全 文 ：假瘤蕨属 (水龙骨科) 植物叶表皮特征的比较研究*
邵摇 文1, 陆树刚1**, 商清春2
(1 云南大学地植物学与生态学研究所, 云南 昆明摇 650091; 2 牡丹江医学院影像系, 黑龙江 牡丹江摇 157011)
摘要: 在光镜和扫描电镜下观察了假瘤蕨属 2 系、 5 亚系的 24 种植物的叶表皮。 结果表明, 叶表皮细胞
形状不规则, 垂周壁波曲状。 下生气孔, 气孔类型有极细胞型、 共环极细胞型、 腋下细胞型、 聚腋下细胞
型和不规则型, 常伴生出现。 在扫描电镜下, 叶表皮气孔器外拱盖内缘为浅波状、 少平滑; 一些种的保卫
细胞两端有 “T冶 型加厚; 角质膜波状有条纹, 有时附有颗粒。 叶表皮的一些特征对该属部分种的区分有一
定的参考价值。 叶表皮的微形态特征, 如气孔器类型和垂周壁类型特征不能用来界定假瘤蕨属的系和亚系。
关键词: 水龙骨科; 假瘤蕨属; 叶表皮; 系统学意义
中图分类号: Q 944摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2011)02-174-09
Comparative Morphology of Leaf Epidermis in the Fern
Genus Phymatopteris (Polypodiaceae)
SHAO Wen1, LU Shu鄄Gang1**, SHANG Qing鄄Chun2
(1 Institute of Ecology and Geobotany, Yunnan University, Kunming 650091, China;
2 Department of Physics, Mudanjiang Medical College, Mudanjiang 157011, China)
Abstract: Leaf epidermises of 24 taxa, representing 2 series and 5 subseries of the genus Phymatopteris (Polypodi鄄
aceae), were examined by light microscopy (LM) and scanning electron microscopy (SEM) to determine the sys鄄
tematically informative characters. The leaf epidermal cells were usually irregular in shape. The patterns of anticlinal
walls were sinuolate, sinuous, and sinuate. The stomatal apparatus were restricted in abaxial epidermises for all spe鄄
cies, and generally could be assigned to polocytic, copolocytic, coaxillocytic, axillocytic, and anomocytic types.
Under SEM observation, the inner margin of the outer stomatal rim was often sinuous and seldom smooth. Strikingly
tall, upright outer stomatal rims occurred in most species. The cuticular membrane of the leaf epidermis was sinuous
and occasionally granular. Leaf epidermal features in Phymatopteris appeared to be constant within species, and thus
could be used for distinguishing some species. The microcharacters of leaf epidermises ( the patterns of anticlinal
walls and the stomatal apparatus types) could not be used as evidence for the classification of the series and sub鄄
series in Phymatopteris.
Key words: Polypodiaceae; Phymatopteris; Leaf epidermis; Systematic significance
摇 摇 Phymatopteris Pichi鄄Serm. (Polypodiaceae) was
a natural group, characterized by simple, trifid,
palmatifid, or pinnatifid leaves with usually charta鄄
ceous texture, glaucous underneath, single鄄rowed
but usually superficial or slightly immersed large so鄄
ri, and the drynarioid type of venation ( Ching,
1964). This genus has been confused with Microso鄄
rum, Crypsinus and Selliguea by Copeland (1947),
Holttum (1954) and Hovenkamp (1998); however
there are several characteristics that can be used to
植 物 分 类 与 资 源 学 报摇 2011, 33 (2): 174 ~ 182
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 3724 / SP. J. 1143. 2011. 09238
*
**
Foundation items: The National Natural Science Foundation of China (30770164, 30970186)
Author for correspondence; E鄄mail: shuganglu@ 163. com
Received date: 2009-11-27, Accepted date: 2010-06-11
作者简介: 邵文 (1979-) 女, 博士研究生, 主要从事蕨类植物系统分类学研究。
distinguish these four genera (Shao and Lu, 2009).
