Dempster. 1990. GENUS GALIUM IN SAmerica III

THE GENUS GALIUM (RUBIACEAE) IN SOUTH AMERICA.
III
Author(s): Lauramay T. Dempster
Source: Allertonia, Vol. 3, No. 3 (December, 1982), pp. 211-258
Stable URL: http://www.jstor.org/stable/23191024 .
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THE GENUS GALIUM (RUBIACEAE) IN SOUTH AMERICA. III.
Lauramay t. Dempster1
The present paper is intended to discuss all of the South American species of
Galium not included in two earlier papers (Dempster, 1980,1981) nor included (with a
few exceptions) by Ehrendorfer ( 1955) in the genus Relbunium. Thus the entire genus,
or pair of genera, is presumed to be comprised in these four papers, insofar as South
American representatives are involved.
The first of this series (Dempster, 1980) dealt with those South American Galia of
which the fruits have long, straight, specialized hairs (sect. Lophogalium); the second
(Dempster, 1981) with those species having fruits with hooked (uncinate) hairs. This
third paper deals with the remaining South American Galia, excluding those species
relegated to the genus Relbunium, discussed below.
Twenty-seven species are here included, all but one having glabrous, tuberculate,
or pubescent fruits but lacking long, specialized, fruit hairs. The exception is Galium
galapagoense, endemic to the Galápagos Islands, which should have been
included in the second paper. Twenty-two of the species are indigenous in South
America (taken to include the Galápagos). One of these occurs northward into Mexi
co, and another extends eastward to islands of the Scotia Arc and to the Kerguélen
Islands. Nineteen of the species are therefore restricted to continental South America.
The remaining five species have been introduced. Most of the species have four
"leaves" to a node (i.e. two leaves and two stipular appendages), but G. magellanicum,
G. surinamense, and G. ascendens among the indigenous species usually have five or
six. The five introduced species have more than four leaves to a node. Although the
genus Galium is usually defined as having opposite leaves and two or more leaflike
stipular appendages, resulting in a whorled condition of four to many (usually four or
six-twelve) "leaves" at each node, the situation in G. lilloi, G. equisetoides, and G.
diphyllum is anomalous in that the stipular appendages are often much smaller than
the true leaves, or are represented only by scales, or even are altogether absent.
The status of the genus Relbunium is seriously open to question, and unless it is
more narrowly defined than heretofore it is unacceptable, as the following discussion is
designed to show.
Endlicher (1839, p. 523), who proposed the name Relbunium as a section of
Galium, employed almost the same words that de Candolle (1830, p. 590) used in
describing his section Involucratae of the genus Rubia. With regard to the inflores
cence, which is the critical point in Relbunium, Endlicher stated: "Pedunculi axillares
apice gerentes bracteas quatuor verticillatas, involucrum constituentes, flores intra
involucrum solitarii v. terni, sessiles v. pedicellati." It can be seen that this definition
'The Jepson Herbarium, Department of Botany, University of California, Berkeley, California 94720.

212 ALLERTONIA 3:3
permits the inclusion not only of involúcrate sessile flowers, but also of involúcrate
inflorescences, although Endlicher in writing of pedicellate flowers was referring to
branchlets instead of true pedicels.
Bentham and Hooker (1873, p. 149), in elevating Relbunium to the status of a
genus, wrote: "Flores... bracteis 4 involucrati," and "inflorescentia involucrata," thus
continuing to accept Endlicher's permissive definition.
Neither Endlicher nor Bentham and Hooker made any binomial combinations
under the proposed section or genus. K. Schuman (1888) was the first to present a
treatment of Relbunium, assigning to the genus 22 species with appropriate binomials.
Ehrendorfer (1955), in his monographic treatment of Relbunium, gave a more
detailed description without altering the meaning: "Flores sessiles vel rarius brevissime
pedicellati, bracteis quaternis vel binis involucrati; bracteae involúcrales nonnun
quam ipsae pedúnculos gerentes." In using the word "pedunculos," Ehrendorfer
showed a probable awareness that these members are branchlets rather than true
pedicels. As type of the genus Ehrendorfer, without discussion, indicated R. hypocar

pium subsp. nitidum (H. B. K.) Ehrend. The nomenclatural basis of this trinomial is
Rubia nitida Η. Β. Κ., the basionym of Relbunium nitidum (H. B. K.) K. Schum.
(1888). It seems therefore that the correct typification of the genus should read:
Lectotype species: Relbunium nitidum (H. B. K.) K. Schum. (Rubia nitida Η. Β. Κ.) =
Relbunium hypocarpium (L.) Hemsl. subsp. nitidum (Η. Β. K.) Ehrend.; vide Ehren
dorfer in Bot. Jahrb. 76: 518, 536. 1955. Index Nominum Genericorum (Farr et al.,
1979) has stated the type species of Relbunium to be "non designatus."
If the definition of Relbunium be restricted so as to include only those species of
which the flowers are solitary, sessile, and hence involúcrate, there results a genus that
is at least discrete, if not monophyletic. If, on the other hand, "involúcrate inflorescen
ces" are included, the matter must be examined more carefully.
The prototype of the Galium inflorescence is the cymule, consisting of a terminal
flower and two branchlets, which may or may not be further proliferated. Furthermore
the flowers, although usually pedicellate, may be sessile, as in G. reynoldsii Dempster,
of Chile, and G. pennella Dempster, of Mexico. However, when we come to the group
of taxa that Ehrendorfer refers to Relbunium richardianum, the flowers are all sessile
and the inflorescences are congested, at least when young, and appear to be involú
crate. As these congested inflorescences develop, however, one or both of the branch
lets sometimes become quite long, and the sessile terminal flower is left alone with the
"involucre." If the branch development is bilateral, the resulting inflorescence is falsely
dichotomous; or if it is unilateral, the result is like the inflorescence of G. reynoldsii,
with sessile and apparently lateral flowers. Both of these situations can and do occur in
the Galium (or Relbunium) richardianum complex. True it is that the ultimate flowers
(and only those), being sessile, are involúcrate in the standard manner of Relbunium.
Ehrendorfer's inclusion of the Galium richardianum complex in Relbunium, fot
which there is plenty of precedent, in addition to blurring the definition of the genus,
ignores the strange and intimate relationship of these plants with Galium suffrutico
sum, which no one would call a Relbunium. In the northern part of their Chilean
range, the only difference between the two taxa lies in whether or not the flowers are
pedicellate. Both share the strange stem character, in which the "sides" of the stem are
nearly or quite reduced to grooves by the expanded "angles" (Figures 24e, 24f). Other

1982 DEMPSTER: GALIUM IN SOUTH AMERICA. III. 213
characters, such as overall size and slenderness, leaf shape and texture, and pubes
cence, occur indiscriminately in both species and are not correlated with pedicel length.
The range of the two is essentially congruent, except that I have seen no pedicellate
plants south of Maule. There are many mixed collections of the two forms or taxa. The
single difference between them is totally inadequate for generic differentiation, and
perhaps even for specific differentiation. The following reasons, however, may be
adduced for giving the two forms specific rank: (1) the difference between them is fairly
obvious at first sight; (2) there are no intermediates; and (3) the material with sessile
flowers apparently extends farther south than that with pedicellate flowers.
The generic segregation of Relbunium from Galium is thus highly questionable and
is, in fact, hard to defend. If there is any justification for a separate genus Relbunium, it
must be based on its solitary, essentially sessile, involúcrate flowers. Any other
interpretation, being vague and highly subjective, is inadmissible. I have therefore
included in the genus Galium, rather than in Relbunium, all species that do not
conform strictly to the definition that flowers are solitary and essentially sessile,
therefore individually involúcrate.
Closely related, and therefore included in the Galium richardianum complex, are
G. richardianum, G. araucanum, G. equisetoides, G. diphyllum, and G. suffruticosum.
Of these, only G. richardianum sometimes fulfills the narrow definition of Relbunium
here enunciated.
The Galium richardianum complex completely bridges the gap between Galium
and Relbunium in the following series: (1) G. suffruticosum has all flowers clearly
pedicellate and in no way suggests the involúcrate condition of Relbunium; (2) G.
araucanum, although related to G. suffruticosum in intimate fashion (see discussion
above), has sessile flowers and looks like a Relbunium, but upon analysis it is seen to
have only a few (i.e. the ultimate) flowers individually involúcrate; and (3) G. richardi
anum is exceedingly like G. araucanum, differing principally in having tuberculate
rather than smooth fruits; it does, however, in the eastern part of its range, include
some plants with all their flowers individually involúcrate.
This group of species thus demonstrates very clearly how the sessile, solitary,
involúcrate flower condition probably arose from the pedicellate, cymulose condition
that is normal and probably primitive in Galium. Two questions must be asked. (1) Did
this condition arise only once, i.e., is Relbunium monophyletic? Almost certainly it is
not. (2) Even if it were monophyletic, is a genus Relbunium justified, based on only one
character, narrowly defined as it must be to make any sense at all?
Although I am persuaded by the evidence that Relbunium as a genus is scarcely
defensible, yet for the purpose of the present paper those species with individually
involúcrate and essentially sessile flowers are considered as belonging in Relbunium.
This was to have been the third and final paper in my South American Galium series,
but it may be desirable to contemplate a fourth to include those presently excluded
species, i.e. those remaining in "Relbunium" sensu str.
The present treatment is a herbarium study, based on collections from the follow
ing herbaria: b, ba, bm, br, c, cge, col, conc, cord, ctes, ds, e, f, g, gb, gh, goet, hal,
JE, Κ, LE, LIL, LP, M, MO, MSC, MVFA, NY, P, R, S, SGO, SI, U, UC, US, VEN, W, and WIS.
Abbreviations are those recommended by Index Herbariorum (Holmgren et al., 1981).

All types and other cited specimens have been seen by me unless otherwise specified.

214 ALLERTONIA 3:3
Key to species
Flowers sessile or essentially so, individually subtended by a usually 4-bracted involucre. "Relbunium"
Flowers sessile or pedicellate, not all individually involúcrate, except sometimes in G. richardianum.
Fruits with specialized hairs.
Fruit hairs straight sect. Lophogalium (Dempster, 1980)
Fruit hairs uncinate.
Mainland South America (Dempster, 1981)
Galápagos Islands 1. G. galapagoense
Fruits glabrous, tuberculate, or merely pubescent.
Plants diminutive, condensed; Andean.
Fruits included or nearly so; leaves lanceolate, not ciliate, narrowed to a slender hyaline point; corolla
4-lobed; Chile 2. G. werdermannii
Fruits well exserted; leaves ovate-elliptic, ciliate, merely acute or obtuse at apex; corolla 3-lobed;
Peru and Ecuador 3. G. pumilio
Plants larger or, if diminutive, not Andean.
Leaves 4 or fewer at each node (rarely 5 in G. antarcticum).
Flowers sessile or essentially so.
Whorls with even-sized leaves, these spreading, linear or lanceolate-oblanceolate to somewhat
acicular.
Ovaries and fruits smooth; west of the Andean crest 4. G. araucanum
Ovaries and fruits tuberculate to short-pilose; east of the Andes 5. G. richardianum
Whorls with very unequal leaf pairs, the leaves strongly ascending to appressed-erect, acicular or
linear, except sometimes near plant base.
True leaves mostly 1.5-2 mm. long, acicular (except sometimes the lowermost); stems for the
most part strongly zigzag and tortuous 6. G. equisetoides
True leaves mostly 4-5 mm. long, linear, not acicular; stems not notably tortuous.
7. G. diphyllum
Flowers pedicellate.
Pedicels 15-30 times as long as fruit, filiform; central Chile 8. G. cotinoides
Pedicels not more than 5 times as long as fruit, usually not filiform.
Stem "angles" at least as large as the "sides ", usually much larger, often eliminating the "sides"
entirely.
Ovaries and fruits smooth; Andean, mostly in Chile 9. G. suffruticosum
Ovaries and fruits tuberculate or pubescent; Uruguay 10. G. uruguayense
Stem angles much smaller than the sides.
Flowers and fruits solitary in main axils (rarely 2 in G. antarcticum).
Bracts lacking below the flower.
Mature fruits consisting of two hard, spherical, glabrous, black mericarps; leaves
fleshy, the apices round or obtuse, not tipped with a hair; Tierra del Fuego and
Magallanes 11. G. antarcticum
Mature fruits various, but mericarps not hard and spherical; leaf apices obtuse or
acute, often tipped with a hair.
Fruit a red or orange, berry, 3-4 mm. wide; leaves 4 per node; Bolivia and
glabrous
Peru 12. G. mandonii
Fruit dry, pubescent, not red, 1-1.5 mm. wide; leaves 2-4 per node; northwestern
Argentina 13. G. lilloi
Bracts present below the flower 14. G. humile
Flowers and fruits variously disposed, but usually not solitary in main axis.
Fruits and flowers solitary at ends of branches, or rarely in main axils.
15. G. inconspicuum
Fruits and flowers in more or less complex inflorescences.
Inflorescence diffuse, terminal; leaves linear.
Plants, particularly the stems, coarse; transmontane in northern Argentina, south
ern Brazil, Bolivia, and Paraguay 16. G. latoramosum
Plants very slender; Coquimbo, Chile 17. G. lep turn
Inflorescence lateral on indeterminate
branchlets; leaves ovate.
Leaves glabrous, 4-8 (-15) mm. long; corolla and fruits glabrous, the fruits fleshy.
18. G. aschenbornii
Leaves hispid on upper surface, less than 3 mm. long; corolla hispid; fruits tubercu
late, dry 19. G. azuayicum

Leaves more than 4 at all or most nodes.

Annuals.
Coarse trailing plant with tuberculate fruits 20. G. tricornutum
Slender erect plant with glabrous fruits 21. G. divaricatum
Perennials; fruits smooth.
Mature fruit consisting of two hard, spherical, glabrous, black mericarps; leaf apices round or
obtuse, never acute nor tipped with a hair.
Leaf margins and stems scabrous with retrorse hairs.
Stems never stout, even when tall, the leaves small and the internodes usually long;
Colombia and Ecuador 22. G. ascendens
Stems stout, with longer leaves, or varying to low and slender, with relatively large leaves
and short internodes; Andes of Chile and Argentina 23. G. magellanicum
Leaves and stems essentially glabrous, or minute hairs, if present, antrorse; aquatic plant,
local near Buenos Aires 24. G. palustre
Mature fruit not as above; leaf apices acute.
Inflorescence a delicate tangle of retrorsely divaricate branchlets and pedicels; Surinam.
25. G. surinamense
Inflorescence not tangled, the branchlets and pedicels not retrorsely divaricate; local introduc
tions.
Plants large and coarse; leaves mostly 10-14 mm. long, not obviously broadest above the
middle; inflorescence large, diffuse 26. G. mollugo
Plants low and slender; leaves mostly less than 10 mm. long, obviously oblanceolate or
obovate; inflorescence on short lateral and terminal branches 27. G. saxatile
1. Galium galapagoense Wiggins in Madroño 20: 250. 1970. Type: ECUADOR:

Galápagos Islands: Santa Cruz Island (Indefatigable): Cerro Copa, Horneman
2 (ds holotype, not seen; isotypes at gh, s).
Slender perennial, growing in tangled masses on other plants; stems indeterminate,
pilose with short retrorse or long spreading hairs, the whorls remote; leaves in fours,
3-nerved, 10-15 (-30) mm. long, lanceolate to obovate, round-apiculate to merely
acute at apex, narrowed to a petiolar base, sparsely pilose above, more or less densely
pilose beneath with slender spreading hairs; inflorescences diffuse, on moderately
long, bracteate, lateral branches; flowers sessile; corolla hispid, the lobes obtuse;
ovaries with short upwardly curved or uncinate hairs; fruits not seen.
Distribution: Galápagos Islands: Santa Cruz Island (Indefatigable), 200-275 m.,
scrambling over shrubs, forming large mats.
Additional collections: ECUADOR: Galapagos Islands: Santa Cruz Island: Trail between Bella
Vista and Ponce Enrique, Fosberg 4478 (mo, uc); trail to Mt. Crocker, Wiggins 18534 (gh).
This species, with uncinate fruit hairs, should have been included in the second
paper of this series (Dempster, 1981). Galium galapagoense shows no very close
relationship to any of the mainland species. The 3-nerved, obovate, basally tapering
leaves distinguish it from G. orizabense, the 3-nerved leaves and obtuse corolla lobes
from G. obovatum, the leaf shape and indûment from G. canescens, and the sprawling,
tangled habit from all continental relatives. Unfortunately, I have been unable to
obtain mature material for better examination of flowers and fruits.
2. Galium werdermannii Standley in Pubi. Field Columbian Mus., Bot. Ser. 8: 398.
1931. Type: CHILE: Cautín: Volcán Llaima, Werdermann 1224 (us holotype,
not seen; isotypes at e, g, gh, lil, m, mo, s, u, uc; fragments at f). Figure 1.
Galium hypnoides Clos in Gay, Fl. Chil. 3: 183. 1848. Type: CHILE: Gay (p holotype; isotype at κ;
fragment at f); non Villars (1787).
Galium closianum Briq. in Ann. Conserv. Jard. Bot. Genève 11 & 12:191. 1908; non Timb.-Lagr. (1862).
"Double emploi accidentel du même nom pour deux espèces complètement différentes . . ."
Low, congested, dioecious (?), herbaceous perennial to 9 cm. high, rooting at

nodes; stems strongly angled, the internodes mostly 0.5-1.5 times as long as leaves;

