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BEN # 227
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No. 227 July 5, 1999
aceska@victoria.tc.ca Victoria, B.C.
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Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2
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BEN # 226 and BEN # 277 are dedicated to
DR. THOMAS CHRISTOPHER (CHRIS) BRAYSHAW,
Curator Emeritus of the Royal British Columbia Museum,
Victoria, British Columbia, Canada,
on the occasion of his 80th birthday, July 2, 1999.
CLASSIFICATION OF SALIX IN THE NEW WORLD
George W. Argus, 1999
The genus Salix is a large, taxonomically complex, genus of
about 450 species worldwide; estimates range from 330-350 to
500. Salix occurs mainly in the Northern Hemisphere. Its center
of abundance is in China, where there are about 270 species, and
in the former Soviet Union, where there are about 120 species.
There are 103 species in North America and 65 species in Europe.
The genus also occurs in Japan, Africa, the Middle East, India,
and Central and South America. In Australasia and Oceania it
occurs only as introductions, some of which are regarded to be
weedy.
In a revised classification of New World species of Salix
(Argus, G.W., 1997, Systematic Botany Monographs No. 52), based
on a numerical taxonomic study, ninety morphological characters
were recorded for each of 103 species. A phenetic distance
matrix was generated using the Gower metric. The data were
analyzed using several clustering methods; the most consistent
results were achieved using ISS FLEX. The New World species were
grouped into four subgenera. Salix subg. Salix is represented by
four sections, S. subg. Longifoliae by one section, S. subg.
Chamaetia by eight sections, and S. subg. Vetrix by 15 sections.
The species composition of each of the sections was compared
with previous taxonomic treatments and problematic taxa were
discussed.
The major problem in the classification of Salix into a linear
arrangement of subgenera and sections is the presence of
polyploidy as well as hybridization and introgression. Many
Salix are polyploid, with ploidal levels of 4x-12x. The
hypothesis was presented (Argus 1997) that intersectional and
intersubgeneric polyploidy is responsible for difficulties in
placing some species into sections and sections into subgenera.
An indirect method used to identify the possible ancestry of
some allopolyploids was to compare species with their nearest
neighbors, based on their Gower dissimilarity value. If the
nearest neighbors belonged to a different section it was
hypothesized that the alloploid may be of intersectional hybrid
origin. For example, Salix maccalliana (dodecaploid) has as its
nearest neighbors members of S. sect. Salicaster (subg. Salix)
and S. sect. Hastatae (subg. Vetrix) and Salix glauca
(tetraploid to hexaploid) has nearest neighbors in S. sect.
Glaucae, S. sect. Myrtilloides, and S. sect. Hastatae (the first
two are in subg. Salix, the third in subg. Vetrix). In the
cluster analysis, using different sets of taxa and characters,
these species, and other high polyploids, often moved between
subgenera. Because the evolutionary pattern of many Salix is
reticulate the placement of intersectional allopolyploids in one
section or another is arbitrary.
The following classification of New World Salix arranges species
into subgenera and sections. Some of the nomenclature in sec-
tions Subalbae, Albae, Glaucae, Cordatae Phylicifoliae and
Geyerianae has been modified from Argus (1997). The names and
citation of introduced and naturalized taxa are marked with
asterisk (*) and appear in Arial font in the website BEN ver-
sion.
Salix subg. Salix
Salix sect. Floridanae Dorn
Salix floridana Chapman, Fl. South. U.S. 430. 1860.
Salix sect. Humboldtianae Andersson
Salix amygdaloides Andersson, Ofvers. Forh. Kongl.
Svenska Vetensk.-Akad. 15: 114. 1858; Salix bonplandiana Kunth
in H.B.K., Nov. Gen. Sp. 2: 24. 1817; Salix caroliniana Michx.,
Fl. bor.-amer. 2: 226. 1803; Salix gooddingii C. R. Ball, Bot.
Gaz. 11: 376. 1905; Salix humboldtiana Willd., Sp. Pl. 4: 657.
1805; Salix laevigata Bebb, Amer. Natu. 8: 202. 1874; Salix
nigra Marshall, Arbust. amer. 139. 1785.
Salix sect. *Subalbae Koidz.
*Salix babylonica L., Sp. Pl. 2: 1017. 1753; *Salix
?pendulina Wenderoth, Schrift. Nat. Ges. Marb. 2: 235. 1831 (S.
babylonica ? S. fragilis).
