DSpace at VNU: Before it had a name: Diagnostic characteristics, geographic distribution, and the conservation of Cupressus tonkinensis (Cupressaceae)

Before it had a name: Diagnostic characteristics, geographic
distribution, and the conservation of Cupressus tonkinensis
(Cupressaceae)
DAMON P. LITTLE1, PHILIP THOMAS2, HIÊP T. NGUYỄN3,4,

AND

LÔC K. PHAN3,4,5

1

Lewis B. and Dorothy Cullman Program for Molecular Systematics, The New York Botanical
Garden, Bronx, NY 10458-5126, USA; e-mail:
2
Royal Botanic Garden, Edinburgh EH3 5LR, Scotland, UK; e-mail:
3
Institute of Ecology and Biological Resources, Vietnam Academy of Natural Science and
Technology, 18 Hoang Quoc Viet, Cau Giay, Ha Noi, Việt Nam; e-mail:
4
Center for Plant Conservation, Vietnam Union of Science and Technology Associations
(VUSTA), No. 25/32 Lane 191, Lạc Long Quan Rd, Nghia Do, Cau Giay, Ha Noi, Việt Nam
5
Hanoi University of Science, 334 Nguyen Trai, Thanh Xuan, Ha Noi, Việt Nam; e-mail:


Abstract. In northern Việt Nam the felling of Cupressus for wood and incense has
resulted in the decimation of wild cypress populations. Ever since specimens of Cupressus from Việt Nam were first sent to Paris in 1905, they have been inappropriately
referred to a variety of names including Cupressus funebris, Cu. torulosa, and synonyms
of Callitropsis lusitanica. This taxonomic disarray was somewhat ameliorated in 1994
when a new species—Cu. tonkinensis—was proposed. Unfortunately, the morphological
differences between Cu. tonkinensis and similar species were not clearly articulated. As a

result, Cu. tonkinensis has generally been ignored or treated as a synonym of Cu. torulosa. These two species can be distinguished by the presence of dimorphic leaves in Cu.
tonkinensis and their absence in Cu. torulosa as well as several continuous morphological
characteristics including the frequency of resin glands (greater in Cu. tonkinensis) on
ultimate segments, and the size of the ovulate cone (smaller in Cu. tonkinensis). The
natural habitat occupied by Cu. tonkinensis in a principal coordinates plot of nine
monthly climate variables and 40 soil variables is remote from other species. Cupressus
tonkinensis habitat is statistically distinct from Cu. chengiana, Cu. gigantea, and Cu.
torulosa, but statistically indistinguishable from the habitats of Cu. funebris.
Key Words: Callitropsis, conifers, Cupressus funebris, Cupressus tonkinensis,
Cupressus torulosa, Thông, Việt Nam, Vietnam.
Abstract.

Brittonia, 63(2), 2011, pp. 171–196
© 2010, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

ISSUED: 1 June 2011


172

BRITTONIA

The conifers of Việt Nam have been the
subject of considerable interest over the last
decade. The most recent taxonomic account
(Nguyễn & Vidal, 1996) described 27 indigenous species from 15 genera. Since then,
six new taxa, including three new species,
have been described (Businsky, 1999, 2004;
Farjon et al., 2002; Averyanov et al., 2004,
2008) and several other genera and species

have been recorded for the first time (Phan &
Nguyễn, 1997, 1999; Nguyễn et al., 2000a,
2002, 2004; Phan et al., 2002). Currently, 32
species from 19 genera are known.
Conifers are at risk throughout Việt Nam
from deforestation and over exploitation of
their valuable timber. A recent review of
conifer conservation status in Việt Nam
found that 90% of taxa were nationally
threatened, and 40% were globally threatened
(Nguyễn et al., 2004). Three species were
listed as nationally or globally Critically
Endangered, with a fourth from the genus
Cupressus L., listed as Data Deficient (Critically Endangered)—due to uncertainty about
its identification, native status, and global
distribution. Since its initial discovery, the
species of Vietnamese Cupressus have been
referred to a variety of names including
Cupressus funebris Endl., Cu. torulosa D.
Don, Cu. tonkinensis Silba, and synonyms of
Callitropsis lusitanica (Mill.) D. P. Little.
Resolution of this taxonomic uncertainty
would warrant a reassessment of Critically
Endangered status due to extremely high
levels of exploitation since the 1950s (Nguyễn
et al., 2004). An assessment of Extinct in the
Wild would make it the first conifer species to
be extirpated in modern times.
The first known collection of Vietnamese
Cupressus was made in 1905 under the

auspices of Souère Forestière du Tonkin
(Castellini 169, P) near Sông Hóa, Hoà Lạc
Lũng district, Lạng
commune,
province in the northern Việt Nam. The
specimen was sterile, with dimorphic ultimate

[VOL 63

segment leaves. The French botanist Philippe
Eberhardt collected a fertile specimen (Eberhardt 5073, NY; P; Fig. 1) of the same
species growing on calcareous substrate at
province. In 1913,
“Kai Kinsh” in Lạng
another French collector, Auguste Chevalier
(Chevalier 29662, P), also collected a
Cupressus from a place called “Cai–khinh”
(variously spelled “Kai Kung” and “Kai
King”; Chevalier, 1919, 1944; Hickel,
1930). In the early taxonomic accounts of
the conifers of Việt Nam (Chevalier, 1919;
Hickel, 1930) the Eberhardt and Chevalier
collection localities are treated as identical—
occurring at, or near, Kilometer 113 of the
railroad. Modern maps of Lạng
Lạng
province include a commune called “Cai
Lũng district. The center of
Kinh” in
Cai Kinh is approximately 110 km (by road)

from Hà Noi.
ˆ: Sông Hóa—the locality of the
Castellini collection—is within 8.5 km of Cai
Kinh. Chevalier’s later account (1944) indicates that his collection came from the
province.
village of “Quan Lan” in Lạng
“Quan Lan” certainly corresponds to Quang
Lang on modern maps, in Chi Lăng district,
province, about 20 km to the
Lạng
northeast of Sông Hoá.
Chevalier (1919) noted that the same
species of Cupressus was found in the karst
dominated province of Thái Nguyên—a
province that adjoins the southwest boundary
and the western boundary of
of Lạng
Giang. During our research we have been
unable to locate any Cupressus specimens
from Thái Nguyên.
Chevalier’s 1944 account stated that the
same species of Cupressus had been collected
Giang
at the village of “Bac–Lé” in
province by the “Société Forestière” (Souère
Forestière du Tonkin) and collected again
from the same locality in 1922 by Mignucci
(s.n.). Hickel (1930) simply listed the province as the locality and cited “Sté. forè no.
8507”. Modern maps of Việt Nam place “Băc

2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

173

FIG. 1. Cupressus tonkinensis: A. Holotype specimen (Eberhardt 5073, NY; image provided by the Digital
Imaging Laboratory at NY). B. Isotype specimen (Eberhardt 5073, P).

Lê” in Hòa
commune,
Lũng district,
province—less than nine kiloLạng
Giang
meters northwest of the Lạng
border. Chevalier (1944) also stated that he
had seen this species of Cupressus in other
localities in northern Việt Nam during his
travels, but did not collect specimens.
have
Recent wild collections from Lạng
much more precise collection localities. From
Chi Lăng district specimens have been colMạc commune (
39, HNU),
lected at
commune (Lo
:ˆ c P-2004, Ptownlet (
2336,
2005, HNU),

et al.
HNU), and Vạn Linh Commune (
1034, LE, MO). In addition, specimens have
Liên Nature Reserve
been collected from
Lũng (North Vietnam Second Darwin
in
Expedition [Thomas & Luu] 3, E; Nguyên et al.
HAL11918, E, HN).
According to Chevalier (1944), the specimens he collected and those collected by the

Souère Forestière du Tonkin were deposited
in the Muséum National d’Histoire Naturelle
in Paris (P). With the exception of the
specimen collected by Castellini, these specimens could not be found among the
collections of P and probably have been lost
(G. Aymonin, pers. comm.). Silba (1998)
indicates that a Cupressus specimen collected
by Chevalier (Chevalier s.n.) was found at P,
but we could not relocate it. The collection
made by Mignucci has not been relocated.
In total, there are nine wild Cupressus
collections known from Việt Nam. The localities are all within the rugged limestone karst
of the Cai Kinh massif on the eastern side of
River in
Lũng and Chi Lăng
province (Fig. 2). The
districts of Lạng
total area of occupancy is 8 km2 (using a 2×
2 km grid). The top ridges of the Cai Kinh

massif are ca. 300–550 m, rarely reaching
Lũng area—near where
650 m. In the


174

BRITTONIA

FIG. 2. Localities of Cu. tonkinensis in
Lũng
province, Việt Nam.
and Chi Lăng districts, Lạng
Grid squares are 2 × 2 km. The extent of natural
occurrence (204 km2) is outlined in grey. Filled stars
represent wild collections. Open stars represent cultivated
collections. This map is centered on the star in Fig. 3
(Albers projection).

the Eberhardt specimen was collected—
rainfall is ca. 1450 mm, with a five month
dry season from November to March (less
than 50 mm/month). The annual average
temperature is 22.8°C and the average
monthly temperature range is 15.3–28.5°C.
Three of the driest months are also amongst
the coldest—December through February
average less than 17°C (Nguyễn et al.,
2000b). Lower slopes are predominantly
evergreen broad-leaved forest. Wild Cupressus

is found on higher slopes, on and near ridge
tops—the vegetation there is closed, or
somewhat open, evergreen broad-leaved forest
dominated by Excentrodendron tonkinense (A.
Chev.) H. T. Chang & R. H. Miao (Tiliaceae;
Averyanov et al., 2003a, 2003b).
The sterile specimen collected by Castellini
was not identified by its collector, but sometime after its arrival at P (19 Jan 1907) it was
annotated (perhaps by Hickel) as Cupressus
torulosa—a species with monomorphic ultimate segment leaves. Eberhardt identified his
collections as Cu. funebris (Eberhardt 5073,
NY)—a species with dimorphic ultimate segment leaves. After the arrival of one of
Eberhardt’s specimens at P (Sep 1919), it
was annotated by Hickel as Cu. torulosa

[VOL 63

(Eberhardt 5073, P). Chevalier (1919) idencollections as Cu.
tified the Lạng
funebris in his annotated check–list of the
forest trees of Tonkin. Hickel’s (1930) annotated check–list of Indo–Chinese conifers
cited the specimens collected by Chevalier,
Lệ
Eberhardt, and the specimen from
collected by the Souère Forestière du Tonkin
as Cu. torulosa—an identification later adopted by Chevalier (1944). Mysteriously, the
collection made by Castellini was not cited by
either Chevalier (1919; 1944) or Hickel
(1930).
Later collections of wild Cupressus from

39, HNU; Lˆo
Việt Nam (
: c P-2004, P2336, HNU;
et al.
2005, HNU;
1034, LE, MO) were also identified as Cu.
torulosa—presumably using Hickel’s identifications as precedent. In the account of the
gymnosperms for the Flore du Cambodge, du
Laos et du Việt Nam (Nguyễn & Vidal,
1996), the collections made by Eberhardt,
et al. are also
Chevalier, Mignucci, and
cited under Cu. torulosa. Castellini’s collection was cited as Cu. funebris.
In addition to Cupressus torulosa, Cu.
funebris is also reported from Việt Nam. All
known specimens of Cu. funebris from Việt
Nam have been collected from cultivated
trees. Hickel (1930) cited a single specimen
of Cu. funebris (Capus s.n., P) and noted that
this species was considered extremely rare—
only occurring at higher elevation (ca.
1000 m) in Tonkin and the south of China.
Chevalier (1944) indicated that the Cu.
funebris specimens from Indo-China that he
had seen were cultivated and thereby
excluded Cu. funebris from his conifer treatment. Besides the Capus collection (precise
locality unknown), Nguyên & Vidal (1996)
listed two additional specimens of Cu. funebris (Castellini 169, P [later identified later as
Cu. tonkinensis]; P. K. Lôc s.n., LE [collected
Văn, Hà Giang

from a cultivated tree at
province]). Collections of apparently cultivated Cu. funebris have been made more
recently in the karst areas of Hà Giang in
northern–most Việt Nam (e.g., Luu s.n., E; P.
K. Loc s.n., LE; P. K. Loc et al. HAL8677,
HAL11342, HN, LE, MO; Averyanov et al.
HAL8479, HN, LE, MO). Almost all of these
collections are from small trees—probably


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

less than 20–30 years old. Cupressus funebris
is widely cultivated in southern and eastern
China and its historic distribution is unclear
(Farjon, 2005).
The literature relating to the habitat and
ecology of wild Cupressus in Việt Nam is
sparse and somewhat confused. In Nguyễn &
Vidal’s (1996) account, they state that it
province at altitudes
occurs in Lạng
between 250–1500 m on semi-arid limestone
hills and is locally associated with semideciduous forests that have a very short
period between leaf shedding and flushing
(i.e., almost evergreen). They also indicate
that Cu. torulosa is cultivated in communal
province (in southern

gardens in Lâm
Khôi 82, HN). Other
Việt Nam; Nguyễn
sources (Vu, 1996; Thái, 2000; Furey et al.,
2002; Nguyễn et al., 2004) indicate that wild
Cupressus is restricted to, and sparsely distributed along, limestone ridges in Lạng
and nearby provinces from 500–700 m.
True Cupressus torulosa is rarely cultivated in Việt Nam—to our knowledge it has
never been recorded in secondary vegetation.
In addition, we have not been able to locate
any specimens from areas adjacent to the Cai
province, from
Kinh massif of Lạng
other provinces of Việt Nam, or from China
that would indicate widespread cultivation or
naturalization. Chevalier (1944) clearly distinguished between cultivated and wild specimens and obviously believed that the
collections from Việt Nam he identified as
Cu. torulosa were of wild origin. The
application of Cupressus torulosa to Vietnamese Cupressus specimens from the Lạng
area has persisted despite the extreme
geographic and morphological disjunction
between the Himalayan (India, Nepal, and
Pakistan) and the supposed Vietnamese populations (compare squares versus the star in
Fig. 3).
In 1994, Silba described Cupressus tonkinensis, designating “Eberhardt 173” (NY) as
the holotype (Fig. 1A). Possibly because the
handwriting on the label is ambiguous, there
is confusion in the literature concerning the
collection number of the Eberhardt specimens: Hickel (1930), Chevalier (1944),
Farjon (2005), and Rushforth (2007) cite

“5073” whereas Silba cites either “173”

