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

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An Erratum to this article was published on 30 March 2012

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.

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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 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).

Author information

Authors and Affiliations

  1. Lewis B. and Dorothy Cullman Program for Molecular Systematics, The New York Botanical Garden, Bronx, NY, 10458-5126, USA

    Damon P. Little

  2. Royal Botanic Garden, Edinburgh, EH3 5LR, Scotland, UK

    Philip Thomas

  3. Institute of Ecology and Biological Resources, Vietnam Academy of Natural Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Ha Noi, Việt Nam

    Hiêp T. Nguyễn & Lôc K. Phan

  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

    Hiêp T. Nguyễn & Lôc K. Phan

  5. Hanoi University of Science, 334 Nguyen Trai, Thanh Xuan, Ha Noi, Việt Nam

    Lôc K. Phan

Authors
  1. Damon P. Little
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Correspondence to Damon P. Little.

Additional information

An erratum to this article can be found at http://dx.doi.org/10.1007/s12228-011-9234-9

Appendices

Appendix 1

Representative specimens examined

1. Callitropsis lusitanica (Mill.) D. P. Little

BELIZE. Cayo: Millinario, 450 m, 1 Jun 1995, Whitefoord 9521 (BM).

COLOMBIA. Cundinamarea: 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 morte, 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 (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 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, 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); 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: 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. (P). Hà Giang: Vn Dist., Thai Phin Tung Commune, 23.26°N, 105.29°E, 1286 m [cultivated], L \( \widehat{{\mathop {o}\limits_\cdot }} \) c et al. HAL8677 (HN); Pho Bang Townlet, 23.25°N, 105.29°E, 1463 m [cultivated], L \( \widehat{{\mathop {o}\limits_\cdot }} \) c et al. HAL11,342 (HN); Vn [cultivated], L \( \widehat{{\mathop {o}\limits_\cdot }} \) c et al. s.n. (LE). Quan Ba Dist., Thanh Vân 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°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

VIỆT NAM. Lạng : Chi Lng Dist., Mạc commune, 21.68°N, 106.52°E, 330 m, n 39 (HNU); commune, 21.70°N, 106.57°E, 350 m, 21 Apr 1976, Lộc P-2004, P-2005 (HNU); townlet, 21.65°N, 106.58°E, 350 m, Ti p 2336 (HNU); Vạn Linh Commune, 21.67°N, 106.48°E, 270 m [cultivated], 19 Dec 2003, North Vietnam Second Darwin Expedition 110 (E); 21.67°N, 106.48°E, 21 Mar 1961, et al. 1034 (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 29,662 (P). Lũng Dist., Cai Kinh Commune, [“Kai 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, Mignucci s.n. (P); Liên Commune, 21.53°N, 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, HAL11,345, HAL11,346, HAL11,347 (HN); Liên 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.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. 01-101-02 (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 99-23-01 (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 99-89-06 (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. 1.

    Length. The length of mature segments were measured from the base of attachment to the apex.

  2. 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. 3.

    Frequency of branching. The number of branches per total number of nodes.

  4. 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. 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. 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

  1. 10.

    Segment length. Measured like character 1, but for ultimate segments.

  2. 11.

    Segment diameter. Measured like character 2, but for ultimate segments.

  3. 12.

    Frequency of branches on the primary plane. The primary plane was defined as the plane with the greatest number of branches (Fig. 10).

    Fig. 10
    figure 10

    Branch segments on the plane of the paper are drawn as solid-fill lines, segments pointing out of the plane of the paper away from the viewer are drawn as dashed lines, and segments pointing out of the plane of the paper towards the viewer are drawn as unfilled lines. The antepenultimate segment is labeled. The penultimate segments are labeled at their apex with “p”. The remaining unlabeled segments are all ultimate segments. Penultimate segments A and D bear ultimate segments on only one plane (character 35) whereas segments B and C bear segments on two planes. The frequency of ultimate segment branches on the primary plane (character 12) for segment A = 100%, B = 60% (6 out of 10), C = 62% (5 out of 8), and D = 100%. The frequency of apical pointing ultimate segments (character 13) for segment A = 50% (5 out of 10), B = 50% (2 out of 4), C = 60% (3 out of 5), and D = 66% (2 out of 3).

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  4. 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

  1. 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.

  2. 15.

    Frequency of resin glands. Scored like character 5, but for the ultimate segment leaves.

  3. 16.

    Frequency of active resin glands. Scored like character 6, but for the ultimate segment leaves.

Pollen Cones

  1. 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.

  2. 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.

  3. 22.

    Pairs of opposite decussate cone scales. The number of pairs of pollen cone scales was counted.

Ovulate Cones

  1. 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.

  2. 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.

  3. 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

  1. 26.

    Seed length. Measured at longest point, from the hilum to the micropyle.

  2. 27.

    Seed width. Measured at the widest point.

  3. 28.

    Seed thickness. Measured at the thickest point.

  4. 29.

    Hilum length. Measured at the longest point.

  5. 30.

    Hilum width. Measured at the widest point.

Discrete morphological characteristics

Antepenultimate Branch Segments

  1. 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

  1. 32.

    Branch arrangement. Scored as character 31, but for penultimate segments.

  2. 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.

  3. 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

  1. 35.

    Arrangement of branches. Scored as character 31, but for ultimate branches.

  2. 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.

  3. 37.

    Leaf form. Scored like character 34, but for ultimate segment leaves.

  4. 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.

  5. 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

  1. 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.

  2. 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.

  3. 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.

Seeds

  1. 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.

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Little, D.P., Thomas, P., Nguyễn, H.T. et al. Before it had a name: Diagnostic characteristics, geographic distribution, and the conservation of Cupressus tonkinensis (Cupressaceae). Brittonia 63, 171–196 (2011). https://doi.org/10.1007/s12228-010-9146-0

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