52 Papaveraceae




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52  Papaveraceae (Drafts: V.V. Petrovsky [VP] & R. Elven [RE])

S Fumariaceae (incl.)

Comments:

(1) Some genera link Papaveraceae s. str. and Fumariaceae, e.g., Hypecoum. I am therefore reluctant to recognise the latter as a separate family in the checklist. (Elven)

(2) Polunin (1956) states that Chelidonium L. (1753), Sp. Pl. 505, with C. majus L. (1753), Sp. Pl. 505, is established in W Greenland. This is neither confirmed nor indicated in any Danish/Greenlandic survey and is extremely improbable. The species is obviously not part of the arctic flora. (Elven)

52.1  Papaver L. (1753), Sp. Pl. 506. [VP]

Comments:

(1) The treatment of the arctic-alpine Meconella (or Scapiflora) group of Papaver must be very tentative and preliminary, for several reasons. (a) There are discrepancies among Russian, North American and Nordic treatments and these partly build on different principles. Nordic treatments have, e.g., placed much emphasis on ploidy differences. These have not been given nearly the same weight in other areas. (b) The typification and circumscription of some of the most widespread entities has been and partly still is unclear. (c) Experimental studies have only been undertaken in the N Atlantic area and the results from the more recent of these do not correspond well with the currently applied taxonomy. (d) The fairly recent monography of the group (Rändel 1974) is based on very few characters. It reaches many conclusions that are unacceptable in the area where experimental studies have been undertaken. Its conclusions in other areas are, therefore, also to be doubted. (e) There may be some duplicate names across the Beringian political boundary. Few of the Asian taxa have traditionally been recognised on the American side, and vice versa.

The edited draft is therefore mostly unchanged from Petrovsky's draft as concerns the Russian taxa. Some changes have been made in the treatment of Nordic, Greenlandic and North American taxa. This implies an unwanted imbalance in the treatment. Some investigation of genetic markers are now undertaken also on material from Greenland (Nordal/Jonsell) and from Canada, Alaska and Siberia (Solstad/Elven). More results may be available for discussion   at least for the North American parts   fairly soon.

The pattern we begin to see is that there probably are only 2-3 major entities in the Atlantic area (very widely spoken) of the Arctic, that is, from Jenisei and Novaya Zemlya westwards through northern Europe, Greenland and Canada to the Mackenzie area. These entities are represented by the species names P. dahlianum, P. lapponicum and P. radicatum but the circumscription of each of them is disputed. The diversity increases dramatically when we cross Jenisei River eastwards in Siberia and Mackenzie River westwards in North America. The highest reported diversity is in NE Asia. The Central and North Asian mountain ranges are probably also the evolutionary center of the group (Rändel 1974 and many other sources).

The most we can hope for in the checklist stage is: (a) To define and circumscribe the three 'major' species as meaningfully as possible; and (b) To group the many taxa in the Siberian   Beringian area into more or less functionl aggregates. A very helpful survey here is Petrovsky (1999) and his aggregates are with few exceptions accepted in this draft.

Remember that my very long comments on some points are part of the discussion and not intended as comments in the final version. (Elven)
The Papaver dahlianum aggregate (calcareum, dahlianum, ?uschakovii)

Comments:

(1) The northern arctic and polar desert plants in all major areas seem to belong in this aggregate. Several names have been applied in different areas. Those most frequently used are: 'polare' (Russia, Canada), 'dahlianum' (Norway), 'radicatum' (in earlier times everywhere, recently mainly Greenland), and 'cornwallisense' (Canada). In the current draft, all the northernmost plants are considered to belong to one species and subspecies. Additional taxa probably occur in the Beringian area. (Elven)
52.1.1  Papaver dahlianum Nordh. (1932), Bergens Mus. Årbok 1931, Naturv. Rekke 2: 46.

S

T Norway: Finnmark, Syltefjord, Austerelva, 04.07.1930, leg. R. Nordhagen (O) lectotype, to be selected by Nilsson [& Elven?].



Comments:

(1) At least four names have been applied on species level within what here is considered as one species: P. dahlianum 1932, P. polare 1936, P. lujarense 1956 and P. cornwallisense 1956. Priority depends on whether one considers the southern marginal and morphologically somewhat deviant population groups as parts of the larger arctic species or not. The earliest species name   P. dahlianum   is based on such populations in Finnmark, northern mainland Norway, as is also P. lujarense from Kola Peninsula.



