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52.1.5 Papaver multiradiatum Petrovsky (1983), Bot. Zhurn. 68: 234.
S
T
Russia: Insula Wrangelii, litus meridionale, ad sinum Somnitelnaja, 23.07.1965, leg.
V.V. Petrovsky (LE) holotype.
2n=
42 (6x).
2nD
Zhukova & Petrovsky (1985 NE As).
G
RFE
Comments:
(2)
A part of the P. radicatum aggregate in Petrovsky (1999, draft). A narrow endemic of
Wrangel Island. There is need for some more extensive morphological and
experimental work before its status and affinity can be decided. Only tentatively
accepted. (Elven)
WARNING! Only tentatively included.
52.1.6 Papaver detritophilum Petrovsky (1985), Bot. Zhurn. 70: 114.
S
T
Russia: Terra Tschuktschorum occidentalis, jugum Anjujensis, in systemate fl. Anjuj
Magnus, in valle fl. Bystrjanka, 25.07.1980, leg. V.V. Petrovsky 80-59 (LE)
holotype.
2n=
42 (6x).
2nD
Zhukova & Petrovsky (1985 NE As).
G
RFE
Comments:
(1)
A part of the P. radicatum aggregate in Petrovsky (1999, draft). An endemic of West
and East Chukotka. Its relationships to Alaskan taxa and to P. radicatum must be
more extensively investigated, morphologically and experimentally, before its status
and final affinity can be decided. Only tentatively accepted. (Elven)
WARNING! Only tentatively included.
The Papaver lapponicum aggregate (chionophilum, hultenii, ?hypsipetes, lapponicum,
minutiflorum, schamurinii, variegatum)
Comments:
(1)
Papaver lapponicum itself (subsp. lapponicum as defined here) is very close to P.
radicatum in genetic markers (see the papers by Solstad et al. referred to above). It is
also close morphologically as demonstrated by the confusion between the two in
Greenland and North America. It is not at all obvious that they should be separated on
two aggregates (or even on two species). The genetic differences between both these
two and the P. dahlianum aggregate are more evident (contrary to the taxonomic
treatments of Rändel 1974 and partly also Kiger & Murray 1997). If we believe in the
evidence of the isoenzymes (as we don't fully do yet), the differences between P.
lapponicum and P. radicatum are less than the variation normally found within
species. (Elven)
52.1.7 Papaver lapponicum (Tolm.) Nordh. (1932), Bergens Mus. Årb. 1931, Naturv. Rekke
2: 45-46.
B
P. radicatum Rottb. subsp. lapponicum Tolm. (1923), Bot. Mater. Gerb. Glavn. Bot.
Sada RSFSR 4, 11-12: 86.
S
T
Russia: Peninsula Kola, Oz. Imandra, 01.08.1911, leg. R. Pohle (LE) lectotype
selected by Egorova (1998, Novosti Sist. Vyssh. Rast. 31: ***). The Pohle lectotype
selected by Löve (1962, Taxon 11: 133) is not unambiguous and he does not seem to
have seen it before selecting it.
Comments:
(1)
The concept of P. lapponicum depends strongly on which concept of P. radicatum is
applied, see above. It is a pity, but unavoidable, that the typification (as also in P.
dahlianum) is made on an isolated, marginal population that deviates from the main
body of the species. In the opinion of Petrovsky (1999, draft) - which we share subsp. lapponicum is restricted to the mountain (and valley) populations of N Norway
and Kola Peninsula where it occurs in three separate areas: (A) The Kvænangen area
in NE Troms province (Norway), where the populations was first described as a
separate var. parviflorum Nordh. (1932), Bergens Mus. Årb. 1931, Naturv. Rekke 2:
46, later proposed as a separate subsp. kvaenangense (Lundström) Nilsson (in draft),
based on P. nudicaule L. subsp. kvaenangense Lundström (1923), Acta Horti Berg. 7:
416. (B) The Alta area in W Finnmark province (Norway), where the populations
were first described as a separate subsp. scandinavicum Knaben (1959), Opera Bot. 2,
3: 56, and later raised to species level as P. norvegicum N. Semen. in Pojark. (1956),
Fl. Murm. Obl. 3: 369. (C) The Chibiny Mts. in central Kola Peninsula from where it
is typified and where previous Russian botanists have separated the material on three
species: P. lapponicum s. str., P. chibinense N. Semen. in Pojark. (1956), Fl. Murm.
