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1 2007, 29 (5) : Acta Botanica Yunnanica 1, 2, 1, 3 (1, ; 2, ; 3, ) : 26 ( 1 2 ) 4 6,,,,, :,,,,,, 3,, : ; ; : Q 944 : A : (2007) Leaf Epidermal Morphology of Lasianthus ( Rubiaceae) and Its Related Genera in Asia * CAI Min 1, 2, ZHU Hua 1 * *, WANG Hong 3 ( 1 Kunming Section of Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming , China ; 2 Graduate University of the Chinese Academy of Sciences, Beijing , China ; 3 Kunming Institute of Botany, Chinese Academy of Sciences, Kunming , Ch ina) Abstract : Leaf epidermal morphology of 26 species ( including 1 subspecies and 2 varieties) of Lasianthus and 6 species of its related 4 genera Morinda, Psychotria, Saprosma and Urophyllum, were investigated under both light microscope ( LM) and scanning electron microscope ( SEM). It was shown that the shapes of upper leaf epidermal cells varies from irregularly sinuolate, nearly elliptic to irregularly polygonal. The lower leaf epidermal cells are usually irregularly sinuous or sinuate. The patterns of anticlinal walls are sinuate irregularly, and the ambits are inconspicuous between the cells. It was found that the cell walls of both upper and lower epidermis are covered with thick corticle. Under SEM, the cuticula membrane of the leaf epidermis is distinct, and granular, reticulate, scaly or papilliform in shape. The stomatal apparatus is of paracytic type and occurs only on the lower surface. The stomatas of some species are sunken. Leaf indumentum of Lasianthus species can be classified into single-celled hairs and uniseriate multi- celled hairs including linear, twisted and straight-walled hairs. The leaf epidermal morphology of the related genera in Rubiaceae is similar to Lasianthus. In conclusion, the leaf micro-morphological characteristics have some significance in identification of Lasianthus species, but show regular varia- : ( ) : : Author for correspondence , : ( ),, Tel: + 86 ( 0) ; minc@ xtbg. ac. cn

2 tion with habitats. The variations of the ratio between stomatal polar axis and outer stomatal ledge aperture, as well as leaf indumentum of Lasianthus species are well correspondent to their habitats. Key words: Leaf epidermal ; Morphology; Lasianthus ( Lasianthus Jack) ( Rubiaceae), 180, ( 160 ) (1 ) ( 20 ) ( 2 ), ( Robbrecht, 1988) 1823,,,, ( Zhu, 2002 ), ( Stace, 1966; Hickey, 1973, 1979; Theobald, 1979) ( Pan, 1990; Shui, 1999; Sun, 1999, 2003; Christophel, 1996), (David, 1974), ( Stace, 1966, 1984; Heywood, 1984),,, (, 1994, 1998, 2001b; Zhu, 2001a, 2002;, 2000),,,, 1 26 ( 1 2 ) 4 6 ( 1), (HITBC), ( KUN), : 2 3, 5 min, 1 cm 1 cm Jefferys ( 10% 10% = 1 1), h,,,, ( ) 1% 12 h, (50 %, 75%, 80%, 95% ),, BX51 BX52 OLYMPUS 3, 20,,, KYKY-1000B, Dilcher (1974) Wilkinson (1979) , ( : 1 5),,, L. biermannii, L. hirsutus, L. japonicus, ( : 2, 3) :,,,, ( : 25 29) 2.2, ( : 9 12),,, ( : 30, 31) ( single-cell hairs ) ( : 20, : 52 ) (line-type hairs ) ( : 19, : 59, 60) ( : 22, 23), ( : 19, 21, 24), ( : ),, m ( : 57, 60)

