Amaranthaceae (Draft: A.A. Korobkov)
Amaranthus L. (1753), Sp. Pl. 989, with Amaranthus retroflexus L. (1753), Sp. Pl. 991, is reported by Korobkov as 'rare-adventive' in Taimyr and as 'rare' in the Hudson Bay area. It is very improbable that any Amaranthus is stable and reproduce anywhere in the Arctic. No Amaranthus is able to produce mature seeds in the boreal or even the boreonemoral zones. If there is no very strong evidence of stability, the species and family must be excluded from the list. (Elven)
46 Portulacaceae (Draft: B.A. Yurtsev)
DRAFT HAS NOT BEEN DISCUSSED SERIOUSLY WITH ANYONE.
(1) This draft is left almost unchanged except for a few comments and some editorial 'shuffling'. (Elven)
46.1 Claytonia L. (1753), Sp. Pl. 204.
(1) Three sections are proposed in Yurtsev's draft. As there only are eight species, these are omitted: sect. Cormosae A. Gray (C. tuberosa), sect. Caudicosae A. Gray (C. acutifolia, C. eschscholtzii, C. arctica, C. joanneana, C. porsildii), and sect. Rhizomatosae A. Gray (C. sarmentosa, C. scammaniana). (Elven)
46.1.1 Claytonia tuberosa Pall. ex Roem. & Schult. (1819), Syst. Veg. 5: 436.
S C. czukczorum Volkova (1960), Bot. Mater. Gerb. Bot. Inst. AN SSSR 20: 139; C. tuberosa Pall. ex Roem. & Schult. var. czukczorum (Volkova) Hultén (1967), Ark. Bot., ser. 2, 7, 1: 49.
T [Described from Kamtchatka: Tigil.]
2n= For the collective species. 30 (6x, x=5).
2nD Zhukova (1966, 1967a 2 counts from N Chuk); Mulligan et al. in Löve (1972c).
(1) Hultén (1967) states that only a few spurious characters separate C. czukczorum from C. tuberosa. He consequently reduced the former to a variety of the latter. He also reported plants with these charactes from the Alaskan side (Little Diomede Isl., Cape Prince of Wales). The reported difference in chromosome numbers is strange in this context but should be back-checked against the sources as the references are taken from Löve & Löve (1975). (Elven)
(2) There is a gradual transition between C. tuberosa and what was described as C. czukczorum Volkova connected with the depth of corms and presence / absence of moss carpet: rosette leaves penetrate the moss cover separately each sometimes; they have long etiolated petioles and are not collected as usual. So, the status of variety seems more appropriate. The chromosome count 2n=16 (Johnson & Packer 1968 for the westernmost Alaska) needs confirmation: in the Chukotkan material both types have identical chromosome numbers. (Yurtsev)
188.8.131.52 Claytonia tuberosa Pall. ex Roem. & Schult. var. tuberosa
2n= (1) 16 (2x, x=8). (2) 30 (6x, x=5).
2nD (1) Johnson & Packer (1968 Ala). (2) Zhukova (1980 2 counts, S Chuk).
G SIB RFE ALA CAN
184.108.40.206 Claytonia tuberosa Pall. ex Roem. & Schult. var. czukczorum (Volkova) Hultén (1967), Ark. Bot., ser. 2, 7, 1: 49.
B C. czukczorum Volkova (1960), Bot. Mater. Gerb. Bot. Inst. AN SSSR 20: 139.
T [Described from N Chukokta: Cape Schmidt.]
2n= 30 (6x).
2nD Zhukova (11967 2 counts, N. Chuk].
G RFE ALA
(1) Probably the ecotype of the arctic coasts of Chukotka and Alaska. (Yurtsev)
(2) Perhaps not worthy of separate rank?
WARNING! The variety might perhaps be reduced to synonymy.
The Claytonia graminifolia aggregate (C. acutifolia, C. eschscholtzii)
(1) Hultén (1944) expressed doubts about the identity between subsp. graminifolia and C. eschscholtzii but fully synonymised them later (Hultén 1967). The separation on two taxa with different ploidy levels, as done by Löve & Löve (1975), is therefore very dubious. If there are two taxa, corresponding to the plody levels, the tetraploid probably needs a new name. (Elven)
46.1.2 Claytonia acutifolia Pall. ex Roem. & Schult. (1819), Syst. Veg. 5: 436.
S C. acutifolia Pall. ex Roem. & Schult. subsp. acutifolia.