Obl. 3: 368, and P. tolmatchevii N. Semen. in Pojark. (1956), Fl. Murm. Obl. 3: 369.
Those three last-mentioned are nearly entirely sympatric, each with 15-30 mapped
'occurrences' within a common area of about 40 x 40 km (Pojarkova 1956, maps 102104). The three Kola Peninsula 'species' are results of the previous 'nonotypic' species
concept and are joined within subsp. lapponicum by Egorova (1998).
An octoploid chromosome number is proved for both Norwegian population
groups (Horn 1938, Knaben 1959).
The investigations of genetic markers have until now been restricted to
material from the Alta populations (subsp. scandinavicum sensu Knaben, subsp.
lapponicum sensu Egorova, Petrovsky and us). Very few markers differentiate this
octoploid from the decaploid 'races' of P. radicatum. From genetic evidence P.
lapponicum subsp. lapponicum could be included in P. radicatum. From a
geographical point of view, too, the N Fennoscandian populations fall inside or just
outside the range of P. radicatum whereas they are well isolated from the rest of what
is named as P. lapponicum in both directions. Until further evidence is available we
should keep P. lapponicum together but we should be aware of the possibility that the
future 'species' might exclude the type subspecies (with some nomenclatural
repercussions) or perhaps be merged with P. radicatum.
Subsp. lapponicum, which was included in Petrovsky's draft, is excluded here
as it does not reach or even closely approach the Arctic as defined. (Elven)
52.1.7.1 Papaver lapponicum (Tolm.) Nordh. subsp. dasycarpum Tolm. (1932), Trav. Mus.
Bot. Acad. Sci. URSS 24: 101.
S
2n=
2nD
G
RUS
Comments:
(1)
According to Tolmachev (1975), this is a local race of Novaya Zemlya where it cooccurs with P. dahlianum subsp. polare. The material deviates distinctly from P.
lapponicum subsp. lapponicum and approaches P. dahlianum in some characters.
This is evident from the significant part of it that Tolmachev annotated as P. polare
(in LE) and which later has been transferred to P. lapponicum subsp. dasycarpum.
Petrovsky (1999) also writes, "This subspecies probably also grows in several islands
of the Siberian sector of the Arctic Ocean", but it is not clear where. The entity needs
experimental comparison with both P. dahlianum subsp. polare and more southern P.
lapponicum s. lat. before it can be fully accepted. (Elven)
WARNING! An ambiguous entity as it stands, see comment.
52.1.7.2 Papaver lapponicum (Tolm.) Nordh. subsp. jugoricum (Tolm.) Tolm. (1932), Trav.
Mus. Bot. Acad. Sci. URSS 24: 101.
B
P. radicatum Rottb. subsp. jugoricum Tolm. (1923), Bot. Mater. Herb. Bot. Gard. 4,
11-12: 86.
S
2n=
56 (8x).
2nD
Zhukova & Petrovsky (1985, 1987 N Russ).
G
RUS SIB
Comments:
(1)
The chromosome counts refer to P. lapponicum subsp. jugoricum from Polar Ural.
(Petrovsky)
(2)
A race of Polar Ural and northwesternmost Siberia (Yamal-Gydan) with a slight
overlap with the next in the Jenisei area. This entity also needs a critical comparison
with neighboring races and species before it can be fully accepted. The split of the
NW Russian and Fennoscandian material of P. lapponicum s. lat. may be an effect of
a less continuous distribution here than elsewhere and be a fairly recent (postglacial)
phenomenon. The results from P. radicatum should warn us against over-description
in such situations. (Elven)
WARNING! An ambiguous entity as it stands, see comment.
52.1.7.3 Papaver lapponicum (Tolm.) Nordh. subsp. orientale Tolm. (1932), Tr. Polar.
Komiss. 13: 131.
S
2n=
(1) 42 (6x). (2) 56 (8x).
2nD
(1) ? (2) Zhukova (1968 ***); Zhukova & Petrovsky (1972 ***).
G
SIB RFE ALA CAN
Comments:
(1)
Considered by Petrovsky (1999, draft) as the major race. Distributed from the Jenisei
area eastwards through Siberia, Alaska and Canada, however, in the text (1999) given
as "West American". Petrovsky's map (1999) also totally excludes Greenland from
the range of P. lapponicum. The problems with this subspecies are therefore mainly
on the North American side.
