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

DSpace at VNU: Before it had a name: Diagnostic characteristics, geographic distribution, and the conservation of Cupres...

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 Lewis B and Dorothy Cullman Program for Molecular Systematics, The New York Botanical Garden, Bronx, NY 10458-5126, USA; e-mail: dlittle@nybg.org Royal Botanic Garden, Edinburgh EH3 5LR, Scotland, UK; e-mail: P.Thomas@rbge.ac.uk 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: ntienhiep@hn.vnn.vn 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 Hanoi University of Science, 334 Nguyen Trai, Thanh Xuan, Ha Noi, Việt Nam; e-mail: pkeloc@yahoo.com 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: 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, Hồ 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: Sơng 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 Sơng Hố 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 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 km2 (using a 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 Localities of Cu tonkinensis in Lũng province, Việt Nam and Chi Lăng districts, Lạng Grid squares are × 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 (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 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 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 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 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 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 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 http://www.daac.ornl.gov) 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 http://www.isric.org) 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 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 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 accounts for 61.7% and PCO 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 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 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 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 POLYMORPHISMS ARE ENCLOSED IN BRACKETS VARIATION IN ULTIMATE SEGMENT LEAF FORM (CHARACTER 37) IS ILLUSTRATED IN FIGS 6, 7, AND VARIATION IN ULTIMATE SEGMENT LEAF APICES (CHARACTERS 38 AND 39) IS ILLUSTRATED IN FIG 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 [12] 2 [12] [12] [12] [12] [12] [12] [12] 1 [12] 2 2 [12] [12] 1 [12] [01] [01] 1 [12] 2 1 [01] [01] 0 0 [01] 1 0 — [01] — 0 — — [01] — [01] — [01] [01] 1 [01] [01] [01] [01] [01] ? [01] [01] ? ? [01] [01] [01] [01] [01] [01] 182 BRITTONIA [VOL 63 FIG 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 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 10 11 12 13 14 15 16 17 18 19 20 21 units 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) ( ) 7– – ( ) – – ( ) – – ( ) – – (0.8)0.8–0.9– (1.6)c (1.2) 1(1.0) (1.5) (1.4)ft ( ) – – 0 (80)83–100– (100)100– (71)71–100– (100)100–100– (100) 100(100) 100–100(100) 100(100) 100(100) ( 3 ) 3 – – 0 (25)33–65– (50)50–67– (0)0–78–100(100) ( ) – – (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) ( ) – – (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) [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; (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 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 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); 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 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 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 : lồi tìm đá vơi` Việt thơng sót lại Nam [Calocedrus rupestris sp nov (Cupressaceae), new relict coniferous species from limestone areas of northern nghiên 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; 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D P LITTLE BELIZE Cayo: Millinario, 450 m, Jun 1995, Whitefoord 9521 (BM) COLOMBIA Cundinamarca: Ubaque, southeast of Botota on trail towards Ubaqe, 4.48°N, 73.93°W, 2928 m [cultivated], 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, 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, 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, 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, Nov 1902, Pringle 11180 (K); Cerro Gordo, 19.15°N, 100.12°W, 3000 m, 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, Sep 1982, Soto 4242 (CAS) Oaxaca: Mun Totontepec, Tepitongo, 17.30°N, 92.03°W, 1700 m, 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) 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, Aug 1958, anon 06469 (SZ); Li Co., 31.47°N, 103.28°E, 2100 m, 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, Aug 1956, anon 5759 (SCFI); Xiaojin Co., Muyaqiao, 31.02°N, 102.38°E, 2800 m, Jul 1958, anon 5852 (CDBI, SCFI); Ma’erkang Co., Baiwan, 31.00°N, 101.82° E, 2380 m, Aug 1975, anon 9554 (CDBI); Ma’erkang Co., Baiwan, 31.00°N, 101.82°E, 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, Nov 1930, Cheng 2066 (DS, E, K); Wenchuan Co., 31.47°N, 103.58°E, 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, Sep 2002, Little et al 861 (BH, CDBI); Lixian Co., Zagunao river, 31.44°N, 103.17°E, 2030 m, 191 Sep 2002, Little et al 862 (BH, CDBI); Dajin Chuang Co., Gana, 31.93°N, 101.88°E, 2250 m, 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, 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) 