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1. Angelica sylvestris L. map ill.
Linnaeus, Sp. Pl.: 251 (1753). – Type: Linnaean Herbarium 354.3 (LINN) lectotype, sel. by Gutiérrez Bustillo, Lazaroa 3: 146 (1981).
D Angelik. F karhunputki. Fa bakkasløkja. I geithvönn. N sløkje. S strätta.
Hemicryptophyte (short-lived hapaxanth). To 150(–200) cm, more or less scentless; taproot 8–20 mm thick. Stem hollow; basal part 5–11(–17) mm thick, usually terete, sometimes red-violet, glabrous, usually glaucous; upper internodes terete to indistinctly sulcate, densely papillose with long papillae. Leaves 2–4 at the base and 3–6 on the stem, the innermost basal one usually the largest; sheath often red-violet, 2.5–4 cm; petiole 14–26(–33) cm, with an adaxial groove (continuing into the rachis); blade 12–32 × 8–24 cm (length/width ratio 1–1.5), often papillose on both sides (with longer papillae above), lower side usually slightly glaucous. Primary leaflets 3–4(–5) pairs; angle leaflet/rachis 40–50°. Ultimate leaflets 1-pinnatifid, usually with one pair of lobes (most leaflets of a leaf-blade with sinuses reaching the midvein); petiolule 0.6–3.5(–4.8) cm; blade (2.8–)4.4–11 × (2.6–)4–10.4 cm (length/width ratio 0.9–1.3); margin not swelled, usually papillose, and serrate to indistinctly doubly serrate (usually with acuminate teeth); base shortly attenuate to cordate; apices of lobes acuminate to obtuse. Apical lobe 2.5–9 × 1.6–3.6(–4.6) cm, with a length/width ratio of 1.5–2.8(–3.3).
Umbels convex, 4–7 cm high and 8.5–17 cm wide; peduncle 6–14 cm; rays ± straight, (3.3–)5.4–10 cm, densely papillose all around (rarely only on the adaxial side). Bracts 0–6. Umbellules (15–)19–50(–56); pedicels 0.8–1.7 cm, usually densely papillose all around (rarely only on the adaxial side). Bractlets (6–)8–16, persistent, (2.5–)4–12 × 0.2–0.6(–0.8) mm, papillose. Flowers 22–52(–66) per umbellule; sepals 0–0.2 mm; petals white or pink, 1.1–2 × 0.7–1.2 mm, apex acuminate; filaments 2–3.7 mm; anthers 0.4–0.6 mm. Fruit oblong to almost rectangular in outline, dorsiventrally distinctly flattened. Mericarps 3.8–6 × 2.5–4.8 × 0.7–1.2 mm (length/width ratio 1–1.6); ridges light brown, the dorsal ones low, close to each other, the lateral ones developed as 0.6–1.5 mm wide wings (measured from the lateral veins); valleculae narrow, each with one dark brown vitta visible on the surface; pericarp thin, not detached from the endocarp at maturity; stylopodium flattened, 0.6–0.9 mm wide; style 1–1.5(–2) mm, directed outwards to deflexed. – Mid-summer to late summer.
2n=22 (S Sk 2, Vg, F A, V 2, U). – [2n=22]
Distribution. Nem–NBor(–LAlp). Alt. N 1100 m. – D common to fairly common throughout (less so in western Jylland). N common to fairly common, but scattered to rare in VFi and ØFi. S common to fairly common, but rare in LL and TL (reaching the low-alpine zone) and in northern Nb. F common to fairly common north to southern PeP and southeastern SoL, further north rare; casual in InL. Fa common. I common to scattered along the coasts and in the lowlands, rare in the central highlands.
Europe (except parts of the Mediterranean), W Siberia.
Habitat. In sun or moderate shade on moist or mesic, nutrient-rich, mull-rich soil. Woodland (e.g. along rivers and creeks), fens, overgrown pastures and meadows, seashores; also roadsides, ditches and moist fields. Favoured by slight overgrowth.
Biology. Life span may vary beween 2 and 5 years (in France; Reduron 2007). The seeds are spread both with wind and water.
