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RUBIACEAE Juss. MADDER
(COFFEE) FAMILY
Linda Mary Reeves
(Latin rub = red or reddish, referring to the flowers
or fruit) Annual or perennial herbs, shrubs, vines or trees. LEAVES
entire, mostly opposite or appearing whorled due to presence of
leaf-like stipules, sometimes forming a sheath. INFL
paniculate, cymose, head-like, clustered or solitary.
FLOWER actinomorphic, perfect, occasionally functionally
imperfect; sepals generally 4-5, united, the lobes sometimes reduced
or absent; petals generally 4-5, united, white, greenish, blue,
purple, yellowish, pink or red, often with striped markings; stamens
4-5, epipetalous, alternate with corolla lobes; ovary inferior,
2-4 loculed, style often lobed. FRUIT 2-4 nutlets,
berries, drupes or schizocarps. About 450 genera with about 6500
species. world-wide, mostly tropical. Many important cultivated
genera and species, including Coffea arabica (coffee),
Gardenia jasminoides (gardenia), Cinchona spp.
(quinine), Ixora, Bouvardia, Pentas and other genera
of ornamental note. The Asian genera Myrmecodia and Hydnophyton
contain noted myrmecophyte species. (Dempster, L.M. 1995 Arizona-Nevada
Acad. of Sci. 29: 29-38).
1. Ovule 1 per carpel; fruit a nutlet, mericarp or berry, flowers
small or inconspicuous.............................(2)
1' Ovules several in each carpel; fruit a capsule, flowers conspicuous,
showy...........................Houstonia
2. Leaves opposite..................................................................................................................................Kelloggia
2' Leaves appearing as in whorls...............................................................................................................Galium
Galium L.
Cleavers, Bedstraw
(Greek: gala = milk, from use of Galium vera
for curdling; the common name bedstraw from old use of vine-like
plants with uncinate hairs to grip and bind up straw used for
bedding) Annual or perennial herbs, small vines, subshrubs or
shrubs. STEM squared in cross-section. LEAVES
generally opposite, but may appear whorled with presence of leaf-like
stipules. INFL a panicle or axillary cyme, often
a subcyme of 3 flowers or solitary in leaf axils. FLOWER
perfect or imperfect; calyx absent; corolla usually rotate, sometimes
campanulate or weakly tubular, usually white, cream, greenish
to pink or dark red, the petal lobes usually 4; style 1, deeply
bifid. FRUIT a schizocarp of 2 nutlets, mericarps,
or 2 berries; variously hirsute to glabrous. The fruits are critical
in identifying the species. About 400 species worldwide, mostly
from temperate regions. Two or more species often grow together
in the same habitat, and are often collected and mounted on the
same herbarium sheet.
1. Fruit of two spherical, hard mericarps, black when mature, weakly
joined; leaf apices rounded..............
..............................................................................................................................................................G.
trifidum
1' Fruit various, but not as above; leaves not round at apex..........................................................................(2)
2. Leaves generally, on a single plant, more than 4 per node.........................................................................(3)
2' Leaves 4 or fewer per node..............................................................................................................................(4)
3. Annual, erect or climbing, leaves linear-spatulate or spatulate.................................................................(7)
3' Perennial, generally prostrate; leaves narrowly to broadly
ovoid, vanilla scented; flowers white to
cream..................................................................................................................................................G.
triflorum
4. All flowers perfect; inflorescence/fruit variable...........................................................................................(6)
4' All or some flowers imperfect; inflorescences narrow and diffuse;
fruits with straight straw colored
hairs.....................................................................................................................................................................(5)
5. Plants dioecious, flowers white, cream or pinkish..............................................................G.
coloradoense
5' Plants polygamous; inflorescence diffuse; flowers generally
dark red, rarely pinkish or striped.............
..............................................................................................................................................................G.
wrightii
6. Inflorescence large with numerous, somewhat congested whitish
flowers..............................G. boreale
6' Inflorescence with solitary flower in leaf axil; pedicel long,
reflexed.........................................G.
bifolium
7. Plants generally climbing; leaves linear-spatulate.........................................................................G.
aparine
7' Plants erect or ascending; lower leaves spatulate....................................................................G.
proliferum
Galium aparine L. (old generic name meaning
"to cling") Bedstraw, catchweed bedstraw, cleavers, cleaverwort,
goose-grass, stickywilly. Clambering or prostrate annuals, scabrous
with prickly hairs. STEM 10-90 cm or more long;
nodes often tomentose. LEAVES 6-8 per node, ovate-spatulate
to linear-oblanceolate, 13-31 mm long, bases narrow, apices mucronate,
margins ciliate. INFL few flowered on indeterminate
lateral branchlets, 1-2 in a small cyme or solitary and axillary.