Papaver polare has not been unambiguously typified until recently (Egorova 1998) but is now typified on materials from Svalbard, within what at least since the 1940ies has been considered as P. dahlianum by Nordic botanists (Nordhagen 1940, Lid 1944, 1952, Knaben 1959a, 1959b, Nilsson in Jonsell in prep., Flora Nordica 2). Kiger & Murray (1997, Fl. N. Amer.: 329-330, 333) and others have been reluctant to accept this identity.

Recent investigations of genetic markers (isoenzymes, Solstad 1998, Solstad et al. 1999, 2000, in prep.) clearly indicate that there are differences between P. dahlianum/-polare in N Norway and Svalbard and the P. radicatum/-lapponicum groups in Norway and Iceland. There are also consistent morphological differences. Papaver dahlianum/polare should therefore not be reduced to a subspecies of P. radicatum as proposed (again) by Rändel (1974) and Kiger & Murray (1997, Fl. N. Amer. 3).

The most recent results also indicate that there are some smaller differences in isoenzymes and morphology between the mainland populations of P. dahlianum s. str. and the Svalbard populations ('polare'). This gives support to recognition as two subspecies, and that solution is proposed here. Some old results to remember: Knaben (1959b) found pollen fertility of 90% and seed developemnt of 95% in crosses between to mainland populations of P. dahlianum (subsp. dahlianum), pollen fertility of less than 80% and seed fertility of 60% in crosses between mainland (subsp. dahlianum) and Svalbard (subsp. polare) populations. Pollen and seed fertility in eight different cross combinations between P. radicatum and P. dahlianum had means of, respectively, 43% and 36%. This is additional support for the proposed solution. (Elven)
52.1.1.1  Papaver dahlianum Nordh. subsp. dahlianum

S P. radicatum Rottb. subsp. brachyphyllum Tolm. (1927), Svensk Bot. Tidskr. 21: 82; ?P. lujarense N. Semen. in Pojark. (1956), Fl. Murm. Obl. 3: 369.

2n= 70 (10x).

2nD Horn (1938 N Norw); Knaben (1959 N Norw). Both from arctic parts.

G NOR

Comments:



(1) Nordhagen (1932) described P. dahlianum at species level to replace Tolmachev's subsp. brachyphyllum (within P. radicatum). Nordhagen indirectly indicated a separate type so the two are not homotypic. I propose that we include the Tolmachev name in the synonymy as the first description of the taxon, even if it is not used in any current sources.

As to P. lujarense, the type and the other available specimens (LE) indicate that it should be included in P. dahlianum subsp. dahlianum (as also indicated by Egorova 1998). The morphological differences between the Kola Peninsula plants ('lujarense') and the Varanger Peninsula plants ('dahlianum' s. str.) are not more than can be expected among isolated and small marginal population groups. There are also differences among the several isolated Varanger populations. I therefore propose to include the name P. lujarense in the synonymy, to show our opinin, even if the name is absent from most current sources and the plants only occur somewhat south of our arctic boundary.

As a curiosity, there is an earlier typification of P. dahlianum, by Löve (1955? ****). One of the Varanger Peninsula populations is yellow-flowered, all the other known populations are white-flowered. Löve typified it on a white-flowered population (as did Tolmachev with his subsp. brachyphyllum). However, as Nordhagen in addition to 'typical' P. dahlianum also described a var. albiflorum, the type must be chosen among material from the single yellow-flowered population and there is a sheet (in O) indicated by Nordhagen as the type. (Elven)
52.1.1.2  Papaver dahlianum Nordh. subsp. polare (Tolm.) comb. nov. needed

B P. radicatum Rottb. subsp. polare Tolm. (1923), Bot. Mater. Gerb. Bot. Glavn. 4, 11-12: 87.

S P. polare (Tolm.) Perfiljev (1936), Flora of North Kray 1-2: 131; P. dahlianum Nordh. subsp. hadacianum (Á. Löve) Hada_ ***; P. cornwallisense D. Löve in Löve & Freedman (1956), Bot. Not. 109: 176.