Variation. Angelica sylvestris is quite variable in leaf characters. The papillae are poorly developed or even absent in some populations (occurring both in coastal and mountain localities); complete leaves or floral parts may be needed to distinguish such plants from A. archangelica. Some coastal populations may resemble A. archangelica in having broader leaflets with more coarsely serrate margins, and stems and leaves of a more whitish green colour; they have been named var. elatior Wahlenb. or var. maior Hartm. However, at least part of the deviating characters probably represent plastic responses to drier conditions; since the morphological delimitation is fairly vague, they are not taxonomically recognized here.
Nomenclature. The lectotype of Angelica sylvestris refers to a Central European taxon at present known as A. sylvestris subsp. bernardiae Reduron (A. sylvestris var. elatior sensu Thell.; A. montana Brot.; cf. Reduron 2007). The name for the race present in Norden is, for the time being, not clear. A proposal for conservation of the name with a type belonging to that race was unfortunately rejected (Jarvis 2007 onwards), but will be attempted again.
Similar taxa. Sterile specimens of Angelica sylvestris may be similar to A. archangelica, but they usually have papillae on leaf margins and veins (but not always, and they are sometimes indistinct). When fresh, A. archangelica is strongly aromatic. – Angelica specimens without inflorescences (both species) may be confused with some other stout species: like A. archangelica, Ligusticum scothicum is aromatic, but its odour is very different; its petioles and the lower parts of the stem are less robust than in either of the Angelica species, the dark-green leaves are ternate and have purplish sheaths, and the proximal half or third of the leaflet margin is entire (leaflets serrate or crenate ± from the base in Angelica). – Heracleum sphondylium differs in having bristles on stems and petioles, and the leaves are 1-pinnate. – Laserpitium latifolium has solid stems, and the venation of the leaflets is palmate (pinnate in Angelica). – Aegopodium podagraria has a creeping rhizome, and its leaves are unequally 1–2-pinnate with 1–2 pairs of primary leaflets (Angelica has ± equally bipinnate leaves).– See also Levisticum officinale and Peucedanum ostruthium.
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2. Angelica archangelica L. map (illustrations see subspecies)
Linnaeus, Sp. pl.: 250 (1753). – Type: Linnaean Herbarium 354.1 (LINN) lectotype, sel. by Reduron, Nordic J. Bot. 22: 83 (2002).
D Kvan. F väinönputki. Fa hvonn. I ætihvönn. N kvann. S kvanne.
Literature. Ojala 1984, Thellung 1926, Weinert 1973.
Hemicryptophyte (long-lived hapaxanth). To 130(–200) cm, with a ± strong aromatic scent (see subspecies); taproot often at least 35 mm thick. Stem hollow; basal part 9–18 mm thick, terete to slightly sulcate, pale greenish or sometimes red-violet, slightly glaucous; upper internodes terete to slightly sulcate, glabrous to sparsely papillose, with short papillae. Leaves 4 at the base and 4 on the stem, the innermost basal one the largest; sheath often red-violet, 4.5–9 cm long; petiole 11–26(–40) cm, terete, without an adaxial groove (but there usually is one on the main rachis); blade 18–60 × 20–60 cm (length/width ratio 0.9–1.1), glabrous, lower side glaucous or not (see subspecies). Primary leaflets 2–3 pairs, angle leaflet/rachis 40–55°. Ultimate leaflets 1-pinnatifid, with 1(–2) pairs of acuminate to obtuse lobes; petiolule 1–5.5 cm; blade 6.5–13 × 8–17 cm (length/width ratio 0.7–1.3); margin usually not papillose, ± swelled, usually indistinctly doubly serrate with acute to acuminate teeth; base shortly attenuate to cordate; at least some ultimate leaflets with shallow sinuses (reaching only halfway to the midvein). Apical lobe 4.5–10 × (2.2–)3.2–7.5 cm (length/width ratio 1.2–2.7).
