即正常植株中仅偶见“异常整齐花”，

或同一个种可同时产生“异常整齐花”植株与正常植株，两者并存。另一种情况是正常型，亦即

只产生“异常整齐花”，而无正常的两侧对称花。后者意味着在进化过程中的一种飞跃，即形成

了新植物。对此最好作为独立的种处理，特别是在它的来源还不清楚的情况下更是如此。目

前所认为的一些所谓“异常整齐花”实体，大都是推测而已，其中有些或许在将来会被证明为正

常的、原始的植物。

关键词: 墨脱毛兰, 足茎毛兰, 兰科, 丝唇美洲兜兰, 无喙兰, 印度无喙兰, 尖唇鸟巢兰, 无距兰属, 缘毛鸟足兰

Abstract:

Eria mêdogensis S. C. Chen et Tsi was recently found in southeastern Tibet, se-

veral specimens of which have been collected by various botanists since 1980. This is a “normal”

entity with its habit very similar to that of Eria coronaria, from which it differs by having a

regular perianth and longer bracts. We think it probable that this new entity is a peloric form

of Eria coronaria.

Peloria (or pelory) is a type of floral abnormality, which is found in many zygomorphic-

flowered taxa. It was first detected by Linnaeus (1744) in Linaria vulgaris, and then by others

in Labiatae, Orchidaceae, etc. However, it is still an open question how to explain it theoretical-

ly and how to treat it taxonomically.

In Orchidaceae, so far as our knowledge is concerned, peloria has been encountered in no

less than 21 genera. In most cases, peloric flowers are found sporadically on an occassional plant,

as seen in Cypripedium reginae and Eria oblitterata. Sometimes, however, peloric form may occur

coexisting with normal-flowered form in one and the same species, as seen in Dendrobium tetro-

don and Epipogium roseum. They are both abnormally peloric forms. It would not result in

naming or renaming a plant taxonomically, whether the appearance of abnormally regular flo-

wers on a normal-flowered inflorescence, or of abnormal-flowered individuals in normal-flower-

ed species. In Phragmipedium lindenii, however, the case is different. It is quite “normal” and

even of wider distribution than its nonpeloric allies P. wallisii and P. caudatum, from which

it has once been considered to be derived. This is a normally peloric form. Whether it is a

reversal or not, the appearance of a “normally” peloric taxon may be taken for a leap in the

process of evolution. Taxonomically, we had better treat it as a separate species, especially when

its origin is uncertain. For example, the entity just mentioned had been treated as a peloric va-

riety of Phragmipedium caudatum (var. lindenii) until 1975, when Dressler & Williams recogniz-

ed it as an independent species based on the fact that its nonpeloric flowers occassionally found

in a peloric population in Jungurahua of Ecuador are dissimilar in lip to those in P. cauda-

tum. Garay (1979) considered it to be a peloric form of P. wallisii but maintained it at the

specific level. This is indeed a good example of taxonomic treatment of normally peloric form.

On the other hand, however, most of the regular-flowered entities in Orchidaceae are not

peloric but rather primitive forms, such as Neuwiedia, Apostasia and Thelymitra, of which no

less than 50 species have been reported since the eighteen century. They have never been regarded

as peloric forms. Unfortunately, this has been neglected by some botanists. For instance, a hypo-

thetically primitive orchid flower designed by Pijl & Dodson (1966) has a distinctly specialized

lip with a short spur. In fact, in addition to the aforementioned genera we have some more ex-

amples of normally regular-flowered orchids. Among them Archineottia is the most interesting.

This is a genus of four species, two of which are regular-flowered. Of special interest is that in

this genus and its ally, Neottia, one can find all steps of column evolution from a simple form

with stamen and style not fully united to a most complicated form in which they have well fused.

Archineottia has a very primitive column, on which neither rostellum nor clinandrium is found

but a terminal and undifferentiated stigma (Fig.2: 2, 4, 6, 8). In addition, there exists on the

back of the column a thick ridge with its upper end joining the filament with which it is of

same texture. It is obviously the lower part of the filament which has been adnate to the style

(column). In Neottia, however, the column is much more advanced and very typical among the

family. It has a very large rostellum and most complicated stigma structure (Fig. 10, 12, 14, 16,

18). One of the most interesting examples is Neottia acuminata, in which the stigma even

becomes lamellate and almost backwards clasps the erect rostellum, but the perianth is more or

less regular with its lip entire and somewhat similar to, but shorter and wider than, the petals.

In these two genera there are altogether three species, namely Archineottia gaudissartii, A mic-

roglottis and Neottia acuminata, possessing regular or nearly regular perianth (Fig. 2: 1, 3, 17).

They are obviously not peloric forms. We can not imagine, indeed, that a complicated form

like Neottia acuminata or its allies would degenerate step by step into a simple form, and finally

into a peloric form. Archineottia belongs to the subtribe Listerinae, which is closely related to

Limodorinae, a rather primitivs subtribe with some genera possessing single pollen grain, relati-

vely few and long chromosomes and monocotyledonous habit. Apparently, there is nothing sur-

prising in the occurrence of some normally regular-flowered taxa, such as Archineottia, Diplan-

drorchis, Tangtsinia and Sinorchis, in these two primitive subtribes.