(A) The 'labradoricum' problem. The relations to Knaben's P. lapponicum
subsp. labradoricum in W Greenland and NE Canada is not made clear in the draft as
the name is missing from Petrovsky's synonymy. The 'labradoricum' entity is
probably included in P. radicatum in Petrovsky's concept of that species (see above).
Kiger & Murray (1997) include the 'labradoricum' entity in their P. radicatum subsp.
radicatum, but that is based on the probably erroneous Greenlandic typification of P.
radicatum. If there is anything at all of importance in capsule shape and leaf
dissection, subsp. labradoricum belongs in P. lapponicum (see Knaben 1959a, Pl. 6).
If the two 'subspecies' are merged, the name 'orientale' 1932 has priority before
'labradoricum' 1958 at level of subspecies. In view of the confusion between P.
lapponicum and P. radicatum, it must be confirmed to which of the 'species' the
'labradoricum' entity should belong.
(B) The 'occidentale' problem. See also P. radicatum. The major body of
non-Dahlianum Canadian plants were placed by Knaben (1959a, 1959b) in her P.
lapponicum subsp. occidentale and her investigations showed it to be fairly
homogeneous as poppies go (see Knaben 1959a, Pl. 5). Her experiments proved that
crosses between Norwegian P. lapponicum subsp. lapponicum and Greenlandic
'occidentale' had c. 90% pollen fertility (the same level as between the two
neighboring Norwegian areas of subsp. lapponicum) and c. 50% good seed (the same
level as between many crosses between Norwegian races of P. radicatum).
As far as experimental and morphological evidence goes, the 'occidentale'
entity is therefore a P. lapponicum as long as subsp. lapponicum is kept inside P.
lapponicum. As I commented under P. radicatum, there might be two major North
Canadian entities besides P. dahlianum, but the 'occidentale' entity then belongs to
the 'lapponicum' part of the variation and not the 'radicatum' part. It should therefore
probably be compared with and perhaps considered synonymous with P. lapponicum
subsp. orientale which then should be extended to include a large part of the
Greenlandic plants. (Elven)
52.1.7.4 Papaver lapponicum (Tolm.) Nordh. subsp. porsildii (Knaben) Á. Löve ***
S
2n=
56 (8x).
2nD
Knaben (1959a, 1959b N Am).
G
ALA CAN
Comments:
(1)
Included in P. radicatum subsp. radicatum (sensu Americans) by Kiger & Murray
(1997). Knaben's (1959b) very scant crossing experiments involving this taxon
involved subsp. labradoricum and what she called a 'intermediate type' from
Greenland, both inside her concept of P. lapponicum. Both crosses gave fairly low
pollen and seed fertility, at the level of crosses between species elsewhere in her
experiments. Capsule (Knaben 1959a, Pl. 6) and leaf shape indicate 'lapponicum'
rather than 'radicatum' in the North European concept. I am inclined to accept it as an
American race but then its relations to the Russian subsp. orientale should be looked
into. Knaben's material of what she named as subsp. porsildii was mainly from Prince
Charles Island in the Foxe Basin, very far east and within the reach of both subsp.
labradoricum and subsp. occidentale. (Elven)
52.1.8 Papaver variegatum Tolm. (1975), Arct Fl. URSS 7: 20.
S
T
Russia: N Siberia: The Putorana plateau: ad litus meridionalis lacu Khantaika, prope
ostium fluminis Mogaddy, 07.08.1970, leg. A. Tolmachev (LE) holotype.
2n=
42 (6x).
2nD
***
G
SIB
Comments:
(1)
A fairly local entity of the Putorana plateau and southern Taimyr. Information
insufficient for full evaluation. Petrovsky (1999) states that: "P. varigatum should be
considered a subspecies of P. lapponicum, since it is only weakly distinguished from
P. lapponicum s. str. Plants of similar appearance used to be referred to P.
lapponicum s. l." I think we should follow that and include it in P. lapponicum, but
that we perhaps should look at the total variation in what will become P. lapponicum
(after the limit between P. lapponicum and P. radicatum is decided) before we assign
rank. A comment is needed in any case. (Elven)
WARNING! Might be reduced to a subspecies of P. lapponicum or to synonymy.