T Svalbard: Spitzbergen, Advent-Bay, 05-30.07.1898, leg. Semenkevich (LE) lectotype, selected by Egorova (1998, Novosti Sist. Vyssh. Rast. 31: 101).

2n= (1) 42 (6x). (2) 56 (8x). (3) 70 (10x). (4) 84 (12x).

2nD (1-2) Zhukova & Petrovsky (1985 NE As). (3) Horn (1938 ***); Flovik (1940 Sb); Holmen (1952 ***); Knaben (1959a, 1959b Sb, N Am); Löve & Löve (1961d ***); Löve (1962b ***); Mosquin & Hayley (1966 Can). (4) Löve (1962b ***, as P. cornwallisense).

G NOR RUS SIB CAN GRL

Comments:

(1) Tolmachev (1923) did not indicate a type specimen. The areas he listed for subsp. polare are partly outside the range now accepted (e.g., Iceland where only P. radicatum is known) and the sheets annotated by him as 'polare' (in LE) also contains many collections of what now is considered as parts of P. lapponicum. These ambiguities had to be solved before the 'polare' name could be of any use at all. Egorova's (1998) typification confirms the usage which has developed in the times after Tolmachev and unambiguously connects the name to the common high-arctic plant as this is the only Papaver taxon known from Svalbard. We support this solution as it will make the name applicable throughout the range we accept today. (Elven)

(2) Papaver cornwallisense is accepted as a species in Petrovsky's draft but is here proposed reduced to synonymy. Its description was mainly based on the assumption of a constant ploidy level in each species of Papaver, a view very prevalent both by Knaben and by the Löves in all their works on Papaver. Papaver cornwallisense was published as dodecaploid (2n=84) whereas P. dahlianum/-polare at that time was known only as decaploid (2n=70).

This view of a constant ploidy level was never fully accepted by the Russians and the North Americans and is also just now weakening among Nordic botanists (see for P. radicatum below). Zhukova & Petrovsky (1985) have later reported lower ploidy levels (6x and 8x) of their P. polare from N Asia. The morphological differences reported by D. Löve between P. cornwallisense and P. dahlianum/-polare were small and insignificant compared with the large variation found in this species elsewhere. We have seen quite a lot of material from the area where P. cornwallisense was described (Cornwallis Island and surrounding high-arctic islands) and only find what we morphologically would name as P. dahlianum/-polare.

As to the dodecaploid number, Kiger & Murray (1997) write: "The report of 2n=84 (as P. cornwallisensis) attributed to G. Knaben (1959) by Á. Löve (1962b) cannot be confirmed." Papaver cornwallisense was reduced to synonymy by Kiger & Murray (1997) and I propose that we follow them. (Elven)

(1) As defined here, P. dahlianum subsp. polare has a nearly continuous distribution from Taimyr across NW Siberia, N Europe, N Greenland and Canada west to the western islands (Banks Island). (Elven)
52.1.2  Papaver calcareum Petrovsky (1983), Bot. Zhurn. 68: 232.

S

T Russia: Insula Wrangelii, ad fl. Gussinaja, 14.07.1969, leg. V.V. Petrovsky (LE) holotype.



2n= 42 (6x). 70 (10x). 84 (12x).

2nD Zhukova & Petrovsky (1985 NE As).

G RFE

Comments:



(1) Populations of P. calcareum needs careful study. (Petrovsky)

(2) A part of the P. dahlianum aggregate in Petrovsky (1999) and his draft. Both this species and the next are narrow endemics of Wrangel Island, isolated geographically from P. dahlianum elsewhere. There is need for more extensive morphological and some experimental work before their status and affinity can be decided. They are therefore only tentatively accepted. (Elven)



WARNING! Needs critical study before full acceptance.
52.1.3  Papaver uschakovii Tolm. & Petrovsky (1973), Bot. Zhurn. 58: 1128.

S

T Russia: Ad sinum Rogersii ad litum meridionale insulae Wrangelii, 27.06.1969, leg. V.V. Petrovsky (LE) holotype.



2n= (1) 42 (6x). (2) 56 (8x).

2nD (1-2) Zhukova & Petrovsky (1985 NE As). See comment.

G RFE

Comments: See also P. calcareum.



(1) [Petrovsky's comment needs translation.]

WARNING! Needs critical study before full acceptance.
The Papaver radicatum aggregate (detritophilum, multiradiatum, radicatum s.l.)