Phymatopteris has more than 60 species and ma鄄
jority of them are from the mainland of Asia. Ching
(1964) recognized 2 series and 5 subseries within Phy鄄
matopteris based on characteristics of the frond shape
and the frond margin, i. e. series Hastatae, including
subseries Griffithianae and subseries Hastatae; and
series Oxylobae, including subseries Oxylobae, sub鄄
series Ebenipedes, and subseries Malacodontes.
Leaf epidermal characteristics such as deposi鄄
tion of wax crystalloids, hair types and stomata types
are of potential taxonomical importance. However,
few studies on the leaf epidermis of this genus have
been done (Wang, 2007). No observations under
SEM have been reported. The aims of this paper are
to present a more detailed study on the leaf epider鄄
mal features of the 24 species of Phymatopteris for
the first time.
1摇 Materials and methods
Adult leaves of 24 taxa were gathered from the
specimens of PYU (Table 1). Samples were taken
from fully expanded sun鄄exposed leaves. The materi鄄
als for LM observation were boiled in water before
being macerated in Jeffery忆s solution. Pieces of leaf
epidermises were stained in a solution of safranin in
50% alcohol before being mounted in glycerin gel.
To check the constancy of epidermal structure, at
least five slides were made from different parts of a
single leaf or from different leaves for each species.
The material for SEM observation was directly
mounted on stubs without any treatment, and sputter
coated with gold鄄palladium. The specimens were ex鄄
amined and photographed using an AMARY鄄1000B
SEM. Descriptive terminology of leaf epidermis fol鄄
lows Zhang (1999), Dilcher (1974), and Fryns鄄
Claessens and van Cotthem (1973).
2摇 Results
The characteristics of leaf epidermis of the Phy鄄
matopteris are listed in Tables 2 and 3. It appears
that stomatal and other epidermal features are con鄄
stant within species and thus may be used in analysis
and discussion.
Table 1摇 Vouchers for leaf epidermis of Phymatopteris
Taxon Locality Voucher
Phymatopteris albopes (C. Chr. et Ching) Pichi鄄Serm. Pingbian, Yunnan, China (云南屏边) W. Shao (邵文) 017
P. chrysotricha (C. Chr. ) Pichi鄄Serm. Lushui, Yunnan, China (云南泸水) W. Shao (邵文) 003
P. crenatopinnata (C. B. Clarke) Pichi鄄Serm. Xinping, Yunnan, China (云南新平) W. Shao (邵文) 004
P. dactylina (Christ) Pichi鄄Serm. Deqin, Yunnan, China (云南德钦) W. Shao (邵文) 009
P. daweishanensis S. G. Lu Pingbian, Yunnan, China (云南屏边) S. G. Lu (陆树刚) 28885
P. ebenipes (Hook. ) Pichi鄄Serm. Lushui, Yunnan, China (云南泸水) W. Shao (邵文) 021
P. engleri (Luerss. ) Pichi鄄Serm. Yakushima Island, Japan (日本屋久岛) S. Mitsuta 267
P. glaucopsis (Franch. ) Pichi鄄Serm. Eryuan, Yunnan, China (云南洱源) W. M. Chu (朱维明) 23013
P. griffithiana (Hook. ) Pichi鄄Serm. Lushui, Yunnan, China (云南泸水) W. Shao (邵文) 001
P. hastata (Thunb. ) Pichi鄄Serm. Pingbian, Yunnan, China (云南屏边) W. Shao (邵文) 002
