全 文 ：Journal of Systematics and Evolution 46 (6): 933–937 (2008) doi: 10.3724/SP.J.1002.2008.07144
(formerly Acta Phytotaxonomica Sinica) http://www.plantsystematics.com
Revision of two species of Araucaria (Araucariaceae) in Chinese
taxonomic literature
1,2Hai-Sang LIU* 1Ci-Quan LIU
1(Modern Biology Research Center, Yunnan University, Kunming 650091, China)
2(Xiamen Overseas Chinese Subtropical Plant Introduction Garden, Xiamen 361002, China)
Abstract Based on the lectotypes of Araucaria cunninghamii and A. heterophylla, the status of those species as
cited in Chinese taxonomic literature, such as Flora of Guangzhou, Dendrology of China, Iconographia Cormo-
phytorum Sinicorum, Flora Reipublicae Popularis Sinicae, Sylva Sinica, Flora of China and Higher Plants of
China, was reviewed. Araucaria heterophylla has linear, acute juvenile leaves held at an angle of less than 45° to
the branchlet and without an obvious adaxial keel and scale-like adult leaves on terminal branchlets; A. cunning-
hamii has bilaterally compressed, subulate, shortly pungent juvenile leaves held at an angle of 45°–90° and with
an obvious keel on both surfaces and tetragonous, subulate adult leaves on terminal branchlets. The names A.
cunninghamii and A. heterophylla have been misapplied in Chinese taxonomic literature and should be A. hetero-
phylla (Salisb.) Franco and A. cunninghamii Aiton ex A. Cunn. respectively.
Key words Araucaria cunninghamii Aiton ex A. Cunn., Araucaria heterophylla (Salisb.) Franco, taxonomic
revision.
The genus Araucaria Juss. includes plants of
great economic and ornamental value. For example, A.
cunninghamii is one of the five major ornamental trees
for warm temperate parks and A. heterophylla is well
known worldwide as a pot or tub plant (Bailey, 1914;
Den Ouden & Boom, 1965; Boutelje, 1980; Everett,
1981; Ying, 1985①). The phylogeny of three genera of
Araucariaceae, Araucaria, Agathis and Wollemia, was
presented by Gilmore and Hill (1997), and by NPWS
(1998).
The use of the name Araucaria cunninghamii
(Hoop pine) in different articles and books has been
inconsistent (Backer & van den Brink, 1963; Ying,
1985①; Huxley, 1992; GRIN, 1994; Hill, 1998; Far-
jon, 1998, 2001; IPNI, 2004). In Chinese taxonomic
literature (How, 1956; Cheng, 1961; Cheng & Fu,
1964; Institute of Botany, Chinese Academy of Sci-
ences, 1972; Institute of Botany, Jiangsu Province,
1977; Cheng & Fu, 1978; Cheng et al., 1982; Fu et al.,
1999; Fu, 2000), the names A. cunninghamii and A.
heterophylla have been confused, because two speci-
mens, Y. Jiang (Y. Tsiang) 16605 and S. Q. Chen (S.
H. Chun) 7347, had been wrongly identified as A.
cunninghamii and then cited in the Flora of Guang-
zhou in 1956. It should be noted that no species of
Araucariaceae are native to China; all trees of the
family are introduced and often of unknown (wild)
provenance.
1 Material and methods
Herbarium material of Araucaria cunninghamii
and A. heterophylla was examined in the following
herbaria: AU, FJSI, HITBC, IBSC, KUN, PE. Field
surveys were conducted in all areas where Araucaria
is grown in China. The lectotypes of Araucaria cun-
ninghamii (Fig. 1) and A. heterophylla (Fig. 2) were
used for comparison. Leaves of representative samples
(Table 1) were embedded in paraffin and sectioned.
The ratio of the width to the thickness of juvenile
leaves of Araucaria cunninghamii and A. heterophylla
was compared.
2 Results and discussion
Araucaria cunninghamii Aiton ex A. Cunn. in
Lamb., Descr. Pinus. 3rd ed. 2: unnumbered pages.
1832. Type: East Coast New Holland 1818–1829, A.
cunningham s.n. (Lectotype=specimen on far left of
sheet, K!).
Araucaria heterophylla auct. non (Salisb.)
Franco. W. C. Cheng, Dend. China 228. 1961; Insti-
tute of Botany, Jiangsu Province, Fl. Jiangsu. 85.
1977; W. C. Cheng & L. K. Fu in Fl. Reip. Pop. Sin.
7: 28. 1978; W. C. Cheng et al. in Sylva Sin. 1: 160.
1982; L. K. Fu et al. in Fl. China 4: 10. 1999; L. K. Fu
in Higher Plants of China 3: 13. 2000.
南洋杉
———————————
Received: 31 October 2007. Accepted: 21 August 2008
* E-mail: .
① Ying S-S. 1985. Coloured illustrated flora of Taiwan with the intro-
duced plants. Published by Author.
Journal of Systematics and Evolution Vol. 46 No. 6 2008 934































