ALLERTONIA
Figure 1. Galium werdermannii; a, habit, x 5/8; b, leaf, lower side, x 6; c, staminate flower, x 12; d,
branchlet with fruit, x 12. All from Werdermann 1224 (type).
leaves in fours, 1-nerved, thick, glabrous except at petiolar base, plane, ca. 3 mm. long,
lanceolate (-ovate), narrowed at base, tapering to a long apical hair, the margins
callose; glands none; flowers solitary in axils, 1 per node (or none), on pedicels 1 mm.
long or less; corolla (reddish?) rotate but not widely spreading, the semierect lobes with
blunt apices; ovary glabrous; fruit fleshy, reddish (fide Clos).
Distribution: Chilean Andes, Cautín to Bio-Bio, 1,100 m. (Figure 2).
Additional collections: CHILE: Malleco: Nahuelbuta National Park, Ricardi el al. 45351 (conc).
Bio-Bio: Copahue, Neger (m); Antuco, Gay 389, possibly isotype of G. hypnoides Clos (photo and frag
ment from Β at f; photos at GH, ny), Philippi 941 (sgo).
This is a very distinctive, mosslike species. The "calyx limb" mentioned by Clos in
his description of G. hypnoides is merely a somewhat prominent corona remaining on
the ovary after the corolla has fallen. Evidence, although scanty, suggests that the
species may be dioecious (see Figure 1).
3. Galium pumilio Standley in Pubi. Field Columbian Mus., Bot. Ser. 4: 298. 1929.
Type: PERU: Pasco: Cerro de Pasco, Macbride 3070 (f holotype; isotypes at F,
G, s, us). Figure 3.
Minute perennial herb with fibrous roots, much condensed and imbricated at base,
but giving rise to vertical shoots 15 mm. long with internodes twice as long as leaves;
stems smooth; leaves in fours, 1-nerved, ca. 2 mm. long, ovate or elliptic, thick,
narrowed to a petiole, acute to obtuse at apex, with 1 or 2 long apical hairs; leaves more
or less ciliate with long fine hairs, the surfaces largely glabrous; glands long and
narrow, in a subapical cluster; flowers perfect, solitary, lateral or terminal, nearly
sessile; corolla glabrous, yellow-green, 3-lobed; fruits glabrous, fleshy, some conspicu
ously exserted on pedicels to 6 mm. long.
Distribution: Peru and Ecuador, 3,700-4,300 m. (Figure 4), in moist grassy
places, the type occurring on limestone.
Additional collections: ECUADOR: Chimborazo: Towards Mt. Chimborazo, Asplund 7916 (s).
Napo: Antisana, Grubb et al. 666 (κ). Pichincha: Volcán Iliniza, Sparre 15630, in small part (s).
Standley and the collectors Grubb et al. have indicated: "peduncle elongating in
fruit." The few flowers that I have seen are nearly sessile, whereas well-exserted fruits
are a conspicuous feature of this diminutive plant.
With few flowers available for observation, it is impossible to say whether three
lobes are truly the norm in this species.

1982 DEMPSTER: GALIUM IN SOUTH AMERICA. III.
Figure 2. Map showing distribution of Galium cotinoides, G. inconspicuum, and G. werdermannii.

ALLERTONIA
riGURE 3. Galium pumilio; a, habit, * 5/8; b, leat, lower side, showing venation, χ 12; c, branchlet with
two leaves removed to show flower, * 12; d, fruit in position, * 6. a from Asplund 7916, b-d from Sparre
15630.
4. Galium araucanum Philippi in Anales Univ. Chile 85: 731. 1894. Type: CHILE:
Malleco: Near Renaico, F. Philippi 2035 (sGO holotype, photo and fragment
seen; isotypes at β and sgo, photos seen). Figure 5.
Galium suffruticosum var. araucanum Reiche, Fl. Chile 3: 147. 1902.
Slender polygamous plants, bunched, decumbent, or sprawling from a woody

base; stems 7-30 (-35) cm. long; "sides" of stems largely eliminated by the expanded
"angles"; branchlets 1-4 per node, i.e. a little fascicled; stems and leaves more or less
densely hispid to nearly or quite glabrous, particularly in the south; leaves in fours,
obviously 1-nerved but sometimes obscurely 3-nerved, 1.5-5 (-8) mm. long, linear or
lanceolate (to narrowly ovate), broad at base, somewhat leathery or fleshy, often shiny,
strongly ascending, often falcate, the callose margins revolute; glands narrow, in a
subapical cluster; inflorescences on short, few-flowered, spreading or ascending
branches, usually consisting of a peduncle topped by 4 leaves, a sessile flower, and 1 or
2 very short or somewhat elongated branchlets ending with an involúcrate flower;
corolla rotate, deeply cleft, more or less hispid externally, the lobes obtuse or merely
acute; fruits glabrous, a little fleshy.
Distribution: Chile: Valdivia to central Coquimbo, 5-1,900 (-2,400?) m. (Figure
6), in rocky places, thickets, and volcanic ash.
Additional collections: CHILE: Valdivia: Gay 133 (p), Krause 942a(sgo), Lechler (m), Philippi (g)\
Corral, Krause (sgo); Pampa de Negrón, Philippi 946 (hal, sgo). Cautín: Near Villarica, Lechler 1394 (g,
goet, ρ). Arauco or Malleco: Nahuelbuta, Philipp il (sgo). Malleco: Near Lonquimay, Merxmuller
24995 (m); Cuesta de las Raíces, Merxmuller 24925 (m). Concepción: Dessauer (m), d'Urville (g), Germain
(g, p, w), King (κ), Philippi (sgo); Tome, "PG." (sgo). Ñuble: Prov. Chillan, Philippi (sgo); Valle del
Renegado, Philipp il (sgo). Linares: Cordillera de Linares, Pablo (sgo). Maule: Ortega 943e (sgo);
Constitución, Reiche (sgo); Cauquenes, Reiche (sgo). Talca: Philippi (p. p., uc, us, not e). Curicó: Llico,
Landbeck 943c (sgo), Philippi? (sgo). Colchagua: Talcaregiie, Gay (p). O'Higgins: Agua de la Vida,
Philippi"! (sgo); Baños de Cauquenes, Dessauer 942d (sgo). Santiago: Dept. Melipilla, Las Vizcachas,
Morrison 16751 (p. p., mo, not others). Valparaíso: Dessauer (m), Gaudichaud24/ (p); Viña del Mar, Behn

Figure 4. Map showing distribution of Galium aschenbornii, G. azuayicum, G. ascendens, and G.

pumilio.

ALLERTONIA
21402 (conc); Curauma, Philippi (sgo); Cerro Campana, Borchers p. p. (bm); near Concón, Pöppig 42 (mo
only, not others); Quillota, Bertero 1014 (g, mo), Borchers 933a (sgo), Germain (g); cuesta del Melón,
Philippi 942c (sgo). Aconcagua: Catapilco, Philippi 943b (sgo); Dept. Petorca, Marlicorena el al. 181
(conc); Fdo. Las Anzal, Behn (ny). Coquimbo: Choapa, Landbeck 933 (sgo); Fray Jorge, Marlicorena et al.
460 (conc).
The relationship of Galium araucanum to G. richardianum is clear and close, but
its relationship to G. suffruticosum is puzzling. All three of these species share the
remarkable stem character that sets them apart as a group. Galium araucanum,
x
Figure 5. Galium araucanum; a, branch with flowers and fruits, 5/8; b, leaves, showing lower surface,
x 6; c, branchlet with * 12; d, branchlet with fruit and staminate flower, * 12; e, staminate
young fruits,
flower, * 12; f, pistillate flower, * 12; g, fruit, * 12. a from Germain, b, c, & g from Landbeck 933, d & e from
Philippi (Talca), f from Philippi 942c.

apparently restricted to Chile, is very similar to the allopatric G. richardianum east of

the Andes, from which it differs mainly in having smooth rather than papillose ovaries
and fruits. From the largely sympatric G. suffruticosum, however, it differs principally
in having sessile rather than pedicellate flowers, although it is also more slender on the
average. In all other respects the two species are remarkably alike: in leaf shape and
size, in range of indumentum, and in flowers. It is a very easy matter to distinguish the
two taxa, even in the several instances where collections are mixed, since presence or
absence of pedicels is immediately apparent without careful examination. For this
reason, despite suspicion of a very close genetic connection, with perhaps as little as a
single gene difference, the two are here considered as separate species.
It appears that Galium araucanum may be "pure" in the south, where G. suffrutico
sum does not occur, and where it also most resembles G. richardianum, and perhaps
genetically mixed farther north where it is sympatric with G. suffruticosum. It is
noteworthy that, although somewhat stouter habit and stems, and greater hairiness,
are apparent in G. araucanum of the north, possibly as a result of introgression, there
seems to be no intermediacy in the matter of pedicels.
5. Galium richardianum (Gillies ex Hook. & Arn.) Endl. ex Walp. Rep. Bot. Syst. 2:
459. 1843.
Very slender, straggling, or tufted polygamous plants 10-30 (-50) cm. high from a
woody base and slender taproot, rarely rooting at nodes; stem "angles" expanded,
often more or less completely eliminating the "sides"; branchlets 1-4 per node (i.e., a
little fascicled); leaves in fours, 1-nerved, 3-8 mm. long, linear or acicular, or rarely
lanceolate-oblanceolate, more or less leathery, tipped with a long hair, the callose
margins revolute, the midrib prominent below; glands none or sometimes in a subapi
cal cluster; inflorescences on short axillary peduncles, with 1 usually sessile flower
subtended by 4 leaves, or more often proliferous, with 1 flower and 1 or 2 long or very
short flowering branchlets; pedicels usually absent, but sometimes merely very short;
corolla small, cream-colored or greenish, rotate, the lobes obtuse or sometimes acute;
fruits dry, from (nearly smooth or) more or less papillose to short-pilose.
Key to subspecies
Stems glabrous or minutely papillose; leaves glabrous or with a few long marginal hairs.
5a. subsp. richardianum
Stems and leaves densely hispid 5b. subsp. haenkeanum
5a. Galium richardianum subsp. richardianum. Figure 7.

Rubia richardiana Gillies ex Hook. & Arn. in Bot. Misc. 3: 363. 1833. Type: ARGENTINA: Mendoza:
Andes, Gillies (κ holotype). There are two specimens in the type collection, almost certainly corres
ponding to the two localities mentioned in the original description, "Magote Aspera" and "Villavicen
zio." Perhaps it is reasonable, in our culture, to assume that the left-hand specimen refers to the first
locality and the right-hand specimen to the second locality. The right-hand specimen (presumably from
Villavacenzio) has been designated as the type by an unidentified previous worker, and I see no reason
to change it.
Rubia pusilla Gillies ex Hook. & Arn. in Bot. Misc. 3: 363. 1833. Type: ARGENTINA: Province of San
Luis, Gillies (κ holotype).
Galium richardianum Endl. ex Walp. Rep. Bot. Syst. 2: 459. 1843.
Galium pusillum Endl. ex Walp. Rep. Bot. Syst. 2:459. 1843. Based on Rubia pusilla Gillies ex Hook. &
Arn., but falsely attributed to Chile; non G. pusillum L. (1753).
Galium chaetophorum Griseb. in Abh. Königl. Ges. Wiss. Göttingen 19: 160. 1874. Type: ARGENTINA:
Córdoba: "Anbei der Calera von Est. Allende," i.e., probably near the lime pit of Estancia
Felsen
Allende, Allende, NNE. of Córdoba,
now Villa Lorentz 439 (goet holotype).
Relbunium chaetophorum K. Schum. in Mart. Fl. Bras. 6 (6): 114. 1888.
Relbunium pusillum K. Schum. in Mart. Fl. Bras. 6 (6): 117. 1888.
Galium ciliatum var. pusillum Kuntze, Rev. Gen. PI. 3 (2): 119. 1898.

ALLERTONIA
Galium ciliatum var. richardianum Kuntze, Rev. Gen. PI. 3 (2): 119. 1898.
Galium chaetophorum var. prostratum Kuntze, Rev. Gen. PI. 3 (2): 119. 1898. Type: ARGENTINA:
Jujuy. (Not seen.)
Galium chaetophorum var. strictum Kuntze, Rev. Gen. PI. 3 (2): 119. 1898. Type: ARGENTINA: Sierra
de Córdoba, Hieronymus. (Not seen.)
Galium patagonicum Kuntze, Rev. Gen. PI. 3 (2): 120. 1898. Type: ARGENTINA: Santa Cruz:
"Patagonia," at 50° 3'S., Moreno & Tonini 364 (NY holotype).
Relbunium patagonicum K. Schum. in Just's Bot. Jahresber. 26 (1): 393. 1900.
Relbunium echinocarpum Hassler in Bull. Herb. Boissier II. 7: 365. 1907. Type: PARAGUAY: Near
Caaguazú, Hassler 9134 (g holotype).
Relbunium richardianum Hicken in Physis (Buenos Aires) 2: 117. 1916. (Not seen.)
Relbunium richardianum subsp. pusillum Ehrend, in Bot. Jahrb. 76: 531. 1955.
Relbunium richardianum subsp. patagonicum Ehrend, in Bot. Jahrb. 76: 531. 1955.
Stems glabrous or minutely papillose; leaves glabrous or sometimes with a few long
marginal hairs; corolla glabrous or often with a few long hairs.
Distribution: Western Argentina (Santa Cruz to Jujuy), eastern Argentina
(Buenos Aires, Corrientes), Uruguay (Montevideo, Colonia), Brazil (Rio Grande do
Sul), Paraguay (Caaguazú), 700-1,500 m. (Figure 6), in grassy fields, on slopes and
hills, or on sandy soil by streams.
Additional collections:
ARGENTINA: Tierra del Fuego: Cabo Comingo, Goodall 3923 (gh).
Santa cruz: Roca, James 76 (bm); "high up river", Darwin 2045 (cge); 50° 3' S., Tonini & Moreno 141
Lago
(ny); Dept. Corpen Aike, south of Comandante L. Piedrabuena, Eyerdam et al. 24035 (uc); west of Puerto
Deseado, Eyerdam et al. 23864 (uc). Chubut: Rio Corcovado, Iliin 121 (bm, uc). Rio Negro: U. S. Expl.
Exped. (ny); San Carlos de Bariloche, Buchtien 1328 (m); Lago Nahuel Huapi, Ljungner 504 (gb), 832 (gb);
Lago Steffen, Pearson 2 (uc); Cerro Leones, Pearson 3 (uc); Estancia San Ramón, Pearson 1 (uc).
Mendoza: Sierra Pintada, Wilczek 400 (g); Villavicencio, Burkart et al. 14307 (us); Las Heras, Μ o lis et al.
380 (lil), Semper 101397 (msc). Córdoba: Lossen 349 (mo); near Córdoba, Kurtz 7243 (ny), Stuckert 16 (G),
3746 (g), 9411 (g); Estancia San Teodoro, Stuckert 21462 (g); Cerro de los Gigantes, Galander 1886 (g); La
Calera, O'Doneil & Rodriguez 463 (gh); Estancia Germania, Lorentz 42b (β photo, F fragment, goet, p);
Barrio S. Martin, Dept. Capital, O'Donell & Rodriguez 331 (gh, uc); San Francisco, Lorentz 325 (goet);
Sierra Achala, Kurtz 8360 (r). Santiago del Estero: Ojo de Agua, Pedro Garcia 910 (msc). Catamarca:
Lorentz & Hieronymus 439 (goet); near Potrero, Schickendantz 173Β (p); near the Ciénaga, Lorentz 55
(goet); Ambato, Tellechea 60 (br); Cuesta de la Viña, Cristóbal562 (ve). Tucumán: Valle del Tafí, de Haas
828 (u). Salta: Lorentz ά Hieronymus 5 (goet). Jujuy: Lorentz & Hieronymus 844 (goet); Calilegua,
Eyerdam & Beetle 22606 (uc); east of Jujuy, Eyerdam & Beetle 22382 (uc). Buenos Aires: Commerson (p).
Entre Ríos: Concepción del Uruguay, Lorentz 1096 (1080) (goet). Corrientes: Bonpland 331 (p); Laguna
Iberá, Schinini & Quarin 14395 (uc); Lavalle, Schinini 6596 (uc).
URUGUAY: Saint-Hilaire 2038 (p); Montevideo, Fruchard (p), Gay (ρ), Gibert 765 (goet); Point
Yeguas, Courbon 989 (p); Ile St. Gabriel, Courbon 1856 (p); Dos Hermanos (probably the islands), Tweedie
(κ).
BRAZIL: Rio Grande do Sul: Saint-Hilaire 1817 (p); Porto Alegre, Reineck 574 (ρ).
PARAGUAY: Caaguazú: Caaguazú, Balansa 1786 (g).
5b. Galium richardiamim subsp. haenkeanum (Gillies ex Hook. & Arn.) Dempster,
comb. nov.
Rubia haenkeana Gillies ex Hook. & Arn. in Bot. Misc. 3: 363. 1833. Type: ARGENTINA: Mendoza,
Gillies (κ holotype).
Galium haenkeanum Endl. ex Walp. Rep. Bot. Syst. 2: 459. 1834. Based on Rubia haenkeana, but
wrongly attributed to Chile.
Stems, leaves, and probably corolla (unknown) densely hispid.

