Elaeagnaceae Taxa treated:
Elaeagnus angustifolia
Elaeagnus commutata
Elaeagnus umbellata
Hippophaë rhamnoides
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by Bengt Jonsell

(6b, 20070529)

 © Flora Nordica

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This treatment is partly outdated - the final version will appear in print.
1 Dioecious; twigs with numerous thorns
- Flowers bisexual; twigs without or with few thorns

1. Hippophaë L.

Linnaeus, Sp. pl.: 1023 (1753)

Hippophaë rhamnoides L.     Map

Linnaeus, Sp. pl.: 1023 (1753) - Type not designated.
D Havtorn. F tyrni. N tindved. S havtorn.
Literature. Palmgren 1912, Rousi 1965, 1971.
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.
Biology. 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.
History. 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 and Brn 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 northern BhG 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 and F. 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 Rousi (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).

2. Elaeagnus L.

Linnaeus, Sp. pl. 121 (1753).
Phanerophytes. Deciduous shrubs or small trees; young shoots and leaves more or less densely covered by silvery, peltate, fringed scales. Leaves alternate, petiolate, undivided; margin entire. Inflorescences axillary, compact clusters of one to rather few flowers. Flowers bisexual, epigynous, usually with a constriction above the tubular hypanthium and a nectariferous disk above the constriction; perianth campanulate, 4-lobed. Stamens 4, inserted inside the upper part of the perianth; filament very short. Ovary superior, with one style and an oblique stigma. Fruit a drupe with an ellipsoid, ribbed stone.
1 Leaves lanceolate, up to 1 cm wide; lower leaf surface without brown scales
- Leaves elliptic or ovate, most of them wider than 1 cm; at least a few brown scales on lower leaf surface
2 Leaves silvery above from dense scales also as old; flowers single or rarely up to 5 few together
- Leaves soon green above; most flowers 5-8 together in axillary clusters

1. Elaeagnus commutata Bernardi ex Rydb.     Map

Rydberg, Fl. Rocky Mountains: 582 (1918). - E. argentea Pursh (1814) non Moench (1794) - Described from North America.

D . F (lännen)hopeapensas. N sølvbusk. S silverbuske.

Shrub with suckers, extensively branched, up to 4 m. Shoots as young with a dense cover of deeply fringed brownish scales and some stellate hairs; old shoots with dark-brown, somewhat pruinose bark. Leaves rather densely arranged along the shoots; petiole 2–9 mm; blade grey above, silvery grey below, narrowly to broadly elliptic or ovate, acute to obtuse, rounded at base, (2.2–)3.5–8 x (0.8–)1.5–5 cm; both surfaces densely covered with silvery scales (on lower surface also single to scattered brownish ones, in particular along the midvein).
Flowers usually 2–3 together in leaf axils, silvery grey from densely covering scales outside, red-brown inside; pedicels 3–5 mm; perianth 10–14 mm (including 2.5–5 mm long triangular lobes), constricted 2–4 mm from the base, with few stellate hairs inside, mainly along the margin; style base surrounded by an inconspicuous disk. Fruit ellipsoid to ovoid, yellow, mealy, c. 1 cm. – Early summer.
[2n = 28]
Distribution and habitat. From the beginning of the 20th century increasingly grown in parks and gardens north to northern Norway and Sweden and central Finland. Sometimes persistent after cultivation or escaped, spreading by means of extensive suckers. Open, rather dry, gravelly to sandy places; roadsides, ditches, railways, waste places, gravel pits, abandoned sites. D scattered in most parts. S scattered or rare north to Vsm and Upl. N a few places in the south (Øf Rakkested, Ak Ås, Bu Drammen, VA Farsund) and in the north NNo Hamarøy. F reported as escaped from dozens of places north to southern PK, but usually a mere relic of cultivation; no confirmed established occurrences.
N North America.

2. Elaeagnus angustifolia L.     Map

Linnaeus, Sp. pl. : 121 (1753). - Type: Clifford Herbarium 38 Elaeagnus 1 (BM) lectotype, sel. by McKean., Regnum Veg. 127: 44 (1993).

D . F idänhopeapensas. N smalsølvbusk. S smalbladig silverbuske.

