Botanical Studies (2012) 53: 509-524.
SYSTEMATICS
The diversity of pistillate flowers and its taxonomic value to the classification of Daphniphyllum (Daphniphyllaceae)
Mo-Shih TANG1,2, Yuen-Po YANG1'3, Chi-Chu TSAI4,5, and Chiou-Rong SHEUE6 *
1Department of Biological Sciences, National Sun Yat-sen University, Kaohsiung 804, Taiwan
2Deaprtment of Pharmaceutical Science and Technology, Chung Hwa University of Medical Technology, Tainan 717, Taiwan
3Department of Bioresources, Dayeh University, Changhua 515, Taiwan
4Kaohsiung District Agricultural Improvement Station, Pingtung 908, Taiwan
5Department of Biological Science and Technology, National Pingtung University of Science and Technology, Pingtung 912, Taiiwan
6Department of Life Sciences, National Chung Hsing University, Taichung 402, Taiwan (Received April 12, 2012; Accepted June 11, 2012)
ABSTRACT. Daphniphyllum is the sole genus of the family Daphniphyllaceae consisting of dioecious spe­cies with inconspicuous reduced unisexual mostly caducous flowers. The paucity of helpful taxonomic char­acters from vegetative and reduced reproductive organs has led to difficulties in intrageneric classification and species identification. The pistillate flowers have characters of higher taxonomic value than those of the vegetative organs, staminate flowers and fruits, but pistillate flower information is rarely reported. Here pistil­late flowers of 19 taxa were studied to reveal pistillate floral diversity and major developmental features with a view to intrageneric classification. Four types of stigma shape were recognized: punctiform, reniform, linear and oblong. Daphniphyllum stigmas, except for the punctiform type, elongate variously after an early stage of anthesis. The pistillate flowers of three taxa are asepalus. The calyces of other taxa were categorized into three types, free, cleft and parted, which are consistent within taxa. The existence and persistence of staminodes are diagnostic characters. This study shows that the characteristics of pistillate flowers provide helpful information for identification at the species and even the variety level. Our results do not support the currently accepted two sections, Daphniphyllum and Lunata. Instead, Huang's original intrageneric classification into three sec­tions, Daphniphyllum, Lunata and Staminodia, is supported, but with two species previously in Daphniphyl­lum reassigned to Staminodia.
Keywords: Caducous; Calyx; Daphniphyllum； Lunata; Staminode; Staminodia; Stigma; Taxonomic value.
INTRODUCTION
Daphniphyllum is the sole genus of the family Daphni­phyllaceae distributed from tropical to subtropical Asia. The genus Daphniphyllum was established by Blume (1826), based on the type species, D. glaucescens. Mono­graphic studies on Daphniphyllum have been made by Muller (1869), Rosenthal (1916, 1919) and Huang (1965, 1966). Currently, this genus comprises about 30 species of evergreen trees or shrubs with a distributional range from tropical to subtropical Asia (East Asia, Southeast Asia, and Indian subcontinent), generally between 46°N to 10°S and 75°E to 150°E (Huang, 1965; Kubitzki, 2007).
Intrageneric classification was first proposed by Huru-sawa (Hurusawa, 1942a, b) who divided Daphniphyllum
*Corresponding author: E-mail: crsheue@gmail.com (Chiou-Rong SHEUE).
into two sections, Staminodia and Calycifera, based on four species in Japan (including Taiwan). He distinguished these two sections based on pistillate flower features: section Staminodia with obvious staminodes; section Calycifera with very obscure or trace of staminodes and persistent or deciduous sepals. Huang (1966) described nine species in his monograph of Daphniphyllum, and divided this genus into three sections, namely Daphniphyl-lum, Lunata and Staminodia. More recently, 16 species of Daphniphyllum were described in the Flora of Malesiana by the same author (Huang, 1997). However, he revised his treatment of this genus by dividing it into two sections (Lunata and Daphniphyllum), and relegated the previous section Staminodia to a subsection of section Daphniphyl-lum (Huang, 1996).
The flowers of Daphniphyllum are unisexual (Heywood et al., 2007), wind-pollinated (Fishbein and Soltis, 2004) and inconspicuous. The flowers of all species are arranged into a racemose inflorescence and are often clustered near
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the apices (Heywood et al., 2007). Bracts at the base of the inflorescence and the bracteole at the base of the flower are almost ovate to elliptic and deciduous (Huang, 1997). Sepals vary from asepalous, aposepalous to connate calyx (Huang, 1997). Petals are lacking in all cases (apetalous) (Huang, 1997; Kubitzki, 2007). The pistil is normally vase-shaped, and has a bi-lobed stigma connected with a short style on the top of a bulbous ovary.
The characters of pistillate flowers of Daphniphyllum display relative higher taxonomic value than others, in­cluding vegetative features (Croizat and Metcalf, 1941; Huang, 1965; Wang, 1981; Kiew and Rafidah, 2008), staminate flowers or drupes (Croizat and Metcalf, 1941; Huang, 1965), which were traditionally used, but have little taxonomic value. Huang (1997) mentioned the diver­sity of stigma morphology: divaricate, revolute, discoid to circinnate or versatile morphology in species of Daphni-phyllum in the Flora Malesiana. In addition, features of the calyx, such as presence, persistence and serration, have been frequently used for taxonomy for this family by various authors (Rosenthal, 1919; Chien, 1933; Huang, 1966; Kiew and Rafidah, 2008; Ming and Kubitzki, 2008). Huang (1996, 1997) used more characters of the calyx, such as number of calyx lobes and calyx articulation, for the taxonomy of the genus. Rosenthal (1916) distinguished taxa by the relative length of the stigma and ovary. Huang (1966) also utilized this character to differentiate three subspecies from the glaucescens subcomplex of subsec­tion Daphniphyllum. Kiew and Rafidah (2008) combined the features of the infructescence, fruit surface, fruit stalk length and stigma persistence to distinguish three varieties of D. glaucescens.
Although the taxonomic value of pistillate flowers of Daphniphyllum was noted (Hurusawa, 1942a, b; Huang, 1966, 1996; Kiew and Rafidah, 2008), several original papers ignored this character altogether, which resulted in difficulties in species identification (Teijsman, 1864; Hooker, 1890; Chien, 1933). Probably this oversight is due to the fact that the stigma and staminode of Daphni-phyllum are often caducous at an early stage of anthesis in most taxa, leading to extreme underreporting of pis-tillate flower information (M. S. Tang, pers. observ.). It is apparent that the morphology of pistillate flowers of Daphniphyllum needs more careful observation. Here, we present a comprehensive comparative study of pistillate flowers, documenting major developmental traits (such as elongation after anthesis, persistence or caducousness) of Daphniphyllum, and floral traits to serve as taxonomic characters. These data are informative to the classification of this poorly understood family.
MATERIALS AND METHODS
A diagram with 34 described taxa of Daphniphyllum (Huang, 1965, 1996; Hatusima, 1971; Ming and Kubitzki, 2008; Kiew and Rafidah, 2008) following Huang's in-trageneric classification (Huang, 1996) is displayed as
Figure 1. In this study, we sampled 19 taxa of Daphniphyl-lum (Figure 1, in bold) during 2006 to 2012, in which two species represented the section Lunata, and 14 species and three varieties represented the section Daphniphyllum. The sources of pistillate flowers included both fresh materials and herbarium specimens (Appendix 1). The fresh materi­als of seven taxa were collected in the field or obtained from botanical gardens in China, Indonesia, Japan, Malay­sia, Philippines and Taiwan (Appendix 1, in parentheses after the species name). The fresh pistillate flower mate­rials (c. 30 samples) include flower buds and flowers at various stages from 3 individuals from each of five popu­lations of native Taiwan taxa. Due to the difficulty of ob­taining materials of pistillate flowers of non-Taiwan taxa, for some of these taxa only one individual was sampled. Voucher specimens were deposited at the Herbarium of the Department of Biological Sciences, National Sun Yat-sen University (SYSU). Herbarium specimens at BM, BO, CDBI, HAST, IBSC, K, KEP, KUN, L, MEL, NTUF, PE, PNH, PPI, PYU, SING, SNP, SZ, TAI, TAIF, TI, TNM and TNS were examined and photographed (Appendix 1, fol­lowing its species name).
Fresh materials were fixed in FPGA (formalin: propi­onic acid: glycerol: 95% ethanol: distilled water=5: 5: 15: 35: 40 in volume) in the field for observation. After fixation, they were examined with a Zeiss Stemi SV11 stereoscope and photographed with a Nikon D70s digital camera. They were also directly mounted on stubs and ob­served with a Hitachi TM-1000 Tabletop microscope. For herbarium specimens, pistillate flowers were photographed with a Nikon CoolPix 5700 or D70s digital camera.
RESULTS
The characters (including style, stigma, staminode and calyx) of pistillate flowers of 19 studied taxa of the genus Daphniphyllum are summarized in Table 1 follow­ing Huang's intrageneric classification (Huang, 1996). To clearly describe stigma and style of a pisllate flower of Daphniphyllum, here we define and distinguish these structures by their cell types. The stigma observed from fresh material is composed of colliculose epidermal cells located on the adaxial surface of the top of the pistil, while the epidermal cells of the style and the ovary of the pistil are flat and elongated, with occasional stomata (Figure 2A). Because the stigma of Daphniphyllum is bi-lobed connected with a short style, the segment of the pistil with colliculose epidermal cells above the fork is defined to be the stigma in this study. The style is the tissue between the ovary and the fork. Based on these results, the characters of the stigma, calyx and the staminode of pistillate flowers can serve as aids for classification (Figures 2-3).
Stigma
Based on our investigation, the features of stigmas at an early stage of anthesis, i.e. before the stigma is mature (before the cells become colliculose in form) or before the
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Figure 1. Diagram with 34 described taxa of Daphniphyllum (Huang, 1965, 1996; Hatusima, 1971; Ming and Kubitzki, 2008; Kiew and Rafidah, 2008) showing a cladogram based on Huang's intrageneric classification (Huang, 1996). Studied species are indicated in bold.
bracteole is lost, are stable in all studied taxa. At this early stage, four types of stigma can be differentiated (Table 1): punctiform (Figure 3A-B), reniform (Figure 3C-G), linear (stigma length longer than the ovary length) (Figure 3H-J), and oblong (stigma length about 2/3 shorter than the ovary length) (Figure 3K-P). Stigmas of the latter three types will elongate after an early stage of anthesis, but only those of the punctiform type are unchanged. However, the stigma elongation tendency varies between these three types. Ob­long stigmas strongly elongate to become longer than the ovary, and become distinctly bent (Figure 3M, O, P), re­sulting in a shape resembling a linear stigma. In addition,
reniform and oblong stigmas expand after an early stage of anthesis. Reniform stigmas show a higher tendency for expansion after anthesis than oblong stigmas. Because of such elongation and expansion, reniform stigmas gradually flatten, elongate and slightly bend after anthesis.
Among the various types of stigma, only linear stigmas can either persist or drop off after anthesis. Stigmas of the other three types are always persistent. Two taxa (D. an-gustifolium and D. glaucescens var. blumeanum), with lin­ear stigmas, were observed to have an articulation appear at the base of the styles at an early stage of anthesis due to the presence of an abscission zone in the base of the style.
