4 BRITTONIA [VOL. 57 FIG. 2. Micromorphology <strong>of</strong> leaves. A–D. Margin <strong>of</strong> <strong>the</strong> leaf. A. <strong>Polygonum</strong> sawatchense subsp. sawatchense (scale bar � 0.3 mm). B. P. utahense (scale bar � 0.3 mm). C. P. sawatchense subsp. oblivium (scale bar � 100 �m). D. P. spergulariiforme (scale bar � 100 �m). E, F. papillae on <strong>the</strong> leaf blade: E. P. spergulariiforme. F. P. utahense (scale bars � 30 �m).
2005] COSTEA & TARDIF: POLYGONUM DOUGLASII COMPLEX 5 lobes are oblong-elliptic to oblong-obovate, <strong>of</strong>ten cucullate and overlapping, naviculate along <strong>the</strong>ir entire length or only in <strong>the</strong> upper 1/4 (Fig. 3). Vasculature <strong>of</strong> tepals consists <strong>of</strong> a single vein, which may be branched or not. EPICUTICULAR WAX PATTERN OF PERIANTH LOBES Epidermis cells are irregularly shaped, elongated or rectangular (Figs. 4, 5). Stomata are usually present. The anticlinal walls are usually sinuate or sometimes straight or weakly undulate (Figs. 4, 5). Epicuticular wax sculpture pattern is variable and complex: Type 1. Epidermis cells with <strong>the</strong> periclinal walls more or less flat. Sculpture pattern is homogenous on <strong>the</strong> same cell. Wax rodlets may be arranged in a dense reticulate, intertwined pattern (Figs. 4C, 5E, F) or <strong>the</strong>y may be more or less transversally oriented (Fig. 4A, B). Type 2. Epidermis cells with <strong>the</strong> periclinal walls evidently convex. Sculpture pattern is heterogeneous on <strong>the</strong> same cell: wax rodlets are � parallel among <strong>the</strong>mselves and perpendicular to <strong>the</strong> long axis <strong>of</strong> cells on <strong>the</strong> lateral inclined faces and arranged in a dense reticulate and intertwined pattern on <strong>the</strong> more or less flat top <strong>of</strong> <strong>the</strong> periclinal walls (Fig. 4D–F, 5A– D). STAMENS Stamens are most commonly eight, arranged in two ‘‘whorls.’’ The outer whorl consists <strong>of</strong> five stamens and <strong>the</strong> inner <strong>of</strong> three stamens. Stamens in <strong>the</strong> outer ‘‘whorl’’ may be reduced, sometimes entirely (e.g., P. sawatchense subsp. sawatchense). The base <strong>of</strong> <strong>the</strong> inner stamens is thickened and associated with a nectarial tissue. An<strong>the</strong>rs are short, 100–550 �m, pink to purple, rarely whitish. POLLEN The pollen belongs to <strong>the</strong> ‘‘Duravia type’’ described by Hedberg (1946). It is prolate, usually tricolporate with <strong>the</strong> exine semitectate-reticulate or microreticulate at <strong>the</strong> mesocolpia and poles, and microreticulate-foveolate or sometimes foveolate with microspinules around ectoapertures (Fig. 6). Two distinct types can be recognized based <strong>the</strong> size <strong>of</strong> <strong>the</strong> pollen grains, <strong>the</strong> reticulum sculpturing at <strong>the</strong> middle part <strong>of</strong> <strong>the</strong> mesocolpium and <strong>the</strong> sculpturing <strong>of</strong> ectoapertures: Type A. Pollen grains 20–26 �m long; mesocolpium is microreticulate with 1–7 rows <strong>of</strong> lumina, 0.5–1(–1.1) �m indiameter, clearly differentiated or undifferentiated from <strong>the</strong> ectoapertures, which are microreticulate to tectate perforate, foveolate-punctate, sometimes with microspinules. ● Subtype A1.—Mesocolpium is microreticulate and not clearly differentiated from <strong>the</strong> ectoapertures. Mesocolpium microreticulum with 4–7 rows <strong>of</strong> lumina, 0.5–0.9 �m in diameter (Fig. 6A–C). ● Subtype A2.—Mesocolpium is reticulate, clearly differentiated from ectoapertures. Mesocolpium reticulum with 1– 5 rows <strong>of</strong> lumina, 0.9–1.1 �m in diameter (Fig. 6D, E). Type B. Pollen grains 28–33 �m; mesocolpium is reticulate with 1–2 rows <strong>of</strong> lumina, 1.5–4 �m in diameter; mesocolpium is sharply differentiated from <strong>the</strong> ectoapertures, which are tectate imperforate, rarely punctate, with a few puncta (Fig. 6F–H). ACHENES Achenes are black, trigonous, ovate, ovate-lanceolate, elliptic, and enclosed in <strong>the</strong> perianth or exserted at maturity (Fig. 7). Faces are concave and subequal; one face is slightly broader than <strong>the</strong> o<strong>the</strong>r two (Fig. 7). The exocarp is usually lustrous and smooth, or sometimes it may be striate-tubercled, especially near <strong>the</strong> edges and apex <strong>of</strong> achenes (Fig. 7C, F). Tubercles may be arranged in longitudinal rows, more or less following <strong>the</strong> anticlinal walls (Fig. 7D). <strong>Taxonomy</strong> Hickman (1984) included within P. <strong>douglasii</strong> five o<strong>the</strong>r species that share a significant area <strong>of</strong> sympatry: P. sawatchense