Umbels globose, 10–14 cm across; peduncle 10–13 cm; rays ± straight, 4.7–6.4 cm, ± papillose all around or only on the adaxial side. Bracts 0–8. Umbellules 22–54; pedicels 0.9–1.5 cm, ± papillose all around (rarely only on the adaxial side). Bractlets 2–13, persistent, glabrous or papillose (size see subspecies). Flowers 34–68 per umbellule; sepals 0–0.2 mm; petals 1–1.9 × 0.7–1.4 mm (excluding the 0.2–0.5 mm long tip); filaments 2.2–4.2 mm; anthers 0.6–0.8 mm. Fruit oblong, broadly oblong or almost rectangular in outline, not or slightly dorsiventrally flattened. Mericarps 4–9 × 3–6 × 1.1–2.2 mm, with a length/width ratio of 1.1–1.6(–2); ridges straw-coloured, the dorsal ones low to high, not very close to each other, the lateral ones developed as 0.2–0.9 mm wide wings measured from the lateral veins; vittae numerous, surrounding the endocarp, not visible on the surface; pericarp thick, detached from the endocarp at maturity; stylopodium flattened, 0.9–1.5 mm wide; style 1–2 mm, directed outwards to deflexed. – Early summer to mid-summer.
2n=22 (S Sk 3, F U 3, EH). – [2n=22]
Distribution. Nem–LAlp(-MAlp). – Native to the area, but also cultivated, escaped and naturalized both in the Scandes and in the lowlands. – D mainly along the coasts, scattered to fairly common in eastern Jylland and the eastern islands, scattered to rare in western Jylland and at Limfjorden. N common in the Scandes (but rare in NT) and rather common along most of the coast (but several gaps from SF to NT); in the north also in lowlands inland. S in the Scandes common to fairly common from northern Dlr to southwestern Jmt, and from LyL to TL, rare to absent in northern Jmt and ÅsL; in the lowlands common to fairly common along the coasts from northern BhG to western Sk, from northern Klm to northern Upl and in Nb; scattered to rare from southern Sk to central Bl, in northern Öl, western Gtl and Ång Nora; inland localities in, e.g., Sk Öved, Dls Gunnarsnäs (field), Vrm Forshaga (farm), Srm Björnlunda (local community house), ÅsL/Vb along Vojmån and Vindelälven, and Nb along Kalix älv and Torne älv (see further under the subspecies). F in the northern mainland fairly common in InL, EnL and eastern SoL; further south scattered to southwestern PeP, northeastern OP and southern Ks, an isolated population in EH Somero; along the coast scattered from PeP to southern KP, further south rather common or common. Fa fairly common. I fairly common throughout.
Native to N and NE Europe, southern Greenland, N and C Siberia, and Himalaya; grown as a spice and vegetable since ancient time.
Habitat. In sun or moderate shade on moist or mesic, nutrient-rich, mull-rich, peaty or sandy soil; late snowbeds, streams, wet meadows, rocky slopes, riversides, sandy, gravelly or rocky seashores, and birdcliffs. Favoured by slight overgrowth. See further under the subspecies.
Biology. All flowers of the primary and secondary umbels are bisexual, or secondary umbels have both bisexual and male flowers (tertiary umbels are hardly developed). The flowers are partly self-compatible (Ojala 1986).
Since the endosperm and perisperm are detached from each other when the seed ripens, they have an air-filled interspace, which enable the seeds to float. They are probably spread mainly with water (Ojala 1984), e.g. along coasts and rivers, and perhaps also in the mountains by water from melted snow. However, wind-spreading may also occur. Seeds from primary umbels have a significantly higher germination rate than seeds from secondary umbels; many of the seeds that do not germinate lack an embryo (Ojala 1985).
Variation and taxonomy. The fruits vary quite extensively in both size, shape and characteristics of the ridges, but it has not been possible to subdivide the whole Nordic material according to fruit characters only (many specimens lack ripe fruits, many intermediates occur, and even within populations there may be extensive variation in fruit characters, e.g. in S Hl Särö and Nb Pajala). Traditionally, Angelica archangelica has been subdivided into two taxa in Norden, one distributed in the mountains (subsp. archangelica, characterized by long bractlets and large fruits with acute ridges) and one distributed on seashores (subsp. littoralis, with short bractlets and small fruits with rounded ridges); however, specimens have usually been classified based on habitat only. A comprehensive investigation of the taxonomy of the species in Norway indicated correlation of habitat and fruit morphology in the south but not in the northern parts (Mehus 1970). Generally, the fruits are larger in subsp. archangelica than in subsp. littoralis, and several further characters can be applied to separate the taxa (see descriptions).