In a species otherwise so consistently glabrous, or merely having a few long
marginal hairs on the leaves, this unusual, densely pilose plant cannot simply be
dismissed as extreme or even aberrant. I have seen no collections other than the type,
nor any intermediates.
Galium richardianum and G. araucanum have been included in the genus Relbu
nium by Ehrendorfer and others, because the ultimate flowers are sessile and involú
crate, but most flowers are apparently lateral, owing to the development and
proliferation of one or both branchlets of the cymule. Such branchlets are sometimes

very short, causing the cymule to remain congested, but often they are long and stout,
leaving the truly terminal flowers or fruits effectively lateral. See discussion of Relbu
nium on p. 212 of the introduction.
Figure 6. Map showing distribution of Galium araucanum, G. diphyllum, G. equisetoides, G. humile,

G. lilloi, G. richardianum, and G. suffruticosum.

ALLERTONIA
Figure 7. Galium richardianum subsp. richardianum; a, branch with flowers and fruits, χ 5/8; b, leaf in
x 6; c, leaf, showing venation, x 6; d, x
position, hermaphrodite flower with abortive ovary beside it, 12; e,
x
part of inflorescence with leaf removed to show young pistillate flower and two young fruits, 12; f, node of
x 12;
unusually acerose plant, showing perfect flower, g, fruit, showing very short pedicel, the leaves
removed, x 12. a from Schickendantz I73B, b from Lorentz & Hieronymus 5, c from Stuckert 16, d from
Wilczek 400, e & g from Galander 1886, f from O'Doneil & Rodriguez 331.
6. Galium equisetoides (Cham. & Schlechtendal) Standley in Field Mus. Nat. Hist.,
Bot. Ser. 11: 212. 1936. Figure 8.
Rubia equisetoides Cham. & Schlechtendal in Linnaea 3: 232. 1828. Type: Southern Brazil, Sellow (b
holotype, photo seen).
Relbunium equisetoides Ehrend, in Bot. Jahrb. 76: 533. 1955.
Wiry, polygamodioecious, scandent perennial, rooting at lower nodes, the long (to
60 cm.), slender, tortuous, apparently leafless stems sprawling or twining on shrubs;
Figure 8. Galium equisetoides; a, habit, showing tortuous branching, x 5/8; b, two nodes with
internode, showing abrupt change in size and shape of leaves,x 6; c, node, showing usual leaves and stipules,
χ 12; d, inflorescence with
hermaphrodite flower and fruits, * 6; e, young hermaphrodite flower with bract
removed to show position, x 12; f, part of inflorescence showing false dichotomy, with abortive ovary in
crotch, x 12; g, cupped corolla flattened, with transverse lines to indicate bending,x 12; h, fruit, showing only
one mericarp, x 12. a & f-h from Dúsén 15560, b from Leite 3709, c from Balansa 1789, d & e from Marchi
35756.

stems glabrous or minutely scabrous, the internodes 1.5-3 cm. long, the angles broad
and prominent but smaller than the sides; leaves mostly reduced to acicular, appressed
erect, glabrous scales, the true leaves 1.5-2 mm. long, the stipular appendages about

226 ALLERTONIA 3:3
half as long or greatly reduced; rarely some lower nodes with 2 ovate, spreading,
leathery leaves and nearly obsolete stipular appendages; inflorescences terminal and
lateral, diffuse, often falsely dichotomous, particularly in staminate inflorescences;
flowers sessile, all terminal but often seeming lateral because of unilateral branch
development, the ultimate flowers individually involúcrate; corolla rotate or the united
portion cupped, glabrous, deeply parted, greenish yellow, the lobes broadly ovate,
obtuse; fruits glabrous, slightly fleshy, the mericarps spherical.
Distribution: Southern Paraguay, Brazil (Rio Grande do Sul, Santa Catarina,
Paraná, Sâo Paulo), and Uruguay, 700-1,400 m. (Figure 6), in marshy places,
climbing among grasses and shrubs.
Additional collections: BRAZIL: Rio Grande do Sul: Sine loco, Leite 3709 (f); Montenegro,
Marchi 35756 (mo); Lagoa Vermelha, Leite 3032 (f). Santa Catarina: "Campo de Capivane" (perhaps Rio
Capivari), Ule 1763 (p); Sâo Joaquim, Reitz & Klein 7405 (uc); Mun. Lebon Regis, Reitz & Klein 13832 (us),
15504 (us); Mun. Campo Alegre, Smith ά Klein 7475 (us); Campo Alegre, Reitz & Klein 5243 (us); Mun.
Agua Doce, Smith & Klein 13472 (us); Santa Cecilia, Reitz & Klein 13473 (us); Curitibanos, Klein 3325 (us),
Reitz & Klein 11843 (us), Smith & Klein 8293 (us); Lajes, Reitz & Klein 14061 (us); Mun. Lajes, Smith &
Klein 8190 (us); Mun. Pòrto Uniäo, Smith ά Reitz 8708 (us); Mun. Caçador, Smith & Reitz 8938 (us), 9128
(us). Paraná: Serinha, Dúsén s.η. (f, g), 15560 (g); Curitiba, Dusén 8719 (f), Saito 451 (us), Dombrowski ά
Kuniyoshi 2422 (us); Tijuco Prato, Hatschbach & Guimaraes 25407 (uc). SÀo Paulo: Sâo Caetano, Brade
6707 (f); Campos do Jordäo, Kuhlmann 32420 (F), Leite 3653 (F).
URUGUAY: Sine loco, Arechavaleta 34 (f).
PARAGUAY: Valley of the stream Y-aca (perhaps Yacaré in Neembucu), Hassler 6817 (g); Caaguazú
or Caazapá: Pastores-mi (probably Pastoreo Mayo), east of the Cordillera of Villa-Rica, Balansa 1789 (f, g,
p). San Pedro: Colonia Nueva Germania, Rojas 2331 (f).
This species is a part of the Galium richardianum complex, which may be said to be
intermediate between Galium and Relbunium, a situation discussed in the introduc
tion. The broad stem "angles", characteristic of the G. richardianum group, are here
moderated, and the basic cymules are more proliferated than in G. richardianum and
G. araucanum.
The habit and appressed acicular leaves with much reduced stipular appendages
are very distinctive. One specimen from Rio Grande do Sul (Leite 3709) is normal for
Galium equisetoides in all respects except that some leaves on the lower nodes are
elliptic, spreading, and opposite, with essentially obsolete stipules (Figure 8b). The
change of leaf form on the stem is abrupt.
7. Galium diphyllum (K. Schum.) Dempster, comb. nov. Figure 9.

Relbunium diphyllum K. Schum. in Mart. Fl. Bras. 6(6): 117.1888. Type: BRAZIL: SÀo Paulo: Campos
da Bocaina at Garrafâo, Glaziou 11604 (p holotype, photo seen; isotype at p).
Rubia diphylla K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. IV. 4: 3. 1891.
Climbing or straggling plant 32 cm. long; stems scabridulous, the angles much
enlarged but not covering the sides, the internodes 3-6 times as long as leaves; leaves in
2 unequal pairs, acicular, glabrous, leathery, semierect, the true leaves 4-5 mm. long,
the stipules 1-1.5 mm. long; inflorescences diffuse, the short floriferous branchlets
more or less divaricate; flowers sessile; corolla rotate, glabrous, deeply divided into
ovate, acute lobes; ovaries glabrous; fruits not seen.
Distribution: Southeastern Brazil, Sao Paulo. The type collection is the only one
known. (Figure 6).
This species shares most characters with Galium equisetoides, but the much longer,
slightly spreading leaves differentiate it sharply from the 14 collections of G. equise
toides that I have seen. The acicular leaves are also in sharp contrast with the elliptic
lower leaves sometimes seen in G. equisetoides (Figure 8). Since such expanded lower

leaves were seen on only two of 14 collections of G. equisetoides, it is clearly possible

that they might be present on some lower stems of G. diphyllum as well. One of the five
stems of the isotype of G. diphyllum, however, has greatly reduced leaves on its lowest
portion, like the normal leaves of G. equisetoides. It is quite possible that more
abundant material would provide intermediates between the two species. It is
unknown whether G. diphyllum, like G. equisetoides, inhabits marshy places, but it
seems probable in view of the location, along the Bocaina River.
χ 3; b,
Figure 9. Galium diphyllum; a & a1, upper portion of branch, showing four leaves to a node,
of inflorescence showing unopened buds and older terminal flower with corolla ready to fall,x 12; c,
portion
x 6. All from Glaziou 11604 (type).
node from lower down, showing one of two leaves, and stipular scale,

ALLERTONIA
8. Galium cotinoides Cham. & Schlechtendal in Linnaea 3: 227. 1828. Type: CHILE:
Concepción: "In vicinia portus Talcaguano" (Talcahuano), Chamisso (hal
holotype; photos of isotype from β at gh, ny). Figure 10.
Slender, probably clambering perennial, the leaves mostly remote; stems scabrous,
the internodes 3-6 times as long as leaves; leaves in fours, 1-nerved, (3-) 7-10 mm.
long, lanceolate-oblanceolate, scabrous on margins and sometimes on upper surface,
acute at apex, tipped with a hair; glands in a subapical cluster; inflorescences merely
bracteate, several times tri- or dichotomous, the branches and pedicels filiform;
Figure 10. Galium cotinoides; a, habit, * 5/8; b, leaves,x 6;c, part of inflorescence,x 3;d, flower, x 12; e,
fruit, χ 12. a & b from Philippi 49, c-e from Reiche.

pedicels and pseudopedicels curved or sinuous, 20-60 mm. long; corolla glabrous, cleft
nearly to base, the ovate lobes semierect, the apices obtuse and inturned; fruits
minutely tuberculate, fleshy.
Distribution: Chile: Near the coast, from Concepción to Curicó, 50-150 m.
(Figure 2), in thickets.
Additional collections: CHILE: Concepción: Concepción, Jaffuel 1917 (gh), Philippi 49 (sgo);
Tumbes, Philippi 48 (sgo). Maule: Constitución, Reiche (sgo); Quebrada Honda, Philippi 53 (sgo).
Curicó: Alto de Vichuquén, Barros 3658 (f). Chile without locality, Eschscholtz (f ex le).
Galium cotinoides, with its many long, slender, curved, and naked pedicels, is one
of the most distinctive of all Galium species. The nearly leafless, much-branched,
filiform inflorescence, with very small flowers and fruits, resembles a mat of curled
horsehair. The frequent dichotomy in the inflorescence probably results from suppres
sion of the terminal flower, but this is not observable.
9. Galium suffruticosum Hook. & Arn. in Bot. Misc. 3: 363. 1833. Type: CHILE:
Valparaíso: Viña de la Mar (sic), Bridges 206 (κ holotype). Figures 11, 24e.
Galium chítense Endl. ex Clos in Gay, Fl. Chil. 3: 180. 1848. Type: CHILE: Valparaiso, Cuming 405 (κ
lectotype; isolectotypes at e); non Galium chilense Hook. (1847) nec Rubia chilensis Molina. The
confusion surrounding the name Galium chilense Endl. can be resolved only by the designation of a
type. It is clear that Rubia chilensis Molina is a Relbunium with "bacche rosse" and "foglie ovali". It is
probable also that de Candolle (Prodr. 4: 590. 1830) referred to the same plant. Hooker and Arnott
(1833), however, in citing the collection Cuming 405 from Valparaiso, were referring not to Rubia
chilensis Molina, as they thought, but to a Galium (G. suffruticosum). The name Galium chilense (i.e.,
excluding G. chilense Hook., which is another species altogether) originated with Endl. ex Clos in 1848,
wherein the Latin diagnosis reads "pedunculis . . . indivisis, apice quadribracteatis, unifloris," and the
discussion that follows, presumably by Clos, reads "pedúnculos... terminados por cuatro bracteas que
envuelven una á tres flores sesiles o muy desigualmente pedicelados." These two statements are
inconsistent, and it is to be presumed that the Latin statement should prevail. The Latin also states,
however, "foliis... lineari-mucronatis;... fructu glabro." Linear-mucronate leaves are appropriate to
G. suffruticosum rather than to Rubia chilensis Molina and, furthermore, no mention is made of
baccate fruits. Since no specimen is cited, one cannot be sure of the identity of Galium chilense Endl.,
although Clos, writing in Spanish, certainly was referring to G. suffruticosum. Philippi ( 1864, p. 4) gives
it as his opinion that Clos was wrong in referring Rubia chilensis, with oval leaves and red berries, to his
(or Endlicher's) Galium chilense, and that he should instead have referred it to G. relbun Endl. Follow
ing Philippus interpretation, I designate Cuming 405, cited mistakenly by Hooker and Arnott under
Rubia chilensis as well as correctly under G. suffruticosum, as the lectotype of Galium chilense.
Galium andinum Philippi in Linnaea 33: 98. 1864. Type: CHILE: Colchagua: Cordillera de Colchagua,
Landbeck (sgo holotype, photo and fragment seen).
Galium peteroanum Philippi in Anales Univ. Chile 85: 733. 1894. Type: CHILE: Curicó: In the Andes,
Vidal (sgo holotype, photo and fragment seen).
Rubia margaritifera Reiche in Anales Univ. Chile 106:978. 1900. Type: CHILE: Santiago: Cordillera de
Santiago, Cajon del Cepo, 2,200 m., Reiche (sgo holotype, photo and fragment seen).
Dioecious or polygamodioecious subshrub 5-30 cm. high, the many branches
tufted from a woody base or from slender woody stems well above ground; branches
stout and dense to slender and lax; stems more or less densely hispid with short or long
retrorse hairs, or at least scabrous, the "angles" greatly expanded, the "sides" more or
less completely invisible; internodes mostly 1.5-2.5 (-4) times as long as leaves; leaves
in fours, obviously 1-nerved but sometimes obscurely 3-nerved, 1.5-5 (-8) mm. long,
linear or lanceolate (to narrowly ovate), broad at base, somewhat fleshy, often shiny,
strongly ascending or sometimes a little arcuate, more or less densely hispid to sparsely
scabrous with short, straight, spreading hairs; glands usually numerous, in subapical
clusters; inflorescences on short, few-flowered, spreading or ascending branchlets
arising from the upper axils; flowering branchlets commonly consisting of a peduncle
and about 3 pedicellate flowers, but often more proliferous; peduncles 5-8 (-14) mm.
long, the true pedicels 1-5 times as long as flowers or fruits; corolla yellowish, rotate,
deeply cleft, more or less hispid externally, the lobes obtuse or merely acute; fruits
glabrous, a little fleshy.