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This caducous stigma in these two taxa often leaves a flat trace on the top of the style in specimens.
Style
Styles in the genus Daphniphyllum are absent or short. The two species of section Lunata, D. calycinum and D. laurinum have absent or short styles (Figure 3A-B). The pistillate flowers of other studied taxa have short styles.
Style length may vary in the same taxon. Therefore, the style length is not helpful for classification in Daphniphyl-lum.
Staminode
There are five to 12 staminodes in a pistillate flower of some taxa of Daphniphyllum, but this number is not generally stable at the individual level and so has little taxonomic value. Based on observation of fresh material in four taxa studied, the staminodes of Daphniphyllum are spoon-shaped (D. glaucescens var. blumeanum), scale-like (D. borneense, Figure 2B), or cylindrical (D. macropodum var. macropodum and D. macropodum var. humile, Figure 2C). The length of staminodes varies in a pistillate flower, which indicates that this feature is not useful for species differentiation. These staminodes appear as scale-like structures of various sizes in desiccated specimens (Figure 3H, M, N).
Staminodes were found consistently in 11 taxa. Only two taxa of section Lunata and five taxa of section Daphn-iphyllum have no staminodes (Table 1). If present, the staminode inserts around the base of an ovary, in which it often occurs intracalycinely. The presence of staminodes in a pistillate flower of Daphniphyllum is stable at the spe­cies level, except in D. macropodum var. macropodum. The pistillate flower of D. macropodum var. macropodum in Japan and China consistently produces staminodes (Fig­ure 2C). However, flowers of this taxon in Taiwan may or may not have staminodes (Figure 2D).
The staminodes are caducous in all studied taxa, except D. macropodum (including the two varieties). The cadu­cous staminode leaves a round scar at the base of the ova­ry. Careful examination is needed to observe this feature, especially in herbarium specimens. However, the presence of staminodes has potential as a taxonomic character.
Calyx
Most of the studied taxa have a calyx. Only three taxa, D. angustifolium, D. longeracemosum and D. macropo-dum var. humile are constantly asepalous. According to the degree of connection of calyx lobes, three calyx types are recognized: free, parted and cleft. This feature is consistent within a species, but the shape and serration of calyx lobes vary intraspecifically.
The free type is found in D. buchananiifolium and D. woodsonianum. It is aposepalous with 2 or 3 sepals and is always lost in early anthesis (Table 1). Eleven studied taxa have a parted calyx. It consists of 4-8 valvate or im­bricate sepals slightly connate at the base and is often lost by anthesis. This parted calyx can be further differentiated into three forms by the length of the calyx lobe: disciform, oblong and linear. The disciform lobe is shorter than 1/3 of the ovary (D. paxianum); the linear lobe is longer than the ovary (D. gracile); the oblong lobe is longer than 1/2 of the ovary, and shorter than the ovary (10 taxa). The cleft type has 4-6 sepals connate at 1/3 to I/2 of the sepal length, forming a calyx tube and often persistent in mature fruit.
Figure 2. Scanning electron micrographs of pistillate flowers of Daphniphyllum. (A) D. oldhamii var. oldhamii. The inset shows colliculose epidermal cells of the stigma; (B) D. borneense with its parted type calyx and staminodes; (C) D. macropodum var. macropodum with its free type calyx and staminodes; (D) D. macropodum var. macropodum without staminodes and asepalous; (E) D. oldhamii var. oldhamii with its parted type calyx and the absence of staminode. Scale bars: A= 50 (im, B-D =1 mm.
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Figure 3. Pistillate flowers ofi Daphniphyllum. A,D. calycinum; B, D. laurinum; C, D. buchananiifolium;muD. macropcfum vmu macropodum; H,D.macropomumYPi. humile;OD. woodsonianum; G, D. glaucescens subsp. oliUhamiimau lanyuense;n,D.umauce-scens var. glaucescens; I, D. glaucescens var. blumeanum; J, D. luzonense; K, D. borneense; L, D. gracile, M, D. neilgherrense, N, D. scortechinii, O, D. oldhamii var. oldhamii, P, D. teijsmannii. Bot534-10-5： punctiform stigma, Bot534-10-6: reniform stigma, Bot534-10-7: oblong stigma, Bot534-10-4: linear stigma, : staminode, Bot534-10-9: cleft sepal, Bot534-10-10: free sepal, Bot534-10-9: parted sepal. A, C, D, E, G, I, K and O were from fresh materials, B, F, H, J, L, M, N and P from herbarium specimens. All the images except M, O and P were before anthesis without elongation.Owasinelongation,andMandP alreadyshrankafterelongation. Arrowsindicatestaminodes. Allscale bars =1mm.
This type includes D. cafycinum (Figure 3A) and D. lauri­num (Table 1). While the populations of D. macropodum var. macvopodum in Taiwan are asepalous type (Figure 2D), the populations of this species in Japan are the free type (Figure 3C) or asepalous.
DISCUSSION
This study is the first comprehensive report on pistil­late flowers of the Daphniphyllaceae including the major developmental events. Of most significance is information
on the varying degree of sti gma elongation afteranthesis in itpee m^pes of stihma. This informationmeans that stig­ma type can only be used as a taxonomic character before early anthesis of a pistillate flower, when the calyx has just opened. This study also shows that persistence of stigma and staminode have taxonomic value. The diverse and in­formative pistillate flowers revealed here provide helpful characters for taxonomy.
Four stigma types with high taxonomic value were rec­ognized in this study. These characters facilitate differen­tiating taxa at the species level or even the variety level.
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For example, two varieties of D. oldhamii present differ­ent types of stigmas: the oblong type in var. oldhamii and the reniform type in var. lanyuense. Although the shape of the leaf apex and fruit size have been used to differen­tiate these two varieties (Huang, 1992), in fact, these fea­tures often vary between populations or are influenced by the environment. In contrast, stigma type can distinguish them reliably.
In addition, persistence of the stigma can serve as an aid to differentiate varieties. Careful observation is needed to apply this feature. According to our observations, the top surface of a caducous stigma usually leaves a smooth and clear flat trace, while that of a broken stigma often dis­plays a rough and irregular trace. Thus, we can distinguish a caducous stigma scar from a damaged stigma scar, which might be due to the process of specimen pressing or col­lecting. Kiew and Rafidah (2008) compared pistillate flow­ers of three varieties of D. glaucescens (var. blumeanum,
var. glaucescens and var. lancifolium). They reported that two of these varieties (var. glaucescens and var. blumea-num of D. glaucescens) have caducous stigmas. These two varieties were examined in this study. However, our assessment is that only one of these two varieties has ca­ducous stigma (var. blumeanum). We found that the stigma trace of D. glaucescens var. glaucescens in all studied speciments was rough and irregular with stigma residue of various lengths, indicating a damaged stigma. Thus, the stigma of D. glaucescens var. glaucescens is not caducous.
The linear, reniform and oblong types of stigma are only stable at early anthesis, because the stigma elongates, bends or forms an articulation, after early anthesis. After that stage, all of three types of stigmas display very similar morphology. Usually, they appear column-like, slightly revolute, in the investigated specimens. This character change leads to misidentification of taxa by some authors. Previously the identification of four species (D. glauce-
Table 1. The comparison of morphological characters of pistillate flowers of Daphniphyllum, the classification followed Figure 1.
Section/
subsection
Stigma
Calyx
Staminode state
Taxon
Style
Type
Elon.
Type
Persistence in fruit
Distribution
Lunata
D. calycinum
n-s
punctiform (P)
-
cleft
P
-
EA
D. laurinum
n-s
punctiform (P)
-
cleft
P
-
SEA
Daphniphyllum
D. buchananiifolium
s
reniform (P)
free
-
C
SEA
Daphniphyllum
D. woodsonianum
s
reniform (P)
+
free
-
C
SEA
D. glaucescens subsp. oldhamii
s
reniform (P)
+
parted
-
-
EA
var. lanyuense
D. oldhamii var. oldhamii (syn. D.
s
oblong (P)
++
parted
-
-
EA
glaucescens subsp. oldhamii var.
oldhamii)
D. teijsmannii
s
oblong (P)
++
parted
-
-*
EA
D. paxianum
s
oblong (P)
++
parted
P
-
EA
D. gracile
s
oblong (P)
++
parted
-
C
SEA
D. borneense
s
oblong (P)
++
parted
-
C
SEA
D. neilgherrense
s
oblong (P)
++
parted
-
C
I
D. scortechinii
s
oblong (P)
++
parted
-
C
SEA
D. glaucescens var. glaucescens
s
linear (P)
+
parted
-
C
SEA
D. glaucescens var. blumeanum
s
linear (C)
+
parted
-
C
SEA
D. luzonense
s
linear (P)
+
parted
-
C
SEA
Staminodia
D. macropodum var. macropodum
s
reniform (P)
+
free/-
-
P/-
EA
D. macropodum var. humile
s
reniform (P)
+
-
P
EA
D. longeracemosum
s
reniform (P)
+
-
-
EA
D. angustifolium
s
linear (C)
-
-
-
C
EA
*Only the type specimen of D. teijsmannii shows staminodes. See Discussion for more detail.
Abbreviations: －: absent; +: slight elongation; ++: elongation; n: lacking; s: short; C: caducous; P: persistent; EA: East Asia; Elon.: elongation; I: Indian subcontinent; SEA: Southeast Asia.
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scens, D. neilgherrense, D. oldhamii and D. teijsmannii) was confused by many authors (Hooker, 1890; Hayata, 1904; Chien, 1933; Hurusawa, 1942b), because these species are similar in vegetative characters and have an elongating stigma. However, this study shows that the ap­plication of stigma types and the presence of staminodes can successfully differentiate these taxa. Additionally, Huang (1965) described D. himalaense subsp. angustifoli-um (currently D. angustifolium) with a short discoid style, which was the same as the punctiform stigma of section Lunata. According to our observation, D. angustifolium actually has a short style with a conspicuous, erect and linear stigma, but the stigma is caducous by forming an abscission zone near the base of the style at early anthesis.
The staminodes in the pistillate flowers had been inter­preted as glands by a few authors, such as Muller (1869), Baillon (1858) and Hooker (1890). However, Rosenthal (1919), in his monograph, argued that these glands actu-ally are staminodes and only occur in three species (D. celebense, D. himalayense and D. macropodum). Decraene and Smets (2001) said "trying to recognize staminodes or any other structures surrounding the ovary, if no structural evidence of their homology with stamens exists, is sense-less." Since there is no study on staminodes of Daphni-phyllum with from ontogenetic point of view, a further developmental study is needed. Here, we tentatively accept these structures as staminodes according to their shape and location. Regardless of their true nature, this study shows that they do have taxonomic value.