175

(Silba, 1994, 1998) or “1073” (Silba, 2005).
Silba (1994, 1998, 2005) identified additional
presumably wild specimens collected from
Guizhou China as Cu. tonkinensis (Cheng
2362, A; Steward et al., 10, A, L; Tsiang
8004, 5123, A) as well as cultivated specimens from Yunnan, China (Chu 51411,
51421, KUN, NY). The description was
self–published and is not widely available: it
was unavailable to Nguyễn & Vidal (1996)
when they were preparing their account. In
his description, Silba (1994) distinguished
between Cu. tonkinensis and Cu. chengiana
S. Y. Hu using the arrangement and relative
thickness of the branches. Silba also distinguished between Cu. tonkinensis and Cu.
lindleyi Klotzsch ex. Endl. (= Callitropsis
lusitanica) using the shape of the leaf apex.
Silba (2005) later treated Cu. tonkinensis as a
subspecies of Cu. funebris. Although Silba
did not clearly articulate the differences
between Cu. tonkinensis and Cu. torulosa,
other researchers (Rushforth et al., 2003;
Rushforth, 2007) noted the contrasting dimorphic leaves of Cu. tonkinensis and the
monomorphic leaves of Cu. torulosa.
Farjon (2005) dismissed Silba’s work and
synonymized Cupressus tonkinensis with Cu.
torulosa—perhaps because the dimorphic leaf

character was not obvious in the literature and
usually cannot be observed on herbarium
specimens without some dissection (dimorphic-leaved species are often mounted such
that all of the facial leaves are parallel to the
herbarium sheet while at the same time all of
the lateral leaves are perpendicular making
the contrast between the two difficult to
detect without some manipulation). Farjon
did not explain the discrepancy in leaf type
between the two species—simply describing
the leaves of Cu. torulosa s.l. (including Cu.
gigantea and Cu. tonkinensis) as monomorphic. Farjon explained the geographic disjunct as the result of human introduction of
Cu. torulosa at “an unknown time”, followed
by naturalization “on semi-arid slopes with
secondary vegetation, mainly on limestone”.
In his monograph, Farjon restricts Cu. torulosa to the dry inner valleys and semi-arid
high mountain environments of the eastern
Himalayas at altitudes between 1560–3670 m
where rainfall is not more than 300 mm; its


176

BRITTONIA

[VOL 63

FIG. 3. The distribution of wild Cupressus chengiana (triangles), Cu. funebris (circles), Cu. gigantea (diamonds),
Cu. tonkinensis (star; with arrow), and Cu. torulosa (squares). Cupressus jiangensis is known only from cultivation
and therefore excluded from this figure. Callitropsis lusitanica is native to the New World (distribution not shown;

Mercator projection).

presence in Yunnan is also attributed to
human introduction at an unknown time.
The distinctiveness of Cupressus tonkinensis from Cu. torulosa (and other species) is
strongly supported by an analysis of DNA
sequence data combined with anatomy, biochemistry, and morphology (Fig. 4; Little,
2006). Cupressus tonkinensis is included
within a clade of Asian species with dimorphic ultimate segment leaves (Cu. chengiana,
Cu. funebris, and Cu. jiangensis N. Zhao)
while Cu. torulosa is nested in a clade of
Asian species with monomorphic (Cu.
duclouxiana Hickel in A. Camus and Cu.
gigantea W. C. Cheng & L. K. Fu) or weakly
dimorphic ultimate segment leaves (Cu. austrotibetica Silba and Cu. cashmeriana Royle
ex Carrière). The clade containing Cu.
tonkinensis is sister to the clade containing
Cu. torulosa plus a clade of Eurasian and
north African species with weakly dimorphic
ultimate segment leaves (Cu. atlantica Gaussen,
Cu. dupreziana A. Camus, and Cu. sempervirens L.).
Regardless of its identity, the Cupressus in
Việt Nam has always been valued for its fine
timber and resin; consequently it has always
been heavily exploited. One indication of this
can be seen in the extraordinary cost of a cubic
,
meter of wood: in 1987 it was 3 million
by 1997 it had increased 10 times to 30 million

. During the same period other high value
timbers and resins such as Fokienia hodginsii
A. Henry & H. H. Thomas (locally known as
to 5 million
“Pomu”) went from 700,000
per cubic meter. After 1997, no commercial quantities of Cupressus were available and
it virtually disappeared from the market (Bay,
2003). Restrictions on its harvest were introduced as early as 1958 (Thái, 2000), but little
was done to stop the exploitation. Between
1986 and 1989 a 10,000 ha nature reserve was
Liên to protect populations
established in
of musk deer, limestone forests, and the
remaining populations of Cu. “torulosa”
(Furey et al., 2002). Cupressus “torulosa”
was listed in the Group IA protected species
under Vietnamese law: this prohibited any
further exploitation or use (Anonymous,
2006). It was listed in Việt Nam’s Red Data
book as Critically Endangered (CR A1a,d;
Anonymous, 2007). In the late 1990s a small
plantation was established by the Forest SciLiên
ence Institute at the edge of the
Reserve. By 2004, what was thought to be the
last known natural tree in the reserve was
destroyed in a fire (Nguyễn et al., 2004). More
recently (October 2008), a single tree was
located in another part of the massif. Apart
from this, no other wild Cupressus trees are
known to occur in Việt Nam. This species is

also represented by about 20 trees remaining in


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

FIG. 4. A phylogenetic hypothesis for the genus
Cupressus (sensu Little, 2006) based on data from
anatomy, biochemistry, morphology, and DNA sequence
(matK, NEEDLY intron 2, nrITS, rbcL, and trnL) redrawn
from Little (2006). Numbers below branches indicate
jackknife values. The number of terminals contained
within each condensed clade is indicated at the clade
base.

the plantation, several cultivated trees growing
in the surrounding villages, and a clonal
collection held by organizations within the
Ministry of Agriculture and Forestry. The
origin, and the identity, of some of these trees
are uncertain (Nguyễn & Thomas, 2004;
Nguyễn et al., 2004).
This paper strives to clearly articulate and
test the morphological and ecological differences between Cupressus tonkinensis and the
species that could potentially be confused
with it. This information should help to
identify additional wild plants, should they
be found, and also identify those held in ex
situ collections.

Methods and materials
Published climate and soil data were used
in combination with georeferenced herbarium
specimens to test for ecological differences
between Cupressus tonkinensis and related
species. Continuous and discrete variation in
morphological characteristics was used to test
for morphological differentiation.
The species sampled include those found in
the least inclusive clade containing Cupressus
tonkinensis (Fig. 4) and the species that have
historically been confused with Cu. tonkinensis or associated taxonomically (Callitropsis

177

lusitanica, Cu. chengiana, Cu. funebris, Cu.
gigantea, Cu. jiangensis, Cu. tonkinensis, and
Cu. torulosa). A complete list of specimens
examined can be found in Appendix 1.
If available, label coordinates from herbarium
specimens were used directly. Specimens without coordinate information were georeferenced
using data downloaded from GNS (http://
gnswww.nga.mil/geonames/GNS/index.jsp).
Georeferencing used the most precise locality
provided on the label that could be found in the
databases. All specimens were considered wild
unless the label explicitly indicated otherwise.
Maps were constructed using GMT version
4.1.1 (Wessel & Smith, 2006).

For wild specimen localities in Asia, nine
monthly climate variables (cloud cover, diurnal temperature range, minimum temperature,
mean temperature, maximum temperature,
ground-frost frequency, precipitation, vapor
pressure, and wet day frequency) were
extracted from the 10-year monthly-mean
(1981–1990) data set of New et al. (2000;
downloaded from ).
As published, the climate data are interpolated and have a gridded resolution of 0.5°.
Forty soil variables (for each soil profile: soil
depth to a physically limiting layer and
available water capacity; separately for topsoil and subsoil: aluminum saturation, base
saturation, total extractable bases, cation
exchange capacity, cation exchange capacity
of the clay fraction, exchangeable sodium, pH
in water, electrical conductivity of a saturated
paste, bulk density, calcium carbonate content, gypsum content, organic carbon, total
nitrogen, organic carbon/nitrogen ratio, clay
content, sand content, silt content, gravel
content, and total porosity) were extracted
for wild specimen localities from the
unbanned data set of Batjes (2005; downloaded from ). As downloaded, the 0.5° gridded unbinned soil dataset
included the variation among soil units within
each grid cell—this variation was reduced to
a single value per variable per grid cell by
calculating a mean weighted by the relative
area within the grid cell occupied by each soil
unit. Because the number of herbarium specimens from a particular region does not
necessarily reflect population size, the data
were filtered for each species prior to analysis

to remove duplicate grid squares.


178

[VOL 63

BRITTONIA

For scanning electron microscopy, field-collected specimens were fixed in FAA (50% or
70% ethanol, 5% acetic acid, and 5% formalin).
After a minimum of 48 h of fixation, samples
were transferred to a solution of 50% or 70%
ethanol and 10% glycerin for storage. Samples
were dehydrated in an ethanol series (70%,
95%, 100%, 100%, 100%) and then transferred
to a 1:1 ethanol:acetone solution. Expulsion of
ethanol was accomplished with three changes of
pure acetone. Specimens were transitioned from
acetone to liquid carbon dioxide and criticalpoint dried in a Denton DCP-1 (Denton
Vacuum, Moorestown, NJ). Specimens were
mounted on aluminum stubs with carbon tape
and sputter coated with gold/palladium for 4
minutes using a Hummer 6.2 sputtering system
(Anatech, Union City, CA). Observations were
made with a JSM-5410LV (JEOL, Tokyo) SEM
using an accelerating voltage of 15 kV.
Measurements of continuously distributed
morphological characteristics and scores for
discrete characters were obtained from herbarium specimens. Characters and scoring techniques are detailed in Appendix 2. We

attempted to gather three measurements for
each continuous characteristic from each specimen examined, but the amount of suitable
material often limited the number of measurements. Prior to statistical analysis intra-individual continuous characteristic variation was
eliminated by using the median value for each
characteristic from each specimen.
Statistical tests and visualizations were
conducted with R version 2.6.2 (R development core team, 2008) and the R packages
LabDSV version 1.3-1 (Roberts, 2008),
pgirmess version 1.3.6 (Giraudoux, 2008),
and vegan version 1.11-4 (Oksanen et al.,
2008). The variation in the climate and soil
data was visualized by principal coordinates
analysis (Gower, 1966) of Manhattan distances calculated from mean-centered and
root-mean-squared scaled data (scale, dist,
and pco functions). A non-parametric multiple-comparison test based on the KruskalWallis (Kruskal & Wallis, 1952; Siegel &
Castellan, 1988) rank sum test (kruskalmc
function) was used to determine if the
species were distinct at the p =0.05 level for
each of the 148 soil and climate variables.
The output was corrected for multiple tests
using the method of Benjamini and Hochberg

(1995; p.adjust function). The variation in
continuous morphological characteristics was
analyzed using the multiple-comparisons
method described above.
Results
For the nine monthly climate variables and
40 soil variables, the two principal coordinates axes together account for 71.5% of the
variation in the original data—PCO 0

accounts for 61.7% and PCO 1 accounts for
9.8% (Fig. 5). The habitat occupied by
Cupressus tonkinensis in the principal coordinates plot is remote from that of Cu.
torulosa (Fig. 5). The area between Cu.
tonkinensis and Cu. torulosa is filled by
habitats occupied by Cu. chengiana, Cu.
funebris, and Cu. gigantea. Cupressus
funebris is geographically widespread and
occupies a wide range of habitats—as a result
it approaches or envelopes every other species
represented in the plot. On a per character
basis, the habitats of Cu. funebris are statistically distinct from those occupied by all other
species except Cu. tonkinensis (Table I).
The habitat of Cu. tonkinensis is statistically
distinct from that of Cu. chengiana, Cu.
torulosa, and Cu. gigantea. The habitats of
Cu. tonkinensis and Cu. torulosa differ statistically for 71 of the 148 soil and climate
variables. Gradients of climate and soil variation generally run parallel to PCO 1. Among
the top 25% best fitting variables
(D2 =0.9570–0.9901) all but three (subsoil
pH in water, subsoil electrical conductivity of
a saturated paste, and subsoil cation exchange
capacity of the clay fraction; Fig. 5D) are
climate variables—mostly relating to the
weather in the months of May through
October (64.8% of the top ranked variables).
The top 25% best fitting variables include
minimum temperature for all twelve months
(e.g., Fig. 5A), nine months worth of vapor
pressure variables (e.g., Fig. 5B), and seven

months worth of mean temperature variables.
Discrete morphological characters can be
used to distinguish Cupressus tonkinensis
from Callitropsis lusitanica, Cu. funebris, Cu.
gigantea, Cu. jiangensis, and Cu. torulosa
(Tables I and II). Of particular interest are the
distribution of dimorphic leaves (Figs. 6–8;
Table II, characters 34 and 37) and mucronate


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

179

FIG. 5. Principal coordinates analysis of nine monthly climate variables and forty soil variables for wild specimen
localities for Cupressus chengiana (triangles), Cu. funebris (circles), Cu. gigantea (diamonds), Cu. tonkinensis (stars),
and Cu. torulosa (squares). Wild populations of Cu. jiangensis are unknown and no native Asian populations of
Callitropsis lusitanica are known therefore these species were excluded from this analysis. Variation in four arbitrary
selected variables are shown as fitted surfaces. A. Minimum September temperature (D2 =0.9894). B. September vapor
pressure (D2 =0.9786). C. September ground frost frequency (D2 =0.9581). D. Subsoil pH in water (D2 =0.9703).

leaf apices (Fig. 8; Table II, character 39). In
addition, continuous morphological variation
can be used to statistically distinguish Cu.
tonkinensis from Cu. chengiana, Cu. funebris,
Cu. gigantea, and Cu. torulosa (Tables I and III,
characters 2, 5, 10, 11, 15, 23, 24, and 25).
Across all taxa, two of the five most statistically

discriminatory characteristics (Table III) are
vegetative: ultimate segment length (character
10) and ultimate segment leaf length (character
14). The remaining three most discriminatory

characters depict features of the ovulate cone:
length (character 23), width (character 24), and
pairs of scales (character 25).
Discussion
Historically Cupressus tonkinensis has been
confused with Cu. torulosa (Hickel, 1930;
Chevalier, 1944; Nguyễn & Vidal, 1996;
Farjon, 2005) despite the extreme geographic
separation (Fig. 3) and marked difference in


180

[VOL 63

BRITTONIA

TABLE I
PERCENTAGE OF PAIR-WISE NON-PARAMETRIC TESTS RESULTING IN SIGNIFICANT DIFFERENCES AT P=0.05 (CORRECTED FOR
MULTIPLE–COMPARISONS AND TESTS) FOR CLIMATE AND SOIL VARIABLES (BELOW THE DIAGONAL) AND CONTINUOUS
MORPHOLOGICAL CHARACTERS (ABOVE THE DIAGONAL) WHEN APPLICABLE DIFFERING DISCRETE MORPHOLOGICAL CHARACTERS ARE GIVEN PARENTHETICALLY. WILD POPULATIONS OF CUPRESSUS JIANGENSIS ARE UNKNOWN AND NO NATIVE ASIAN
POPULATIONS OF CALLITROPSIS LUSITANICA ARE KNOWN, THEREFORE STATISTICAL TESTS OF CLIMATE AND SOIL VARIABLES ARE
NOT APPLICABLE (NA) FOR THESE SPECIES.