Figs. 1–4. 1. The lectotype of Araucaria cunninghamii (A. cunningham s.n., K) (only seed cone and the branchlet attaching seed cone in the left of
picture). A, keeled on the abaxial of leaves; B, seed cone ovoid. Photo: A. Farjon. 2. The lectotype of A. heterophylla (P. G. King s.n., G). A, linear
and falcate leaves; B, acute apex being never pungent. Photo: A. Farjon. 3. Two forms of subulate leaves of A. cunninghamii (S. H. Chun 8280,
IBSC). A, bilaterally compressed leaves held at right angle to branchlet; B, adaxially depressed leaves encircling branchlet. Photo: H. S. Liu. 4.
Specimen of A. heterophylla (S. H. Chun 7347, IBSC) misidentified as A. cunninghamii in Flora of Guangzhou. A, scale-like leaves without keel on
the abaxial surface; B, subglobose seed cone. Photo: H. S. Liu.
LIU & LIU: Revision of two species of Araucariaceae in Chinese taxonomic literature

935
Juvenile leaves on terminal branchlets of Arau-
caria cunninghamii are subulate, shortly pungent and
held at an angle to the stem of 45°–90° (Fig. 3). The
ratio of thickness to width of juvenile leaves is usually
more than 1.7–3.1 to 1. Transverse paraffin sections
show the leaves to be elliptical in outline. Adult leaves
on the terminal branchlets of A. cunninghamii are
tetragonous and subulate (Figs. 1, 3). Both juvenile
and adult leaves have an obvious keel on both surfaces
(Fig. 3). Table 2 gives a morphological comparison of
A. cunninghamii and A. heterophylla.
Araucaria cunninghamii is endemic to Australia
and New Guinea. The oldest tree of Araucaria in
China, A. cunninghamii (misidentified as A. hetero-
phylla in The Marvellous Spectacle of Trees of China
(State Forestry Administration, P. R. China, 2003)),
was introduced in 1864 and is now 35 m tall and 123
cm in diameter.
Araucaria heterophylla (Salisb.) Franco in An. Inst.
Super. Agron. 19: 11. 1952. Type: Australia, Norfolk
Island, P. G. King s.n. (lectotype, G!).——Eutassa
heterophylla Salisb. in Trans. Linn. Soc. London 8:
316. 1807.
Araucaria cunninghamii auct. non Aiton ex A.
Cunn. F. C. How. Fl. Guangzhou. 68. 1956; W. C.
Cheng, Dend. China. 227. 1961; W. C. Cheng & L. K.
Fu in Fl. Hainan. 1: 214. 1964; Institute of Botany,
CAS, Icon. Cormophyt. Sin. 1: 316. 1972; W. C.
Cheng & L. K. Fu in Fl. Reip. Pop. Sin. 7: 28. 1978;
W. C. Cheng et al. in Sylva Sin. 1: 160. 1982; L. K.
Fu et al. in Fl. China 4: 9. 1999; L. K. Fu in Higher
Plants of China 3: 13. 2000.
诺福克南洋杉
The juvenile leaves on the terminal branchlets of
Araucaria heterophylla are falcate and linear. The
ratio of the thickness to width of the juvenile leaves is
1:1 or less. Transverse paraffin sections show the
lower portion of juvenile leaves to be quadrangular
and the upper portion to be obdeltoid. Both juvenile
and adult leaves have no obvious adaxial keel. The
adult leaves on the terminal branchlets of A. hetero-
phylla are scale-like. Bailey (1933) for Araucaria
cultivated in North America and Hill (1998) for species
of Araucaria of Australia, did not mention the growth
angle of the leaves on terminal branchlets. The adult
leaves, except on terminal branchlets, of A. hetero-
phylla are not obviously scale-like (Fig. 4). The apices
of the bracts of A. heterophylla are upcurved or re-
flexed (Fig. 4).
The bark of A. cunninghamii and A. heterophylla
is shown in Fig. 5 and Fig. 6.
Araucaria heterophylla is endemic to Norfolk
Island and it is the most famous plant of the island
(Green, 1994).
The specimen, J. J. Hong 235, was identified in-
correctly as A. cunninghamii in Flora Reipublicae
Popularis Sinicae, and specimens Y. Tsiang 16605
and S. H. Chun 7347 were identified wrongly as A.
cunninghamii.
As ornamental plants, A. heterophylla and Cycas
revoluta are common in southern China.

