Hippophaë rhamnoides L. Map
Linnaeus, Sp. pl.: 1023 (1753) - Type not designated.
D Havtorn. F tyrni. N tindved. S havtorn.
Dioecious phanerophyte. Deciduous shrub, often forming dense, extended stands, or rarely a small tree (up to 6 m); all green parts densely to moderately densely covered with peltate, fringed, silvery or copper-brown scales. Roots vertical, to 1 m depth (sometimes also horizontal roots, creeping extensively near the surface). Stems usually angled and curved, ascending or patent, to 15(-30) cm thick, richly and densely branched with many dead branches remaining below the leaf-bearing ones; living branches thorny from transformed side-shoots and apical shoots. Leaves alternate, densely set along this year’s long-shoot and on short-shoots along last year’s long-shoot; petioles 1-3(-5) mm; blade rather thick, lanceolate, 35-65 x 5-8 mm, grey-green above and silvery grey beneath, with subacute apex, attenuate base and entire margin.
Male inflorescences axillary at buds of last year’s shoots, short, spike-like with up to 12 flowers; tepals 2, ovate, greenish, 2–3 mm (at pollen dispersal still almost appressed to each other with only a narrow slit between them; stamens 4 with short filaments and 1.2–1.7 mm long anthers. Female flowers epigynous, single in leaf axils on this year’s shoots; hypanthium tubular with two small tepals; ovary superior with one style and an elongated, yellow stigma. Fruit a juicy drupe formed by the swollen hypanthium, broadly ellipsoid to ovoid, 9.5-12 x 8.5-10 mm, orange with scattered brownish scales, densely clustered along the young shoots; stone narrowly ovoid, flattened, 4-7 x 2-3 mm, finely rugose. - Spring.
2n = 24 (F OP 2, PeP, St, U N Op 2, SNo). [2n = 24]
Distribution. Nem–NBor(–LAlp). Alt. N Op 1140 m. D fairly common along the North Sea from VJy Blåvands Huk to NJy Skagen, and at Limfjorden from VJy Tyborøn to NJy Ålborg; ØJy rare, mainly at Århus Bugt; FyL Weddelsborg Hoved; LFM locally abundant in eastern Falster and eastern Møn; Sjæ rare, mainly in the northwest and in Stevns. N scattered on sandy shores and scree slopes from ST Melhus, Orkdal and Hitra to Tr Ibestad; rare on inland scree and in willow swamps in Op Lom, Skjåk and SF Stryn; often escaped from cultivation and spreading in sandy places along the coast from Øf to Ro and inland to Bu Nore and Uvdal. S common along most of the coast from central Upl to eastern Nb but almost lacking from northern Gst to central Hls and from northern Mpd to central Ång; in the coastal areas scattered occurrences inland, more or less ephemeral (relictual or bird spread). BhG rare in screes in the central and northern part. Öl a few coastal localities with one to a few individuals, possibly spontaneous there and in western Sk (Öresund). Elsewhere planted and escaped or persisting, most frequently in sandy coastal areas, in the rest of Sk, Gtl, Hl and BhG Göteborg. F common in A and along the Bothnian coast from the SW archipelago of V to KP, less frequent from OP to western PeP; elsewhere introduced and escaped on seashore (at least in U Espoo and Helsinki and EK Kotka) or recorded from dumps, filling earth or along railways (V Nummi-Pusula and Salo, EH Hämeenlinna, Kärkölä and Tampere, ES Kouvola and Lappeenranta and PeP Kemijärvi).
NW Europe, C European mountains, Black Sea area, Caucasus, C Asia, S Siberia, the Himalaya, N China.
Habitat. Sun-exposed places with sand, gravel or stones, and without closing vegetation; calciphilous; very sensitive to competition and shade. Seashores, in particular when subject to land uplift with slight to moderate exposition to the sea; coastal sand-dunes; scree. Usually more or less ephemeral in man-made habitats: gravel- and sand-pits, road cuttings, railways, sandy ditches, edges of fields, dump heaps. From the 1990’s increasingly common in cultivation.
. On land uplift shores the species may form scrub coherent along vast stretches of coast line, keeping pace with rise of new formed land. New shoots and roots grow outwards towards light and empty, surged soil, while the inner, old and high parts of the scrub successively die from competition and shade (Palmgren 1912
) combined with fungal infestations, especially by Phellinus
species (Jahn 1965
). A Hippophaë
stand must slowly shift its position to remain alive, as it is constantly depending on open soil.
The running roots of Hippophaë
bear outgrowths formed by an actinomycete able to bind atmospheric nitrogen into nitrates (Gatner & Gardner 1970
), which considerably improves the nutritional status of the soil. Hippophaë
thus prepares for a luxuriant vegetation in the next phase of succession on the land left behind the moving scrub.
Hippophaë is wind-pollinated; observations by Palmgren (1912) indicate that the wind does not carry the pollen very far, fruit-set being much reduced in females only 10 to 20 m away from males; isolated females, even with males only 30–40 m away, set no fruits.
The fruits are chiefly spread by birds (Darmer 1952
), in Norden especially by late migrating species as crows, thrushes and waxwings (Rousi 1971
). Inland occurrences in the Baltic area have that background rather than being escapes from cultivation.
. Numerous finds of fossil pollen and macrofossils indicate that Hippophaë
was among the earliest and most important late-glacial invaders into Norden. Fossil pollen has been found in Norden from D SJy
in the south to F KiL
in the north (e.g. Sandegren 1943
, Hafsten 1966
). The populations in Denmark, in Norway from ST
north to Tr
and in the Bothnian area are most probably relictual from the pioneer phase, while those in S
have been considered of more recent date (Palmér 1920
) or even very recent (Magnusson 1918
Variation and taxonomy
. There is obvious variation in plant height within Norden; in D
Jylland the shrubs are low and do not reach tree dimensions as may be the case at the Bothnian Sea in S
. This variation was confirmed in cultivation experiments by Rousi (1971)
. Plants from scree and shore localities in N SNo
have narrower leaves than the Baltic population.
According to Rousi (1971)
, the Nordic populations belong to subsp. rhamnoides
(northern and northwestern Europe), which is distinguished by broader leaves from subsp. fluviatilis
Soest (the Alps) and by tortuous shoots and cylindrical rather than spherical fruits from subsp. carpatica
(the Carpathians and southeastern Germany). The genus Hippophaë
has its centre of diversity in Asia; within the widespread H. rhamnoides
five Asian subspecies have been distinguished, and there are a few more species in the Himalaya, Tibet and northwestern China (Swenson & Bartish 2003