3 5 : ,, ( : 46) m, m, : ( : ( : 11, 12, : 14, 15 ) : ( : 33, 36, : 38, 42),,, ( : 34, 33, 35, 36, : 37, 41 45) ( : 39, 41 ) : 34, : 38 40, 47) ( : 37) 1 Table 1 Origin of materials Species Locality Voucher Lasianthus attenuatus Jack Mengyang, Yunnan ( K. M. Feng) (KUN) L. austrosinensis H. S. Lo Dinghucaotangken, Guangdong ( K.Q. Ding) 553 ( KUN) L. biermannii King ex Hook. f. Lincangdaxueshanqu, Yunnan ( T. P. Zhu ) 79 ( KUN) L. calycinus Dunn. Locality Unknown ( X.Q. Liu) ( KUN ) L. chevalieri Pitard Hainan Hainan Exped. 317 (1955 ) ( KUN) L. chinensis Benth. Longjin, Guangxi ( S.Q. Chen ) 1324 ( KUN) L. chrysoneurus Miq. Cangyuan, Yunnan ( K. D. Tao ) (HITBC) L. fordii Hance Lingshui, Hainan ( L. Deng) 3011 ( KUN) L. formosensis Matsum Bubeng, Mengla, Yunnan ( H. Zhu ) 2522 ( HITBC) L. hainanensis Merr. Qiongzhong, Hainan (S. Q. Chen) ( KUN) L. henryi Hutch Mengsong, Jinghong, Yunnan ( H. Zhu ) ( HITBC) L. hirsutus ( Roxb.) Merr. Bawanglin, Hainan ( K.D. Tao) ( KUN) L. hirsutus ( Roxb.) Merr. Sumatra K. lwatsuki, et. 926 ( HITBC) L. hispidulus ( Drake) Pitard Dahuangmaoshan, Vietnam ( H.D. Zeng) ( KUN) L. hookeri C. B. Clarke Cangyuan, Yunnan ( Y.H. Li) ( KUN) L. hookeri var. dunnaiana H. Zhu Xishuangbanna, Yunnan ( H. Wang) ( HITBC) L. inodorus Blume Kinabalu, Borneo, Malesia ( H. Zhu ) ( HITBC) L. japonicus Miq. Rongjiang, Guizhou Qinnan Team 2933 (KUN) L. japonicus var. latifolius H. Zhu Locality Unknown ( Q.A. Wu) 7500 ( KUN) L. lancifolius Hook. f. Maguan, Yunnan ( H. Sun) 1466 ( KUN) L. lucidus Bl. Qingmai, Thailand ( H. Zhu ) ( KUN) L. micranthus Hook. f. Locality Unknown ( H. Zhu ) 2508 ( HITBC) L. rhinocerotis Bl. Kinabalu, Borneo, Malesia ( H. Zhu ) ( HITBC) L. rhinocerotis subsp. xishuangbanna- Middle of Thailand ( H. Zhu ) ( KUN) ensis H. Zhu & H. Wang L. rigidus Miq. Cangyuan, Yunnan ( K.D. Tao) ( KUN) L. schmidtii K. Schum. Simao, Yunnan Unknown 6445 (HITBC) L. sikkimensis Hook. f. Mengsong, Jinghong, Yunnan ( H. Zhu ) ( HITBC) L. trichophlebus Hemsl. var. Hainan Hainan Exped. 520 latifolius H. Zhu (1955 ) ( KUN) L. verticillatus ( Lour.) Merr. Menglun, Banna, Yunnan ( Y.H. Li) 1507 ( HITBC) Morinda umbellata Linne. Dulong River, Yunnan ( K. M. Feng) (KUN) Morinda hainanensis Merr. et How Locality Unknown ( K. Z. Hou) ( KUN ) Psychotria prainii L vl. Luchun to Lixianjiang, Yunnan ( T. D. Tao) 573 ( KUN) Psychotria henryi L vl. Luchun, Yunnan (Y. M. Shui) ( KUN) Saprosma ternatum Hook. f. Mengla, Yunnan (X.W. Li ) ( KUN) Urophyllum chinensis Merr. et Chun Bubeng, Mengla, Yunnan ( K.D. Tao) 9178 (KUN)

4 500 29

5 5 : 501

6 502 29

7 5 : , ( : ), ( : 61) :,, ( : 61 69),,, : ( single-cell hairs),,,,, 52) ( : 20, : ( uniseriate multi-cell hairs),,, 2, 30 ( : ), ( : 22, 23, : 57, : 62, 63 ),,, ( : 61, 63, 67),, ( : 52, 57, : 62) ( : 57, 58, 60) : (1) ( twist hairs ) :,, 2 10, 10 ( : 24, : 57, 58) (2 ) (line-type hairs ) :, 2 5, ( : 19, : 59, 60) (3) ( straight-wall hairs ) :, 10,, ( : 22, 23, : 61, 63, 65) 3 3.1,,, ;,,,,,,, (, 1991 ), -,,,,,,,, ( L. fordii, L. henryi, L. hirsutus, L. hispidulus, L. japonicus var. latifolius, L. sikkimensis ),, : ( L. hirsutus ),,,,,,, 3.2