ALLERTONIA
Figure 11. Galium suffruticosum; a, branch with fruits, x 5/8; b, leaves, χ 6; c, congested inflorescence
with pistillate flowers, x 12; note that stem surfaces are really "angles"; d, lax inflorescence with probably
hermaphrodite flowers, x 12; e, staminate flower, χ 12; f, branchlet with fruit, sterile ovaries, and sterile
flower, x 12. a from Buchtien 1202, b from Mahu 4250, c from Eyerdam 10114a, d from Worth & Morrison
16576, e from Morrison 17018, f from Looser 3607.
Distribution: Chile: Talca to southern Atacama, 200-2,700 m., and in Argentina

one collection from Mendoza near the Chilean border (Figure 6); rocky slopes and
outcrops.

Additional collections: ARGENTINA: Mendoza: Sta. Rosa de los Andes to Uspallato Pass,
Moseley (bm).
CHILE: Talca: Philippi (e, us). CuRicó: Los Queñes, Barros 3665 (f); Potrero Grande, Barros 3595 (f);
Cordillera El Planchón, Barros 3659 (f). Colchagua: Landbeck 942b (sGo), Philippi (g, w), Ricardi 1050
(CONC, lil), Simons 93 (sgo). O'Higgins: Tepidaria Cauquenes, Ball (e, gh), Dessauer (m), Philippi (m);
Agua de la Vida, Borchers (goet). Santiago: Monte Cantillana, Barros 3666 (f); Rio Yeso, Biese 104 (lil),
659 (lil); San Gabriel, Montero 542 (κ); El Volcán, Looser 3670 (f); Cerro de San Cristóval, Phillipil (sgo);
Cajón del Maipó, Mahu 4250 (uc), Villarroel 45340 (CONC); Peñalolén, Looser 3607 (f); La Obra, Montero
477 (f); near Santiago, Claude Joseph 630 (us), 2906 (us), Gay 428 (p), 429 (p), Philippi (f), Philippi?373 (g,
goet, k, p, s), Schlegel 45338 (conc), U. S. Expl. Exped. (us); Cordillera de Santiago, Hastings 189 (ny, uc,
us), Looser 5174 (gh), Philippi (w), Pirion 1738 (gh), Werdermann 480, a mixed collection (g, gb, m, ny, s,
u); Cerro Abanico, Grandjot 989 (conc); Rio Mapocho, Gersh 64 (wis); near Pérez Caldera, Marticorena &
Matthei 687 (conc); Dept. Melipilla, Morrison 16751 (g, gh, mo, uc); Cerro Manguchue, Looser 3606 (f);
Paso Cruz, Kuntze (ny, us); Altos de Tiltil, Philippi? (sgo). Valparaíso: Cerro Las Vizcachas, Eyerdam
10114a (uc), Hutchison 106 (f, sgo, uc), West 5147 (gh, uc); Viña del Mar, Bultman 23512 (conc);
Hacienda de Cauquenes, Dessauer (m). Aconcagua: Juncal, Buchtien 1202 (gh, lil, us), Spegazzini 426
(lp); Valle de Marga-Marga, Jaffuel & Pirion 3148 (gh). Coquimbo: Dept. Illapel, Morrison 17018 (uc),
Worth & Morrison 16576 (gh, uc); Quilmenco, Geisse (sgo). Atacama: Vallenar, Volckmann 54 (sgo).
Galium suffruticosum is often confused with non-fruiting plants of G. philippia

num Dempster. The stems will easily distinguish them, since in G. philippianum the
sides are much larger than the angles. See also the discussion under G. araucanum, as
to the relationship with that species.
10. Galium uruguayense Bacigalupo in Darwiniana 19: 515.fig. 4. 1975. Type: URU
GUAY: Maldonado: Sierra de las Animas, Marchesi 6493 (mvfa holotype not
seen; isotype at si, fragment seen).
Galium uruguayense var. echinulatum Bacigalupo in Darwiniana 19:51 i.fig. 5. 1975. Type: URUGUAY:
Maldonado: Sierra de las Ánimas, Marchesi 6493 bis (mvfa holotype not seen; isotype at si,
fragment seen).
Low, prostrate, perennial herb, rooting at lower nodes; stems glabrous or with
occasional hairs at nodes, the "angles" larger than the sides; leaves in fours, essentially
glabrous, 5-11 mm. long, narrowly elliptic or lanceolate to nearly linear, coriaceous,
1-nerved, acute at apex, with terminal hair, the margins notably callose; flowers solita
ry in axils of short leafy branchlets; pedicels 0.5-0.7 mm. long, reflexed in fruit; ovary
tuberculate or pubescent; corolla (ex char.) rotate, hispid, brownish red; fruit tubercu
late or pubescent.
Distribution: Uruguay: Maldonado, Lavalleja, and Tucuarembó. (Figure 12).
Of the two varieties described by Bacigalupo, I have seen only fragments of the
isotypes, supplemented by her excellent illustrations.
Galium uruguayense, of eastern South America, bears a relationship to G. richar
dianum similar to that of G. suffruticosum to G. araucanum in the far west. The
principal difference between the pairs in both instances lies in the presence or absence
of pedicels, and consequently of involucres. In both cases a close genetic relationship is
strongly suspected. The stems with conspicuously broad "angles" are characteristic of
all four species mentioned above, as are also the more or less callose leaf margins.
Galium richardianum in the eastern part of its range has ovaries and fruits that vary
from barely tuberculate to definitely pilose, whereas the type specimens of G. uru
guayense and its variety echinulatum have tuberculate and pilose ovaries respectively.
It seems highly probable that more material would show a graded series in G.
uruguayense similar to that seen in G. richardianum. Galium uruguayense appears to
be a somewhat lower and more condensed plant than G. richardianum.

ALLERTONIA
11. Galium antarcticum Hook. f. FI. Antarct. 303 (bis). 1846. Type: ARGENTINA:
Τ ierra del Fuego: Good Success Bay, Banks & Solander in 1769 (bm lectotype;
isolectotypes at GH, s, us). This first-mentioned collection, an appropriate
choice, was indicated by Moore (1968) as the lectotype. Figure 13.
Galium debile Banks & Solander ex Hook. f. Fl. Antarct. 303 (bis), nom. nud. pro syn. 1846.
Low plants 4-12 (-22) cm. high, much branched from fibrous roots, spreading
among grass, rooting at nodes; stems minutely scabrous or sometimes glabrous; leaves
in fours, glabrous or sometimes minutely scabrous, 1-nerved, 4-12 mm. long, fleshy,
oblanceolate to obovate, tapering to base, the apex round or at most obtuse; flowers
rarely more than 1 per node, on pedicels 1-3 times as long as flowers or fruit; flowers
perfect; corolla white, sometimes tinged with pink, the lobes 3 or 4; fruits dry, hard,
glabrous, the halves spherical.
Distribution: Argentina (Tierra del Fuego, southern Santa Cruz) and Chile
(Magallanes); Falkland Islands; South Georgia; Kerguélen Islands. Sea level to 170 m.
(Figure 14), in moist or wet places in grassy areas (Festuca) or bogs, or in sand, rock,
or gravel, from the seashore (above the tide line) or lake shore to comparatively dry
places at top of cliffs.
Figure 12. Map showing distribution of Galium latoramosum, G. leptum, G. mandonii, and G.
uruguayense.

Additional collections: ARGENTINA: Tierra del Fuego: Staten Island, Rodriguez 84 (f), Webster
(G); Isla Observatorio, Castellanos 12968 (g, gb); Bahia Aquirre, Moore 1888 (c, κ); Harberton, Castellanos
7913 (f), Goodall 645 (lp, ny, uc), Moore 1403 (κ); Remolino, Gusinde (K); Ushuaia, Alboff 347 (lp),
Castellanos 7909 (f), Goodall 3974 (gh), Skottsberg 145 (s); Lago Fagnano, Goodall3992 (gh), Moore 2617
(c); Río del Fuego, Holmberg & Calcagnini 3888 (f); Cabo Domingo, Moore 1488 (κ); San Sebastián,
Alboff (f, lp); Gable Island, Goodall 372 (lp). Santa Cruz: Rio Coy (Coig, Coyle), Hatcher (ny).
CHILE: Magallanes: Andersson 325 (f, s); Cape Horn, Eights (us); Hermite Island, Hooker (k);
Orange Harbor, U. S. Expl. Exped. (gh, us); Vicuña, Ricardi & Matthei 223 (conc); Punta Arenas, Dúsén
154 (s), Cunningham (?) (κ); Cabo Negro, Lechler 1092 (br, g, goet, s); Gregory Bay, Cunningham (κ);
Possession Bay, Blake (gh); Nicolas Bay, Andersson 324 (s); Str. Magellan, Whinnie (κ).
FALKLAND ISLANDS: Gaudichaud (g), Hamilton 69 (bm), Hennis (bm), Hooker (bm, k, u), Nicol
(bm), Vallentin 35 (bm). West Falkland: Vallentin (gh); West Point Island, Sladen J Β123/9 (bm); French
Bay, Lesson (k); Shallow Bay, Vallentin (us); Roy Cove, Vallentin (k); New Island, Dempster 4470 (uc);
Fox Island, Skottsberg (gb, sgo); Port Stephens, Moore 785 (c, us). East Falkland: Birger (s), Darwin
(cge), Greene 29/1 (κ), Hamilton (bm), Hill (κ), Lechler 121 (s), Moore 553 (gh, κ, lp, s), Skottsberg 50 (s),
104 (S), Sladen Fa53/49 (bm), Fa6/49 (bm).
SOUTH GEORGIA: Bonner 46 (bm), 47 (bm), 84 (bm), 94 (bm, k), 113 (κ), Greene 1070 (κ), 1485 (κ),
1502 (gh, κ), 1627 (κ), 1992 (κ), 2884 (gh, κ, s), 3045 (κ), Skottsberg (s), Sladen JB14/1 (bm), M. 1004 (κ),
M. 1013 (κ).
KERGUÉLEN ISLANDS: A üben de La Rüe (br), B.A.N.Z. Exped. (us), Coulter (f), Kidder (ny, us),
Morley (uc), Ott (f).
As the name suggests, this species has the southernmost distribution of all Galium
species, exceeding even G. fuegianum Hook, f., not only in respect to latitude, but
especially in its tolerance for a rigorously cold habitat, largely beyond the range of
trees. It is clearly a member of the G. trifidum complex, typified in North America (G.
Figure 13. Galium antarcticum; a, habit, χ 5/ 8; b, node with leaves,x 6; c, flower with 4-lobed corolla, «
12; d, 3-lobed corolla, χ 12; e, fruit, χ 12. a from Skottsberg 50, b & d from Dempster 4470, c from Moore
785, e from Andersson 325.

ALLERTONIA
Figure 14. Map showing distribution of Galium antarcticum and G. magellanicum.

trifîdum Michaux from Canada) but occurring in all hemispheres. Although very
closely related to G. magellanicum, G. antarcticum is distinguished by the constantly
four leaves per node and the usually solitary flowers. The leaves are also fleshy and
truly 1-nerved, the corollas frequently 3-parted. The solitary flower condition proba
bly represents a reduction of the ideal 3-flowered cymule, imperfect examples of which
occur rarely in this species.
I have seen aberrant corollas with three large lobes and one small; with three perfect
lobes and one bifid; and with four lobes and bearing only three stamens.
12. Galium mandonii Britton in Bull. Torrey Bot. Club 18: 263, as G. mandoni. 1891.
Type: BOLIVIA: Larecaja: Near Sorata, Rusby 1831 (= Mandón 334) (ny
lectotype; isolectotypes at bm, e, f, g, k, mo, msc, ny, p, s, us, w).
Figures 15, 24a.
Climbing or sprawling perennial with slender, annual, scabrous stems to 60 cm.
long from a diffuse fibrous root system; leaves in fours, 1-nerved, 4-20 mm. long,
oblanceolate, acute or apiculate, narrowed to a petiole, but the base broader, erect and
clasping, densely hispid, persistent; leaf surfaces nearly glabrous or the upper pubes
cent, the margins and midribs usually with moderately long, mostly retrorse, hairs;
flowers solitary in axils, mostly 1 at a node, the pedicels filiform, 2-8 mm. long, often
recurved in fruit; corolla glabrous or short-pilose, about as large as ovary, cleft nearly
to base, the lobes ovate, obtuse to merely acute; ovary 8-sulcate, glabrous; fruit a red or
orange berry, 3-4 mm. in diameter when dry, sulcate when turgid.
Distribution: Bolivia and Peru, from Tarija to Apurimac, 2,400-3,800 m.
(Figure 12), in wet places at edges of fields.
Additional collections: BOLIVIA: Tarija: Canaleta, Krapovickas et al. 18973 (ctes, wis).
Cochabamba: Cochabamba, Julio 11.16 (us); near Choro, Brooke 6022 (bm, ny), 6159 (bm); Liriuni,
Adolfo 338 (us). La Paz: Near Quime, Brooke 5291 p. p. (bm, ny); White 163 (ny); Illimani, Buchtien 3270
(f, ny, us); Unduavi, Rusby 1839 (ny, us, wis), Mandón 333 (ny, p, w); La Paz, Buchtien 4445 (f, gh, mo,
us); Pflanz 51 (f); Sorata, Mandón 334 (photo from b), Weddell (p).
PERU: Puno: Lake Titicaca, Manheim M108/ M96 (f). Apurimac: Valley of Rio Colcachaca, Iltis &
Ugent 545 (wis). Cuzco: Cuzco to Pisac, Iltis & Ugent 973 (ctes, gh, mo, msc, uc, us, wis). Huancavelica:
Churcampa, Antunez de Mayolo 248 (uc). Peru without locality, Haenke (ρ).
* 6; c,
Figure 15. Galium mandonii; a, fruiting branch, χ 5/8; b, leaf, upper side, showing venation,
* with leaves and fruit, * 6. a from litis & Ugent 973, b-d from Brooke 6022.
flower, 12; d, node,

ALLERTONIA
a c
Figure 16. Galium lilloi; a, branch showing mostly four leaves per node, * 5/8; b, branch showing two
leaves per node, * 5/ 8; c, node, showing lower side of leaf, and young fruit, * 6; d, node, showing flower, leaf
bases and stipular scale, x 12; e, fruit, χ 12. a from Venturi 1319, b & d from Ragonese 296, c from Pierotli
1127, e from Wall 344.
13. Galium lilloi Hicken in Darwiniana 1: 140. 1924. Type: ARGENTINA: Tucu
mán: Las Tranquitas, Dept. Burruyacú, Lillo 2560 (si holotype; fragment at uc).
Figure 16.
Slender creeper, rooting at nodes; stems strongly angled, usually gla
perennial
brous (or a little scabrous); leaves 2-4 at a node, often in unequal pairs, or the stipular
leaves reduced to a more or less minute scale; leaves 1-nerved, elliptic or oval to
oblanceolate, abruptly apiculate, truly petiolate, glabrous or often scabrous on midrib,
margins, and upper surface, the margins callose; flowers solitary in axils, rarely more
than 1 at a node, on slender pedicels 1-1.5 mm. long; corolla rotate, white, deeply
parted, the lobes ovate, shortly apiculate, minutely pubescent externally; ovary pubes
cent, about as large as corolla; fruits softly pubescent, 1-1.5 mm. broad, on reflexed
pedicels, the mericarps reniform.

Distribution: Argentina: Santiago del Estero to northern Salta, 700-3,000 m.
(Figure 6), in moist places among rocks.
Additional collections: ARGENTINA: Santiago del Estero: Grácián, Castellanos 33975 (f).
Tucumán: Anfama, Monetti (Osten 10971) (f); Dept. Tafí, Venturi 1319 (f, us), 6090 (us); Dept. Monteras,
Wall 344 (s). Catamarca: Fuerte de Andalgalá, Schickendantz309 (goet). Jujuy: Dept. Anta, Sierra Cresta
del Gallo, Ragonese 296 (ba, f); Dept. Capital, Sierra de Zapla, Cabrera et al. 23684 (si, fragment at uc);
Dept. Santa Bárbara, Santa Clara-El Fuerte, Cabrera et al. 27964 (si, fragment at uc); Dept. Ledesma, road
to Valle Grande, Cabrera et al. 27821 (si, fragment at uc). Salta: Dept. Capital, Quebrada de San Lorenzo,
Borsini 505 (lil); Dept. Orán, Pierotti 1127 (uc), 1308 (ny).
This is one of the most distinctive species of Galium, extraordinary in this genus in
having fewer than four leaves to many nodes. In fact, in half of the collections that I
have seen, two leaves per node is the commoner number. Galium lilloi is therefore, in
this one respect, probably among the most primitive of all Galium species, having
imperfectly developed the stipular leaflike appendages that have been considered
diagnostic for the genus.