Comments:

(1) The concept of this aggregate depends both on the concept of P. radicatum Rottb. and on the concept of P. lapponicum, see below. As defined by Petrovsky, the aggregate contains hexaploids in the Beringian sector, octoploids in the continental sectors in Siberia   Russia and Canada and in Greenland, and mainly decaploids in the European part of the Atlantic sector.

It was shown by Solstad (1998), Solstad et al. (1999, in prep.) that there is a close similarity in isoenzymes between the diploid P. alpinum (subsp. sendtneri) and the North Atlantic octoploids (P. lapponicum) and decaploids (P. radicatum). It is therefore probable that the mainly European P. alpinum or a related extant or extinct diploid has furnished at least one of the original diploid genomes. (Elven)


52.1.4  Papaver radicatum Rottb. (1770), Skr. Kiøbenhavnske Selsk. Lærd. Elsk. 10: 455.

S P. relictum (Lundstr.) Nordh. (1932), Bergens Mus. Årb. 1931, Naturv. Rekke 2: 45; P. laestadianum (Nordh.) Nordh. (1939), Bot. Not. 1939: 623; P. nordhagenianum Á. Löve (1955), nom. illeg., Nytt Mag. Bot. 4: 15.

T Rottbøll (1770), Tab. VIII, No XXIV, lectotype to be formally selected by Nilsson [& Elven?] (in prep.), a NW Icelandic epitype will also be proposed.

Comments:

(1) Papaver radicatum Rottb. (1770) is the first name coined for a northern Papaver after P. nudicaule L. (1753). The ambiguous concept of P. radicatum has been a major source of confusion for two reasons: (a) Typification; and (b) Ploidy level(s) and circumscription. No stability can be reached   in names and species concepts of the northern papavers   before the 'riddles' of this name are solved. An important task of the checklist might then be to propose a solution which leads to more stability and less ambiguity.

(a) Typification. A short survey is given by Solstad et al. (1999) and here, a fuller survey by Solstad (1998) and Solstad et al. (in prep.). Rottbøll (1770) included plants from S Norway (Dovrefjell Mts.), Iceland and Greenland in his protologue. He referred to Oeder's illustration in Flora Danica from the Dovrefjell Mts. (now considered as P. radicatum subsp. ovatilobum), to Egede's collection from Greenland, and to material collected from Iceland by almost all visiting collectors. His very nice habitus illustration (reproduced by Solstad et al. 1999) was probably drawn from a live plant. It corresponds with NW Icelandic plants and not with Greenlandic ones. His drawing of a fruit is more ambiguous.

Hultén (1945) indicated that the illustration was a natural choice as a type, but he did not formally select it as a lectotype. He interpreted the illustrastion to represent a Greenland plant. That view has later influenced especially North American treatments.

A long and amusing discussion took place between Löve (1955, 1962a, 1962b) on one side and Knaben (1958) and Knaben & Hylander (1970) on the other side. Löve argued for a Greenlandic typification based first on the fruit illustration, later on a find of an Egede specimen which he identified as the 'holotype'. The drawn fruit is narrow and obconical, which corresponds with Greenland rather than Iceland plants. It was interpreted as an immature and unidentifiable gynoecium by Knaben and Knaben & Hylander, a very evident interpretation when one looks at the drawing. Löve's proposed 'holotype' is very badly preserved and does not fit either the Rottbøll drawing or his diagnosis. As Rottbøll cited several elements in his protologue there can be no 'holotype'.

The typification is of importance as the Iceland plants belong to P. radicatum in the concept of Knaben and Hylander, the Greenland ones to P. lapponicum in their concept. This concept of two species was also shared by Löve, and in accordance with his views he 'transferred' the name P. radicatum to the Greenland (and North American) plants whereas he renamed the Icelandic and Scandinavian ones, first as P. nordhagenianum, later as P. relictum as this is an earlier name at species level within the N European group.

Löve was influential among North American (and Greenland) and partly also Russian botanists. His views have largely been accepted there, as late as by Kiger & Murray (1997) and Petrovsky (1999). North European botanists have been much more sceptical and has rejected his opinion throughout in this case. This means that, at present, the name P. radicatum is applied quite differently, and for different plants.

