Panarctic Flora

370136 Papaver radicatum subsp. radicatum Rottb.

• Rottb., Skr. Kiøbenhavnske Selsk. Lærd. Elsk. 10: 455 (1770). Lectotype: Rottbøll, Skr. Kiøbenhavnske Selsk. Lærd. Elsk. 10: t. VIII, No. XXIV, right illustration. 1770. (Knaben 1958: 62); epitype (UPS!): Iceland, Bardastrandarsýsla, Brjánslækur, 23. July 1962, leg. J.A. Nannfeldt 17564 (Nilsson and Elven in Jonsell 2001a: 520).
• Papaver radicatum subf. islandicum C.E. Lundstr., Acta Horti Berg. 7, 5: 411 (1923). Type (C?): Iceland: Gufudalsháls, leg. S. Stefánsson. - Papaver nordhagenianum subsp. islandicum (C.E. Lundstr.) Á. Löve, Nytt Mag. Bot. 4: 16 (1955). - Papaver radicatum subsp. stefanssonii Á. Löve, Ízlenzk. Jurtir: 149 (1945) nom. nud.. - Papaver nordhagenianum subsp. stefanssonii (Á. Löve) Á. Löve, Taxon 11: 137 (1962). comb. illeg.. - Papaver stefanssonianum Á. Löve, Nytt Mag. Bot. 4: 14 (1955). Holotype (ICEL): Iceland: Gufudalsháls, 08. Aug. 1893, leg. S. Stefánsson and O. Davidsson. See note.
• Papaver nudicaule subsp. relictum C.E. Lundstr., Acta Horti Berg. 7, 5: 415 (1923). Lectotype (S!): Norway: Oppland: Vang (Vasendlifjeld), 15. Aug. 1870, leg. Söderén and Eisen (Elven and Nilsson in Jonsell 2001a: 521). - Papaver relictum (C.E. Lundstr.) Nordh., Bergens Mus. Årbok 1931, Naturv. r. 2: 45 (1932).
• Papaver nordhagenianum Á. Löve, Nytt Mag. Bot. 4: 15 (1955) nom. illeg.. Holotype (UPS): Sweden: Torne lappmark, Nissontjåkko, leg. H. Smith (see Nordhagen 1932: 30, fig. 13).
• We also synonymize with this species (and subspecies) the numerous other names for non-arctic and decaploid races described from Iceland, the Faeroes, and Scandinavia (see Nilsson 2001b): subspp. avkoënse Knaben (northern Norway), dovrense Nordh. (central Norway), faeroënse C.E. Lundstr. (the Faeroes), gjaerevollii Knaben (central Norway), groevudalense Knaben (central Norway), hyperboreum Nordh. (northern Scandinavia), intermedium (Nordh.) Knaben (southern Norway), macrostigmum Nordh. (northern Norway), oeksendalense Knaben (central Norway), ovatilobum Tolm. (central Norway), steindorssonianum (Á. Löve) Knaben ex Ö. Nilsson (Iceland), and subglobosum Nordh. (northern Norway).

2n= 70 (10x). - Europe (N). - At least five reports and more than 40 counts. Decaploid in FCM, Solstad (2009, one plant).
Not included: Reports under the name P. radicatum that with certainty belong to other species, e.g.: 2n = 42 (6x) from the Yukon Territory (Mulligan and Porsild 1969b, perhaps P. kluanense) and Chukotka (Zhukova and Petrovsky 1985); 2n = 56 (8x) from Prince Patrick and Ellesmere islands in northern Canada (Mosquin and Hayley 1966, probably P. lapponicum subsp. occidentale), western Greenland (Dalgaard 1988, P. lapponicum s. lat. or P. labradoricum), and Chukotka (Zhukova 1965); and 2n = 70 (10x) from northern Canada (Mosquin and Hayley 1966, P. cornwallisense or P. dahlianum).

Geography: European (NW): ICE.

Notes: We follow Knaben (1958, 1959a, 1959b, and elsewhere) in recognizing Papaver radicatum as a European North Atlantic species restricted to Iceland, the Faeroes, and Scandinavia. We include all described decaploid races in subsp. radicatum, whereas we accept an octoploid (Horn 1938; Solstad 2009, one plant in FCM) subsp. laestadianum Nordh., Bergens Mus. Årbok 1931, Naturv. r. 2: 49 (1932) [P. laestadianum (Nordh.) Nordh., Bot. Not. 1939: 623 (1939)] from a few mountains northern Scandinavia. Subspecies laestadianum and Scandinavian subsp. radicatum do not reach the Arctic, whereas Icelandic subsp. radicatum does.

The currently accepted type for the name P radicatum is from Iceland (Solstad et al. 1999, Nilsson and Elven in Jonsell 2001a). There has been a long discussion concerning the type, whether from Iceland or from Greenland (and then belonging to current-day P. lapponicum or P. labradoricum), see Hultén (1945a: 805-806) and Löve (1955, 1962a, 1962b) advocating a Greenland type and Knaben (1958) and Knaben and Hylander (1970) advocating an Icelandic one. The Hultén-Löve approach was followed by, e.g., Tolmachev (1975a), Kiger and Murray (1997), and Petrovsky (1999), making the usage of both names P. radicatum and P. lapponicum ambiguous until 2000 or later.

The results from morphology, cytology, and molecular data (isozymes, RAPD-DNA, AFLP) all support Knaben's view of a geographically restricted P. radicatum. The alternative is to merge P. radicatum and P. lapponicum as one species (under the former priority name), with several subspecies and subsp. radicatum as one of these. Still, this merged species would be absent from the Beringian regions in Asia and North America.

A single Icelandic-Scandinavian race, subsp. radicatum, finds support in the molecular data. Icelandic plants were found identical with Scandinavian plants in isozymes and RAPD-DNA markers (Solstad 1998; Solstad et al. 1999, 2003), and Icelandic plants from all parts of the country where the species occurs (east, north, northwest, west) were closely connected to southern and central Norwegian plants in AFLP markers (Solstad 2009). The plants are also generally similar in morphology but the 12-15 described subspecies differ somewhat, mostly by combinations of quantitative characters (Nordhagen 1932; Knaben 1959a, 1959b; Selin and Prentice 1988; Selin 2000; Nilsson 2001b). All these subspecies have small and isolated or semi-isolated ranges supporting inbreeding. The investigations of Solstad (1998, 2009) and Solstad et al. (1999, 2003) throw doubts on their relevance as races. Only some small molecular variation was found in isozymes and RAPD-DNA markers and that within proposed subspecies and partly within populations. No difference whatsoever was found between the subspecies. The AFLP markers closely connected Icelandic and southern and central Norwegian populations and races. This collective cluster was more uniform than found for most other similarly investigated species (and subspecies) of Papaver sect. Meconella. Northern Scandinavian plants (previously as P. radicatum subsp. hyperboreum and subsp. macrostigmum, and the octoploid subsp. laestadianum) constituted another cluster in AFLP markers with some distinctions among the mentioned entities. However, it will be difficult to recognize this cluster and its entities as one or more separate races on morphological grounds.