THE GENUS GALIUM (RUBIACEAE) IN SOUTH AMERICA.

II
Author(s): Lauramay T. Dempster
Source: Allertonia , July, 1981, Vol. 2, No. 8 (July, 1981), pp. 393-426
Published by: National Tropical Botanical Garden

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 THE GENUS GALIUM (RUBIACEAE) IN SOUTH AMERICA. II.
Lauramay t. Dempster1

The quality for which the genus Galium (bedstraw) is popularly known is i
tendency to stick to clothing, although many species in fact lack this character. Tho
species that are adhesive are so by virtue of either or both of two mechanisms:
minute, retrorse, more or less aculeate hairs on salient portions of the stems and leave
and (2) hooked (uncinate) hairs on the fruits. Both of these mechanisms are frequent
and worldwide throughout the genus.
The first paper in this South American series (Dempster, 1980) dealt with tho
species of Galium having straight, rather than uncinate, fruit hairs (sect. Lophagaliu
K. Schum.). The present paper deals with those species of which the fruits ha
uncinate hairs, and a third paper is planned to deal with the remaining species, i
those that lack specialized fruit hairs.
The present group includes 16 species (one of which is divided into two subspecies
all but three of them being perennial and native to South America. Of the th
annuals, two (Galium murale and G. parisiense) are introduced, and one (G. aparine) is
cosmopolitan, although apparently absent from southern Mexico and Central Am
ica. Two species (G. orizabense and G. mexicanum, both occurring in Colombia a
Venezuela) are believed to be conspecific with Mexican and Central American plan
Another, G. canescens, is morphologically very close to its Central American counter
part, G. uncinulatum DC. (see discussion under G. canescens). The other ten spec
are distinct and are limited to South America. Among these, four new species, one n
subspecies, and one new name are here proposed.
Like the species of sect. Lophogalium (Dempster, 1980), the species with uncin
hairs are principally monticolous plants, with altitudes ranging from 1,000 to 4,400
except for the annual species (see discussion under Galium aparine) and G. diffu
ramosum, of Chile, which grows nearer the coast at 100 to 700 m. The territ
occupied by the group (excluding the annuals) extends from the tip of Tierra del Fue
(G. fuegianum) northward in the Andean region into the mountains of Venezuel
far east as Monagas (G. canescens). In addition, G. orizabense extends into Centr
America and far into Mexico. Galium mexicanum subsp. mexicanum also occurs
Central America, Mexico, Arizona, and western Texas.
Although hermaphroditic flowers are the rule in this group, female-sterile flowe
have been observed in several species, viz. Galium boyacanum, G. canescens, G
diffusoramosum, and G. weberbaueri.
The grouping together of Galium species having uncinate fruit hairs is practical a

■The Jepson Herbarium, Department of Botany, University of California, Berkeley, California 947

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 ALLERTONIA

Figure 1. Some inflorescence patterns in Galium·, a, flowers solitary in axils, each w

peruvianum)·, b, flowers in threes in axils (G. fuegianum and G. obovatum); c, flow
huancavelicum)·, d, branchlets indeterminate, with many flowers of various ages (G. ca
num, G. weberbaueri, etc.).

convenient, but probably not "natural". The perennial species with fo

node are probably more closely related to each other, regardless of the t
the fruits, than they are to the annual species G. aparine, or perha
perennial species G. pseudotriflorum and G. mexicanum. This opinio
support by the discovery of both straight and uncinate hairs on th
ovalleanum (Dempster, 1980). Although the annual G. aparine and th
pseudotriflorum are similar enough to be easily confused, it is doubtful
similarity is indicative of very close relationship rather than convergen

TAXONOMIC METHODS AND CRITERIA

This is an herbarium study, based on material from the following inst

bm, br, c, col, conc, cord, e, f, g, gb, gh, goet, hal, hut, K, lil, lp, m, mo, ny, p, pr,
r, rb, s, sgo, u, uc, us, usm, ven, and wis. Abbreviations for herbaria are those

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

recommended by Index Herbariorum (Holmgren & Keuken, 1974). Tha

the administrators of the listed herbaria for the indispensable use of spe
their care. Very little field work was done by me in South America, a
some counts were made. All holotypes and specified isotypes have b
unless otherwise stated.
In this group of species several characters have been found to be especially useful
for taxonomic purposes. Several of the species have a definite and quite consistent
pattern of inflorescence branching (viz. Galium obovatum, G.fuegianum, G.peruvia
num), which helps to distinguish them from other, somewhat similar species with less
clearly definable inflorescences. In connection with reduction in complexity of flower
ing branchlets, there is a concomitant tendency toward fewer branchlets, as illustrated
by Figure la and lb. This character is, however, not constant, and usually takes the
form of alternate rather than opposite branchlets.
The direction of the apical curvature of the fruit hairs, i.e., whether upward toward
the corolla or downward away from the corolla, is remarkably constant in most
species. An exception is Galium pseudotriflorum, in which the tips of the hairs usually
turn downward; but when the tips turn upward, as in some plants they do, the bases of
the hairs turn downward, so that the effect is still that of a turning away from the
corolla. Galium huancavelicum also shows some ambiguity in the direction of ovary
hair curvature. It should be mentioned that mature fruits of various species are
sometimes less clearly in one category or the other than the ovaries from which they
were derived.
The number of actual or visible veins in the leaves is useful. "Visibly 3-nerved"
means that three separate nerves can be seen running entirely to the base of the leaf.
"Obscurely 3-nerved" means that the midrib is obvious, but the lateral nerves connect
ing directly to the stem rather than to the midrib can rarely be observed without a
microscope and clearing techniques. "Cryptically 3-nerved" is more extreme, with
clearing definitely required to see the lateral nerves. One-nerved leaves have all the
secondary vascular bundles arising from the midrib.
Statements as to corolla color must be considered with skepticism. They are based
on scarce collectors' data, plus assessments of color from dried specimens. The latter
can be deceiving.

Key to species
Leaves 4 at each node.
Mature inflorescence falsely dichotomous, with short pedicels and much longer branchlets; Coqu
Chile 1. G. ovalleanum
Mature inflorescence var
Tips of ovary hairs usual
Flowering branchlets det
tapering to a petiolate base, appearing 1-nerved 2. G. obovatum
Flowering branchlets more complex (i.e., indeterminate); leaves ovate or elliptic, usually na
suddenly at base, obviously 3-nerved 3. G. weberbaueri
Tips of ovary hairs turned upward (somewhat inconsistent in no. 4).
Flowering branchlets determinate, with 1-5 flowers.
Leaves 3-nerved; flowering branchlets mostly 5-flowered; Peru 4.
Leaves mostly 1-nerved.
Branchlets with 1 internode and 3 pedicellate flowers; leaves commonly 5-10
southern Chile and Argentina, O'Higgins and southward 5. G. fuegianum
Branchlets with 1 flower above a usually solitary bract; leaves less than 4 mm. long; Peru.
ή η nor,.s,i ην,,,m

Flowering branchlets more complex, indeterminate.

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 ALLERTONIA

Leaves 1-nerved, or at least not obviously 3-nerved.

Corollas mostly red, the filiform tips spreading; Peru and Ecuador 7
Corollas green, yellow, or white, the obtuse or acute tips inturned.
Lobes of corollas not widely spreading (i.e., semierect), shortly hispid exter
none, or a few subapical; Chile 8. G. diffusoramosum
Lobes of corollas rotately spreading, glabrous; glands subapical and dist
surface of leaves; Venezuela and Colombia 9. G. orizabense
Leaves more or less obviously 3-nerved.
Corollas definitely campanulate; Colombia 10. G. boyacanum
Corollas more or less rotate; widespread in mountains from central Peru to nor

Leaves more than 4 at each node.

Fruits with elongated mericarps; leaves mostly less than 6 mm. long; annuals.
Mericarps sausage-shaped, the hairs unevenly distributed between them 12. G. mu
Mericarps reniform, the hairs evenly distributed 13. G. parisiense
Fruits with round, more or less isodiametric mericarps; leaves most often longer than
Annual; corollas rotate; ovary and fruit hairs upturned; leaves 6-8 at each node. ..
Perennials.
Corollas rotate; leaves 5 or 6 at each node; ovary hairs often, but not always, downturned.
15. G. pseudolriflorum
Corollas campanulate; leaves 6-12 at each node; ovary hairs always upturned. 16. G. mexicanum

1. Galium ovalleanum Philippi in Anales Univ. Chile 85: 734. 1894. Type: CHILE:
Coquimbo: Dept. Ovalle, Tulahuén, Geisse (sgo holotype, photo seen; isotype
at bm).
Polygamous subshrub to 40 cm. high, the fertile branches springing from slender,
woody stems above a stout rootcrown; entire plant canescent (stems, leaves, corollas);
internodes 1.5-3 times as long as leaves; leaves in fours, 4-7 (-9) mm. long, lanceolate
to narrowly ovate, obtuse at apex, narrowed to a broad insertion, obviously 1-nerved
but obscurely 3-nerved, the margins revolute; glands none; inflorescence diffuse,
many-flowered, falsely dichotomous when mature, the branchlets widely divaricate;
pedicels mostly very short, nearly obsolete to as long as flowers or fruits; corollas
rotate, greenish yellow, glandular-dotted along veins above, externally hispid, the
lobes spreading, blunt or more or less elongated; fruit hairs moderately fine, dense, a
little longer than fruit body, uncinate, curved or straight at ends.
Distribution: Chile: Coquimbo, 1,120-2,600 m. (Figure 11), in rocky ground.
This species was treated at greater length as a member of sect. Lophogalium,
although some individuals have uncinate fruit hairs. For illustration and detailed
discussion of this puzzling taxon that straddles a morphological line previously
thought to be perfectly clear, see Dempster, 1980, p. 256.