Salix sect. *Salix
*Salix alba L., Sp. Pl. 2: 1021. 1753; *Salix alba var.
vitellina (L.) Stokes, Bot. Mat. Med. 4: 506. 1812 (Basionym:
Salix vitellina L., Sp. Pl. 1016. 1753); *Salix alba var.
caerulea (Sm.) Sm. Engl. Fl. 4: 231.1828 (Basionym: Salix
caerulea Sm. Engl. Bot. 34: t 2431. 1812); *Salix fragilis L.,
Sp. pl. 2: 1017. 1753 (Probably equals S. ?rubens Schrank,
Reidar Elven, pers. com. June 1999); *Salix ?sepulcralis Simonk.
Oest. Bot. Zeitschr. 40: 424. 1890 (Salix alba L. ? S.
babylonica L.); *Salix ?sepulcralis Simonk. nothovar. chrysocoma
(Dode) Meikle, Watsonia 15: 274. 1985 (Basionym: S. chrysocoma
Dode, Bull. Soc. Bot. Fr. 55: 655. 1909; *Salix alba var. vitel-
lina ? S. babylonica).
Salix sect. Salicaster Dumort.
Salix lucida Muhl., Ges. Naturf. Freunde Berlin II. 4:
239. 1803; Salix lucida subsp. caudata (Nutt.) E. Murray, Kalmia
15: 11. 1984 "1985" (Basionym: Salix pentandra L. [var.] caudata
Nutt., North Am. Sylva 1: 61. 1842); Salix lucida subsp.
lasiandra (Benth.) E. Murray, Kalmia 15: 11. 1984 "1985"
(Basionym: Salix lasiandra Benth. Pl. Hartweg. 335. 1857); Salix
lucida subsp. lucida; *Salix pentandra L., Sp. pl. 2: 1016.
1753; Salix serissima (L. H. Bailey) Fernald, Rhodora 6: 6. 1904
(Basionym: Salix lucida Muhl. var. serissima L. H. Bailey, Geol.
& Nat. Hist. Surv. Minn. Bull. 3: 19. 1887).
Salix sect. Maccallianae Argus
Salix maccalliana Rowlee, Bull. Torr. Bot. Club 34: 158.
1907.
Salix subg. Longifoliae (Andersson) Argus
Salix sect. Longifoliae (Andersson) Andersson
Salix exigua Nutt. N. Amer. Sylv. 1: 75. 1842; Salix
interior Rowlee, Bull. Torrey Bot. Club 27: 253. 1900; Salix
fluviatilis Nutt. N. Amer. Sylva 1: 89. 1842; Salix melanopsis
Nutt. N. Amer. Sylva 1: 78. 1842; Salix microphylla Schldl. &
Cham. Linnaea 6: 354. 1831; Salix sessilifolia Nutt. N. Amer.
Sylva 1: 68. 1842; Salix taxifolia Kunth in H.B.K. Nov. gen. sp.
2: 18. 1817. The nomenclature proposed for this section by Dorn
(Brittonia 50: 193-210. 1998) has not yet been evaluated.
Salix subg. Chamaetia (Dumort.) Nasarov
Salix sect. Chamaetia Dumort.
Salix nivalis Hook. Fl. bor-amer. 2: 152. 1838; Salix
reticulata L., Sp. pl. 2: 1018. 1753; Salix reticulata subsp.
glabellicarpa Argus, Canad. J. Bot. 43: 1021. 1965; Salix
reticulata subsp. reticulata; Salix vestita Pursh, Fl. Amer.
Sept. 2: 610. 1814.
Salix sect. Setchellianae Argus
Salix setchelliana C. R. Ball, Univ. Calif. Publ. Bot.
17: 410. 1934.
Salix sect. Herbella Ser. (S. sect. Retusae)
Salix herbacea L., Sp. pl. 2: 1018. 1753; Salix num-
mularia Andersson in DC. Prodr. 16(2): 298. 1868.
Salix sect. Myrtosalix A. Kerner
Salix arctophila Cockerell ex A. Heller, Cat. N. Amer.
Pl., ed. 3, 89. 1910; Salix chamissonis Andersson in DC. Prodr.