T [Described from Kamtchatka.]
2n= (1) 30 (6x, x=5). (2) 32 (4x, x=8).
2nD (1) Sokolovskaya (1963 Kamtch). (2) Sokolovskaya (1963 Kamtch); Zhukova (1966 C Chuk); Zhukova & Petrovsky (1977 W Chuk); Zhukova (1980 S Chuk).
G SIB RFE
(1) It was not found in Alaska. The count 2n=30 needs confirmation. (Yurtsev)
(2) Yurtsev reports it from 'BOa 13' in his draft, i.e. the Beringian Sea islands. This, in combination with his comment above, must mean that it is found in Big Diomede but not in Little Diomede or Seward Peninsula. True? (Elven)
46.1.3 Claytonia eschscholtzii Cham. (1831), Linnaea 6: 561.
S C. acutifolia Pall. ex Roem. & Schult. subsp. graminifolia Hultén (1944), Lunds Univ. Åsskr., n. f., avd. 2, 40, 1: 634.
T [Described from Russian Far East: St. Lawrence Bay. Subsp. graminifolia is typified from Alaska: Nome, leg. Anderson 3747 (LD) holotype?
2n= 32 (4x).
2nD Johnson & Packer (1968 Ala); Zhukova in Yurtsev (1980 E Chuk).
G RFE ALA
(1) The record of 2n=16 for C. eschscholtzii (Zhukova 1966) in reality belongs to [the non-arctic] C. soczaviana. (Yurtsev)
(2) Strictly amphi-Beringian tetraploid (2n=32) arctic taxon, replaced in mountainous areas of subarctic NE Siberia and southern E Siberia by the diploid (2n=16) C. soczaviana Jurtz. In Chukotka, C. eschscholtzii is spread only in the eastern part of Chukchi Peninsula, so the species is predominantly American.
In eastern Chukchi Peninsula C. eschscholtzii is sympatric with C. acutifolia and does not show any transitional forms. It replaces C. acutifolia (an acidophilous plant) on basic rocks, first of all on limestones. The most remarkable differences are traced in structure and shape of leaves which in C. eschscholtzii are almost linear and without petioles (sessile); blade gradually transforms into dilatated membranaceous sheath (whereas even narrow-leaved forms of C. acutifolia have a true petiole). Each individual of both taxa is identifiable to species, which does not justify the status of subspecies. Despite the similarity of C. eschscholtzii and C. acutifolia in shape of inflorescence, size, and white or pink colour of flowers, the closer relative of the former seems to be C. soczaviana.
Together with C. acutifolia, C. eschscholtzii and C. soczaviana belong to the group of claytonias with bracts at the base of the spike and a membranaceous sheath of leaves. Leaves are born in a monopodial rosette on taproot. The following three species lack bracts at the base of the inflorescence. (Yurtsev)
The Claytonia arctica aggregate (C. arctica, C. joanneana, C. porsildii)
(1) The Taimyr population, reminding C. arctica in general appearance, differs from the latter in wider, well-developed hyaline sheaths of leaves, pinkish or white petals (not yellow or white, yellow at base), blades of rosette leaves larger, lanceolate, acutish (not obovate) and was referred in the "Arctic Flora of the USSR" to the South Siberian alpine C. joanneana, which viewpoint is accepted here, despite the opposite opinion in the "Flora of Siberia".
Porsild (1974) records C. arctica from anumber of localities in North America from Ogilvie Mts. and Brooks Range westwards to Cape Lisburn. However, it becomes clear from the rexr and photograph that the Alaska Yukon plant differs from true C. arctica from E Siberia and Chukotka in pinkish petals, oblanceolate leaves of rosette, and stems nearly as long as rosette leaves. They were described as C. porsildii. Planst similar in many respects occur in the Aleutian Commander Islands as well as Central Kamtchatka, They were recorded as C. arctica by Hultén (1968) but not included in the latter in the "Arctic Flora of the USSR". One should check the record of C. arctica from St. Lawrence I. by Young (1971). (Yurtsev)
46.1.4 Claytonia arctica Adams (1817), Mém. Soc. Nat. Moscou 5: 94.
T [Described from Siberia: Lower Lena R., probably the mountainous right bank where the type yellow-flowered form is common.]