Ca. lusitanica

Cu. chengiana

Cu. funebris

Ca. lusitanica

—

0% (34)

Cu. chengiana
Cu. funebris

NA
NA

—
77%

Cu. gigantea

NA

3%

25% (32, 23% (36, 38,
34, 37)
44)
13% (39) 0% (34)

—
0% (32, 34,
35, 36, 37, 39,
43, 44, 45)
74%
—

Cu. jiangensis

NA

NA

NA

NA

55%
17%

0%
80%

55%
5%

Cu. tonkinensis NA
Cu. torulosa
NA

Cu. gigantea

habitat preference (Fig. 5; Table I). However,
these species are easily distinguished by the
presence of dimorphic leaves in Cu. tonkinensis
and their absence in Cu. torulosa (Tables I and
II, characters 34 and 37; Figs. 6E–G, 7E–G) as
well as many continuous characteristics including the frequency of resin glands on ultimate
segments (Table III, character 15) and the size
of the ovulate cone (Table III, characters 23, 24,
and 25).
Cultivated Callitropsis lusitanica has
also been confused with Cupressus tonkinensis. The presence of dimorphic leaves in
Cu. tonkinensis is the most diagnostic
characteristic (Figs. 6E–F, 7H, and 8H;
characters 34 and 37), but penultimate
branches arranged on a single plane (character 32) and obtuse leaf apices (character

Cu. jiangensis

Cu. tonkinensis

0% (34,
37, 38)
0% (39)
0%

0% (32, 34,
37, 38)
19%

4% (39, 41)

Cu. torulosa

4% (36, 38)
13% (34)
17% (34, 36,
37, 39)

0% (34, 35, 10% (32, 34, 14%
36, 37, 39)
35, 37)
—
0% (39)
0% (34, 36,
37, 39)
NA
—
24% (34, 37)
NA
48%
—

38; Fig. 8F and 8D; Silba, 1994) are also
useful (Table II).
Silba (1994) used stem diameter (characters
2 and 11) to distinguish between Cupressus
tonkinensis and Cu. chengiana—the differences in these characters are statistically significant (Table III). Additional significant
continuous characters include ultimate branch
segment length (Table III, character 10), the

length of the ovulate cone (Table III, character
23), and the number of ovulate cone scales
(Table III, character 25).
Cupressus tonkinensis is perhaps most
easily confused with Cu. funebris. These
two species can be distinguished by the
absence of a mucronate tip on the apex of
the ultimate segment lateral leaves in Cu.
tonkinensis (Fig. 8F) and the presence of such

TABLE II
CHARACTER DESCRIPTIONS AND STATES ARE GIVEN IN APPENDIX 2.
POLYMORPHISMS ARE ENCLOSED IN BRACKETS. VARIATION IN ULTIMATE SEGMENT LEAF FORM (CHARACTER 37) IS
ILLUSTRATED IN FIGS. 6, 7, AND 8. VARIATION IN ULTIMATE SEGMENT LEAF APICES (CHARACTERS 38 AND 39) IS ILLUSTRATED
IN FIG. 8.
SCORES FOR DISCRETE MORPHOLOGICAL CHARACTERS.

character:

Ca.
Cu.
Cu.
Cu.
Cu.
Cu.
Cu.

lusitanica
chengiana
funebris

gigantea
jiangeensis
tonkinensis
torulosa

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

2
2
[12]
2
2
[12]
[12]

2
[12]
1
2
[12]
1
[12]

1
[12]
1
[12]
1
1
[12]

1

2
2
1
2
2
1

[12]
[12]
1
2
1
1
[12]

0
[01]
0
1
0
[01]
1

1
[12]
2
1
2
2
1

1
[01]
[01]
0
0
0
0

[01]
0
1
0
1
0
0

—
[01]
0
—
0
0
—

—
[01]
1
—
[01]

0
—

[01]
1
[01]
1
1
[01]
[01]

[01]
[01]
0
1
[01]
?
[01]

1
[01]
1
0
?
?
[01]

[01]
[01]
1

0
[01]
[01]
[01]


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

181

FIG. 6. Ultimate segments with attached leaves. A, B. Cupressus funebris (Little et al. 862, BH, CDBI). A. Facial
leaves. B. Lateral leaves. C, D. Cupressus gigantean (anon. 2002 Sep 11, BH). C. Leaves. D. Rotated 90° from that
shown in C—indicating that the leaves are not dimorphic. E, F . Cupressus tonkinensis (NY-HN 110, HN, NY). E.
Facial leaves. F. Lateral leaves. G, H. Cupressus torulosa (Little 632, BH, TUCH). G. Leaves. H. Rotated 90° from
that shown in G—indicating that the leaves are not dimorphic. I, J. Cupressus jiangensis (Little & Sun 869, BH,
CDBI). I. Facial leaves. J. Lateral leaves. K, L. Cupressus chengiana (Little et al. 862, BH, CDBI). K. Facial leaves.
L. Lateral leaves.

a tip on the ultimate segment lateral leaves of
Cu. funebris (Fig. 8B).
Although the habitat occupied by Cupressus tonkinensis is distant from the habitats

occupied by other species in the principal
coordinates plot (Fig. 5), the habitat is not
statistically distinct in all cases (Table I). All
of the verified Cu. tonkinensis collections

182

BRITTONIA

[VOL 63

FIG. 7. A–G, I–L. Ultimate segment leaf margins at the mid point of the leaf, adaxial view. A, B. Cupressus
funebris (Little & Chen 847, BH, KUN). A. Facial leaf. B. Lateral leaf. C. Cupressus gigantea, leaf (anon. 2002 Sep
11, BH). D. Callitropsis lusitanica, leaf (Little & Ochoterena 795, BH, MEXU). E, F. Cupressus tonkinensis (NY-HN
110, HN, NY). E. Facial leaf. F. Lateral leaf. G. Cupressus torulosa, leaf (Little 632, BH, TUCH). H. Callitropsis
lusitanica, an ultimate segment with attached leaves (Little & Ochoterena 795, BH, MEXU). I, J. Cupressus
jiangensis (Little & Sun 869, BH, CDBI). I. Facial leaf. J. Lateral leaf. K, L. Cupressus chengiana (Little et al. 862,
BH, CDBI). K. Facial leaf. L. Lateral leaf.

are from an area less than 0.5° square,
therefore Cu. tonkinensis is represented in
the habitat analysis by just two data points
whereas other species are represented by 5–

85 data points each. The Kruskal-Wallis type
test (Kruskal & Wallis, 1952; Siegel &
Castellan, 1988) used for the analysis is
robust to unequal variance and sample size,


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

183

FIG. 8. A–G, I–L. Ultimate segment leaf margins and apices, adaxial view. A, B. Cupressus funebris (Little &
Chen 847, BH, KUN). A. Facial leaf. B. Lateral leaf. C. Cupressus gigantean, leaf (anon. 2002 Sep 11, BH). D.
Callitropsis lusitanica, leaf (Little & Ochoterena 795, BH, MEXU). E, F. Cupressus tonkinensis (NY-HN 110, HN,
NY). E. Facial leaf. F. Lateral leaf. G. Cupressus torulosa, leaf (Little 632, BH, TUCH). H. Callitropsis lusitanica, an
ultimate segment with attached leaves, rotated 90° from that shown in Fig. 7H—indicating that the leaves are not
dimorphic (Little & Ochoterena 795, BH, MEXU). I, J. Cupressus jiangensis (Little & Sun 869, BH, CDBI). I. Facial
leaf. J. Lateral leaf. K, L. Cupressus chengiana (Little et al. 862, BH, CDBI). K. Facial leaf. L. Lateral leaf.

but results obtained using such small sample
sizes should be treated with caution. Increasing the sample size for Cu. tonkinensis in the
habitat analysis would likely increase the

number of significant comparisons, but this
is not possible given the biological reality—
Cu. tonkinensis is a narrow endemic
whereas the other species range from wide-


mm

mm

%
mm

%

%
mm

%
%
mm

mm

%

%

mm

%

%
mm

%
%
mm

mm

2

3
4

5

6
7

8
9
10

11

12

13

14

15

16
17

18
19
20

21

units

1

character

(6.1)7.8–17.4–
29.9(32.1)
(0.7)0.8–1.0–
1.2(1.3)tr
(3)4–13–32(39)
(0.9)1–1.1–1.8(2.1)

Cu. chengiana

(10.6)10.6–
19.8–39(58.5)
(0.9)0.9–1.2–
1.4(1.5)
(9)9–22–42(43)l
(1)1–1.4–2.2(2.3)

Cu. funebris

(8.2)8.2–18.4–
71.7(71.7)
(0.9)0.9–1.1–
1.5(1.5)
(2)2–6–23(23)
(0.9)0.9–1.3–
1.6(1.6)
(8)8–69–78(78)

Cu. gigantea

Cu. jiangeensis

(14.8)14.8–21.9–
27.7(27.7)
(0.8)0.8–0.9–1
(1)
(8)8–11–18(18)
(1.1)1.1–1.1–
1.4(1.4)
(27)29–78–
(0)31–76–93(93)
(57)57–64–
100(100)
100(100)
(0)0–0–41(55)
(0)0–0–29(31)
(0)0–0–0(0)
(0)0–0–0(0)
(0)0–0–0(0)
—
(0.9)1–1.2–1.9(2.2) (1.0)1.1–1.5–
—
(1.1)1.1–1.2–
2.3(2.3)
1.4(1.4)
—
(0)0–49–100(100) (0)0–18–42(44)
—

(0)0–0–45(45)
—
(0)0–0–0(13)
(0)0–0–0(0)
—
(0)0–0–0(0)
(4.5)4.7–8.2–
(4.2)4.4–
(6.1)6.1–11.9–
(6.7)6.7–8.4–
(4.1)4.2–7.2–
fg
t
l
l
13.3(15.0)
6.6–10.8(13.1)
23.4(28.4)
18.1(18.1)
14.4(19.5)
( 0 . 7 ) 0 . 7– 0 . 9 – 1 . 1 ( 0 . 7 ) 0 . 7 – 1 – 1 . 2 ( 0 . 9 ) 1 – 1 . 2 – 1 . 6 ( 0 . 8 ) 0 . 8 – 1 – 1 . 1 (0.8)0.8–0.9–
(1.6)c
(1.2)
1(1.0)
(1.5)
(1.4)ft
( 5 0 ) 5 0 – 7 1 – 1 0 0 (80)83–100–
(100)100–
(71)71–100–
(100)100–100–

(100)
100(100)
100–100(100)
100(100)
100(100)
( 3 3 ) 3 3 – 5 7 – 1 0 0 (25)33–65–
(50)50–67–
(0)0–78–100(100) ( 6 0 ) 6 0 – 6 7 – 7 5
(100)
100(100)
100(100)
(75)
(0.8)0.8–1–
(0.7)0.8–1.1–
(0.8)0.8–0.8–
(0.8)0.8–1–1.3(1.3) (0.7)0.8–1–
r
r
r
1.3(1.4)
1.4(2.5)
1.1(1.1)
1.2(1.3)
(0)0–54–93(100)
(0)0–76–100(100) (29)40–80–
(0)0–47–91(100) (0)0–50–70(70)
100(100)
(0)0–0–0(33)
(0)0–0–75(91)
(0)0–0–0(0)

(0)0–0–0(0)
(0)0–0–0(0)
—
(0.7)0.9–1.0–
(0.8)0.8–1.1–
—
(0.8)0.8–0.9–
1.4(1.5)
1.4(1.4)
1.1(1.1)
—
(0)0–0–0(0)
—
(0)0–64–100(100)f (0)0–0–25(40)c
—
(0)0–0–20(57)
(0)0–0–0(0)
—
(0)0–0–0(0)
( 2 . 6 ) 2 . 6 – 3 . 6 – 4 . 3 (1.7)2.1–2.8–
(3.3)3.3–3.4–
unknown
(2.6)2.6–3–
(4.5)
3.4(4.2)
3.5(3.5)
3.5(3.5)
(1.6)1.6–1.7–
unknown
(1)1–1.3–1.6

(1.6)1.6–2–2.3(2.3) (1.0)1.1–1.4–
1.9(1.9)
(1.6)
1.8(2)r

(8.6)9–20.4–30.1
(34)
(0.7)0.8–1.1–1.4
(1.5)
(8)9–19–30(31)f
(1.1)1.1–1.4–1.7
(1.9)
(0)0–67–95(96)

Ca. lusitanica

CU. JIANGENSIS, L = CALLITROPSIS LUSITANICA, T = CU. TONKINENSIS, AND R = CU. TORULOSA.