All the taxa distributed in Southeast Asia of subsec­tion Daphniphyllum have caducous staminodes, but the taxa distributed in East Asia have no staminodes. Huang (1966) reported that the section Daphniphyllum (currently a subsection of section Daphniphyllum) has prominent staminodes in tropical Asia and this character is reduced in temperate Asia. Our observations confirm his report. The plants of D. teijsmannii in Japan have no staminodes, but the type specimen has distinct staminodes. This type specimen was collected from a cultivated individual, which was initially introduced from Japan to the Bogor Botanical Garden in Java (Teijsman and Binnendijk, 1864; Huang, 1965). It is possible that abnormal flowers with staminodes might have been induced by its cultivated environment. Furthermore, the variety, D. teijsmannii var. amamiense, was named based solely on a single type specimen possessing minute staminodes collected from the Ryukyu Islands (Hurusawa, 1942b, 1954). However, this variety was soon synonymized as D. teijsmannii (Ha-tusima, 1971), since the presence of staminodes in the type specimen appears to be a rare exception. Staminodes have not been found in other populations of D. teijsmannii. Sev­eral specimens of D. macropodum with abnormal bisexual flowers were observed in the herbarium of Kew, which were collected from introduced plants cultivated at Kew Garden (Tang, personal observation). Based on these phe­nomena, the presence of staminodes, and their size, might be occasionally influenced by environmental factors.
Many authors have regarded the morphology of the calyx (presence, caducous or persistent, degree of con­nection) as providing the most effective characters for taxonomic recognition of species within Daphniphyllum (Muller, 1869; Rosenthal, 1919; Chien, 1933; Croizat and Metcalf, 1941; Hurusawa, 1942a, b; Huang, 1966; Ming, 1980; Wang, 1981; Huang, 1996; Ming and Kubitzki, 2008). Huang (1965) mentioned that the serration of sepals is useless in species distinction. Hurusawa (1942b) classi­fied six varieties of D. teijsmannii (included D. oldhamii) with the shapes of sepals and leaves. Our observation sup­ports Huang's opinions. Except in D. macropodum var. macropodum, calyx type was found to be stable intraspe-cifically in this study. The pistillate flower of D. macropo-dum var. macropodum was found to be either asepalous or have a free calyx. Because the free type of sepal is always caducous at early anthesis, it recommended that fresh materials with calyx features be obtained before the loss of the bracts from the inflorescence (caducous, free sepals and bracts will drop off around the same time). Because they are caducous, the free calyx is the most difficult fea­ture to observe in herbarium specimens. Therefore, many taxonomists, except Hurusawa (1942a) and Huang (1996), have ignored this feature and have seldom used it in tax­onomy of Daphniphyllum.
This study shows that the types of calyx are important for taxonomy, especially for the distinction of section levels of Daphniphyllum (Table 1). Therefore, the care­ful examination of pistillate materials is required. Section Lunata (D. calycinum and D. laurinum) has only the cleft calyx. Section Daphniphyllum has three types of calyx. The asepalous or free calyx is possessed by the subsection Staminodia. Moreover, the other two taxa of this subsec­tion (D. parvifolium, D. chartaceum) (Huang, 1966; Ming and Kubitzki, 2008) are found to have free calyces. The majority (12 studied taxa) of subsection Daphniphyllum has the parted calyx. Only two taxa (D. buchananiifolium and D. woodsonianum) show free calyces. They also have reniform stigmas. These two features are also present in subsection Staminodia. Consequently, D. buchananiifo-lium and D. woodsonianum may have a closer relationship with subsection Staminodia than with subsection Daphni-phyllum.
The informative pistillate flowers (including stigma type, persistence of stigma, stigma elongation, calyx type, persistence of calyx in fruit, prescence of staminode) of Daphniphyllaceae contribute greatly to identification and classification of this family of dioecous plants. However, cautious efforts to examine these inconspicuous features with attention of developmental change, and to distinguish caducous traces from damaged organs in a pistillate flow­er, are necessary. The results of this study support Huang's original intrageneric classification of this genus into three sections (Huang, 1966), Daphniphyllum, Lunata and Sta-minodia. However, two taxa of section Daphniphyllum as mentioned above, D. buchananiifolium and D. woodso-nianum, are suggested to be reassigned to section Stamino-
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dia. We provide a Key below to our revised intrageneric classification of Daphniphyllum. As for intrasectional classification of Daphniphyllum, these results do not show consistent grouping relationships in accordance with Huang's intrasectional classification (1996). Further
studies should be done to address species that were not obtained for the present study. Moreover, a combination of multiple data will be needed for a full understanding of species diversity, intraspecific taxa, and the obscure in-trageneric relationships of Daphniphyllaceae.
A key to the sections and studied taxa of Daphniphyllum based on the features of pistillate flowers
1a. Sepals absent or free.............................................................................................................................Section Staminodia
2a. Stigma linear.......................................................................................................................................D. angustifolium
2b. Stigma reniform
3a. Staminodes absent .................................................................................................................. D. longeracemosum
3b. Staminodes present
4a. Staminodes persistent .........................................................D. macropodum (var. macropodum, var. humile)
4b. Staminodes caducous ...................................................................... D. buchananiifolium, D. woodsonianum
1b. Sepals connate
5a. Calyx cleft; stigma punctiform..............................................................Section Lunata (D. calycinum, D. laurinum)
5b. Calyx parted; stigma not punctiform.....................................................................................Section Daphniphyllum
6a. Stigma reniform........................................................................... D. glaucescens subsp. oldhamii var. lanyuense
6b. Stigma linear or oblong
7a. Stigma linear, longer than ovary after anthesis
8a. Stigma caducous .................................................................................... D. glaucescens var. blumeanum
8b. Stigma persistent............................................................D. glaucescens var. glaucescens, D. luzonense
7b. Stigma oblong, about the same length of ovary after anthesis
9a. Staminodes absent ........................................... D. oldhamii var. oldhamii, D. teijsmannii, D. paxianum
9b. Staminodes present ..................................... D. gracile, D. borneense, D. neilgherrense, D. scortechinii
Acknowledgements. The authors thank the curators of the herbaria BM, BO, CDBI, HAST, IBSC, K, KEP, KUN, L, MEL, NTUF, PE, PNH, PPI, PYU, SING, SNP, SZ, TAI, TAIF, TI, TNM, TNS for allowing access to their col-lections, Tutie Djarwaningsih (Indonesia), Jean W. H. Yong (Singapore), T. M. Balangcod, R. Hipol (Philippines), C.-h. Li, Y. He, Z.-R. He (China), Y. Ito, N. Tanaka (Japan), Ra-fidah, H. Mujih, A. Bium, R. C. K. Chung (Malaysia), M.J. Wu, S. H. Yen and L. C. Shih (Taiwan) for contributing or helping to collect plant materials, T.C. Huang for pro-viding valuable discussion, P. Chesson for improving the manuscript, and D. Boufford and two anonymous referees for providing comments and advices on the manuscript. The project was supported by the National Science Coun-cil, Taiwan (NSC 95-2621-B-110-002).
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Appendix 1. Specimens observed and voucher specimens of fresh pistillate flowers of Daphyniphyllum in this study.
D. angustifolium Hutch.
CHINA. Guangxi: Cangwu, S.H. Chen 10372 (PE). Guizhou: Longtanhu, S.H. Chen 1386 (IBSC). Shensi: Cheng-kuhsien, T.N. Liou 11354 (SZ). Hubei: Badon, G.X. Fu & Z.S. Jhang 972 (IBSC, PE). Fang Hsien, Wilson 2959 (BM, K). Szechuan: Chengkou, T.L. Tai 107049 (SZ), 107504 (CDBI, PE). Fanjingshan, D.H. To 3809 (IBSC). Nanchuan, W.P. Fang 956 (IBSC, K), 957 (IBSC). Omei Mount, D.H. To 158 (IBSC), 221 (IBSC, PE, SZ); W.P. Fang 14039 (PE). Wushan, G.H Yang 59021 (CDBI, IBSC). Wuxi, G.H Yang 58691 (IBSC), 59418 (CDBI, IBSC); Y.D. Chen 2009 (PE). Yunjin, J.W. Yao 2263, 3753 (PE, SZ).
D. borneense Stapf (fresh materials: A. Biun et al. T970, T971, T972, T973, T1550 (SYSU)).
MALAYSIA. Sabah, Anthea & Tan 1601; B. Perumal 105 (SNP); B.E. Snythies S.10645; C.E. Carr 27544, 27633 (SING); Chew & E.J.H. Corner RSNB5886 (TI, K); D. Hou 250 (BO, L); G.D. Haviland 1070K (L, BM); J.M.B. Smith 476 (L); J. Nais & Dolois S.3285; J. Sinclair (SING); K. Kitayama 4910 (SNP); M.S. Clemens 10656; M.S. Clemens
et al. 27821 (BO); M. Togashi 6861 (TI); Nooteboom 4607 (BO); Pns 2333 (SNP); Rao 80 (SING); RSNB 761 (BO);
S. Kokawa & M. Hotta 3378 (TI); W.L. Chew et al. 761 (SING), 5886 (L, SING, TI); W. Meijer SAN21051 (BO, KEP, PNH, SING); Y.F. Lee et al. SAN69984 (L). Sarawak, I. Paie S26536 (K, SING).
D. buchananiifolium Hallier f. (fresh materials: M.S. Tang T917, T918 (SYSU)).
PHILIPPINES. Agusan, D. Mendoza & P. Convocar 861 (BM, PNH); M. Ramos & Convocar 83733 (K, SING). Benguet, Williams 1337 (K). Biliran Is., M.D. Sulit 5573 (L, PNH). Leyte, M. Ramos 15235 (BM, K). Luzon, A.D.E.
Elmer 8538 (BO, K, L); Barbon et al. 8278 (KEP); E.D. Merrill 1748 (BO, BM, L); G. Edano 40478 (L), 28662 (BM, BO, K); M. Jacobs 7252, 7288 (K, L, PNH, SING), 7556 (K, SING); M. Ramos 1503 (BM, L, SING), 23478 (BM, K,
L); M. Ramos & G. Edano 28705 (K). Mindanao, A.D.E. Elmer 11344 (BO, L), 11671 (BM, BO, L); D. Mendoza & P. Convocar 10814 (K, PNH). Without precise locality, M. Jacobs q252, q556 (L); M. Ramos & P. Convocar 83733
(NTUF, PNH).