2n= 10 (2x).
2nD Zhukova (1966 Wrangel I.).
G SIB RFE
(1) If Claytonia arctica is synonymized with C. scammaniana, the 'arctica' name has priority at species level. In any case, we should therefore look for a valid name or combination in Montia. Hultén (1944) considered C. arctica very close to C./M. sarmentosa and not to M. scammaniana. (Elven)
(2) The idea of proximity (or at least relationship) of C. arctica to C. scammaniana Hultén (Montia scammaniana (Hultén) Welsh) has no morphological grounds. These are rather contrasting species, even in their growth form (one-headed monocephalous tap-rooted herb versus a herb with a filiform rootstock forming loose clumps).
The arctic-Yakutian populations are eurychoric montane tundra plants populating mesic and mesic-dry tundras and demonstrating significant variability. Whereas nthe Chukotka mainland populations (except for those from Wrangel I.) are rare and restricted mostly to screes or rubble ridges with sparse vegetation. (Yurtsev)
46.1.5 Claytonia joanneana Roem. & Schult. (1819), Syst. Veg. 5: 434.
S C. arctica Adams var. joanneana (Roem. & Schult.) Trautv. (1877), *** 5, 1: 56.
2n= (1) 16 (2x, x=8). (2) 20 (4x, x=5).
2nD (1) Malachova (1971 S Sib); Krasnoborov (1976 S Sib). (2) Krogulevich (1971 S Sib); Zhukova et al. (1973 S Sib).
46.1.6 Claytonia porsildii Jurtz. (1981), Bot. Zhurn. 66: 1043.
S C. arctica auct. amer., non Adams (1817)
T Canada: Yukon Terr.: Ogilvie Mts., near mile post 50-54 on Dempster Highway, 5800' a.s., 15.08.1966, leg. R.T. Porsild B447 (CAN) holotype.
2n= 10 (2x).
2nD Mulligan & Porsild (1969 Yukon Terr.).
G ALA CAN
46.1.7 Claytonia sarmentosa C.A. Mey. (1829), Nouv. Mém. Soc. Naturalistes Moscou 1: 137.
S Montia sarmentosa (C.A. Mey.) Robins. in A. Gray (1897), Synopt. Fl. N. Amer. 1, 2: 272.
T [Described from Alaska: St. George Isl., leg. Eschscholtz.]
2n= 10 (2x).
2nD Löve & Löve (1975) list two counts, one Russian as arctic; Zhukova & Tikhonova (1971 E Chuk).
G RFE ALA CAN
(1) Counts of 2n=40 (8x) refer to C. udokanica Zuev (1990, Bot. Zhurn. 75, 1: 99), a trans-Baicalian alpine octoploid endemic plant. Thus, the situation here is just opposite to that in the species pair C. soczaviana (alpine diploid) and C. eschscholtzii (arctic polyploid). (Yurtsev)
46.1.8 Claytonia scammaniana Hultén (1939), Bot. Not. 1939: 826.
S Montia scammaniana (Hultén) S.L. Welsh (1968), Great Basin Nat. 28: 154.
T C Alaska: Mt. Porcupine Dome, near Miller House, Steese Highway, 12.07.1937, leg. E. Scamman 749 (GH) holotype.
2n= 10 (2x).
2nD Mulligan & Porsild (1969b NW Can).
(1) This alpine subarctic species is probably absent from the Arctic, though almost reaches its southern boundary in Kuskokwim Mts. (see Hultén 1968). Indications of its presence in the lower reaches of Noatak R., W Alaska, in field report by S. Young (1975 unpubl.) shoudl be checked; it could well belong to C. porsildii. (Yurtsev)
(2) Ascribing chromosome number 2n=10 to C. scammaniana by Elven (in his drafts) is based on the identifying it with C. arctica s. lat. (incl. C. porsildii Jurtz.), which cannot be accepted. Record of 2n=10 in Zhukova (1966) certainly refers to the Wrangel I. population of C. arctica s. str. and the same count for the Ogilvie Mts. plant in Mulligan & Porsild (1969), to the type population of C. porsildii. (Yurtsev)
(3) Ascribing the chromosome number of 2n=10 to C. scammaniana was (and is) based on Cody's (1996) synonymisation of C. porsildii Jurtz. and C. arctica auct. amer (non Adams) with C. (Montia) scammaniana. This is still a valid viewpoint as it was shared by competent North American botanists like Welsh and Cody. (Elven)
46.2 Claytoniella Jurtz. (1972), Bot. Zhurn. 57: 644.
S Montiastrum auct., non (A. Gray) Rydb. (1917).
