Empetrum nigrum subsp. hermaphroditum (Lange ex Hagerup) Böcher
Crowberry, Curlewberry.
Empetraceae, crowberry family.
Medd. Grønl. 149, 9: 81. 1952.
Nomenclatural section used by Flora of North America project Empetrum
hermaphroditum Hagerup, Dansk Bot. Ark. 5, 2: 1. 1927.
Empetrum
eamesii Fernald and Wiegand subsp. hermaphroditum (Hagerup)
D.Löve, Rhodora, 62: 289. 1960.
Empetrum hermaphroditum Lange ex Hagerup, Dansk Bot. Archiv. 5(2):
16.1927.
Empetrum nigrum var. hermaphroditum (Lange ex
Hagerup) T.J. Sorensen, Medd. Gronl. 101(3): 95. 1933.
Empetrum
eamsii subsp. hermaphroditum (Lange ex Hagerup) D. Löve,
Rhodora, 62: 289. 1960.
Vegetative morphology. Plants shrubs; dwarf shrubs, or low shrubs; 5–10(–30) cm high; prostrate, matted, freely branching, evergreen, with trailing branches. Ground-level or under-ground stems not developed horizontally or vertically (but branches are usually prostrate). Aerial stems prostrate (usually), or ascending (occasionally growing somewhat erect to 20–30 cm high in warmer sites, or by rocks); with glandular hairs; sparsely hairy, or densely hairy; stem hairs spreading, or erect. Branches epidermis flaky (older stems). Leaves distributed along the stems; alternate (close together and approaching whorled); simple; evergreen. Petioles absent. Leaf blade bases attenuate. Blades 2–6 mm long; 0.5–1 mm wide. Blades spreading, or divaricate; leathery, or succulent (deeply furrowed on the lower side, the groove almost closed by the overlapping marginal glandular hairs, forming a central longitudinal cavity (see image library)); straight; linear; revolute; with inconspicuous veins. Blades adaxial surface glabrous (superficially the leaves appear glabrous), or hairy (glandular hairs seen at 40x). Blades adaxial surface hairs glandular (tiny). Blades abaxial surface hairy. Blade margins with non-glandular hairs (these overlapping along the revolute leaf margins).
Reproductive morphology. Plants monoecious, or dioecious (usually), or bisexual. Flowering stems absent. Flowering stems with leaves (single flowers are on "short shoots" that arise laterally from the main axis and bear only scale leaves below the flower). Flowers solitary; small, less than 5 mm in diameter or length (1–2 mm). Perianth present. Calyx sepals 3; free; 0.2–0.3 mm long; 0.08–0.12 mm wide. Calyx green, or purple (green at base, purple towards the apex); petaloid; without sessile glands; glabrous. Petals free; 3; purple; 1–2 mm long. Flowers bisexual. Stamens 2. Anthers ovoid (without appendages); 0.6–0.8 mm long. Nectaries absent. Gynoecia superior. Carpels syncarpous; (2–)4–9. Ovaries subglobose; glabrous. Styles variable; partially fused (hollow, the cavity fluted in alignment with the carpels). Placentation axile. Ovules 2–5. Fruit sessile. Fruit with calyx persisting; fleshy; a drupe; spherical; indehiscent. Fruit 3–8 mm long; 3–8 mm wide; black, or purple, or blue; glabrous; surface appearing veinless. Seeds 2–9; 1.6–1.8 mm long; brown (to pale reddish); with surfaces verrucose (minutely).
Chromosome information. 2n = 52. Hagerup (1927 Greenland?); Arwidsson (1938 northern Europe); Flovik (1940 Svalbard); Löve in Arwidsson (1943 northern Europe); Löve and Löve (1956b Iceland, 1959 northeastern North America, 1961d northern Europe, 1966b north eastern USA); D. Löve (1960 north eastern North America); Hedberg and Hedberg (1964 Sweden); Ulsaker in Engelskjøn (1979 Norway); Engelskjøn (1979 Svalbard); Löve and Löve (1982a central Canada). Several more southern counts. Supposed basic chromosome number of family 4x.
