This draft has not been discussed




Дата канвертавання27.04.2016
Памер48.74 Kb.
67  Violaceae (Draft: N.N. Tzvelev)

THIS DRAFT HAS NOT BEEN DISCUSSED

67.1  Viola L. (1753), Sp. Pl. 933.

Comments:

(1) Tzvelev's draft includes three subgenera and in the first subgenus five sections. As there are about 13 species, a subgeneric classification is acceptable. The three subgenera are too coarse a division as all except two species belong to only one subgenus (Viola). Use of sections only is problematic because sectional affinity of the two last species is not indicated. I have therefore tentatively kept all of Tzvelev's subgeneric entities. (Elven)
Subg. Viola
Sect. Trigonocarpaea Godr. (1857), Fl. Lorr., ed. 2, 1: 88.
67.1.1  Viola riviniana Rchb. (1823), Iconogr. Bot. Pl. Crit. 1: 81.

S

2n= 40 (8x).



2nD Löve & Löve (1975) list numerous counts, one Icelandic.

G ICE


Comments:

(1) Added to Tzvelev's draft. It reaches the Arctic in Iceland, acc. to Kristinsson's list and maps. Nordal & Jonsell (1998) have convincingly documented that the basic chromosome number of this group is x=5. (Elven)


67.1.2  Viola canina L. (1753), Sp. Pl. 935.

S

Comments:



(1) Added to Tzvelev's draft; see subspecies. The two subspecies are more or less generally accepted. They are parapatric and clearly different in some few features (e.g., shape of lamina, shape and size of stiples) when well separated geographically. Where they meet, as they do over most of Fennoscandia, they are almost impossible to keep apart. It may be that nearly all of C Fennoscandia is a transition zone. (Elven)
67.1.2.1  Viola canina L. subsp. canina

S

2n= 40+0-5B (8x).



2nD Lövkvist & Hultgård (1999 S Sweden); **Vigdis?**

G ICE?


Comments:

(1) Tentatively added to Tzvelev's draft. It is uncertain whether subsp. canina reaches the Arctic in Iceland. It is mapped as common by Hultén & Fries (1986) but not separated from subsp. montana by Kristinsson in his maps. In his list, Kristinsson gives subsp. canina as certainly arctic in Iceland whereas subsp. montana is indicated as a border case. This is opposite to the relative distribution of the two subspecies in Fennoscandia. (Elven)



WARNING! Needs to be confirmed from Iceland before acceptance in the list.
67.1.2.2  Viola canina L. subsp. montana (L.) Hartman (1841), Bot. Not. 1841: 82.

B V. montana L. (1753), Sp. Pl. 935.

S

2n= 40 (8x).



2nD Löve & Löve (1975) list five counts. one Icelandic.

G ICE? NOR RUS GRL

Comments:

(1) Added to Tzvelev's draft as it reaches the Arctic in S Greenland, N Norway, N Russia (Kola Peninsula) and perhaps in Iceland. (Elven)


67.1.3  Viola rupestris F.W. Schmidt (1791), Abh. Königl. Böhm. Ges. Wiss., ser. 2, 1: 60.

S

Comments:



(1) A polymorphic and genetically heterogeneous species in Europe. Two of the entities are comparatively well differentiated, both morphologically and genetically ('rupestris' s. str. and 'relicta') and these are also allopatric. A third entity proposed as a subspecies in Tzvelev's draft ('glaberrima') is here reduced to synonymy of the first subspecies, see comment below. (Elven)
67.1.3.1  Viola rupestris F.W. Schmidt subsp. rupestris

S ?V. rupestris F.W. Schmidt subsp. arenaria (DC.) Tzvelev in Tolm. (1980), Fl. Arct. URSS 8: 33; V. rupestris F.W. Schmidt subsp. glaberrima (Murb.) V. Nikit. in Tzvelev (1996), Fl. Vost. Evr. 9: 190 [basionym: V. rupestris F.W. Schmidt var. glaberrima Murb. (1887), Bot. Not. 1887: 186]; V. glaberrima (Murb.) C. Serg. (1961), Bot. Mater. (Leningrad) 21: 283, non House (1906).