Another instance is Aceratorchis, a genus formerly included in Orchis, from which it is dis-

tinguished by the entire lip which is more or less similar to the petals. Strictly speaking, howe-

ver, its flowers are not truly regular. Two species have been described in this genus, but they

were recently considered as conspecific. Aceratorchis tschiliensis is widely distributed from

Hebei through Qinghai and Sichuan to northwestern Yunnan. It is cross-pollinated and produces

seeds efficiently. All these indicate its normally primitive taxon, instead of peloria. It may be

noted here that Asia is rich in members of Orchidioideae, as well as its primitive representatives.

The occurrence of a normally regular-flowered form in Asia, whether representing primitive

form of Orchis or Orchidioideae, is imaginable.

In Orchidaceae, as mentioned above, regular flowers are not only found in some primitive

taxa and peloric forms, but also in a few advanced groups. For example, a close investigation

by the senior author (Chen 1979) on Satyrium ciliatum revealed that this species has hermap-

hrodite, staminate and pistallate forms, for which no less than nine names have been published.

The flowers of its pistallate form are almost regular, in which nothing is found but three

similar petals and an elongate style with three stigmatic lobes at its top (Fig. 2: 19).

It is interesting to note that floral reversions in Orchidaceae are not always in connection

with peloria. For example, Epidendrum triandrum of North America represents another kind

of reversion. It is a reversal to abnormal polymery of stamens and not to abnormal regularity

of perianth. Like Phragmipedium lindenii, it is also hereditary. We may give it a new name

“Polyandrism” or something else, but, in fact, there is no essential distinction of this kind of re-

version from peloria.

It deserves mentioning that most of the regular-flowered entities, including primitive, ad-

vanced and peloric ones, occur in Asia and Australasia, where the Orchidaceae may have ori-

ginated as pointed out by some botanists. We have good reason to verify the primitiveness and

normality of many regular-flowered entities, but there exists no sufficient evidence for the im-

possible existances of normally regular-flowered species in those like Dendrobium, Eria, Lecanor-

chis, etc. For instance, Lecanorchis javanica, Dendrobium atavus and the new species described

here are considered to be peloric forms, but it is only a conjecture, for no reason can be given

for it. It is not impossible that some so-called peloric forms may prove to be truly primitive ones

in the future. Of course, a closer investigation is needed.

Summarizing the above, we may come to the following conclusions:

1. Regular or nearly regular perianth is a normal characteristic of orchids. It is chiefly

found in some primitive taxa and sometimes also in certain peloric forms and advanced groups.

Regular-flowered entities may not necessarily be peloric forms.

2. There exist two different types of peloria in Orchidaceae. One is abnormal form, with

its peloric flowers appearing at random. The other is “normal” form, with its individuals all

possessing peloric flowers. The latter is inheritable and can produce seeds efficiently, It would

be best to treat it as an independent species taxonomically, especially when its origin is uncertain.

3. Although peloria has been considered to he a reversal as a whole, conditions vary from

plant to plant. Some peloric forms have petal-like lip, and others have labellum-like petals.

Sometimes the same plant produces different kinds of peloric flowers in different years, sometimes

peloric flowers do not reappear upon the same plant. A few species can produce both peloric

and normal individuals, but others produce peloric forms only. Peloria is in fact a term only

used to cover the phase in which lip becomes similar to the petals. It is never all-embracing.

We recognize the existance of peloria in Orchidaceae, but great care must be taken to distinguish

truly peloric form from normally primitive one. It must be admitted that what causes peloria

and even what is peloria are still problems awaiting solution.

Acknowledgments: Our heartfelt thanks are due to Dr. Leslie A. Garay, Curator of the

Orchid Herbarium of Oakes Ames, Botanical Museum of Harvard University, for his valuable

suggestions during the preparation of this paper. We are also indebted to the artists, Mrs. Chun-

rung Liu and Mr. Chao-zhen Ji of our department, for their preparing the fine drawings.

Key words: Eria mêdogensis, Eria coronaria, Orchidaceae, Phragmipedium lindenii, Archineottia gaudissartii, Archineottia microglottis, Neottia acuminata, Aceratorchis, Satyrium ciliatum

陈心启, 吉占和. 墨脱毛兰，一个可能的“异常整齐花”型，兼论兰科中“异常整齐花”现象[J]. 中国科学院大学学报.

Chen Sing-Chi, Tsi Zhan-Hue. Eria mêdogensis, a Probably Peloric Form of Eria Coronaria, with a Discussion on Peloria in Orchidaceae[J]. .

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墨脱毛兰，一个可能的“异常整齐花”型，兼论兰科中“异常整齐花”现象

Eria mêdogensis, a Probably Peloric Form of Eria Coronaria, with a Discussion on Peloria in Orchidaceae

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