Distribution. Northern hemisphere distribution: amphi-Atlantic; Greenland, Canada, Eurasia. Northwest Territories Islands, Continental Northwest Territories, Nunavut Islands, Continental Nunavut, Northern Québec. Arctic Islands: Baffin, Ellesmere, Victoria (literature record), Axel Heiberg, Somerset (literature record), Southampton.
Ecology and habitat. Substrates: slopes, ridges, seashores; dry;
acidic; rocks, gravel, sand; with low organic content. Habitats: Often reported
on rocky or gravely slopes, E. nigrum subsp. hermaphroditum is
also found on sandy or gravely upper beaches, or on flats near the seashore. It
is associated with Vaccinium and Cassiope, and also occurs with
Pyrola grandifolia.
Growth responses of Empetrum nigrum and
Vaccinium vitis-idaea to environmental manipulation in the Finish
sub-Arctic were investigated by Shevtsova et al. (1997).
Indigenous knowledge. Inuit name, paurngait, blackberries.
Ootoova, et al. (2001) reported that the word paurngait means "which
looks like pauq (soot) because they are black in colour. They are very
healthy to eat, juicy and crunchy. The word paurngaqutiti is used for
crowberry patches. The berries were recommended for a person who had diarrhea.
Paurngait are more effective in hardening watery stools than
Kigutangirnait (blueberries, bilberries). If you eat a lot of
Paurngait berries the stool hardens. It is okay if you eat them with
meat, but if you just eat the berries alone, you'll get constipated. They are
delicious when mixed with caribou fat. Today paurngait are also eaten as
jam. As the weather gets colder, crowberries and other berries start to freeze
on the ground. When it starts melting again, berries from the year before become
visible and can even be eaten then. Augujjiaq refers to picking berries
with a bowl with holes in it to drain the snow out. The bowl was used as a
filter. People would dig in the snow, and put the berries into the bowl. The
berries were not dried up. To store the berries, a hole could be made in the
sand and some fat poured in. When the fat hardened the berries were put in, and
the hole covered with seal skin. After the hole was covered it was further
buried for a winter supply. The berries were found to keep very fresh."
Crowberry branches made nice mattresses for iqluit (Loosely woven
summer mattress. Interpretation by N. Hallendy 2003).
The branches with the
leaves attached were used to clean gun barrels (Ootoova et al. 2001).
The
purplish-black shiny fruits are very juicy and sweet but contain a number of
large, hard seeds (Porsild 1957).
In Alaska, large numbers of crowberries
were picked in late summer and stored in seal oil for use in fall and winter
(Ager and Ager 1980).
Andre and Fehr (2000) noted that the Gwich'in elders
reported these berries make good jam and are tasty when eaten alone or when
mixed with other berries. They are used to make it'suh, a desert prepated
from pounded dryfish. (Recipe: take the fish broth from boiled white fish, add a
pail of blackberries, enough sugar to sweeten it up and a dipper of fish blood,
heart and liver. Cook this mixture until it is just like jam).
Andrew and
Fehr (2000) noted that a tea was made by collecting and boiling the roots,
berries and stems of this plant and that some Gwich'in people considered it as
good as spruce gum tea for stomach aches and bad colds.
Porsild (1950)
stated that because of its abundance and hardiness, the crowberry, although not
as well-flavoured as some other berries, is easily the most important fruit in
Arctic regions.
Rink (1857) claimed that a sparkling white wine may be
produced by fermentation of the juice.
Notes. Jordal (9952) recognized a forma ciliatrum naming a
specimen collected from Old John Lake, in the Brooks Range of Alaska. He
commented that this form has strongly villous-ciliate leaves and is undoubtedly
only of sporadic occurrence. The type is Jordal 2415, deposited in the
university of Michigan herbarium.
Cody (1996) noted that flowers in this
subspecies are polygamous.
Kuvaev et al. (1996) studied the anatomy,
morphology and chemical composition of four Empetrum species in northern
Siberia, E. hermaphroditum, E. nigrum, E. sibiricum, and
E. subholarcticum V. Vassil and recognized three groups E. nigrum,
E. hermaphroditum, and E. subholarcticum, on the basis of the
flower sexuality.