FLOWER perfect; corolla rotate, lobes obtuse
or acute, white or yellowish, especially with age. FRUIT
with short, uncinate, upturned hairs. 2n = 20 (typical
in our area), 22, 42, 44, 63, 64, 66, 86, 88. [Galium spurium
L., Galium vaillantii DC] Moist places in blackbrush,
sagebrush, willow, piñon-juniper, aspen in shade or edges of forests;
talus. Relatively uncommon in our area. COLO: Arc, Hin, Min, Mon;
UTAH. 1130-2600 m (3700-8400'). Flowering: Apr-Jun. Fruit: Jun-Jul.
Western North America from southern British Columbia and Montana
to southern California and central Colorado. Possibly native to
Europe.
Galium bifolium S. Watson (two leaved).
Low mountain bedstraw, twinleaf bedstraw. Erect, slender, glabrous
annuals. STEMS 5-18 cm high. LEAVES
lanceolate or narrowly elliptic, acute, mainly 4 per node, in
unqual pairs, the upper leaves occasionally in pairs; the larger
leaves 0.4-2 cm long. FLOWER perfect; solitary
in leaf axil; pedicel long, reflexed below the fruit; corolla
3 lobed, lobes glabrous, cleft about 2/3 of length, white, ascending,
obtuse, shorter than ovaries. FRUIT with short
uncinate hairs; mericarps nearly separate at maturity. Shaded
slopes and meadows. Rare, but may be mixed and/or misidentified
with other Galium collections. AZ: Apa; COLO: Mon. 1700-2800m
(5500-9200') Flowering: May-Jun. Fruit: Jun-Aug. Western North
America from British Columbia and Montana to southern California
and southwest Colorado.
Galium boreale L. (northern) Northern
bedstraw. Erect perennials, 3-6 dm high, nearly glabrous. LEAVES
4 per node, 13-31 mm long, linear to broadly lanceolate, 3-veined,
minutely scabrous, apex obtuse. INFL a congested
terminal, nearly leafless, more or less pyramidal panicle of cymules.
FLOWER perfect, numerous; corollas cream or white,
sometimes with faint purplish markings which fade in dried specimens,
rotate or a little cupped at base, the 4 lobes ovate, the apices
blunt; ovaries generally densely hairy with short upwardly-curved
hairs. FRUIT dry, with short bristles, the mericarps
becoming reniform. 2n = 44, 66. [G.septentrionale
Roemer & Schultes, G. utahense Eastw.] Sagebrush, meadows,
aspen, spruce-fir streamsides in moist shade. ARIZ: Apa; COLO:
Arc, Hin, LPl, Min, Mon, RGr; NMEX: RAr, SJn. 1770-3050 m (5800-10000')
Flowering: May-Sep. Fruit: Jul-Oct. Montane North America, east
to New Jersey, east Asia, Europe. One of our most common species
of Galium. Plants from North America and east Asia are
hexaploid (2n = 66), differing from those tetraploids of Europe
and west Asia (2n = 44). Ours are subsp.septentrionale
(Roem. & Schult.) Iltis. Roots used as a dye by Great Basin tribes.
Galium coloradoense W. Wight (of Colorado)
Shrubby bedstraw, Colorado bedstraw. Plants perennial, mostly
erect, sometimes shrubby to 30 cm. STEMS slender,
several from woody base, arising from creeping rhizomes, to about
40 cm long, often woody and darker toward the base. LEAVES
mostly 4 in whorl-like clusters, sessile, mostly linear to lanceolate,
apex acute, often reflexed, 5-25 mm long, 2-3 mm wide, with prominent,
often lighter midrib below, glabrous to scabrous. INFL
small and few flowered, borne on short axillary branches, occasionally
larger and more congested. FLOWERS dioecious,
pedicellate; corolla greenish-yellow, inconspicuous, sometimes
with purplish markings which fade in dried specimens, 2-4 mm wide,
often scabrous-hispid externally. FRUIT with
long, spreading, flattened, white or straw-colored hairs 1-3 mm
long. 2n = 22. [G.multiflorum Kellogg var. coloradoense
(W. Wight) Cronq] Shaded rocky or sandstone crevices and cliffs
in desert scrub, sagebrush, mountain brush and piñon-juniper.