518
Botanical Studies, Vol. 53, 2012
D. calycinum Benth. (fresh material: S.H. Yen T957 (SYSU)).
CHINA. Fujian: Fuzhou, G.D. Ye 1622 (PE). Huaiji, Y.G. Liou 3014 (PE). Liancheng, R. Lin 3236 (PE). Nanjing, D. Sia 268; G.D. Ye 2277; R. Lin 4607 (PE). Ninghua, L.L. Lin 878 (PE). Pinghe, G.S. He 9603 (PE, TAIF). Xianyou, Y. Lin 458 (PE). Guangdong: Baiyunshan, T.J. Lai 68 (IBSC). Baoan, T. ZhongM123 (IBSC). Beijiang, H.U. Chen 6117 (IBSC). Chaoan, F.Y. Zeng et al. 1273 (TNM). Conghua, S.Q. Chen 17422 (TNM). Deqing, Y.G. Liou 919 (IBSC, PE). Dinghushan, G.L. Shih 12781; H.U. Chen 6428 (IBSC). Enping, M.S. Huang 110308 (PE). Fengkai, C. Huang 164130 (IBSC, PE); K.C. Ting & L. Yu 6418 (CDBI). Fengshun, H.G. Ye et al. 4242 (TNM). Gaoming, M.S. Huang 110700 (PE). Gaoyao, C. Huang 162287 (IBSC). Guangzhou, F.W. Sing et al. 436 (TNM). Heping, C.F. Wei 120249 (IBSC, PE, SZ); C.M. Tan 99571 (IBSC, TAIF). Heshan, M.S. Huang 110854 (PE). Huadu, C. Huang 164423 (PE). Huaiji, H.G. Ye et al. 3087 (TNM). Huiyang, C.F. Wei 121540; T.M. Tsui 117 (IBSC, PE). Jiaoling, L. Deng 4534 (IBSC, PE); Nanling Exped. 2255 (TNM). Jiexi, H.G. Ye et al. 4576 (TNM). Kechew, Y. Tsiang 2241 (SING). Lechan, Nanling Exped. 3423 (IBSC, TNM); W.T. Tsang 20879 (PE). Lian, T.P. Tan 59901 (IBSC, PE). Liannan, T.P. Tan 59410 (PE). Lianping, C.F. Wei 120012 (IBSC, PE). Lianshan, F.Y. Zeng et al. 286; H.G. Ye 3507 (TNM); T.P. Tan 58424 (PE), 59054 (SZ). Lianyang, Y.C. Cheng 3371 (IBSC). Lingnan, Tsang 16518 (IBSC). Lokchong, C.L. Tso 20409 (BO, TAI). Longchuan, B.H. Chen 1017 (IBSC). Longmen, C.F. Wei 121933 (IBSC, PE). Luoding, H.U. Chen 9512 (PE). Maoming, L. Deng 1892 (PE). Mei, Nanzhidi Exped. 7326 (IBSC). Nanxiong, L. Deng 6179 (PE). Pingyuan, S.G. Li 201951 (PE). Poklo, 2366 Exped 3170 (PE); L.S. Chen 41685 (IBSC, PE); Levine 7005A; S.P. Ko 52401 (PE); T.M. Tsui 79 (IBSC, PE); Y. Tsiang 1687 (BO, IBSC, PE). Qingyuan, C.K. Huang 30173; S.C. Liou 28120 (PE). Raoping, S.G. Li 200561 (IBSC, PE). Renhua, L. Deng 7469 (PE), 8329 (PE, SZ), 8730 (IBSC, PE). Ruyuan, S.P. Ko 52570 (PE); Yue 73198 (IBSC, PE). Sanwui, M.S. Huang 110933 (PE). Shenzhen, F.W. Sing 11540; S.W. Wang et al. 527 (IBSC, TNM). Shixing, S.W. Wang 367 (IBSC). Sinfung, Y.W. Team 1066 (HAST). Taishan, B.H. Chen 902 (IBSC). Wantongshan, T. M. Tsui 353 (PE). Wengyuan, S.C. Liou 24123 (PE). Wuhua, S.G. Li 201659 (IBSC, PE). Xinfeng, H.G. Ye 923; L. Deng 7968 (PE, SZ). Xinging, H.G. Ye 9880 (TNM); Y.G. Liou 2070 (IBSC, PE). Xinyi, Z. Huang 31080 (PE). Xuwen, Xuwen Exped. 163 (PE). Xuwenshan, S.R. Leung 65616 (PE). Yangchun, H.G. Ye 5348 (IBSC, TNM). Yangjiang, L.S. Chen 42764 (PE). Yangshan, L. Deng 1335 (PE); T.M. Tsui 431 (IBSC, PE). Yingde, C. Huang 163825 (IBSC, PE); S.R. Leung 60358 (PE); Y. Tsiang 1983 (IBSC, PE). Yunann, N. Liou et al. 2781 (IBSC, TNM). Yunfon, C. Wang s.n. (TAI). Yunfu, Z. Huang 489 (IBSC). Zengcheng, C.F. Wei 122059 (IBSC, PE, SZ). Zhanjiang, Z.X. Li 1907 (TNM). Zijin, C.F. Wei 120853 (IBSC, PE).
Guangxi: Cangwu, J.J. JhongA60907 (PE). Ceheng, Anonymous 82-0095 (PE). Daxin, Anonymous 235 (PE). Dayaos-han Yao Autonomous Zizhixian, C.H. Lyu 4376 (PE). Fusui, S.Q. Chen 12305 (PE). Gui, S.C. Jhong 50693 (PE). Le­chan, Z.X. Li 1529 (PE). Maoqiaolinchang, Guangxi Exped. 194 (PE). Rong, S.Q. Chen 9809 (PE). Wuming, J.J. Jhong 20922 (PE). Xingan, C.H. Lyu 2103; G.J. Li 15141 (PE); Guangxi Exped. 816; S.L. Yu 900483 (PE); Xingan Exped. 76 (PYU). Yanshan, S.L. Yu 900521 (PE). Yung Hsien, A.N. Steward & H.C. Cheo 769 (BO, SING). Zhaoping, Anony­mous 604185 (PE). Guizhou: Ceheng, Ceheng Exped. 82-0095 (PE). Hainan: Chingmai, C.I. Lei 61 (BO, NTUF, PE). Dingan, Y.G. Liou 1613 (PE). Manning, P.C. How 73028 (PE, SING, TAI); F.W. Xing et al. 6029 (TNM). Ren Hsien, E. Hainan Exped. 279 (CDBI, PE). Hongkong: Emheip 2770 (BO); Hance 1523; Lamont 674 (BM); M.Y. Chun 6850; S.Y. Hu 7296 (PE); Seemann 2448 (BM). Hunan: Jianghua, H.G. Ye et al. 2780 (TNM); B.G. Li 5183 (PE). Junshan, P.C. Tsoong 850 (PE). Mangshan, B.Z. Xiao 3459 (HAST, PE). Nanyue, Z.H. Hu 321 (PE, TAIF). Shangsi, Beijing Youth Exped 754 (PE). Yizhang, B.H. Liang 84633; S.Q. Chen 2747 (PE). Jiangxi: J.B. Yang 1468 (PE). Anyuan, Q.M. Hu 1880 (PE). Dayu, J.S. Yue 1034; G. Yao 1223; Z.R. Yu 1539 (PE). Huichang, Q.M. Hu 3005 (PE). Longnan, Jiangxi Exped 436; No. 236 Exped 1011 (PE). Ningdu, J.S. Yue 1940; Q.M. Hu 5528 (PE). Quannan, J. Syong 513 (PE). Ruijin, Q.M. Hu 3803 (PE). Shangyou, Jiangxi Exped 27 (PE). Shicheng, Q.M. Hu 4474 (PE). Suichuan, No. 236 Exped 222; X.S. Hu 618 (PE). Xingguo, Sci. Institute ofW. China 933 (PE). Xunyang, Q.M. Hu 1202 (PE).
VIETNAM. Tonkin, B. Balansa 4425, 4679 (K).
D. glaucescens var. blumeanum (Baill. ex Mull. -Arg.) Smith (fresh materials: M.S. Tang T1095, T1097 (SYSU)
*pictures from trees cultivated in Cibodas Botanical Garden).
INDONESIA. Bali, Sarip 395 (BO, L). Bali Timur, McDonald & Ismail 4775 (KEP, L). Celebes, Teysmann 14064 (BO). Java, Blume s.n. (k); Brinkman 748; Bruggeman 1251, 3740 (BO); C.A. Backer 22544 (BO, L); C.G.G 502; Djamhari 243 (BO); Kuhl & van Hasselt s.n. (L); H. Nutra 1 (K). L.R. Lanjouw 75 (BO, L); R.F. Doclitrsch Ja3606; Reenis 4133; S.H. Koorders 2123, 9970B, 2166B (BO), 37928 (BO, K), 42061 (BO); van Balgooy 2930 (L); van Leeuwen-Reijnvaan 12479; van Probolinggo Ja3000 (BO, SING); van Steenis 5013, 5027; W.S. Hoover et al. 32783, ARS145 (BO); Zollinger 2140 (BM). Lombok, J. Ebert 2140 (BO, L, SING), 2302 (BO, K, L, PNH, SING). Moluccas, A. Kostermans 1229 (BO, PNH, SING).
D. glaucescens Blume var. glaucescens
INDONESIA. Java, Blume 235 (BM, L); R. Preanger 13833B, 13947B; S.H. Koorders 2132 (BO), 3466 (L, MEL), 33319 (BO, L, K), 27998B (BO); Zollinger 5 (L), 1087 (MEL). Babaoria, S.H. Koorders 24359 (BO, L, K), 24365B (BO, K). Banjumas, S.H. Koorders 39031B (BO, L). Brichensorg, Lam 3782 (BO, L, SING). C. Kerala, A. Kostermans
TANG et al. ― Pistillate flowers of Daphniphyllum
519
s.n. (K, KEP). G. Salak, Blume 1908 (L); van Balgooy 5243 (L); W.S. Hoover & D. Girmansyah 31921 (BO). Mt. Ha-limun, W.S. Hoover et al. 32812 (BO). Nirmala, C.A. Backer 11191 (BO). Preanger, C.A. Backer 22544 (BO, K); S.H. Koorders 26557, 26523B (BO, L). Scerakarta, R. Bringmna 748 (SING). W. Java, van Balgooy 2930 (K), 5195 (BO, L). Cult. in Hort. Bog., S.H. Koorders 39027B (BO); Anonymous IX82 (BO, L, SING), IX87 (BO), IXC.107 (BO, L). With­out precise locality, Kalshoren XI (BO).
D. gracile Gage
INDONESIA. Irian Java, J.M. Mangen 1839 (BO). Papua, R.J. Johns 9213 (BO, L).
PAPUA NEW GUINEA. Central, C.E. Ridsdale NGF36814 (BO); L.A. Craven 2992 (K, L); P. van Royen NGF30051 (BO), NGF30165 (SING); Vink & Schram BW8828 (BO). Chimbu, J. Vandenberg NGF39512 (PNH, L), NGF39653 (BO); J.C. Regalade & Katik 1148 (K). Eastern Highland, A.N. Millar & C.D. Sayers NGF23715 (SING); J.S. Womera-ley NGF37369 (BO, SING); R.D. Hoogland & R. Pullen 5521 (BM, L, MEL), 5730 (BO, BM, L, MEL). Hubrecht, G.M. Versteeg 2439 (L). Idenburg River, L.J. Brass 12188 (BO). Lake Habbema, L.J. Brass 9525 (BO, K). Morbe, P. van Royen 11086 (K, L); R.D. Hoogland 9522 (BM, BO, L); T.G. Hartley 12785 (SING). Mt. Amungwiwa, J.S. Womeraley NGF17942 (BO). Nettoti, P. van Royen & H. Sleumer 7867 (L). Southern Highlands, A. Vinas 116; C. Kalkman 4994 (L), 5129 (L, SING); J. Wiakabu & J. Veldkamp LAE73727 (L); P.F. Stevens & D.B. Foreman LAE55789 (K); R. Johns et al. 1578 (L); R. Schodde 1712 (L, PNH), 1900 (L); J.S. Womeraley LAE55296 (L, SING); W. Vink 17060 (SING). West Sepik, J.R. Croft & Y. Lelean LAE65826 (K, L, SING); J.E. Veldkamp 6388, 6476 (L). Western Highlands, A. Pulle 5030 (L); J.S. Womeraley NGF11236 (BM, SING); R. Schodde 2057 (L); R.D. Hoogland. & R. Pullen 6112 (BM, L, MEL); W. Vink 16042 (L), 16495 (K). Without precise locality, A. Pulle 1029 (L); C.E. Carr 13857 (BM), 15094 (BM, SING), 15235 (BM); F.A.W. Schram BW13415 (L); J.R. Croft LAE61811 (BM, PNH); R. Pullen 2439 (K); R.D. Hoogland & R. Schodde 6778 (PNH).