An Icelandic typification will soon be made final in connection with Flora Nordica 2 (probably by Nilsson & Elven). This will imply that the majority of the material referred to by Rottbøll (1770) will be kept within P. radicatum (Iceland, Norway). It will be in accordance with his diagnosis and illustration, and an epitype very similar to his illustration will be chosen.

(b) Ploidy level(s) and circumscription. The acrimony between Löve and Knaben/Hylander was based on the assumption that there was a strict correspondence between ploidy levels and species limits in northern Papaver. The relevant Greenlandic plant was octoploid (2n=56) whereas the Icelandic and Scandinavian one was decaploid (2n=70). Knaben (1958, 1959a, 1959b) restricted the name P. radicatum to Nordic plants: Iceland, the Faeroes, S Norway and N Scandinavia north to W Finnmark province. The only place her species reaches the Arctic as we define it is in N Iceland.

Knaben described all the Greenlandic and North American plants of the group as three subspecies of P. lapponicum: subspp. labradoricum (Fedde) Knaben, occidentale (Lundström) Knaben, and porsildii Knaben. Her classification of all Greenlandic and North American plants as P. lapponicum seems to have been made more out from ploidy level than from overall morphology.

This is also evident from Nordhagen's and Knaben's treatment of the narrow (non-arctic) N Scandinavian endemic P. laestadianum (Nordh.) Nordh. This entity is very close morphologically to P. radicatum sensu Knaben and was first described as a subspecies of that species. It was raised to species when Horn (1938) found it to be octoploid. For morphological reasons it was not possible to include it in the octoploid P. lapponicum. The investigations of Solstad (1998) and Solstad et al. (1999) showed that it is identical with P. radicatum sensu Knaben in all interpretable enzymatic markers and that it might differ only in single versus double dose of one of its original diploid genomes. If one accepts several ploidy levels in a species, as one should do, P. laestadianum must be re-included in P. radicatum sensu Knaben, perhaps as a subspecies because it has a few differentiating characters and as it occurs in several populations in a distinct (although small) area.

North Americans (Kiger & Murray 1997) and Russians (Petrovsky 1999, draft) are convinced that there are two major widespread arctic entities in addition to P. dahlianum subsp. polare and that both are at last predominantly octoploid. Their naming of one of them as P. radicatum is, however, based on the concept of a Greenlandic typification. It must therefore be critically evaluated in view of the Icelandic typification. A preliminary experimental investigation of a fairly large Canadian material is under way (Solstad/Elven). This material includes populations of P. radicatum subspp. labradoricum and occidentale (sensu North Americans), P. lapponicum (sensu North Americans) and the P. dahlianum complex. We might have some more data available fairly soon for this evaluation as we already have much data for S Norwegian P. radicatum and some data for E Icelandic P. radicatum and for P. lapponicum subsp. lapponicum. (Elven)

(2) Also P. radicatum sensu Knaben, i.e., in a narrow concept, has been split into numerous, morphologically weakly differentiated, and geographically very narrow subspecies: 3-4 in Iceland, one in the Faeroes, 5-6 in S Norway, and 4-5 in N Scandinavia. They are all morphologically identifiable and recognisable (Nordhagen 1932, Knaben 1959a, 1959b, Selin & Prentice 1988, Selin 1999, Nilsson in Jonsell in prep., Flora Nordica 2), although mostly by combinations of quantitative characters. They were interpreted by Nordhagen and Knaben as results of splitting and isolated survival on nunataks and coastal forelands of a more continuous pre-Weichselian (or perhaps pre-Quaternary) distribution.

The investigations of Solstad (1998) and Solstad et al. (1999, in prep.) throw strong doubts on that interpretation. She investigated all the S Norwegian races, one N Norwegian and one Icelandic. In enzymatic markers and RAPD-DNA, there is no difference between the subspecies. The small variation found is rather within subspecies and partly within populations. It is improbable that these 'races' have the indicated high age. A much more probable interpretation is that the variation is a result of postglacial genetic drift in small, semi-isolated and fluctuating populations in unstable habitats, and in predominantly inbreeding plants. Interpreted in this way, the variation is what normally could be expected in a species with this reproductive system and population structure. The subspecies concept should not be applied for this situation. As the entities mainly are very local, with few differences, and presumed to be fairly recent, variety might be applied. However, no ecotypical differentiation is involved. (Elven)
52.1.4.1  Papaver radicatum Rottb. subsp. radicatum