2. Galium obovatum Kunth in H.B.K. Nova Gen. et Sp. 3: 336. /. 277. 1818. Type:
ECUADOR: "Locis temperatis Regni Quitensis, prope Chillo et Cachabamba
(sic)," at 2,500 m., Humboldt 2225 (p holotype, photo seen). Figure 2.
Galium lappaceum Ruiz & Pavon, Fl. Peru 1: 59. 1798. Type: PERU: "In circuitu Huanuci urbis ad
Puelles," Ruiz & Pavon (ma holotype, photo seen; isotype at f).
Galium rotundifolium L. var. uncinulatum Kuntze, Rev. Gen. PI. 3: 120. 1898. Type: BOLIVIA:
Cochabamba: Tunari Mountains, Kuntze (ny holotype; isotypes at f, us).
Decumbent or climbing perennial, herbaceous above ground; stems weak, slender,
sparsely set with long slender spreading hairs, or sometimes glabrous, the internodes
2-3.5 times as long as leaves; leaves in fours, thin, 1-nerved or sometimes obscurely
3-nerved, mostly 7-15 mm. long, broadly obovate, acuminate, often tipped with a long
slender hair, narrowed to a petiole, sparsely set above and below with long spreading
or antrorse hairs; glands subapical and often distributed; inflorescence on lateral
branches usually consisting of 1 internode topped by 1-4 reduced leaves and 3 flowers
on subequal pedicels; pedicels 2-7 times as long as flowers, becoming 6-19 mm. long in

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 2. Galium obovatum·,a, flowering and fruiting branch, χ 5/8; b, leaf, lower side, χ 6;
12; d, fruit, * 12. a from Knight 505, b & c from Steinbach 9450, d from Daniel 2645.

fruit; flowers perfect; corollas rotate, glabrous, (green or) reddish to dark
apices filiform, to as long as lobes; ovaries thickly set with very short
uncinate hairs; hairs becoming much longer in fruit.
Distribution: Andes from central Bolivia (Cochabamba) to northern
(Norte de Santander), (1,000-) 2,000-3,700 m. (Figure 3), creeping on for
climbing to 60 cm. (5 m.) high, in moist shady places.
Additional collections: BOLIVIA: Cochabamba: Frov. Chapare, Steinbach 9155 (bm, e, f, gh, k,
ny, s), 9450 (bm, F, G, gh, κ, LIL, mo, NY, s, uc, us), 9817 ( g); Choro, ca. 100 mi. NW. of Cochabamba, Brooke
6267 (bm). La Paz: Cordillera Real, Tate 236 (ny); Unduavi, Buchtien 671 (w),3004 (f, ny, us), Rusby 1841
(gh, ny); Prov. Larecaja, near Sorata, Mandón 331 (G, ρ), Rusby 1832 (bm, gh, κ, mo, ny, p, us, w, wis).
PERU: Cuzco: Herrera 2588 (f). Apurímac: Prov. Andahuaylas, N. of Chincheros, Stork & Horton
10772 (F, g, gh, mo, uc). Ayacucho: Ccarrapa, between Huanta and Río Apurímac, Killip & Smith 22511
(us). Huánuco: 16 mi. SE. of Huánuco, Macbride & Featherstone 2112 (f, g, s, us); Cani, Macbride 3386
(bm, f, g, s, us); Carpish, Antunez de Mayolo 402 (uc), Asplund 12897 (g, s); Huacachi, Macbride 4133 (f,
gh, us); Muña, Macbride 3916 (f, us).
ECUADOR: Loja: Cerro Villonaco, Knight 505 (wis). Azuay: Eastern Cordillera, Ν. of Sevilla de Oro,
Camp E-4652 (s), E-5191 (s). Cañar: Ν. rim of valley of Río de Cañar, G Her (Camp E-2838) (ny).
Chimborazo: Riobamba-Guayaquil, Iltis E-570c (uc, wis). Tungurahua: Near Patate, Asplund 7967 (κ,
s); Baños, Rorud (f). León: Headwaters of Rio Macuchi, above Pilaló, Steere 8084 (f, us). Pichincha:
Benoist (p), Asplund 6721 (s), Fagerlind & Wibom (s), Sodiro 871 (w), Sparre 13698 (s). Imbabura: Asplund

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 ALLERTONIA

VENEZUELA ζ

Equator

o=Galium obovatum

□ =G. ferrugineum
ssp. ferrugineum

■ =G. ferrugineum
ssp. jamesonii
A = G. weberbaueri

Β =G. boyacanum
H = G. huancavelicum
L1 G. orizabense

P= G. peruvianum

BOLI VI A

Figure 3. Map showing distribution of several species of Galium.

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

7146 (br, g, κ, lil, νυ, ρ, R, s), Drew & Wiggins 13 (f). Carchi: Asplund 16869 (f, s). Napo-Pa
8005a (F, νυ, us), 8006 (f).
COLOMBIA: Nariño: Near Túquerres, Karsten (w), Triana 1595 (p),3100 (col). Cauca: R
Core 1030 (νυ, us); Río Santo Domingo, Fosberg 21290 (uc, us). Valle del Cauca: Cual recasa
20693 (f). Tolima: Pennell 3133 (f, gh, mo, ny, us). Cundinamarca: Cualrecasas 6688 (f), Fo
(ny). Antioquia: Salto de Guadalupe, Daniel 2645 (f, us); near Boquerón, Hodge 6596 (gh
Santander: Alston 7261 (bm, s, us), Killip & Smith 20201 (f, ny, us).

Figure 4. Galium weberbaueri\ a, flowering and fruiting branch, x 5/8; b, flowering branch
leaf, upper side, x 6; d, flower, χ 12; e & f, fruits, showing variation in hair length, x 12. a & c fr
14519, b & d from Ferrevra 7196, e from Weberbauer 5800, f from Weberbauer 7516.

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 ALLERTONIA

The inflorescence is basically the same as that of Galium fuegianum

consistent in pattern, i. e. some cymules are reduced to one or two flowe
are slightly proliferated.

3. Galium weberbaueri Krause in Bot. Jahrb. 40: 349. 1908. Type: PERU: Near
railroad from Lima to Oroya, 2,370-3,000 m., Weberbauer 204 "& 205" (
holotype, photo seen; isotype at g). Figure 4.
Galium andicolum Krause in Bot. Jahrb. 40: 350. 1908. Type: PERU: Ancash: Below Hacienda
Cajabamba at 3,000-3,500 m., Weberbauer 3123 (b holotype, photo seen; isotype at usm
Galium límense Standi, in Pubi. Field Columbian Mus., Bot. Ser. 4: 298. 1929. Type: PERU: Lima:
Canta, Penneil 14351 (f holotype; isotypes at g, ny, s, us, usm).
Climbing, trailing or pendent, polygamous, perennial herb, or suíírutescent below;
stems to 50 cm. long, scabrous, or hispid with retrorse hairs; internodes 2-4 times as
long as leaves; leaves in fours, 6-12 (-18) mm. long, elliptic or ovate, acute or apiculate,
usually narrowed rather abruptly to a broad base, obviously 3-nerved, the nerves
prominent beneath; leaves set above with long or short antrorse hairs, (glabrous or) set
beneath with retrorse hairs, those on ribs aculeate; glands none or few, subapical and
scattered; inflorescence diffuse, on lateral branches, the pedicels long (1.5-5 times as
long as flowers), slender to filiform, well exserted in fruit, variously curved but not
regularly nodding at apex; corollas usually greenish yellow but sometimes dull
maroon, glabrous or rarely with a few hairs externally, much larger than ovaries with
hairs, the lobes ovate, the apices often much prolonged but shorter than lobes,
occasionally filiform; ovaries set with short downturned hairs; fruits with stoutish
hairs, usually much shorter than fruit body.
Distribution: Peruvian Andes from Moquegua to Huanuco and Ancash, 2,300
3,500 m. (Figure 3), on rocky slopes with shrubs or shade.
Additional collections: PERU: Moquegua: Carumas, Weberbauer 7294 (bm, f, g, s, us). Ayacucho:
Prov. Lucanas, Allpaja, Ferreyra 7196 (uc, us); Prov. Huanta, mountains NE. of Huanta, Weberbauer 7516
(f, s, us). Huancavelica: Prov. Castrovirreina, above Huaytará, Weberbauer 5420 (f, gh, us); Prov.
Tayacaja, 3 km. N. of Salcabamba Village, Stork & Horton 10321 (F, uc). Lima: Matucana, Macbride &
Featherstone 87 (F, gh, us), 556 (F, s), Martinet 51 (f), Asplund 11082 (G, R, s); Surco, Ferreyra 9640 (uc);
Prov. Canta, below Canta, Ferreyra 9016 (uc); San Buenaventura, Pennell 14519 (F, bm, GH, NY, S); San
Pedro de Casta, Duncan, Dempster, et al. 2731 (mo, uc, usm), 2732 (mo, uc, usm). Ancash: Ocros,
Weberbauer 5800 (f, gh, us); Prov. Bolognesi, Rumipuquio, Cerrate et al. 6519 (uc). Huánuco: Llata,
Macbride & Featherstone 2234 (BM, F, us).
Specimens of this species are widely attributed to Galium ferrugineum, and the two
species must be thought of as closely related. The polygamous sexual system, unusual
in this group, the conspicuously long pedicels, the habit, and the inflorescence pattern
are important characters that both species share. However, the sharp difference in
orientation of the ovary hairs, together with their geographic separation, distinguish
the two species well; other characters, such as flower color, leaf venation, and length of
corolla apices, are less decisive.

