370104 Papaver dahlianum Nordh.
Distribution
Northern Fennoscandia: Rare
Svalbard - Franz Joseph Land: Frequent
Polar Ural - Novaya Zemlya: Frequent
Yamal - Gydan: Scattered
Taimyr - Severnaya Zemlya: Frequent
Anabar - Onenyo: Presence uncertain
Central Canada: Frequent
Hudson Bay - Labrador: Scattered
Ellesmere Island: Frequent
Western Greenland: Rare
Eastern Greenland: Rare
Polar desert: Frequent
Northern arctic Tundra: Frequent
Mid Arctic Tundra: Frequent
Southern Arcti Tundra: Scattered
Shrub Tundra: Rare
Bordering boreal or alpine areas: Rare
- Nordh., Bergens Mus. Årbok 1931, Naturv. r. 2: 46 (1932). Lectotype (O!): Norway: Finnmark, Båtsfjord, Syltefjord, Østerelven, 04. July 1930, leg. R. Nordhagen (Elven and Nilsson in Jonsell 2001a: 521).
- Papaver radicatum subsp. brachyphyllum Tolm., Svensk Bot. Tidskr. 21: 82 (1927). Syntypes (S!): Norway: Finnmark (Varanger Peninsula), Tana: "Troldfjord (oder Gulgofjord) in Tana", 16. July 1920, leg. Th. Svedberg fl. albi; Båtsfjord: "Syltvikfjord", 1857, leg. A. Landmark fl. flavi.
- ?Papaver lapponicum subsp. dasycarpum Tolm., Trudy Bot. Muz. 25: 101 (1932). Described from Novaya Zemlya (European Russia). Not lectotypified.
- ?Papaver lujaurense N. Semenova, Fl. Murmansk. Obl. 3: 369 (1956). Holotype (LE!): European Russia: the Murman area, "Paeninsula Kola, in montibus Lovoserskye-Tundry, prope pag. Revda", 25. Aug. 1955, leg. N. Semenova-Tjan-Schanskaja 185.
2n=
70 (10x). - Europe (N), Canada (N), Greenland (NE). - At least five reports, as "dahlianum" and "polare". Decaploid in FCM, Solstad (2009, two plants from two localities).
Some of the chromosome counts may refer to P. cornwallisense. The decaploid FCM estimates are made on plants confirmed as P. dahlianum by molecular means.
Not included: Reports of 2n = 70 (10x) from northern Greenland (Jørgensen et al. 1958), Melville Island in northern Canada (Mosquin and Hayley 1966, as P. radicatum), and Löve (1962b), and a report of 2n = 84 (12x) from northern Greenland (Jørgensen et al. 1958). These counts could belong to either P. dahlianum or P. cornwallisense, the dodecaploid count most probably to the latter. For reports of 2n = 42 (6x, six counts) from Wrangel Island and West Chukotka and 2n = 56 (8x, two counts) from Wrangel Island (Zhukova and Petrovsky 1985a), see notes.
Geography: North American (N) - amphi-Atlantic - European (N) - Asian (NW): NOR RUS SIB CAN GRL.
Notes: Papaver dahlianum has a predominantly arctic, broadly amphi-Atlantic range reaching west to the western Canadian Arctic Archipelago and east to Taimyr. We have no molecular support for presence in northeastern Siberia or Beringia, as proposed by Rändel (1974) and by most Russian authors (e.g., Tolmachev 1975a; Bezdeleva 1987; Petrovsky 1999).
Nordhagen (1932) described P. dahlianum from the Varanger Peninsula in northeastern mainland Norway, where a small population group is isolated from the main range in the Arctic. Nordhagen's P. dahlianum corresponds with Tolmachev's (1927a) P. radicatum subsp. brachyphyllum, also described from the Varanger Peninsula where only one species of Papaver is present.
The majority of northwestern European authors have included both the mainland Norwegian and the more widespread arctic plant in P. dahlianum, whereas Russian authors have kept them separate and have applied the name P. polare for the arctic plants. There are some minor morphological differences, summarized by Nilsson (2001b), and some minor differences in isoenzymes (Solstad et al. 2003). Knaben (1959b) found pollen fertility of 90% and seed development of 95% in crosses between two mainland populations of P. dahlianum, pollen fertility of less than 80% and seed fertility of 60% in crosses between mainland and Svalbard populations. For comparison, pollen and seed fertility in eight different crosses between P. radicatum (several 'subspecies') and P. dahlianum had means of, respectively, 43% and 36%, at levels with her other crosses between taxa at species level. These crosses are among decaploid (2n = 70) plants. All evidence from morphology, hybridization, and molecules assign the arctic plants to P. dahlianum (and/or P. cornwallisense) and not to P. radicatum (Knaben 1959a, 1959b; Solstad 1998, 2009; Solstad et al. 1999, 2003; Nilsson 2001b). Acceptance of two subspecies of P. dahlianum - a mainland Norwegian subsp. dahlianum and a high arctic subspecies (proposed as subsp. polare by Egorova 1998; Elven and Nilsson in Jonsell 2001a; Nilsson 2001b; Elven et al. 2005) - is not supported by the AFLP data. The mainland Norwegian plant joins in molecular markers in a dense cluster with plants from Svalbard and Franz Joseph Land (Solstad 2009). The morphological distinctions of the mainland populations can be due to rapid postglacial divergence in an isolated group of small and fluctuating populations.
Papaver lujaurense is a local species of the central Murman area. Both the type and the other available specimens (LE) indicate that it should be included in P. dahlianum, an option considered also by Egorova (1998). The morphological differences between P. lujaurense and the Varanger Peninsula plants of P. dahlianum (ca. 350 km distant) are not more than could be expected among small, marginal, and semi-isolated populations of a species.
Papaver lapponicum subsp. dasycarpum, described from Novaya Zemlya (Tolmachev 1932c), is more problematic to assign. It combines features of P. lapponicum and P. dahlianum. However, the material we have studied (LE) deviates from P. lapponicum subsp. lapponicum and subsp. jugoricum (see below) and approaches P. dahlianum in most characters. This is also evident from the significant part of subsp. dasycarpum (in LE) that Tolmachev himself first annotated as P. polare and later transferred to subsp. dasycarpum.
Papaver dahlianum is probably an allopolyploid including several different parental genomes. Among the more than thirty Papaver species analysed for AFLP markers, P. dahlianum has the most scored fragments (more than 50) and shares these with a broad span of other species.
North European chromosome counts of unambiguous P. dahlianum are all decaploid. The few FCM measures are most likely decaploid as well, but decaploids and dodecaploids could be difficult to separate with FCM. Whether other ploidy levels are present in this species is undecided. The few dodecaploid counts reported from northern Greenland and northern Canada may belong to either P. dahlianum or P. cornwallisense. The plants reported with lower ploidy levels from northeastern Asia probably belong to other species in this affinity, perhaps P. "murrayi".
Higher Taxa
- Papaver [3701,genus]