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, 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, 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, Nov 1958, Li & Qinghua 2087 (KUN) Guizhou: Jiangkou Co., Shaoxiding, 27.70°N, 108.85°E, 500 m, 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, 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, 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, Nov 1962, Liu & Linhan 13270 (KUN); Xining Co., 26.40° N, 110.80°E, 300 m, 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, 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, Jun 1960, anon XI-1 (CDBI), anon XII-1 (CDBI); Jiange Co., Jianmen, 32.20°N, 105.56°E, 18 Mar 1977, anon (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, 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, May 1959, anon 5043 (CDBI); Qiaomei Co., Daping, 29.90°N, 105.25°E, 800 m, May 1959, anon 5149 (CDBI); Maowen Co., Tumen, 31.77°N, 104.10°E, 1100 m, Aug 1975, anon 8611 (CDBI); Zhaoyuan Co., Zhuhe, 28.07°N, 102.91°E, 2150 m, Jul 1976, anon 12898 (CDBI); Mibo Co., Huanglang, 28.40°N, 103.80°E, 1400 m, 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, 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, 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, Feb 1939, Fang 13206 (KUN); Mount Omei, 29.53°N, 103.35°E, 19 Dec 1939, Fang 13740 (KUN), 450 m, 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, 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, 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, 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, Sep 1994, Liu 14656 (BM, CAS, E, K, P), 900 m, Feb 1994, Liu 15212 (BM, CAS, E, K); Jiange Co., Jianmenguan, Liangshan Temple, 32.07°N, 105.43°E, 1200 m, 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, 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 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, 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, 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 (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, 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, 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, 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, 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, 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], Nov 2002, North Vietnam First Darwin Expedition 138 (E) 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, Sep 1989, Dickoré 4862 (K); Xizang Co., Nang Xian, 29.98°N, 93.22°E, 3400 m, Aug 1994, Dickoré 10560 (K); Xizang Co., Nang Xian, 29.00°N, 93.23°E, 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, 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, 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, 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) 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], 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) 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], 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); Hồ Lạc commune, Sơng Hóa, 21.57°N, 106.48°E, 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, Nov 2003, North Vietnam Second Darwin Expedition [Thomas & Luu] (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) Cupressus torulosa D Don INDIA Balan, 2287 m, 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, 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, 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, Dec 1818, Webb 6046A (K, P), Webb 6046B (K, P), Webb 6046C (K) NEPAL Barbung Khola, 28.87°N, 83.30°E, 3355 m, 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, 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, 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, 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, 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, Sep 2001, Miehe et al 01101-02 (MB); Mustang Dist., Chalungpa, 28.90°N, 83.75°E, 3410 m, 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, 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, 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, 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 CONTINUOUS MORPHOLOGICAL CHARACTERISTICS PENULTIMATE BRANCH SEGMENTS Length The length of mature segments was measured from the base of attachment to the apex 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 Frequency of branching The number of branches per total number of nodes 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 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 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 196 BRITTONIA 44 First year cone scale color Scored as rich glossy brown at the end of the first year (0) or as green or green–blue at the end of the first year (1) This character can only be scored accurately from living plants, as the colors of the scales change unpredictably upon drying [VOL 63 SEEDS 45 Seed wing resin pustules Scored as absent (0) or present (1) Usually manifest as distinct bubbles on the surface of the wing, mostly aggregated towards the seed corpus ... 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... 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. .. example, Acacia pennata Willd (Fabaceae), Aspidopterys oligoneura Merr (Malpighiaceae), Bauhinia oxysepala Gagnep (Fabaceae), Bauhinia wallichii J F Macbr (Fabaceae), Beaumontia pitardii Tsiang