4. Galium huancavelicum Dempster, sp. nov. Figure 5.

Herba perennis caespitosa vel semiprostrata, caulibus ca. 20 cm. lo
quam foliis 2-3.5-plo longioribus; caulibus foliisque pilos dispe
gerentibus; foliis in quoque verticillo quatuor lanceolatis 3-nerviis 5
insertionem latam angustatis, apice obtusis, pilo apicali si praesenti
glandulis subapicalibus; inflorescentiis 3-5-floris, ramulis brevib
internodio longo typice constantibus, foliorum verticillo apice in
fructum unicum necnon ramulos duos laterales bifloros subtendent
lobis obtusis vel apice haud productis ascendenti-patentibus, extu
validis ornatis, intus glandulis multis clavatis praeditis; ovarii

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 5. Galium huancavelicum; a, flowering and fruiting branch, χ 5/8; b, leaf,

flower, χ 12; d, fruit, * 12. All from Tovar 257 (type).

paulo brevioribus pro parte maxima sursum uncinatis sed inconsta

sub fructu et manifestis 5-10 mm. longis, statim infra fructum nutan
fructibus pilis gracilibus quam corpore paulo brevioribus ornatis.
Type: PERU: Huancavelica: Dept. Huancavelica, Motcca, 4 km. S
Tovar 257 (us holotype).
Distribution: Peru: Huancavelica, 3,400-3,450 m. (Figure 3), in rocky soil,
among shrubs. The holotype is the only available specimen; it includes immature,
flowering, and fruiting material.
Except for Galium pseudotriflorum, this is the only species of the group in which
the ovary hairs are not clearly either downwardly or upwardly uncinate. Although in
G. huancavelicum most ovary hairs curve upward at the tips, some unquestionably do
not. The exceptional length of the hairs might in theory account for some disorienta
tion, but I am inclined to look to the regionally sympatric G. weberbaueri as a possible
source of downcurved hairs in G. huancavelicum. The two species have 3-nerved
leaves and very long pedicels in common, although the indeterminate flowering
branchlets and the elongated corolla tips of G. weberbaueri, as well as the clearly
downturned ovary hairs, distinguish it from G. huancavelicum.

5. Galium fuegianum Hook. f. FI. Antarct. 302 (bis). 1846. Type: CHILE: Port Famine

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 ALLERTONIA

Figure 6. Galium fuegianum·,a, fruiting branch, χ 5/8; b, leaf, lower side,x 6; c, flower, * 12; d, fruit,x
12. a from Dúsén 5622, b & c from Godley 1079, d from Buchtien 48.

(i.e., just S. of Point Santa Ana at mouth of San Juan River), Strait of Magellan,
King (κ holotype). Figure 6.
Loosely tufted, decumbent peren
sprawling, herbaceous above groun
lower nodes, glabrous or with scatte
times as long as leaves; leaves in four
5-10 (-20) mm. long, elliptic, apicu
nearly glabrous except midrib, the u
downwardly rolled margins ciliate;
lets consisting usually of 1 long in
flowers on subequal pedicels 3-30 m
perfect; corollas glabrous or with fe
hairs, greenish to pale yellow, ting
widely spreading; ovaries and fruits d
Distribution: Andes of Chile an
Patagonia at 60-500 (-2,000?) m.; C

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DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 7. Map showing distribution of Galium fuegianum.

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 ALLERTONIA

central Chile from Malleco to O'Higgins at 1,000-1,500 m. (Figure 7).

somewhat disjunct distribution is probably unreal. Moist places in Notho
or in scrub.
Additional collections: CHILE: Magallanes: Navarino Island, Godley 1079 (κ, sgo), Skottsberg
285 (s); Basket Island, Spegazzini 18632 (lp); Isla Grande, Moore 2468 (c, κ), Moore & Goodall 103 (c); near
Punta Arenas, Düsen (MO, s), Lechler 1206 (G, goet, κ, m, ρ, s, w), Savatier 70 (ρ), Spegazzini 18629 (lp);
Laredo Bay, Lee (us); Seno de Skyring, Skottsberg (sgo); near Puerto Natales, Behn 21414 (conc), Eyerdam
et al. 26324 (gh, uc); Última Esperanza, Magens 11469 (conc). Malleco: Volcán Lonquimay, Sparre &
Constance 17584 (conc, uc); Termas de Tolhuaca, Gunckel 16369 (us), 16370 (us). Ñuble: Chillán, Jaffuel
3865 (gh), Philippi? (sgo). Rancagua: Mont. La Leona, Bertero 287 (ρ).
ARGENTINA: Tierra del Fuego: Harberton, Goodall 1113 (us), Moore 1329 (κ); Vallee d'Olivoia,
Atboff (lp); Estancia Punta Segunda, Moore 1945 (c, κ). Santa Cruz: Lago Argentino region, Cantino 136
(gh), 176 (gh), Dempster 4472 (uc), Dusén 5622 (s), Hauthal 498 (f, lp), James 145 (bm), 814 (bm), 1304
(bm); Estancia Cerro Fitzroy, Sleumer 1325 (s, us). Chubut: Lago Fontana, Castellanos 6211 (f); Los
Alerces National Park, Pérez Moreau 49631 (ba). Río Negro: Vicinity of San Carlos de Bariloche, Agrasar
MC 2985 (mo), Buchtien 48 (gh, m, s, us), 1330 (gh, s, us, w), Fabris 2240 (lp), Fabris & Solbrig 844 (lp);
Sierra Mamuil Malal, Comber 1100 (κ).
Throughout most of the range of Galium fuegianum, its lower leaf surfaces are
essentially glabrous. However, half of the specimens from Malleco and all specimens
from farther north have leaves with long hairs on both surfaces. They also vary from
the type in other, inconsistent, ways, e.g. leaves broader and very large (Bertero 287,
Philippi (?)), or leaves tapering at apex (Gunckel 16369 and 16370). If more of the
northern material were available, and if chromosome counts were made, these collec
tions might prove to represent a separable subspecies. Indeed, the Philippi specimen
bears an unpublished name.
Apparently in this species, as in G. obovatum, the inflorescence is truly determi
nate. Only in very rare instances does a bract occur on a "pedicel", indicating the
presence of a node.

6. Galium peruvianum Dempster & Ehrendorfer, sp. nov. Figure 8.

Herba perennis parvula in petrarum fissuris verosimiliter caespitosa; caulibus
foliisque pilis patulis canescentibus, internodiis quam foliis 1.5-3-plo longioribus;
foliis in quoque verticillo quatuor ovatis haud 7 mm. longis 1-nerviis (abdite 3-nerviis),
margine reflexis, apice subobtusis, basi in petiolum brevem angustatis; glandulis sub
apicalibus; floribus hermaphroditis, pro parte maxima axillaribus, ramulis plerumque
cum bractea unica infra florem enatis, pedicello vero igitur brevissimo; corolla glabra,
lobis ovatis haud patentibus acutis sed appendice apicali deficienti; ovarii pilis apice
sursum anfractis; fructus pilis corpore subaequalibus.
Type: PERU: Junín: La Oroya, eastern limestone cliffs, ca. 12,000 ft. (3,600 m.),
Macbride & Feather stone 937 (f holotype; isotype at g).
Distribution: Peru: Junin. Besides the type, two other collections are known, one
a mere fragment from the road between Jumn and the mountain (la Montaña),
Raimondi 11891 (f); the other from ca. 18 km. SSE. of Tarma, Iltis 134 (mo, uc, us,
ven, wis) (Figure 3).
Although Galium peruvianum is essentially a 4-leaved species (i.e., with 2 leaves
and 2 leaflike appendages per node), there is a strong tendency in the type material for
one of the false leaves to split, giving rise, occasionally, to a "5-leaved" condition. In the
absence of more material, it cannot be indicated whether this is an aberration or a
specific character. Note that in Figure 8, which is a faithful portrayal, one node has
five leaves, and each of the other nodes has one bifurcated false leaf. Note also that
each branchlet has only one flower, and that all but one have only a single bract. The
exception has an extra internode and an extra bract. I have also seen a solitary instance

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 8. Galium peruvianum\ a, branch, χ 5/8; b, portion of branch, showing flowers,

branching pattern, « 3; c, leaf, lower side, x 6; d, flower, χ 12; e, immature fruit, with bract and o
carpel, * 12. All from Macbride & Featherstone 937.

where three fruits terminate a branchlet, suggesting further variability in the

cence.

7. Galium ferrugineum Krause in Bot. Jahrb. 40: 349. 1908.

Slender, lax, climbing, trailing or pendulous, polygamous perennial,
50 cm. or longer, the internodes 3-5 times as long as leaves; leaves in f
long, apparently 1-nerved but often obscurely 3-nerved, elliptic t
apiculate, narrowed to a short petiole; glands none or few, subapic
inflorescence diffuse on few-flowered axillary branches; pedicels fi
ously exserted, becoming 10-15 mm. long in fruit and slightly n
corollas reddish, the widely spreading lobes narrowly ovate, the filifor
as lobes to quite short; ovaries much smaller than corollas, covered
upturned hairs; fruits covered with stout uncinate hairs much shorter
This species has often been confused with the more widespread Galiu
eri, to which it has a remarkable overall resemblance. It differs, howev
important respects: (1) the ovary hairs are upturned instead of dow
leaves are mostly 1-nerved instead of obviously 3-nerved; and (3)
apparently red instead of usually green or yellow. In addition, the plan
more slender.

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 ALLERTONIA

Figure 9, a-f. Galium ferrugineum subsp. ferrugineum·, a, flowering and fruiting

lower side, χ 6; c, leaf, upper side, χ 6; d, leaf apex, lower side, glands omitted, * 12;
unnaturally flattened, x 12; f, fruit, χ 12. g & h. Galium ferrugineum subsp. jamesoniv, g
with leaves, * 3; h, leaf apex, lower side, glands omitted, « 12. a-f from Lopez M. 104
from Sparre 17732.

Key to subspecies
Stems scabrous; leaves sparsely hairy, principally on upper surface; northern Peru. 7a. subsp.ferrugine
Stems mostly hirsute with long hairs; leaves abundantly hairy on both surfaces; Ecuador.
7b. subsp. jamesonii

7a. Galium ferrugineum subsp. ferrugineum. Figure 9, a-f.

Galium ferrugineum Krause in Bot. Jahrb. 40: 349. 1908. Type: PERU: Cajamarca: Above San
2,400-2,700 m., Weberbauer 3807 (b holotype, photo seen; fragments at f, usm).

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1981 DEMPSTER: GALIUM IN SOUTH AMERICA. II. 407

Stems finely scabrous with aculeate hairs; leaves with few scattered an
above, essentially glabrous beneath, the reflexed margins and sometimes
set with retrorse, mostly aculeate hairs; corollas glabrous, the filiform api
Distribution: Northern Peru in Cajamarca and La Libertad, 1,6
(Figure 3).
Additional collections: PERU: Cajamarca: Chorillos, Raimondi 6870 (f). La Libertad: Prov.
Otuzco, road to Paranday (Sinsicap), Lopez M. 1045 (lil, us).