Kudo et al. (1999) studied the effects of snow-melt timing
on leaf traits (for five deciduous and five evergreen species), shoot growth,
and leaf production (for five evergreen species) of tundra plants along a
snow-melt gradient in an alpine snowbed in northern Sweden. In deciduous plants,
leaf life-span and leaf mass per area (LMA) decreased, and nitrogen
concentration (leaf N) increased with decreasing growing season, whereas in
evergreen plants, both leaf life-span and leaf N increased with decreasing
growing season. By extending leaf life-span, evergreen plants are able to have a
large leaf mass, which may contribute to maintain net annual carbon gain in
short snow-free seasons. In two predominantly boreal evergreen species,
Empetrum hermaphroditum and Vaccinium vitis-idaea, leaf life-span
was negatively correlated with both annual leaf production and shoot growth, but
there were no similar significant correlations for the other three, strictly
arctic-alpine evergreen species that these authors studied.
Phytochemistry. The content and composition of sugars in the juice of
the northern crowberry, E. nigrum subsp. hermaphroditum were
studied by Kallio et al. (1984). The sugars in the pressed juice were purified
with ion exchange resins, and were analyzed with specific gravity analyses. The
content of glucose was 2.8 &#;+&#;none 0.1 g/100 ml, fructose 2.6&#;+ &#;none0.1
g/100 ml and the sugar alcohol inositol 0.02 g/100 m. Traces of galactose were
identified. Sucrose was absent in all samples (less than 0.01 mg/100 ml).
Hethering et al. (1984) studied the lipid composition of the upland E.
nigrum subsp. hermaphroditum and E. nigrum subsp.
nigrum with a lowland distribution and found no evidence to support the
suggestion that lipids (triacylglycerols) play a major energy storage role in
the leaves of the alpine taxon. On a quantitative basis the triacylglycerols
constituted less than 5% of the total lipid in both species and the lowland
species possessed the higher levels of total lipid, neutral lipid, and
triacylglycerols. These authors suggested that the presence of a well-developed
waxy cuticle might account for the high total lipid levels encountered in dwarf
evergreen shrubs.
Puntari et al. (1985) extracted volatiles from the black
berries of the crowberry, Empetrum nigrum with steam distillation and
capillary GLC-MS analysis. Thirty-three volatile compounds were identified,
among which benzoic acids and benzyl alcohol were the main ones. Fifteen
compounds known to exist in blueberries (bilberry, bog blueberry and high-bush
blueberry) were found in crowberry. Esters of hydroxy acids, which are most
important and characteristic of bilberries were totally absent which explains
the very mild aroma of crowberry that is otherwise similar to that of
blueberries. The aromatic components of the two subspecies studied, E.
nigrum subsp. nigrum and subsp. hermaphroditum, were very
similar to each other.
Saratikov et al. (1991) found anticonvulsive activity
in an ethanol extract from the above ground part of E. nigrum and
compared it with that of benzonal in experiments carried out on mice and rats.
The extract exhibited anticonvulsive properties in the models of corazole-,
camphor-, strychnine-, and nicotine-induced convulsions and at maximal
electroshock. Benzadiazepine tranquilizers increased the potency of the
anticonvulsive effect of the extract. The extract prolongs the effects of
sedative and exhibits moderate H-cholinolytic, hypothermic and analgesic
activity.
Matsuura et al (1995) documented the occurrence of antibacterial
and antifungal compounds in Empetrum nigrum.
Pollen
morphology. The pollen grains of Empetrum were studied using both
light and electron microscopy by Warner and Chinnappa (1990). They found that
the grains are the typical tetrad form of most Ericales and that E.
nigrum has tetrahedral tetrads with 12 compound costae. The tetrads of E.
nigrum subsp. hermaphroditum are larger than those of subsp.
nigrum and possess 12–16 compound costae per tetrad, which the
authors suggested is related to polyploidy.
Cytology. Inorthern
Europe the distribution of the cytotypes E. nigrum L. and E.
hermaphroditum (Lange) Hagerup were mapped based on literature references
and original data (Velluti et al. 1995). These authors showed that the diploid
taxon, E. nigrum, 2n = 26, is mainly distributed in central Europe
and is rare in the Alps. The tetraploid taxon, E. hermaphroditum ,
2n = 52, has a more northern distribution and is frequent in the Alps and
northern Apennines. In the geographic areas where both taxa are present, they
are never simultaneously present in the same habitats. The diploid taxon is
almost exclusively linked to peat-bogs and the tetraploid taxon to windy peaks
with scarce and short-term snow cover.
