全 文 ：Journal of Systematics and Evolution 46 (1): 62–72 (2008) doi: 10.3724/SP.J.1002.2008.06036
(formerly Acta Phytotaxonomica Sinica) http://www.plantsystematics.com
Pollen morphology of the genus Lasianthus (Rubiaceae) and
related taxa from Asia
1,2Min CAI 1Hua ZHU 3Hong WANG*
1(Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming 650223, China)
2(Graduate University of Chinese Academy of Sciences, Beijing 100049, China)
3(Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China)
Abstract Pollen morphology of 16 Asian species including two subspecies and one variety of the genus Lasian-
thus and five species from five related genera was examined by light microscopy and scanning electron micros-
copy. Lasianthus Jack is eurypalynous. The pollen grains are small to medium in size and spheroidal or subprolate
in shape, with a few being prolate. Pollen morphology is remarkably diverse, particularly in aperture and exine
ornamentation. The apertures are usually (3–)4–(–5)-zonoaperturate, being porate and pororate. The number of
apertures varies from three to five, with the majority of species being 3–4 zonoaperturate. The exine is usually
perforate, finely reticulate or coarsely reticulate. Pollen data appear to be morphologically informative and useful
for distinguishing among species of Lasianthus, and for elucidating the relationships among some species.
Key words Lasianthus, pollen morphology, Rubiaceae.
Lasianthus Jack of the family Rubiaceae is a
large genus with more than 180 species. It has a
pantropical distribution, occurring in Asia, Africa and
America, with one species in Australia (Zhu, 2001a).
The greatest number of species is found in tropical
Asia (Robbrecht, 1988).
The genus Lasianthus was first described by Jack
in 1823 (Jack, 1823). Most Lasianthus species are
shrubs, including some small trees. They grow almost
exclusively in the understory of primary forests, with
occasional records from secondary or seriously dis-
turbed forests or forest edges. The genus is an ecol-
ogically important element in tropical forests of Asia
(Zhu, 2001a), making up a quantitatively important
component of the vegetation.
Hooker (1880) recognized some 52 species from
British India, which was the first regional floristic
work in Asia. Subsequent regional floristic accounts
have been published for the former “Indo-Chine”
(Pitard, 1924) and Thailand (Craib, 1934). Later
Yamazaki (1964) revised the genus for the Ryukyu
Islands and Liu & Chao (1964) treated the Taiwanese
species. Deb & Gangopadhyay (1989, 1991) pub-
lished a revision of the genus for India, and Ridsdale
(1998) for Sri Lanka as well as Zhu (2001b) for
Thailand. Zhu (1994) revised the Chinese species of
the genus, and Lo (1999) treated the genus for Flora
of China, recognizing 34 species and ten varieties.
Zhu (1998, 2002) revised the Chinese species again
and recognized a total of 33 species, four subspecies
and three varieties in China.
Wight (1846) attempted the first infrageneric
classification of Lasianthus, dividing the genus into
two groups (i.e., Bracteosae and Nudiflorae), which
were not typified or given a specific rank. Subsequent
infrageneric classifications were made by Hooker
(1880), Ridley (1923), and Gangopadhyay and Chak-
rabarty (1992). Hooker’s system, comprising four
sections (i.e., Stipulares, Bracteatae, Nudiflorae and
Pedunculatae), was commonly accepted, despite the
absence of exclusive characters distinguishing any of
these sections. Deb and Gangopadhyay (1989) re-
moved sect. Pedunculatae Hook. f. and treated it in
the genus Litosanthes Bl., but later Gangopadhyay &
Chakrabarty (1992) returned the taxa that had previ-
ously been in the sect. Pedunculatae from Litosanthes
back to Lasianthus again. Lo (1999) and Zhu (1994,
1998, 2001b) treated the genus without recognizing
any infrageneric taxa, and Zhu (2002) treated the
genus Litosanthes as a synonym of Lasianthus.
