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[00]  Solanaceae (Draft: R. Elven)

THIS DRAFT HAS NOT BEEN DISCUSSED
Probably only casual occurences in the Arctic, e.g., Solanum tuberosum L. The family should probably be excluded. (Elven)

89  Scrophulariaceae (Drafts: L.I. Ivanina, Castilleja by O.V. Rebristaya [OR])

THIS DRAFT HAS ONLY BEEN SUPERFICIALLY DISCUSSED

Comments:

(1) The following two genera and species probably occur as casuals only and should not be entered: Chaenorhinum minus (L.) Lange in Willk. & Lange (1870), Prodr. Fl. Hisp. 2: 577 [basionym: Antirrhinum minus L. (1753), Sp. Pl. 617] from CAN, and Verbascum thapsus L (1753), Sp. Pl. 177 from GRL. (Elven)

89.1  Linaria Mill. (1754), Gard. Dict. Abr., ed. 4, 2.
89.1.1  Linaria vulgaris Mill. (1768), Gard. Dict., ed. 8, no. 1.

B Based on Antirrhinum linaria L. (1753), Sp. Pl. 616.

S

Comments:



(1) The two races included below are treated as separate species by Ivanina in Tolmachev & Yurtsev (1980, Fl. Arct. URSS 8) but fully united by Chater et al. (1976, Fl. Eur. 3). I ahve followed a middle path. (Elven)
89.1.1.1  Linaria vulgaris Mill. subsp. vulgaris

S

2n= 12 (2x).



2nD Gadella & Kliphuis (1966 W Eur); Laane (1969 Norw); Lövkvist & Hultgård (1999 Sweden)

G NOR* RUS? GRL*?

Comments:

(1) Subsp. vulgaris only occurs as a vegetatively remaining garden escape without seed propagation in arctic Norway and probably also in Greenland. It is therefore debatable whether it should be included or not. The only reported native Pechora occurrence should be checked as to status as subsp. vulgaris is comparatively thermophilous. The Pechora occurrences are mapped as subsp. acutiloba by Hultén & Fries (1986). (Elven)



WARNING! Might be excluded as non-stable.
89.1.1.2  Linaria vulgaris Mill. subsp. acutiloba (Fisch. ex Rchb.) Hultén (1971), Kungl. Sv. Vetensk.-Akad. Handl. 13, 1: 401.

B L. acutiloba Fisch. ex Rchb. (1827), Pl. Crit. 5: 14, t. 611.

S L. vulgaris auct., non Mill. (1768).

2n= 12 (2x).

2nD ?

G SIB RFE



Comments:

(1) The same combination of subspecies was later made by D.Y. Hong (1979), Fl. Reipubl. Popul. Sin. 67, 2: 208, 400. Why? As far as I can see the Hultén combination is valid. (Elven)



89.2  Limosella L. (1753), Sp. Pl. 631.
89.2.1  Limosella aquatica L. (1753), Sp. Pl. 631.

S

2n= 40 (4x).



2nD Löve & Löve (1975) list several counts, one Greenlandic and two Icelandic as arctic.

G ICE RUS SIB ALA CAN GRL

Comments:

(1) For RUS: Present in arctic parts of Kola Peninsula. Approaches the Arctic also in N Norway. (Elven)



89.3  Mimulus L. (1753), Sp. Pl. 634.
89.3.1  Mimulus guttatus DC. (1813), Cat. Pl. Horti Monsp. 127.

S

2n= 48. **Check British counts, Stace, for ploidy level**



2nD Maude (1939 W Eur).

G ALA?


Comments:

(1) Genus and species added to Ivanina's draft. The species occurs, acc. to Hultén (1968), in W Alaska in the Yukon Delta and Kuskokwim Mts and should be accepted as a border case. (Elven)



89.4  Pentstemon Mitchell (1769), Diss. 36.
89.4.1  Pentstemon gormani Greene (1902), Ottawa Naturalist 16: 39.

S

T Canada: Yukon Territory, Dry Gulch, leg. Gorman 1011 (US, CAN).



2n=

2nD


G ALA?