Extensively branched shrub, to 7 m in cultivation. Shoots as young with a dense cover of silvery scales; old shoots with violet-brown, somewhat pruinose bark. Leaves rather densely arranged; petioles 2-7 mm; blade grey-green above, silvery grey beneath, lanceolate, subacute, cuneate at base, (2.5-)3-7 x 0.4-1 cm; upper surface with scattered scales, lower surface with a dense cover of silvery scales (and sometimes also with very few brownish ones along the midvein).
Flowers 1-3 together in leaf axils, silvery grey with dense scales outside, red-brown inside; pedicels 3-4 mm; perianth 8.5-13 mm (including 3-4 mm long, triangular lobes), constricted 2-4 mm from the base, with scattered stellate hairs inside; style base surrounded by a prominent conical disk. Fruit ellipsoid, yellow, mealy, 1-2 cm. - Early summer.
[2n = 28]
Distribution and habitat. Rather little cultivated in D and southern S; rarely escaped on sandy ground, e.g. in waste places and harbours. D LFM Nakskov 1972, Brn Sandvig 1971. S Sk Bjärred 1991, Glimåkra 1995, Kristianstad 1998, Malmö 2002.
W and C Asia, the Himalaya, Mongolia.

3. Elaeagnus umbellata Thunberg     Map

Thunberg, Fl. Jap.: 66 (1784).- Type: Thunberg Herbarium 3725 (UPS) holotype.
D S japansk silverbuske.
Shrub, to 4 m in cultivation. Shoots as young brown to silvery grey with scattered to dense silvery scales; old shoots with grey to olive-green bark. Leaves densely arranged; petioles 5–8 mm; blade green above, silvery grey below, elliptic to oblanceolate, subacute to acute, cuneate at base, 2–9 x 1–4 cm; upper surface with scattered scales, lower surface with a dense cover of silvery scales (and very few scattered brown ones).
Flowers usually 5–8 together in leaf axils, silvery grey with dense scales outside, lobes yellow inside; pedicels 0.5–4 mm; perianth 8–9.5 mm mm (including c. 4 mm long lobes) without distinct constriction; style base surrounded by a prominent conical disk. Fruit spherical, red, juicy, 0.5–1 cm. – Early summer.
[2n = 28]
Distribution and habitat. A fairly rare ornamental in D, occasionally escaped: NJy Farø 2001, VJy Holstebro 1995, Nørre Snede 2000.
Japan, Korea, China.

References   To top

Darmer, G. 1952: Der Sanddorn als Wild- und Kulturpflanze. 89 pp. Leipzig.
Gatner, E.M.S. & Gardner, I.C. 1970: Observations on the fine structure of root nodule endophyta of Hippophaë rhamnoides L. - Arch. Microbiol. 70: 183-196.
Hafsten, U. 1966: Den senkvartere forekomst av tindved (Hippophaë rhamnoides L.) i Sør-Norge. - Blyttia 24: 196-215.
Jahn, H. 1965: Die Phellinus robustus var. hippophaës - Ph. contiguous -Ass., eine Pilzgesellschaft auf Sanddorn. - Westf. Pilzbr. 5: 139-141.
Magnusson, A.H. 1918: Om Hippophaës rhamnoides L. i Bohuslän. - Svensk Bot. Tidskr. 12: 471-476.
Palmér, J.E. 1920: Hippophaës rhamnoides L. i Bohuslän. - Svensk. Bot. Tidskr. 14: 88-90.
Palmgren, A. 1912: Hippophaë rhamnoides auf Åland. - Acta Soc. Pro Fauna et Flora Fenn. 36 (3): 1-188.
Rousi, A. 1965: Observations on the cytology and variation of European and Asiatic populations of Hippophaë rhamnoides. - Ann. Bot. Fennici 2: 1-18.
Rousi, A. 1971: The genus Hippophaë L. A taxonomic study. - Ann. Bot. Fennici 8: 177-227.
Sandegren, R. 1943: Hippophaë rhamnoides L. i Sverige under senkvartär tid. - Svensk Bot. Tidskr. 37: 1-26.
Swenson, U. & Bartish, V. 2002: Taxonomic synopsis of Hippophaë. Nordic J. Bot. 22: 369-374.