The current classifications of Lasianthus are
mainly based on traditional macromorphology. Recent
molecular work revealed the phylogenetic relation-
ships of Lasianthus and other related genera (Bremer,
1996; Andersson & Rova, 1999; Bremer & Manen,
2000; Piesschaert et al., 2000; Xiao & Zhu, 2007). A
group of genera (Lasianthus, Saldinia and
Trichostachys) was recognized at tribal level as
Lasiantheae Bremer & Manen on a strongly supported
branch (Bremer & Manen, 2000), which was sup-
ported by further molecular work (Xiao & Zhu, 2007),

———————————
Received: 8 March 2006 Accepted: 13 December 2007
* Author for correspondence. E-mail: .
CAI et al.: Pollen morphology of Lasianthus from Asia


63
and is currently accepted.
Determining infrageneric relationships in
Lasianthus is extremely difficult because the delimita-
tion of some species is obscure (Zhu, 1994, 2002). In
addition, flowers and fruits are small and are often
missing from herbarium specimens, leaving taxono-
mists few distinct diagnostic characters. The majority
of the characters used to determine species are quanti-
tative features of the vegetative organs, such as leaves,
stipules and bracts. Except for L. fordii Hance (Wang
et al., 1997), no palynological data is available for this
genus. The present study, therefore, aims to examine
the major pollen characteristics of the genus and
provide additional support to classification treatments
of difficult species or section division currently recog-
nized within Lasianthus.
1 Material and methods
Pollen of 16 species including two subspecies
and one variety of Lasianthus and five species of
related genera were included in this study, and exam-
ined and summarized in Table 1. Specimens were
examined from the following herbaria: Xishuang-
banna Tropical Botanical Garden (HITBC) and Kun-
ming Institute of Botany, Chinese Academy of Sci-
ences (KUN) (abbreviations according to Holmgren et
al., 1990). A list of voucher specimens for pollen
samples is given in Appendix I.
For light microscopy (LM), pollen samples were
acetolysed (Erdtman, 1960), mounted in glycerol jelly
and sealed with paraffin. Measurements of equatorial
diameter (E) and polar axis (P) of individual pollen
grains were based on 20 grains per sample.
For scanning electron microscopy (SEM), the
acetolysed samples were mounted on glass cover slips
and attached to aluminium stubs. After sputter coating
with gold, pollen grains were examined and photo-
graphed under SEM (KYKY-10000B, Science In-
strument Company, Beijing) at 15 kV. Descriptive
terminology of the pollen follows Punt et al. (1994)
and Lens et al. (2000).
2 Results
2.1 General description of Lasianthus pollen
grains
Pollen grains of Lasianthus were generally iso-
polar, but some showed a slight heteropolarity (e.g., L.
rhinocerotis Bl.).
The size of pollen grains varied from small to
medium. The most common size had P (a polar axis) =
40–42 µm and E (an equatorial axis) = 37–39 µm. The
smallest grains (L. formosensis Matsum.) had P =
19.56 µm and E = 15.75 µm. The largest grains (L.
japonicus Miq.) had P = 53.13 µm and E = 45.38 µm.
Three distinct shape classes were recognized:
prolate (P/E: 1.33; Figs. 1, 2), spheroidal (P/E: 1.11;
Figs. 4, 5) and subprolate (P/E: 1.17; Figs. 15, 20), of
which spheroidal pollen grains were the most com-
mon, making up 74% of the specimens investigated.
Prolate pollen grains were only found in L. formosen-
sis, while subprolate pollen grains were observed in
three taxa: L. chinensis (Champ.) Benth., L. attenuatus
Jack, and L. japonicus ssp. longicaudus C. Y. Wu &
H. Zhu. The polar views of the grains were usually
circular, e.g. in L. hookeri (Fig. 7), triangular, quad-
rangular or pentangular with convex sides in L. hain-
anensis Merr. (Figs. 11, 12), L. biermannii King ex
Hook. f. (Fig. 23) and L. rhinocerotis (Fig. 26).
Pollen grains were usually zonoaperturate, most
of which were pororate. Aperture number varied from
three up to five. Approximately 63% of the pollen
were 3- and 4-aperturate in the same sample exam-
ined. Three-porate pollen was seen in L. rhinocerotis
ssp. xishuangbannaensis H. Zhu & H. Wang. Colpo-
rate pollen with long colpi, extending nearly to the
two poles, was observed in L. formosensis.