Relatively common. ARIZ: Apa, Nav; COLO: Dol, LPl, Mon; NMEX:
SJn: UTAH. 1460-3100 m (4800-10,100') Flowering: May-Aug. Fruit:
May-Oct. Eastern Utah, western Colorado, northeastern Arizona,
northwestern New Mexico. A highly variable species.
Galium proliferum A. Gray (bearing many
offspring). Annuals to 30 cm high, erect to stiffly ascending,
glabrous to hairy. LEAVES mostly 4 per node;
3-9 mm long; the lower petiolate and spatulate; the upper more
or less sessile, linear to obovate, more or less remote. INFL
a small cyme with short branches at nodes or flowers solitary.
FLOWER perfect, minute; short-pedicelled; corolla
tiny; lobes erect, apices rounded; white, often tipped with pink.
FRUIT with 2 dark, kidney-shaped mericarps, downturned
from two erect leaves; hairs long, straight or weakly uncinate.
Damp locations near streambeds, washes, hanging gardens or under
shrubs. Uncommon. UTAH. 1620 m (5300') Flowering: Mar-Jun. Fruit:
Jun-Sep. Southern California to Nevada, Arizona, New Mexico, the
Edwards Plateau and Trans-Pecos region of Texas south through
Baja California and Sonora.
Galium trifidum L. (three-parted). Threepetal
bedstraw. STEMS usually perennial, thin, lax;
internodes widely spaced, 6-12 cm. long; glabrous. LEAVES
2-6 per node, mostly 4, sessile, apex rounded, 4-12 mm long; linear
to obovate; glabrous. INFL with 1-4 flowers,
sometimes subtended with a minute bract or scale. FLOWER
perfect; pedicel slender, long; corolla rotate, white to greenish-white,
rarely striped with pink, generally 3 lobed, lobes ovate, ascending.
FRUIT with 2 mericarps, spherical, glabrous or
with a few short, flat hairs or scales, black or dark brown. 2n
= 24. [G. brandegei A. Gray, G. columbianum
Rydb.] Margins of lakes, ponds, streams, mountain brush, ponderosa
pine, spruce-fir. May be locally common at higher elevations.
Often grows with or near G. boreale. ARIZ: Apa; COLO:
Arc, LPl, Hin, Min, Mon, SJn; NMEX: SJn. 2470-3400 m (8100-11100').
Flowering: Jun-Jul. Fruit: Jun-Oct. Circumpolar, extending southward
in North America along the Rocky Mountains. Our plants belong
to var. pusillum A.Gray.
Galium triflorum Michx. (three-flowered).
Fragrant bedstraw, sweet-scented bedstraw. STEMS
perennial, herbaceous, 10-50 cm long, decumbent or viney, growth
radiating from a central rhizome or root. LEAVES
generally 5- 6 per node, 5-40 mm long, elliptic to ovate-obovate,
base gradually narrowing, apex apiculate or mucronate, glabrous
to scabrous. INFL 1-5 flowered, usually 3, pedicellate
in axillary, open, often divaricate cymules with two subtending
bracts. FLOWER perfect; corolla rotate, cream-colored
or greenish, ovary about as long as or longer than corolla. FRUIT
covered with soft white, cream or golden brown hooked hairs; mericarps
dark papillate or furrowed, curved. 2n = 22, 44, 66.
[Galium brachiatum Pursh] Shady forest floors or moist
streamsides under brush, ponderosa pine, Douglas fir and aspen.
A common species in our area. Often grows with or near G.
boreale. ARIZ: Apa, Nav; COLO: Arc, Hin., LPl, Hin, Min,
Mon, SJn; NMEX: RAr, SJn; UTAH. 1650-3290 m (5400-10800'). Flowering:
May-Aug. Fruit: Jun-Sep. Circumboreal, south to California, Mexico
and Florida in North America.