52.1.9 Papaver minutiflorum Tolm. (1960), Bot. Mater. Herb. Bot. Inst. Ac. Sc. USSR 20:
180.
S
T
Russia: Yakutia, in regione alpina jugi montium Verchojanski, in ditione fluminis
Tempo, 05.07.1957, leg. I. Kildjuschevski (LE) holotype.
2n=
2nD
G
SIB RFE
Comments:
(1)
A disjunct entity known from the Kharaulakh area (northern Yakutia) and southern
Chukotka. Information insufficient for final evaluation. Petrovsky (1999) states:
"Papaver minutiflorum may only represent a small-flowered variety of P. lapponicum
subsp. orientale." Its disjunct occurrence indicates the same and small-flowered
plants of P. lapponicum are known from several areas, partly as population groups.
Rändel (1974) was of a similar opinion [translated from German through Norwegian]:
"As can be evaluated from the available specimens P. indirkense and P. minutiflorum
have the morphological characteristics of P. lapponicum and can be included in P.
lapponicum". Perhaps just synonymy, with a comment. (Elven)
WARNING! Might be reduced to synonymy within P. lapponicum subsp. orientale.
52.1.10 Papaver hultenii Knaben (1959), Opera Bot. 3, 3: 49.
S
T
USA: Alaskan Copper Mine River *** (O) holotype, see comment.
2n=
42 (6x).
2nD
Löve & Löve (1975) list several arctic counts from NW America and NE Asia under
this name.
G
RFE ALA
Comments:
(1)
The type specimen is a plant raised from seed and cultivated in the Botanical Garden,
University of Oslo. Maybe P. hultenii is the Alaskan form (variety) of P. lapponicum
subsp. occidentale. (Petrovsky)
(2)
Kiger & Murray (1997) synonymize P. hultenii with P. lapponicum in a fairly
restricted sense which excludes the 'occidentale' entity. There might, however, be a
Beringian race in this complex but Petrovsky (1999) rather interprets P. hultenii as
part of P. lapponicum subsp. orientale. We should perhaps follow that, with a
comment. (Elven)
WARNING! Will probably be reduced to synonymy within P. lapponicum.
52.1.11 Papaver schamurinii Petrovsky (1985), Bot. Zhurn. 70: 116.
S
T
Russia: Insula Wrangelii, sinus Somnitelnaja, ad litus lacunae Bazovaja, 12.07.1971,
leg. V.V. Petrovsky 71-200 (LE) holotype.
2n=
42 (6x).
2nD
Zhukova & Petrovsky (1985 NE As).
G
SIB RFE ALA CAN
Comments:
(1)
Mapped by Petrovsky (1999) from three different areas: Taimyr, Wrangel Island
(typification) and N Alaska, in the text also mentioned for the Canadian Arctic
Archipelago but not included from here in the map. It is mentioned for North
America in a comment under P. lapponicum by Kiger & Murray (1997). The text of
Petrovsky (1999) indicates that he also could accept this as a race (subspecies) of P.
lapponicum. (Elven)
WARNING! Might be reduced to a race of P. lapponicum, or perhaps to synonymy.
52.1.12 Papaver chionophilum Petrovsky (1983), Bot. Zhurn. 68: 233.
S
T
Russia: Insula Wrangelii, ad sinum Somnitelnaja, ad fl. Somnitelnaja, 05.08.1971,
leg. V.V. Petrovsky (LE) holotype.
2n=
56 (8x).
2nD
Zhukova & Petrovsky (1985 NE As).
G
RFE
Comments:
(1)
A local endemic of Wrangel Island. Petrovsky (1999) indicates it as possibly a
variation within P. lapponicum subsp. orientale, but as octoploid in opposition to the
otherwise hexaploid P. lapponicum (subsp. orientale) on Wrangel Island. However,
the majority of P. lapponicum elsewhere (Greenland, Canada, western Russia,
Norway) is also octoploid so this argument is not very strong. (Elven)
WARNING! Might be reduced to a race of P. lapponicum or to synonymy.
52.1.13 Papaver hypsipetes Petrovsky (1985), Bot. Zhurn. 70: 113.
S
T
Russia: Terra Tschuktschorum occidentalis, districtus Bilibinskij, montes Anjujensis,
jugum Hirnejensis, ad lacus Hirnej Superior, 18.07.1973, leg. V.V. Petrovsky 73-26
(LE) holotype.