7b. Galium ferrugineum subsp. jamesonii Dempster, subsp. nov. Figure 9, g & h.
Herba perennis, ramulis pilis longis plerumque hirsutis sed aliis inferioribus pilis
aculeatis aliquando instructis; foliis utrinque pilis longis ornatis; corolla glabra vei pilis
longis paucis extus induta, apicibus saepe brevibus.
Type: ECUADOR: Pichincha: Ravines towards the base of Pichincha, June,
1859, Jameson s. η. (κ holotype; probable isotypes at bm, c, e, g, gh, goet, k, p).
Distribution: Ecuador, 3,200 m. (Figure 3), in ravines.
With one exception (Sparre 17732, s), all specimens at hand were collected by

Figure 10. Galium diffusoramosum\a, flowering branch, x 5/8; b, leaf in normal position, upper side, *
6; c, flower, χ 12; d, corolla from same, spread open, χ 12; e, fruit, « 12. a from Behn 8279, b-d from Worth &
Morrison 16251, e from Johnston 5621.

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 ALLERTONIA

Figure 11. Map showing distribution of Galium diffusoramosum and G. ovallean

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Jameson. Except for the holotype, most of these bear only the
Andibus Quitensibus", 1859 (bm, c, e, g, gh, goet, k, p). It is probable,
that all of these, including a specimen in the Hooker Herbarium (
number 148, are isotypes. A specimen at w, from "environs of Alausi,
Jameson at a later date (1864).
This taxon has been treated as a subspecies rather than a species for
abundant material. I have seen only three collections ofsubsp.ferrugin
the type, and the number of collections of subsp.jamesonii may also b
(see above). The conspicuously different indûment is probably stable. A
of subsp. ferrugineum has long corolla apices, whereas some of that o
nii has short (although not blunt) corolla apices.

8. Galium diffusoramosum Dempster & Ehrendorfer, nom. nov. Figure 10.

Galium diffusum Clos in Gay, Fl. Chil. 3: 180. 1848; non Galium diffusum Gilibert(1781)necDon(1821)
nec Schräder (1821 or 1822). Type: CHILE: Coquimbo: On road to Arqueros, Gay 233 (p holotype;
probable isotypes at f, g, gh, k).

Sprawling, prostrate or clambering, polygamous perennial, the long, slender, often

purple stems arising from a woody base; internodes 2.5-8 times as long as leaves,
scabrous with short, stout, aculeate hairs; leaves in fours, clearly 1-nerved (or some
times obscurely 3-nerved), 5-12 mm. long, ovate or elliptic, more or less obtuse at
apex, narrowed above the swollen base, commonly reflexed at maturity and breaking
above the persistent base, more or less scabrous on both surfaces, and particularly on
the margins, with very short, stout, often aculeate hairs; glands none, or a few
subapical; inflorescence diffuse, on lax, few-flowered, lateral and terminal branchlets;
pedicels 1-3 (-5) times as long as flowers; corollas sordid white or yellowish, hispid
externally, elongated in bud, usually much longer than ovary, the lobes ovate, blunt at
apex, ascending or semierect, the tips turned inward; fruit hairs many and very fine,
shorter than fruit body, the tips upwardly uncinate.
Distribution: Chile: Central Coquimbo northward to southern Antofagasta,
100-700 m. (Figure 11), at base of rocks and shrubs and on talus slopes, in hills and
arroyos back of the coast.
Additional collections: CHILE: Coquimbo: Region of Ovalle, Jiles 2711 (conc), Barros 3679 (f),
Marticorena et al. 1452 (conc), Behn 8279 (conc); Fray Jorge, Ricardi 12797 (conc, lil); La Serena, Cerro
Grande, Barros 3602 (F); road from La Pelicana to Arqueros, Marticorena & Matthei 295 (conc); Dept. La
Serena, Los Choros, jiles 3959 (conc). Atacama: Dept. Huasco, ca. 4 km. S. of Huasco, Worth & Morrison
16251 (g, gh, mo, uc). Antofagasta: Taltal, Darapsky 2036a (sgo); Cerro Perales near Taltal, Johnston
5621 (gh).

9. Galium orizabense Hemsl. Diagn. PI. Nov. Mexic. 54. 1880. Type: MEXICO:
Veracruz: Orizaba, Botteri 838 (κ holotype; isotype at bm). Figure 12.
Galium canescens f. alstonii Steyermark in Acta Bot. Venez. 8:253. 1973. Type: VENEZUELA: Mérida:
Between Timotes and Chachopo, Alston 6533 (ny holotype, not seen; isotype at bm).
Trailing perennial herb; stems with few to many short, very fine, mostly down
curved hairs; internodes 1-5 times as long as leaves; leaves in fours, mostly 12-17 mm.
long, narrowly elliptic or oblanceolate, round-apiculate or obtuse at apex, narrowed to
a slender petiole, lightly pubescent with fine antrorse hairs or nearly glabrous, the
largest visibly, but not obviously, 3-nerved; glands subapical and thickly distributed on
lower leaf surface; inflorescence on mostly few-flowered indeterminate lateral
branches, the pedicels and branchlets widely spreading, subequal, 3-5 times as long as
fruit; flowers perfect, very small (2 mm. across); corollas glabrous, yellowish white,
rotate, the blunt tips inturned; ovaries and fruits with upturned uncinate hairs; fruits
very small, the body 2 mm. broad or less.

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 ALLERTONIA

Figure 12. Galium orizabense\ a, branch in young fruit, χ 5/8; b, leaf, lower side,
glands, χ 6; c, flower, » 12; d, fruit, * 12. a from Oberwinkler 12743b, b-d from P

Distribution: Mountains of northwestern Venezuela and adjace

from Distrito Federal to Santander, 1,000-3,000 m. (Figure 3), o
Mexico and Central America.
Additional collections: VENEZUELA: Distrito Federal: Mountains above Caracas, Jahn (us);
Los Venados de Galipán, Pittier 10451 (f, g, gh, ny, ven); cuenca del Rio Macarao, Montes 87828 (ven); La
Guayra-Caracas, Kuntze 1318 (ny). Barinas: Barinitas-St. Domingo, Oberwinkler 12743b (m).
COLOMBIA: Santander: Near Tona, Killip & Smith 19509 (gh, ny, us).
I have observed no significant difference between Galium orizabense of Mexico
and Central America (Dempster, 1978) and the plants of Colombia and Venezuela
described in 1973 by Steyermark as G. canescens f. alstonii. These South American
plants, in fact, form a segment in a geographic arc that begins at southern Tamaulipas,
Mexico, passes through northern Colombia and Venezuela, the Caribbean Sea (His
paniola), and into Florida. The plants that compose this arc should probably be
thought of as conspecific, although a subspecific break can be recognized between
South American and Mexican plants on the one hand and the Caribbean and Floridán
plants on the other.
Some adjustments of nomenclature are required in the recognition of these rela

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

tionships. The earliest name is Galium orizabense Hemsl., typified by a

Veracruz, Mexico. The Venezuelan and Colombian plants were erroneousl
to G. canescens as a form by Steyermark. Plants of Hispaniola of this relati
all labeled G. pilosum Ait. var. puncticulosum (Michx.) Torrey & Gray.
thus of a relationship to G. pilosum is valid, but the Floridán and Caribbean
much more closely related to G. orizabense than they are to G. pilosum. Th
ships here stated categorically are discussed in more detail below.
That Galium orizabense is more closely related to G. pilosum than to G. c
is supported principally by the strikingly abundant glandular cells distribu
lower surface of the leaves of both G. orizabense and G. pilosum, and b
approach to the 1-nerved rather than the 3-nerved condition of the lea
orizabense differs from G. pilosum pincipally in habit, position of inflores
shape, and indûment. In contrast with the sprawling habit, indeterminate
lateral inflorescences of G. orizabense, plants of G. pilosum are erect, with
terminal, merely bracteate inflorescences. Their leaves are broadly ellip
rather than usually narrowly elliptic or oblanceolate, and both the stems an
have much more indûment than those of G. orizabense.
Although the differences between Galium orizabense of Mexico and South Amer
ica and the plants of Hispaniola and Florida are slight, they are, together with the
geographic distance, sufficient for infraspecific differentiation. The stems of the Carib
bean and Floridán plants are glabrous, the leaves a little wider on the average, acute or
obtuse at apex, the fruits a little larger, and the inflorescences concentrated near the
terminus. The two lateral nerves on the larger leaves of the Latin American plants are
more visible than on those of Hispaniola and Florida or on those of G. pilosum·, they
can usually be seen under magnification without clearing and are probably always
present. In the Caribbean material and in G. pilosum, lateral nerves are more often
impossible to see and are probably often altogether lacking. All of these differences are
slight, but are in the direction of G. pilosum, of North America.
The Floridán plants that have been called Galium pilosum var. laevicaule by
Weatherby & Blake (in Rhodora 18: 194. 1916. Type: U. S. Α.: Florida: Near
Jacksonville, Curtiss 6420; GH holotype; isotype at uc) I propose to call Galium
orizabense Hemsl. subsp. laevicaule (Weatherby & Blake) Dempster, comb, et stat.
nov.