A. archangelica has an uneven distribution in Norden; there are large gaps in the inland, both in southern and central F and southern S. There is also a smaller, but distinct gap in S northern Jmt, ÅsL, and N NT. Subsp. archangelica occurs in most of the distribution area of the species, but seems to be rather common northwards (S LL, TL, N Tr, NNo and F EnL); only subsp. archangelica occurs in I and Fa. Subsp. littoralis, on the other hand, is mainly restricted to the southern coasts where it, however, occurs sympatrically with subsp. archangelica in several provinces. Intermediates occur throughout the distribution area, but seem to be less common in the northern parts. The distribution information given for the subspecies was based only on specimens examined by the author.
A. archangelica has also been cultivated for a very long time especially in the mountains. Most of the specimens studied that derive from cultivation are intermediates between the subspecies. It has been suggested that A. archangelica immigrated into Norden from the northeast after the last glaciation, and then differentiated into the two subspecies (Mehus 1970). However, since both subspecies occur in NW Russia, subsp. littoralis e.g. along the White Sea and subsp. archangelica e.g. on the Kola Peninsula, it seems more plausible that both subspecies of A. archangelica have immigrated from the east, but along different paths. Subsp. littoralis, with Nordic distribution only in the southern coastal districts in Norden, may have immigrated along a southern path. Subsp. archangelica, occurring both in the south and in parts of the Scandes, may have immigrated both along a southern and a northern path; this could also explain the distribution gap in central S and N. The fact that intermediates seem to be more frequent south of the gap than north of it, supports the idea of these migration paths. Later cultivation of the species may have altered earlier distribution ranges further.
Similar taxa see Angelica sylvestris (1).
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2A. subsp. archangelica map ill.
D Fjeld-Kvan. F väinönputki. N fjellkvann. S fjällkvanne.
Angelica officinalis Moench (1794).
Plant with an aromatic taste, not glaucous. Stem and petioles rather easily compressed. Apical leaflet elongated (length/width ratio 1.5–2.7). Bractlets 8–13, 4–18 × 0.3–1(–1.5) mm. Flowers with green-grey to olive-green petals. Fruit with 6–9 × 4–6 mm large mericarps; median ridges with a keeled edge.
Distribution and habitat. Nem–LAlp(-MAlp). – Probably native both in lowlands (mainly on seashores) and mountain areas (e.g. brooks and rivers, mainly in Betula woodland).
Verified occurrence: D NJy Læsø (Storedal), ØJy Anholt, Hørning, Randers, FyL Odense, Horne (possibly from cultivation), Sjæ 4 localities in the north, LFM Kragenæs. N in the south mainly in mountains (in most of the range of the species except ST and northern He), but also confirmed lowland records from Øf Hvaler and Fredrikstad, Ak Asker and Eidsvoll (lake shore), He Ringsaker (brook), Te Kragerø, Ro Hå, Ho Stord and Voss, SF Eid (roadside), and MR Haram; from southern SNo to ØFi Kirkenes scattered both in the lowlands and the mountains. S BhG Göteborg area 5 localities, Sk western and southern coasts and Billeberga (inland), Bl Edestad (Skaftö), Klm Misterhult (Blå Jungfrun) and Västervik, Öl Borgholm, Gtl Visby (outside botanical garden) and Lummelunda, Ög Jonsberg, and from Srm Torö to Upl Österlövsta, Hls Los (inland, saw mill and brook), Vb Degerfors, Nb 6 localities (inland in the northern part), ÅsL Vilhelmina (Stalon, roadside); in the mountains in most of the range of the species but fairly rarely collected (possibly more frequent in TL near Torne Träsk). F A Föglö, from V Turku to EK Kotka, and EP Mustasaari; in the north rare throughout the range of the species. Fa confirmed from Eysturoy, Streymoy and Suðuroy (probably fairly common also elsewhere). I confirmed from all provinces (probably fairly common).