2n=
28 (4x).
2nD
Zhukova & Petrovsky (1985 NE As).
G
RFE
Comments:
(1)
A local endemic of West Chukotka mountains. The information is insufficient for
final decision. As it is the most low-ploid member of the P. lapponicum aggregate
(tetraploid) it should not be included in the otherwise similar subsp. orientale without
careful evaluation and some experimental work. (Elven)
52.1.14 Papaver alaskanum Hultén (1937), Fl. Aleut. Isl. 190.
S
P. radicatum Rottb. subsp. alaskanum (Hultén) J.P. Anders. (1959), Fl. Alaska 244;
P. microcarpum DC. subsp. alaskanum (Hultén) Tolm. (1975), Fl. Arct. URSS 7: 30.
2n=
42 (6x).
2nD
Mulligan & Porsild (1969b NW Can).
G
ALA
Comments:
(1)
Without studying type material a relation of P. alaskanum to P. radicatum or P.
microcarpum is not clear. (Petrovsky)
(2)
As long as we don't have available the data needed to make a decision - as subspecies
of P. radicatum (as preferred by Kiger & Murray 1997), subspecies of P.
microcarpum (as preferred by Tolmachev 1975), or as separate species - we have to
keep it as species. Tolmachev's proposal should perhaps be taken as an indication that
he saw the P. microcarpum entities as comparatively closely related to P. radicatum?
(Elven)
(3)
The material annotated as P. alaskanum or P. radicatum subsp. alaskanum from
Alaska in ALA is heterogeneous. A re-evaluation of this entity is urgently needed
before full acceptance and assignment to aggregate. (Elven & Solstad)
The Papaver pulvinatum aggregate (?angustifolium, anjuicum, leucotrichum, mcconnellii,
nivale, pulvinatum)
Comments:
(1)
The circumscription of this aggregate is still very unclear. Petrovsky (1999) included
P. macounii subsp. discolor and P. angustifolium with a question mark. As he there
only listed the Russian taxa, we also have to include the Americans P. mcconnellii
and P. macounii subsp. macounii, the latter as questionable. Petrovsky (1999)
excluded P. keelei from this complex but that is a little problematic as Kiger &
Murray (1997) synonymised P. keelei with P. macounii subsp. discolor. They also
stated that specimens of P. paucistamineum from Chukotka fell inside the range of
variation they accepted for P. macounii subsp. discolor. I have tentatively made a
change from Petrovsky (1999, draft) by transfer of P. macounii and P. keelei to an
extended P. macounii - paucistamineum aggregate.
The problems seem at least partly to be linked with the current concept of P.
macounii. This 'species' includes a mainly Beringian Sea Island large-flowered plant
(subsp. macounii) that resembles the central parts of the P. pulvinatum aggregate and
a mainland amphi-Beringian but mainly American and much more small-flowered
plant (subsp. discolor) that much more resembles Petrovsky's (1999) P. keeleipaucistamineum aggregate. Kiger & Murray (1997) also - by an elegant
understatement - indicated doubts about whether subsp. macounii and subsp. discolor
really belong within the same species by stating that P. keelei would be the valid
species name if subsp. discolor was accepted as a species of its own.
(2)
Very, very tentative results from some isoenzyme investigations in Oslo
(Solstad unpubl.) might support a separation of subsp. macounii and subsp. discolor.
However, the two plants become more similar in comparative cultivation than they
are in field-collected material; 'discolor' becomes very large-flowered and 'macounii'
becomes tall-grown.
We should therefore discuss the possible re-structuring of the aggregates
where the P. pulvinatum aggregate would contain P. pulvinatum, P. leucotrichum (as
a race of P. pulvinatum?), P. angustifolium (perhaps only as a comment), P.
anjuicum, P. mcconnellii, P. nivale, and P. macounii s. str. (possibly); the P.
paucistamineum aggregate would contain P. paucistamineum, P. atrovirens, P.
macounii subsp. discolor, and perhaps as a separate entry: P. keelei. (Elven)
The map of the taxa of the aggregate presented by Petrovsky (1999) does not
correspond with his text. There must have been an interchange of legends between P.
angustifolium and P. leucotrichum, and P. pulvinatum subsp. interius is not in the
map. (Elven)