Comments:

(1) Genus and species added to Ivanina's draft. According to Hultén (1968) it occurs in the southern parts of Brooks Range (N Alaska-Yukon) and thereby becomes an acceptable border case. (Elven)

89.5  Lagotis P. Gaertn. ***

Comments:

(1) The data concerning the two entities listed below are conflicting. Hultén (1968) considered them as subspecies in Alaska as he reported that "transitions between the two races occur frequently where the ranges overlap". He also recognised L. hultenii as a separate species in NE European Russia and NW Siberia. This opinion was not shared by Webb (1976, Fl. Eur. 3) who restricted L. minor to NE European Russia and Siberia, an opinion also reflected by Hultén & Fries (1986) as they mapped L. minor (as species) only in some smaller areas of European Russia and Siberia and excluded it from North America. It would perhaps be better to treat the two listed entities as subspecies as done by Petrovsky in Tolmachev & Yurtsev (1980) and I have changed accordingly. (Elven)
89.5.1  Lagotis glauca P. Gaertn. (1770), Nov. Comment. Acad. Sci. St. Peterb. 14: 534.

S

Comments:


89.5.1.1  Lagotis glauca P. Gaertn. subsp. glauca

S

2n= 22 (2x).



2nD Löve & Löve (1975) list five counts, one American and one Russian as arctic.

G RFE ALA

Comments:
89.5.1.2  Lagotis glauca P. Gaertn. subsp. minor (Willd.) Hultén (1967), Ark. Bot., ser. 2, 7, 1: 118.

B Gymnandra minor Willd. (1811), Mag. Ges. Naturf. Fr. Berlin 5: 393.

S L. minor (Willd.) Standl. (1931), Publ. Field Mus. Bot. (Chicago) 8: 325; L. hultenii Polunin (1951), Rhodora 53: 228; L. glauca P. Gaertn. subsp. stelleri (Cham. & Schltdl.) Gjaerev. (1967), Bot. Invest. Centr. Alaska 3: 30.

2n= (1) 22 (2x). (2) 44 (4x).

2nD (1) Löve & Löve (1975) list six counts, all as arctic. (2) Löve & Löve (1975) list one count (Sokolovskaya & Strelkova 1960) under the name L. hultenii.

G RUS SIB RFE ALA CAN

Comments:

(1) In view of the opinions of Webb (1976) and Hultén & Fries (1986), the relationship between the North American and Siberian plants should be discussed. (Elven)



89.6  Synthyris Benth. in DC. (1846), Prodr. 10: 454.
89.6.1  Synthyris borealis Pennell (1933), Proc. Acad. Nat. Sci. Philad. 85: 88.

S

T Alaska: Mc Kinley Park, Double Mt., leg. Dixon 20 (Cal U, Hb. Philad.).



2n=

2nD


G CAN

Comments:

(1) Genus and species added to Ivanina's draft. Occurs within the Arctic as defined in the northern Richardson Mts, both in Yukon Territory and Mackenzie District (see Cody 1996). (Elven)

89.7  Veronica L. (1753), Sp. Pl. 9.

S Pseudolysimachion Opiz (1852), Seznam 80; Veronicastrum Moench ***

Comments:

(1) Pseudolysimachion is sometimes treated as a separate genus due, e.g., to a different basic chromosome number. If these species are segregated, also other species groups need to be considered. We prefer to keep the genus in a wide sense until further. (Ivanina & Elven)

(2) Excluded species: V. arvensis L. (1753), Sp. Pl. 13), casual, GRL; V. persica Poir. in Lam. (1808), Encycl. Méth. Bot. 8: 542, casual, GRL. (Elven)
Sect. Pseudolysimachion Koch ***
89.7.1  Veronica longifolia L. (1753), Sp. Pl. 10.

S Pseudolysimachion longifolium (L.) Opiz (1852), Seznam 80; V. septentrionalis Boriss. (1955), Bot. Mater. Gerb. Bot. Inst. AN SSSR 17: 341; Pseudolysimachion longifolium (L.) Opiz subsp. septentrionale (Boriss.) Holub ***; Pseudolysimachion septentrionale (Boriss.) Á. Löve & D. Löve (1976), Bot. Not. 128: 518; Pseudolysimachion maritimum (L.) Á. Löve & D. Löve (1976), Bot. Not. 128: 518.

2n= (1) 34 (2x). (2) 68-70 (4x).

2nD (1) Löve & Löve (1975) list four counts for Veronicastrum septentrionale and V. maritimum; Laane (1965 N Norw); Kukkonen (1986 N Finl, 2 counts for V. longifolia s. str., S Finl, 5 counts for var. maritima); Lövkvist & Hultgård (1999 Sweden, 5 counts). (2) Raitanen (1967 Finl, 2n=68), see comment.