2n= 20 (4x).

2nD ?


G RUS SIB

Comments:

(1) Murbeck's Scandinavian 'glaberrima' falls, morphologically and genetically, within the variation pattern of N European subsp. rupestris and should not be accepted as a separate race (Nordal & Jonsell 1998, Bot. J. Linn. Soc. 128: 105-122). (Elven)
67.1.3.2  Viola rupestris F.W. Schmidt subsp. relicta Jalas (1950), Ann. Bot. Soc. Zool.-Bot. Fenn. 'Vanamo' 24, 1: 70.

S

2n=



2nD

G NOR


Comments:

(1) A morphologically and genetically distinct Fennoscandian race which reaches the Arctic in North Norway (Nordkapp). Recently studied by Nordal & Jonsell (1998). (Elven)


67.1.4  Viola adunca J.E. Sm. in Rees (1817), Cycl. 37, no. 63.

S V. labradorica Schrank (1818), Denkschr. Bot. Ges. Regensb. 1, 2: 12; V. adunca J.E. Sm. var. minor (Hook.) Fern. ***

T [Described from the west coast of N America.]

2n=


2nD

G CAN GRL

Comments:

(1)
Sect. Arcion Juz. in Kom. (1949), Fl. SSSR 15: 437.


67.1.5  Viola langsdorffii Fisch. ex DC. (1824), Prodr. 1: 296.

S

T [Described from Alaska: Unalaska.]



2n= c. 120 (24x?).

2nD Taylor & Mulligan (1968).

G RFE ALA

Comments:


Sect. Violidium (C. Koch) Juz. in Kom. (1949), Fl. SSSR 15: 408.

B Subg. Violidium C. Koch (1841), Linnaea 15: 251.


67.1.6  Viola selkirkii Pursh ex Goldie (1822), Edinb. Philos. J. 6: 324.

S

T [Described from Canada: near Montreal.]



2n= 24 (2x? 4x?).

2nD Löve & Löve (1975) list five non-arctic counts.

G ALA? CAN GRL

Comments:

(1) Strongly approaches our Arctic also in SW Alaska, Kuskokwim Mts. (Elven)
67.1.7  Viola pallens (Banks) Brainerd (1905), Rhodora 7: 247.

B V. rotundifolia Michx. var. (b) pallens Banks in DC. (1824), Prodr. 1: 295.

S V. macloskeyi Lloyd var. pallens (Banks) Hitchc. ***

2n=


2nD

G CAN


Comments:
67.1.8  Viola renifolia A. Gray (1870), Proc. Amer. Acad. 8: 288.

S V. renifolia A. Gray var. brainerdii (Greene) Fern. *** [basionym: V. brainerdii Greene (1902), Pittonia 5: 89; described from Canada, near Ottawa].

2n=

2nD


G CAN

Comments:

(1) The arctic and near-arctic material is usually named as var. brainerdii. The differential characters are weak (glabrous or glabrescent vs. permanently villous) and the 'variety' occurs more or less throughout the range of the species. This is not a variety as we think it. I propose that we reduce it to synonymy as done here. (Elven)
Sect. Plagiostigma Godr. (1857), Fl. Lorr., ed. 2, 1: 90.

Comments:

(1) If the sections are omitted from the checklist, this is a natural V. palustris aggregate. (Elven)

(2) It is difficult to reconcile Tzvelev's treatment and Nordic treatments and experiences. We have two generally accepted entities in the Nordic area. One of them is tetraploid (2n=48), comparatively little varied, and corresponds with the Linnaean V. palustris. The other is diploid (2n=24), more varied, and is customarily named V. epipsila by us. The two hybridise frequently. The hybrids are mostly triploid (2n=36) but numbers between 20 and 48 have been found. Some good seed-production is observed but the majority is seed-sterile. They may, however, occur in large clones due to vegetative spread. We don't recognise these hybrids as a taxon.