In regard to the Caribbean plants labeled Galium pilosum Ait. var

(Michx.) Torrey & Gray, I have seen the microfiche of the type of G
from "Basse Carolina", as well as "No. 2" of the same name, mou
(microfiche #20). Both specimens have the broad leaves and stouter
tic of G. pilosum, rather than the narrow leaves and slender stems u
Caribbean specimens. The epithet puncticulosum seems to have be
color. One collection from Haiti (Ekman 4632) indicates "fl. redd

10. Galium boyacanum Dempster, sp. nov. Figure 13.

Herba perennis serpens poly gama; caulibus pilis multis longis ri
angulis latis scabridis, internodiis quam foliis 2.5-5-plo longior
verticillo quatuor lanceolatis vei ellipticis 3-nerviis pro parte
longis, maturitate reflexis, apice gradatim vel abrupte acutis,
angustatis; petiolis nodisque dense pilosis, foliorum paginis inferni
pilis multis grossis patulis vestitis, superficiebus pilis dispersis

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 ALLERTONIA

Figure 13. Galium boyacanum\ a, flowering and fruiting branch, χ 5/8;b, leaf, lower
χ 12; d, corolla spread open, χ 12; e, fruit, x 12. All from Cuatrecasas 1159 (type

instructis; glandulis paucis subapicalibus; inflorescentiis (lateralibu

nalibus proliferis tantum bracteatis, pedicellis ramulisque late d
glabra campanulata ad medium fissa et late patenti, apicibus obtus
acutis; ovarii pilis sursum curvatis; fructus pilis crassis sursum curvat
multo brevioribus.
Type: COLOMBIA: Boyacá: Between Soatá and Cocuy, Valle de la Uvita, El
Hatico, Cuatrecasas 1159 (us holotype; isotype at f).
Distribution: Colombia: Boyacá, 2,900 m. (Figure 3), in thickets and forest. The
type is the only known collection.
The range of variation, particularly in quantity of indûment, size and shape of
leaves, and length of fruit hairs, cannot be known from one collection. Other charac

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 14. Galium canescens\ a, flowering and fruiting branch, showing congested habit, x
flowering and fruiting branch, showing lax habit, χ 5/8; c, leaf, lower side, showing few hairs and lar
χ 6; d, leaf, lower side, showing many hairs and small size, * 6; e, branchlet, showing staminate flo
young fruits, χ 6; f, perfect flower,x 12; g, fruit,x 12. a from Asplund 6729, b írom Asplund 17065, c
Baruch 102, d from Fassen 25501, e from Steyermark 61922, g from Hutchison 6968.

ters, such as the campanulate corollas, the inflorescence type, the 3-nerved leave
the very coarse indûment, are probably constant.

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 ALLERTONIA

11. Galium canescens Kunth in H.B.K. Nova Gen. et Sp. 3: 336. 18

DOR: Near the city of Quito at 2,700 m., Humboldt (b holoty
isotype at p). Figure 14.
Galium trianae Wernham in J. Bot.
commonly called G. trianae, and so fil
has glabrous fruits and is related to t
fruits of G. trianae are hispid with un
undoubtedly the holotype and that is
irifidum. Although this specimen has
its taxonomic segregation from G. ca
group of plants.
Trailing or scandent perenni
intern odes commonly 3-5 times
set with long coarse hairs (canes
beneath; leaves in fours, mostly
ovate or elliptic, plane or rolle
broad or frequently petiolar b
abundantly distributed, or some
lateral branches; branches inde
equal or unequal proliferating
from almost none to 6 (-11) mm
or sometimes with a few long
reddish externally, the lobes mo
fruits set with upwardly uncina
Distribution: Peru (Prov. Lim
Ecuador and Colombia to Monagas in northeastern Venezuela, 1,300-4,400 m.
(Figure 15), growing in tangled masses, sprawling, pendulous, or climbing among
shrubs, in open grassland, thickets, damp cliffs, or open forest.
Additional collections: PERU: Lima: Callao, Gaudichaud 99 (g), 100 (G). Ancash: Between
Tallenga and Pachapaque, Cerrate & Cevasco 704 (uc). Cajamarca: Between Celendi'n and Cajamarca,
Antiinez de Mayolo 360 (uc). Amazonas: Prov. Chachapoyas, above Leimebamba, Ferreyra 15493 (uc),
Hutchison & Wright 6968 (F, NY, uc).
ECUADOR: Cañar: NE. of Azogues, Prieto (Camp E-2465)(s). Chimborazo: Fagerlind & Wibom 786
(s), 877 (s). Pichincha: Asplund 6729 (s), Barclay & Juajibioy 7964 (ny), Holmgren & Heilborn 183 (f, s),
Sparre 17431 (s), 17780 (s), Steere & Camp 8321 (F, s), Jameson 196 (w), Sodiro (w). Imbabura: So lis 14222
(f), Sparre 13568 (s). Carchi: Asplund 17065 (s), Steere 8091 (f, ny).
COLOMBIA: Nariño: Near Túquerres, André 3192 (F, κ, ny), Karsten (w), Triana 3100 (ρ). Cauca:
Río Blanco, Core 913 (ny), east of Tacueyó, Fosberg 21290 (ρ). Valle del Cauca: Páramo de Sumapaz,
Cleef 7681 (u); Loma de Barragán, Cuatrecasas 20727 (f), 20857 (F). Caldas: Páramo del Quindío, Pennell
& Hazen 10051 (F, gh, ny, us). Cundinamarca: Barkley et al. 17C770 (us), Camilo 2a (f), Cleef5423 (υ),
Cuatrecasas 367 (f), 5122 (F, col), 5606 (col), Duque-Jaramillo 2767 (col), Ewan 15587 (p), Fosberg 19703
(ny), Goudot (κ), Haught 5677 (F, ny, s), 6501 (uc, ven), Holton 412 (gh, κ, ny), 413 (κ), 415 (F, ny), Kme
4514 (c), Pennell 1838 (f, GH, ny), Schiefer 313 (GH), 530 (GH, col), Schuttes 18569 (τ), S. G. Smith et al. 1044
(uc), Triana 3100 (col). Antioquia: Between Río Negro and St. Helena, Garcia-Barriga 11101 (col).
Boyacá: Municipio de Ráquira, Ranghel Galindo 88 (col); Sierra Nevada del Cocuy, Grubb et al. 72 (κ, us),
757 (κ, us), Barclay & Juajibioy 7299 (ny), Cuatrecasas 1591 (f), 1622 (f, us), 1631 (col, f, us). Santander:
Páramo de Romeral, Killìp & Smith 18549 (F, gh, ny, us), 18556 (F, gh, ny, us); near La Baja, Killip & Smith
18148 (bm, f, ny, us); Zapatoca, Fassett 25501 (p, uc, us). Norte de Santander: Near Pamplona, Fassen
25941 (ρ, uc, us), Killip & Smith 19764 (f, gh, ny, s, us); Pica-Pica Valley, Killip & Smith 20201 (f).
Magdalena: Sierra de Perijá, Cuatrecasas & Romero Castañeda 25129 (us, ven); Sierra Nevada de Santa
Marta, Cuatrecasas & Romero Castañeda 24449 (us), 24499 (us), Simons (bm), Weston 10303A (uc),
10303B (uc), 10344 (uc), 10370 (uc).
VENEZUELA: Tachira: Near international border, Terän 87486 (ven), Steyermark & Dunsterville
73083 (ny, ven). Mérida: Breteler 3357 (ny, us, ven), Fend 1er 537 (ΐ, G, gh, κ, μο,νυ, ρ), Gehriger 200 (f, g,
ny, us, ven), Jahn 802 (f, gh, us, ven), 1060 (f, us), López-Palacios 83901 (ven), Oberwinkler 12381 (m,
ven), 13016 (m, ven), Ruiz-Teran 6836 (ven), 7826 (ven), 9452 (ven), Schulz & Rodríguez 293 (ven),
Vareschi 2144 (m), 7330 (ven). Trujillo: Aristeguieta 3482 (ny, ven), Ruiz-Teran 7582 (ven). Distrito
Federal: Baruch 102 (ny, ven), Juzepczuk 299 (f), Kuntze 1647 (ny), Manara (ven), Pittier 8354 (us).

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DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 15. Map showing distribution of Galium canescens.

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 ALLERTONIA

Miranda: Pico de Naiguatá, Pittier 6267 (us). Monagas: Cerro de la Cueva de Doña
61922 (F, NY, ven); Cerro Negro, Steyermark 62115 (f, ny, ven).
Galium canescens is a variable and complex species occupying a l
mountains of northwestern South America. It varies principally i
ment and size of parts, although the shape of the leaves also va
Nevertheless, despite its variability, the usually more or less canescen
or less ovate, always (at least obscurely) 3-nerved leaves, mostly
flowers, and fruits on indeterminate branchlets is, as thus broad
distinct from other species in South America. The often glabro
surprising contrast to the usually hairy herbage.
Galium uncinulatum DC. (Prodr. 4: 600. 1830), of Mexico and Ce
closely related to G. canescens, and if both were found in South Amer
difficult, if not impossible, to separate them. As far as the available e
however, they are allopatric, the nearest approach being about
uncinulatum at Chiriqui, Panama, and G. canescens in southeaste
quia, Colombia). The differences between these two large and poly
tions are subtle and perhaps scarcely adequate to justify their spe
Galium uncinulatum is, however, a usually much more lax plant, part
inflorescence, where pedicels and branchlets are longer and leaves
The habit of G. uncinulatum is in many instances lower, inclined
decumbent rather than climbing; the plants are more slender and deli
than most (but not all) plants of G. canescens·, the indumentum is les
most (but not all) plants of G. canescens; and the leaves tend to be lar
canescens is in general a more hairy species, the reverse is true of the
more often glabrous. There is much overlapping of characters betwee
to such a degree that, although many specimens of G. uncinulatum ar
from G. canescens, many more are doubtfully distinguishable exc
grounds. A thorough study of the chromosome numbers of these
probably shed much light on the problem. Numerical comparisons of
of the two species may be summarized as follows:
Galium canescens Galium uncinulatum
Longest stem 25-54.0-106 cm. 15-57.5-106 cm.
Longest leaf 5-9.Ä-16 mm. 5-/2.7-30 mm.
Longest pedicel 0.5-5./-11 mm. 1-6.2-12.5 mm.
Corollas hispid: 32% hispid: 66%
slightly hispid: 15% slightly hispid: 10%
glabrous: 53% glabrous: 24%
The graph (Figure 16) shows a comparison based

Figure 16. Graphic comparison of Galium canescens and G.

discussion of G. canescens.