Wood anatomy. Carlquist (1989)
analyzed the wood and bark of 12 collections of the Empetraceae and found that
the family has vessels that are somewhat angular in cross section, with
scalariform perforation plates and scalariform to opposite vessel-ray pitting.
Imperforate trachery elements are all trachieds. Axial parenchyma is sparse and
not subdivided. Rays are characteristically uniseriate and composed of upright
cells. Older stems have rays with both upright and procumbent cells. These
features ally Empetraceae closely to Ericaceae and Epacridaceae. The narrow
vessels are quite numerous per mm squared and denote a high degree of wood
xeromorphy. The growth rings and tracheids may also be indicative of adaptation
to drought or physiological drought due to cold. The woods demonstrate the
accepted criteria for paedomorphosis with rays composed of upright cells,
nonconversion of the uniseriate rays to multiseriate or heterocellular rays, and
a decrease in vessel element and tracheid length with age.
Phytogeography. Th elimit in Canada between the amphi-Atlantic subsp.
hermaphoroditum and the amphi-Beringian subsp subholarcticum is
not clear. The ellesmer, Baffin Hudson Bay, and Ungava plants are subsp.
hermaphroditum and it is most probable that the plants of Southampton and
SOmerset belong here. If a voucher for the literature record from Victoria
Island can be found it should be investigated as plants that occur there may be
subsp. subholarcticum.
Illustrations. • Habitat of Empetrum and Cassiope. Empetrum (in front of white marker) growing beside Cassiope with the grass Hierochloe alpina (background) on a rocky hillside. Nunavut, Baffin Island, at Iqaluit. Aiken 97–005 and 97–006. Scale bar in cm. • Undersurface showing branching stem. Underside of an Empetrum plant that had the soil washed out from around the stem and roots. Found at high water level, on the banks of the Soper River, Nunavut, Baffin Island. Aiken 2002. No voucher. • Close-up of stem apex. The underface of leaves at the stem apex. The upper adaxial surface is a darker green and rolled over the lower surface that appears as a pale green stripe when the leaves are young and is less obvious in mature leaves. Nunavut, Baffin Island. S. Aiken 4 July, 2002. No voucher. • Close-up of flowering stem. Flowering stem with a flower borne in the leaf axil. Nunavut, Baffin Island, Iqaluit. Aiken 4 July, 2002. No voucher. • Plant in habitat. Plant tending to die in the centre as seen by the brown leaves. Note immature green fruit. Plant growing on rocky till. Nunavut, Baffin Island, Iqaluit, July 22, 1982, J.M. Gillett 18987. CAN. • Close-up of plants. Note dying brown leaves in the centre of the shrub and immature green berries. Nunavut, Baffin Island, Iqaluit. July 22, 1982, J.M. Gillett 18987. CAN. • Habit. Late-season plants with mature blue berries. Note needle-like leaves that are evergreen. Nunavut, Baffin Island, Iqaluit. Aiken 97–005. CAN. • Close-up of fruit. Mature blue, Iqaluit, "blackberries". The leaves are cylindrical. The leaf margins almost meet and surround an internal air space. Nunavut, Baffin Island, Iqaluit. Aiken 97–005. • Underside of fruits. Underside of fruits showing three creamy sepals, three browning red petals and narrow brown stamen filaments some with the remains of anthers. These indicate this is subspecies hermaphroditum, c.f. subsp. nigrum. Photographed by A. Brysting in Manitoba, Churchill. No voucher. • Arctic Island Distribution.
Cite this publication as: ‘S.G. Aiken, M.J. Dallwitz, L.L. Consaul, C.L. McJannet, L.J. Gillespie, R.L. Boles, G.W. Argus, J.M. Gillett, P.J. Scott, R. Elven, M.C. LeBlanc, A.K. Brysting and H. Solstad. 1999 onwards. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 29th April 2003. http://www.mun.ca/biology/delta/arcticf/’. Dallwitz (1980) and Dallwitz, Paine and Zurcher (1993, 1995, 2000) should also be cited (see References).