16(2): 290. 1868; Salix fuscescens Andersson, Monogr. Salicum
97. 1867; Salix phlebophylla Andersson in DC., Prodr. 16(2):
290.1868; Salix polaris Wahlenb. Fl. Lap. 261. 1812; Salix
rotundifolia Trautv. Nouv. Mem. Soc. Nat. Mosc. 2: 304. 1832;
Salix rotundifolia subsp. dodgeana (Rydb.) Argus, Canad. J. Bot.
47: 795. 1969 (Basionym: Salix dodgeana Rydb. Bull. N. Y. Bot.
Gard. 1: 277. 1899); Salix rotundifolia subsp. rotundifolia;
Salix uva-ursi Pursh, Fl. Amer. Sept. 2: 610. 1814 .
Salix sect. Ovalifoliae (Rydb.) C. K. Schneider
Salix jejuna Fernald Rhodora 28: 177. 1926; Salix
ovalifolia Trautv. Nouv. Mem. Soc. Mosc. 2: 306. 1832; Salix
ovalifolia var. arctolitoralis (Hulten) Argus, Canad. J. Bot.
47: 795. 1969 (Basionym: Salix arctolitoralis Hulten, Sv. Bot.
Tidskr. 34: 373. 1940); Salix ovalifolia var. cyclophylla
(Rydb.) C. R. Ball, Proc. Nat. Acad. Sci. 21: 184. 1935
(Basionym: Salix cyclophylla Rydb. Bull. N. Y. Bot. Gard. 1:
274. 1899); Salix ovalifolia var. glacialis (Andersson) Argus,
Canad. J. Bot. 47: 798. 1969 (Basionym: Salix glacialis An-
dersson, Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 15: 131.
1858); Salix ovalifolia var. ovalifolia; Salix stolonifera
Coville, Proc. Wash. Acad. Sci. 3: 333. 1901.
Salix sect. Diplodictyae C. K. Schneider
Salix arctica Pall., Fl. Ross. 1: 86. 1788; Salix cas-
cadensis Cockerell, Muhlenbergia 3: 9. 1907; Salix petrophila
Rydb. Bull. N. Y. Bot. Gard. 1: 268. 1899; Salix sphenophylla A.
K. Skvortsov in Tomatchev, Fl. Arct. URSS 5: 62. 1966.
Salix sect. Myrtilloides (Borrer) Andersson
Salix athabascensis Raup, Rhodora 32: 111. 1930; Salix
chlorolepis Fernald Rhodora 7: 186. 1905; Salix pedicellaris
Pursh, Fl. Am. Sept. 2: 611. 1814; Salix raupii Argus, Canad. J.
Bot. 52: 1303. 1974.
Salix sect. Glaucae (Fries) Andersson
Salix brachycarpa Nutt. N. Am. Sylva 1: 69. 1842; Salix
brachycarpa var. brachycarpa; Salix brachycarpa var. psammophila
Raup, J. Arnold Arb.17: 230. 1936; Salix glauca L., Sp. Pl. 2:
1019. 1753; Salix glauca subsp. acutifolia (Hook.) Hulten, Ark.
f. Bot. 7: 40. 1967 (Basionym: Salix villosa Andersson ? [var.]
acutifolia Hook. Fl. bor.-am. 2: 145. 1839); Salix glauca subsp.
callicarpaea (Trautv.) Bocher, Meddel. Groenl. 147: 19. 1952
(Basionym: Salix callicarpaea Trautv. Nouv. Mem. Soc. Nat. Mosc.
2: 295. 1832); Salix glauca subsp. stipularis (Flod. ex Hayren)
Hiitonen, Suomen kasvio, 272. 1933 (Basionym: Salix glauca var.
glauca auct.; Salix stipulifera Flod. ex Häyren, Mem. Soc. Faun.
Fl. Fenn. 5: 133. 1929); Salix glauca var. villosa Andersson,
Proc. Amer. Acad. Arts Sci. 4: 68. 1858; Salix niphoclada Rydb.
Bull. N. Y. Bot. Gard. 1: 272. 1899.
Salix subg. Vetrix (Dumort.) Dumort.
Salix sect. Hastatae (Fries) A. Kerner
Salix arizonica Dorn, Canad. J. Bot. 53: 1499. 1975;
Salix ballii Dorn, Can J. Bot. 53: 1501. 1975; Salix barclayi
Andersson, Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 15: 125.
1858; Salix boothii Dorn, Canad. J. Bot. 53: 1505. 1975; Salix
commutata Bebb, Bot. Gaz. 13: 110. 1888; Salix cordata Michx.