There has been a fairly extensive combined investigation of morphology and cytology (ploidy) in Norway and we are fairly sure about these facts and ploidy levels of both parents and hybrid. The parents are largely sympatric in our area but with different ecological demands. They behave very well as species. **REFS**

Tzvelev indicates three species for Fennoscandia and NW Russia: V. palustris with 2n=48, V. epipsila with 2n=40, and V. epipsiloides with 2n=24. If these chromosome counts really relate to these taxa (or rather names) it is clear that we mean the same when we talk about V. palustris but that the current Nordic application of the name 'V. epipsila' compares with the current Russian application of 'V. epipsiloides'.

The strange chromosome numbers of 2n=40 are listed by Löve & Löve (1975) from seven sources. We need to go back to the original papers to check whether the Löves really refer the exact numbers of these authors: Clausen (1926) treated V. arvensis and V. tricolor; Clausen (1927) the genus in general; Clausen (1931) subgenus Melanium, i.e., the V. tricolor group; Löve & Löve (1956) the flora of Iceland where both 2n=24 and 2n=48 entities occur; Sokolovskaya & Strelkova (1960, 1962) northern Russia where both 2n=24 and 2n=48 entities occur; and Sorsa (1965) referred the aberrant number from Finland in connection with hybridisation between V. epipsila and V. palustris. It is therefore at least probable that all records of 2n=40 either are mentioned only in passing or refer to hybrids.

It will be impossible to decide whether Ledebour's V. epipsila, described in a list from the botanical garden of Dorpat (Tartu), had 2n=24 or 2n=40 (or 2n=48). No justification is given by the Löves for their identification of the 2n=40 plants with Ledebour's V. epipsila. It might, however, be possible to decide with some certainty whether an eventual type specimen of V. epipsila belongs to the 'species' or to the hybrid between the two N European species. It would not be very nice to have to reject the name V. epipsila for the species that has gone by that name throughout Eurasia for 180 years.

Tzvelev in Tolmachev & Yurtsev (1980, Fl. Arct. URSS 8) and in his draft synonymises V. episiloides with V. epipsila subsp. repens. This is the race or entity accepted for most of W Canada, Alaska, NE Asia and Siberia. It should, from the ploidy levels and from Tzvelev's distributional draft, be the same as the Nordic plant. It is not. As Löve & Löve (1976) based the description of their V. epipsiloides on V. repens Turcz. ex Trautv. & C.A. Mey., Turczaninov's opinion decides the meaning of V. epipsiloides. It is described from S Siberia (Baikal area and Dahuria) and most probably represent the Siberian   NW American entity.

In the present state of knowledge, the most reasonable interpretation is that we have a widely amphi-Atlantic tetraploid species (V. palustris) and a Eurasiatic and NW American diploid species (V. epipsila) in 2-3 races, i.e., subspecies. The draft and chromosome information has been changed accordingly. (Elven)


67.1.9  Viola palustris L. (1753), Sp. Pl. 934.

S

2n= 48 (4x? 8x?).



2nD Löve & Löve (1975) list numerous counts, one Greenlandic and one Icelandic. **Oslo-oppgave!**

G ICE NOR RUS CAN GRL

Comments: Subsp. palustris in the Arctic.
67.1.10  Viola epipsila Ledeb. (1820), Ind. Sem. Horti Dorpat. 5.

S

Comments:


67.1.10.1  Viola epipsila Ledeb. subsp. epipsila

S

2n= 24 (2x? 4x?).



2nD Sorsa (1968 Finl); **OSLO**

G ICE NOR RUS SIB

Comments:
67.1.10.2  Viola epipsila Ledeb. subsp. repens (Turcz. ex Trautv. & C.A. Mey.) W. Becker (1917), Bot. Centralbl. 34, 2: 406.

B V. repens Turcz. ex Trautv. & C.A. Mey. in Middend. (1856), Reise Sibir. 1, 2, 2: 18, non Schwein. (1822).

S V. epipsiloides Á. Löve & D. Löve (1976), Bot. Not. 128: 516.

2n= 24 (2x? 4x?).

2nD Löve & Löve (1975) list six counts, four as arctic, that with some certainty refer to this subspecies.

G SIB RFE ALA CAN

Comments:
67.1.10.3  Viola epipsila Ledeb. subsp. palustroides W. Becker (1921), Feddes Repert. 17: 72.