D. laurinum (Benth.) Baill.
INDONESIA. Bangka, H.A.B. Biinnemeijer 1481 (BO); A. Kostermans 1059 (BO, L), 1079 (BO); A. Kostermans & Anta 661 (BO, K, PNH, SING), 1059 (BO, L); T.D. Kobus s.n. (BO); Tejsmann 3316A, 3453 (BO); Meidang 81 (BO). Central East Kornea, F.H. Endert 1576 (BO). Kalimantan: Central Kalimantan, J.P. Mogea & W.J.J.O. 3784 (BO, K, KEP); Mogea 3784 (BO). East Kalimantan, Ambri & Arifin AA129 (K, L), AA130 (L). South Kalimantan, Kessler et al. PK1721 (K). West Kalimantan, A.C. Church et al. 2488 (K, L); M. Shea 23485, 28000 (BO). Kembayang, M. Shea 26286 (BO). Pulau Sebuku, Ambriansyah 2456 (L). Java, Bakh & B. Yun 3010; C.A. Backer 26011 (BO), Teysmann s.n. (K). Sumatra, A.H.G. Aliston 13835; D. Arbain 543, 558; H.F. Sun 9857; H.S. Gusdarff 38 (BO); M. Jacobs 4621 (BO, K, PNH, SING); R.S. Bocea 6891, 7381 (SING); Teysmann 1006 (BO); W.J.J.O de Wilde & B.E.E. Wilde-Duyfjies 20732 (L). Cult. in Hort. Bog., III-G 41 (BO); IX. C. 93 (BO, IBSC); IXC102 (BO, SING). Without precise locality, H.A.B. Biinnemeijer 1465 (BO).
MALAYSIA. Malay Peninsula: Kelantan, E. Soepadmo & Mahmud 1193 (BO, K); M. Shah MS2540 (KEP, SING).
Kepong, C.F. Symington 45913; J.H. Smith 63710 (KEP); K.M. Kochummen FRI2273, 26247 (KEP, L); T.C. Whitmore FRI20703 (K). Kuantan, K.M. Kochunmen KEP94910 (KEP). Malacca, Griffith 4915 (K); Ncoutter s.n. (MEL); van. M. Abris 28, 1769. (SING). Negri Sembilan, J.N. Ruekill 2523 (BO, SING); van. M. Abris 1252 (SING). Pahang, Burkill & Haniff 16804 (K); D. Hou 758 (BO, K, KEP, SING); Hauiff 16804 (SING); Kiah 23935 (BO, SING); L. Chua et al. FRI40652 (KEP); M.R. Henderson 21653 (BO, K, SING); R. Ismail 97948, 98934 (KEP, SING); Ruekill 16804 (BO); S.L. Guan FRI34070 (KEP); T.C. Whitmore FRI3789 (KEP); Thiah 23935 (BO). Perak, F. Sentiasium s.n. (K); H.N. Ridley 555 (SING); King 4648 (BO), 5707 (L, SIGN), 10116 (SING); S.B. Kiah SK497 (KEP, SING); Scortechini 226.; T.C. Whitmore FRI0883 (SING). Selangor, H. Asnah et al. FRI135588; T.C. Whitmore FRI15965 (KEP). Ulu Gombak, C.F. Symington KEP48608 (KEP). Borneo: Sahah, D.D. Wood 1767 (BO); G. Majawat 125769 (K); Geofarry & Suga-rah 7780 (SNP); H.C. Tang 1643 (SING); H. Hallier 691 (BO); J. & M.S. Clemens 30674 (BM, BO, L); J. Nais et al. 4957 (SNP); K. Ogata 11496 (L); Korthals s.n. (L); O. Posthumus 2140 (BO); R. Rimi et al. 7936 (SNP); Valera 4831 (L, SING). Sarawak, I. Paie S.28080 (SING), S.46069 (L, KEP); L.C.J. Julaihi et al. S.83560 (KEP).
SINGAPORE. I. H. Burtill 524 (SING); H.N. Ridley 1816 (MEL, SING), 3436A (SING), 3437 (MEL); R.W. Hullelt 169 s.n. (SING); Wallich 8020 (BM, K).
D. longeracemosum Rosenth.
CHINA. Guangxi: Damiaoshan, S.H. Chen 15088, 15733 (PE). Huanjiang, H.N. Qin et al. 894300 (HAST, PE). Guiz­hou: Anlong, S.G. Wu et al. 101046 (KUN). Tibet: Gangde, H. Sun et al. 3701 (KUN). Gelin, H. Sun et al. 2700 (KUN). Motuo, Qinghai-Xizang Exped. 74-1811, 74-5011 (KUN, PE). Taipingshan, H. Sun et al. 2728 (KUN). Yunnan: Der-gong, H. Sun et al. 2451, 2495 (KUN). Eryuan, Sino-Ross. Exped 3353 (PYU). Gongshan, Dulongjiang Exped. 3930; Qinghai-Tibet Exped. 9487; P.I. Mao 487 (KUN). Jinping, S.K. Wu et al. 3297, 4495 (KUN); Sino-Ross. Exped 3040 (KUN, PE). Luchun, S.K. Wu et al. 249, 604 (KUN). Maguan, Q.A. Vvu 8614 (KUN). Marlipo, K.M. Feng 13121, 22680; Z.W. Lin 499 (KUN). Mengzi, T.N. Liou 18728 (PE). Pingpien, H.T. Tsai 60821 (SZ); K.M. Feng 4819 (KUN, PE, SZ); P.I. Mao 2436 (KUN), 4293 (SZ, PE), 8936; S.H. Chen 548 (KUN); Sino-Ross. Exped 2453 (PE). Sichou, C.W.
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Wang 85861 (KUN, PE); K.M. Feng 12076 (KUN, PYU); Q.A. Vvu 9983; S.J. Wang 387; Wenshan Exped. 101785; Z.H. Yang 90-1431; Z.W. Lin 667 (KUN). Wenshan, K.M. Feng 11199 (KUN). Yuanyang, Luchun Exped. 1736 (KUN).
VIENAM. Tonkin, Chapa, M. Poilane 12590 (K). Hacoi, W.T. Tsang 29517 (BO, K, L, SING). Without precise loca­lity, Sino-Vietnam Exped. 109 (KUN).
D. luzonense Elmer
PHILIPPINES. Bayninan, H.C. Conklin & Buwaya I978 (K, PNH). Biliran Is., M.D. Sulit 21554 (K, PNH, SING). Bukidnon, M.D. Sulit 3489 (BM, PNH). Ilocos Morte, M. Ramos 32838 (L, SING). Luzon, A.D.E. Elmer 6290 (K, PNH, TI), 8615 (K, L); D.R. Mendoza 97535 (L, PNH); H. Conklin 78 (K, PNH), 42703 (L); M. Ramos 33273 (SING); M. Ramos & G. Edano 45660 (BM, K). Mindanao, A.D.E. Elmer & Merrill 5658 (K); Caerlan et al. 10953, 19873 (l); M. Ramos 14623 (BM, K); M. Ramos & G. Edano 38507 (K), 38516 (BM, L), 38743 (BM); C.E. Ridsdale 963; C.E. Ridsdale et al. 5535 (K). Palawan, E.D. Merrill 9494 (BM, K, SING). Poitan, H.C. Conklin & Buwaya I868 (PNH). Sibuyan Is., G. Raynoso et al. 21469 (L).
D. macropodum Miq. var. humile (Maxim.) Rosenth. (fresh materials: M.S. Tang T1200, T1201, T1341, T1342 (SYSU) from trees cultivated in Tsukuba Botanical Garden).
JAPAN. Hokkaido, H. Hoshi 1500 (TAI); T. Tanaka 15359 (NTUF). Hakodate, Maximowicz s.n. (MEL); U. Fau-rie 6804 (BM); J. Ohwi 4793 (SING). Hidaka-shico, H. Yamaji 4474 (HAST). Horoizumi, H. Izumi 704171 (TNS). Ishikari, H. Iuramoto 59698; M. Kusahu 38-458 (TNS); Y. Kudo 1229 (TAI); S. Saito 1126 (TNM). Sapporo, K. Ku-wabara 255716; N. Hiratsuka s.n. (TNS); S. Saito 280 (TI). Sha-ri-cho, T. Kohyyama et al. 196 (TI). Shiribeshi, Y. Kudo 2625 (TAI). Teshio, Y. Kudo 4532 (TAI). Honshu: Aomori, K. Yonekura 11546 (HAST); M. Mizushima 2542 (TI). Fukushima, S. Kunio 108802 (TNS). Gifu, H. Takahashi 5953, 20150; H. Takahashi & H. Takano 1639, 7888 (TNS). Gunma, H. Ohba et al. 73113 (TI); S. Kurata & T. Nakaike 10033; S. Okuyama 18398 (TNS). Hyogo, G. Murata 20401 (TNS). Ishikawa, H. Sakurai 7732; K. Deguchi & S. Tsugaru 18912; S. Tsugaru et al. 15993 (TNS). Iwate, S. Okuyama 1103, 12700; T. Okawa 426874 (TNS). Miyagi, K. Sugawara et al. s.n. (TI); T. Kajita & H. Iketani 295 (PPI). Niigata, F. Konta 2902; K. Hisauti 93900; M. Togasi 1352; S. Kitamura & G. Murata 2841; S. Okuyama 52956 (TNS). Shiga, G. Murata 20801 (TNS). Toyama, S. Oura 889 (TI). Yamagata, A. Matsui et al. 3027 (HAST); H. Okuyama 36335; S. Tsu- garu & T. Takahashi 13561 (TNS); Y. Yuhki 123938; U. Nishiokitama 14694 (TNS). Without precise locality, Y. Ikoma 2167 (TNS).
D. macropodum Miq. var. macropodum (fresh materials: C.K. Liao s.n.; M.S. Tang T778, T793, T897, T898, T899, T1141, T1259, T1260, T1314, T1315; C.K. Yang T1198 (SYSU) from Taiwan, M.S. Tang T1202, T1204, T1340 (SYSU) from Japan).