Fl. bor.-amer. 2: 225. 1803; Salix eastwoodiae Cockerell ex A.
Heller, Cat. N. Am. Pl. ed. 3 89. 1910; Salix farriae C. R. Ball
in Standley, Contr. U. S. Natl. Herb. 22: 321. 1921; Salix
hastata L., Sp. pl. 2: 1017. 1753; Salix monticola Bebb in
Coulter, Man. Bot. Rocky Mt. Reg. 336. 1885; Salix myricoides
Muhl. Ges. Naturf. Fr. Neue Schr. 4: 235. 1803; Salix myricoides
var. albovestita (C. R. Ball) Dorn, Canad. J. Bot. 54: 2782.
1976 (Basionym: Salix glaucophylla Bebb var. albovestita C. R.
Ball, Jour. Wash. Acad. Sci. 29: 429. 1939); . Salix myricoides
var. myricoides; Salix myrtillifolia Andersson, Ofvers. Forh.
Kongl. Svenska Vetensk.-Akad. 15: 132. 1858; Salix orestera C.
K. Schneider, J. Arnold. Arb. 1: 164. 1920; Salix pseudomon-
ticola C. R. Ball in Standley, Contr. U. S. Natl. Herb. 22: 321.
1921; Salix pseudomyrsinites Andersson, Ofvers. Forh. Kongl.
Svenska Vetensk.-Akad. 15: 130. 1858; Salix pyrifolia Andersson,
Monogr. Salicum 162. 1867; Salix wolfii Bebb in Rothr. Rep. Bot.
U. S. Geogr. Surv. 6: 214. 1878.
Salix sect. Cordatae Barratt ex Hook.
Salix eriocephala Michx. Fl. bor.-amer. 2: 225. 1803;
Salix eriocephala var. eriocephala; Salix eriocephala var.
famelica (C.R. Ball) Dorn, Brittonia 47: 165. 1995 (Basionym: S.
lutea var. famelica C.R. Ball, Bot. Gaz. 71: 426. 1921); Salix
ligulifolia (C. R. Ball) C. R. Ball in E. D. Smith, Amer. Midl.
Nat. 27: 236. 1942 (Basionym: Salix lutea Nutt. var. ligulifolia
C. R. Ball, Bot. Gaz. 71: 428. 1921); Salix lutea Nutt. N. Amer.
Sylva 1: 63. 1842; Salix monochroma C. R. Ball, Bot. Gaz. 71:
431. 1921; Salix prolixa Andersson, Monogr. Salicum 94. 1867;
Salix turnorii Raup, J. Arnold Arb. 17: 234. 1936.
Salix sect. Cinerella Ser. (S. sect. Vetrix)
*Salix atrocinerea Brotero, Fl. Lusit. 1: 31. 1804;
*Salix aurita L., Sp. pl. 2: 1019. 1753; *Salix caprea L., Sp.
pl. 2: 1020. 1753; *Salix cinerea L., Sp. pl. 2: 1021. 1753;
Salix discolor Muhl. Ges. Naturf. Freunde Berlin II. 4: 234.
1803; Salix hookeriana Barratt ex Hook. Fl. bor.-amer. 2: 145.
1838; Salix humilis Marshall, Arbust. amer. 140. 1785; Salix
humilis var. humilis; Salix humilis var. tristis (Aiton) Griggs,
Proc. Ohio Acad. Sci. 4: 301. 1905; Salix paradoxa Kunth in
Humboldt and Bonpland, Nov. Gen. Pl. 2: 20. 1817; Salix
scouleriana Barratt ex Hook. Fl. Bor.-amer. 2: 145. 1838.
Salix sect. Fulvae Barratt
Salix bebbiana Sarg., Gard. & For. 8: 463.1895.
Salix sect. Phylicifoliae (Fries) Andersson
Salix drummondiana Barratt ex Hook. Fl. bor.-am. 2: 144.
1838; Salix pedunculata Fernald, Rhodora 28: 188. 1926; Salix
pellita Andersson, Monogr. Salicum 139. 1867; Salix planifolia
Pursh, Fl. Am. Sept. 2: 611. 1814; Salix pulchra Cham. Linnaea
6: 543. 1831; Salix tyrrellii Raup, J. Arnold Arb. 17: 231. 1936
Salix sect. Arbuscella Ser.
Salix arbusculoides Andersson, Monogr. Salicum 147.
1867.