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

four characters in such a way as to give them roughly equal value (leaf le
length + longed stem + coro^a indûment on a basis of 0 to 10). Meas
inevitably rough, particularly of stem length, since collectors tend to fav
of convenient size. In measuring pedicels, care was taken to measure only
rather than apparent pedicels (i.e., lateral branchlets of the cymules)
crudity of the data, however, the picture that emerges shows a mark
between the two populations, with individuals of G. canes cens running f
with peaks from 20 to 34, whereas those of G. uncinulatum run from 26
peak at 45 to 48.

12. Galium murale (L.) Allioni, Fl. Pedem. 1: 8. pl. 77, fig. I. 1785.
Sherardia muralis L. Sp. PI. 103. 1753. "Habitat in Italia," type not seen.
Diminutive annual 1.2-7 cm. high, simple or branched at base, erect or
stems glabrous, the internodes 1.5-3 times as long as leaves; leaves com
each node, 2-5 mm. long, obovate or oblanceolate, tapered gradually to ba
abruptly to apex, nearly glabrous on surfaces, the margins with antr
apex tipped with a long slender hair; glandular cells none; flowers perfect
usually 1 or 2 in axils; corolla cleft nearly to base, green, becoming yellow
glabrous, the lobes ovate, ascending, obtuse, less than half as long as
ovaries; fruiting pedicels stout, recurved, somewhat shorter than fru
blackish, narrow, curved (sausage-shaped) mericarps, spreading apar
variously set with stout uncinate hairs, most commonly one mericarp hai
to apex, the other at apex only.
Distribution: Adventive in Chile: Concepción, 1949, Curicó, 1937,
1951, Santiago, 1879, Valparaiso, 1927, Coquimbo, 1952. Locally abund
of the mossy ground cover in damp places, and as an undergrowth on gra
600-650 m. Dates refer to the earliest known collection from each Province.
Collections seen: CHILE: Concepción: San Pedro, Ricardi9015(conc). Curicó: Rauco, Barros3653
(f); Cerro Condell, Barros 13967(cONC). Colchagua: Santa Cruz, Barros 13968 (conc); La Rufina, Ricardi
10011 (conc). Santiago: Cerro de Bravo, Philippi 940a (sgo); Apoquindo, Philippi 940 (sgo); San Cristóbal,
Philippi 940b (sgo), Hastings 377 (ny, uc, us); Santiago, Philippi (κ, uc, us); Maipú, Schlegel 1696 (sgo);
Peñaflor, Montero 43 (gh). Valparaíso: Marga-Marga, Jaffuel 1425 (gh), Limache, Looser 7436 (gh).
Coquimbo: Illapel-Salamanca, Barros 14008 (conc); Salamanca-Coquimbo, Barros 13969 (conc).

13. Galium parisiense L. Sp. Pl. 108. 1753. European, type not seen.
Slender annual, the erect stems 15-25 cm. high, simple or branched from base;
stems scabrous, the internodes 2.5-4 times as long as leaves; leaves in whorls of six, 4-6
mm. long, linear-oblanceolate, narrowed to a slender apical hair; glandular cells none;
inflorescence diffuse; flowers hermaphrodite, the peduncles and pedicels filiform;
corollas whitish or purplish, the lobes erect, rounded, more or less hispid externally;
ovaries somewhat elongated, the short hairs upwardly uncinate; fruits very small, the
mericarps at maturity very narrowly reniform and attached only at base; 2η = 44, 66
(Europe).
Distribution; Adventive in Chile but apparently rare there. Introduced from
Europe.
Collection seen: CHILE: Santiago: Cerro San Cristóbal, Gersh 80, in 1959 (wis).

14. Galium aparine L. Sp. Pl. 108. 1753. European; for discussion of possible typifica
tion see Moore (1975, pp. 879-881). The complex nomenclature and synonymy
that have grown up around this species or species complex in other continents are

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 ALLERTONIA

here omitted. The following names refer to South American plan

complete synonymy see Moore (1975).
Galium chítense Hook. f. FI. Antarct. 302 (bis). 1846. Type: CHILE: (probably) Aisén
lago, Darwin (κ).
Galium chonosense Clos in Gay, Fl. Chil. 3: 189. 1848. Based on G. chítense Hook.
chonosense (sic) Clos is not listed in Index Kewensis (Jackson, 1893-1895), although i
published; non G. chonoense Hook. f. (1846). N. B.: Index Kewensis, Reiche (1896-1
& Cowan (1976) all credit Clos with authorship of the Galium species published by G
his introduction to Vol. 1 (1845) Gay acknowledges the help of various botanists but
Clos. In indicating Clos as the author, I am assuming that the accreditation is cor
Galium aparine var. microphyllum Clos in Gay, Fl. Chil. 3:190. 1848. Type: CHILE:
(P)·
Galium pseudoaparine Griseb. in Abh. Königl. Ges. Wiss. Göttingen 6: 125. 1854. Type: CHILE:
Magallanes: "Pr. Sandy Point in silvis," i. e. near Punta Arenas, Lechler 1207 (goet holotype;
ISOTYPES at BR, G, K, P).
Galium aparine var. pseudoaparine Speg. in Revista Agron. Univ. Nac. La Plata 3:526. 1897. (Not seen.)
Galium australe Reiche, Fl. Chile 3: 149. 1900. Based on G. chítense Hook. f.
Galium larecajense Wernham in J. Bot. 50: 244. 1912. Type: BOLIVIA: La Paz: Prov. Larecaja, vicinity
of Sorata, between Munaypata and Challasuyo River, Mandón 339 (bm holotype; isotype at ny).
Clambering or prostrate annual, 10-90 cm. long or longer, climbing high among
shrubs, or more or less densely sprawling on other plants, or erect and depauperate in
grasslands; stems brittle and very scabrous with retrorse aculeate hairs; internodes
2.5-4 times as long as leaves; leaves 6-8 at each node, 13-31 mm. long, the earliest often
spatolate and petiolate, the later linear-oblanceolate to oblanceolate, tapering to a
narrow base, tapering abruptly or gradually to a mucronate apex; glandular cells none;
inflorescences few-flowered on well-exserted leafy indeterminate lateral branches, 1 or
2 in a cymule, or often solitary in axils; pedicels often very long (3-45 mm.), straight or
reflexed at tip; corollas rotate, white or sometimes yellowish; ovary and fruit hairs
upturned, the fruit hairs usually shorter than fruit body; 2η = 20, 22, 42, 44, 63, 64, 66,
ca. 86, 88 (published counts from Europe and North America).
Distribution: From Tierra del Fuego northward in the Andes to southern Colom
bia; northern Colombia in the Santa Marta Range; eastern Argentina (Buenos Aires
and Santa Fe) and southern Uruguay; Juan Fernández Islands (Figure 17), on rocky
or grassy slopes or in clay soil, and in moist places in thickets, open forest, grassland, or
on stream banks.
Additional collections (representative): ARGENTINA: Tierra del Fuego: Stateri Island, Cast
nos 12967 (f, gb); Isla Grande, Alboff (cord, lp), Banks & Solander(BM, gh, mo, s, us), Castellanos 79
7911 (F), 7914 (f), 7915 (f), 45754 (f), 45758 (f), Goodall 480 (lp, ny, us), 3994 (gh), Gusinde 178pp
Holmberg & Calcagnini 3946 (f), Moore 1542 (c, κ), 1333 (κ), 1784 (κ), 2614 (κ), Rousson & Wille
Santa Cruz: Ager TA A 525 (us), Anliot 6147 (sgo), Donat 188( F, G, GH, κ, mo, ny, s, u, uc), Eyerdam
24092 (f, g, gh, mo, s), James 83 (bm), Hünicken (cord), O'Doneil 3998 (c, p), Prichard (bm), Ljungne
(gb), Scolnik 370 (lil), Tweedie 351 (κ, lp). Chubut: Coastal, Aurelius 26 (s), Eyerdam et al. 23814
Hosseus (cord), O'Doneil 3552 (lil), Riggs 80 (F, gh, mo, us); Andean, Beetle & Bignoli256 (mo), I
(s), 477 (lp), 18145 (G), Koslowsky 208 (κ), Skottsberg 950 (s). Rio Negro & Neuquén: Region of N
Huapí, de Agrasar et al. 12375 (mo), De Barba 7^9 (lil), Bòcheret al. 1824(c), Bridarolli 2129 (lp), Bu
(us), Burkart 12373 (f), Cabrera & Job 157 (f, ny), 5839 (f, lp), Cordini 217 (f, us), Hosseus 358 (
Kurtz 6366 (cord), Meyer 7661 (gh), Otto 5758 (f), Parodi 11556 (F), 11716 (mo), Pérez Moreau 504
Mendoza: Molis et al.'381 (lil), Ruiz Leal 1128 (f), 1725 (f), 2314 (f), 2327 (f), 4140 (lil), 1132
Buenos Aires: Gamerro 1181 (lp), Hunziker 1020(ul), Krapovickas 2528(lil, us), Miers 1423(us), P
8979 (gh), Spegazzini 234 (lp), Venturi 249 (cord, f). Santa Fe: Molfino 5757 (ba). San Juan: Flor
Tucumán: Villa Cerenzo 3426 (lil). Salta: Venturi 9860 (f, gh, s, us). Jujuy: Venturi 10142 (us
BOLIVIA: Cochabamba: Adolfo 386 (us), Vgent 4659 (gh, mo, wis). La Paz: Buchtien 673 (ny, us
4444 (f, gh, us), Mandón 335 (f, g, gh, k, p, s), Rusby 1829 (ny), Tate 233 (ny).
CHILE: Magallanes: Clarence Island, Exp. Antarct. Beige398(BR);Grevy Island, Vervoorst 30
Navarino Island, Skottsberg 174 (s); other localities, Alboff 140 (sgo), Andersson 326 (s), 327 (s), B
Blake (c, gh), Commerson (G, p), Coppinger (bm), Cunningham (κ, ny), Dudley et al. 71 (sgo), Dús
(mo, s), 477 (s), Eyerdam et al. 24150 (F, G, gh, mo, s), Hombron (p), King (κ), Le Guillou (p), Lechler
only), Magens 55 (CONC), Marivauit (p), Moore & Goodall 24 (c, κ), Moore 2137 (c, κ), Ortega 936
Pfister & Ricardi 171 (lil), 11875 (conc), 11985 (conc), 12022 (conc), 12066 (conc), 12102 (conc), Pis

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DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 17. Map showing distribution of Galium aparine.