CHINA. Anhwei: Chengkou, T.L. Tai 10633, 107057 (PE). Huang Mount, E. China Bot. Station 6211; Fordham Uni. Biological Sci. Team 157 (KUN); K.J. Guan 75096; M. Chen 1204; M.J. Wang 3680; P.C. Tsoong 3441 (PE). Hwan-gshan, T.N. Liou & P.C. Tsoong 2045 (PE). Jinzhai, K. Yao 8975 (MEL). She Hsien, K.J. Guan 75096; M.B. Deng 4667 (PE). Songsi, W.C. Cheng 3976 (PE). Fujian: Chongan, Wuyi. Exped 594, 2292 (PE). Longxishan, Longxishan Exped. 1546 (HAST). Wuyi Mount, M.J. Wang 3283 (PE). Yongan, M.J. Wang 3445 (PE). Guangdong: Lechan, L.S. Chen 42905 (PE); Z. Huang 31405 (PE, SZ). Lianshan, Nanling Exped. 536 (TNM). Ruyuan, S.C. Liou 28947; X.M Gao 53582 (PE, SZ); Y. Li 2081; Z. Huang 44302 (PE). Yangmingshan, S.H. Chen 694 (PE). Yangshan, S.G. Li 201162 (PE). Renhua, L. Deng 7469 (SZ). Guangxi: Cangwu, S.H. Chen 10103 (CDBI). Damiaoshan, Q.H. Lu 2588 (PE). Long-sheng Gezu Zizhixian, S.C. Jhong 51054; Y.C. Chen 600486 (PE). Luocheng, H.N. Qin et al. 894909 (HAST). Xingan, S.C. Jhong 51208; S.L. Yu 900048 (PE). Zhongshan, J.T. Li 604044 (PE). Ziyuan, S.C. Jhong 51863 (PE). Guizhou: Fanjingshan, C.Y. Chiao 525; N. Guizhou Exped. 491; Y. Tsiang 7632 (PE). Jiangkou, B. Bartholomew 28724 (PE). Kai-li, S. Guizhou Exped. 1053, 2165 (PE), 3442 (PE, SZ). Leigongshan, Anonymous 85-0500 (PE). Leishan, S. Guizhou Exped. 843 (PE). Rongjiang, S. Guizhou Exped. 3227 (PE). Songtao Miaozu Zizhixian, B. Bartholomew 20565-9 (PE). Yinjiang Tujia and Miao Autonomous Zizhixian, S. Guizhou Exped. 491, 623 (PE). Hubei: Badong, C. Y. Chen 1985 (PE). Enshi, Anonymous 1522 (PE). Guidong, Anonymous C-151 (PE). Hefeng, H.J. Li 6320 (PE). Jianshi, L.Y. Dai 41 (PE). Lichuan, G.X. Fu & S.Z. Jhang 1696 (PE, SZ); S. Zhang 1693; W.B. Lin 272 (PE). She皿ongjia, Sino-Amer. Ex­ped. 1468 (PE). Xianfeng, H.J. Li 5743, 8474, 9069, 9323 (PE). Xingshan, Y. Liou 402 (PE). Xuanen, H.J. Li 5425 (PE). Hunan: Chanziping, Hunan Exped. 253 (PE). Cili, Hunan Exped. 429, 690 (PE). Dayong, Hunan Exped. 1261 (PE). Dongan, Y. Liu 929 (PE). Heng Mount, Anonymous 5886 (PE). Ling Hsien, L. Jiang 199 (PE). Mangshan, B.Z. Xiao 3640 (PE), 3766 (HAST); M.S. Huang 111931 (CDBI). Nanyue, K.J. Guan K169; M.H. Li 862 (PE). Sinning, C.S. Fan & Y.Y. Li 616 (BO). Wugang, L.H. Liou 16273 (PE). Xinning, L.B. Luo 1109; S.C. Luo 621 (PE). Yizhang, S.H. Chen 2692 (PE, SZ). Yongshun, Hunan Exped. 118 (PE). Yuanling, G.K. Chang 589 (TNM). Jiangsu: Yifeng, H.N. Zou 1165 (TNM). Jiangxi: Jingan, C.M. Tan 971297; C.S. Ye 2367 (TNM). Jinggangshan, J. Syong 2350; J.S. Yue 5236 (PE). Lichuan, W.Y. Yang 650057 (PE); S.S. Lai & H.R. Shan 1230 (TNM). Mingyueshan, J.S. Yue 3314 (PE). Shangrao, Anonymous 4868 (PE). Shangyou, Jiangxi Exped 503 (PE). Suichuan, J.S. Yue 4355 (PE). Wugongshan, J.S. Yue 2993; Jiangxi Exped 402, 1249 (PE). Wuning, C.M. Tan 941238 (PE, TNM); C.S. Ye 1445 (TNM); J.H. Zhang 1052 (TAIF);
TANG et al. ― Pistillate flowers of Daphniphyllum
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L.H. Liou 15037; W.S. Lai 7781 (PE); Tangshan bot. garden 68 (PE); Y. Zou 63319 (TNM). Xunyang, Anonymous 1226 (PE). Yanshan, C. Chen 42; J.S. Yue 401305; M.J. Wang 2319; M.S. Nie 4349 (PE). Szechuan: Chengkou, G.H. Yang 57211 (PE); T.L. Tai 101920 (CDBI, PE, SZ), 107057 (CDBI, PE). Dazhu, Z.Z. Chang 232 (CDBI). Fengjie, Szechuan Uni. Exped. 107927, 108018; W.P. Fang 24267 (SZ); X.M. Zhang 108018 (PE). Kuan Hsien, Chuan 84 (PE); D.E. Boufford & B. Bartholomew 24235 (HAST, L); D.P. Ho 47077; L.C. Hu 5753; S.S. Chien 5626 (SZ); W.P. Fang 2030 (PE). Nanchuang, H.F. Chang 20888; J.H. Syong 90845 (PE); J.H. Syong & G.F. Li 90845 (SZ); N.L. Chii 927; T.H. Tu 91285 (PE). Omei Mount, B. Luo et al. 1909 (SZ); C.H. Li 96-158 (PE); C.J. Cho 6161; C.L. Chaw 6387 (SZ); D.H. Zhu et al. 626 (TAIF); F.T. Wang 23149 (PE); G.F. Li 52701 (SZ); G.H. Yang 54140 (SZ), 56980 (PE); H.L. Tsiang & S.S. Chang 30131; H.L. Tsiang et al. 30866 (SZ); J.H. Syong 784, 32711 (PE); J.H. Syong & G.F. Li 93976; J.H. Syong et. al. 30130 (SZ), 33207; J.W. Yao 3199 (PE, SZ), 4871 (PE); R.K. Jhao 604 (PE); S.J. Yu 50025; S.L. Sun 133 (SZ); S.N. Syi 675; S.S. Chang 1953, 33607 (PE); S.Y. Hu 311; Sino-Ross. Exped 1895; Szechuan Exped. 2648; T.C. Lee 4555; T.H. Tu 559 (SZ), 32928 (PE); T.L. Dai 421 (CDBI, PE); T.N. Liou 1315; T.T. Yu 423 (PE), 2663 (SZ); W.P. Fang 2084 (SZ), 30866 (PE); Y.H. Tao 52092 (SZ). Qianjiang, Anonymous 4170 (PE). Qingchengshan, J.L. Wu 33810; T.P. Wang 10002; W.P. Fang 19362 (PE). Shixhu, J.A. Wang 173 (CDBI); K. Chen 840 (CDBI, SZ). Tianquan, D.P. Ho 43175 (SZ). Wushan, Szechuan Uni. Exped. 108018 (SZ). Wuxi, G.H. Yang 65452 (CDBI, PE). Xianfeng, W.C. Cheng 1072 (PE). Xiyang, Anonymous 2545 (PE). Yunnan: Chenggong, N.E. Yunnan Exped. 1113 (KUN). Laojianshan, Anonymous 4819 (PE). Longling, J.Y. Wu 1283 (PE). Suijiang, B.X. Sun Exped. D394 (KUN, PYU). Wenshan, K.M. Feng 32595 (KUN). Zhejiang: Chunan, Anonymous 30279 (PE). Kaihua, Zhejiang PL. Resour. Exped. 26206 (PE). Linan, Deng et al. 4536; F.Y. He 23030 (PE). Longquan, S.Y. Jhang 322; Y.Y. Ho 1612 (PE). Niutoushan, S. Chen 797 (PE). Ruian, S.Y. Jhang 5461 (PE). Suichang, J.S. Wang 2038; Zhejiang PL. Resour. Exped. 25707 (PE). Tianmushan, F.Y. He 21885; K.K. Tso­ong 18570; L.B. Deng et al. 4105; M. Chen 706; T.N. Liou L7274; W.C. Cheng 3652 (PE) Tiantai Mount, R.C. Ching 2090; S. Chen 412, 20909 (PE). Taishun, S.Y. Jhang 3770 (PE).
JAPAN. Honshu: Gifu, H. Takahashi 5907; H. Takahashi & H. Takano 7667 (TNS). Hondo, M. Togasi 1352 (PE). Hyogo, G. Murata 22182 (TNS). Yaku Is., T. Yahara et al. 10575 (TI). Kyoto, S. Tsugaru 2715; S. Tsugaru et al. 23198; T. Takahashi & T. Katsuragawa 25986 (TNS). Mie, G. Murata 19761 (TNS). Miyagi, S. Junji s.n. (TNS). Nagano, S. Ito 61 (TNS). Nagasaki, Maximowicz 14816 (BO). Shiga, H. Kanai& T. Morita 239; N. Fukuoka et al. 1023 (TNS). Tokyo, H. Hara 3470 (TI); K. Inoue 1576 (PE, TI),. Tottori, G. Murata et al. 322 (TNS). Ueno, M. Mizushima 296 (Ti). Yamaguchik, Anonymous 47623 (TNS). Yokohama, Maximowicz s.n. (MEL). Without precise locality, M. Mizushima 1139; K. Kondo 6580; M. Togashi 10229; T. Yamazaki 9735; Kamitsuke 381 (TI).
KOREA. Anmyon Is., B.R. Yinger et al. 2359 (K). Jejudo, I.C. Chung 1204 (TNS). Without precise locality, E.H. Wil­son 8541, 9449 (K).
TAIWAN. Chiayi, C.E. Chang 1234 (PPI); C.M. Wang 5886 (HAST, TNM); W.T. Chao et al. 2206 (TNM). Hsinchu, C.H. Chen et al. 3525 (TAIF, TNM); J.C. Wang et al. 11388; K.C. Yang et al. 2770 (TNM); S.C. Liu & H.M. Chang 1506 (HAST, TAIF). Hualien, C.I. Peng 5984, 16522 (HAST); Liu et al. 78; S.S. Yin 1327; T. Tanaka 5360 (NTUF). Ilan, B. Hayata s.n. (TI); C.C. Liao 902 (HAST); C.H. Chen et al. 3525 (HAST); C.I. Peng 17436; J.C. Wang et al. 11388 (HAST); M. Tamura & T. Shimizu 20614; S. Suzuki 1070 (TAI); T.Y.A. Yang et al. 12238 (TNM); W.L. Chiou 14117 (TAIF). Kaohsiung, C.M. Wang 6658, 6659 (TNM); Y. Yamamoto YM608 (TAI). Miaoli, N. Fukuyama 152 (TAI). Nanou, C.C. Chen 4330 (HAST, PPI, TNM); C.C. Kuo 163 (NTUF); J.C. Wang et al. 8130 (HAST, TAIF); J.W. Hsu 2419; T.Y. Yang 3692 (TNM). New Taipei, C.C. Liao 372; C.I. Huang 1399 (HAST); M.F. Loa 217 (TAIF). Taitung, Y. Yamamoto 2061 (TAI).
D. neilgherrense (Wight) Rosenth.
CEYLON. Central, A. Moon 549 (BM); Davidse 7978 (L); Nooteboom 3230A (KEP, L); R.D. Hoogland 11488 (L). Garadenya, Thwaites 491 (BM, BO, K, MEL). Namana Kula, T.B. Worthiogton 6181 (BM). Sabaragamuwa, A. Kos-termans 28196 (L). Uva, S. Waas 1623 (L); T.B. Worthington 5423 (BM). Without precise locality, Frnastes s.n. (MEL).