—
—
(1.8)1.8–2.6–
5.2(5.2)
(1.2)1.2–1.6–
2.1(2.1)c

(0)0–0–0(0)
—

—
—

(5.1)5.5–9.9–
17.6(19.2)
(0.6)0.7–0.9–
1.1(1.3)
(50)62–100–
100(100)
(0)25–60–
100(100)
(0.7)0.8–0.9–
1.1(1.2)cfg
(0)0–0–78(82)t

(0)0–0–0(0)
—

(10.3)11.7–20–
27.4(27.6)
(0.8)0.9–1–
1.4(1.4)c
(3)3–12–20(22)
(0.5)0.9–1.2–
1.5(1.6)
(0)0–36–82(88)t

Cu. torulosa

BRITTONIA

unknown

(38)44–78–
100(100)r
(0)0–0–0(47)
(0.7)0.7–0.9–
1.2(1.3)
(0)0–25–100(100)
(0)0–0–17(17)
unknown

(44)48–87–
96(100)r
(0)0–0–0(18)
(0.9)0.9–1.2–
1.6(1.6)
(0)0–17–100(100)
(0)0–0–0(0)
(3.2)4.2–7–
12.3(13.5)c
(0.6)0.8–1.0–
1.2(1.2)c
(54)54–100–
100(100)
(33)33–67–
100(100)
(0.6)0.6–0.8–1(1)

(8)11.2–17.7–
31.3(32)
(0.9)0.9–1.1–
1.3(1.3)c

(3)4–11–23(28)
(0.5)0.8–1–1.4(1.6)

Cu. tonkinensis

TABLE III
VARIATION IN CONTINUOUS MORPHOLOGICAL CHARACTERS. DATA ARE GIVEN IN THE FORM OF (MINIMUM)5TH PERCENTILE–MEDIAN–95TH PERCENTILE(MAXIMUM). SUPERSCRIPT LETTERS
INDICATE STATISTICALLY SIGNIFICANT DIFFERENCES AT P =0.05 (CORRECTED FOR MULTIPLE–COMPARISONS AND TESTS): C = CUPRESSUS CHENGIANA, F = CU. FUNEBRIS, G = CU. GIGANTEA, J =

184
[VOL 63


LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)
(7.5)8.6–14.3–
16.8(18.0)fgt
(9.2)9.6–13.6–
16.1(16.4)fgt
(3)4–5–6(6)flt
(2.2)2.9–3.6–
4.4(4.9)
(1.8)2–3.0–
4.5(4.7)
(0.8)0.8–1–
1.3(1.4)
(0.3)0.5–0.9–
1.4(1.6)
(0.1)0.2–0.4–
0.7(0.7)

(4)4–6–11(11)

6 [only one cone,
not used for
statistical
analysis]
(15.1)15.1–15.3– (6.7)6.7–8–
15.9(15.9)
9.5(9.5)cgr
(12.1)12.1–13.4– (7.5)7.5–9.1–
13.9(13.9)
10.1(10.1)r
(5)5–6–6(6)
(3)3–3–4(4)cfgr
(3.5)3.5–3.8–
(3.0)3.0–3.4–
4.2(4.2)
3.7(3.7)
(2.3)2.3–3.2–
(1.8)1.8–3.1–
4.1(4.1)
3.7(3.7)
(0.8)0.8–1–
(0.7)0.7–1.1–
1.3(1.3)
1.4(1.4)
(0.7)0.7–0.8–
(0.6)0.6–0.9–
1.0(1)
1.1(1.1)

(0.3)0.3–0.3–
(0.3)0.3–0.4–
0.5(0.5)
0.5(0.5)
(13.2)14.4–16.5–
21.8(28.5)ltr
(12.1)12.8–15.6–
18.3(19.1)lr
(4)4–5–7(7)lt
(3.5)3.5–4.1–
4.4(4.4)l
(2.2)2.2–3.0–
3.6(3.6)
(0.6)0.6–0.9–
1.2(1.2)
(0.2)0.2–0.4–
1.0(1)
(0.1)0.1–0.2–
0.4(0.4)
(5.4)6.3–8.9–
11.9(12.8)cr
(6.1)6.2–9.5–
12.0(12.1)cr
(3)3–3–4(4)ltr
(2.5)2.6–3.3–
3.5(4.0)
(2.0)2.3–2.7–
3.8(4.5)
(0.6)0.6–1.1–
1.3(1.3)l

(0.3)0.4–0.7–
0.9(1)l
(0.1)0.2–0.4–
0.6(0.7)l
mm

mm

pairs
mm

mm

mm

mm

mm

23

24

25
26

27

28

29

30

(9.9)10.9–13.3–
16.8(19.2)g
(8.3)9.6–13.6–
19.4(19.8)g
(2)3–3–4(4)fgr
(2.4)2.8–3.9–
4.8(4.9)g
(2.3)2.5–3.1–
4.0(4.2)
(0.7)0.7–1.1–
1.5(1.5)f
(0.5)0.7–1.1–
1.6(1.7)f
(0.3)0.4–0.5–
0.8(1.4)f

(10.7)11.6–15.4–
18.7(18.8)ft
(10.2)10.5–14.1–
17.0(18.4)f
(4)4–4–5(6)t
(2.4)2.4–3.6–
4.5(4.5)
(2.2)2.2–3.3–
4.5(4.5)
(0.8)0.8–1–

1.4(1.4)
(0.8)0.8–1–
1.2(1.2)
(0.2)0.2–0.3–
0.5(0.5)

(7)7–8–10(10)
(4)4–6–8(8)
(6)6–7–8(8)
(5)5–7–8(8)
(5)5–7–8(9)
pairs
22

Table III Continued

Cu. gigantea
units
character

Ca. lusitanica

Cu. chengiana

Cu. funebris

Cu. jiangeensis

Cu. tonkinensis

Cu. torulosa

2011]

185

spread endemics (e.g., Cu. gigantea) to
broadly distributed generalists (e.g., Cu.
funebris).
Although additional Cupressus tonkinensis data points could be manufactured by the
use of a higher resolution soil and climate
data set (e.g., Hijmans et al., 2005), the error
inherent in georeferenceing historic specimens with imprecise localities would be
magnified—rendering the resulting analysis
even more suspect. The use of individual
herbarium specimens or collection events as
terminals rather than grid squares in habitat
analysis would also increase sample size for
all species, but the relationship between
population size and number of collection
events or herbarium specimens is suspect at
best.
Taxonomic treatment
Cupressus tonkinensis Silba, J. Int. Conifer
Preserv. Soc. 1: 23. 1994. Type: Việt Nam.
province,
Lũng district, Cai
Lạng
Kinh Commune [“Kai Kinsh, Lausoy”],
21.55°N, 106.40°E, Dec 1913, Eberhardt

5073 (holotype: NY; isotype: P).
Cupressus funebris Endl. subsp. tonkinensis Silba, J. Int.
Conifer Preserv. Soc. 12: 67. 2005.

Trunk solitary; 10 m or more tall, DBH
40 cm or more. Outer bark on the main trunk;
grey-brown; exfoliated in linear strips
(Fig. 9A). Antepenultimate branch segments
arranged on one plane or two planes. Penultimate branch segments (8)12.3–30.7(32) mm
long; (0.9)1–1.3 mm in diam.; penultimate
branchlets arranged on one plane; (3)4–22
(28)% of the nodes bearing branches; branchlets strictly one per node; leaves externally
dimorphic. Adult penultimate segment facial
leaves (0.5)0.8–1.4(1.6) mm long; (44)56–95
(100)% of the leaves have resin glands; 0
(18)% of the resin glands produce resinous
exudates. Adult penultimate segment lateral
leaves 0.9–1.5(1.6) mm long; (0)5–86(100)%
of the leaves with resin glands; none of the
resin glands produce resinous exudates. Ultimate branch segments (3.2)4.3–9.5(13.5) mm
long; (0.6)0.8–1.1(1.2) mm in diam.; branchlets arranged on one plane; (54)100% of
ultimate segment branchlets on the primary
plane; (33)50–100% of the ultimate branch-


186

BRITTONIA

[VOL 63

FIG. 9. Cupressus tonkinensis. Mature cultivated specimens (A–C: NY-HN 115, NY, HN; D–H: Phan et al.
HAL11,344, HN). A. Trunk of a mature tree with grey–brown bark exfoliating in linear strips. B. Ultimate branch
segments with characteristic glossy green appearance. C. Penultimate branch segments arranged on a single plane. A
majority of the ultimate segments point towards the apex of the antepenultimate segment that ultimately bears them.
D. Pollen cone just past time of pollen shed. E. An ovulate cone at the time of pollination. F. Ovulate cones after
pollination, but before seed maturation. G. Mature (second year) ovulate cones with mature seeds. H. Mature seeds.

lets point towards the apex of the antepenultimate segment that ultimately bears them
(Fig. 9C); juvenile (needle–like) leaves produced only by young individuals; stem

rectangular or sometimes isodiametric; leaves
externally dimorphic, glossy green (Figs. 6E–
F, 9B). Adult ultimate segment facial leaves
(0.6)0.7–1 mm long; (38)50–94(100)% of the


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

leaves have resin glands; 0(47)% of the resin
glands produce resinous exudates; dorsal
resin glands with oblong (slot-like) external
faces; marginal leaf band extending from the
leaf margin, denticulate, extending across the
leaf apex; leaf apex acute, unadorned
(Figs. 6E, 7E, and 8E). Adult ultimate segment lateral leaves (0.7)0.8–1.1(1.3) mm
long; 0–80(100)% of the leaves with resin
glands; 0(17)% of the resin glands produce

resinous exudates; dorsal resin glands with
oblong (slot–like) external faces; marginal
leaf band extending from the leaf margin,
denticulate, extending or not extending across
the leaf apex; leaf apex acute, unadorned
(Figs. 6F, 7F, and 8F). Pollen cone oblong,
with 6 pairs of opposite decussate cone scales
(Fig. 9D). Ovulate cone spheric (6.7)7.1–9.1
(9.5) mm long; (7.5)7.8–10 (10.1) mm wide;
3(4) pairs of opposite decussate cone scales;
mucro on the boss of apical cone scales entire
or ciliate; peduncle leaves externally monomorphic; cones non–serotinous, the scales
opening and dying upon seed maturation
(Fig. 9E–G). Seeds irregular in shape; 3–
3.7 mm long; 1.8–3.5(3.7) mm wide; 0.7–
1.4 mm thick; seed coat not glaucous, dull
red–brown; seed wings well–developed
(Fig. 9G–H); 2 years required for seed
maturation. Hilum 0.6–1.1 mm long; 0.3–
0.5 mm wide (Fig. 9H). Cotyledons
unknown. Chromosome number unknown.
Current distribution.—Known from northeastern Việt Nam; Lạng Son province; Cai
Lũng and Chi
Kinh limestone massif;
Lăng districts (Fig. 2). The last known wild
Cupressus tonkinensis is located within the
Liên Reserve (Nguyên et al. HAL11,918,
E, HN). All of the collections cited by Silba
(1994, 2005) from Guizhou China have been
identified as Cupressus funebris with the

exception of Cheng 2362 (A), which cannot
be traced. In addition, the cultivated specimens from Yunnan (Chu 51411, 51421,
KUN, NY) cannot be located.
Habitat.—Restricted to the upper parts of
slopes and ridges of karst limestone formations;
(350)–454–(650) m. At higher altitudes, and
further north, records of Cupressus refer to
cultivated Cupressus funebris.
Associated species.—Where it has not been
heavily disturbed by logging or by mining for

187

Cupressus roots, the upper ridges of the
Liên Reserve support a very stunted forest.
Liên
The species composition in the
Reserve as well as in other Cai Kinh localities
is very similar. The forest where Cu. tonkinensis is sometimes found is composed of
broad-leaved, mainly evergreen, large and
medium-sized tree species. Excentrodendron
tonkinense (A. Chev.) H. T. Chang & R. H.
Miao (Tiliaceae) is dominant. Other common
evergreen large-sized trees include Acer tonkinense Lecomte (Aceraceae), Garcinia
fagraeoides A. Chev. (Clusiaceae), Garcinia
poilanei Gagnep. (Clusiaceae), two unidentified species of Magnolia (Magnoliaceae),
Pterospermum truncatolobatum Gagnep.
(Sterculiaceae), and some unidentified species
belonging to Anacardiaceae, Burseraceae,
and Meliaceae. Common small-sized trees

include Albizia kalkora Prain (Fabaceae;
deciduous), Lagerstroemia micrantha Merr.
(Lythraceae; deciduous), Microdesmis caseariifolia Planch. ex Hook. (Pandaceae),
Platycarya strobilacea Siebold & Zucc.
(Juglandaceae), Sinosideroxylon wightianum
(Hook. & Arn.) Aubrév. (Sapotaceae), Sterculia parviflora Roxb. (Sterculiaceae), Streblus ilicifolius (Vidal) Corner (Moraceae),
and Streblus tonkinensis (Eberhardt &
Dubard) Corner (Moraceae). Shrub species
are abundant and mostly evergreen, including
Illicium difengpi B. N. Chang (Illiciaceae),
Memecylon edule Roxb. (Melastomataceae),
Phyllanthus dongmoensis N. N. Thìn
(Euphorbiaceae), Pistacia cucphuongensis T.
Ð. Ðai (Anacardiaceae), Pistacia weinmannifolia Poiss. ex Franch. (Anacardiaceae),
Schefflera pesavis R. Vig. (Araliaceae), Syzygium levinei (Merr.) Merr. (Myrtaceae), and
Tirpitzia sinensis Hallier f. (Linaceae).
Woody vines and scandent shrubs are common, for example, Acacia pennata Willd.
(Fabaceae), Aspidopterys oligoneura Merr.
(Malpighiaceae), Bauhinia oxysepala Gagnep. (Fabaceae), Bauhinia wallichii J. F.
Macbr. (Fabaceae), Beaumontia pitardii
Tsiang (Apocynaceae), Cardiopteris quinqueloba Hassk. (Cardiopteridaceae), Dalbergia
rimosa Roxb. (Fabaceae), two unidentified
species of Elaeagnus (Elaeagnaceae), Erythropalum scandens Blume (Erythropalaceae),
Maclura cochinchinensis (Lour.) Corner


188

[VOL 63

BRITTONIA

(Moraceae), Maclura fruticosa (Roxb.) Corner (Moraceae), Pegia sarmentosa (Lecomte)
Hand.-Mazz. (Anacardiaceae). In addition
there are many lithophytes, epiphytes, and
terrestrial herbs belonging to genera such as
Aglaonema (Araceae), Asplenium (Aspleniaceae), Bulbophyllum (Orchidaceae), Carex
(Cyperaceae), Cleisostoma (Orchidaceae),
Colysis (Polypodiaceae), Dendrobium
(Orchidaceae), Dischidia (Asclepiadaceae),
Eria (Orchidaceae), Hoya (Asclepiadaceae),
Impatiens (Balsaminaceae), Pyrrosia (Polypodiaceae), and Scleria (Cyperaceae).
Associated conifers.—A rich and distinct
assemblage of conifers dominates many karst
ridges in northern Việt Nam: at higher altitudes
up to ten species may be present. Nageia fleuryi
(Hickel) de Laub. (Podocarpaceae) and Podocarpus neriifolius D. Don (Podocarpaceae) are
district, near
found in the understory. In
Liên Reserve,
to the northern end of the
Pseudotsuga sinensis Dode (Pinaceae)—the
most widespread and characteristic conifer on
limestone ridge tops—dominates the canopy at
450 m (North Vietnam Second Expedition 2, E)
on ridges just outside of the reserve.
Conservation status.—CR A2acd, B2ab
(i-v), D. This species has undergone a population reduction of more than 80% within the last
three generations and its population size is
estimated to be less than 50 mature individuals.

The extent of occurrence is 204 km2 and the
actual, current area of occupancy is less than
8 km2. It is known from one area (Cai Kinh
limestone massif; Fig. 2).
Cultivated trees and ex situ collections.—
Several trees of Cupressus tonkinensis are
cultivated in the villages around and within
Liên Reserve, particularly in
Liên
the
Lũng district. The largest tree
commune,
(up to 10 m high, DBH to 40 cm) was planted
by one of the original settlers and left as a
family legacy. The seedling was collected from
one of the nearby ridges (N. V. Thang, North
East Forest Enterprise, pers. comm.). The
Liên Reserve area date
villages in the
from the mid 1960s (Furey et al., 2002)—prior
to this, the area was undisturbed. The Center for
Plant Conservation (CPC) in cooperation with
Liên Reserve planted
the residents of the
110 Cu. tonkinensis saplings in 2007 and 2008
as part of an ex situ conservation effort.