The compound apertures consisted of an ec-
toporus and an endoporus. In L. rhinocerotis ssp.
xishuangbannaensis (Fig. 5). the endoporus and
ectoporus were congruent with a circular ecto-aperture
present (termed congruent apertures by Lens et al.,
2000).
The ectopori were usually elongated and elliptic.
The ends of the ectopori were obtuse (Figs. 31, 35), or
acute (Figs. 23, 27, 30). Sometimes the margin of the
ectoaperture was irregular with sexine bulges (Figs.
28, 31), or was very smooth (Figs. 30, 32). Ectoaper-
ture membranes were often covered with irregular
microverrucae (Figs. 27, 28), or could not easily be
observed due to closing of ectopori. The endoaperture
was irregular in shape, and was difficult to observe
under both LM and SEM.
All grains were tectate with either perforate or
reticulate tecta. Perforate tecta were present only in L.
attenuatus (Fig. 36) and L. japonicus ssp. longicaudus
(Fig. 37). Perforations were variable in size and shape
and were smaller at the poles. The reticulate sexine
was divided into two types according to variations in
the diameter of the lumina. They were finely reticulate
(lumina less than 1 µm in diameter) and coarsely
reticulate (lumina greater than 1 µm in diameter). The
Journal of Systematics and Evolution Vol. 46 No. 1 2008

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CAI et al.: Pollen morphology of Lasianthus from Asia


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Figs. 1–12. SEM of Lasianthus pollen. 1–3. L. formosensis. 1. Equatorial view. 2. Equatorial view indicating 3-zonocolporate aperture (arrow). 3.
Polar view. 4, 5. L. rhinocerotis ssp. xishuangbannaensis. 4. Polar view. 5. Equatorial view with 3-zonoporate aperture. 6–12. Polar view and
3-zonopororate aperture. 6. L. rhinocerotis. 7. L. hookeri. 8. L. rigidus. 9. L. chinensis. 10. L. attenuatus. 11, 12. L. hainanensis. Scale bars = 10 µm.
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Figs. 13–24. SEM of Lasianthus pollen. 13–18. Polar view and 3-zonopororate aperture. 13. L. inodorus. 14. L. japonicus. 15. L. japonicus ssp.
longicaudus. 16. L. fordii. 17. L. chrysoneurus. 18. L. sikkimensis. 19–24. Polar view and 4-zonopororate aperture. 19. L. hookeri var. dunniana. 20. L.
chinensis. 21. L. attenuatus. 22. L. inodorus. 23. L. biermannii. 24. L. austroyunnanensis. Scale bars = 10 µm.
CAI et al.: Pollen morphology of Lasianthus from Asia


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Figs. 25–36. SEM of Lasianthus pollen. 25. L. lucidus, polar view and 4-zonopororate aperture. 26, 27. L. rhinocerotis. 26. Polar view with
5-zonopororate aperture. 27. Same under higher magnification, showing mesocolpium. 28, 29. L. schmidtii. 28. Higher magnification of the mesocol-
pium. 29. Equatorial view. 30–35. Higher magnification of the mesocolpium. 30. L. rigidus. 31. L. chinensis. 32. L. inodorus. 33. L. biermannii. 34. L.
fordii. 35. L. sikkimensis. 36. L. attenuatus, higher magnification of the exine sculpture. Scale bars = 10 µm for whole pollen grains; =5 µm for details
of the exine sculpturing.
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Figs. 37–48. SEM of Lasianthus pollen. Features of exine ornamentation. 37. L. japonicus ssp. longicaudus, higher magnification of the exine
sculpturing. 38. L. formosensis. 39. L. rhinocerotis ssp. xishuangbannaensis. 40. L. hookeri. 41. L. hookeri var. dunniana. 42. L. hainanensis. 43. L.
inodorus. 44. L. japonicus. 45. L. austroyunnanensis. 46. L. chrysoneurus. 47. L. lucidus. 48. L. sikkimensis. Scale bars = 5 µm.