SOUTH INDIA. Kerala, A. Kostermans 26123 (K, KEP). Tamil Nadu, Bourdillon s.n.; C. Saldanha CS5198 (K); Clarke 10752 (BM); Fischer s.n.; Frauaucme s.n. (K); Goughia 10749, 10752 (BM, K); J.S. Gamble 11970, 14468 (K), 17985 (MEL); K.M. Matthew RHT26754 (L), RHT41101 (K); M.A. Eversheds.n. (BM); Subramanian 47, 401 (L); Wight 2649 (K, MEL). Without precise locality, Beddome 7171 (BM); Bourdillon 322,; Saldanha CS4964; Perrottete
s.n. (K).
D. oldhamii (Hemsl.) Rosenth.var. oldhamii (syn. D. glaucescens Bl. subsp. oldhamii (Hemsl.) T. C. Huang var. oldha­mii) (fresh materials: C.C. Chen T1196; C.S. Liu T1134; M.S. Tang T697, T698, T713, T769, T847, T887, T888, T961, T962, T1000, T1001, T1002, T1118, T1120, T1157, T1219, T1220, T1244, T1255, T1275, T1299, T1320, T1321, T1326, T1420, T1421, T1512, T1513, T1514; C.K. Yang T601, T1109 (SYSU)).
CHINA. Chekiang: Foochow, H.N. Chung 2273 (PE, SING). Jieukaishan, R.C. Ching 2610 (PE). Jingning, S.Y. Chang 4377 (PE). Kaihua, G.R. Chen 2457; J.X. Wang 1850 (PE). Longquan, S.Y. Chang 4608; Y.Y. Ho 1612 (PE). Ningbo,X.Y. Ho 27113 (PE). Niutoushan, S. Chen 797 (PE). Qingyuan, S.Y. Chang 3390 (PE). Simingshan, B.Q. Zhong 182 (PE).
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Suichang, Zhejiang PL. Resour. Exped. 25679 (PE). Taishun, S.Y. Chang 3731 (PE). Tiantaishan, S. Chen 412; Zhejiang PL. Resour. Exped. 28416 (PE). Yandangshan, J.X. Wang 1761 (PE). Yinhsien, Y.Y. Ho 1029 (PE). Fujian: Chongan, M.J. Wang 3068 (PE). Dehua, Tsoong 1007 (PE). Jiangjun, Lunghishan Exped. 2038 (HAST). Jiangle, Anonymous 3045 (PE). Jiangning, Z.Y. Li 10730 (PE). Kushan, H.H. Chung 2273 (PE, SING). Liancheng, H.B. Chen 702 (PE). Nanjing, R.C. Lin 3787 (PYU). Shanghang, L.G. Lin 7447 (PE). Wuyi Mount, M.J. Wang 3183 (PE). Zhangdian, Q.M. Hu 3654 (PE). Zhangping, R. Lin 4527 (PE). Guangdong: Fengkai, K.C. Ting & L. Yu 6262 (CDBI). Guangzhou, P.C. Zhou 10413 (PE). Hedoukou, C. Huang 164546 (PE). Hulishan, L. Deng 4338 (PE). Jiaoling, L. Deng 4713 (PE). Jiufengs-han, Z. Huang 31246 (PE). Lechang, Nanling Exped. 3644 (TNM). Lianshan, P.C. Tam 58340 (PE). Lechan, W.T. Tsang 20795 (BO, SING). Lokchong, Y. Tsiang 2314 (SING). Qujiang, X.M Gao 50823 (PE). Raoping, N.Q. Chen 42675 (PE). Ruyuan, Y.G. Liu 318 (PE). Sinfung, Y.W. Team 496, 597 (HAST). Tsengshing, W.T. Tsang 20204 (PE). Wenkuangshan, C. Huang 163719 (PE). Xinyi, Z. Huang 31783 (PE). Zhaoquing, Y.G. Lin 2541 (PE). Guangxi: Damiaoshan, Q.H. Lu 2422; S.H. Chen 15980 (PE). Hsinan, Z.Z. Chen 51365 (PE). Huanjiang, H.N. Qin et al. 894011 (HAST, PE). Lingui, C.F. Liang 31609 (PE). Lungsen, Guangfu. For. Exp. 75, 403, 438 (PE). Pingna, Z. Huang 39128 (PE). Rongshui, Z.Y. Li et al. 891917 (HAST, PE). Tseunghsien, Z. Chuang 39368 (PE). Guizhou: Leigongshan, K.M. Lan 85-0239; S. Guiz­hou Exped. 4004 (PE). Tiantaishan, B.Q. Zhong 231 (PE). Xishui, Bijie 1477 (PE). Zunyi, S. Guizhou Exped 1031 (PE). Hainan: Niulanjiao, S.R. Leung 61276 (PE). Hongkong, N.K. Chun 40267 (SING); S.K. Wah 10915; S.Y. Hu 8446; W.Y. Chun 5202 (IBSC); Wright 421 (K). Hubei: Lichuan, L.Y. Dai & C. H. Qian Hubei-889 (PE). Hunan: Chengbu, B. Q. Zhong 1584 (PE). Cili, W. Hunan Exped. 1894 (PE). Hsuehfeng, Z.T. Li 2103 (PE). Longhui, L.D. Duan 5740 (PE). Mongshan, M.S. Huang 112616 (CDBI). Qiyang, S.H. Chen 825 (PE). Shaoyang, F.J. Huang 146 (PE). Tiexiang, Anjiang Agric. School 1135 (PE). Wuyi Mount, M.J. Wang 3183 (PE). Xining, C.C. Luo 1283; L.B. Luo 858 (TNM); Z.C. Luo 304 (PE). Yangmingshan, S.H. Chen 489 (PE). Yizhang, B.H. Liang 85157; L.H. Liou 10250 (PE). Jiangxi: Jingan, C.M. Tan 9611200 (PE, TNM); C.S. Ye 2141 (TNM). Shangyou, Jiangxi Exped 94, 179 (PE). Suichuan, J.S. Yue 4507 (PE). Wuyuan, Q.H. Li & C. Chen 322 (PE). Xiushui, C.M. Tan 951067 (HAST, TNM). Yongxin, W.S. Lai 1110 (PE). Zixi, M.L. Nie 3434 (PE). Szechuan: Changning, Anonymous 15219 (CDBI). Kuen Hsien, F.T. Wang 20678 (PE). Lichuan, W.C. Cheng & C.T. Hwa 1948; C.T. Hwa 155 (PE). Nachuan, G.F. Li 60895; J.H. Syong & Z.Z Lin 91827; G. F. Li 63295 (PE). Omei Mount, C.Y. Chiao & C.S. Fan 442 (HAST); G.H. Yang 54797; J.H. Syong et. al. 32982; S.S. Chang 32504; W.P. Fang 15118, 31620 (PE). Pa Hsien, T.T. Yu 2893 (PE). Pingshan, S.L. Sun 5413 (PE). Xiushan, Ano­nymous 4103 (PE). Yunnan: Fooning, C.W. Wang & Y. Liu 88699 (PE). Libo, Z.M. An 2-184 (PE). Zhejiang: Huang-keng, S.Y. Jhang 4011 (PE). Jiangshan, P.S. Siang 2170; S.Y. Jhang 4884 (PE). Linan, F.Y. He 22894 (PE). Linglongyan, S.Y. Jhang 9283 (PE). Lishui, S.Y. Jhang 6059, 6291, 6471 (PE). Longquan, S.Y. Jhang 2826, 7340 (PE). Qingyuan, S.Y. Jhang 3390 (PE). Taishun, S.Y. Jhang 3731, 5596 (PE). Tiantaishan, F.Y. He 28219 (PE).
TAIWAN. Hsinchu, C.I. Peng 14846 (HAST); K. Odashima 17827 (MEL); T. Tanaka & Y. Shimada 17827 (K, MEL, PNH, SING). Keelung, C.I. Peng 12898 (HAST); H.L. Chiang 154 (HAST, TAIF). Miaoli, H.Y. Chen & S.C. Wu 18 (TAIF, TNM). New Taipei, B. Bartholomew 7841 (HAST); C.C. Liao 1250; C.I. Peng 7498; C.L. Huang 75 (HAST); H. L. Chiang 154 (HAST, TAIF, TNM); R. Oldham 497 (BM, K); S.C. Hsiao 1305 (HAST); S.C. Wu et al. 974 (HAST, TNM); T.Y. Liu et al. 208; T.Y. Yang et al. 1657 (HAST, TNM); Y. Tateishi & T. Nemoto 20478; Y.C. Lu 378 (HAST). Taichung, B. Bartholomew 7760; C.I. Peng 9963 (HAST); C.M. Wang 3775 (TNM); J.C. Huang 1094 (TAIF); K.Y. Wang 274 (HAST, TNM); Y. Tateishi et al. 21301 (HAST); Y.P. Cheng 4352 (HAST, TAIF). Taipei, B.L. Shih 1277 (TAIF); C.C. Wang 934; C.C. Liao 393 (HAST); C.I. Huang 199 (TNM); C.I. Peng 6421, 12972 (HAST); C.M. Wang 1847 (HAST, TNM); C.M. Wang & Y.H. Tsai 3021 (HAST, TAIF, TNM); C.Y. Kuo 8 (HAST, TAIF); H.L. Chiang 330 (TNM); J.C. Liao 10818; Laio et al. K61 (NTUF); R.S. Weng et al. 233 (TNM); S. Sasaki s.n. (TAIF); S.C. Liu et al. 549 (HAST, TAIF, TNM); S.H. Su 813; S.W. Chung 3873; S.Y. Lu 25034 (HAST); T.C. Huang 1553 (NTUF); T.S. Liu & C.C. Kuo 580 (NTUF); T.Y. Liu 1183 (HAST); T.Y. Yang et al. 1657 (HAST, TNM), 12253 (TNM); W.C. Leong 1149 (HAST); W.F. Ho 1179; W.P. Leu 1423 (HAST); W.P. Leu et al. 1839 (HAST, TNM); W.S. Tang 1397 (TAI); Y.C. Chan et al. 8 (HAST, TNM); Y.H. Chang 3931 (HAST, TAIF); Y.J. Lai 198, 241 (HAST). Taoyuan: C.M. Wang & H.M. Lin 2190 (HAST, TAIF, TNM); T. Tanaka & Y. Shimada 13531 (L, SING), 13582 (L); T. Tanaka et al. 13532 (MEL, L, SING); Y. Shimada s.n. (NTUF).
D. glaucescens Bl. subsp. oldhamii (Hemsl.) T.C. Huang var. lanyuense T.C. Huang (fresh materials: L.C. Shih s.n.; M.S. Tang T800, T801, T803, T805, T806, T810 (SYSU)).