P. majoensis (C. Chr. ) Pichi鄄Serm. Yongshan, Yunnan, China (云南永善) W. M. Chu (朱维明) 4898
P. nigrovenia (Christ) Pichi鄄Serm. Deqin, Yunnan, China (云南德钦) W. M. Chu (朱维明) 26458
P. omeiensis (Ching) Pichi鄄Serm. Daxiangling, Sichuan, China (四川大相岭) X. X. Kong (孔宪需) 3971
P. oxyloba (Wall. ex Kunze) Pichi鄄Serm. Xinping, Yunnan, China (云南新平) W. Shao (邵文) 013
P. pianmaensis W. M. Chu Lushui, Yunnan, China (云南泸水) W. M. Chu (朱维明) 11350
P. rhynchophylla (Hook. ) Pichi鄄Serm. Pingbian, Yunnan, China (云南屏边) W. Shao (邵文) 029
P. shensiensis (Christ) Pichi鄄Serm. Kangding, Sichuan, China (四川康定) X. X. Kong (孔宪需) 6235
P. stewartii (Bedd. ) Pichi鄄Serm. Lushui, Yunnan, China (云南泸水) W. Shao (邵文) 019
P. stracheyi (Ching) Pichi鄄Serm. Lushui, Yunnan, China (云南泸水) W. Shao (邵文) 018
P. subebenipes (Ching) Pichi鄄Serm. Dali, Yunnan, China (云南大理) W. Shao (邵文) 005
P. tenuipes (Ching) Pichi鄄Serm. Mt. Jinfoshan, Chongqing, China (重庆金佛山) Z. Y. Liu (刘正宇) 3547
P. tibetana (Ching) W. M. Chu Deqin, Yunnan, China (云南德钦) W. M. Chu (朱维明) 23725
P. trisecta (Baker) Pichi鄄Serm. Kunming, Yunnan, China (云南昆明) W. Shao (邵文) 024
P. yakushimensis (Makino) Pichi鄄Serm. Yakushima Island, Japan (日本屋久岛) Murata s. n.
5712 期摇 摇 摇 摇 SHAO Wen et al. : Comparative Morphology of Leaf Epidermis in the Fern Genus Phymatopteris摇 摇 摇 摇 摇 摇
Table 2摇 Leaf epidermal characters of Phymatopteris under LM (surface view)
Taxa
Adaxial epidermis Abaxial epidermis
Shape
of
cells
Pattern of
anticlinal
walls
Shape
of
cells
Pattern of
anticlinal
walls
Size range
of stomata
(滋m2)
Stom鄄
atal
index
Stomatotype (% )
Pol Axi Coa Ano Cop
摇 Plate玉
Series Hastatae
Subseries Griffithianae
P. griffithiana irr sinuous irr sinuous 48. 2伊42. 3 25. 8 46. 7 39. 4 — 13. 9 — 1,2
P. chrysotricha irr sinuous irr sinuolate 46. 1伊38. 9 27. 8 61. 6 25. 7 — 5. 7 7 3,4
Subseries Hastatae
P. rhynchophylla irr sinuate irr sinuous 43. 6伊36. 7 19. 0 45. 5 45 — 9. 5 — 5,6
P. dactylina irr sinuous irr sinuolate 46. 6伊38. 9 14. 8 25 50. 6 — 24. 4 — 11,12
Series Oxylobae
Subseries Oxylobae
P. oxyloba irr sinuate irr sinuate 47. 9伊41. 7 27. 1 48. 6 26. 9 — 16. 5 8 7,8
P. trisecta irr sinuolate irr sinuous 49. 5伊45. 6 22. 4 43. 8 41. 4 — 9. 2 5. 4 9,10
Subseries Ebenipedes
P. albopes irr sinuous irr sinuous 48. 7伊39. 8 24. 3 44. 6 32. 7 — 12. 7 10 13,14
P. ebenipes irr sinuous irr sinuous 48. 9伊44. 6 29. 7 32. 7 37 5. 8 14. 8 9. 7 15,16
Subseries Malacodontes
P. crenatopinnata irr sinuous irr sinuate 52. 1伊39. 9 24. 3 50. 6 29. 4 4 10 6 17,18
P. stewartii irr sinuolate irr sinuous 48. 8伊43. 4 30. 1 39 37 — 34 — 19,20
irr = irregular; Pol =polocytic; Axi = axillocytic; Coa=coaxillocytic; Ano=anomocytic; Cop=copolocytic
Table 3摇 Leaf epidermal characters of Phymatopteris under SEM (surface view)
Taxa Cuticular layer Shape of guard cells Outer stomatal rims Plates
Series Hastatae