It is interesting to note that in other parts of
Việt Nam, species with high commercial value
may be locally cultivated as the resource

becomes scarce. Recently, two 10 m tall trees
of the narrowly distributed endemic Callitropsis vietnamensis (Farjon & Hiep) D. P. Little
were reported from a village garden in Hà
Giang during a survey of wild populations
(Nguyễn et al., 2006). The primary sources of
material for such plantings are wild collected
plants (Nguyễn & Thomas, 2004).
A sample from the VaÁ n Linh trees (North
Vietnam Second Darwin Expedition 111, E,
NY) was included in a DNA analysis (Little,
2006) and matched the profile of the single
sample collected from natural (not secondary)
habitat (North Vietnam Second Darwin Expedition 3, E). These cultivated trees should be
used for propagation and as a reference for
the identification of other collections—
particularly those in the plantation at the edge
of the reserve. The trees in the plantation
originated from a government nursery and at
least one is Cupressus funebris rather than
Cu. tonkinensis. All of the trees in the
plantation need to be verified against the
cultivated trees in the VaÁ n Linh village before
they are used for further propagation. The
clonal collection held at the North East Forest
Enterprise also needs to be verified against
the VaÁ n Linh trees.
Acknowledgments
We thank D. Barrington, L. Campbell, L.
Kelly, and an anonymous reviewer for constructive comments on earlier drafts of this
manuscript; D. Stevenson for color Cupressus

tonkinensis photographs. Specimens were
collected with help from K. Armstrong, D.
Atha, R. Chaudhary, Chen B., Chen W., Luu,
N. D. T., K. Shrestha, Sun H., Tan L. D.,
Tang Y., N. V. Thang, Tran B., Tran T. T.,
Wang C., Zhou Z., and numerous others
whom we have inadvertently omitted. C.
Martin generously translated various texts
from French and located critical specimens
at P. Nguyễn H. translated from Vietnamese.
Dong H. Y. kindly translated and transliterated label data from Chinese. H. Stevens


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

provided GIS assistance. Funding to DPL
from the American Society of Plant Taxonomists, the H. E. Moore Fund (BH), the Lam
Family Travel grant (Einaudi Center, Cornell
University), the National Science Foundation
(DEB 0206092), and the South East Asia
Travel grant (Einaudi Center, Cornell University) are gratefully acknowledged. PT’s
field work in Việt Nam was supported by
Darwin Initiative of the Department of Environment, Fisheries and Agriculture (DEFRA,
United Kingdom) project 162/10/017—a
project also supported by the North East
Forest Enterprise Company and the Central
Forest Seed Company of the Ministry of
Agriculture and Rural Development (MARD,

Việt Nam). Support for in situ and ex situ
conservation research as well as fieldwork
for LKP and HTN was provided by the
Basic Research Program in Natural Science
(MOST—Ministry of Science and Technology, Việt Nam; grant 6.077.06), Department
for Environment, Food and Rural Affairs
(UK), Global Tree Campaign (Fauna & Flora
International, UK; grant FSF-Defra-07-11),
and the National Foundation for Science &
Technology Development (NAFOSTED, Việt
Nam; grants 106.11.17.09 and 106.11.20.09).
Literature cited
Anonymous. 2006. Vietnamese governmental decree
number 32/2006/ND-CP dated 30 March 2006: IAGroup IA- Plant species or group of plant species for
which exploitation and commercial use is prohibited.
Việt Nam,
II,
Vật.
———. 2007. Sách
nhiên và Công nghệ. Hà Nˆoi
Nxb Khoa học
:
[Vietnam Red Data Book, part II: plants]. Publishing
House for Science and Technology, Hanoi.
Averyanov, L., P. Cribb, L. K. Phan & H. T. Nguyễn.
2003a. Slipper orchids of Vietnam. Royal Botanic
Gardens Kew, Richmond.
———, H. T. Nguyễn, T. V. Phạm & L. K. Phan. 2004.
Calocedrus rupestris sp. nov. (Cupressaceae), mˆot
: loài

tìm
đá vôi`
Việt
thông sót lại
Nam [Calocedrus rupestris sp. nov. (Cupressaceae), new
relict coniferous species from limestone areas of northern
nghiên
trong
Vietnam]. In:
. Báo cáo khoa học, H i nghị toàn
khoa học
[Issues of Basic Research in Life Sciences with direction
in upland agriculture and forestry. Proceedings. National
Conference on Life Sciences, Thai Nguyen University.
Publishing House for Science and Technology, Hanoi]:
40–44.
———, ———, L. K. Phan & T. V. Pham. 2008. The
genus Calocedrus (Cupressaceae) in the Flora of
Vietnam. Taiwania 53: 11–22.

189

———, L. K. Phan , H. T. Nguyen & D. K. Harder.
2003b. Phytogeographic review of Vietnam and the
adjacent areas of eastern Indochina. Komarovia 3: 1–
83.
Batjes, N. H. 2005. ISRIC–WISE global data set of
derived soil properties on a 0.5 by 0.5 degree grid
(version 3.0; report 2005/08). ISRIC–World Soil
Information, Wageningen.

Bay, A. V. 2003. Report on forest–based incense Cao
. Report commissioned by Cao
Rural
Development Project and Darwin Initiative Project
162/10/017.
Benjamini, Y. & Y. Hochberg. 1995. Controlling the
false discovery rate: a practical and powerful
approach to multiple testing. Journal of the Royal
Statistical Society, Series B (Methodological) 57:
289–300.
Businsky, R. 1999. Study of Pinus dalatensis Ferré and
of the enigmatic “Pin du Moyen Annam”. Candollea
54: 125–143.
———. 2004. A revision of the Asian Pinus subsection
Strobus (Pinaceae). Willdenowia 34: 209–257.
Chevalier, A. 1919. Premier inventaire des bois et autres
produits forestiers du Tonkin. Bulletin Économique
de L’Indochine 131–132: 1–228.
———. 1944. Notes sur les conifères de l’Indochine.
Revue Botanique Appliquée et d’Agriculture Tropicale 24: 7–34.
Farjon, A. 2005. A monograph of Cupressaceae and
Sciadopitys. Royal Botanic Gardens Kew, Richmond.
———, H. T. Nguyen, D. K. Harder, L. K. Phan & L.
V. Averyanov. 2002. A new genus and species in
Cupressaceae (Coniferales) from northern Vietnam,
Xanthocyparis vietnamensis. Novon 12: 179–189.
Furey, N., L. X. Canh & E. Fanning (eds.). 2002.
Liên Nature Reserve: Biodiversity Survey and Conservation Evaluation. Frontier Vietnam Forest
Research Programme Report No. 17.
Giraudoux, P. 2008. pgirmess: data analysis in ecology.

. R package version 1.3.6.
Gower, J. C. 1966. Some distance properties of latent
root and vector methods used in multivariate analysis.
Biometrika 53: 325–338.
Hickel, R. 1930. Les conifères d’Indo–Chine. Bulletin
de la Société Dendrologique de France 76: 73–78.
Hijmans, R. J., S. E. Cameron, J. L. Parra, P. G.
Jones & A. Jarvis. 2005. Very high resolution
interpolated climate surfaces for global land areas.
International Journal of Climatology 25: 1965–1978.
Kruskal, W. H. & W. A. Wallis. 1952. Use of ranks in
one–criterion variance analysis. Journal of the American Statistical Association 47: 583–621.
Little, D. P. 2006. Evolution and circumscription of the
true cypresses (Cupressaceae: Cupressus). Systematic
Botany 31: 461–480.
New, M., M. Hulme & P. Jones. 2000. Representing
twentieth–century space–time climate variability. Part
II: development of 1901–96 monthly grids of
terrestrial surface climate. Journal of Climate 13:
2217–2238.
Nguyễn, D. T. L. & P. Thomas. 2004. Cây lá kim Vi e:ˆ t
. Hà Nˆo
Nam. Nxb
: i. [Conifers of Vietnam: an
illustrated field guide. World Publishing House,
Hanoi.]


190

[VOL 63

BRITTONIA

Nguyên, H. T. & J. Vidal. 1996. Gymnospermae In:
Flore du Cambodge, du Laos et du Vietnam: 28.
Muséum National d’Histoire Naturelle, Paris.
. 2000a. Mot
ˆ:
———, L. K. Phan & L. V.
loài
cho
vật Việt Nam thu
núi
[Some new species for the flora of
đá vôi Cao
Vietnam collected from the limestone mountains of
province]. Tạp chí sinh học [Journal of
Cao
Biology], Hanoi 22: 1–11.
———, D. T. Đỗ & L. K. Phan. 2002. Tính đa dạng của
vật Việt Nam 9. Taiwania Hayata Bách tán
hệ
đài loan và T. cryptomerioides Hayata Bách tán đài
loan kín (họ Bụt mọc Taxodiaceae), chi và loài
cho hệ
vật [The diversity of the flora of Vietnam
9. Taiwania Hayata and T. cryptomerioides Hayata
(Taxodiaceae): new genus and species for the flora.]
học và

dụng [Journal of Genetics
TC Di
and Applications] 1: 32–40.
———, L. K. Phan, D. T. L. Nguyễn, P. Thomas, A.
Farjon, L. Averyanov & J. Regalado Jr. 2004.
Vietnam Conifers: Conservation Review 2004. Flora
and Fauna International, Vietnam Programme,
Hanoi.
———, T. V. To, L. K. Phan, K. S. Nguyễn, T. V.
Pham, S. T. Nguyễn & L. V. Averyanov. 2006.
Preliminary survey of Golden Cypress Xanthocyparis
vietnamensis in Ha Giang Province, northern Vietnam
within the limits of Quan Ba, Yen Minh, Dong Van
and Meo Vac districts. Report for Fauna and Flora
International—Vietnam Programme.
Nguyễn, V. K. (ed.), H. T. Nguyễn, L. K. Phan & H. T.
sinh khí hậu Việt Nam
Nguyễn. 2000b. Các
[Bioclimatic diagrams of Vietnam]. Nxb Đại học
gia Hà Nội [Vietnam National University
Publishing House, Hanoi].
Oksanen, J., R. Kindt, P. Legendre & R. B. O’Hara.
2008. Vegan: community ecology package version 1.82. R package version 1.11-4.
Phan, L. K. & H. T. Nguyễn. 1997. Các loài hạt
Vệt Nam bị đe dọa tiêu diệt và
có mặt
chúng
[Endangered gymnosperms and
trong các khu
their occurrence in nature reserves]. Tạp chí Lâm

Nghiệp [Forestry Magazine] 1: 18–20.
——— & ———. 1999. Cunninghamia konishii Hayata
có mọc hoang dại Việt Nam hay không và tên khoa
cây Sa mộc
là gì? [Is there Cunninghọc
hamia konishii Hayata growing in wild in Vietnam,
?].
and what is the scientific name of the Sa mộc
tập công trình hội
Đa dạng
Pp. 61–64. In:
Nxb Đại học
sinh học
gia, Hà Nội. [Selected papers of seminar on
north Truong Son biodiversity. Vietnam National
University Publishing House, Hanoi].
———, ——— & L. V. Averyanov. 2002. Keteleeria
davidiana (Bertrand) Beissn. var. davidiana, a gymnosperm newly identified in Northern Vietnam. Pp.
37–45. In: Sustainable Development and Protection
of the Forest and Biodiversity on Limestone Mountains of Vietnam. Forest Inventory and Planning
Institute, Hanoi.
R development core team. 2008. R: a language and
environment for statistical computing. R Foundation
for Statistical Computing, Vienna, Austria.

Roberts, D. W. 2008. LabDSV: Laboratory for Dynamic
Synthetic Vegephenomenology. http://cran.r-project.
org/. R package version 1.3-1.
Rushforth, K. 2007. Notes on the Cupressaceae in
Vietnam. Tạp chí sinh học [Journal of Biology] 29: 32–39.

———, R. P. Adams, M. Zhong, X. Q. Ma & R. N.
Pandley. 2003. Variation among Cupressus species
from the eastern hemisphere based on Random
Amplified Polymorphic DNAs (RAPDs). Biochemical Systematics and Ecology 31: 17–24.
Siegel, S. & N. J. Castellan Jr. 1988. Nonparametric
statistics for the behavioral sciences. MacGraw Hill,
New York.
Silba, J. 1994. The trans–Pacific relationship of Cupressus in India and North America. Journal of the
International Conifer Preservation Society 1: 1–28.
———. 1998. A monograph of the genus Cupressus L.
Journal of the International Conifer Preservation
Society 5: 1–98.
———. 2005. A monograph of the genus Cupressus L.
in the twenty–first century. Journal of the International Conifer Preservation Society 12: 31–103.
nhiệt
Việt
Thái, T. V. 2000. Các hệ sinh thái
Nam [The tropical forest ecosystems in Vietnam].
Chí
Nxb Khoa Học & Kỹ Thuật, Chi nhánh tại tp
Minh [Publishing House for Science and Technology,
Filial at Ho Chi Minh City].
Vu, D. V. (ed.) 1996. Vietnam Forest Trees. Agricultural
Publishing House, Hanoi.
Wessel, P. & W. H. F. Smith. 2006. GMT (Generic
Mapping Tools). Computer program distributed by the
authors />
Appendix 1
REPRESENTATIVE SPECIMENS EXAMINED
1. Callitropsis lusitanica (MILL.) D. P. LITTLE

BELIZE. Cayo: Millinario, 450 m, 1 Jun 1995,
Whitefoord 9521 (BM).
COLOMBIA. Cundinamarca: Ubaque, southeast of
Botota on trail towards Ubaqe, 4.48°N, 73.93°W, 2928 m
[cultivated], 7 Jan 1945, Little et al. 9227 (UC).
Santander: La Granja, Mesa de los Santos, 5.93°N,
73.92°W, 1630 m, 27 Jun 1953, Langenheim 3122 (UC).
COSTA RICA. Cerro de la muerte, 4 Jan 2001, Little
608 (BH), 2300 m, Little 609 (BH), 1700 m, Little 610
(BH); Volcan Irisu, 2045 m, 12 Jan 2001, Little 613 (BH).
EL SALVADOR. Ahuachapan: Ahuachapan, 13.86°
N, 89.91°W, 1340 m [cultivated], 19 Jul 2000, Monro et
al. 3431 (BM). Chalatenango: Los Eacsmiles, 14.35°N,
89.15°W, 2250 m, 25 Mar 1942, Tucker 1130 (K). Jujutla:
Dist. Finca L’esparanza, Ahuachapan, 13.82°N, 89.87°W,
1030 m, 19 Jun 1999, Monro et al. 2995 (BM). San
Salvador: San Salvador, El boqueron, 13.76°N, 89.27°W,
1800 m, 2 Apr 1998, Monro et al. 2173 (BM).
GUATEMALA. Huehuetenango: Mun. Soloma,
Soloma, Cuchumatanes, 15.72°N, 91.45°W, 2500 m, 12
Mar 1985, McCarter & Styles 390 (K).
HONDURAS. Dept. Yoro, Cerro El Cypresal, 15.15°N,
87.22°W, 1300 m, 15 Mar 1983, McCarter & Styles 117