G NOR RUS SIB

Comments:

(1) There is an evident and large variation but probably not in the plants that reach the Arctic. A correlation between ploidy level and morphology remains to be proved. The tetraploid Finnish count (Raitanen 1967, Ann. Bot. Fenn. 4) has been critically checked by Kukkonen (1986, Symb. Bot. Upsal. 27, 2) and is dubious. (Elven)
89.7.2  Veronica incana L. (1753), Sp. Pl. 10.

S Pseudolysimachion incanum (L.) Holub (1967), Folia Geobot. Phytotax. 2: 424; V. spicata L. subsp. incana (L.) Walters (1972), Bot. J. Linn. Soc. 65: 269.

2n= (1) 34 (2x). (2) 68 (4x).

2nD (1) ? (2) Graze (1933 C Eur).

G SIB*

Comments: Subsp. incana in the Arctic.


89.7.3  Veronica spicata L. (1753), Sp. Pl. 10.

S Pseudolysimachion spicatum (L.) Opiz (1852), Seznam 80.

2n= (1) 34 (2x). (2) 68 (4x).

2nD (1) ? (2) Engelskjøn (1979 Norw); Kukkonen (1986 Finl, 10 counts); Lövkvist & Hultgård (1999 Sweden, 18 counts).

G RUS

Comments: Subsp. spicata in the Arctic.


Sect. Veronicastrum Koch ***
89.7.4  Veronica alpina L. (1753), Sp. Pl. 11.

S

Comments: See subsp. pumila.


89.7.4.1  Veronica alpina L. subsp. alpina

S

2n= (1) 18 (2x). (2) 36 (4x).



2nD (1) Löve & Löve (1975) list numerous counts, several as arctic; Engelskjøn (1979 N Norw, 3 counts). (2) Knaben & Engelskjøn (1967 Norw).

G ICE NOR RUS SIB CAN GRL

Comments:
89.7.4.2  Veronica alpina L. subsp. pumila (All.) Pennell (1921), Rhodora 23: 14.

B V. pumila All. (1785), Fl. Pedem. 1: 75.

S V. alpina L. var. (beta) australis Wahlenb. (1814), Fl. Carpath. 5.

2n= 18 (2x).

2nD Löve & Löve (1975) list eight non-arctic counts; Engelskjøn (1979 N Norw).

G ICE? GRL?

Comments:

(1) The 'pumila' (or 'australis') entity is enigmatic. The differences from 'alpina' s. str. race are small but constant in stem, bract and especially in fruit indumentum, there are also differences in leaf shape and denture, and the stem is almost never straight as it usually is in 'alpina' s. str. In Fennoscandia it differs both ecologically and geographically from subsp. alpina. It is restricted to open calcareous soil at high altitudes whereas subsp. alpina is indifferent and often grows in closed vegetation at all altitudes. It is restricted to the two central mountain areas in the south and north whereas subsp. alpina is more ubiquitous, but this might partly be an effect of its high-altitude preference. Intermediates have not been documented.

This race 'crops out' very disjunctly in most major mountain areas in Europe (the two names originate from W Alps and the Carpathians). According to Hultén & Fries (1986) it is the only race south of Britain and the Nordic area. Most authors treat it as a variety, but as it is a major geographic and partly allopatric entity differing in several unrelated characters and without documented intermediates, the choice is rather between subspecies and species.

Subsp. pumila is reported to occur in Iceland but I do not know the distribution. In Norway it occurs in the mountain range south of the Arctic. Occurrence in Greenland is disputed but the entity is accepted from Greenland by Böcher et al. (1978, as var. australis). (Elven)



WARNING! Might be excluded as non-arctic.
89.7.5  Veronica wormskjoldii Roemer & Schult. (1817), Syst. 1: 101.

S V. alpina L. var. wormskjoldii (Roemer & Schult) ***

T [Described from Greenland.]

2n= For the collective species. (1) 18 (2x). (2) 36 (4x).

2nD (1) ? (2) Löve & Löve (1975) list four counts, three as arctic.