Subseries Griffithianae
P. majoensis apophysis and granules, many in Ab; few in Ad suborbiculate sinuolate 域:25,26
P. chrysotricha many granules (Ad & Ab), few apophysis (Ab) suborbiculate sinuolate 域:27,28
Subseries Hastatae
P. engleri few sinuous stripes (Ad), few apophysis and granules (Ab) suborbiculate sinuolate 域:21,22
P. yakushimensis many sinuolate stripes (Ad), many sinuous stripes (Ab) suborbiculate nearly smooth 域:23,24
P. tenuipes many apophysis and granules (Ad & Ab) suborbiculate sinuolate 域:29,30
P. hastata many granules (Ad & Ab), few apophysis (Ab) wide elliptic unknown 域:31,32
P. omeiensis few rounded apophysis (Ada & Aba) suborbiculate nearly smooth 域:33,34
P. dactylina many granules (Ad & Ab), few apophysis (Ad) suborbiculate unknown 域:35,36
Series Oxylobae
Subseries Oxylobae
P. trisecta many sinuous stripes and many hairs (Ad & Ab) suborbiculate unknown 域:37,38
P. oxyloba many sinuous stripes (Ad & Ab) suborbiculate unknown 域:39,40
Subseries Ebenipedes
P. tibetana many sinuous stripes (Ad), many granules (Ab) suborbiculate unknown 芋:41,42
P. subebenipes few sinuous stripes (Ad), many granules (Ab) suborbiculate unknown 芋:43,44
P. pianmaensis few sinuolate stripes and many hairs (Ad & Ab) suborbiculate unknown 芋:45,46
P. nigrovenia few sinuous stripes and many hairs (Ad & Ab) suborbiculate sinuolate 芋:47,48
P. daweishanensis few sinuolate stripes (Ad & Ab) wide elliptic sinuolate 芋:49,50
Subseries Malacodontes
P. shensiensis few sinuous stripes (Ad), few sinuolate stripes (Ab) wide elliptic sinuolate 芋:51,52
P. stracheyi few sinuous stripes (Ad & Ab) wide elliptic unknown 芋:53,54
P. glaucopsis few sinuous stripes (Ad & Ab) suborbiculate sinuolate 芋:55,56
P. crenatopinnata few sinuous stripes and hairs (Ad & Ab) wide elliptic unknown 芋:57,58
P. stewartii few sinuous stripes (Ad), many apophysis (Ab) wide elliptic nearly smooth 芋:59,60
Ad=adaxial epidermis; Ab=abaxial epidermis
671摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
2. 1摇 Characteristics of leaf epidermis under LM
Epidermal cells. The epidermal cells of the
Phymatopteris as seen under LM are usually irregular
in shape, with anticlinal walls sinuolate, sinuous or
sinuate. The patterns of anticlinical cells may vary
in different species or between adaxial (Ad) and aba鄄
xial (Ab) epidermises of the same species: sinuo鄄
late walls are found in P. chrysotricha and P. trisecta
(Ad & Ab, Plate 玉: 4, 9), sinuous walls in P.
griffithiana and P. albopes (Ad & Ab, Plate 玉: 1,
2, 13, 14), and sinuate walls in P. oxyloba and P.
crenatopinnata (Plate 玉: 7, 8, 18).
Stomatal apparatus. All species studied here
have the stomatal apparatus restricted in abaxial epi鄄
dermises. The shape of the stomata is usually subor鄄
biculate. The stomata can be polocytic, copolocytic,
axillocytic, anomocytic or less often coaxillocytic.