(1) The genus Claytoniella Jurtz. (1972) was included by Nilsson (1966, 1967) and Kubitzki et al. in Montiastrum (A. Gray) Rydb. (1917), Fl. Rocky Mts., ed. 1, 265, 1061. Yurtsev strongly opposes that solution. The case should be discussed further. (Elven)
(2) The plants much resemble Claytonia species in the appearance and size of inflorescence and to some extent in their ecology as alpine or arctic montane plants. However, the peculiar type of pollen grains (panto-colpate, with a tholus, spinulose at apex, in each hexagon) and some macromorphoilogical features (alternate stem leaves, cristate seeds) testify to the proximity of the alpine plants to the two annual vernal plants of hemiarid intermontane depressions in W North American Cordilleras, rather contrasting to the alpine plants in many other respects morphological, cytological (basic chromosome number), phenological, biological as well as phytogeographic. These numerous distinctions give serious grounds to separate the alpine amphi-Beringian allogamic perennial species at generic level as the genus Claytoniella Jurtz. The peculiar fragments of pollen grains of Claytoniella have kept fossil in many localities in NE Asia since late Pliocene and may serve as a stratigraphic indicator (Grichuk 1973, 1976, Yurtsev 1974). There are good reasons to consider Montiastrum s. str. as descendant of Claytoniella having persisted the glacial maxima south of the Cordilleran Ice Sheet and transformed at one of the interglacials into biological type of a vernal annual plant.
The chromosome numbers of Montiastrum s. str. are 2n=14 (2x, x=7) for M. dichotomum (Nutt.) Rydb. and 2n=28 (4x) for M. lineare (Dougl.) Rydb. In Claytoniella vassilievii three numbers have been counted: 2n=40 (8x, x=5), 60 (12x, x=5) and 64 (8x?, x=8?). The 2n=60 plants from Wrangel I. and the 2n=40 plants from the Ola plateau (Kolyma-Okhotsk divide) have some accompanying morphological and micro-morphological distinctions and were suggested to be described as a subspecies (subsp. petrovskii M. Grichuk & Jurtz.) or a species (C. kolymensis Jurtz.), but these still remain nomina nuda.
The fragments of fossil pollen grains of Claytoniella are easily identifiable and were recorded by 1973 from 23 localities in NE Asia (Grichuk 1976) from Late Pliocene up to Holocene. The range of the genus previously covered the interfluves of the Kolyma and Indigirka Rivers (i.e., the continental interior parts of Verkhoyan-Kolyma mountainous country) and W Chukotka. The earliest (Late Pliocene) pollen of Claytoniella (after Grichuk loc. cit.) combined some features of C. kolymensis and the diploid Montiastrum dichotomum. (Yurtsev)
46.2.1 Claytoniella vassilievii (Kuzen.) Jurtz. (1972), Bot. Zhurn. 57: 645.
B Claytonia vassilievii Kuzen. in Kom. (1936), Fl. SSSR 6: 383, 880.
S Montiastrum vassilievii (Kuzen.) Ö. Nilsson (1967), Grana Palyn. 7: 355, 360.
T Siberia: Anadyr area, Rarytkin Range, Mt. Irgunei, 18.08.1933, leg. V. Vassiliev (LE) holotype.
2n= For the collective species. (1) 40 (8x, x=5). (2) 60 (12x, x=5). (3) 64 (8x?, x=8?).
2nD (1) Yurtsev & Zhukova (1972, NE Sib, one count). (2) Löve & Löve (1975) list three Russian counts, all as arctic. (2-3) See below.
(1) The Ola basaltic Plateau race probably deserves status as a separate species, Claytoniella kolymensis Jurtz. (1974), nom. nud., Problems of Botanical Geography of Northeastern Asia 111-112. The plant is rather common on the plateau but was not met outside it. It could be considered as a subspecies of C. vassilievii, but some characters in common with the Alaska-Yukon plant C. bostockii pinkish petals, floriferous stems with several (vs. one) leaves, narrow basal part of leaves of creeping vegetative shoots are in favour of aquiring it the status of a species. It has the lowest known chromosome number among claytoniellas (2n=40, x=5), but we still do not know the chromosome number(s) of C. bostockii, the only American (also non-arctic) species and the presumed ancestral type of Claytoniella.
