Flora of North America species comparison

Populus trichocarpa has been mistakenly reported from North Dakota based on specimens of P. ×jackii (P. balsamifera × P. deltoides). It hybridizes with P. balsamifera to form P. ×hastata Dode along the northern Rocky Mountain axis (Alaska, Alberta, British Columbia, and Idaho). Hybrids have capsules with 2–4 glabrous or sparsely hairy valves. The extent of hybridization has led to treatment of P. trichocarpa as a subspecies of P. balsamifera (T. C. Brayshaw 1965; L. A. Viereck and J. M. Foote 1970); these two balsam poplars are more closely related to Asian members of sect. Tacamahaca than they are to each other (J. E. Eckenwalder 1996). Mountain ranges of the Intermountain Region (California, Idaho, Nevada, Oregon, and Utah) have trees intermediate between P. trichocarpa and P. angustifolia. These hybrids have narrower leaves with shorter petioles and sparsely hairy capsules with 2–3 valves.

In addition to hybridizing with other North American species of sect. Tacamahaca, Populus trichocarpa also hybridizes with both native species of sect. Aigeiros. Populus ×generosa A. Henry (synonym P. ×interamericana van Broekhuizen), a hybrid between P. trichocarpa and P. deltoides, is rare in the far western area of the range for P. deltoides subsp. monilifera, where it overlaps with the more drought-tolerant inland P. trichocarpa (Idaho, Montana, Washington, and Wyoming) (J. E. Eckenwalder 1984). This hybrid has also been grown artificially, and such hybrids between coastal P. trichocarpa and P. deltoides subsp. deltoides are becoming increasingly important plantation trees in the Pacific Northwest from northern Oregon to British Columbia, as well as in Europe. They are perhaps the fastest growing of all poplars in volume, with the rapid height growth of P. trichocarpa added to the steady diameter growth of P. deltoides (R. F. Stettler et al. 1988).

Populus ×parryi Sargent, a hybrid between P. trichocarpa and P. fremontii, is commonly found in a wide variety of mesic habitats throughout the region of sympatry between its parents in California and Nevada (and beyond the range of P. trichocarpa in Mohave County, Arizona; J. E. Eckenwalder 1992). It can be found particularly in canyons where its parents are elevationally separated but overlap as permanent streams spill out into lower elevations (Eckenwalder 1984, 1984b). A morphologically and ecologically distinctive race of P. trichocarpa in coastal southern California with heart-shaped leaves may have arisen through this kind of hybridization (Eckenwalder 1984c). This race includes the type of P. trichocarpa from Ventura County.

Populus maximowiczii A. Henry is an Asian balsam poplar that is sometimes cultivated as an ornamental, but usually as a plantation tree or parent of plantation hybrids. It is distinguished from P. trichocarpa and P. balsamifera by its elliptic leaves with rugose adaxial surfaces.

(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.)