Comments: **Check Pennell (1921, Rhodora 23 p. 15 ca) and Fernald (1939, Rhodora 41: 447-457)**

(1) The taxonomic consequences of the variation in this species is disputed. Hultén (1968) recognised two entities in NW America: the wide-ranging subsp. wormskjoldii from Greenland west to Alaska and the Cordilleran subsp. alterniflora from California to Alaska where it reaches the Arctic in Brooks Range. These were not accepted by Cody (1996). Hultén synonymised Bongard's V. nutans with subsp. wormskjoldii. However, after a ploidy variation was proved in the 1960ies, Löve & Löve (1975) separated the NW American plants as a diploid V. nutans from the NE American tetraploid V. wormskjoldii s. str. They do not mention the 'alterniflora' entity and it is not clear whether all the NW American plants belong in their V. nutans.

The differences reported between the races or species are small and mainly quantitative. We have to decide whether we shall enter them formally or keep a (very) collective species. The options for separate entities seem to be:

(a) As an eastern ('wormskjoldii') and western species ('nutans'), the former restricted to GRL and eastern CAN, the western to western CAN and ALA. The western entity then includes both subsp. alterniflora and western subsp. wormskjoldii in Hultén's meaning.

(b) Similar, but as two subspecies. Then subsp. wormskjoldii is the name for the eastern entity, subsp. alterniflora (with the 'nutans' names as synonyms) for the western entity. Both solutions (a) and (b) neglect the morphological variation found in the western plants.

(c) As Hultén (1968) with a wide subsp. wormskjoldii and a Cordilleran subsp. alterniflora. This solution probably neglects the ploidy difference.

(d) As three entities, perhaps represented by the names 'wormskjoldii' (eastern tetraploid entity, Greenland typification, CAN and GRL), 'nutans' (western coastal, perhaps diploid entity, SE Alaskan typification, ALA), and 'alterniflora' (western Cordilleran, perhaps tetraploid entity, Canadian Rockies typification, ALA).

Until further I have only entered the two entities from Hultén (1968). (Elven)

WARNING! Subspecies might be omitted.
89.7.5.1  Veronica wormskjoldii Roemer & Schult. subsp. wormskjoldii

S ?V. nutans Bong. (1833), Mem. Acad. Petersb. 2 ['Veg. Sitcha']: 157; ?V. wormskjoldii Roemer & Schult. var. nutans (Bong.) Pennell (1921), Rhodora 23: 15.

2n=

2nD


G ALA CAN GRL

Comments:

(1) Veronica nutans is described from Alaska: Sitka. In the 'western' sense, the possible 'nutans' entity reaches the Arctic in Seward Peninsula and perhaps in SW Kuskokwim Mts. whereas 'wormskjoldii' reaches it in S Greenland, NE Canada N to S Baffin Island, and perhaps a few sites W of Hudson Bay. (Elven)
89.7.5.2  Veronica wormskjoldii Roemer & Schult. subsp. alterniflora (Fern.) Pennell in Abrams (1951), Ill. Fl. Pacific States 3: 793.

B V. alpina L. var. alterniflora Fern. (1939), Rhodora 41: 455.

S

T [Described from Canada: Rocky Mts.]



2n=

2nD


G ALA

Comments:

(1) In the restricted sense it reaches the Arctic only in Brooks Range. (Elven)
89.7.6  Veronica fruticans Jacq. (1761), Enum. Stirp. Vindob. 2: 200.

S

2n= 16 (2x).



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

G ICE NOR GRL

Comments: Subsp. fruticans in the Arctic.
89.7.7  Veronica serpyllifolia L. (1753), Sp. Pl. 12.

S Veronicastrum serpyllifolium (L.) Fourr. (1869), Ann. Soc. Linn. Lyon, n. s., 17: 128.

Comments: See subsp. humifusa.
89.7.7.1  Veronica serpyllifolia L. subsp. serpyllifolia

S Veronicastrum serpyllifolium (L.) Fourr. subsp. serpyllifolium.

2n= 14 (2x).

2nD Löve & Löve (1975) list numerous counts, two as arctic; Lövkvist & Hultgård (1999 Sweden).

G ICE NOR RUS GRL*

Comments:

(1) May also be introduced in Iceland and N Norway but then an archaeophyte. (Elven)
89.7.7.2  Veronica serpyllifolia L. subsp. humifusa (Dickson) Syme in Sowerby (1866), Engl. Bot., ed. 3, 6: 158

B V. humifusa Dickson (1794), Trans. Linn. Soc. London 2: 288.

