39. Galium paradoxum Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg. 19: 281. 1874.
林猪殃殃 lin zhu yang yang
Herbs, perennial, ascending from filiform rhizomes. Stems erect, slender, 4-25 cm tall, 4-angled and narrowly winged, glabrous and smooth, only nodes slightly shortly hairy. Middle stem leaves opposite and with 2 leaflike but clearly smaller stipules in whorls of 4, at lower nodes stipules linear, 1.5-3 mm; petiole 1.5-10 mm; leaf blade membranous, suborbicular, broadly ovate to ovate-lanceolate, or elliptic-oblong, (5-)6-30(-40) × (3.5-)5-15(-23) mm, adaxially with scattered, ± appressed, short hairs, abaxially glabrescent, base attenuate, obtuse to truncate, margins antrorsely hispidulous-ciliolate, apex acute to rounded; single principal vein with 2-4 pairs of pinnate lateral veins. Inflorescences terminal and in axils of upper leaves with 3-11-flowered cymes; axes trichotomous and ± divaricate; bracts narrowly elliptic or ligulate, 0.8-3 mm; pedicels 1-3 mm. Ovary ovoid, ca. 0.5 mm, with undeveloped uncinate hairs. Corolla white, rotate, 2.5-3 mm in diam., lobed for 1/2-2/3; lobes ovate, obtuse, subapiculate to acute or acuminate. Mericarps ovoid, 1-2 mm, densely covered with uncinate yellowish brown trichomes 0.8-1 mm, on pedicels thickening and elongating up to 11 mm. Fl. May-Aug, fr. Jun-Sep.
Forests, meadows, near water, on shady (sub)alpine rocks; 1200-4000 m. Anhui, Gansu, Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jilin, Liaoning, Qinghai, Shanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Russia].
Galium paradoxum is a rather unusual species of the genus because of its broad, petiolate, and pinnately nerved opposite leaves, forming whorls of 4 with 2 smaller leaflike stipules. Therefore, it can be confused with other genera, in particular Kelloggia, which differs by its calyx teeth and its never-leaflike stipules. Because of its isolated position, G. paradoxum was made the type species of G. sect. Cymogalia and placed into a monotypic series, G. ser. Paradoxa (see Pobedimova et al., Fl. URSS 23: 326. 1958). As shown by Ehrendorfer et al. (Fl. Iranica 176: 232. 2005) and unpublished DNA analyses, it is only distantly related to G. kamtschaticum, G. rotundifolium, and G. elegans in G. sect. Platygalium s.l. or to members of G. sect. Hylaea, as G. hoffmeisteri.
Schönbeck-Temesy and Ehrendorfer (in Tan et al., Davis & Hedge Festschrift, 111-114. 1989) commented on the morphological variation and biogeography of Galium paradoxum and recognized three subspecies, of which subsp. paradoxum and subsp. duthiei occur in China as keyed out and described below. The third, G. paradoxum subsp. franchetianum Ehrendorfer & Schönbeck-Temesy, is restricted to Japan and can be recognized by its relatively larger leaves with relatively short trichomes and often acute apices, and by its relatively large flowers. A survey of many new collections now available from the herbaria PE, KUN, MO, and WU shows that ranges of variation in all differential features used overlap and that many intermediates occur. With respect to the two Chinese taxa one can suspect not only an allopatric but also an altitudinal differentiation pattern.