CAI et al.: Pollen morphology of Lasianthus from Asia


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Figs. 49–60. SEM pollen morphology of other five related genera. 49, 50. Brachytome wallichii. 49. Polar view with 3-zonopororate aperture. 50.
Finely reticulate with sexine. 51, 54. Saprosma ternatum. 51. Polar view with 5-zonopororate aperture. 54. Finely reticulate with sexine. 52, 53.
Psychotria henryi. 52. Polar view with 3-zonopororate aperture. 53. Coarsely reticulate with sexine. 55–57. Prismatomeria sp. 55, 57. Polar view with
(3–)4–(–5)-zonopororate aperture. 56. Coarsely reticulate with sexine. 58–60. Urophyllum chinensis. 58. Equatorial outline view. 59. Sunken colpori.
60. Polar view with 3-zonopororate aperture. Scale bars = 10 µm for whole pollen grains; =5 µm for details of the exine sculpturing.
Journal of Systematics and Evolution Vol. 46 No. 1 2008

70
lumina were commonly elliptical, circular, or angular
in shape (Figs. 39, 41, 44, 48). Some species exhibited
granules on the inside of the lumina, such as L. hain-
anensis (Fig. 42), L. japonicus (Fig. 44) and L. chry-
soneurus (Fig. 46). The muri usually lacked supratec-
tal ornamentation, except in L. chinensis, which had
dense microverrucae (Fig. 31).
2. 2 Pollen description of the related genera
2.2.1 Brachytome wallichii Pollen grains were
medium (P: 49.06 µm×E: 39.88 µm) in size and
spheroidal (P/E: 1.13) in shape. Apertures were 3- or
4-colporate. The polar outline view was circular, with
colpori narrow and slightly sunken (Fig. 49). The
ectoaperture was usually elongated. The ends of the
ectopori were acute. The sexine was finely reticulate
(Fig. 50), with the muri protruding.
2.2.2 Saprosma ternatum Pollen grains were
medium (P: 47.13 µm×E: 40.19 µm) in size and
subprolate (P/E: 1.17) in shape. Apertures were
(3–)4–(–5)-pororate. The polar outline view was
circular with the colpori wide and short (Fig. 51). The
ends of the ectopori were acute or obtuse. The sexine
was finely reticulate (Fig. 54), with some mi-
cro-granules on the muri.
2.2.3 Psychotria henryi Pollen grains were large
(P: 51.75 µm×E: 45.19 µm) in size and spheroidal
(P/E: 1.10) in shape. Apertures were 3-pororate. The
polar outline view was circular (Fig. 52). The ends of
the ectopori were obtuse. The sexine was coarsely
reticulate (Fig. 53), and the lumina were very large
(mean 1–2.5 µm).
2.2.4 Prismatomeria sp. Pollen grains were
medium (P: 42.25 µm×E: 37.94 µm) in size and
spheroidal (P/E: 1.10) in shape. Apertures were
(3–)4–(–5)-pororate. The polar outline view was
circular (Figs. 55, 57). The ectoaperture was often
elongated. The ends of the ectopori were acute. The
sexine was coarsely reticulate (Fig. 56), with the
irregular lumina in shape and size.
2.2.5 Urophyllum chinensis Pollen grains were
medium (P: 22.50 µm×E: 19.50 µm) in size and
subprolate (P/E: 1.20) in shape. Apertures were
3-colporate. The polar outline view was circular to
almost triangular (Fig. 60), with the colpori sunken
(Fig. 59). The equatorial outline view was elliptic with
comparatively acute apices (Fig. 58). Ectopori were
very long, sometimes narrower at the center (Fig. 59).
The ectocolpus margins were irregular. The ends of
the ectopori were obtuse. The sexine was perforate,
with the lumina irregular in shape and size, and the
perforations present in the amb and near ectocolpus
margins.
3 Conclusions and Discussion
Pollen grains showed a considerable morpho-
logical and structural polymorphism within the genus
Lasianthus and among related genera. Pororate 3–4-
zonoaperturate pollen grains, the most dominant type
in Lasianthus, have been reported otherwise in the
Rubiaceae. They have previously been seen in Gar-
denieae-Gardeniinae (Persson, 1993), Isertieae
(Huysmans et al., 1993), Coptosapelteae (Huysmans,
1998) and Vanguerieae (Havard & Verdcourt, 1987).
