Flora of North America species comparison

Salix orestera is so similar to S. glauca var. villosa that it is sometimes included in S. glauca. It is phenetically most similar to S. eastwoodiae and S. lemmonii (G. W. Argus 1997), and the possibility that it is a hybrid or an alloploid involving those species needs study.

Salix orestera can be distinguished from S. glauca var. villosa in having largest medial blades usually narrower, 3.4–7.1 times as long as wide, abaxial surfaces distinctly silky with appressed hairs pointing toward apex, hairs sometimes ferruginous, secondary veins raised abaxially and adaxially, branches often strongly glaucous, and staminate flowers without abaxial nectaries; S. glauca var. villosa has largest medial blades usually broader, 2.2–3.9 times as long as wide, abaxial surfaces usually glabrescent or, if hairy, hairs unkempt, slightly spreading, hairs always white, secondary veins raised abaxially, flat or impressed adaxially, branches not glaucous, staminate flowers often with abaxial nectaries.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera 50+, species ca. 1000 (4 genera, 123 species in the flora).

Taxonomic placement of the Salicaceae and the genera included in it have varied greatly. Some botanists (H. G. A. Engler and K. Prantl 1887–1915) treated it as a primitive member of the Dicotyledoneae and grouped it with other families having simple, apetalous, unisexual flowers arranged in catkins, the “Amentiferae.” At about the same time, others (C. E. Bessey 1915) took a different view, regarding the simple flowers as the result of reduction, and placed the taxa in Caryophyllales. As early as 1905, H. Hallier could see that there were similarities between Salicaceae and Flacourtiaceae; at the time, he was vigorously challenged by E. Gilg (1915). A. D. J. Meeuse (1975) summarized evidence for a close relationship between these families, including wood anatomy, phytochemistry, host-parasite relationships (including rust fungi), and morphology. He concluded that the Salicaceae could be combined with the Flacourtiaceae, “perhaps as a tribe.” A. Cronquist (1988) and R. F. Thorne (1992b) placed the Salicaceae, in a narrow sense, in Violales near Flacourtiaceae.

Molecular studies support a close relationship between Salicaceae and Flacourtiaceae in Malpighiales and show that Flacourtiaceae, in a broad sense, is paraphyletic. Based on a study of plastid rbcL DNA sequences, Salix and Populus were nested within a subset of 52 genera of Flacourtiaceae (M. W. Chase et al. 2002). Chase et al. proposed moving some genera from broadly circumscribed Flacourtiaceae to Salicaceae. Other studies, based on different gene sequences, came to the same conclusion (O. I. Nandi et al. 1998; V. Savolainen et al. 2000; K. W. Hilu et al. 2003; Angiosperm Phylogeny Group 2003). The discovery of the extinct fossil genus Pseudosalix (L. D. Boucher et al. 2003), from the Eocene Green River Formation of Utah, provided further support for placing some members of Flacourtiaceae in Salicaceae. The well-preserved Pseudosalix fossils, in which reproductive structures are directly associated with the leaves, occur intermixed with Populus fossils. The leaves are slender and have salicoid teeth, inflorescences are cymose, flowers are unisexual, pedicellate, tetrasepalous, and 3- or 4-carpellate, and seeds are comose, i.e., having characteristics intermediate between Salicaceae and Flacourtiaceae.

The presence, in both families, of salicoid teeth is often cited in support of their close relationship (W. S. Judd 1997b; O. Nandi et al. 1998; M. W. Chase et al. 2002; H. P. Wilkinson 2007). Salicoid teeth were first recognized and defined as having the tip of the medial vein (seta) of the tooth retained as a dark, but not opaque, non-deciduous spherical callosity fused to the tooth apex and were reported to occur in Salicaceae and Idesia of the Flacourtiaceae (L. J. Hickey and J. A. Wolfe 1975). Nandi et al. reported that a broad survey of angiosperm leaves showed that salicoid teeth occur outside of Flacourtiaceae and Salicaceae only in Tetracentraceae.

Isozyme and cytological evidence show that Populus and Salix are ancient polyploids (D. E. Soltis and P. S. Soltis 1990; Wang R. and Wang J. 1991). All Salix and Populus species contain salicin (R. T. Palo 1984).

The genera often included in Salicaceae, in the narrow sense, are Chosenia, Populus, Salix (A. K. Skvortsov 1999), and, sometimes, Toisusu. Molecular studies (E. Leskinen and C. Alström-Rapaport 1999; T. Azuma et al. 2000) show that Chosenia is nested within Salix. H. Ohashi (2001) treated Toisusu as Salix subg. Pleuradinea Kimura and Chosenia as Salix subg. Chosenia (Nakai) H. Ohashi.

(Discussion copyrighted by Flora of North America; reprinted with permission.)