However, these types of stomata occur often in a sin鄄
gle leaf.
The size range of stomata is (43. 6 -52. 1) 伊
(36. 7-45. 6) 滋m2 . The stomatal density is diffe鄄
rent among species. The stomatal index ranged from
30. 1% in P. stewartii, with the stomata being dense鄄
ly distributed, to 14. 8% in P. dactylina, with the
stomata being sparsely distributed.
2. 2 摇 Characteristics of leaf epidermis under
SEM
Guard cells. The outlines of the pair of guard
cells of the Phymatopteris are variable, from usually
suborbiculate to wide elliptical as seen in the surface
view, with length / width (L / W) ratio 1. 1-1. 3 颐 1.
Guard cells are often thickened to a certain extent,
are made up of outer stomatal ledges or rims, and
are nearly smooth or sinuolate. Nearly smooth outer
stomatal rims occur in P. stewartii and P. omeiensis
(Plate 域: 34, Plate 芋: 60), and sinuolate outer
stomatal rims are present in P. glaucopsis and P. ni鄄
grovenia (Plate 芋: 48, 56).
Strikingly tall and upright outer stomatal rims
occur in all species, except for P. omeiensis, P. tri鄄
secta, and P. glaucopsis ( Plate 域: 34, 38, Plate
芋: 56), in which the stomatal rims are almost at
the same level with the epidermis. Furthermore, the
cuticular thickening of the common wall between the
guard cells is found in some species, such as P.
chrysotricha and P. dactylina (Plate 域: 28, 36).
Cuticular and wax ornamentation. Stripes,
hairs, and wax granules are present in both sides of
epidermises of the specimens investigated. Two
types of the cuticular stripes are found: sinuolate
and sinuous. The outer surface of the cuticle is
smooth to sinuolate in P. pianmaensis and P. daweis鄄
hanensis ( Plate 芋: 45, 46, 49, 50); while the
sinuous stripes are found in P. stracheyi (Plate 芋:
53, 54). Wax granules (d抑20 滋m) are present in
P. dactylina and P. tibetana ( Plate 域: 35, 36,
Plate 芋: 41, 42). A few hairs are distributed on
the abaxial epidermis of P. crenatopinnata and the
adaxial epidermis of P. nigrovenia (Plate 芋: 47),
also also both sides of epidermises of P. trisecta.
2. 3摇 Key to the Phymatopteris species studied with special reference to foliar morphological characters
1a. Fronds simple, trilobed or palmately divided 2………………………………………………………………………………
1b. Fronds pinnate or pinnatifid 11………………………………………………………………………………………………
2a. Fronds simple 3………………………………………………………………………………………………………………
2b. Fronds trilobed or palmately divided 10………………………………………………………………………………………
3a. Fronds venation is simple, costal areoles have no forked veinlets, and the excurrent included veinlets dominate in most
areoles 4………………………………………………………………………………………………………………………
3b. Fronds venation is variable, veinlets are often forked, rarely simple on costal areoles, and the recurrent included veinlets
dominate in most areoles 6……………………………………………………………………………………………………
4a. Fronds dimorphic, the sterile much shorter, ovate, obtuse at apex; the fertile much longer, lanceolate
P. rhynchophylla……………………………………………………………………………………………………………
4b. Fronds subdimorphic, sterile fronds rarely present, fertile parts similar to sterile parts 5…………………………………
5a. Rhizome few pruinose, frond membranaceous, obtuse at apex, rounded at base P. tenuipes………………………………
7712 期摇 摇 摇 摇 SHAO Wen et al. : Comparative Morphology of Leaf Epidermis in the Fern Genus Phymatopteris摇 摇 摇 摇 摇 摇
5b. Rhizome pruinose, frond chartaceous, acute at apex, cordate at base P. chrysotricha……………………………………
6a. Fronds dimorphic, the sterile much shorter, ovate, obtuse at apex; the fertile much longer, lanceolate P. griffithiana…
6b. Fronds monomorphic, with no differentiation of fertile and sterile fronds 7…………………………………………………