TAIWAN. Lanyu, C.C. Liao 1205 (HAST); C.E. Chang 3074 (PPI), 16822 (HAST, PPI), 19668 (PPI); C.H. Tsou 509;
C. I. Peng 10704 (HAST); C.K. Yang 295; C.M. Wang 3562 (TNM); C.Y. Kuo 437 (TAIF); G.S. Wang & C.N. Koh 169 (TNM); J.C. Wang 5579 (HAST); M.C. Ho & H.C. Huang 114 (NTUF); M.F. Lao 215 (TAIF); Mamoru 9910 (TAI); P.F. Lu 1063, 6269 (HAST, TAIF); Q.L. Ye 5784 (PPI); S.F. Huang 2670 (TAI); S.H. Liu et al. 901 (TAIF); S.T. Chiu & J.N. Chen 4158 (HAST, TAIF, TNM); S.W. Chung 6545 (HAST, TAIF); S.Y. Lu 17604 (TAIF); T.C. Huang 5132 (TAI); T.Y. Yang & C.H. Chu 8004; T.Y. Yang et al. 3876 (TNM); W.L. Chiou 10119 (HAST, TAIF); W.L. Wagner 6709 (HAST); W.P. Leu 1395 (HAST); Y.C. Lu 1596 (HAST, TAIF, TNM); YC. Lu & C.C. Liao 1596 (TAIF, TNM); Y.C. Lu et al. 920 (TAIF); Y.P. Yang s.n. (HAST, TAIF).
TANG et al. ― Pistillate flowers of Daphniphyllum
523
D. paxianum Rosenth.
CHINA. Guangxi: Baise, Baise Exped.1899 (PE). Jinxiuyao Autonomous Zizhixian, Anonymous 424 (PE). Lingyun, A.N. Steward & H.C. Cheo 181 (BO, SING). Longjin, S.H. Chen 11763 (PE). Nanning, R.C. Cing 8208 (PE). Rongshui, S.H. Chun 14198 (PE); Z.Y. Li et al. 897314 (HAST, PE). Yaoshan, NE Exped. 175 (PE). Guizhou: Jiangkou, B. Bar­tholomew 291 (PE). Libo, Lipo Exped. 81-4-0053 (PE); X.G. Song 936 (K). Yongcong, P.C. Jhong 1105 (PE). Hainan: Baoting, Diaoluoshan Exped. 3246 (PE). Chimfungshan, S.K. Lau 5264 (HAST). Dingan, Z. Chuang 35708 (PE). Dongfang, H.Y. Liang 65264 (PE). Lingshui, Z. Chuang 36591 (PE). Pehsar, S.K. Lau 25449 (TAI). Poting, F.C. How 73071 (BM). Qionghai, Hainan Exped. 322 (PE). Wanning, F.W. Sing et al. 6238 (TNM). Yaichow, H.Y. Liang 63112 (KPE, PE). Szechuan: Omei Mount, G.H. Yang 54823; J.W. Yao 5179; R.K. Jhao 1011; S.S. Chang 138, 33615; T.T. Yu 506; W.P. Fang 3352 (PE), 3874 (K, PE); X. Jiang & S.S. Chang 30029; Y.L. Qiao 598 (PE). Yunnan: Amherst, A. Henry 11903 (K). Fooning, C.W. Wang 89062 (KUN, PE), 89619 (KUN, PE). Lungling, H.T. Tsai 55595 (BO, KUN), 55680 (BO, PE). Mianing, T.T. Yu 18175 (PE). Shweli, G. Forrest 24089 (BM, K). Sichou, Feng, K.M. 12289 (PE). Szemes, A. Henry 12064, 12657; G. Forrest 8494 (K). Tengchong, D.Y. Xia BG010 (KUN); W.Q. Yin 60-1001 (KUN). Tengyueh, G. Forrest 7697 (BM, K). Tunchong, J. Chen 1048 (PE).
VIETNAM. Tonkin: Baie, B. Balansa 557 (K). Damha, W.T. Tsang 30270 (L, SING), 30381 (BO, K, L, SING).
D. scortechinii Hook. f..
MALAYSIA. Malay Peninsula: Cameron Highlands, T.C. Whitmore FRI15473 (K, KEP, L). Pahang, C.F. Symington & Kiah 28850 (BO, K, KEP); J. Wyatt-Smith KEP78820 (K, KEP, L, SING); L. Chua et al. 39042 (K, KEP), FRI40519; P. Wilkie et al. FRI52910 (KEP); S. & A. Mohel 2970 (KEP). Perak, B. Perumal et al. FRI41697; C.F. Symington 32121 (KEP); King 7326 (BM), 8054 (SING, K); L. Wray 324 (SING), 3847 (BM), 3926 (SING); L.S.L. Chua FRI40421 (KEP); R.H. Yapp 523 (K); Scortechini 830 (K), 1347 (K). Selangor, Robinson s.n. (BM, SING); S. Carlquist 4404 (K); van Balgooy 7149 (KEP, L).
D. teijsmannii Zoll. ex Teijsm. & Binn. (fresh material: M.S. Tang T1338 (SYSU))
INDONESIA. Cult. in Hort. Bogor: Zollinger 1029 (TI); Anonymous s.n. (L).
JAPAN. Honshu: Chiba, T. Nakaike 342840 (TNS). Fukui, G. Murata 154982; T. Takahashi et al. 84335; Z. Tashiro 42339 (TNS). Gifu, H. Takahashi 714801 (TNS). Hiroshima, K. Oka 40529 (TNS). Hyogo, G. Murata 154991 (TNS). Kyoto, K. Katsu 37153 (TNS); S. Tsugaru & T. Sawada 28450 (HAST); S. Tsugaru et al. 82379 (TNS). Mie, H. Saku-rai 7729 (TNS); S. Tsugaru et al. 33770 (HAST). Shiga, G. Murata 238442; S. Seko 225707 (TNS). Shimane, S. Takaki 45376 (TNS). Shimoagata, H. Ohashi & K. Sohma s.n. (TAI). Shizuoka, E. Aihara 31513; F. Chikada et al. 32300; F. Konda 8393, 18148 (TNS); F. Yamazaki 2543; F.G. Meyer & H. Ohba 31444 (TI); H. Funakoshi 336 (TI); I. Yamashita 20497; K. Hisauchi 59432 (TNS); K. Inoue 79354 (TAIF); Rinken 51042 (TNS); S. Saito 4842 (PE); T. Okawa 388854 (TNS). Tokyo, H. Kanai 10806 (L); K. Inoue 1808 (L, PE); N. Kasil 10806 (SING). Wakayama, H. Murakami et al. 287 (HAST), 77309; K. Nishiyama 2374; N. Kurosaki & S. Miyake 692569 (TNS). Yamaguchi, A. Minami 440029 (TNS); J. Nikai 47622; K. Oka 262769; N. Fukuoka 336857 (TNS); S. Saito 638 (TNM, TI). Kyushu: Kagoshima, C. Whight 275 (L); F. Konda 9088; H. Migo 373885; H. Takahashi 4193; K. Deguchi 31592, 31878 (TNS); K. Inoue 77079 (TAIF); S. Saito 3440 (TI, TNM); T. Kamiya 28585 (TNS); T. Yahara et al. 6143 (TI); Y. Shimada 131157 (TNS). Kanagawa, Z. Tashiro 30407 (TNS). Nagasaki, N. Fujita & S. Nitsuta 128 (PE); R. Oldham 740 (K, L), 951 (K). Oita, K. Maetaro 180236 (TNS). Yakushima, M. Furuse 215695 (TNS). Shikoku: Ehime, T. Nakai s.n. (TI). Tokushima, J. Nikai 260177 (TNS). Okinawa, E.H. Walker 162107 (TNS); E.H. Walker et al. 5920 (L); E.H. Wilson 275 (K), 8056 (SING); F.R. Fosberg 37178, 38010 (L, TI); G. Murata & H. Tabata 323712; H. Kanai et al. 318301; H. Koyama et al. 544; H. Mi-gou 26997 (TNS); H. Noguchi 3501 (L); H. Ohba et al. 384 (PPI); H. Ohba & S. Akiyama 8903064 (TI); H. Okada & K. Ueda 2733 (PPI, TI); Hatusima & Sako 30967; I. Sasaki 2523 (TI); J.C. Wang 11796 (HAST); J.C. Wang & C.H. Chen 9818 (HAST); J.C. Wang et al. 11796 (HAST, TAIF); K. Iwatsuki et al. 262 (L); K. Oka 376220 (TNS); K. Shimabuku & K. Ohgami 4677 (TI); K. Yonekura 11218 (HAST); M. Kato et al. 166 (PPI, TI); R. Moran 4985 (L); S. Hatusima 17750 (L), 18802 (TI); S. Saito 1587 (TNM), 1816 (PE, TI); S. Tsugaru 341124 (TNS); S.W. Chung 5413 (TAIF); T. Amano 6960 (TNS); T. Fujii TWTw-12834 (TI); T. Kawasaki 4547; T. Kinjo 229114 (TNS); T. Nakai 11669 (TI); T. Nakaike 614 (L, MEL, PE); T. Yamazaki 6427 (PE, TI); W.P. Leu & C.C. Wang 1922 (HAST, TNM); Wright s.n. (K, L); Y. Kimura & I. Furusawa 5719 (TI); Y. Taateishi & J. Murata 4899 (PE)
.
D. woodsonianum T.C. Huang
INDONESIA. N. Sumatra, Gaju & Alas Lends, C.G.G.J. van Steenis 8362 (L, K, BO), 8654 (BO, L). Gunung Banda-hara, W.J.J.O. de Wilde & B.E.E. Wilde-Duyfjies 15442 (L, K, KEP). Gunung Leuser Nature Reseve, W.J.J.O. de Wilde & B.E.E. Wilde-Duyfjies 16002 (L, K).
MALAYSIA. Sabah, Mt. Kinabalu, Mt. Trus Madi, A. Bium SNP 05043 (SNP).
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Botanical Studies, Vol. 53, 2012
虎皮楠科植物雌花的多樣性其及分類價值
唐默詩1,2 楊遠波1,3 蔡奇助4,5 許秋容6
1國立中山大學生物科學系
2中華醫事科技大學製劑製造工程學系
3大葉大學生物資源學系
4高雄區農業改良場
5屏東科技大學生物科技系
6國立中興大學生命科學系
虎皮楠科植物僅具虎皮楠一屬，雌雄異株；花單性，不顯著，結構返化且往往早落。其葉、雄花和
果實可資應用的分類特徵十分有限；與前者相較，雌花的特徵較多且穩定，對其分類極有幫助。然而雌
花的相關報導卻仍相當稀少，諸此原因，造成本屬物種鑑定之困難和屬下分類位階--節或系等的不確
定性。本研究觀察19個分類群，報導虎皮楠屬雌花的多樣性和其發育之主要變化，除了提供本科植物
分類特徵之參考外，也有助於屬下的分類評估。虎皮楠的雌花柱頭可分成四型：點狀、腎形、線形和長
橢圓形。除了點狀柱頭外，其他三型的柱頭在開花的過程中都會有延長的現象。在研究的19個分類群
中,三個分類群的雌花無花萼，其他分類群的花萼可區分成三型：萼片離生，合生深裂和合生淺裂。不
孕性雄蕊的存在和宿存與否亦為鑑別特徵。本研究結果支持本屬的屬下分類系統依黃氏原先之處理，分
為三個節：虎皮楠節、半月形藥節和不妊性雄蕊節，並建議將原屬於虎皮楠節其中的二物種歸屬為不妊
性雄蕊節。
關鍵詞：早落的；花萼；虎皮楠節；半月形藥節；不孕性雄蕊；不妊性雄蕊節；柱頭；分類價值。