Galium aparine L.

Abstract

Galiun L., Section KOLGYDA Dumort. (Section Aparine Koch). An annual herb with prostrate or, more usually, scrambling-ascending diffusely branched stems 15-300cm, stout and hairy at the nodes, the four angles very rough with downwardly directed prickles. Leaves 12-60 x 3-8 mm, in whorls of 6-9, narrowly to widely oblanceolate or elliptical, mucronate, 1-veined, glabrous or with hooked bristles above, margins rough with backward-pointing prickles. Flowers 1.5 mm diameter in 2-5-flowered axillary cymes, their peduncles topped by a whorl of 4-8 bracts; hermaphrodite, sometimes male (andromonoecious). Calyx a minute annular ridge; corolla 1.5-1.7 mm in diameter, whitish, with four acute lobes. Stamens 4, exserted; styles 2, short, connate below; stigmas capitate. Two-seeded fruits with length of longest axis 3-5 mm (excluding setae), wider than the corolla, olive or purplish, covered with white hooked setae with tuberculate bases, their stalks divaricate. Mature fruit a dry mericarp with length of longest axis 2-3 mm, and a mean airdry mass of 9.65 mg ? 0.33 (10 samples of 100 from a hedgerow population). Salisbury (1974) reports the average mass of seed from shade-tolerant woodland and hedgerow plants as 3.7 mg, and Grime et al. (1988) record a mean seed mass of 7.25mg for a hedgerow population. Galiumn iparine is a highly variable species owing to a combination of phenotypic plasticity and genecological variation. The species has different responses to the environment corresponding to different environmentally conditioned life cycles [see VI (e)]; plasticity in the timing of seed germination, productivity, growth form, freezing tolerance, and the light and temperature dependence of photosynthesis enable G. apa7ine to establish itself in a wide variety of habitats. Potential variability concealed in the higher polyploid conditions, especially in the widespread hexaploids [see VI (d)], enabling them to respond to changing environmental conditions, is probably also the basis of their success. Among plants of G. apariine raised from seed collected from various sites in continental Europe, B6cher etal. (1955) distinguished at least five different climatic ecotypes or races that either may be discrete or more probably ranges of a large clinal variation: mediterranean, summer-annual (seed germinates in the spring) with accelerated development from Portugal; mediterranean, biennial from Spain; southern, winterto summer-annual from France; northern, rather late flowering winterto summerannual from Denmark and Poland; and northern, very late flowering winter-annual (seed germinates in autumn) from Denmark and Sweden. Ecotypes of G. aparine have been demonstrated by Berkefeld (1988), differing in shape of cotyledons, hairiness of stems, colour of stem edges and length of fruit spines in plants grown under uniform garden conditions from seed collected in several different habitats. In Britain, two discrete ecotypes of G. apa7i71e exist, one found in arable fields or open situations and the other growing in hedgerows (FroudWilliams 1985); the two ecotypes are very similar morphologically but differ markedly in seed dormancy and germination requirements [see VIII (d)]. Bain & Attridge (1988) provide evidence of genotypic differences in shade strategies between hedgerow and field populations [see VI (e)]. Two forms have been recognized, the rarer G. aparine f. interimediwan Bonnet, with smooth or tuberculate fruits without spiny hairs, and the common form G. apar-ine f. apari71e with spiny fruits (Moore 1975). Variants of G. spur-ium with setose fruits, apart from the flower characters and the diploid chromosome number, are often not easy to distinguish from G. caparine and have been described as G. spuritini ssp. infestlin (Hanf 1983). Galiuiml apariine is native on maritime shingle beaches, in primary tall-herb fen communities, and in primary alderwood in natural hydrarch successions, but is also a very variable cosmopolitan weed found in hedges, waste places, drained fen peat, on limestone scree, roadside verges, river and stream banks, and is a major arable weed, especially of cereal crops. *Abbreviated references are used for standard works, see Jolurnal of Ecology (1975), 63, 335-344. Nomenclature of vascular plants follows Florca Euri0opaea and Stace (1997) for British species.