1
[00] 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.
Comments:
(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.
Comments:
(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).
Comments:
(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 backchecked 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)
46.1.1.1 Claytonia tuberosa Pall. ex Roem. & Schult. var. tuberosa
S
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
Comments:
2
46.1.1.2 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.
S
T
[Described from N Chukokta: Cape Schmidt.]
2n=
30 (6x).
2nD
Zhukova (11967 2 counts, N. Chuk].
G
RFE ALA
Comments:
(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)
Comments:
(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
S
T
2n=
2nD
Claytonia acutifolia Pall. ex Roem. & Schult. (1819), Syst. Veg. 5: 436.
C. acutifolia Pall. ex Roem. & Schult. subsp. acutifolia.
[Described from Kamtchatka.]
(1) 30 (6x, x=5). (2) 32 (4x, x=8).
(1) Sokolovskaya (1963 Kamtch). (2) Sokolovskaya (1963 Kamtch); Zhukova (1966 C
Chuk); Zhukova & Petrovsky (1977 W Chuk); Zhukova (1980 S Chuk).
G
SIB RFE
Comments:
(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
Comments:
(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);
3
blade gradually transforms into dilatated membranaceous sheath (whereas even narrowleaved 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)
Comments:
(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.
S
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
Comments:
(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
S
2n=
2nD
Claytonia joanneana Roem. & Schult. (1819), Syst. Veg. 5: 434.
C. arctica Adams var. joanneana (Roem. & Schult.) Trautv. (1877), *** 5, 1: 56.
(1) 16 (2x, x=8). (2) 20 (4x, x=5).
(1) Malachova (1971 S Sib); Krasnoborov (1976 S Sib). (2) Krogulevich (1971 S Sib);
Zhukova et al. (1973 S Sib).
4
G
SIB
Comments:
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
Comments:
46.1.7
S
T
2n=
2nD
Claytonia sarmentosa C.A. Mey. (1829), Nouv. Mém. Soc. Naturalistes Moscou 1: 137.
Montia sarmentosa (C.A. Mey.) Robins. in A. Gray (1897), Synopt. Fl. N. Amer. 1, 2: 272.
[Described from Alaska: St. George Isl., leg. Eschscholtz.]
10 (2x).
Löve & Löve (1975) list two counts, one Russian as arctic; Zhukova & Tikhonova (1971 E
Chuk).
G
RFE ALA CAN
Comments:
(1)
Counts of 2n=40 (8x) refer to C. udokanica Zuev (1990, Bot. Zhurn. 75, 1: 99), a transBaicalian 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).
G
ALA?
Comments:
(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).
Comments:
5
(1)
(2)
46.2.1
B
S
T
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)
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)
Claytoniella vassilievii (Kuzen.) Jurtz. (1972), Bot. Zhurn. 57: 645.
Claytonia vassilievii Kuzen. in Kom. (1936), Fl. SSSR 6: 383, 880.
Montiastrum vassilievii (Kuzen.) Ö. Nilsson (1967), Grana Palyn. 7: 355, 360.
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.
G
RFE
Comments:
(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 AlaskaYukon 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
6
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)
46.2.1.1 Claytoniella vassilievii (Kuzen.) Jurtz. subsp. vassilievii
S
2n=
64 (8x, x=8).
2nD
Zhukova (unpubl. NE Sib, Pekulnei Range, 6 counts).
G
RFE
Comments.
46.2.1.2 Claytoniella vassilievii (Kuzen.) Jurtz. subsp. petrovskyi M. Grichuk & Jurtz. (1974), nom.
nud., Problems of Botanical Geography of Northeastern Asia 111-112.
S
2n=
60 (12x, x=5).
2nD
Zhukova & Petrovsky (1971 Wrangel I., 10 counts).
G
RFE
Comments:
(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.
6220'N, 13730'W, 1933, leg *** (N 19) (CAN) holotype.
2n=
2nD
G
ALA
Comments:
(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.
Comments.
(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)
7
46.3.1
S
2n=
2nD
Montia fontana L. (1753), Sp. Pl. 87.
M. lamprosperma Cham. (1831), Linnaea 6: 564.
(1) 18 (2x, x=9). (2) 20 (4x, x=5).
(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)