THELIGONACEAE [Draft]

假繁缕科  jia fan lu ke

Chen Jiarui (陈家瑞 Chen Chia-jui)[1]; A. Michele Funston[2]

Annual or perennial monoecious herbs. Stems somewhat succulent. Leaves petiolate, somewhat succulent, the lower ones opposite, upper ones alternate, entire; stipules membranaceous, uniting at the base of petioles. Flowers unisexual, rarely bisexual, axillary in upper nodes, small 1-3 flowered cymes. Male flowers: sessile; perianth lobes 2–5, valvate, closed in bud, spreading or reflexed at anthesis, 3–5 parallel nerves; stamens (2–)6–30, anthers linear, straight in bud, dorsifixed at anthesis, free slender filaments, opening lenghthwise. Female flowers: subsessile; perianth tubular, oblique, the upper part an elongate constricted mouth, 2–4 denticulate-lobes; ovary one carpel; style simple, extended from tube; ovule solitary, basal. Fruit a nutlike drupe, compressed; seed U-shaped, endosperm fleshy.

One genus and about four species: Atlantic islands, Mediterranean region, and E Asia; three species (two endemic) in China.

Traditionally Theligonaceae has been treated as a monogeneric family of controversional relationship, Rubiales (Cronquist), Myrtales (Engler), Caryophyllales (Dalla Torre & Harms); however, recent reliable molecular data places it within a highly derived group of Rubiaceae (along with Galium, Hedyotis, and Rubia; Pl. Syst. Evol. 225: 43-75. 2000).

Wan Wenhao. 2000. Theligonaceae. In: Chen Chiajui, ed., Fl. Reipubl. Popularis Sin. 53(2): 148–151.

1. THELIGONUM Linnaeus, Sp. Pl. 2: 993. 1753.

假繁缕属  jia fan lu shu

Cynocrambe Gagnebin.

Morphological characters and geographic distribution are the same as those of the family.

1a.       Stamens 5–7; stem often ca. 10 cm........................................................ 1. T. formosanum

1b.       Stamens ca. 20; stem 15-50 cm.

2a.       Annual herbs; leaf-blades 2–5 × 1.5–3 cm; stamens more than 20....... 3. T. macranthum

2b.       Perennial herbs; leaf-blades 0.7–3 × 0.7–1.5 cm; stamens 20–25 or 16–20  2. T. japonicum

1. Theligonum formosanum (Ohwi) Ohwi & T. S. Liu in H. L. Li et al., Fl. Taiwan 3: 904. 1977.

台湾假繁缕tai  wan jia fan lu

Cynocrambe formosana Ohwi, Acta Phytotax. Geobot. 2: 157. 1933.

Small perennial herbs, becoming black when dry. Stem erect, 10–15 cm, branched from base, creeping below, somewhat hairy. Leaf blade broadly ovate, 1.2–2 × 1–1.5 cm, base subcordate, subtruncate or rounded, decurrent to petiole, apex acute, pilose adaxially, puberulent on veins abaxially; petioles 0.5–1.2 cm; stipules membranaceous, connate, deltoid-ovate, 2–3 mm, ciliate. Flowers white, unisexual. Male flowers: perianth membranaceous, 3-parted, lobes oblanceolate, 2.5–3 mm, obtuse, reflexed, 3–5 veins; stamens 5–7. Female flowers: smaller; perianth 2-lobed, lobes obtuse. Fruit obliquely obovoid, ca. 4 mm, hairy. Fl. Mar–Jul, fr. Jun–Aug.

* Wet places in forests, along mountain trails; ca. 2700 m. Taiwan (Pingdong).

2. Theligonum japonicum ôkubo & Makino, Bot. Mag. (Tokyo) 8: 348. 1894.

日本假繁缕  ri ben jia fan lu

Cynocrambe japonica (ôkubo & Makino) Makino.

Fetid perennial herbs. Stem erect or ascending, 15–36 cm, branched often from base, creeping below, pubescent above. Leaf blade ovate or elliptical, 0.7–3 × 1–1.5 cm, base rounded or broadly cuneate, apex acute, pubescent on both surfaces, lateral veins 3 each side; petioles 0.3–1.5 cm; stipules membranaceous, connate, ovate or deltoid-ovate, 2–3 mm, ciliate. Flowers unisexual. Male flowers: perianth membranaceous, 3-parted, lobes oblanceolate, 2.5–3 mm, obtuse, reflexed, 3–5 veined; stamens 20–25 or 16–20, pendulous. Female flowers: minute; perianth 3–4-toothed. Fruit obliquely ovoid, ca. 5 mm, hairy. Fl. Mar–Jun, fr. Jun–Aug.

Wet places in valleys, by streams; 900–1200 m. Anhui, Shaanxi, Zhejiang [Japan].

3. Theligonum macranthum Franchet, Nouv. Arch. Mus. Hist. Nat., sér. 2, 10: 71 [Pl. David. 2: 109]. 1887.

假繁缕  jia fan lu

Cynocrambe macrantha (Franchet) Poulsen.

Fetid annual herbs. Stem erect, 30–50 cm, ferrugineously pubescent. Leaves herbaceous, leaf blade broadly ovate, ovate-lanceolate or suboblong, 2–5 × 1–3 cm, base subcuneate or subrounded, somewhat decurrent to petiole, apex acuminate, sparsely pubescent on both surfaces, glabrescent; petioles 0.5–1.8 cm; stipules membranaceous, connate, ovate or deltoid-ovate, 4–6 mm, ciliate. Flowers unisexual. Male flowers: perianth membranaceous, 2-lobed, tube 2 mm, lobes 3–5 mm, lanceolate, apex obtuse, reflexed, 5–7 veined; stamens more than 20. Female flowers: smaller; perianth 2 lobed, apex obtuse. Fruit ovoid, ca. 5 mm. Fl. Apr–Jul, fr. Jun–Aug.

* Wet places in forests; 1800–2400 m. W Hubei, Sichuan, Zhejiang.



[1] Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

[2] Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A.