8 504 29,, : ( L. hainanensis),, ( Saprosma merrillii) (, 1999 ),,,,, : ( L. hookeri) ( L. hookeri var. dunnaiana), ( L. japonicus) ( L. japonicus var. latifolius ), ( L. rhinocerotis ) ( L. rhinocerotis subsp. xishuangbannaensis),, ( L. formosensis), (, 2007), ( Morinda L.) ( M. hainanensis), ( Psychotria Linn.) ( P. prainii) ( P. henryi) ( Urophyllum Merr.) ( U. chinensis), ( Saprosma Bl.) ( S. ternatum), 3.3,,,, (, 1991),,, ( Prabhakar, 2004 ),,,, ( ), ( ),,, ( Wilkinson, 1979), ( Theobald, 1979),,,, ( : 54, 59), Cai M ( ), Zhu H ( ), Wang H ( ), Pollen morphology of the genus Lasi anthus ( Rubiaceae) and related taxa in Asia [ J]. Acta Phytotax Sin ( ) ( in press) Christophel DC, Kerrigan R, Rowett AI, The use of cuticular features in the taxonomy of the Lauraceae [ J]. Ann Missouri Bot Gard, 83: David LD, Approaches to the identification of Angiosperm leaf r e- mains [ J ]. The Botanical Review [ C ], Vol. 40, January- March, No.1 Dilc her DL, Approaches to the identification of Angiosperm leaf r e- mains [ J ]. The Botanical Review [ C], 40 : Heywood VH, Stace CA, Moore DM, Current concepts in plant taxonomy [ C], Systematics Association Special Volume No. 25 [ M]. London and Orlando: Academic Press Hickey LJ, Classification of the architecture of Dicotyledonous leaves [ J]. Amer J Bot, 60: Hickey LJ, A revised classification of the architecture of Dicotyledonous leaves [ A]. In: Metcalf CK, Chalk L. Anatomy of the Dicotyledons ( 2nd edition ), Vol. 1 [ M]. Oxford: Clarendon Press, Lo HS ( ), Lasianthus Jack [ A ]. In: Flora Reipublicae Popularis Sinicae [M]. Beijing: Science Press, 71 (2 ) :

9 5 : 505 Lu SW ( ), Xu XS ( ), Shen MJ ( ), Botany (2nd edition) [M]. Beijing: Higher Education Press, Pan KY ( ), Lu AM ( ), Wen J ( ), Characters of leaf epidermis in Hamamelidaceae ( S. L) [ J]. Acta Phytotax Sin ( ), 28 ( 1) : Prabhakar M, Structure, delimitation, nomenclature and classification of stomata [ J]. Acta Bot Sin ( ), 46 (2 ) : Robbrecht E, Tropical woody Rubiaceae [ J]. Opera Bot Belgica, 1: 132 Shui YM ( ), Li QR ( ), Huang SH ( ), Observation of leaf Epidermis and its hair of Begonia from Yunnan [ J ]. Acta Bot Yunnan ( ), 21 ( 3) : Stace CA, The use of ep idermal characters in phylogenetic considerations [ J ]. New Phytologist, 65 : Stace CA, Cuticular studies as an aid to plant taxonomy [ J]. Bulletin British Museum ( Natural History) Botany, 4: 3 8 Sun TX ( ), Hu YX ( ), Lang KY ( ), A Study on micromorphological characters of leaf epidermis of Neottianthe in China [ J ]. Acta Bot Yunnan ( ), 21 ( 1 ) : Sun TX ( ), Yao H ( ), Wu H ( ) et al Comparative anatomy on leaves of Polyalthia (Annonaceae) [ J]. Wuhan Bot Res ( ), 21 ( 1) : 9 17 Theobald WL, Krahulik JK, Rolli ns RC, Trichome description and classification [ A], Anatomy of the Dicotyledons ( 2nd edition ) [ M]. Oxford: Clarendon Press, Wilkinson HP, The plant surface ( mainly leaf ) [ A], Anatomy of the Dicotyledons ( 2nd edition ) [ M ]. Oxford: Clarendon Press, Zhu H ( ), A taxonomic study on the genus Lasianthus Jack in China [ J ]. Acta Phytotax Sin ( ), 32 (1) : Zhu H ( ), Notes on the genus Lasianthus Jack ( Rubiaceae) from Asia [ J ]. Acta Bot Yunnan ( ), 20 ( 2 ) : Zhu H ( ), Wang H ( ), Li BG ( ), A new subspecies of the genus Lasianthus Jack ( Rubiaceae) from Yunnan with a reference to its biogeographical implication [ J ]. Acta Phytotax Sin ( ), 38 (3 ) : Zhu H, 2001a. New plants of Lasianthus Jack ( Rubiaceae) from Kinabalu, Borneo and its biogeographical implication [ J]. Blumea, 46: :,, 100 m 1 5 : 1. ; 2. ; 3. ; 4. ; : 6. ; 7. ; : ; 11. ; 12. :,, 100 m : 13. ; 14. ; 15. ; 16. ; 17. ; : 19. ; 20. ; ; 23. ; 24. :,, m, 33, 34, m : 25. ; 26. ; 27. ; 28. ; : 30. ; 31. ; 32. ; 33. ; 34. ; 35. ; 36. :,, m, 10 m 37. ; 38. ; 39. ; 40. ; 41. ; 42. ; 43. ; 44. ; 45. ; ; 48. :,, m, 50 m : 49. ; 50. ; : 52. ; 53. ; 54. ; ; 57. ; 58. ; 59. ; 60. :,, m, 50 m ; 63. ; 64. ; 65. ; 66. ; 67. ; 68. ; 69. ; 70. ; 71. ; 72. Explanation of Plates Zhu H ( ), 2001b. A taxonomic revision of the genus Lasianthus Jack ( Rubiaceae) from Thailand [ J ]. Acta Phytotax Sin ( ), 39 ( 2) : Zhu H, A revision of the genus Lasianthus Jack ( Rubiaceae) from China [ J ]. Syst Geogr Plant, 72: Plate : 1-12 Under LM, the epidermal cell morphology of Lasianthus and some related genera on the upper and the lower leaf surfaces. Bars, 100 m. 1-5 Epidermal cell morphology and thick corticle of Lasianthus on the upper leaf surface : 1. L. attenuatus; 2. L. micranthus; 3. L. f- ordii; 4. L. lancifolius; 5. L. hispidulus. 6-8 Epidermal cell morpholo-