In addition, L. formosensis and L. rhinocerotis ssp.
xishuangbannaensis had unique 3-colporate and
3-porate pollen grains, respectively.
There were three general types of exine orna-
mentation in the genus Lasianthus: perforate, finely
reticulate and coarsely reticulate. Perforate pollen
grains were less frequent, and seen only in L. attenu-
atus and L. japonicus ssp. longicaudus.
The pollen morphological characteristics of the
five related taxa examined here were similar to those
of Lasianthus, providing little diagnostic characteris-
tics for distinguishing Lasianthus from these related
genera.
However, the pollen studies did provide distinc-
tive characters for some taxonomically difficult
species. Lasianthus formosensis was previously
distinguished from L. fordii only by the indumentum,
which is usually with dense and much spreading
tomentose hairs in the former and with thin or sparse
and appressed pubescence in the latter. Indumentum,
however, often found to be environmentally variable.
The former species has even been reduced to a pu-
berulent variety of the latter (Yamazaki, 1964). Zhu
(2002) treated them as two separate species based
upon differences in the length of lanceolate calyx
lobes and the dense or sparse indumentum. Our
palynological results indicate that L. formosensis has
the less common 3-colporate pollen grains, which is
conspicuously different from L. fordii, supporting the
separation of the two species. Aperture or exine types
can also be quite different in infra-specific taxa. For
instance, L. rhinocerotis ssp. rhinocerotis has more
diversified pororate pollen grains (3, 4 to 5-apertu-
rate), whereas ssp. xishuangbannaensis has only
3-porate apertures. Lasianthus japonicus ssp. japoni-
cus has common reticulate pollen tecta, while the ssp.
longicaudus has uncommon perforate pollen tecta.
Additionally, L. chinensis has unique pollen grains
with dense microverrucae on the muri. Our results
suggest that more detailed palynological work could
be useful in resolving taxonomic issues within the
genus Lasianthus.
CAI et al.: Pollen morphology of Lasianthus from Asia


71
Acknowledgements This project was funded by
The National Natural Science Foundation of China
(Nos. 30770158, 30670160). The authors thank the
herbaria of Xishuangbanna Tropical Botanical Garden
(HITBC) and Kunming Institute of Botany (KUN),
Chinese Academy of Sciences for providing pollen
samples from collections. Special thanks are given to
senior Engineer Xi-Kai FAN of KUN for his technical
assistance in SEM, and also the great help of
Chun-Fang LI in preparing the figures. We are very
grateful to Dr. Chuck CANNON from Texas Techni-
cal University, USA, for reading and commenting on
the manuscript. We also thank reviewers very much
for their important and constructive comments.
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Appendix Ⅰ Voucher information for taxa used in this study
Lasianthus attenuatus Jack, Mengla, Yunnan (云南勐腊), H. Zhu (朱华) 2059 (HITBC).