7a. Many wax granules on abaxial and adaxial epidermis P. majoensis…………………………………………………………
7b. Few or no granules on abaxial or adaxial epidermis 8………………………………………………………………………
8a. Obtuse at apex, few rounded apophysis on both sides of epidermis P. omeiensis……………………………………………
8b. Acuminate at apex, no rounded apophysis on abaxial or adaxial epidermis 9………………………………………………
9a. Lamina oblanceolate, outer stomatal rims sinuolate P. engleri………………………………………………………………
9b. Lamina lanceolate, outer stomatal rims nearly smooth P. yakushimensis……………………………………………………
10a. Fronds palmately divided P. dactylina………………………………………………………………………………………
10b. Fronds trilobed, many wax granules on both sides of epidermis P. hastata…………………………………………………
11a. Fronds trilobed to pinnatifid 12………………………………………………………………………………………………
11b. Fronds pinnatifid to pinnate 13………………………………………………………………………………………………
12a. Anticlinal walls of epidermis sinuolate or sinuous, more hairs on both sides of epidermis P. trisecta……………………
12b. Anticlinal walls of epidermis of fronds sinuate, few hairs on both sides of epidermis P. oxyloba…………………………
13a. Notches usually present, with a shallowly serrate at each pair of vein 14…………………………………………………
13b. Notches usually absent, with biserrate at each pair of vein 20……………………………………………………………
14a. Many wax granules on abaxial and adaxial epidermis 15……………………………………………………………………
14b. Few or no wax granules on abaxial or adaxial epidermis 17…………………………………………………………………
15a. Lateral lobes all ascending P. tibetana………………………………………………………………………………………
15b. Lowest lobes deflexed or slightly deflexed 16………………………………………………………………………………
16a. Lamina obtuse at apex, pinnae 3-5 (7) pairs P. subebenipes……………………………………………………………
16b. Lamina acuminate at apex, pinnae 4-10 (13) pairs P. ebenipes…………………………………………………………
17a. Many hairs on adaxial epidermis 18…………………………………………………………………………………………
17b. Few or no hairs on abaxial or adaxial epidermis 19…………………………………………………………………………
18a. Few sinuolate stripes on both sides of epidermis, more hairs on both sides of epidermis P. pianmaensis…………………
18b. Few sinuous stripes on both sides of epidermis, more hairs on adaxial epidermis P. nigrovenia…………………………
19a. Lowest lobes ascending P. daweishanensis…………………………………………………………………………………
19b. Lowest lobes deflexed or slightly deflexed P. albopes………………………………………………………………………
20a. Lateral lobes acuminate or caudately acuminate at apex 21…………………………………………………………………
20b. Lateral lobes obtuse or acute at apex 22……………………………………………………………………………………
21a. Lowest lobes deflexed, few apophysis on abaxial epidermis P. stracheyi……………………………………………………
21b. Lowest lobes ascending, many apophysis on abaxial epidermis P. stewartii………………………………………………
22a. Few sinuolate stripes on abaxial epidermis, pinnae 2-3 (4) pairs P. shensiensis…………………………………………
22b. Few sinuous stripes on abaxial epidermis, pinnae 2-5 (7) pairs 23………………………………………………………
23a. Lobes crenate or undulate at margin, shape of guard cells wide elliptic P. crenatopinnata………………………………
23b. Lobes diserrate at margin, shape of guard cells suborbiculate P. glaucopsis………………………………………………
3摇 Discussion
The leaf epidermal cells in the fern genus Phy鄄
matopteris are usually irregular in shape, and the sto鄄
matal apparatus are restricted in abaxial epidermises.
The main stomatal apparatus types in all species exami鄄
ned in this paper are polocytic and axillocytic. The
cuticular membrane of the leaf epidermis is generally
sinuolate to sinuous. The leaf epidermal features a鄄
mong species are fairly similar to each other, which
indicate the affinity of Phymatopteris (Sen, 1981).