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 ALLERTONIA

2470 (CONC), 2575 (conc), Ricordi & Matthei 154 (conc), Safford 29 (ν, us), Savali
Skoltsberg 177 (s), U. S. Expl. Exped. (gh, mo, ny, p, us). Aisén: Bruzzone 205 (lp), Esp
18385 (conc), 18430 (conc), 18513 (conc). Chiloé: Dilfin (sgo), Hooker (κ), Jung
Marticorena 1728 (conc), Miers (bm), Pennell 12563 (f, gh, ny, s, sgo, us), Ricardi &
(conc). Osorno: Sparre <5t Smith 294 (conc). Valdivia: Behn 21409 (conc), Bridges 649
(sgo). Cautín: Sparre 3414 (s). Bío-Bío: Barros 3661 (f). Concepción: Álvarez 10569 (co
(f), Holway 148 (ny, us), Macrae (g, κ), Martius (br), Palma-Inostroza 35191 (conc), Pa
(conc). Ñuble: Junge 3211 (conc), Pfister 6567 (conc). Maule: Reiche (sgo). Curicó
O'Higgins: Bertero 285 (a), 286 (bm, g, p), Marticorena & Weldt 649 (conc), Pfister
13064 (conc), 13080 (conc), Re id (κ). Santiago: Biese 226 (lil), Claude Joseph 2901 (
(conc), Garaventa 1279 (gh), Guzman (sgo), Hastings 83 (us), Killip & Pisano 39670 (us
3608 (f), 3609 (f), Mahu 4364 (conc), Nazarre (ny), Pérez Moreau 23410 (f), Schleg
(conc), Sparre 10980 (conc). Valparaíso: Barros 3648 (f), Buchtien 12092 (us), Claude
Gaudichaud 2612 (p), 2613 (p), Gunckel 18976 (lil), Holway 16 (us), Landbeck 936b (sgo
247 (this collection was given a name by Kunze which, however, was apparently never va
see Ehrendorfer, 1955, p. 537). Aconcagua: Salazar 11625 (conc). Coquimbo: Bailey (sgo
3598 (f), 3601 (f), 3606 (f), 3612 (f), 3613 (f), 3614 (f), 13790 (conc), Borchers (goet), C
(us), 4478 (us), 4479 (us), Granjot 377 (sgo), Jiles 345-b (conc), 1450 (conc), 3815 (conc), M
169 (conc), 1409 (conc), 1555 (conc), 1582 (conc), Montero 2831 (gh), Muñoz & Co
Skottsberg 780 (gb, s), Sparre 2820 (s), 2877 (s), 2900 (s), 3046 (κ, s), Torres 14129
Morrison 16309 (gh), 16434 (G, gh). Atacama: Barros 3617 (f), 3618 (f), Geisse 141
Ricardi & Marticorena 3885 (conc), Rivera 936c (sao). Antofagasta: Barros 3620 (f)
1050 (gh), Werdermann 863 (f, g, gh, lil, mo, ny, s, us), Worth & Morrison 16121 (g, g
Ricardi et al. 155 (conc), 198 (conc), 1326 (conc), Werdermann 763 (bm, f, g, gh, lil, m
Juan Fernández Islands: Masafuera, Meyer 9357 (mo, s), Solbrig et al. 3652 (gh, ny, u
(gh, s, us).
COLOMBIA: Nariño: Triana 3100 (col, ρ), Wiggins 10587 (f, ny, us). Magdalena: Cuatrecasas &
Romero 24453 (us, ven).
ECUADOR: Azuay: Camp E-2135 (s). Cañar: Camp E-2437 (ny). Chimborazo: Soli's 7170 (f).
Tungurahua: Asplund 8308 (s), Solis 8699 (f). Pichincha: Asplund 6749 (s). Imbabura: Penland &
Summers 823 (f, gh). Carchi: Steere 8161 (f, us), 8164 (f, ny), 8166 (f, s), Wiggins 10642 (f).
PERU: Arequipa: Pennell 13270 (f, gh, ny, us). Puno: Metcalf 30482 (f, mo). Cusco: Herrera 534 (f,
us), 2077 (f, us), Marin 164 (F, lil), Ugent 4089 (usm, wis). Ayacucho: Killip & Smith 22261 (f, ny).
Apurímac: Iltis & Ugent 635 (usm, wis), 709 (mo, usm, wis), 710 (wis). Huancavelica: Tovar 162 (us, usm).
Junîn: Esposito 11076 (uc, usm), Killip & Smith 21901 (F, ny, us), Soukup 1984 (us). Lima: Asplund 11303
(s), 13749 (s), Bail (gh, k, ny), Ferreyra 3973 (uc, usm), 7026 (uc, us, usm), 8680 (uc, usm), 11288 (uc, usm),
14141 (usm), Ferreyra & Cenate 16551 (uc, usm), Grant 7424 (gh, ny, us), Killip & Smith 21596 (f),
Macbride 5893 (f, s, us), Macbride & Featherstone (F, gh, s, us), Pennell 14613 (F, gh, ny, s, us), Raimondi
12383 (f), Rose 18607 (ny, us). Huánuco or Pasco: Ferreyra 9507 (uc, usm), Reente-Horsch 19254 (F).
Ancash: Ferreyra 7339 (uc, us, usm). La Libertad: Ferreyra 8607 (uc, usm). Amazonas: Mathews (bm, g).
URUGUAY: Montevideo: Herter 60364 (mo, ny, us), Legrand 264 (f). Canelones: Fruchard 919 (p).
As might be expected, Galium aparine is found at generally lower elevations in the
far south than closer to the equator; yet as far north as Lima (12° S.), collections have
been made at elevations of only 150-250 m.
The question of where Galium aparine is native and where introduced will proba
bly always remain moot. It is highly probable that the South American plants are for
the most part native, but that the species has also been introduced from Europe early
and repeatedly in grain, on clothing, and in animal fur. The highly effective dissemina
tion mechanism, as well as the early maturity and ready germination of the seeds, have
made this species one of the most cosmopolitan of plants. It is extremely adaptable as
to soil quality, plant association, altitude, human interference, and light. Although it
does not grow in deep shade, it thrives in moderate shade or in the open, providing that
there is sufficient moisture, at least during the spring. An extremely prolific annual, its
seeds will germinate in forest, scrub-land, rock slides, road-cuts, stream banks, and
grassy or ploughed fields. If conditions are favorable, the vigorous shoots climb many
decimeters high on whatever scaffold presents itself. Under less favorable conditions
the plants are often less than 10 cm. high, but nonetheless soon flower and set seed. As a
garden weed, this species is one of the most persistent.

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

d
Figure 18. Galium pseudotriflorum·, a, part of branch, x 5/8; b, leaf, under side, x 6; c, branchlet,
showing flower and ovary, with bracts, x 12; d, fruit, x 12. a from Cazalel & Pennington 5396, b from
Langenheim 3647 (type), c from Core 1625, d from Barclay & Juajibioy 7965.

There are no collections of Galium aparine between Colombia (Nariño, Magda

lena) and Nuevo León, Mexico. With such a discontinuity of distribution, one would
expect some morphological difference in the two populations; yet the same great range
of variability exists in South America as in North America. Although the possibility
must be recognized that G. aparine, with its extraordinary ability to disperse and to
adapt, is an introduction in South America, this interpretation can probably be
rejected, especially considering that a similar discontinuity exists between the two
closely related but clearly different species G. trißorum Michx., of the Northern
Hemisphere, and G. pseudotriflorum, of South America.
It is possible that intensive study might eventually demonstrate that South Ameri
can Galium aparine is specifically different from G. aparine of Europe and North
America. In consideration, however, of the great likelihood that South American
plants have been hybridizing for nearly five centuries with plants repeatedly intro
duced from Europe, it seems unlikely that this tangle will ever be unraveled.