14. Galium humile Cham. & Schlechtendal in Linnaea 3: 226. 1828. Type: Southern
Brazil, Sellow (Sello 2979) (β holotype, photo seen; fragment at f).
Relbunium humile K. Schum. in Mart. Fl. Bras. 6(6): 103. 1888.
Diminutive matted perennial, ca. 7 cm. high from fibrous roots; stems with very
prominent slender angles, these sparsely to densely set with long spreading hairs or
rarely glabrous; leaves in fours, mostly 3-5 mm. long, 1-nerved, elliptic or ovate
obovate, petiolate, acute at apex and tipped with a hair, sparsely to abundantly set with
long spreading hairs or rarely glabrous; glands in a subapical cluster; flowers solitary in
axils, on peduncles 1-2 mm. long, short-pedicellate or sessile above 2-4 bracts; corolla
greenish white, externally hispid; fruits dry, pubescent (or tuberculate), the mericarps
cerebriform.
Distribution: Southern Uruguay, Brazil, Paraguay, and adjacent Argentina,
2-1,650 m. (Figure 6), in damp places among grass or moss; meadows and bogs. For
excellent illustrations, see Bacigalupo (1975, p. 514).
Additional collections: ARGENTINA: Corrientes: Concepción, Petersen 8692 (c).
BRAZIL: Rio Grande do Sul: Porto Alegre, Reineck & Czermak 573 (f); Vila Oliva, Rambo 30875
(lil); Cachoeirinha, Rambo 39538 (lil); Sao Leopoldo, Leite 1634 (f). Santa Catarina: Mun. Säo
Joaquim, Smith & Reitz 10150 (us); Mun. Bom Retiro, Smith & Reitz 10349 (us), Smith, Reitz, & Klein
7725 (us); Mun. Pòrto Uniäo, Smith & Klein 12162 (us), Smith & Reitz 8908 (us); Mun. Chapecó, Smith &
Klein 11574 (us); Mun. Caçador, Smith & Reitz 9067 (us); Palhoça, Reitz & Klein 386 (us), 987 (us); Ilha de
Santa Catarina, Klein & Bresolin 8802 (us). Sao Paulo: Butantan, Krause 374 (f), 923 (f), 2569 (f).
PARAGUAY: Itapua: Villa Encarnación, Rojas (Osten 7979) (f). Misiones: Santiago, Pedersen 6585
(uc). Paraguarî: Ibitimi (Ybytimi), Balansa 1916 (f). Cordillera: San Bernardino, Rojas 1762 (f, lil).
Caaguazu: Yhu, Sparre & Vervoorst 2034 (lil).
URUGUAY: San José: Barra Santa Lucia, Osten 22451 (f).
Galium humile is very closely related to G. sphagnophilum (Greenman) Dempster,

of Mexico, with smooth fruits, and to its var. mazocarpum (Greenman) Dempster,
with tuberculate fruits, the only obvious difference being that the fruits of G. humile
are pubescent. Bacigalupo (1975) mentions three collections of Relbunium mazocar
pum (= G. sphagnophilum var. mazocarpum) from Entre Ríos and Corrientes, Argen
tina, and it is to be presumed that their fruits are tuberculate, although I have not seen
the material. In a former paper (Dempster, 1973b), I expressed some doubt as to the
taxonomic importance of the difference between smooth and tuberculate fruits in the
Mexican plants mentioned above. Similar doubt may be expressed with regard to the
South American plants, i.e. as to whether the difference between tuberculate and
pubescent fruits is taxonomically important, at least in regionally sympatric material
(but see G. richardianum and G. araucanum, which are allopatric).
In view of the great geographic gap between the South American and Mexican
plants, and in consideration of the fact that fruits of the South American plants vary
from tuberculate to pubescent, whereas those of the Mexican plants vary from
tuberculate to smooth, it seems best to keep the two populations as two separate
species, G. humile in South America and G. sphagnophilum in Mexico, even though

some individuals from the two populations, i.e. those with tuberculate fruits, may be
nearly indistinguishable.
In some Paraguayan material, the stems and leaves are essentially glabrous, yet the
corollas are hispid and the fruits pubescent.
15. Galium inconspicuum Philippi in Linnaea 33: 99. 1864. Type: CHILE: Talca:
Meneses, Volckmann (sgo holotype, photo and fragment seen). Figure 17.
Galium hypnoides var. inconspicuum Reiche in Anales Univ. Chile 106: 990. 1900.
Low slender plants 5-15 cm. high, tufted from fibrous roots; stems prominently

ALLERTONIA
angled, essentially glabrous or at most minutely scabrous; internodes about as long as
leaves; leaves in fours, more or less thick, 1-nerved, mostly 3-10 mm. long, linear,
tapered at base, acute but not pungent at apex, scabrous on the usually revolute
margins, reflexed above the erect base; flowers few, solitary, terminal or axillary;
pedicels 1.5-4 mm. long, reflexed at maturity; corolla glabrous or scabrous, deeply
cleft, the lobes obtuse, semierect; ovary glabrous; fruit fleshy.
Distribution: Andes from southern Neuquén, Argentina, northward in Chile to
Talca; doubtfully in Santa Fe, Argentina, 1,500-2,000 m. (Figure 2).
Additional collections: ARGENTINA: Neuquén: Lago Nahuel Huapí, Pérez Moreau 35515 (f).
Santa Fe: Carcarañá, Berndt 5217 (cord), a doubtful locality.
CHILE: Cautín: Cerro Malalco, Neger (m); Volcán Llaima, Sparte &. Smith 229 (conc), Sparre &
Constance 10636 (conc). Malleco: Volcán Lonquimay, Sparre & Constance 10899 (conc); San Carlos,
Barros 1152 ( f). Bío-Bío: Bureo, Claude Joseph 4113 (us).ÑuBLE:Chillán, Grau 1471 (m), Jäffuel 2798 (gh),
Pfister 963 (conc); San Carlos, Barros 3676 (f). Talca: El Picazo, Barros 3656 (f).
The many available specimens from Chile and the one from Andean Argentina are
morphologically indistinguishable from the Berndt specimen of east-central Argen
tina. The latter is an excellent specimen and well documented, but the locality should
nevertheless be questioned.
The leaves in this species are 1-nerved, but derivation from the 3-nerved condition
seems fairly obvious (see Figure 17b).
Figure 17. Galium inconspicuum; a, branchlet, « 5/8; b, leaf, lower side, showing venation, x 6; c,
flower, χ 12; d, fruit showing seeds through fleshy covering, x 12. a from Berndt 5217, b & d from Neger, c
from Pérez Mor eau 35515.
16. Galium latoramosum Clos in Gay, Fl. Chil. 3: 187. 1847. Based on Rubia intricata
Hook. & Arn. Figures 18, 24b.
Rubia ephedroides Cham. & Schlechtendal in Linnaea 3:231.1828. Type: "Brasilia meridionalis," Sellow
(hal holotype). Non Galium ephedroides Willk. (1852).
Rubia intricata Hook. & Arn. in Bot. Mise. 3: 362. 1833. Type: ARGENTINA: Buenos Aires: "Among
hedges of cactus, at San Isedro," (i.e. San Isidro), Tweedie (bm holotype). The label on the type reads:
"on cactus hedges Santecidro." Non Galium intricatum Reuter (1839).
Galium intricatum Endl. in Walp. Rep. Bot. Syst. 2: 459. 1843. Based on Rubia intricata Hook. & Arn.
Non Galium intricatum Reuter (1839).
Galium chapi Wedd. Chloris Andina 2: 36. 1859. Type (sterile): "In prov. boliviensis Azero," i.e.
BOLIVIA: Chuquisaca: Prov. Azero, Weddell 3699 (ρ holotype; isotype at f).
Galium tetragonum Griseb. in Abh. Königl. Ges. Wiss. Göttingen 19: 161. 1874. Type: ARGENTINA:
Sierra de Cordoba, near Ascochinga, Lorentz 162 (goet holotype).

Figure 18. Galium latoramosum; a, habit, x 5/8; b, fruiting branch, * 5/8; c, leaf, lower side, χ 6; d,
portion of pistillate inflorescence, showing short, stout stems, x 12; e, pistillate flower with long, stout
x 12; f, staminate flower seen from above, χ 12; x 12;
pedicel, exceptionally large, g, same, seen from below,
h, fruit, x 12. a & d from Cast ilion 7285, b from Schinini & Quarin 14357, c from Boelcke 4993, e from Kurtz
6566, f & g from Ar iza 1643, h from Lutz 78261.

240 ALLERTONIA 3:3
Rubia tetragona Κ. Schum. in Mart. Fl. Bras. 6 (6): 120. 1888.

Galium ephedrodes (sic) Kuntze, Rev. Gen. PI. 3 (2): 119. 1892. Based on Rubia ephedroides Cham. &
Schlechtendal. Non Galium ephedroides Willk. (1852).
For a careful discussion of the nomenclature and synonymy of this species, see Bacigalupo (1973).
Perennial, climbing (to 4 m.), tangled, dioecious herb (or woody at base); stems
conspicuous, stoutish, somewhat brittle, prominently 4-angled, more or less minutely
scabrous on angles, the internodes 4-9 times as long as leaves; leaves in fours,
inconspicuous, essentially glabrous or minutely scabrous, mostly lanceolate (to ovate
obovate), 5-10 mm. long, often reflexed at maturity, 1-nerved, the nerve prominent
below; glands in a subapical cluster; inflorescences diffuse, usually many-flowered;
flowering pedicels usually 1-4 mm. long but sometimes longer (to 8 mm.), those of
staminate flowers slender, those of pistillate flowers stouter, accrescent in fruit; corolla
rotate, divided nearly to base, green or cream-colored (or sometimes reddish?), gla
brous, the ovate lobes blunt at apex or shortly tipped; pistillate flowers with small
corollas, developed ovaries, protruding styles, and abortive stamens with short fila
ments; staminate flowers with larger corollas, fertile stamens with longer filaments,
abortive ovaries, and negligible styles; fruits fleshy, glabrous, pearly white when fresh,
brown and wrinkled when dried, far fewer than flowers.
Distribution: Argentina (Buenos Aires and Cordoba) northward to Brazil (Santa
Catarina), Paraguay (Guaira and Alto Paraná), and Bolivia (Chuquisaca), 40-1,450
m. (Figure 12), climbing on bushes, in ravines or at forest edges.
Despite the neat geographical separation (see map, Figure 12), there is no observa
ble morphological difference between the eastern and western populations. Recorded
elevations, of which there are too few, are somewhat higher for the western population
(500-1,450 m.) than for the eastern (40-600 m.).
Additional collections: ARGENTINA: Buenos Aires: Punta Piedras, Castellanos 24/1994 (f);
Punta del Indio, Cabrera 12318 (lp), 24277 (lp); Ensenada, Spegazzini 512 (f); Buenos Aires, Hooker (ny);
San Isidro, Burkart 4583 (f), Pérez Moreau 13581 (f); San Andrés, Lorentz & Hieronymus 263 (goet); Dept.
Zarate, Boelcke 4935 (ny), Cabrera 17237 (lp), Krapovickas 3539 (lil); Isla Martín García, Bacigalupo &
Crespo SI25995 (us), Boelcke 4993 (us), F abris 6071 (lp). Entre Ríos: Gualeguay, Lefebure (br), Lutz
78261 (r); Gualeguaychú, Burkart 22764 (F, us); Concepción del Uruguay, Burkart 18048 (f, us), Lorentz
438 (goet), 444 (goet), 522 (w), 1111 (goet, k, p, us, w); Concordia, Bur kart 22762 (f), Castellanos 31 /1363
(f). Corrientes: Dept. Esquina, Krapovickas et al. 27449 (ctes, mo); Dept. Mercedes, Schinini & Quarin
14357 (uc). Misiones: El Paraíso, Torres 77 (lp); Bonpland, Jorgensen 2133 (f), Lilliesköld (F, s); Santa
Ana, Rodriguez 529 (ba, f), Spegazzini 404 (f); Dept. San Ignacio, Denis 5758 (f), Schmidt 4995 (lil),
Schwarz 8021 (lil), 10527 (lil); Dept. Iguazú, Schwindt 2104 (lil); Tobuna, Pérez Moreau 21222 (f);
"Obraje Caburei", Montes 27450 (ny); Loreto, Burkart 1553 (f), Pérez Moreau 31 /2273 (f). Córdoba: San
Javier, Castellanos 11767 (f), 31255 (f), Conrad & Dietrich 2537 (mo); Dept. Calamuchita, Stuckert 24106
(cord), A. Hunziker 7147 (cord, uc), 7159 (cord, uc), Krapovickas & Cristóbal 14704 (c, uc); Sierra de
Cordoba, Fester (f), Stuckert 2123 (g), 2871 (cord, g, uc), 4276 (g), 17254 (g); Dept. Colón, Ariza 345
(cord, uc), 1643 (cord, uc), Kurtz 6593 (cord, uc), 12825 (cord, uc), Meyer 13459 (c, p), 22565 (lil),
O'Donell & Rodrigues 699 (gh), 879 (gh), Stuckert 23736 (cord), Subils et al 1483 (cord, uc), Villa
Carenze 2141 (lil), 3606 (lil); Cordoba, Escuela de Agricultura 14894 (f), Spegazzini 242 (bm); La Falda,
Holway 2023 (us); Dept. Cruz del Eje, A. Hunziker 7899 (cord, uc); Dept. Ischilín, Bacigalupo 9972 (c);
Dept. Punilla, Krapovickas 6626 (ctes, lil), Kuntze 7284a (cord, ny), Kurtz 6566 (cord, lp, ny), Meyer &
Sleumer 15653 (lil), Pedersen 136 (c, us), Stuckert 19271 (cord); Sierra Chica, near the Caleras, Hierony
mus (κ, ρ), Lorentz 435 (goet); Sierra Grande, Burkart 7431 (f), Río Primero (river), Hieronymus 183
(goet). Tucumán: Dept. Tafí, Castillon 7285 (f), Legname 9 (lil), Schreiter 1223 (f); Quebrada de Lulis,
Cristóbal 248 (lil); La Florida, Legname 4379 (lil); Dept. Monteros, Sparre 5222 (gb), Venturi 7032 (us).
Jujuy: Dept. San Pedro, Venturi 9547 (gh, us).
URUGUAY: Montevideo, Anderson 151 (bm, f, k); Dept. Maldonado, Lombardo (lil), Osten 7788 (f,
gh); Dept. Canelones, Giben 11 (κ), Osten 8722 (f); San José, H. M. Smith 96 (f, us); Santa Lucia,
Arechavaleta 50 (g), 111 (g), Fruchard, in 1867 (f, p), in 1872 (f, p), in 1874(f, p), Gibert 911 (goet, k), 1006
(goet, κ), Kuntze (ny), Safford (f, us); Dept. Lavalleja, Bartlett 20946 (f, gh, uc, us), Krapovickas &
Cristóbal 16196 (ctes); Dept. Florida, Gallinal et al. PE-4857 (us), Herter 1303c (υ, us); Dept. Colonia,
Doello Jurado 31356 (f), Fabris & Gebhard 60 (bm); Dept. Durazno, Osten 3844 (f); Mercedes, Osten 3074
(f); La Nigritas, Tweedie (k); Soriano, Gallinal et a!. PE-4884 (us); Banda Oriental, Saint-Hilaire 2130 (F, p);
Las Piedras, Fruchard 911 (f).

BRAZIL: Rio Grande do Sul: Gaudichaud 1084 (p); Neu-Württemberg, Bornmüller 132 (GH, u);
Ivoti, Rambo 42104 (lil). Santa Catarina: Liso, Klein 5732 (us); Lebon Regis, Klein 3125 (us); Xaxim,
Klein 5566 (us); Coquiro, Klein 5672 (us).
PARAGUAY: Guaira: Iturbe, Montes 12442 (lp); Isla Guaira, Montes 15989 (lil). Alto Paraná: Sta.
Teresa, Bertini 1660 (lil); "in regione fluminis", Fiebrig 5415 (gh, k, lil, us).
I have seen no indication that this species is other than perfectly dioecious. The
small seed set is not due to any visible imperfection in the pistillate apparatus.
17. Galium leptum Philippi in Anales Univ. Chile 85: 734. 1894. Type: CHILE:
Coquimbo: Ovalle, Rio Torca, "ad originum flum.", Geisse, in December, 1890
(sgo lectotype; isolectotype at sgo, photo seen; possible isolectotype at bm).
There are five specimens at sgo that answer to the type specifications, but the
collection dates given on the labels are different. One was supposedly collected in
November, 1889, the others in October, November, and December of 1890.
Although the two November specimens are a little larger, one of the December
specimens, which have more flowers, is here designated as lectotype. If the other
three specimens do indeed represent other collections, they cannot be considered
as isolectotypes. The specimen at bm is without date. Figure 19.