The Wrangel Island plant is very close to the type race in its morphology but differs in the chromosome number (60 vs. 64, basic number x=5 vs. 8). It also differs from the type race in greater size of pollen grains. It is also the only species of Claytoniella adapted to the clomate of zone B (Somnitelnaya Bay, rare at coast, more common in the mountains) and C (upper reaches of Neizvestnaya R.). All the other races of Claytoniella occur in zones D and E or in their altitudinal analogs. (Yurtsev)
220.127.116.11 Claytoniella vassilievii (Kuzen.) Jurtz. subsp. vassilievii
2n= 64 (8x, x=8).
2nD Zhukova (unpubl. NE Sib, Pekulnei Range, 6 counts).
18.104.22.168 Claytoniella vassilievii (Kuzen.) Jurtz. subsp. petrovskyi M. Grichuk & Jurtz. (1974), nom. nud., Problems of Botanical Geography of Northeastern Asia 111-112.
2n= 60 (12x, x=5).
2nD Zhukova & Petrovsky (1971 Wrangel I., 10 counts).
(1) In "Vascular Plants of Soviet Far East" Vol. 2 (Probatova & Rudyko 1987: 79) there is a wrong statement that the Ola Plateau (southern) race is named C. vassilievii subsp. petrovskyi. (Yurtsev)
46.2.2 Claytoniella bostockii (A.E. Porsild) Jurtz. (1972), Bot. Zhurn. 57: 645.
B Claytonia bostockii A.E. Porsild (1951), Bull. Natl. Mus. Can. 121: 160-161, pl. 17, 3, 4.
S Montiastrum bostockii (A.E. Porsild) Ö. Nilsson (1971), Bot. Not. 124: 92; Montia bostockii (A.E. Porsild) Welsh ***; Claytonia vassilievii auct., non ***.
T Canada: Yukon Terr., Dawson Range, south of Selkirk near source of Klaza R., approx. 6220'N, 13730'W, 1933, leg *** (N 19) (CAN) holotype.
(1) Metaarctic (arctic-alpine subarctic) species. The main area of distribution is easternmost Alaska and SW Yukon (high elevation areas), but with one locality on the northern main slope of Brooks Range near Toolik Lake (Murray & Lipkin 1987) in the Arctic.
The only species of Claytoniella with distinctively petiolate leaves, petioles being twice as long as the oblanceolate blades (see also comments above). The most resembling race in NE Asia grovs on the Ola Basaltic Plateau outside the Arctic and probably deserves recognition as a separate species, C. kolymensis Jurtz. ined. (Yurtsev)
46.3 Montia L. (1753), Sp. Pl. 87.
(1) Large, highly variable genus. Nevertheless, one cannot agree with the including of a number of species of Claytonia with 3-colpate pollen grains in it, as it was made by Welsh (1974) and accepted by some other botanists [e.g., Cody 1996]. This would much narrow the volume of the genus Claytonia at the expense of expanding that of Montia. Serious morphological ground for this have not been presented.
The type genus of subfam. Montioideae, including also the genus Claytonia and a number of mono and oligotypic genera (see Nilsson 1967). (Yurtsev)
46.3.1 Montia fontana L. (1753), Sp. Pl. 87.
S M. lamprosperma Cham. (1831), Linnaea 6: 564.
2n= (1) 18 (2x, x=9). (2) 20 (4x, x=5).
2nD (1) Veselukhina (1976 E Chuk). (2) Löve & Löve (1975) list several counts, four as arctic from W Chuk, W Ala, Grl and Icel.
G ICE NOR RUS SIB RFE ALA CAN GRL
Comments: Subsp. fontana in the Arctic.
(1) The reasons for treating M. lamprosperma Cham. [described from Alaska: Unalaska] as a synonym of M. fontana L. were presented by Moore (1963, Bot. Not. 116: 16-30) and confirmed by the monographic studies of Nilsson (1967).
The count of 2n=18 by Veselukhina should be checked against specimen. (Yurtsev)