N and E Europe, Greenland and NW Asia. Other subspecies further east.
Variation. Specimens from populations in the mountain region generally have more elongated leaf-lobes than those in the lowlands. – A deviating morphotype from N Ho Voss, with solid stems/petioles (var. maiorum Fægri, N vossakvann), probably belongs to subsp. archangelica, but since it is only known in vegetative state, the inclusion in this taxon has not been verified; it has been collected for cultivation (Øvstedal, pers. comm.).
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2B. subsp. littoralis (Wahlenb.) Thell. map ill.
Thellung, Ill. Fl. Mitteleur. 5: 1342 (1926). –A. archangelica var. littoralis Wahlenb., Flor. Carp.: 84 (1814). – A. littoralis (Wahlenb.) Fr. (1817). – Type: S Hl Ölmevalla, Åsa, 29.VII.1953 Ö. Nilsson (LD) neotype, sel. by Fröberg, Nordic J. Bot. 27: 139 (2009).
D Strand-Kvan. F meriputki. N strandkvann. S strandkvanne.
Plant with a sharp taste, usually distinctly glaucous. Stem and petioles hard. Apical leaflet rounded (length/width ratio1.2–1.6). Bractlets 2–12, 2.5–5 × 0.2–0.4 mm. Flowers with white to greenish white petals. Fruit with 5–6.5 × 3.5–4.5 mm large mericarps; median ridges with a rounded edge.
Distribution and habitat. Nem-BNem (SBor). Native on the southern coasts, very few inland occurrences. Stony or sandy seashores, seashore meadows; also rivers and ditches close to the sea.
Verified occurrence: D NJy Skagen and Gudum, ØJy Anholt, SJy both coasts (3 localities in all), FyL east coasts of Fyn and Langeland (5 localities), LFM (Lolland and Falster), Sjæ west and east coasts (5 localities), Brn Hasle. N along the very coast and in some isolated northern islands, but not in the fjords; scattered from Øf Hvaler to VA Kristiansand; Ro 3 localities, Ho Bømlo, SF Solund, MR Haram, SNo Vega and Træna, and NNo Bodø, Steigen, Hadsel and Røst. S scattered on the west coast south to Sk Bunkeflo, and from Ög Gryt to Upl Gräsö; Bl Mjällby, Klm 3 localities near Oskarshamn, Öl Högby, and Gtl 4 localities on the west coast; inland in Sk Bjärshög and SmI Tranås (riverside). F scattered on the south coast west to V Kustavi; A 2 localities and St Pori. – Records from I and Fa (e.g. Löve 1970, Mossberg & Stenberg 2003) could not be confirmed; intermediate specimens have been seen from I, but no subsp. littoralis, and all material from Fa which could be classified belonged to subsp. archangelica.
Verified occurrence: D NJy Skagen and Gudum, ØJy Anholt, SJy both coasts (3 localities in all), FyL east coasts of Fyn and Langeland (5 localities), LFM (Lolland and Falster), Sjæ west and east coasts (5 localities), Brn Hasle. N along the very coast and in some isolated northern islands, but not in the fjords; scattered from Øf Hvaler to VA Kristiansand; Ro 3 localities, Ho Bømlo, SF Solund, MR Haram, SNo Vega and Træna, and NNo Bodø, Steigen, Hadsel and Røst. S scattered on the west coast south to Sk Bunkeflo, and from Ög Gryt to Upl Gräsö; Bl Mjällby, Klm 3 localities near Oskarshamn, Öl Högby, and Gtl 4 localities on the west coast; inland in Sk Bjärshög and SmI Tranås (riverside). F scattered on the south coast west to V Kustavi; A 2 localities and St Pori. – Records from I and Fa (e.g. Löve 1970, Mossberg & Stenberg 2003) could not be confirmed; intermediate specimens have been seen from I, but no subsp. littoralis, and all material from Fa which could be classified belonged to subsp. archangelica.
Endemic; outside Norden on the Baltic Sea from Russia to Germany, and on the White Sea in NW Russia.
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