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BEN # 227

Adolf Ceska aceska at VICTORIA.TC.CA
Mon Jul 5 21:44:46 EST 1999

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No. 227                              July 5, 1999

aceska at victoria.tc.ca                Victoria, B.C.
 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2

            BEN # 226 and BEN # 277 are dedicated to


     Curator Emeritus of the Royal British Columbia Museum,
              Victoria, British Columbia, Canada,
      on the occasion of his 80th birthday, July 2, 1999.

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

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

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-

        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.

        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.

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 at sympatico.ca

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
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