Polocytic and axillocytic stomatal apparatus
types are indicated as more primitive types ( Pant,
1965), but as more evolutional types in Polypodi鄄
aceae (Thurston, 1969). Furthermore, Wang (2007)
accepted Thurston忆s point, and found the evidence
in Phymatopteris crenatopinnata, which also has
been found in this paper (Plate 玉: 18).
The patterns of anticlinal walls and the stomata
871摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 33 卷
types are constant within species, and thus can be
used to distinguish closely related species. We also
find these two characteristics distributed in all sub鄄
series randomly, and thus cannot be used as evi鄄
dence for the classification of the series and sub鄄
series in this genus.
The cuticular and wax ornamentation ( stripes,
hairs, and wax granules) are present in some epi鄄
dermises of the specimens investigated. Wax gran鄄
ules are in P. dactylina, P. subebenipes and P. tibet鄄
ana, hairs in P. crenatopinnata, P. nigrovenia and
P. trisecta, and small granules in P. tenuipes, P.
hastate, P. majoensis. Though it may be affected by
the living environment in some degree, the wax and
hair appendages are constant within species and thus
provide some anatomical evidence for the classifica鄄
tion of some species.
The original function of stomata is the limitation
of water loss in land plants and the maintenance of
homoiohydry, while allowing gas exchange, which
places constraints on stomatal patterning within the
epidermises. The guard cells of stomatal apparatus
in this paper are all slightly sunken, and most stoma
pores are closed, which may be suitable to their epi鄄
phytic habitus.
Our results suggest that the characteristics of the
leaves, i. e. the shape of frond, the margin and ap鄄
pendage character of frond are constant within species
and thus can provide some anatomical evidence for the
classification of Phymatopteris in some species. How鄄
ever, the microcharacters of leaf epidermises, such as
the patterns of anticlinal walls and the stomatal ap鄄
paratus types cannot serve as a criterion of distin鄄
guishing the series and subseries in Phymatopteris.
Acknowledgements: We would like to thank Prof. W. M.
Chu for providing access to the specimens cited in this paper,
and Prof. R. X. Wang and engineer X. K. Fan for assistance
on the observation of LM and SEM. The first author would al鄄
so thank Prof. Jerry Li and Prof. Damian Shea of North Caro鄄
lina State University for helpful suggestions and critical read鄄
ing of the manuscript.
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Explanation of Plates
Plate 玉摇 1 -20. The characters of leaf epidermis of Phymatopteris
under LM. 摇 1-2. Phymatopteris griffithiana, 3-4. P. chrysotricha,
5-6. P. rhynchophylla, 7-8. P. oxyloba, 9-10. P. trisecta, 11-12.
P. dactylina, 13-14. P. albopes, 15 -16. P. ebenipes, 17 -18. P.
crenatopinnata, 19-20. P. stewartii.
Plate 域摇 21-40. The characters of leaf epidermis of Phymatopteris
under SEM. 摇 21 -22. Phymatopteris engleri, 23 -24. P. yakushi鄄
mensis, 25-26. P. majoensis, 27-28. P. chrysotricha, 29-30. P. te鄄
nuipes, 31-32. P. hastata, 33-34. P. omeiensis, 35-36. P. dactyli鄄
na, 37-38. P. trisecta, 39-40. P. oxyloba.
Plate 芋 摇 41-60. The characters of leaf epidermis of Phymatopteris
under SEM. 摇 41-42. Phymatopteris tibetana, 43-44. P. subebeni鄄
pes, 45-46. P. pianmaensis, 47-48. P. nigrovenia, 49-50. P. da鄄
weishanensis, 51-52. P. shensiensis, 53-54. P. stracheyi, 55-56.
P. glaucopsis, 57-58. P. crenatopinnata, 59-60. P. stewartii.
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