S V. palustroides (W. Becker) Tzvel. in Tolm. & Jurtz. (1980), Fl. Arct. URSS 8: 36.

2n=

2nD


G RFE

Comments:

(1) A fourth entity in the V. palustris affinity needs some very good justification before it can be accepted. I also wonder about the relations between this narrow Russian Far East plant and the V. achyrophora Greene (1902), Pittonia 5: 33, described from Pribilof Islands: St. Paul, 18.07.1897, leg. Macoun (CAN) holotype. Hultén (1947 p. 1130) accepted it but later (Hultén 1968) included it in the synonymy of subsp. repens. (Elven)

WARNING! In need of good justification before accepted.
Sect. Dischidium Ging. (1823), Mem. Viol. 28.
67.1.11  Viola biflora L. (1753), Sp. Pl. 936.

S

2n= For the collective species. 12 (2x).



2nD Löve & Löve (1975) list numerous counts, two Russian as arctic.

Comments:

(1) Most arctic plants probably belong to subsp. biflora. However, I am not at all certain that the Beringian entity (seen in Alaska 1998 and treated as subsp. biflora in Tzvelev's draft) is the same as the Atlantic entity. (Elven)

(2) Tzvelev proposes three subspecies. They are all listed below, but with some doubts. More documentation is needed. (Elven)



WARNING! Subspecies need some additional justification before they can be fully accepted, see comments.
67.1.11.1  Viola biflora L. subsp. biflora

S

2n=



2nD

G NOR RUS SIB RFE ALA

Comments:
67.1.11.2  Viola biflora L. subsp. microceras (Rupr.) Tzvel. comb. nov. proposed.

B V. microceras Rupr. (1845), Distr. Pl. Vasc. Ross. 22; id. (1846), Symb. Pl. Ross. 22, 223.

S

2n=


2nD

G RUS


Comments:

(1) Acc. to Tzvelev (1996, Fl. Eur. Orient. 9), this is a local plant of Kolguyev Isl. We need some more information about how it differs from other northern marginal populations of the species before we accept it. (Elven)


67.1.11.3  Viola biflora L. subsp. avatschensis (W. Becker & Hultén) Tzvel. in Tolm. & Jurtz. (1980), Fl. Arct. URSS 8, 1: 39.

B V. avatschensis W. Becker & Hultén (1928), Ark. Bot. 22A, 3: 1.

S

2n=


2nD

G RFE


Comments:
Subg. Chamaemelanium (Ging.) Juz. in Kom. (1949), Fl. SSSR 15: 146.

B Sect. Chamaemelanium Ging. (1823), Mém. sur la Fam. Viol. 1.


67.1.12  Viola uniflora L. (1753), Sp. Pl. 936.

S

2n= (1) 12 (2x). (2) 24 (4x).



2nD (1) ? (2) ?

G SIB?


Comments:

(1) A border case in the Dudinka area, lower Jenisei. (Elven)


Subg. Melanium (Ging.) Peterm. (1846), Deutschl. Fl. 65.

B Sect. Melanium Ging. (1823), Mem. Viol. 28.

Comments:

(1) Tzvelev includes in his draft only V. arvensis Murray (1770), Prodr. Stirp. Gött. 73, from this subgenus - as adventive in Polar Ural and W Greenland. In the latter case it is obviously ephemeral and I assume it is the same in Polar Ural. It has therefore been omitted. However, V. tricolor is stable in the Arctic and has been added. (Elven)


67.1.13  Viola tricolor L. (1753), Sp. Pl. 935.

S V. tricolor L. subsp. faeroeensis W. Becker ***

2n= 26 (2x).

2nD Löve & Löve (1956b Icel); Lövkvist & Hultgård (1999 S Sweden).

G ICE NOR*

Comments:



(1) A border case as presumed native in Iceland; more or less stable as introduced in arctic Norway. Löve & Löve (1975) treated all the 'arctic', N Atlantic plants as a subsp. faeroeensis but this subspecies needs much additional justification before general acceptance. (Elven)


База данных защищена авторским правом ©shkola.of.by 2016
звярнуцца да адміністрацыі

    Галоўная старонка