Table 1 Origin of materials
Taxon Locality Voucher
Araucaria cunninghamii Aiton ex A. Cunn. Xiamen, Fujian, China (福建厦门) H. S. Liu (刘海桑) 2008101 (AU)
A. heterophylla (Salisb.) Franco Xiamen, Fujian, China (福建厦门) H. S. Liu (刘海桑) 2008102 (AU)

Table 2 Comparison of Araucaria cunninghamii and A. heterophylla
Character Araucaria cunninghamii (Hoop Pine) Araucaria heterophylla (Norfolk Island Pine)
Crown of mature tree Wide conical. Narrow conical, esp. pagoda-shaped because of branches
on stem in regular wheels.
Bark Peeling in layers; exposing smooth inner bark
after outer bark is exfoliated.
Peeling in flakes; exposing rough inner bark after outer
bark is exfoliated.
Juvenile leaves on terminal
branchlets
8–16 mm long, 1–2 mm wide. Subulate, straight
or slightly curved; short pungent; keeled on both
adaxial and abaxial surfaces, bilaterally com-
pressed. Arranged loosely, at an angle of
45°–90° from their branchlet.
6–12.5 mm long, 1 mm wide. Linear, falcate; acute, not
pungent; slightly or not keeled on adaxial surface, never
bilaterally compressed. Arranged close, at an angle less
than 45° from their branchlet.
Adult leaves on terminal
branchlets
6–10 mm long, 2–3 mm wide. Subulate,
acuminate, obviously keeled on abaxial surface.
4–8 mm long, 3–6 mm wide. Scale-like, horny apex,
without obvious keel on abaxial surface.
Pollen cone 2–4 cm long, 0.5–0.7 cm in diameter. 4–9 cm long, 0.8–2 cm in diameter.
Seed cone 6–10 cm long, 5–7 cm in diameter. 7.5–12 cm long, 6–10 cm in diameter.
Seed size and 1000 seed weight 1.5–2 cm long, 0.7–1 cm wide; 150–300 g. 2.5–3 cm long, 1.2–1.5 cm wide; 1750–2500 g.
Journal of Systematics and Evolution Vol. 46 No. 6 2008 936




















Figs. 5, 6. 5. Bark of Araucaria cunninghamii (FJISB). A, slightly rough, peeling in layers; B, bright, smooth inner bark after exfoliation of outer
bark. Photo: H. S. Liu. 6. Bark of A. heterophylla (FJISB). A, rough, peeling in flakes. B, dull, rough inner bark after exfoliation of outer bark. Photo:
H. S. Liu.

Key to Araucaria cunninghamii and A. heterophylla
1a. Juvenile leaves bilaterally compressed, subulate, shortly pungent, diverging from branchlet at 45°–90°, transverse sections
elliptic, thickness to width ratio greater than 1.7 : 1; adult leaves on terminal branchlets tetragonous, subulate …A. cunninghamii
1b. Juvenile leaves linear, acute, diverging from branchlet at less than 45°, transverse sections quadrangular to obdeltoid, thickness to
width ratio not more than 1 : 1; adult leaves on terminal branchlets scale-like ……………………….……………A. heterophylla

3 Conclusions
The names Araucaria cunninghamii and A. het-
erophylla have been misapplied in China. Thus A.
cunninghamii and A. heterophylla, as cited in Chinese
literature, are A. heterophylla (Salisb.) Franco and A.
cunninghamii Aiton ex A. Cunn. respectively.
Acknowledgements We thank Aljos FARJON (K)
for photos of the lectotypes of A. cunninghamii and A.
heterophylla, Val STAJSIC and Josephine MILNE
(MEL) for information about the type of A. hetero-
phylla, Yong LI (SZG) and Li-Xiu GUO (IBSC) for
their help during our investigation, Yun LIN (FNU)
for discussing the revision of the paper, and Zhen-Yu
LI (PE) and David E. BOUFFORD (HUH) for cor-
recting the revision paper.
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中国分类学文献中南洋杉属两种之订正
1, 2刘海桑* 1刘次全
1(云南大学现代生物学研究中心 昆明 650091)
2(厦门华侨亚热带植物引种园 厦门 361002)
摘要 通过调查, 对Araucaria cunninghamii和A. heterophylla的选模式和中国分类学文献(如《广州植物志》、《中国树木学》、
《中国高等植物图鉴》、《中国植物志》、《中国树木志》、《Flora of China》、《中国高等植物》)的A. cunninghamii和A. heterophylla
(或A. excelsa)作比较, 证实中国分类学文献中的A. cunninghamii和A. heterophylla (或A. excelsa)分别是A. heterophylla (Salisb.)
Franco和A. cunninghamii Aiton ex A. Cunn.。这两种植物的叶形变化较大, 但有明显的区别: 诺福克南洋杉A. heterophylla幼树
的末级小枝的叶细长呈线形、叶尖急尖, 生长角小于45°, 腹面无明显的脊, 成龄株的末级小枝的叶扁平呈鳞片状; 南洋杉A.
cunninghamii幼树的末级小枝的叶呈两侧扁的钻形、叶尖短刺状, 生长角45°–90°, 腹背两面具明显的脊, 成龄株的末级小枝的
叶呈四棱状钻形。
关键词 南洋杉; 诺福克南洋杉; 分类修订