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

(K). Intibuca: Esperanza, Parque Central, 14.30°N, 88.18°

W, 2000 m, 3 Oct 1983, Enamorado 100 (NY). Lempira:
Montana de Celaque, 14.54°N, 88.68°W, 2670 m, 29 May
1991, Davidse 34883 (BM); Montana de Celaque, El
Mojon Peak, 14.55°N, 88.68°W, 2840 m, 15 Apr 1994,
Farjon & Mejia 299 (K).
MEXICO. Between Anganguco and Tlalpuxahua,
1839, Hartweg 437 (K). Chiapas: Mun. San Cristobal
Las Casas, San Cristobal Las Casas, Cerro San Cristobal,
16.75°N, 92.63°W, 2225 m, 16 Feb 1966, Laughlin 64
(DS). Mexico: Ozumba, 19.05°N, 98.8°W, 2592 m, 8 Nov
1902, Pringle 11180 (K); Cerro Gordo, 19.15°N, 100.12°W,
3000 m, 4 Jul 1965, Rzedowski 20072 (DS). Michoacan:
Maravatio, 19.81°N, 100.35°W, 2150 m, 23 Apr 2002,
Little & Ochoterena 795 (BH, MEXU), Little & Ochoterena 796 (BH, MEXU), 19.80°N, 100.33°W, 2255 m, Little
& Ochoterena 797 (BH, MEXU), 19.76°N, 100.26°W,
2380 m, Little & Ochoterena 798 (BH, MEXU), 19.77°N,
100.22°W, 2505 m, Little & Ochoterena 799 (BH, MEXU);
Dieta, 19.45°N, 2685 m, 1 Sep 1982, Soto 4242 (CAS).
Oaxaca: Mun. Totontepec, Tepitongo, 17.30°N, 92.03°W,
1700 m, 1 Jan 1988, Velasco et al. 0172 (MO). Veracruz:
Agua Blanca, 20.41°N, 98.45°W, 2550 m, 26 Apr 2002,
Little et al. 809 (BH); Cofre De Perote, 19.54°N, 97.23°W,
2415 m, 27 Apr 2002, Little et al. 818 (BH); 19.52°N, 97.2°
W, 3300 m, 21 Jan 1984, Taylor et al. 193 (CAS).
PORTUGAL. Mata Do Bussaco, 40.38°N, 8.37°W
[cultivated], Nov 1881, Muller 1471 (DS).
2. Cupressus chengiana S. Y. Hu
CHINA. Gansu: Chu Kun, 18 Oct 1914, Meyer 1981
(NY, MO); Wuto ho, between Kaichow and Minchow,
between 34.44°N, 104.03°E, and 33.40°N, 104.92°E, Apr

1925, Rock 12073 (E, UC). Sichuan: Pao–Hsin Co.,
Mupin, Tzu–pu Soong, 30.38°N, 102.83°E, 9 Aug 1958,
anon. 06469 (SZ); Li Co., 31.47°N, 103.28°E, 2100 m, 5
Sep 1963, anon. 10604 (SCFI); Kangding Co., Kongyu,
30.05°N, 102.03°E, 1850 m, 29 Feb 1974, anon. 5325
(CDBI); Li Co., Laisugou, 31.47°N, 103.28°E, 2400 m, 3
Aug 1956, anon. 5759 (SCFI); Xiaojin Co., Muyaqiao,
31.02°N, 102.38°E, 2800 m, 9 Jul 1958, anon. 5852
(CDBI, SCFI); Ma’erkang Co., Baiwan, 31.00°N, 101.82°
E, 2380 m, 1 Aug 1975, anon. 9554 (CDBI); Ma’erkang
Co., Baiwan, 31.00°N, 101.82°E, 1 Aug 1975, anon.
9650 (CDBI); Xiaojin Co., Mengguqiao, 31.02°N,
102.38°E, 2400 m, 27 Aug 1975, anon. 9822 (CDBI);
Xiaojin Co., Xiaojin, 15 km from Xiaojin to Maerkang,
31.02°N, 102.38°E, 2500 m, 30 May 1989, Chamberlain
et al. 4044 (E); Wenchuan Co., Chekiang, 31.47°N,
103.58°E, 2 Nov 1930, Cheng 2066 (DS, E, K);
Wenchuan Co., 31.47°N, 103.58°E, 2 Nov 1930, Cheng
2073 (E, DS, NY, P); Xiaojin Co., Hubian River, 31.06°
N, 102.40°E, 2445 m, 19 Sep 2000, Howick &
McNamara 2285 (CAS, MO); Xiaojin Co., Shuangbai,
31.02°N, 102.38°E, 2500 m, 24 May 1964, Kaitai et al.
10730 (SCFI); Li–fan–hsien, 30.00°N, 100.27°E,
2550 m, 14 Aug 1943, Kuo 32 (SZ); Maoxian Co.,
Shigu (Zongqu), Minjiang river, 31.64°N, 103.81°E, 31
Aug 2002, Little et al. 858 (BH, CDBI), Little et al. 859
(BH, CDBI); Lixian Co., Muka, Zagunao river, 31.58°N,
103.36°E, 1780 m, 1 Sep 2002, Little et al. 861 (BH, CDBI);
Lixian Co., Zagunao river, 31.44°N, 103.17°E, 2030 m, 1

191

Sep 2002, Little et al. 862 (BH, CDBI); Dajin Chuang Co.,
Gana, 31.93°N, 101.88°E, 2250 m, 1 Oct 1994, Miehe et al.
94-470-7 (MB), Miehe et al. 94-470-8 (MB); Min Jiang Co.,
Li Xian, 31.43°N, 103.03°E, 2180 m, 7 Oct 1994, Miehe et
al. 94-534-3 (MB); Daocheng Co., Kasi, 29.05°N, 100.17°
E, 3800 m, 28 Jun 1973, Sichuan Plant Survey Team 2258
(CDBI, KUN); Maowen Co., Fengyi, 31.68°N, 103.87°E,
2300 m, 21 Jun 1959, Sichuan Plant Survey Team 2823
(CDBI), 2500 m, Sichuan Plant Survey Team 2843 (CDBI);
Xiangcheng Co., 29.00°N, 99.77°E, 2700 m, 26 Jul 1973,
Sichuan Plant Survey Team 2999 (KUN); Sikang Co.,
Vaszeko, 30.05°N, 102.03°E, 1500 m, 13 Nov 1934, Smith
13387 (E, MO, NY); Fu–Pien Hsien, 31.30°N, 102.45°E,
2750 m, 13 Jun 1930, Wang 21323 (KUN); Wenchuan Co.,
Min River, 31.47°N, 103.58°E, Nov 1908, Wilson 798a
(K); valle Tung Fluminis, 1907–1909, Wilson 2106 (BM,
E); Sikang Co., Shiangcheng, Tungzung, 30.05°N,
102.03°E, 3000 m, 20 Sep 1937, Yu 13407 (BM);
Kangding Co., 30.05°N, 102.03°E, 2200 m, 24 Apr
1981, Zhao 113376 E, SZ).
3. Cupressus funebris Endl.
CHINA. Fujian: Zhangzhou Co., Liqiao, 24.52°N,
117.67°E, 150 m, 20 Aug 1972, Cai & Guoliang 125
(KUN). Gansu: Huazhuang Zhan, 36.20°N, 103.12°E,
2700 m, 8 Aug 1993, Lian et al. 93-169 (MO); Wen Xian
Co., Mulinli, 32.98°N, 104.67°E, 1000 m, 18 Apr 1992,
Wang 71 (MO). Guangdong: Lechang Co., 25.13°N,
113.33°E, 30 Dec 1930, Chung 42480 (MO); Lechang

Co., 25.13°N, 113.33°E, 12 Nov 1931, Huang & Zhi
31282 (KUN); Lokchong Co., Lokchong, 25.13°N,
113.33°E, 8 Jun 1929, Tso 21017 (K, NY); Ruyuan,
24.77°N, 113.27°E, 12 Nov 1956, Wang 42583 (MO);
Tangnan, 23.72°N, 116.20°E, 1550 m, 15 Mar 1940,
Wang 87867 (KUN); Lokchong Co., Jui–Feng, Hwan–
kun, 417 m, 25.13°N, 113.33°E, 11 Oct 1928, Ying 1277
(P, UC). Guangxi Zhuang: Kwangsi, Nee Bai, 549 m,
28 Jun 1928, Ching 6281 (NY); Quan Co., Miaotou,
26.18°N, 111.22°E, 3 Nov 1958, Li & Qinghua 2087
(KUN). Guizhou: Jiangkou Co., Shaoxiding, 27.70°N,
108.85°E, 500 m, 2 Sep 1986, Bartholomew et al. 746
(CAS, NY); Jiangkou Co., Daiyenpeng, Kaitu River,
27.70°N, 108.83°E, 750 m, 10 Sep 1986, Bartholomew et al. 1047 (CAS, MEXU); Tsunyi Hsien, Liang
Feng Yah, 28.28°N, 106.83°E, 1100 m, 17 Jul 1931,
Steward et al. 10 (A, K, NY, P); Ta Ho Yen, Fanjing
Shan (Fan Ching Shan), 27.95°N, 108.83°E, 1000 m, 15
Oct 1931, Steward et al. 825 (K, NY, P); Tungtze, 28.14°
N, 106.82°E, 17 May 1930, Tsiang 4928 (E, K), 450 m,
26 May 1930, Tsiang 5123 (NY, UC); Kiangkow, 27.70°
N, 108.85°E, 450 m, 7 Dec 1930, Tsiang 7505 (BM);
Szenan, 27.94°N, 108.25°E, 200 m, 25 Jan 1931, Tsiang
8004 (A, E); Fanjingshan Co., Furong, 27.91°N, 108.71°E,
1100 m, 12 May 1959, Zhu & Liu 862 (KUN). Hubei: Pa–
Tung Hsien, 31.03°N, 110.33°E, 25 May 1934, Chow 322
(E, NY); Shennongjia Co., Nongdongwan, 31.73°N,
110.73°E, 30 Jul 1957, Fu & Zhang 1178 (KUN); Lichuan
Co., Shuishan, 30.30°N, 108.85°E, 1100 m, 4 Oct 1957, Fu
et al. 1779 (KUN); Ichang, 30.90°N, 110.80°E, Nov 1907,
Wilson 798 (E, K). Hunan: Sing–Ch’u, Sintien–hsien,

25.93°N, 112.22°E, 600 m, 28 Jul 1935, Fan & Li 371
(BM, P); Yongshun Co., Liandong, Gaofeng Hill, 28.67°N,


192

BRITTONIA

110.18°E, 860 m, 13 Oct 1982, Li & Peng 1683 (KUN);
Xining Co., Mount Ziyun, 26.60°N, 111.20°E, 700 m, 8
Nov 1962, Liu & Linhan 13270 (KUN); Xining Co., 26.40°
N, 110.80°E, 300 m, 2 Mar 1996, Luo 1249 (E), 320 m, Luo
1250 (E). Shaanxi: Mien–Hsien Co., Fangehiapa,
33.16°N, 106.67°E, 12 Apr 1942, Fu 3527 (CDBI);
Yang Co., Huayang, 34.41°N, 109.99°E (or 33.59°N,
107.54°E), 1200 m, 19 Aug 1952, Guo 1990 (CDBI);
Xixiang Co., Cha, 32.98°N, 107.76°E, 500 m, 3 Sep
1952, Guo 2115 (CDBI); Mienhsien Co., Fang–Chia–
T’a, 38.28°N, 110.10°E, 12 Apr 1942, Fu 3627 (KUN).
Shanghai: Jui–Feng, Lokchang, 31.43°N, 121.60°E, 11
Oct 1928, Ying 1277 (E). Sichuan: Tongjiang Co.,
Yong’an, 29.22°N, 104.77°E, 860 m, 7 Jun 1960, anon.
XI-1 (CDBI), anon. XII-1 (CDBI); Jiange Co., Jianmen,
32.20°N, 105.56°E, 18 Mar 1977, anon. 5 (CDBI);
Huayun Co., Mount Huayun, Liujiajing, 30.23°N,
106.67°E, 1280 m, 12 May 1959, anon. 414 (CDBI);
Pingshan Co., Longhua, 28.78°N, 104.02°E, 600 m, 2
Jun 1969, anon. 608 (CDBI); Qionglai Co., 30.42°N,
103.48°E, 710 m, 13 Jul 1959, anon. 1629 (KUN);
Pengshui Co., Daya, 29.28°N, 108.22°E, 850 m, 20 May

1959, anon. 3298 (CDBI); Baoxing Co., Longdong,
30.50°N, 102.72°E, 1600 m, 19 May 1958, anon. 4847
(CDBI), 1800 m, 20 May 1958, anon. 4877 (CDBI);
Xiangshu, 32.03°N, 105.47°E, 850 m, 5 May 1959,
anon. 5043 (CDBI); Qiaomei Co., Daping, 29.90°N,
105.25°E, 800 m, 9 May 1959, anon. 5149 (CDBI);
Maowen Co., Tumen, 31.77°N, 104.10°E, 1100 m, 8 Aug
1975, anon. 8611 (CDBI); Zhaoyuan Co., Zhuhe, 28.07°N,
102.91°E, 2150 m, 8 Jul 1976, anon. 12898 (CDBI); Mibo
Co., Huanglang, 28.40°N, 103.80°E, 1400 m, 7 Aug 1976,
anon. 13367 (CDBI); Dujiangyan Co., Qingcheng Mountain, 30.97°N, 103.52°E, 900 m, 26 Aug 1988, Boufford et
al. 24268 (CAS, MEXU); Luding Co., Moya, 29.93°N,
102.20°E, 1800 m, 19 Apr 1984, Cao & Li 36 (CDBI);
Shizhu Co., Huanghe, 29.98°N, 108.12°E, 1050 m, 2 Aug
1978, Chen 3143 (CDBI); Yuchi, 26.10°N, 101.83°E,
1150 m, 25 Apr 1978, Chen 68 (CDBI); Fengdu Co.,
Shiping Forest Farm, 29.88°N, 107.72°E, 675 m, 18 May
1996, Chen et al. 960289 (MO); Hupei Co., Modaoxi,
30.40°N, 108.07°E, 1948, Cheng & Hwa 828 (K); Ya–An,
29.98°N, 103.08°E, 1000 m, 31 Jul 1939, Chiao 1291
(CAS); Omei, 29.53°N, 103.35°E, Jul 1938, Chiao & Fan
138 (CAS, E, P); Chien–yang–hsien, 30.40°N, 104.55°E,
21 Apr 1925, Chien 5241 (E, UC); Kuan–hsien, 31.00°N,
103.62°E, 28 Feb 1937, Chien 5847 (E, UC); Lu–shan–
hsien, 30.17°N, 102.98°E, 1100 m, 16 Oct 1936, Chu 3994
(E), Chu 3995 (K); Tianquan Co., Shiyang, 29.93°N,
105.62°E, 780 m, 1 Jul 1982, Dingyi 45269 (CDBI);
Lushan Co., Shuangshi, 29.35°N, 104.67°E, 1300 m, 10
May 1983, Dingyi 47751 (CDBI); Mount Omei, 29.53°N,
103.35°E, 1067 m, 25 Aug 1928, Fang 3356 (NY, P);