Very slender perennial, to 30 cm. high; stems strongly angled; minutely scabrous;
leaves in fours, 3-5 mm. long, linear, thick, shortly hispid on margins and upper
surface with short, stout, spreading hairs; glands none; flowers several on short
indeterminate branchlets; flowering pedicels and ultimate branchlets filiform; corolla
a little campanulate at base, the lobes obtuse or apiculate; fruit fleshy, on strongly
recurved pedicels.
Distribution: Chile, in west central Coquimbo, 1,800 m. (Figure 12).
Additional collection: CHILE: Coquimbo: Dept. La Serena, Cerro Tololo, Jiles 5114-a (conc).
Although scarcity of material prevents careful analysis, it seems probable that the
species is dioecious or polygamodioecious. Its closest affinityis with Galium latoramo
sum.
x 6; c, flower with probably

Figure 19. Galium leptum; a, flowering branch, χ 5/8; b, leaf in position,
sterile ovary, χ 12; d, fruit with one abortive mericarp, and flower bud, x 12. a & c from Geisse (type), b & d
from Jiles 5114-a.

ALLERTONIA
18. Galium aschenbornii Schauer in Linnaea 20: 701. 1847. Type: MEXICO:
Hidalgo: Zimapán, Aschenborn (je holotype).
Rubia debilis Kunth in Η. Β. Κ. Nova
Gen. et Sp. 3: 340. 1818. Type: ECUADOR: "Juxta Chillo et
Cataractam Ichubamba, Bonpland (holotype b, photo seen; isotype at ρ). Non Galium debile
Hoffmansegg & Link (1813-1820) nec Desv. (1818).
Galium fraserii Wernham in J. Bot. 50:244.1912. Type: ECUADOR: Fraser (bm holotype; isotype at g).
For full synonymy, with discussion, see Dempster (1973a, 1973c); for illustration see Dempster (1973a).
Perennial, polygamous, trailing or climbing plant, with long, slender, wiry stems
arising singly or in clumps from a rootstock; stems glabrous or usually more or less
scabrous on angles, the internodes 1-5 times as long as leaves; leaves in fours,
ovate-lanceolate or elliptic, 4-8 (-15) mm. long, 1-nerved or often obscurely 3-nerved,
apiculate, narrowed to a short petiole, the surfaces usually glabrous, the callose
margins often with aculeate hairs; glands few, distributed and in a subapical cluster;
inflorescences on short, leafy, determinate branches having 1-several or sometimes
many flowers; flowers staminate, pistillate, or perfect; corolla glabrous, rotate, the
lobes prolonged at apex; fruits fleshy, glabrous, black and wrinkled when mature.
Distribution: Ecuador and western Colombia (Figure 4), through Central
America to Jalisco and San Luis Potosí, Mexico; 2,650-2,750 m., on moist slopes or
streambanks.
Additional collections: ECUADOR: Pichincha: Base of Pichincha, Benoist251l (p). COLOMBIA:

Valle del Cauca: Cordillera Central, Loma de Barragán, Cuatrecasas 20726 (f). Antioquía: Boquerón,
Daniel 441 (f).
There seems no adequate reason to recognize a taxonomic separation between the
Central American plants and those of northwestern South America. The few South
American specimens observed, however, have smaller and broader leaves than do
many of the Central American and Mexican specimens.
19. Galium azuayicum Dempster, sp. nov. Plate 1; Figure 20.

Planta perennis gracilis scandens 1-3 m. alta, ramis pilis brevibus retrorsis scabri
dis, internodis quam foliis 2-8-plo longioribus; foliis in quoque verticillo quatuor
minus quam 3 mm. longis saepe remotis ovatis, subtus glabris, supra pilis antrorsis
longe pubescentibus; glandulis apicem versus fasciculatis; floribus pedicellatis in
extremitatibus vel in axillis ramulorum brevium foliiferorum lateralium solitariis,
pedicellis quam ovariis fructibusve 0.5-3-plo longioribus; corolla glabra rotata autem
non late patenti albida vel rosea tincta, lobis obtusis vel tantum acutis; fructibus siccis
tuberculatis.
Type: ECUADOR: Azuay: At kilometer 91 on Pan American Highway north of
Loja (3° 25' S., 79° 10' W.), Holm-Nielsen, Jeppesen, Lfljtnant, & 0llgaard 5114 (s
holotype).
Distribution: The type is the only specimen seen; scandent in dry scrub at 2,900 m.
(Figure 4).
The new species is most closely related to Galium pumilio, of much higher alti
tudes. It differs not only in its strikingly distinct habit, but also in having scabrous
stems, leaves that are pilose on the upper surface, 4-lobed corollas, and tuberculate
fruits.
The wall of the mericarps, when softened, fragments into individual cells or groups
of cells. The cells have thin walls and are filled with a granular material not yet
investigated, perhaps cellulose.
The appearance of hairs on corollas and anthers in the type material is owing to the
presence of a fungus, presumably of the Dematiaceae (I. Tavares, personal communi
cation).

HniAfltw L'nivihuty
JfttON o» Caiiimnia
HOLOTT
PB
Gallun azua;
Lauiamav
T. Dmnru ly8?
• OTAMlSK INSTITUT . AARHUS UNIVERSITET

tAAOIAartiua
(NtrtMr JurUndteum Ctonmark)
Flora of Ecuador
colteci««!
1973
byL " ··**n.S JoppeMn.Β lo|tn«nt
C Β Ollqeard
AZUAV
Km9» Amanean Ν utLot·Dryscruti1-3mhigh«li 2900m.
Highway
ίΤβ-10W y 25'S)
no. Ί14
;c«incîent in scrub. l lovera white.
Plate 1. Galium azuayicum; type.
20. Galium tricornutum Dandy in Watsonia 4: 47. 1957. Type European.

Galium tricorne Stokes in With. Bot. Arr. Brit. PI. ed. 2. 1: 153, nom. illeg. 1787. (See Dandy, loe cit.)
Coarse sprawling annual; stems retrorsely scabrous; leaves 6-9 at a node, oblan
ceolate, strongly scabrous on margins; flowers pedicellate, usually 3 on a peduncle, the
pedicels strongly reflexed in fruit; corolla rotate, white; fruits dry, tuberculate, the
mericarps nearly spherical.

ALLERTONIA
Figure 20. Galium azuayicum; a, branchlet, χ 5/8; b, node, showing upper surface of leaves, * 6;c, node,
x 6; d, fruit, x 12; e, flower, χ 12. All from Holm-Nielsen
showing lower surface of leaves, et al. 5114 (type).
Distribution: Locally introduced in cultivated fields.
Collections seen: ARGENTINA: Chaco: Col. Benitez,ßchulz 503, in 1931 (f).

BRAZIL: Pernambuco: (Probably Recife), Gardner (e).
CHILE: Ñuble: Portezuelo-Chillán, Matthei 27110, in 1959 (CONC).
21. Galium divaricatum Pourret ex Lam. Encycl. Méth. Bot. 2: 580. 1788. Type
European.
Galium tenellum Clos in Gay, Fl. Chil. 3: 188. 1856. Type: CHILE: Gay (p holotype).
Galium aridicola Briq. in Ann. Conserv. Jard. Bot. Genève li & 12:193. 1908. Based on G. tenellum Clos;
non Jordan (1846). "Double emploi accidentel du même nom pour deux espèces complètement
différentes . . ."
Diminutive erect annual similar to G. parisiense L.; stems and branches slender to
filiform; leaves in whorls of 6, 4-6 mm. long, the whorls remote; flowers perfect,
whitish, the peduncles and pedicels filiform; fruits dry, glabrous.
Distribution: European introduction, found by Gay somewhere in Chile.
22. Galium ascendens Willd. ex Spreng. Syst. Veg. 1:390. 1824. Type: South America,
Bonpland (b holotype, microfiche seen). Figures 21, 24c.
Rubia scabra Kunth in Η. Β. Κ. Nova

Gen. et Sp. 3:340. 1818. Type: COLOMBIA: Between Cartagoand
Ibagué, Bonpland (b holotype, photo seen; fragment at f). Non Galium scabrum L. (1753).
Galium scabrum Wedd. Chloris Andina, 39. 1857; non Galium scabrum L. (1753).
Galium cuatrecasasii Standley in Pubi. Field Mus. Nat. Hist., Bot. Ser. 22:113. 1940. Type: COLOMBIA:
Cundinamarca: Sabana de Bogotá, Cuatrecasas 468 (us holotype; isotypes at f, p).
Slender, polygamodioecious, herbaceous perennial, decumbent or usually erect,

the stems 9-30 (-60) cm. long; stems scabrous on angles, the internodes 1.2-5 times as
long as leaves; leaves 1-nerved, mostly 6 per node, 4-14 mm. long, narrowly to broadly
oblanceolate or elliptic, scabrous on margins, the apices obtuse and without apical
hair; flowers usually few, on leafy lateral branchlets, the pedicels as long as flowers or
fruits or sometimes longer; corolla white or lilac-tinged, usually somewhat cupped at
base, the lobes ovate or triangular, obtuse or acute, somewhat urceolately spreading;
fruits of 2 hard, spherical, black mericarps, or often 1 aborted, wrinkled on surface
when dry, in clusters of 1-5 on short pedicels at ends of long peduncles.

Figure 21. Galium ascenderti; a, branch, x 5/8; b, node with leaves, x 6; c, leaf showing venation, x 6;d,
staminate flower, x 12; e, hermaphrodite flower, x 12; f, same flower seen from above, x 12;
g, fruiting
branchlet, x 3; h, fruit, x 12. a from Triana 1601, b & e from Cuatrecasas & Garcia Barriga 9902, c from Killip
34122, d from Antonio Camilo 4a, f from Haught 5932, g & h from COL 18524.
Distribution: Colombia: Cordillera Oriental, from Cundinamarca to Santander,

2,400-3,900 m. (Figure 4), in wet or at least moist places, páramos, savannas,
streamsides, and lake shores. Puff (unpublished manuscript) also attributes the species
to Ecuador (Pichincha).
Additional collections: COLOMBIA: Cundinamarca: Bogotá: Antonio Camilo 4a (f), Apollinaire
(s, ven), Ariste-Joseph A268 (f), Guevara Amórtegui 219 (f, us), Goudot (κ), Triana 3100 (f, p); Sabana de
Bogotá, Schiefer 698 (gh, uc), Uribe 332 (col); Las Cruces, Penneil 2161 (f, ny); San Cristóbal, Pennell
2310( F, gh, κ, mo, ny); Facatativá, André 579 (κ); Tequendama, Ariste-Joseph Α149(ΐ), Triana3101 (p);
Laguna Negra, Cuatrecasas & Jaramillo 25873 (ven); Represa de Neusa, Cleef & Jaramillo 4177 (u, uc);
Páramo de Chisacá, Barclay & Juajibioy 6149 (ny); Sesquile, Haught 5932 (f, lil); Páramo de Sumapaz,

246 ALLERTONIA 3:3
Cleef2596 (υ), 5188 (υ), 6555 (u); Laguna de Fúquene, collector unknown (col 18524, F 1092384); Páramo
de Guasca, Killip 34122 (bm, f, gh, lil, mo, ny, p, s, uc); Cerro de Suba, Duque-Jaramillo 2798 (col).
Boyacá: Páramo de Güina, Cuatrecasas & García Barriga 9760 (us); between Belén and Susancón, Barclay
& Juajibioy 7626 (ny). Santander: Páramo del Almorzadero, Cuatrecasas & García Barriga 9902 (f).
Department?: Verjon, Ariste-Joseph Λ159 (f, us); New Granada, Triana 1598 (bm, f, ny, us), 1601 (f, k,
ny).
Although this species has been generally called Galium trianae Wernham, this is
clearly an error, since the original description of G. trianae states "fructu . . . pilis
uncinulatis hispidissimo," corresponding to a Τriana specimen at bm that I equate with
G. canescens Kunth. For further discussion see under Galium canescens (Dempster,
1981, p. 414).
Galium ascendens is very closely related to G. magellanicum, of the southern
Andes. There is a geographic gap of 32 degrees of latitude between them, and although
the two species are not in every instance clearly distinguishable on morphological
grounds, there are some differences. Galium ascendens is, for the most part, a slender
plant, with small leaves and relatively long internodes, whereas in G. magellanicum the
plants are either low, with slender stems, relatively large leaves, and short internodes,
or they are tall with stoutish stems and relatively long leaves.
Leaves of Galium magellanicum, when cleared and stained, are seen to be crypti
cally 3-nerved, whereas those that I have observed of G. ascendens show only remnants
of the two slender lateral nerves, nearly always interrupted above the leaf base.
23. Galium magellanicum Hook. f. FI. Antarct. 302 (bis). 1846. Type: CHILE:
Magallanes: Cape Negro, Darwin (κ holotype). Figures 22, 24d.
Galium chonoense Hook. f. Fl. Antarct. 303 (bis). 1846. Type: CHILE: Chiloé: Chonos Archipelago,
Darwin (κ holotype).
Galium nigricans Clos in Gay, Fl. Chil. 3: 188. 1848. Type: CHILE: Valdivia: Gay 132 (p holotype not
seen; photo and fragment at f; isotype at p).
Galium valdivianum Philippi in Linnaea 28:655. 1856. Type: CHILE: Valdivia: San Juan, Philippi (bm
lectotype; isolectotype photo and fragment from β at f).
Galium forsteri Philippi in Anales Univ. Chile 85: 735. 1894. Type: CHILE: Strait of Magellan, Philippi
(sgo holotype, photo and fragment seen).
Galium chonoense var. valdivianum Reiche in Anales Univ. Chile X06: 990. 1900.
Slender, polygamodioecious, herbaceous perennial 7-60 cm. tall, matted or tufted

from fibrous roots; stems weak or strong, minutely scabrous, the internodes mostly
3-5 as long as leaves;
times leaves 5-7 (-8) to a node, apparently 1-nerved but obscurely
3-nerved or nearly so, mostly 6-17 mm. long, linear or oblanceolate to narrowly
obovate, scabrous on margins (and sometimes midribs), tapered at base, round or
obtuse at apex, without a terminal hair; flowers 3-many in more or less proliferous
cymules; corolla white, rotate, or slightly cupped at base, the lobes usually 4, rarely 3,
ovate, obtuse; fruits consisting of 2 hard, spherical, black mericarps.
Distribution: Chile and western Argentina from Magallanes and Tierra del
Fuego to Valparaiso and southern Neuquén, with a real or apparent gap between 50°
40' S. and 45° 20'S., 3-1,200 m. (Figure 14), along river banks and lake shores, in bogs
and moist meadows.
Additional collections: ARGENTINA: Santa Cruz: Patagonia, 50° 53' S., Moreno 765 (ny); Lago
Argentino, Dúsén 5680 (s), Hauthal 8842 (f). Chubut: Río Senguer, Spegazzini ? 452 (f); Lago Fontana,
Kozlowsky (f, lp); Río Pico, Roth (f, lp); Carrenleofu, Iliin 469 (f, lp); Corcovado, Iliin 478 (f, lp). Rio
Negro or Neuquén: Near Nahuel Huapi, Buchtien s. n. (s), 49 (us), Castellanos 22091 (f), Corte 278 (lp),
Fabris & Solbrig 550 (lp), Job 2455 (lp), Ljungner 700 (gb), 989 (gb), Pérez Moreau 35512 (f), 48765 (f),
Roth (lp). Neuquén: Near Nahuel Huapi, Cabrera 6015 (lp), Cabrera & Job 176 (f, lp), Fabris & Solbrig
272 (lp), 283 (lp), 1135 (lp), Pérez Moreau 49633 (f); Lake Quillén, Villas (m); Parque Nacional Lanín,
Eskuche & Klein 1443-15 (ctes).