S P. radicatum Rottb. subsp. stefanssonii A. Löve (1945), Izlenzkar Jurtir 149 [type: Iceland: Gufudalshals, 08.08.1893, leg. S. Stefansson & O. Davidsson (ICEL) lectotype, selected by Löve (1955, Nytt Mag. Bot. 4: 14)]; P. nordhagenianum Löve subsp. stefanssonii (Löve) Löve (1962), Taxon 11: 137; P. radicatum Rottb. subf. islandicum Lundström (1923), Acta Horti Berg. 7: 411; P. radicatum Rottb. subsp. islandicum (Lundström) Á. Löve (1955), ***; P. stefanssonianum Löve (1955), Nytt Mag. Bot. 4: 14.

2n= 70 (10x).

2nD Löve (1955 Icel, for subsp. stefanssonii). See comment (1).

G ICE

Comments:



(1) If the plants elsewhere in Iceland, the Faeroes, and at least the majority of the SCandinavian plants are included in this subspecies, the number and areas of chromosome counts will increase dramatically as all races have been counted, partly many times. (Elven)

(2) The NW Icelandic plants, from which that of Rottbøll's illustration probably came and from where the epitype will be chosen, have not been available yet for experimental investigations. It is therefore not possible to tell whether these also belong to the large and 'genetically homogeneous' complex that now encompasses the E Icelandic plants, all the S Norwegian ones, and at least parts of the N Scandinavian ones (including P. laestadianum). As none of the Scandinavian 'subspecies' reach the Arctic as defined, they don't matter for the checklist. However, two 'subspecies' are described from the arctic parts of Iceland. I he propose that we until further include the second race (subsp. stefanssonii) only in synonymy, as done above.


52.1.4.2  Papaver radicatum Rottb. subsp. labradoricum (Fedde) Fedde in Engler & Prantl (1936), Natürl. Pflanzenfam. 2, 17b: 120.

B P. nudicaule L. subsp. radicatum *** var. (gamma) labradoricum Fedde in Engler (1909), Pflanzenreich 4, 104: 377.

S P. lapponicum (Tolm.) Nordh. subsp. labradoricum (Fedde) Knaben (1958), Blyttia 16: 78;

2n= 56 (8x).

2nD Löve & Löve (1975) list several counts, all as arctic.

G CAN GRL

Comments:

(1) Petrovsky (1999, draft) separates the Greenlandic   Canadian material of the P. radicatum/-lapponicum group(s) on three entities: 'lapponicum orientale' and 'radicatum occidentale' (in Petrovsky 1999) and 'radicatum labradoricum' (added in draft). The 'labradoricum' entity is indicated for Greenland and NE Canada whereas both the two others are indicated with a very wide range from NW Siberia through NE Siberia and Russian Far East, Alaska and all of northern Canada. This is probably one too many.

Kiger & Murray (1997) accept 'lapponicum' (probably synonymous with Petrovsky's 'orientale') throughout Alaska and Canada and a little more tentative for Greenland while they unite 'labradoricum' and 'occidentale' in their P. radicatum subsp. radicatum (sensu Löve). Pending the results of the investigations mentioned above, the taxa of Petrovsky's draft are very tentatively retained. If 'labradoricum' and 'occidentale' are merged in one subspecies, 'occidentale' has priority. It is probable that the 'labradoricum' entity will land in P. lapponicum after a revision. (Elven)

WARNING! Might be transferred to P. lapponicum.
52.1.4.3  Papaver radicatum Rottb. subsp. occidentale Lundström (1923), Acta Horti Berg. 7, 5: 413.

S P. lapponicum (Tolm.) Nordh. subsp. occidentale (Lundström) Knaben (1959), Opera Bot. 2, 3: 413.

2n= 56 (8x).

2nD Löve & Löve (1975) list several arctic counts under P. radicatum subsp. radicatum which they synonymise with subsp. occidentale.

G RUS SIB RFE ALA CAN GRL

Comments:



(1) Iceland is indicated for this race by Petrovsky (1999) but there are no records of it from there; omitted. Greenland material should also be critically checked. **Check KNABEN** (Elven)

WARNING! Might be transferred to P. lapponicum.


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