Chungking, 30.63°N, 103.67°E, 15 Aug 1928, Fang 6043
(P); Chengdu, 30.67°N, 104.07°E, 3 Feb 1939, Fang 13206
(KUN); Mount Omei, 29.53°N, 103.35°E, 19 Dec 1939,
Fang 13740 (KUN), 450 m, 6 Jun 1940, Fang 13775
(KUN); Xinmin Co., 29.30°N, 102.25°E, 500 m, 24 May
1958, Fang & Mingyuan 24008 (KUN); Fengjie Co.,
Yongxing, 31.12°N, 106.98°E, 1200 m, 16 Jun 1958, Fang
& Mingyuan 24279 (KUN); Fengjie Co., Shuangdian,
31.05°N, 109.52°E, 800 m, 7 Aug 1958, Fang & Mingyuan
24976 (KUN); Chengkou Co. (Tchen Keou Tin), 32.06°N,
105.43°E, 1200 m, 19 May 1911, Farges 1216 (K, P);

[VOL 63

Jiange Co., Wen Chang Temple, 32.15°N, 105.54°E,
820 m, 18 Sep 1996, Kirkham et al. 1702 (CAS); Hejiang
Co., Tiantang, Dacao River, 28.80°N, 105.83°E, 700 m, 25
May 1981, Lan et al. 2-80 (SCFI); Nanchuan Co.,
Banxijing, 29.12°N, 107.27°E, 23 Sep 1957, Li 60799
(SCFI); Nanchuan Co., Sanquan, 29.12°N, 107.27°E,
770 m, 25 Mar 1957, Li & Guofeng 60199 (KUN);
Nanchuan Co., Niuxijing, 29.12°N, 107.27°E, 500 m, 26
Apr 1957, Li & Guofeng 60794 (KUN); Hejiang Co.,
Tiantang, 28.80°N, 105.83°E, 710 m, 26 May 1981, Liao et
al. 1-91 (SCFI); Jiange Co., Jianmenguan, 32.03°N,
105.47°E, 585 m, 10 Aug 1939, Liou & Wang 299
(KUN); Mianyang, 31.44°N, 104.59°E, 720 m, 2 Sep
2002, Little & Sun 863 (BH, CDBI); Zhitong, 31.74°N,
105.20°E, 960 m, 12 Sep 2002, Little & Sun 868.1, 868.2,
868.3, 868.4, 868.5, 868.6 (BH, CDBI); Lixian, Zagunao

river, 31.58°N, 103.36°E, 1780 m, 1 Sep 2002, Little et al.
860 (BH, CDBI); Jiange Co., Jianmenguan, Xujiadian,
32.30°N, 105.57°E, 10 Aug 1939, Liou & Wang 299
(KUN); Nanchuan Co., 29.10°N, 107.00°E, 860 m, 3 Sep
1994, Liu 14656 (BM, CAS, E, K, P), 900 m, 2 Feb 1994,
Liu 15212 (BM, CAS, E, K); Jiange Co., Jianmenguan,
Liangshan Temple, 32.07°N, 105.43°E, 1200 m, 9 Sep
1980, Mo 294 (SCFI); Nanjiang Co., 32.35°N, 106.83°E,
950 m, 15 May 1981, Mo 403 (SCFI); Tongjiang Co.,
Zhonghe, 30.33°N, 106.67°E, 850 m, 12 Jun 1981, Mo 529
(SCFI); Chin–fu Shan, 29.03°N, 107.22°E, 12 Aug 1945,
Pei 10164 (K); Tianquan Co., Yongxing, 30.43°N, 106.70°
E, 920 m, 14 Jun 1982, Peng 45538 (CDBI); Tianquan Co.,
30.07°N, 102.83°E, 890 m, 5 Jul 1982, Peng 45726
(CDBI); Mianyang Co., 31.47°N, 104.77°E, 520 m, 20
Sep 1985, Sichuan Plant Survey Team 445 (CDBI);
Baoxing Co., Yulong, 29.94°N, 105.46°E, 1500 m, 14
May 1958, Sichuan Plant Survey Team 4684 (CDBI);
Baoxing Co., 30.38°N, 102.83°E, 1600 m, 28 May
1958, Sichuan Plant Survey Team 5056 (CDBI);
Beichuan Co., Guangrong, 31.90°N, 104.30°E,
1350 m, 25 Aug 1984, Tang & Changlin 478 (CDBI);
Hechuan Co., Qingping, 30.23°N, 106.20°E, 460 m,
11 May 1959, Team 2 1582 (CDBI); Tongjiang Co.,
Mashi, 31.93°N, 107.25°E, 820 m, 17 Jun 1963, Wang
58 (SCFI); Jiulong Co., Zhongshan, 30.00°N, 103.30°
E, 1000 m, 9 May 1959, Wang 7707 (CDBI); Kuan,
31.00°N, 103.62°E, 1200 m, 13 May 1930, Wang
20606 (KUN); Wenchuan, 31.47°N, 103.58°E,
2000 m, 21 Jul 1930, Wang 21788 (KUN); Tongjiang

Co., Mashi, 31.93°N, 107.25°E, 820 m, 17 Jun 1963,
Wang & Jin’ao 58 (CDBI); Shizhu Co., Sanjiang,
30.53°N, 103.78°E, 1100 m, 9 Jul 1978, Wang &
Wenhua 1791 (CDBI); Guan Co., Lianghe, 31.22°N,
105.30°E, 650 m, 23 Jul 1987, Wang et al. 870159
(CAS, E, K, MO); Jiange Co., Liangshan, Lanmaqiang, ancient messenger route, 28.03°N, 102.87°E, 10
Oct 1984, Xi 4 (SCFI); Luding Co., 29.93°N, 102.20°
E, 2100 m, 28 Aug 1981, Xu 25871 (CDBI); Wushan
Co., Liziping, 28.99°N, 102.31°E, 1600 m, 2 Oct
1958, Yang 5553 (CDBI, KUN); Wuxi Co., Chenglongtouzui, 30.97°N, 100.22°E, 450 m, 22 Sep 1958,
Yang 59632 (CDBI); Hejiang Co., Fubao Ranch,
28.80°N, 105.83°E, 900 m, 20 May 1981, Yuan et
al. 3-30 (SCFI); Dechang Co., Wuxing, 27.58°N,
102.17°E, 1400 m, 5 Jun 1978, Zhao & Hu 5097
(CDBI); Xuyong Co., Shuiweiduizi, 28.17°N, 105.43°
E, 21 May 1959, Zhong 126 (CDBI). Yunnan:


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

Kunming, 25.04°N, 102.72°E, 1908, Dueloux 1818
(UC); Naxi, Dongde, 23.75°N, 104.80°E, 1680 m,
Little s.n. (BH, KUN); Xingcui, 23.44°N, 104.59°E
[cultivated], 27 Aug 2002, Little & Chen 842 (BH),
Little & Chen 843 (BH), Little & Chen 844 (BH),
Little & Chen 845 (BH); Naxi, Dongde, 23.75°N,
104.80°E, 1680 m, 27 Aug 2002, Little & Chen 846.1
(BH, KUN), Little & Chen 846.2 (BH, KUN), Little &

Chen 846.3 (BH, KUN), Little & Chen 846.4 (BH,
KUN), Little & Chen 846.6 (BH, KUN); Sanhe,
23.47°N, 104.25°E, 1790 m, 28 Aug 2002, Little &
Chen 847 (BH, KUN), Little & Chen 849 (BH, KUN),
23.46°N, 104.26°E, 28 Aug 2002, Little & Chen 851
(BH, KUN); Yan–Shan Co., To–Kou–Tzai, 23.61°N,
104.34°E, 1200 m, 7 Oct 1939, Wang 84218 (KUN);
Si–chou, 25.82°N, 100.17°E, 1300 m, Dec 1939,
Wang 85694 (KUN); Malipo, 23.15°N, 104.73°E,
1000 m, 4 Feb 1940, Wang 86681 (KUN); Kunming,
25.04°N, 102.72°E, 27 Oct 1991, Yang 91-1007 (KUN);
Fengqing Co., 24.58°N, 99.90°E, 22 Apr 1962, Yang &
Zenghong 8938 (KUN); Weixin Co., Shuanghe, Tiandi
District, 27.85°N, 105.20°E, 1460 m, 4 Jun 1960, Zhai &
Ping 1081 (KUN). Zhejiang: Yen–tang Shan, 28.00°N,
120.42°E, 10 Aug 1927, Chiao 14809 (K, NY, UC);
Changhua, 30.17°N, 119.22°E, 122 m, 29 Jun 1927, Keng
575 (UC).
NEPAL. Bagmati: Kathmandu Dist., Thamel, 27.70°N,
85.30°E, 1350 m [cultivated], 13 Jun 2001, Little 663 (BH).
VIỆT NAM. Tonkin, haute region, 1000 m, Capus s.n.
Văn Dist., Thai Phin Tung Commune,
(P). Hà Giang:
oc et al.
23.26°N, 105.29°E, 1286 m [cultivated], Lb
Á
HAL8677 (HN); Pho Bang Townlet, 23.25°N, 105.29°E,
Văn
oc et al. HAL11,342 (HN);
1463 m [cultivated], Lb

oc et al. Ás.n. (LE). Quan Ba Dist., Thanh Vân
[cultivated], Lb
Á
Commune, 23.07°N,
104.99°E [cultivated], 10 Dec 2001,
Luu, s.n. (E). Lạng Son: Chi Lang Dist., Hũu Liên
Commune, 21.53°N, 106.35°E, 300 m [cultivated], 2 Nov
2002, North Vietnam First Darwin Expedition 138 (E).
4. Cupressus gigantea W. C. Cheng & L. K. Fu
CHINA. Tibet: Lingshi, 29.62°N, 94.40°E, 3090 m,
11 Sep 2002, anon. s.n. (BH); Yarlung Tsangpo, Km 192,
29.00°N, 93.00°E, 3100 m, 30 Sep 1993, Bolton & Pendall
93-1 (CAS); Xizang Co., Tsela Dzong, 29.40°N, 94.42°E,
3080 m, 8 Sep 1989, Dickoré 4862 (K); Xizang Co., Nang
Xian, 29.98°N, 93.22°E, 3400 m, 4 Aug 1994, Dickoré
10560 (K); Xizang Co., Nang Xian, 29.00°N, 93.23°E, 3
Aug 1994, Dickoré 10508 (K); Lang Co., 29.05°N,
93.20°E, 3300 m, 21 Aug 1977, Guo & Benzhao
23061 (KUN); Nge, Kongbo, Tsangpo Valley, 3050 m,
21 Oct 1947, Ludlow et al. 13345 (BM, E); Xizang Co.,
Domar, Yarlung Zhangbo, 29.03°N, 93.15°E, 3390 m, 14
Sep 1997, Miehe et al. 97-080-01 (K); Xizang Co.,
Gyemdong, 29.00°N, 93.23°E, 3280 m, 4 Aug 1994, Miehe
& Wundisch 94-141-1 (K); Lang Co., 29.05°N, 93.20°E,
3400 m, 25 Jul 1975, Qing–Zang et al. 450763 (KUN);
Xizang Co., Nang Xian, Sannan Region, 1 km west of Nang
Xian, 29.05°N, 93.20°E, 3500 m, 17 Oct 1995, Rushforth
3350 (E); Xizang Co., Yarlung Tsangpo, 13–15 km below
Nang, 29.05°N, 93.20°E, 3000 m, 19 Oct 1995, Rushforth
3355 (E); Yarlung Tsangpo, station 19, 29.46°N, 94.06°E,

3200 m, 1 Oct 1997, Rushforth 4792 (E); Nyingshi Co.,
between Nyingshi and Bayi at station 66, 29.62°N, 94.40°

193

E, 3100 m, 20 Oct 1997, Rushforth 5787A (E); Medog Co.,
Gedang, 29.32°N, 95.32°E, 2200 m, 22 Mar 1993, Sun et
al. 5032 (KUN).
5. Cupressus jiangensis N. Zhao
CHINA. [cultivated] 10 Aug 1933, anon. 01105 (SZ).
Sichuan: Jiange Co., Hanyang, Cuayulang park, 32.13°N,
105.51°E, 1000 m [cultivated], 3 Sep 2002, Little & Sun
869 (BH, CDBI), Little & Sun 869a (BH, CDBI); Jiange
Co., Hanyang, Cuayulang park, 32.13°N, 105.51°E,
840 m [cultivated], 10 Sep 1980, Mo 295 (SCFI).
6. Cupressus tonkinensis Silba
: Chi Lăng Dist.,
Mạc
VIỆT NAM. Lạng
39 (HNU);
commune, 21.68°N, 106.52°E, 330 m,
commune, 21.70°N, 106.57°E, 350 m, 21
townlet,
Apr 1976, L c P-2004, P-2005 (HNU);
2336 (HNU); Vạn Linh
21.65°N, 106.58°E, 350 m,
Commune, 21.67°N, 106.48°E, 270 m [cultivated], 19
Dec 2003, North Vietnam Second Darwin Expedition 110
et al. 1034
(E); 21.67°N, 106.48°E, 21 Mar 1961,

(LE, MO); Hoa, field near farmer’s house, 21.64°N,
106.44°E, 271 m [cultivated], 5 Nov 2002, North
Vietnam Second Darwin Expedition 137 (NY); Quang
Lang commune, 21.62°N, 106.57°E, 320 m, Chevalier
Lũng Dist., Cai Kinh Commune, [“Kai
29,662 (P).
Kinsh, Lausoy”], 21.55°N, 106.40°E, Eberhardt 5073
(NY, P); Hoà Lạc commune, Sông Hóa, 21.57°N,
106.48°E, 2 Oct 1905, Castellini 169 (P); Hoà
commune,
Lệ, 21.53°N, 106.42°E, Jun 1922,
Liên Commune, 21.53°N,
Mignucci s.n. (P);
106.33°E, 270 m [cultivated], 21 Feb 2003, North Vietnam
Second Darwin Expedition 78 (E); 21.67°N, 106.38°E,
200 m [cultivated], 24 Jun 2006, Nguyên NTH6231 (HN);
24 Sep 2007, Phan et al. HAL11,220, HAL11,221,
HAL11,222 (HN); 15 Nov 2007, Phan et al. HAL11,290
(HN); 23 Feb 2008, Phan et al. HAL11,343, HAL11,344,
Liên
HAL11,345, HAL11,346, HAL11,347 (HN);
Nature Reserve, 21.69°N, 106.43°E, 464 m, 3 Nov 2003,
North Vietnam Second Darwin Expedition [Thomas & Luu]
3 (E); Lân Mai Mountain near Tân Lai village, 21.67°N,
106.39°E, 437 m, 18 Oct 2008, Nguyên et al. HAL11,918
(E, HN); Lũng Na, 21.63°N, 106.43°E, 250 m [cultivated],
19 Dec 2003, North Vietnam Second Darwin Expedition
111 (E, NY).
7. Cupressus torulosa D. Don
INDIA. Balan, 2287 m, 8 May 1948, Koelz 20324

(E); Himachal Pradesh: Chamba Dist., Brahmaur,
32.45°N, 76.53°E, 14 Sep 1896, Gammie 18581 (K);
Kullu Dist., Malana, 29.83°N, 78.92°E, 2135 m, 7 Jun
1976, Stainton 7593 (E); Simla Dist., Simla, 31.11°N,
77.15°E, 2440 m, 1847/8, Thomson s.n. (K). Punjab:
Jheri Kulu, Malaua, 25 Oct 1916, Cooper 5793 (E).
Uttarakhand: Dehradun Dist., Mussoorie Hill station,
30.45°N, 78.08°E, 1982 m, May 1977, Page 10715 (E);
Dehradun Dist., Mussoorie, 30.45°N, 78.08°E, 1830 m,
1 Mar 1967, Rodin 8089 (K); Almora Dist., Chulkot,
29.77°N, 79.77°E, 2286 m, 29 May 1951, Sahire 20407
(CAS); Dehradun Dist., Landour, 30.47°N, 78.10°E,
2135 m, Jul 1938, Stewart 16632 (NY); Kumaon Dist.,
29.83°N, 79.50°E, 1982 m, Strachey & Winterbottom s.