CHILE: Magallanes: Ortega 937 (sgo); Isla Observatorio, Castellanos 12968 p. p. (specimen perhaps
adulterated) (f); Isla Grande between San Sebastián and Bahía Inútil, Kunze 12351 (cord); Cabo Negro,
Strait of Magellan, Darwin (cge), Hatcher (ny); Port Famine, Andersson 43 (bm, k), Cunningham (k); San
dy Point, Lechler 1206 (s), 1206a (goet), 1207 (s), 1208 (br, g, goet, k, p, s); Strait of Magellan, Mc fVhinnie
(k); Isla Riesco, Pfister & Ricardi 11924 (coNc); Gruta del Milodón, Pfister & Ricardi 12050 (coNc), 12492
(conc). Aisén: Valley of Río Aisén, Dusén (s); Coihaique, Andreas 552 (u); Puyuhuapi, Schwabe 55 (CONC,
ny). Aisén or Chiloé: Chonos Archipelago, Darwin (cge). Chiloé: Reed (κ), King Expedition (bm, k, w);
Dept. Castro, Marticorena et al. 131 (conc), Weldt-Rodriguez 742/37 (conc); Cucao, Borchers (bm);
Chepu, Godley 326 (κ, sgo); Talcahue (a reef), Reed (sgo). Llanquihue: Maullin, Vidal (sgo); Nueva
Braunau, Sparre & Smith 318 (conc); Rio Manso, Reiche (sgo); Puntiagudo, Andreas 266 (u); Peulla,
Pfister 13540 (conc). Osorno: Lake Llanquihue, Elwes (k); Volcán Osorno, Reed (κ). Valdivia: Bridges
(bm, ny), 648 (ε, κ), Buchtien (us), Gunckel (s), Junge 561 (conc), Krause 928b (sgo), Ochsenius (fragment)
(f), Philippi (goet, hal, k, s, sgo, uc, us), Philippi & Grisebach (p); Valdivia to Puerto Montt, Calvert (bm);
Corral, Krause (sgo), Philippi (hal, κ), Reed 140 (bm), Sparre & Smith 398 (conc); Cordillera de Raneo,
Philippi 928e (sgo); Magedehue, Hollermayer 553 (m), Werdermann 1377 (leg. Hollermayer) (f, g, gh, s, u);
Croconn, Lechler 742 (gh, goet, k, p); Cordillera Chaihuin, Gunckel 3049 (gh); Quinchilco, Hollermayer
138 (uc). Cautín: Budi, Philippi 928d (sgo); Villarrica, Beh η & Pfister 5998 (conc); Paso de Malalco,
Figure 22. Galium magellanicum; a, branch, northern type, x 5/8; b, branch, southern type, * 5/8; c,
leaf, northern type, x 6; d, leaf, southern type, x 6; e, staminate flower, x 12; f, 3-lobed corolla, x 12; g,
x 12; h, fruit, x 12. a from Behn &
pistillate flower, Pfister 5998, b from Dúsén 5680, c & e from Landbeck
928c, d from Buchtien 49, f from Fabris & Solbrig 550, g from Ricardi 9257, h from Pérez Moreau 49633.

248 ALLERTONIA 3:3
collector? (m). Malleco: Palobotado, Philippit (sgo); Laguna Malleco, Behn 7173 (conc); Nahuelbuta,
Eyerdam 10268 (f, ny, uc); Marimenuco, Pfister 7384 (conc); Lonquimay, Pfister 7254 (conc), 7955
(conc), 8041 (conc). Arauco: Contulmo, Ricardi 9257 (conc). Arauco or Malleco: Nahuelbuta,
Volckmann (sgo). Bio-Bio: Los Angeles, Pöppig 43 (Diar. 809) (bm, br, f, g, hal, p); Negrete, Philippic
(sgo). Concepción: Coronel, Ochsenius (br, goet); Isla Santa Maria, Eights (us). Curicó: Llico, Landbeck
928c (sgo), Philippil (sgo, κ). Valparaíso, King (κ).
Galium magellanicum belongs to the G. trifidum complex, which is worldwide and
notoriously difficult to treat taxonomically. As here broadly construed, it includes low
plants with weak stems and few flowers, as well as tall plants with stout, strong stems
and many flowers. Leaves are highly variable in shape and size. There are some elinal
tendencies, mostly quantitative, but the north-south trend is very inconsistent, so that
it is impossible to know from what latitude a specimen derives. Forms that are
common in the north (i.e., tall, with sturdy stems and many flowers) are not to be found
far south, but the reverse is not true: scattered through the northern part of the range
are plants that could easily be supposed to be from Magallanes.
There is a geographic disjunction among the available collections between 45° 30'
S. and 50° 30' S. It is therefore tempting to call the southern plants Galium magellani
cum and the northern plants G. chonoense, or at least to consider them as subspecifi
cally different. Aside, however, from the fact that the geographic disjunction may not
be real, there is unfortunately no consistent morphological difference between the
northern and southern plants that would justify such a classification. Similarly, it is
tempting to differentiate the Chilean plants from the trans-Andean plants of Argen
tina, but again this cannot be justified on morphological grounds. It seems preferable,
therefore, to keep all these varying plants as a single species, rather than to divide them
into several morphological varieties having neither clear differences nor geographic
unity.
Plants of the south are generally lower, with fewer flowers and broader leaves,
whereas northern plants are often tall, with strong stems, a greater number of flowers,
and sometimes very narrow leaves. Such differences could be partly based on ecologi
cal conditions, with high altitude substituting in the north for high latitude in the
south. Unfortunately, few altitudes are given on the many specimens at hand, and few
localities are sufficiently precise so that altitudinal figures could be derived.
The taxonomic separation of Galium antarcticum and G. magellanicum on one
hand and of G. magellanicum and G. ascendens on the other is probably already
somewhat artificial. Further to subdivide G. magellanicum (i.e., to accept G. cho
noense, G. nigricans, and G. valdivianum) would be to make the situation even worse.
It is a far cry from "typical" G. antarcticum to "typical" G. ascendens, but when one
includes G. magellanicum as a bridge between them, sharp morphological discontinui
ties are hard to find. We can arbitrarily state that G. antarcticum has solitary flowers
and four leaves to a node, whereas G. magellanicum has flowers in clusters and more
than four leaves to a node. Nevertheless, in southern Magallanes, and even in Tierra
del Fuego, they are sometimes mixed, and one suspects that they interbreed, at least to
some extent.
The geographic gap between Galium magellanicum, of Chile and Argentina, and
G. ascendens, of Colombia, seems ample justification for separating them taxonomi
cally, although some specimens of the latter would be hard to diagnose as such without
knowledge of their provenance.
Ehrendorfer, Puff, Bacigalupo, and others have used the name Galium chonoense

extensively in annotating specimens, limiting its use to specimens from north of Aisén.
This name, typified as it is by a collection from the middle of the geographic range of
the species, and therefore representative of moderation in morphological characters,
might seem a better choice for the taxon than G. magellanicum. Two considerations
prevail, however, in the selection of the latter name. Not only does it have page priority
but, unlike G. chonoense, it is clear of nomenclatural confusion. Unfortunately, Clos
(1847) published a Galium chonosense that I equate with G. aparine (Dempster, 1981,
p. 418).
Plate 2. Galium surinamense; type.

ALLERTONIA
24. Galium palustre L. Sp. Pl. 105. 1753. Type European.

Aquatic perennial; stems many, slender, flexible, essentially glabrous; leaves 4-6 at
a node, 10-30 mm. long, linear to narrowly oblanceolate, glabrous or minutely and
antrorsely scabrous on margins, round or obtuse at apex; flowers numerous in
proliferated cymules at ends of slender peduncles, or inflorescence diffuse; corolla
white, campanulate at base, the lobes 4, ovate or triangular, obtuse; fruit of 2 spherical,
hard mericarps.
Distribution: Argentina: Buenos Aires. Locally introduced, 10-50 m., aquatic.
Earliest collection in 1939.
Collections seen: ARGENTINA: Buenos Aires: Punta Lara, Burkart 22402 (ny, u), Castellanos
33246 (ba, f), Dawson 1015 (lp), 1137 (f, lil, mo, u, us), Fabris 2462 (c), 3207 (ny), Gamerro 1027 (s);
between Villa Elisa and Punta Lara, Cabrera 9823 (gh); between La Plata and Ensenada, Cabrera 16416
(lp); between Rio Santiago and Palo Blanco, Rodriguez 604 (lil).
Figure 23. Galium surinamense; a, portion of inflorescence, with one node and leaves,x 3; b, small leaf,
χ 6; c, portion of large leaf, lower surface,
showing inrolled margins (from soaked material), x 6; d, portion of
inflorescence, showing fruit, one mericarp, vacated receptacle, and geniculate character of pedicel, χ 12. All
from Wright (type).
25. Galium surinamense Dempster, sp. nov. Plate 2; Figure 23.

Planta perennis humilis ad 14 cm. alta, caulium parte inferiore lignosa autem
gracili; caulibus pilis brevibus varie curvatis plus minusve lanuginosis; foliis linearibus
in quoque verticillo sex 5-10 mm. longis supra scabridis margine recurvatis; inflores
centia solummodo bracteata multiflora, ramis ramulisque et pedunculis filiformibus
subaequalibus ca. 2 mm. longis retrorse divaricatis caulium extremitatibus nodum
subtilem facientibus; fructibus glabris ca. 1.5 mm. longis; floribus non visis.
Type: SURINAM: Wright (w holotype and isotype).
The only available collection lacks flowers, but the distinctive tangled inflores
cence, the linear leaves in sixes, and the fleshy fruits set it well apart from other species.
No data are given beyond the collector's surname and the country of origin. The
specimens were acquired by w in 1889 from the collection of Reichenbachyi/iMS. There
is clear quartz sand clinging to the lower portion of the largest specimen, suggesting a
beach habitat.
26. Galium mollugo L. Sp. PL 107. 1753. Type European.

Perennial from subterranean stolons; stems erect or procumbent, 30-150 cm. long,
glabrous; leaves 6-8 per node, 10-14 mm. long, lanceolate-oblanceolate, scabrous on
margins; inflorescences diffuse, many-flowered, terminal and lateral, merely bracteate,

the flowers perfect; corolla rotate, white, the lobes ovate, acute; fruits dry, glabrous.
Distribution: European introduction, collected in Argentina and Uruguay.
Collections seen: argentina: Buenos Aires: Bettfreund21912, in 1910(g); Stuckert 22314, in 1911
(cord); Parodi 8769, in 1928 (f, gh).
uruguay: Rivera, Herter 99206, in 1923 (ny).
Figure 24. Stem cross sections of several species, somewhat stylized, and not drawn to scale, a, Galium
mandonii from Buchtien 3270; b, G. latoramosum from Pérez Moreau 31/2273; c, G. ascendens from
Cuatrecasas & García Barriga 9902; d, G. magellanicum from Vidal; e, G. suffrulicosum from Morrison
16751; f, G. araucanum from Philippi (Talca).

252 ALLERTONIA 3:3
27. Galium saxatile L. Sp. PI. 106. 1753. Type European.

Low perennial 15-20 cm. high, forming large patches by rooting at nodes; leaves
mostly 6 or 7 at nodes, 4-8 mm. long, oblanceolate-obovate, with prolonged hyaline
tip, more or less scabrous on margins; inflorescences terminal and lateral on short
branches, merely bracteose; corolla white, rotate, the lobes ovate, merely acute; fruits
glabrous, dry.
Distribution: European introduction, collected in Falkland Islands.
Specimens seen: FALKLAND ISLANDS: Carcass Island, West Falklands, Hansen in 1950 (bm); San
Carlos, East Falkland, Moore 648, in 1964 (gh, k, lp, s, us).
Excluded species
Galium brasiliense Veil. Fl. Flum. 43. 1829, Icon. 1: t. 109. 1831. Type locality:
Brazil. This species is represented only by a brief description and poor illustration.
From J. P. P. Carauta, who has researched the matter (Carauta, 1969, 1973), I
have the following personal communication (my translation): "The holotypes of
Bro. Vellozo, for the most part, were lost, therefore we consider as lectotypes the
prints of the iconographie volumes."

Galium octophyllum Larrañaga, Escritos D. A. Larrañaga 2:61. 1923. Type localty:
Uruguay. The Larrañaga names were published after the author's death, with
inadequate descriptions, no illustrations, and apparently no types.
Disposition of names (accepted epithets are in boldface type)

The following list is intended to include all Galium names used for South American
plants, including those previously listed by me (Dempster, 1980, 1981), as well as
selected names from Rubia and Relbunium.
Galium albicans Wedd. = Relbunium (R. hypocarpium (L.) Hemsl. subsp. hypocar
pium, fide Ehrendorfer, 1955)
Galium andicolum Krause = G. weberbaueri
Galium andinum Philippi = G. suffruticosum
Galium antarcticum Hook. f.
Galium aparine L.
Galium charoides Rusby in Phytologia 1: 80. 1934. Type (not seen): BOLIVIA: Pongo, Tate 234. Syno
nym inadvertently omitted from Dempster (1981, p. 418).
Galium repugnans Kunze, a name unpublished except as referred to by Ehrendorfer, 1955, p. 537. Men
tioned obliquely in Dempster (1981, p. 420, lines 15, 16)
Galium aparine var. microphyllum Clos = G. aparine
Galium aparine var. pseudoaparine = G.
Speg. aparine
Galium apiculatum Roemer & Schultes = probably Relbunium (R. croceum (Ruiz &
Pavón) Κ. Schum. subsp. croceum, fide Ehrendorfer, 1955)
Galium apricum Veli. = Relbunium (questionably R. megapotamicum subsp. campo
rum (Pohl ex DC.) Ehrend., fide Ehrendorfer, 1955)
Galium araucanum Philippi
Galium aridicola Briq. -G. divaricatum
Galium ascendens Willd. ex Spreng.
Galium aschenbornii Schauer
Galium atherodes Spreng. = Relbunium (R. atherodes (Spreng.) Κ. Schum., fide
Ehrendorfer, 1955)

Galium australe Reiche = G.

aparine
Galium azuayicum Dempster, sp. nov.
Galium benthamianum Walp. = Relbunium (R. croceum subsp. involucratum (H. B.
K.) Ehrend., fide Ehrendorfer, 1955)
Galium bigeminum Griseb. = Relbunium (R. bigeminum (Griseb.) K. Schum. fide
Ehrendorfer, 1955)
Galium boyacanum Dempster
Galium brasiliense Veil. See under excluded species in the present paper.
Galium brasiliense Wawra = Relbunium (R. indecorum (Cham. & Schlechtendal)
Ehrend., fide Ehrendorfer, 1955)
Galium brevifolium Philippi = G. eriocarpum
Galium Pohl ex DC. = Relbunium
camporum (R. megapotamicum subsp. camporum
(Pohl ex DC.) Ehrend., fide Ehrendorfer, 1955)
Galium canescens Kunth
Galium canescens f. alstonii = G. orizabense

Steyermark
Galium Kunth = G. mexicanum
caripense
Galium chaetophorum Griseb. = G. richardianum subsp. richardianum
Galium var. Kuntze = G. richardianum
chaetophorum prostratum subsp. richardia
num
Galium var. strictum Kuntze = G. richardianum

chaetophorum subsp. richardianum
Galium chamissonis Hook. & Arn. = G. trichocarpum
Galium = G. latoramosum
chapi Wedd.
Galium charoides Rusby = G. aparine (q. v. in this list)
Galium chilense Endl. ex Clos = G. suffruticosum
Galium chilense Hook. f. = G. aparine
Galium chonoense Hook. f. = G. magellanicum
Galium chonoense var. valdivianum Reiche = G. magellanicum
Galium chonosense Clos = G. aparine
Galium ciliatum Larrañaga = Relbunium (questionably R. megapotamicum subsp.
camporum (Pohl ex DC.) Ehrend., fide Ehrendorfer, 1955)
Galium ciliatum Ruiz & Pavon = Relbunium croceum involucratum
(R. subsp. (H. B.

