492 
Telopea 8(4): 2000 
with cuticular spinules arising from the angles of palisade-like, very slender and 
radially elongated cells of the outer layer of the testa; the cuticle is concave over the 
outer surface of each cell. The flange encircling the seed in the median vertical plane 
appears to be a projection of the cuticle and not a cellular structure. 
Reproductive biology: the flowers are wind-pollinated and dioecious, with the seeds 
serving as disseminules. The pollen is released explosively and normally completely 
expelled from the anther. Flowering is autumn or spring (bimodal) (Meney & Pate 
1999). Germination is hypogeal with the epicotylar rhizome developed (Pate & Meney 
pers. comm.) Lyginiaceae occur in fire-prone environments, and the presence of an 
obligate seeder species as well as resprouters, with distinctive patterns of resource 
allocation, was reported and discussed by Pate and colleagues (Bell & Pate 1993; 
Meney, Dixon & Pate 1999; Pate, Meney, Dixon, Bell & Hickman 1999). 
Fungal associates: Lyginiaceae resemble Restionaceae in their resistance to 
Phytophthora infection but a culm smut recorded as infecting all Lyginia species 
(Websdane et al. 1994; Sieler et al. 1999) is sufficiently distinctive to have been 
described as the new monotypic genus Websdanea (Vanky 1997). This smut has not 
been recorded on other hosts. Meney et al. (1993) recorded the presence of mycorrhizal 
associations for a brief period during winter in the roots of Lyginia ; mycorrhizae have 
rarely been reported in Restionaceae. 
Phytochemistry: Lyginia differs from members of related families in the presence of 
the unusual sugar allose and of allosides, while resembling Restionaceae in producing 
some simple flavonol glycosides (Williams et al. 1997). Quercetin and isorhamnetin 
were reported in most samples; proanthocyanidins and flavonoid sulphates are 
absent. Williams et al. also note that, as in some Restionaceae, 'the more chemically 
advanced 8-hydroxy flavonol gossypetin ... is present in at least one species'. Unlike 
Restionaceae, Lyginia produces significant amounts of fructan-type oligosaccharides in 
its rhizomes and lacks starch (Pate, Meney & Dixon 1991). 
Systematics and relationships: Lyginia has until now been included in Restionaceae. 
As noted above, cladistic analysis of DNA sequence data from the chloroplast gene 
) cL, the /n/L intron and the fnzL-F intergenic spacer (Briggs et al. 2000) do not 
associate Hopkinsia and Lyginia with Restionaceae but show a robust clade of these two 
genera with Anarthria. Within this clade Anarthria is sister to ( Hopkinsia , Lyginia). 
As with Hopkinsia, the general similarity of Lyginia to Restionaceae in gross 
morphology and anatomy, and its occurrence in a region of great diversity of 
estionaceae, have led to its distinctiveness being overlooked. Lyginiaceae possess 
various conditions that are considered plesiomorphic within Poales, in the 
in oiescence, flowers (unreduced tepals, tetrasporangiate anthers, gynoecium), pollen 
morp o ogy, ovule and flavonoids. However, they show distinctive apomorphies in 
1 oresc fnce structure, connate staminal filaments, stomatal position, structure of the 
chlorenchyma, and seed surface. 
Distribution and habitats: the single genus Lyginia is widespread on well drained or 
seasona y moist oligotrophic soils in the southern half of Western Australia, in regions 
o mo erate to low rainfall. One species is rare but two are wide-ranging and common. 
Lyginia 
Lyginia R. Br., Prodr.: 248 (1810). 
Lectotype. L. barbata R. Br. (typ. cons.) (In 1910, Lyginia R. Br. was conserved against 
c toeno urn Labill., following a proposal by Janchen 1908), but this was subsequently 
recognised as a superfluous conservation (Rickett & Stafleu 1959, p. 231). Schoenodum 