S Veronicastrum serpyllifolium (L.) Fourr. subsp. humifusum (Dickson) W.A. Weber (1987), Phytologia 62, 5: 437; V. tenella All. (1785), Fl. Pedem. 1: 75.

2n= 14 (2x).

2nD Rutland (1941 W Eur); Löve & Kjellqvist (1974b SW Eur?).

G ALA CAN?

Comments:

(1) The 'humifusa' entity is geographically absurd and does not at all fit our 'rule' that an accepted entity shall have a consistent and at least theoretically explainable distribution. It occurs scattered in mountainous (but not alpine) areas nearly everywhere on the northern hemisphere (perhaps except for Russia   Siberia) and also in a very few arctic or nearly arctic areas. In addition it is found in some isolated islands (the subtropical Azores are indicated by Hultén & Fries 1986) and probably south America. The three epithets applied to it are based on plants from the British Isles (Dickson's 'humifusa'), the western Alps (Allioni's 'tenella') and North Fennoscandia (Laestadius' 'borealis'). It is not at all clear that they refer to the same taxon.

The entity is generally characterised by lower growth than subsp. serpyllifolia, efficiently rooting branches (forming carpets), shorter racemes (also in fruit), a dark brownish indumentum, presence of dark glandular hairs in inflorescence (vs. eglandular in subsp. serpyllifolia), pedicels longer than the calyx, flowers slightly larger and sky blue (vs. pale blue to white with bluish marks), and capsule with a cuneate base (vs. rounded). These differences refer at least to the North Fennoscandian and Alaskan plants. The differences are too many and presumed independently inherited to be explained by a recurrent mutation. Hybrids are fairly frequent in intermediate habitats in the few places in Fennoscandia where the entities meet and here they behave like races (subspecies). There is also a distinct eco-geographical difference, at least in Fennoscandia. Subsp. serpyllifolia is more or less ruderal and occurs throughout the area except for northern boreal woodlands and mountains. Subsp. humifusa is a specialist of northeastern boreal forest swamps, mountain swampy willow thickets and cold springs, mostly found well outside the range of subsp. serpyllifolia.

Until more evidence is available, I propose that we accept subsp. humifusa as a subspecies. Its rank has varied in more recent decades from species to variety. As to name, I am more in doubt. In Fennoscandia this is a distinctly northeastern continental and boreal taxon and we should check the British typification of 'humifusa'. (Elven)


Sect. Veronica
89.7.8  Veronica chamaedrys L. (1753), Sp. Pl. 13.

S

2n= 32 (4x).



2nD Gadella & Kliphuis (1963 W Eur); Laane (1969 Norw); Lökvist & Hultgård (1999 Sweden, 3 counts).

G NOR*


Comments:

(1) Occurs in arctic mainland Norway (Nordkapp), most probably as an archaeophyte but I can't completely exclude the possibility that it is native here. (Elven)


89.7.9  Veronica officinalis L. (1753), Sp. Pl. 11.

S

2n= (1) 18 (2x). (2) 36 (4x).



2nD (1) Böcher (1944 Denm, Sweden). (2) Löve & Löve (1975) list numerous counts, one Icelandic; Lövkvist & Hultgård (1999 Sweden, 4 counts).

G ICE


Comments:

(1) Also approaches the arctic boundary in N Norway (Porsanger). (Elven)


89.7.10  Veronica scutellata L. (1753), Sp. Pl. 12.

S

2n= 18 (2x).



2nD Löve & Löve (1975) list numerous counts, one Icelandic.

G ICE RUS CAN

Comments:
Sect. Pocilla Dumort. ***
89.7.11  Veronica peregrina L. (1753), Sp. Pl. 14.

S

Comments:


89.7.11.1  Veronica peregrina L. subsp. xalapensis (Humb., Bonpl. & Kunth) Pennell (1919), Torreya 19: 167.

B V. xalapensis Humb., Bompl. & Kunth (1818), Nov. Gen. Sp. 2: 167.

S

2n=


2nD

G CAN


Comments:

(1) Added to the Russian drafts as it is reported from the Arctic at York Factory, Hudson Bay (Scoggan 1978 and other sources).



According to Hultén (1968), subsp. xalapensis is the widespread race whereas subsp. peregrina as indigeneous is restricted to eastern North America. The two entities seem to be well characterised and acceptable as subspecies. (Elven)


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