Galium ciliatum var. normale Kuntze = Relbunium
(R. croceum subsp. involucratum
(H. B. K.) Ehrend., fide Ehrendorfer, 1955)

Galium ciliatum var. pusillum Kuntze = G. richardianum subsp. richardianum
Galium ciliatum var. richardianum Kuntze = G. richardianum subsp. richardianum
Galium ciliatum var. weddellianum Kuntze = Relbunium
form, fide (intermediate
Ehrendorfer, 1955)
Galium closianum Briq. = G. werdermannii
Galium cochabambense = G.
Rusby plumosum subsp. plumosum
Galium comberi Dempster
Galium corymbosum Ruiz & Pavon = Relbunium (questionably R. croceum subsp.
involucratum (H. B. K.) Ehrend., fide Ehrendorfer, 1955)
Galium cotinoides Cham. & Schlechtendal
Galium croceum Ruiz & Pavon = Relbunium (R. croceum (Ruiz & Pavon) K. Schum.
subsp. croceum, fide Ehrendorfer, 1955)
Galium cuatrecasasii Standley = G. ascendens

254 ALLERTONIA 3:3
Galium dasycarpon Kunze = G. philippianum

Galium debile Banks & Solander ex Hook, f., nom. nud. pro syn. = G. antarcticum
Galium diffusoramosum Dempster & Ehrendorfer
Galium diffusum Clos = G. diffusoramosum
Galium diphyllum (K. Schum.) Dempster, comb. nov.
Galium divaricatimi Pourret ex Lam.
Galium ecuadoricum Dempster
Galium = G. latoramosum
ephedrodes (sic) Kuntze
Galium equisetoides (Cham. & Schlechtendal) Standley
Galium ericoides Lam. = Relbunium (R. ericoides (Lam.) K. Schum., fide Ehren
dorfer, 1955)
Galium ericoides var. atherodes Cham. & Schlechtendal = Relbunium (R. atherodes
(Spreng.) Κ. Schum., fide Ehrendorfer, 1955)
Galium ericoides var. intermedium Cham. & Schlechtendal = Relbunium (R. ericoides
(Lam.) K. Schum., fide Ehrendorfer, 1955)
Galium eriocarpum Bartling ex DC.
Galium var. Kuntze = G.
eriocarpum leiocarpum eriocarpum
Galium var. normale Kuntze = G.
eriocarpum eriocarpum
Galium ferrugineum Krause
Galium ferrugineum Krause subsp. ferrugineum
Galium ferrugineum subsp. jamesonii Dempster
Galium = Relbunium (R. atherodes
filiforme Larrañaga (Spreng.) Κ. Schum., fide
Ehrendorfer, 1955)
Galium flossdorfii Hicken = G. plumosum subsp. flossdorfii
Galium fluminense Veil. = Relbunium (R. hypocarpium subsp. fluminense (Veil.)
Galium forsten Philippi = G. magellanicum
Galium fraseri Wernham = G. aschenbornii
Galium fuegianum Hook. f.
Galium galapagoense Wiggins
Galium gardneri Walp. = Relbunium (R. noxium (St.-Hil.) K. Schum., fide Ehren
dorfer, 1955)
Galium geminum Larrañaga = Relbunium (R. hirtum (Lam.) K. Schum., fide Ehren
dorfer, 1955)
Galium gilliesii Hook. & Arn.
Galium gilliesii Hook. & Arn. subsp. gilliesii
Galium gilliesii subsp. telanthos (Philippi) Dempster
Galium gilliesii var. β Hook. = G. gilliesii subsp. gilliesii
Galium gracilicaule Bacigalupo & Ehrendorfer
Galium haenkeanum Endl. = G. richardianum subsp. haenkeanum
Galium hirsutum Ruiz & Pavon = Relbunium (R. hirsutum K. Schum., fide Ehren
dorfer, 1955)
Galium hirtum Lam. = Relbunium (R. hirtum (Lam.) K. Schum., fide Ehrendorfer,
1955)
Galium huancavelicum Dempster
Galium humile Cham. & Schlechtendal
Galium hypnoides Clos = G. werdermannii

Galium hypnoides var. inconspicuum Reiche = G. inconspicuum

Galium hypocarpium Clos = Relbunium (R. hypocarpium (L.) Hemsl. subsp. hypo
carpium, fide Ehrendorfer, 1955)
Galium inconspicuum Philippi
Galium intricatum Endl. = G. latoramosum
Galium involucratum H. B. K. = Relbunium croceum involucratum B.

(R. subsp. (H.
Galium killipii Dempster & Ehrendorfer
Galium kunthii Wedd. = Relbunium (R. hirsutum K. Schum., fide Ehrendorfer, 1955)
Galium lanatum Kuntze = G. gilliesii subsp. gilliesii
Galium Ruiz & Pavon = G. obovatum
lappaceum
Galium Wernham = G.
larecajense aparine
Galium lasiocarpum = G.
Philippi trichocarpum
Galium latoramosum Clos
Galium laxum Philippi = G. trichocarpum

Galium leptum Philippi
Galium leucocarpum DC. = probably Relbunium (R. croceum (Ruiz & Pavon) K.
Schum. subsp. croceum, fide Ehrendorfer, 1955)
Galium lilloi Hicken
Galium límense Standley = G. weberbaueri
Galium magellanicum Hook. f.
Galium mandonii Britton
Galium masafueranum Skottsberg
Galium = Relbunium
megapotamicum Spreng. (R. megapotamicum Spreng.)
Ehrend, subsp. megapotamicum, fide Ehrendorfer, 1955)
Galium mexicanum Kunth
Galium mollugo L.
Galium monanthum ex DC. = Relbunium croceum involucratum
Bartling (R. subsp.

montanum ex Steud. = Relbunium croceum involucratum
Galium Bartling (R. subsp.
Galium mucronatum Ruiz & Pavon = Relbunium croceum &
probably (R. (Ruiz
Pavon) K. Schum. subsp. croceum, fide Ehrendorfer, 1955)

Galium murale (L.) Allioni
Galium mycrosphyllum Larrañaga = Relbunium (R. ericoides (Lam.) K. Schum., fide
Ehrendorfer, 1955)
Galium Clos = G.
nigricans magellanicum
Galium obovatum Kunth
Galium octophyllum Larrañaga. See under excluded species in the present paper.
Galium orinocense Humb. ex Spreng. = Relbunium (R. hypocarpium (L.) Hemsl.
subsp. hypocarpium, fide Ehrendorfer, 1955)
Galium orizabense Hemsl.
Galium ovale Ruiz & Pavon = Relbunium (R. hypocarpium (L.) Hemsl. subsp.
hypocarpium, fide Ehrendorfer, 1955)
Galium ovalleanum Philippi
Galium palustre L.

256 ALLERTONIA 3:3
Galium paratyense Veil, (sterile) = Relbunium noxium (St.-Hil.) K. Schum., fide

Ehrendorfer, 1955
Galium parisiense L.
Galium Kuntze = G. richardianum richardianum
patagonicum subsp.
Galiumpauciflorum Willd. ex DC., nom. nud. pro syn. = Relbunium (R. hypocarpium
(L.) Hemsl. subsp. hypocarpium, fide Ehrendorfer, 1955)
Galium peruvianum Dempster & Ehrendorfer
Galium = G.
peteroanum Philippi suffruticosum
Galium philippianum Dempster
Galium = G.
philippii Briq. eriocarpum
= G. mexicanum
Galium piliferum Kunth
Galium plumosum Rusby
Galium plumosum Rusby subsp. plumosum
Galium plumosum subsp. flossdorfii (Hicken) Dempster
Galium pseudoaparine Griseb. = G. aparine
Galium pseudotriflorum Dempster & Ehrendorfer
Galium pumilio Standley
= G. richardianum
Galium pusillum Endl. ex Walp. subsp. richardianum
=
Galium quítense Wedd. Relbunium (R. hypocarpium subsp. nitidum (H. B. K.)
Galium radicosum Steud. = Relbunium
(R. megapotamicum subsp. camporum (Pohl
ex DC.) Ehrend., fide Ehrendorfer, 1955)
Galium reflexum Pohl ex DC., nom. nud. pro syn. = Relbunium (R. megapotamicum
subsp. reflexum (K. Schum.) Ehrend., fide Ehrendorfer, 1955)
Galium relbun Clos = Relbunium (R. hypocarpium subsp. fluminense (Veil.) Ehrend.,
fide Ehrendorfer, 1955)
Galium repugnans Kunze, nom. nud. = G. aparine (q. v. in this list)
Galium reynoldsii Dempster
Galium richardianum (Gillies ex Hook. & Arn.) Endl.
Galium richardianum (Gillies ex Hook. & Arn.) Endl. subsp. richardianum
Galium richardianum subsp. haenkeanum (Gillies ex Hook. & Arn.) Dempster, comb,
nov.
Galium var. uncinulatum Kuntze = G. obovatum

rotundifolium
Galium = Relbunium
rupestre Walp. (R. rupestre (Gardner) Ehrend., fide Ehren
dorfer, 1955)
Galium saxatile L.
Galium scabrum Wedd. = G. ascendens
Galium scandens Philippi = G. trichocarpum
Galium sellowianum Walp. = Relbunium (R. sellowianum (Cham.) K. Schum., fide
Ehrendorfer, 1955)
Galium sessiliflorum Willd. ex Roemer & Schultes, nom. nud. pro syn. = Relbunium
(R. croceum subsp. involucratum (H. B. K.) Ehrend., fide Ehrendorfer, 1955)
Galium sessilifolium Willd. ex K. Schum., nom. nud. pro syn. = Relbunium (R.
croceum subsp. involucratum (H. B. K.) Ehrend., fide Ehrendorfer, 1955)
Galium suffruticosum Hook. & Am.

Galium var. araucanum Reiche = G. araucanum

suffruticosum
Galium surinamense Dempster, sp. nov.
Galium tarmense Spreng. = probably Relbunium (R. croceum (Ruiz & Pavon) K.
Schum. subsp. croceum, fide Ehrendorfer, 1955)
Galium telanthos Philippi = G. gilliesii subsp. telanthos
Galium tenellum Clos = G. divaricatum
Galium tenue Larrañaga = Relbunium (questionably R. megapotamicum (Spreng.)
Galium Griseb. = G. latoramosum
tetragonum
Galium trianae Wernham = G. canescens
Galium trichocarpum DC.
Galium trichocarpum var. telanthos Reiche = G. gilliesii subsp. telanthos
Galium tricorne Stokes = G. tricornutum
Galium tricornutum Dandy

Galium uruguayense Bacigalupo
Galium var. echinulatum = G.
uruguayense Bacigalupo uruguayense
Galium valdivianum Philippi = G. magellanicum
Galium volckmanni Philippi = G. gilliesii subsp. telanthos
Galium weberbaueri Krause
Galium werdermannii Standley
Relbunium chaetophorum K. Schum. = G. richardianum subsp. richardianum
Relbunium diphyllum K. Schum. = G. diphyllum
Relbunium echinocarpum Hassler = G. richardianum subsp. richardianum
Relbunium equisetoides Ehrend. = G. equisetoides
Relbunium humile K. Schum. = G. humile
Relbunium mazocarpum (South American usage only) = G. humile
Relbunium patagonicum K. Schum. = G. richardianum subsp. richardianum
Relbunium pusillum K. Schum. = G. richardianum subsp. richardianum
Relbunium richardianum Hicken = G. richardianum subsp. richardianum
Relbunium richardianum subsp. patagonicum Ehrend. = G. richardianum subsp.
richardianum
Relbunium richardianum Ehrend. = G. richardianum richardi
subsp. pusillum subsp.
anum
Rubia debilis Kunth = G. aschenbornii

Rubia diphylla Κ. Schum. = G. diphyllum
Rubia Cham. & Schlechtendal = G. latoramosum
ephedroides
Rubia equisetoides Cham. & Schlechtendal = G. equisetoides
Rubia haenkeana Gillies ex Hook. & Arn. = G. richardianum subsp. haenkeanum
Rubia intricata Hook. & Arn. = G. latoramosum
Rubia margaritifera Reiche = G. suffruticosum

Rubia pusilla Gillies ex Hook. & Arn. = G. richardianum subsp. richardianum
Rubia richardiana Gillies ex Hook. & Arn. = G. richardianum subsp. richardianum
Rubia scabra Kunth = G. ascendens
Rubia K. Schum. = G. latoramosum

tetragona
Sherardia muralis L. = G. murale

ALLERTONIA
LITERATURE CITED
Bacigalupo, Ν. M. 1973. Aclaración sobre la verdadera patria y la sinonimia de "Galium latoramosum"
Clos. Darwiniana 18: 279-280.
1975. Rubiaceas nuevas para la flora argentina y uruguaya. Darwiniana 19: 510-519.
Bentham, G., & J. D. Hooker. 1873. Genera Plantarum 2 (1).
Candolle, A. P. de. 1830. Rubiaceae, Prodromus Systematis Naturalis Regni Vegetabilis 4: 341-622.
Carauta, J. P. P. 1969. A data efectiva de publicaçâo da Flora Fluminensis I. Vellozia 7: 26-33.
1973. The text of Vellozo's Flora Fluminensis and its effective date of publication. Taxon 22:
281-284.
Dempster, L. T. 1973a. The fleshy-fruited Galiums (Rubiaceae) of Mexico and Central America. Brittonia
25: 15-20.
1973b. Transfer of Relbunium sphagnophilum (Rubiaceae) and R. mazocarpum to Galium.
Phytologia 26: 221-223.
1973c. Note on the type of Rubia acuminata Mart. & Gal. Brittonia 25: 313.
1980. The genus Galium section Lophogalium (Rubiaceae) in South America. Allertonia 2:
247-279.
1981. The genus Galium (Rubiaceae) in South America. II. Allertonia 2: 393-426.
Ehrendorfer, F. 1955. Revision of the genus Relbunium (Endl.) Benth. et Hook. (Rubiaceae-Galieae).
Bot. Jahrb. 76: 516-553.
Endlicher, S. 1836-1840. Genera Plantarum Secundum Ordines Naturales Disposita.
Farr, E. R., J. A. Leussink, & F. A. Stafleu. 1979. Index Nominum Genericorum (Plantarum) 3.
Holmgren, P. K., W. Keuken, & E. K. Schofield. 1981. Index Herbariorum. I. Ed. 7.
Hooker, W. J., & G. A. W. Arnott. 1833. Contributions towards a flora of South America and the islands
of the Pacific. Bot. Misc. 3: 129-212.
Moore, D. M. 1968. The vascular flora of the Falkland Islands. British Antarctic Survey Scientific Reports
60.
Philippi, R. A. 1864. Commentar zu den von Molina beschriebenen chilenischen Pflanzen. Bot. Zeitung
(Berlin) 22: Suppl. 2: 1-24.
Schumann, K. 1888. Rubiaceae. In: Martius, C. F. P. von. Flora Brasiliensis 6 (6): 1-466.


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THE GENUS GALIUM (RUBIACEAE) IN SOUTH AMERICA.

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pedicels. As type of the genus Ehrendorfer, without discussion, indicated R. hypocar

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Although I am persuaded by the evidence that Relbunium as a genus is scarcely

Abbreviations are those recommended by Index Herbariorum (Holmgren et al., 1981).

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Leaves more than 4 at all or most nodes.

1. Galium galapagoense Wiggins in Madroño 20: 250. 1970. Type: ECUADOR:

Low, congested, dioecious (?), herbaceous perennial to 9 cm. high, rooting at

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species may be dioecious (see Figure 1).

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Figure 2. Map showing distribution of Galium cotinoides, G. inconspicuum, and G. werdermannii.

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Slender polygamous plants, bunched, decumbent, or sprawling from a woody

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Figure 4. Map showing distribution of Galium aschenbornii, G. azuayicum, G. ascendens, and G.

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apparently restricted to Chile, is very similar to the allopatric G. richardianum east of

5a. Galium richardianum subsp. richardianum. Figure 7.

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Stems, leaves, and probably corolla (unknown) densely hispid.

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nium on p. 212 of the introduction.

Figure 6. Map showing distribution of Galium araucanum, G. diphyllum, G. equisetoides, G. humile,

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7. Galium diphyllum (K. Schum.) Dempster, comb. nov. Figure 9.

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leaves were seen on only two of 14 collections of G. equisetoides, it is clearly possible

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Distribution: Chile: Talca to southern Atacama, 200-2,700 m., and in Argentina

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Galium suffruticosum is often confused with non-fruiting plants of G. philippia

to the relationship with that species.

Galium uruguayense, of eastern South America, bears a relationship to G. richar

dianum similar to that of G. suffruticosum to G. araucanum in the far west. The

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Figure 14. Map showing distribution of Galium antarcticum and G. magellanicum.

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pedicels, the mericarps reniform.

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Galium humile is very closely related to G. sphagnophilum (Greenman) Dempster,

species, G. humile in South America and G. sphagnophilum in Mexico, even though

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angled, essentially glabrous or at most minutely scabrous; internodes about as long as

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