15. Galium pseudotriflorum Dempster & Ehrendorfer, sp. nov. Figure 18.

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 ALLERTONIA

Planta perennis herbacea caulibus repentibus vel in fruticibus ad 3

bus; caulibus subglabris vel angulis brevissime scabridis; internodii
2-4(-5)-plo plerumque longioribus; foliis in quoque verticillo quinqu
parte maxima 8-20 mm. longis, vulgo margine et costa pilis brevibus vali
aculeatis scabridis, oblanceolatis (raro obovatis), basim versus gradatim
saepe subpetiolatis, apice abruptius angustatis et pilo subrigido term
centiis ramulis brevibus lateralibus enatis, unaquaque cymulis 1-3 c
imperfectis composita, cymula omni bracteis 2 vel pluribus vulgo subten
ultimis ramulisque subaequalibus; ovarii pilis saepe longis plerumque g
uncinatis, sursum deorsum flexis vel basi deorsum et apice sursum c
rotata glabra subviridi vel cremea vel aliquando subrubella, lobis ova
apicibus brevibus obtusatis; fructus pilis quam corpore plerumque br
Type: COLOMBIA: Boyacá: Páramo de Guantiva near km. marker
Belém and Susacón, Langenheim 3647 (uc holotype; isotype at us).
Distribution: Andes of Bolivia, Ecuador, Colombia, and western V
2,000-4,000 m., in moist places along streams and among shrubbery
have seen one specimen of this species from Costa Rica {Weston 10062
from Peru, where it might be expected to occur.
Additional collections: BOLIVIA: La Paz: Prov. Larecaja, vicinity of Sorata, Mandó
s); Prov. Nor Yungas, Unduavi, Rusby 1835 (ny), Buchtien 672 (w), 3003 (f, ny, us),
ν y), Eyerdam 25130 (f, uc).
COLOMBIA: Putumayo: Encano, Fosberg 20417 (p, us). Nariño: Chiles, Wiggins 10
ny, uc). Cauca: Quebrada del Duende, Cuatrecasas 18936 (f, gh). Tolima: Páramo de R
(ny), 3112 (ny); Auxilio, Core 1625 (ny, us). Valle del Cauca: Quindió La Linea, Dr
Páramo de Bavaya, Cuatrecasas 20222 (f). Meta: Río Arroz, Fosberg 20922 (ny, us
Above Bogotá, Pennell 1930 (f, gh, mo, ny). Santander: Vetas, Killip & Smith 17382
Smith 19509 (f). Magdalena: Sierra de Perijá, east of Manaure, Cuatrecasas & Romer
(us); Santa Marta Range, Weston 10374 (uc).
ECUADOR: Loja: Casapiamba, André 2999 (κ). Azuay: Páramo del Castillo, C
Hoyada de Galápagos, Steyermark 53474 (f). Cañar: Ventanillas, Fosberg & Priet
Chimborazo: Between Culebrillas and Yanayacu, Solis 7613 (f). Tungurahua: Minza, Penland &
Summers 342 (f, gh). Cotopaxi: Pilaló, Holm-Nielsen & Jeppesen 1272 (F, ny). Pichincha: Fosberg 23228
(us), Asplund 18985 (s), Barclay & Juajibioy 7965 (ny), Solis 8328 (f), Steere & Camp 8323 (F), Firmin 644
(f). Napo: Cerro Antisana, Grubb et al. 649(κ, ny). Imbabura: Cayambe, Drew & Wiggins27(f), Cazalet &
Pennington 5396 (κ, ny, uc). Carchi: Tufiño, Steere 8028 (F, ny, us); Salado, Steere 8105 (f, us), Tulcán,
Asplund 17066 (s), La Rinconada, Hitchcock (gh, ny).
VENEZUELA: Mérida: Distr. Rangel, above Las Piedras, Ruiz-Terán et al. 8248 (ven); Páramo de
Santo Domingo, Merxmïiller 22941 (m).
As the name indicates, this species has some of the basic characteristics of Galium
triflorum Michx., of the Northern Hemisphere. Galium pseudotriflorum is, however, a
longer plant with smaller organs, and the leaves, besides being generally smaller, are
definitely narrower and often coriaceous, with a much more pronounced tendency to
be broadest in the apical region. The plants climb and clamber aggressively, rather
than sprawling or spreading radially on the forest floor like those of G. triflorum.
Consonant with the climbing habit, G. pseudotriflorum has retrorse hairs on its leaves,
whereas the leaf hairs of G. triflorum are antrorse.
Considering these quite striking and obvious differences between the two species,
their resemblances are remarkable and surely indicative of a common ancestry more
recent than that which links them with other Galium species. Common characteristics
include the inflorescence pattern, the long ovary hairs that tend to turn downward
rather than the more usual upward and that give an appearance of great width to the
ovaries, and not least the almost perfect regularity with which the leaves are five or six
to each node, rarely exceeding or falling short of those numbers.
The northernmost collection of Galium pseudotriflorum that I have seen is in

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Figure 19. Map showing distribution of Galium mexicanum and G. pseudotriflorum

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 ALLERTONIA

Costa Rica, and the southernmost collection of G. triflorum is in Ver

separating distance of about 1,700 km.
Although the ovary hairs are most often turned downward at the ap
always so. Sometimes they are recurved at the base and turned upwar
either case, the result is an effect of flatness at the top of the ovary
after the corolla has fallen.
From herbarium specimens it is sometimes not easy to distinguish perennial
Galium pseudotriflorum from annual G. aparine. The fewer number of leaves (five or
six in a whorl), the usually smaller leaves, the often downturned ovary hairs, and the
often conspicuous triads of pedicels, particularly in old material, are indicative of G.
pseudotriflorum. Indicative of G. aparine are a greater number of leaves per whorl,
leaves that are often very long, albeit sometimes very narrow, round-appearing ovaries
with invariably upturned hairs, more amorphous inflorescences, and sometimes very
large fruits.

16. Galium mexicanum Kunth in H. B. K. Nova Gen. et Sp. 3: 337. 1818. Type:
Original type presumed lost. Neotype (Dempster, 1976): MEXICO:
Guanajuato: 18 miles east of Guanajuato along road to Dolores Hidalgo,
Solbrig & Ornduff 4540 (uc). Figure 20.

Figure 20. Galium mexicanum\ a, flowering branch, x 5/8; b

side, χ 6; d, flower, * 12; e, fruit, x 12. a from Pursell et al.
Steyermark 61785.

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 DEMPSTER: GALIUM IN SOUTH AMERICA. II.

Galium caripense Kunth in H. B. K. Nova Gen. et Sp. 3: 337. 1818. Type: VENEZUE
"Prope Caripe CumanensiumHumboldt (p holotype, photo seen; isotype at p).
Galium piliferum Kunth in H. B. K.. Nova Gen. et Sp. 3: 337. 1818. Type: COLOMB
Andibus Quinduensium, prope Los Gallegos et Quebrada de Tochecito," Humbold
photo seen).
Scandent or trailing perennial herb, the stems 10-90 cm. long, set with aculeolate
hairs; internodes 2-6 times as long as leaves; leaves mostly 6-12 at each node, more or
less coriaceous, often revolute, mostly 7-17 mm. long, 1-nerved, linear to oblanceolate,
tapered gradually to base and abruptly at apex, armed with a stoutish terminal hair,
the surfaces glabrous, the margins and midribs set with coarse aculeate hairs; glandular
cells none; flowers very small, few to many on complex lateral branches with reduced
leaves, the branchlets divaricate and very slender, the filiform pedicels several times as
long as flowers or fruits; flowers hermaphrodite; ovaries covered with fine upwardly
appressed curved hairs; corollas white (or pink?), glabrous or shortly hispid externally,
campanulate, cleft about halfway, the lobes more or less erect, the apices often strongly
reflexed; stamens usually included; fruits densely set with very short, upwardly curved
or uncinate hairs.
Distribution: Venezuela (Distrito Federal and Monagas) and Colombia (Cundi
namarca and Tolima), 900-2,800 m. (Figure 19), on mountain slopes among
limestone boulders, or in wet places among grass or open forest.
Additional collections: COLOMBIA: Cundinamarca: Rio San Francisco, above Bogotá, Perirteli
1927 (F, GH, mo, ny); near Bogotá, Ricordi 10a (f).
VENEZUELA: Distrito Federal: Colonia Tovar, Fendler 539 (G, gh, goet, k, mo, ny, p), Moritz 467
(GH, p); Avila, Pittier 9522 (f, gh, ny, us, ven), Köhl 124 (ven); Los Venados, H. C. 149a (ven). Monagas:
Vicinity of Caripe, Funcke 128 (bm, g, κ, p), Pursell et al. 8456 (ny, us, ven), 9389 (ny, us, ven), Moritz 467
(bm, hal), Steyermark 61785 (f, gh, ny, us, ven).
There seems no reason to consider the South American plants as other than Galium
mexicanum subsp. mexicanum, although the range of morphological variation is less
in South America than in Mexico and Central America.

Disposition of names (accepted epithets are in boldface type)

Galium andicolum Krause = G. weberbaueri
Galium aparine L.
Galium aparine var. microphyllum Clos = G. aparine
Galium aparine var. pseudoaparine Speg. = G. aparine
Galium australe Reiche = G. aparine
Galium boyacanum Dempster, sp. nov.
Galium canescens Kunth
Galium canescens f. alstonii Steyermark = G. orizabense
Galium caripense Kunth = G. mexicanum
Galium chilense Hook. f. = G. aparine
Galium chonosense Clos = G. aparine
Galium diffusoramosum Dempster & Ehrendorfer, nom. nov.
Galium diffusum Clos = G. diffusoramosum
Galium ferrugineum Krause
Galium ferrugineum Krause subsp. ferrugineum
Galium ferrugineum subsp. jamesonii Dempster, subsp. nov.
Galium fuegianum Hook. f.
Galium huancavelicum Dempster, sp. nov.
Galium lappaceum Ruiz & Pavón = G. obovatum
Galium larecajense Wernham = G. aparine

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 ALLERTONIA

Galium límense Standi. = G. weberbaueri

Galium mexicanum Kunth
Galium murale (L.) Allioni
Galium obovatum Kunth
Galium orizabense Hemsl.
Galium orizabense subsp. laevicaule (Weatherby & Blake) Dempster, comb, et sta
nov.

Galium ovalleanum Philippi

Galium parisiense L.
Galium peruvianum Dempster & Ehrendorfer, sp. nov.
Galium piliferum Kunth = G. mexicanum
Galium pilosum var. laevicaule Weatherby & Blake = G. orizabense
Galium pseudoaparine Griseb. = G. aparine
Galium pseudotriflorum Dempster & Ehrendorfer, sp. nov.
Galium rotundifolium var. uncinulatum Kuntze = G. obovatum
Galium trianae Wernham = G. canescens
Galium weberbaueri Krause
Sherardia muralis L. = G. murale

LITERATURE CITED
Dempster, L. T. 1976. Galium mexicanum of Central America and western North
378-386.
1978. The genus Galium in Mexico and Central America. Univ. Calif. Pubi. Bot. 73: 1-33.
1980. The genus Galium section Lophogalium (Rubiaceae) in South America. Allertonia 2:
247-279.
Ehrendorfer, F. 1955. Revision of the genus Relbunium (Endl.) Benth. et Hook. (Rubiaceae-Galieae).
Bot. Jahrb. 76: 516-553.
Gay, e. 1845-1854. Historia Fisica y Politica de Chile. Botánica [Flora Chilena].
Holmgren, P. Κ., & W. Keuken. 1974. Index Herbariorum. I. Ed. 6.
Jackson, B. D. 1893-1895. Index Kewensis.
Moore, R. J. 1975. The Galium aparine complex in Canada. Canad. J. Bot. 53: 877-893.
Reiche, e. F. 1896-1911. Flora de Chile.
Stafleu, F. Α., & R. S. Cowan. 1976. Taxonomic Literature. Ed. 2, Vol. 1.

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