Galium wrightii A. Gray. (for Charles
Wright, botanist). Wright's bedstraw. Polygamous erect subshrubby
perennials. STEMS suffrutescent, several to many,
5-50 cm high, glabrous to scabrous. LEAVES 4
per node, two often below flowers, 7-20 mm long, linear to oblanceolate
with broad insertions, apex acute; glabrous, scabrous or hispid,
with hookless hairs. INFL diffuse, branchlets
spreading. FLOWER sometimes subtended by a single
bract; corolla dark red to pink, rarely pale or striped, rotate,
lobe tips slender, acute. Helpful to see flowers in this species.
FRUIT often with reddish papillae, hairs few
to many, long, straight. 2n = 22. [Galium rothrockii
A. Gray]. Moist, semi-shady slopes under ponderosa pine, Douglas
fir and aspen, hanging gardens, cliffs. Uncommon, but may be locally
abundant. ARIZ: Apa; NMEX: SJn. 1980-3820 m (6500-9900'). Flowering:
Jun-Sep. Fruit: Jul-Oct. Texas to California, south to Baja California,
Coahuila and Sonora, Mexico.
Houstonia
L. Bluet
(For William Houston, who collected in tropical America). Small
annual or perennial herbs. STEMS upright. LEAVES
opposite with interpetiolar stipules. INFL cymose
or flowers solitary. FLOWER perfect, calyx 4-lobed;
corolla 4-lobed, funneliform or salveriform; stamens 4; stigmas
bifid. FRUIT a capsule, bilocate, inferior. SEED
crateriform. 50 species, U. S. and Canada to Mexico. Closely allied
to the more tropical genus Hedyotis.
Houstonia rubra Cav. (red). Red bluet.
Caespitose perennials with small woody taproots. STEM
1-10 cm tall, slender. LEAVES linear to oblanceolate,
5-30 mm long, 0.5-4 mm wide, erect or ascending. FLOWER
usually one per node, sessile or with a short pedicel to 4 mm
long, corolla 8-40 mm long, salveriform, red or pink to purple
or rarely, white; heterostylous. FRUIT 2-3.5
mm long, 2-5 mm wide, walls thick on recurved pedicel. 2n
= 22. [Hedyotis rubra (Cav.) A. Gray, Houstonia saxicola
Eastw]. Blackbrush, rocky grassland, pin~on-juniper, washes. ARIZ:
Apa, Nav; UTAH. 1680-2650 m (5500-8700'). Flowering: Apr-Oct.
Fruit: May-Oct. Texas to Utah, south to southern Mexico. Puebloan
tribes used an infusion of the plant for sore eyes and stomach
upsets. Navajos used a decoction of the plant for menstrual problems.
Kelloggia
Torr. ex Benth. Kelloggia
(For Albert Kellogg, nineteenth century California botanist). Perennial
herbs. LEAVES opposite with small interpetiolar
stipules. INFL a loose terminal cyme. FLOWER
4-5 merous; corolla funnelform; ovary 2-loculed, each with one
ovule. FRUIT a schizocarp splitting into two
mericarps. 2 species, one from western North America, the other
from China.
Kelloggia galioides Torrey (resembling
Galium). Milk kelloggia. STEMS erect, several,
15-40 cm long. LEAVES opposite, fascicled, 19-38
mm long, lanceolate to narrowly ovate, minutely serrulate, apex
acute; stipules hyaline, 0.5-2 mm long, lanceolate, bifid or erose.
INFL few flowered. FLOWER with
pedicel filiform, 6-25 mm long, swelling below the junction with
flower, becoming divergent; calyx curved inward toward the corolla;
corolla 4-8 mm long, pink or white, outer surface pubescent, inner
surface with tufts of hairs, tube slender, lobes spreading. FRUIT
obovoid, 3 mm long, covered with flat transparent hooked hairs.
Moist slopes, under piñon-juniper, sagebrush, mountain brush,
Douglas fir, ponderosa pine or in meadows. ARIZ: Apa; NMEX: McK,
SJn (Falling Iron Cliffs); UTAH. 1770-2670 m. (5800-8700'). Flowering:
Jun-Sep. Fruit: July-Oct. Washington south to Baja California
along the coast; Rocky Mountains from Arizona and New Mexico to
Wyoming.
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