L. austroyunnanensis H.Zhu, Funing, Yunnan (云南富宁), Q. A. Wu (武全安) 9556 (KUN).
L. biermannii King ex Hook. f., Yongde, Yunnan (云南永德), T. P. Zhu (朱太平) 0079 (KUN).
L. chinensis (Champ.) Benth., Mengla, Yunnan (云南勐腊), G. D. Tao et al. (陶国达等) 0114 (HITBC).
L. chrysoneurus Miq., Cangyuan, Yunnan (云南沧源), G. D. Tao & Y. W. Li (陶国达, 李锡文) 40052 (HITBC).
L. fordii Hance, Lingshui, Hainan (海南陵水), L. Deng (邓良) 3011 (KUN).
L. formosensis Matsum., Mengla, Yunnan (云南勐腊), H. Zhu (朱华) 2522 (HITBC).
L. hookeri C. B. Clarke ex Hook. f., Mengla, Yunnan (云南勐腊), H. Zhu (朱华) s.n. (HITBC).
L. hookeri var. dunniana H. Zhu, Mengla, Yunnan (云南勐腊), S. J. Pei (裴盛基) 59-9361 (HITBC).
L. inodorus Bl., Kinabalu, Borneo, H. Zhu (朱华) s.n. (HITBC).
L. japonicus Miq., Rongjiang, Guizhou (贵州榕江), Qiannan Team (黔南队) 2933 (KUN).
L. japonicus ssp. longicaudus C. Y. Wu & H. Zhu, Guangnan, Yunnan (云南广南), C. W. Wang (王启无) 88109 (KUN).
L. japonicus var. lancilimbus C. Y. Wu & H. Zhu, Yangshan, Guangdong (广东阳山), L. Deng (邓良) 1272 (KUN).
L. lucidus Bl., Jinghong, Yunnan (云南景洪), H. Zhu (朱华) 2508 (HITBC).
L. rhinocerotis Bl., Kinabalu, Borneo, H. Zhu (朱华) s.n. (HITBC)
L. rhinocerotis ssp. xishuangbannaensis H. Zhu, Jinghong, Yunnan (云南景洪), H. Zhu (朱华) 981101 (HITBC).
L. rigidus Miq., Cangyuan, Yunnan (云南沧源), G. D. Tao & X. W. Li (陶国达, 李锡文) 40023 (KUN).
L. schmidtii K. Schum., Yunnan (云南), C. W. Wang & Y. B. Zhang (王启无, 张音波) 86022 (KUN).
L. sikkimensis Hook. f., Mengla, Yunnan (云南勐腊), H. Zhu (朱华) 2180 (HITBC).
L. verticillatus (Lour.) Merr., Mengla, Yunnan (云南勐腊), H. Zhu (朱华) 1315 (HITBC).
L. hainanensis Merr. = Saprosma merrilii Lo, Hainan (海南), S. K. Lau (刘心祈) 26545 (KUN).
Brachytome wallichii Hook. f., Lüchun, Yunnan (云南绿春), Lüchun Team (绿春队) 838 (KUN).
Saprosma ternatum Hook. f., Xishuangbanna, Yunnan (云南西双版纳), Anonymous 8234 (KUN).
Prismatomeria sp., Yunnan (云南), Anonymous 623 (HITBC).
Psychotria henryi Lévl., Mengla, Yunnan (云南勐腊), G. D. Tao et al. (陶国达等) 0268 (HITBC).
Urophyllum chinensis Merr. & Chun., Thailand, Sino-Thai Botanic Exp. (中国-泰国植物调查队) 1726 (KUN).
亚洲茜草科粗叶木属及其相关属植物花粉形态研究
1,2蔡 敏 1朱 华 3王 红*
1(中国科学院西双版纳热带植物园 昆明 650223)
2(中国科学院研究生院 北京 100049)
3(中国科学院昆明植物研究所生物多样性与生物地理学重点实验室 昆明 650204)

摘要 利用光学显微镜和扫描电镜研究了茜草科粗叶木属Lasianthus 16种2亚种、1变种及相关的5属5种的花粉形态。粗叶木
属的花粉属于广孢型, 单粒。一般中等大小, 绝大多数为圆球形, 少数为近长球形或长球形。花粉形态特征, 特别是在萌发孔
和外壁纹饰上表现出多样化。根据孔沟的数目或是否具有内孔, 可以将萌发孔分为(3–)4–(–5)孔沟和3孔。在所观察的这些种
中, 萌发孔以3–4孔沟为主要类型, 比例为62.4%。外壁纹饰可分为细网状、粗网状和穴状。有部分种的花粉极面有穴状纹饰,
其余均为网状纹饰。网眼一般椭圆形、近圆形、三角形或者不规则形。少数外壁纹饰网脊上有颗粒状雕纹或模糊的颗粒, 网
脊轮廓线呈波浪形, 一般凸出且平滑。大部分种的花粉具有沟膜, 沟膜上具有瘤状突起或小颗粒状, 沟边缘一般较平滑, 或粗
糙, 有的种具有沟桥。
关键词 粗叶木属; 花粉形态; 茜草科