10 gy of the related genera on the upper leaf surface: 6. Psychotria prainii; 7. Morinda umbellata; 8. Morinda hainanensis Epidermal cell morphology and the stomata of Lasianthus on the lower leaf surface : L. henryi; 11. L. micranthus ; 12. L. hispidulus. Plate : Under LM, the epidermal stomata and the indumentum morphology of Lasian thus and its related genera on the lower leaf surface. Bars, 100 m Stomata morphology: 13. L. sikkimensis; 14. L. hookeri; 15. L. chrysoneurus; 16. L. rhinocerotis subsp. xishuangbannaensis; 17. Saprosma ternatum; 18. Urophyllum chinensis Indumentum morphology: 19. L. hookeri var. dunniana; 20. L. chrysoneurus; L. hirsutus; 23. L. attenuatus ; 24. L. hookeri. Plate : Under SEM, the epidermal cell corticle and the stomata morphology of Lasianthus and its related genera on the upper and the lower leaf surfaces. Bars, 50 m for and 35, 10 m for 33, 34, Epidermal corticle on the upper leaf surface: 25. L. henryi; 26. L. sikkimensis; 27. L. chevalieri; 28. L. japonicus; 29. L. hai nanensis Epidermal cell morphology and the stomata on the lower leaf surface: 30. L. rhinocerotis; 31. L. fordii; 32. Urophyllum chinensis ; 33. L. hirsutus ; 34. L. chinensis ; 35. L. hispidulus; 36. L. henryi. Plate : Under SEM, the stomata morphology of Lasianthus and its related genera. Bars, 50 m for 37 and 46, 10 m for the others. 37. L. sikkimensis; 38. L. hookeri; 39. L. chrysoneurus; 40. L. hookeri var. dunniana; 41. L. japonicus; 42. L. rhinocerotis subsp. xishuangbannaensis; 43. L. formosensis ; 44. L. hainanensis ; 45. L. schmidtii; L. biermannii; 48. Psychotria henryi. Plate : Under SEM, the stomata and the indumentum morphology of Lasianthus and its related genera. Bars, 10 m for 49 and 51, 50 m for the others Stomata morphology: 49. Morinda hainanensis; 50. Saprosma ternatum; 51. Urophyllum chinensis Indumentum morphology: 52. L. hirsutus; 53. L. formosensis; 54. L. hispidulus; L. hirsutus; 57. L. sikkimensis; 58. L. hookeri; 59. L. chrysoneurus; 60. L. hookeri var. dunniana. Plate : Under SEM, the indumentum morphology of Lasianthus and its related genera. Bars, 10 m for 62 and 68, 50 m for the others L. hirsutus; 63. L. hispidulus; 64. L. verticillatus; 65. L. chevalieri; 66. L. lancifolius; 67. L. japonicus; 68. L. biermannii; 69. L. rhinocerotis subsp. xishuangbannaensis; 70. Psychotria prain ii; 71. L. schmidtii; 72. Urophyllum chinensis.

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Ps22Pdf 2006, 28 (6) : 587 592 Acta Botanica Yunnanica ( ) 1, 2, 1 (1, 650091; 2, 541004) : 5 16,,, : ; ; : Q 944, Q 949 : A : 0253-2700( 2006)06-587 - 06 Morphological and Anatomical Studies of Subfam. Polypodioideae