194

[VOL 63

BRITTONIA

n. (P); Kumaon Dist., 29.83°N, 79.50°E, 1500 m, 1 Dec
1818, Webb 6046A (K, P), Webb 6046B (K, P), Webb
6046C (K).
NEPAL. Barbung Khola, 28.87°N, 83.30°E, 3355 m,
4 Jun 1952, Polunin et al. 1063 (BM). Dhawalagiri:
Thini Khola, 28.73°N, 83.78°E, 3500 m, 15 Apr 1969,
Dobremez 40 (E); Mustang Dist., Kali Gandaki Valley,
28.63°N, 83.63°E, 2600 m, 21 May 1974, Dobremez &

Nomauther 3108 (E); Mustang Dist., Maroha, Kali
Gandaki Valley, 27.70°N, 84.42°E, 3500 m, 9 Jun
1973, Greyhilson & Phillips 771 (K); Mustang Dist.,
Lo Mantang, 29.18°N, 83.97°E, 13 Dec 1992, Kafle 013
(K); Mustang Dist., Kali Gandaki Valley, 28.68°N,
83.62°E, 2500 m, 4 Jun 2001, Little 630, 631 (BH,
TUCH), 28.69°N, 83.62°E, Little 632, 633 (BH, TUCH),
28.68°N, 83.63°E, Little 634, Little 635, 636, 637 (BH,
TUCH), 28.70°N, 83.63°E, Little 639, 640 (BH, TUCH),
28.70°N, 83.64°E, Little 641, 642, 643, 644, 645 (BH,
TUCH), 28.73°N, 83.67°E, 8 Jun 2001, Little 652, 653,
654 (BH, TUCH), 28.72°N, 83.67°E, Little 655, 656
(BH, TUCH); Mustang Dist., Syang, 28.78°N, 83.70°E,
3263 m, 4 Sep 1977, Miehe 598 (BM); Mustang Dist.,
lower Chalungpa Khola, 28.90°N, 83.77°E, 3670 m, 28
Aug 2001, Miehe et al. 01-086-01 (MB), 83.78°E,
3300 m, 28 Aug 2001, Miehe et al. 01-087-01 (K,
MB), 3670 m, 28 Aug 2001, Miehe et al. 01-089-01
(MB); Mustang Dist., south bank of lower Chalungpa,
28.88°N, 83.77°E, 3160 m, 1 Sep 2001, Miehe et al. 01101-02 (MB); Mustang Dist., Chalungpa, 28.90°N,
83.75°E, 3410 m, 8 Sep 2001, Miehe et al. 01-119-01
(MB); Mustang Dist., Tuckche, Yamkin Khola valley,
28.69°N, 83.63°E, 20 Sep 1995, Mikage & Yonekura s.n.
(E); Mustang Dist., Tukuche, between Yak Kharka and
Tukuche, 28.70°N, 83.65°E, 2810 m, 22 Aug 1994,
Noshiro et al. 9455337 (E); Mustang Dist., Tukuche,
28.70°N, 83.65°E, 3010 m, 24 Aug 1994, Noshiro et al.
9455353 (E); Mustang Dist., Larjung, Kali Gangaki
Valley, 28.68°N, 83.62°E, 2592 m, 27 May 1954,
Stainton et al. 726 (BM, E); Mustang Dist., Taglung,

Kali Gandaki Valley, 28.65°N, 83.63°E, 10 Jul 1954,
Stainton et al. 1673 (BM, E); Mustang Dist., Maikot,
28.68°N, 82.88°E, 2440 m, 26 Jun 1954, Stainton et al.
3273 (BM, E); Mustang Dist., Kali Gandaki Valley,
29.67°N, 81.00°E, 2440 m, 10 May 1965, Stainton 4848
(BM); Mustang Dist., Larjung, Kali Gangaki Valley,
28.68°N, 83.62°E, 2440 m, 3 Jun 1954, Stainton et al.
5555 (BM, E); Mustang Dist., Kalopani, 29.58°N, 83.58°
E, 2460 m, 29 Aug 1988, Suzuki et al. 8881565 (BM).
Karnali: Mugu Dist., Rara, 29.57°N, 82.07°E, 2800 m,
anon. s.n. (KATH); Dolpa Dist., Rimi, 29.13°N, 82.57°E,
2890 m, anon. s.n. (KATH); Mugu Dist., Lumsa, 29.57°
N, 82.25°E, 2050 m, Jun 1999, Miehe 99-21-01 (MB);
Mugu Dist., east of Mangri, 29.57°N, 82.37°E, 2300 m,
8 Jun 1999, Miehe 99-022-01, Miehe 99-022-02 (K); Terr
Gompa, 29.58°N, 82.43°E, 2400 m, Jun 1999, Miehe 9923-01 (MB); Dolpa Dist., upper Bauli Khola between
Polam and Sumduwa, 29.12°N, 82.92°E, 3250 m, Jun
1999, Miehe 99-88-04 (MB); Dolpa Dist., upper Suli
Khola southwest of Sumduwa Gorge, 29.12°N, 82.88°E,
3000 m, Jun 1999, Miehe 99-89-05 (MB); Miehe 99-8906 (MB); Dolpa Dist., lower Suli Khola south of Hanke,
29.00°N, 82.88°E, 2500 m, Jun 1999, Miehe 99-91-01
(K); Dolpa Dist., Suligad, Suli Khola, 29.00°N, 82.88°E,
2200 m, Jun 1999, Miehe 99-92-01 (MB); Dolpa Dist.,

Ringmo, southwest end of Poskumd, 29.17°N, 82.93°E,
3650 m, Jun 1999, Miehe 99-93-01 (MB); Dolpa Dist.,
Rimi, roadside between saddle and Rimi, 29.00°N,
82.50°E, 2910 m, 29 Sep 1991, Minaki et al. 9106095
(E); 29.50°N, 81.75°E, Oct 1975, Shrestha & Mamamdhar 311 (E). Mahakali: Darchula Dist., Laum, 29.70°N,
82.67°E, 2450 m, anon. s.n. (KATH); Darchula Dist.,

Khadang, 29.00°N, 82.67°E, 1700 m, anon. s.n.
(KATH); Launi, 29.70°N, 81.00°E, 2450 m, 6 May
1971, Dobremez 2090 (BM, E); Darchula Dist., Khadang, 29.00°N, 82.67°E, 1700 m, 24 Apr 1974,
Dobremez & Nomauther 2719 (E).
PAKISTAN. Punjab: Chamata, 33.24°N, 73.36°E,
Oct 1888, Lace 1870 (E).
Appendix 2
CONTINUOUS

MORPHOLOGICAL CHARACTERISTICS

PENULTIMATE BRANCH SEGMENTS
1. Length. The length of mature segments was measured
from the base of attachment to the apex.
2. Diameter. The diameter of the segment (stem plus
attached leaves) was measured in the medial position. If
a branch occurred at the medial position, the measurement
was taken one node up (towards the apex). For flattened
stems, the larger of the two possible measurements was
used.
3. Frequency of branching. The number of branches
per total number of nodes.
4. Leaf length. Cupressus species have small, tightly
appressed leaves that are difficult to directly measure
(Fig. 6). To estimate the leaf length, the length of the
segment was measured and the number of leaves
(including immature leaves) on one side of the four–
sided penultimate axis was counted. Segments with
overlapping leaves were avoided. Characters 4–6 were
scored from the facial leaves of dimorphic species,

and the leaves of monomorphic species. The comparison of facial leaves to monomorphic leaves was made
because these two leaf types are very similar; in
contrast the lateral leaves are markedly different from
monomorphic leaves (e.g., Fig. 6A and 6C vs. 6B and
6C). These characters were also scored for the lateral
leaves of dimorphic species as characters 7–9.
5. Frequency of resin glands. Resin glands (also
referred to as resin pustules in some literature) are
manifest as a single indentation (sporadically with a
slightly exerted center) on the abaxial surface of the leaf
(e.g., Figs. 6L, 7H). Glands were considered present if
they were clearly visible at low magnification (7.5–10×).
On some specimens leaf glands are present only as a
darkened, slightly sunken area. Data were collected from
one side of the four–sided main axis of a penultimate
segment.
6. Frequency of active resin glands. Resin glands were
considered active if they produced some exudate.
Color of the exudate was not scored because it could
not be easily quantified.
ULTIMATE BRANCH SEGMENTS
10. Segment length. Measured like character 1, but for
ultimate segments.


2011]

LITTLE ET AL.: CUPRESSUS TONKINENSIS (CUPRESSACEAE)

11. Segment diameter. Measured like character 2, but

for ultimate segments.
12. Frequency of branches on the primary plane. The
primary plane was defined as the plane with the
greatest number of branches (Fig. 10).
13. Frequency of apical pointing ultimate segments.
The number of ultimate segments pointing
towards the apex of the antepenultimate segment
that ultimately bears them per the number of
ultimate segments on that plane. The plane is
defined by the antepenultimate segment, the
penultimate segment, and ultimate segments
pointing towards the apex of the antepenultimate
segment (Fig. 10).
ADULT ULTIMATE–SEGMENT LEAVES
14. Leaf length. Scored like character 4, but for the
ultimate segment leaves. Characters 14–16 were
scored from the facial leaves of dimorphic species,
and the leaves of monomorphic species. In addition
these characters were scored from the lateral leaves
of dimorphic species as characters 17–19.
15. Frequency of resin glands. Scored like character 5,
but for the ultimate segment leaves.
16. Frequency of active resin glands. Scored like
character 6, but for the ultimate segment leaves.
POLLEN CONES
20. Cone length. The length of the pollen cone along
the axis from the attachment point on the axis of
the first cone scale to the top of the last cone scale
was measured. In the case of irregular cones the
largest possible value was used.

21. Cone width. The medial width of the pollen cone,
perpendicular to the axis, was measured. In the case
of irregular cones the largest possible value was
used.
22. Pairs of opposite decussate cone scales. The
number of pairs of pollen cone scales was counted.
OVULATE CONES
23. Cone length. The length of the ovulate cone along
the axis from the attachment point on the axis of the
first woody cone scale to the top of the last cone
scale—including the boss—was measured. In the
case of irregular cones the largest possible value was
used.
24. Cone width. The width of the ovulate cone,
perpendicular to the axis, was measured. If present
the boss(es) were included. In the case of irregular
cones the largest possible value was used.
25. Pairs of opposite decussate cone scales. The
number of pairs of ovulate cone scales was counted.
Cases in which the terminal pair consisted of a single
scale (due to the failure of the scales in the pair to fully
differentiate) were counted as a full pair.
SEEDS
26. Seed length. Measured at longest point, from the
hilum to the micropyle.
27. Seed width. Measured at the widest point.
28. Seed thickness. Measured at the thickest point.

195

29. Hilum length. Measured at the longest point.
30. Hilum width. Measured at the widest point.
DISCRETE MORPHOLOGICAL
CHARACTERISTICS
ANTEPENULTIMATE BRANCH SEGMENTS
31. Branch arrangement. Scored as all branches on one
plane (1) or branches on two planes (2). Rare observations (occurring less than 5% of the time) were excluded.
PENULTIMATE BRANCH SEGMENTS
32. Branch arrangement. Scored as character 31, but
for penultimate segments.
33. Multiple branches per node. Scored as never more
than one ultimate segment per node (1) or as
sporadically two ultimate segments per node (2)—
most nodes have only one ultimate segment, but
several instances of multiple ultimate segments per
node can usually be found on an average herbarium
specimen of a species that has this character state.
34. Leaf form. Leaves were considered dimorphic if
alternating pairs differed significantly in size and/or
shape (e.g., Fig. 6I vs. 6J and Fig. 8A vs. 8B). In
Cupressus most differences in leaf shape are due to
bending around a non–radially symmetrical stem. In
many instances the lateral leaves of a dimorphic
species have a lower frequency of resin gland
occurrence, and in some a different marginal leaf
band. Scored as monomorphic (1) or dimorphic (2).
ULTIMATE BRANCH SEGMENTS
35. Arrangement of branches. Scored as character 31,
but for ultimate branches.
36. Stem cross–sectional shape. Scored as isodiametric

(0) or rectangular (1). The state of a given specimen can
usually be determined by careful external observation
rather than actually making a cross section.
37. Leaf form. Scored like character 34, but for ultimate
segment leaves.
38. Leaf apex. Scored as obtuse (0; Fig. 8K) or acute (1;
Fig. 8A). Characters 38 and 39 were scored from the
facial leaves of dimorphic species, and the leaves of
monomorphic species. These characters were also
scored for the lateral leaves of dimorphic species as
characters 40 and 41.
39. Leaf apex adornment. Scored as unadorned (0; e.g.,
Fig. 8C) or with a white mucronate tip (1; e.g., Fig. 8A).
OVULATE CONES
42. The mucro on the boss of the apical cone scales.
Scored as entire (0) or ciliate (1) at the time of seed
maturation. Older cones are often weathered to the
point that it is difficult, if not impossible, to
accurately score this character. In some species
different pairs of ovulate cone scales have different
mucro types. For standardization the terminal pair
of scales was arbitrarily selected to score this
character for all taxa.
43. Cones abscission. Scored as abscised upon maturity
(0) or as remaining attached to the tree for an extended
period (1). Abscission of mature cones was scored from
field observations where possible.