Salix sect. Candidae C. K. Schneider
Salix candida Flugge ex Willd. Sp. Pl. 4: 708. 1806;
Salix wiegandii Fernald, Rhodora 35: 243. 1933 (Treated here as
a species of hybrid origin, S. calcicola ? S. candida.).
Salix sect. Lanatae (Andersson) Koehne
Salix calcicola Fernald & Wiegand, Rhodora 13: 251.
1911; Salix richardsonii Hook. Fl. bor.-amer. 2: 147; Salix
tweedyi (Bebb ex Rose) C. R. Ball in J. M. Coulter, Bot. Gaz.
40: 377. 1905 (Basionym: Salix barrattiana Hook. var. tweedyi
Bebb ex Rose, Contr. U. S. Natl. Herb. 3: 572. 1902).
Salix sect. Villosae (Andersson) Rouy
Salix alaxensis (Andersson) Coville, Proc. Wash. Acad.
Sci. 2: 280. 1900 (Basionym: Salix speciosa Hook. & Arn. [var.]
alaxensis Andersson in DC. Prodr. 16(2): 275. 1868); Salix
alaxensis var. alaxensis; Salix alaxensis var. longistylis
(Rydb.) C. K. Schneider, J. Arnold Arb. 1: 225. 1919 (Basionym:
Salix longistylis Rydb. Bull. N. Y. Bot. Gard. 2: 163. 1901);
Salix barrattiana Hook. Fl. bor.-amer. 2: 146. 1838; Salix
silicicola Raup, J. Arnold. Arb. 17: 236. 1936.
Salix sect. *Viminella Ser.
*Salix viminalis L. Sp. Pl. 1021. 1753.
Salix sect. *Canae A. Kerner
*Salix elaeagnos Scop. Fl. Carn. 2 ed. 2: 257. 1772.
Salix sect. Argyrocarpae Fernald
Salix argyrocarpa Andersson, Monogr. Salicum 107. 1867.
Salix sect. Geyerianae Argus
Salix cana Mart. & Gal. Bull. Acad. Roy. Sci. Bruxelles
10: 344. 1843 (This seems to be the proper section for this
taxon according to usage, but the type specimen has not yet been
studied.); Salix geyeriana Andersson, Ofvers. Forh. Kongl.
Svenska Vetensk.-Akad. 15: 122. 1858; Salix lemmonii Bebb in S.
Watson, Bot. California 2: 88. 1879; Salix petiolaris Sm. Trans.
Linn. Soc. 6: 122. 1802.
Salix sect. Mexicanae C. K. Schneider
Salix irrorata Andersson, Ofvers. Forh. Kongl. Svenska
Vetensk.-Akad. 15: 117. 1858; Salix lasiolepis Benth. Pl. Hartw.
335. 1857; Salix lasiolepis var. lasiolepis; Salix lasiolepis
var. bigelovii (Torr.) Bebb in S. Watson, Bot. Calif. 2: 86.
1879 (Basionym: Salix bigelovii Torr. Pacific Railway Reports 7:
139. 1857); Salix mexicana Seemen, Bot. Jahrb. 21 (52): 9. 1896;
Salix riskindii M. C. Johnston, Madrono 28: 150. 1981.
Salix sect. Brewerianae C. K. Schneider
Salix breweri Bebb in S. Watson, Bot. Calif. 2: 88. 1879
Salix sect. Griseae (Borrer) Barratt ex Hook.
Salix sericea Marshall, Arbust amer. 140. 1785
Salix sect. Sitchenses (Bebb) C. K. Schneider
Salix delnortensis C. K. Schneider, J. Arnold. Arb. 1:
96. 1919; Salix jepsonii C. K. Schneider, J. Arnold Arb. 1: 89.
1919; Salix sitchensis Sanson ex Bong. Mem. Acad. St. Petersb.
6. 2: 162. 1833.
Salix sect. *Helix Dumort.
*Salix purpurea L., Sp. pl. 2: 1017. 1753.
George W. Argus
R.R.3-310 Haskins Rd.
Merrickville, Ontario
Canada K0G 1N0
argus@sympatico.ca
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I apologize to Dr. Argus and to all of you who receive BEN via
e-mail for the loss of formatting and special characters in this
article. Adolf Ceska
For the definitive version of this contribution, please consult
BEN archives at
http://www.ou.edu/cas/botany-micro/ben/ben227.html
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