Review of The Genus Diplostephium

Review of the Genus Diplostephium
Author(s): S. F. Blake
Source: American Journal of Botany , Jan., 1928, Vol. 15, No. 1 (Jan., 1928), pp. 43-64
Published by: Wiley
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REVIEW OF THE GENUS DIPLOSTEPHIUM
S. F. BLAKE
(Received for publication May 28, I927)
In 1922 the writer' published a revision of Diplostephium, a genus

closely allied to Aster but distinguished by habit and by its double pappus.
Forty species were recognized, in addition to one of uncertain position,
divided into five series on characters of inflorescence and foliage. In
consequence of the fact that a third of the species were known to me only by
description, one or tWo were misidentified, and at least three were redescribed
under new names. The receipt, through the kindness of Dr. L. Diels and Dr.
J. Mattfeld, of a set of fragments of all the species described by Hieronymus,
and the opportunity to examine various other types in European herbaria in
1925, have made it possible to rectify most of the errors of the earlier
treatment and to present a new review of the genus.
The taxonomic history of the genus was sufficiently sketched in my
earlier paper, but its distribution calls for more discussion than was there
given it. Diplostephium now consists of 43 species, with their center of
abundance in Colombia, where 24 species are found; II or I2 are found in
Ecuador, IO or ii in Peru, 2 in Chile, and i each in Costa Rica, Venezuela,
and Bolivia. One species, D. ochraceum, is of doubtful habitat, but is
probably either Colombian or Ecuadorian. All the species are nearly or
quite confined to the paramo or its lower bordering shrub zone, the paramillo,
usually at altitudes of 3000 to 4270 meters, or perhaps even higher. Several
species occur down to 1830 meters in Peru, and D. costaricense was collected
by Pittier in humid woods at La Palma, Costa Rica, altitude I500 meters,
but also occurs in paramo thickets at 2700 to 3000 meters.
The local distribution of the 24 species definitely known from Colombia,
although as yet imperfectly known, offers some points of interest. Four
species only (D. anactinotum, D. parvifolium, D. cyparissias, and D. weddellii)
are known from the Sierra Nevada of Santa Marta, where all of them are
endemic. At least two species (D. schultzii and D. floribundum) occur in
both the central and western cordilleras, and one, D. baccharideum, is found
in both the eastern and western cordilleras, even to the extreme northern
limit of the latter, but is not known from the central cordillera.
The genus may be regarded as an Andean development of Aster, which
has taken on the shrub habit and further adapted itself to the extreme
conditions prevailing on the paramos by the development of tomentum on
the branches and under surface of the leaves. Specific differences are
furnished chiefly by the character of the inflorescence, the shape, size, and
1 Contr. U. S. Nat. Herb. 24: 65-86, pIs. 2I-28. 1922.
43
4

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44 AMERICAN JOURNAL OF BOTANY [Vol. I5,
pubescence of the leaves, and the size of the involucre and heads. Of the
five series distinguished in my earlier paper, the three last (Rupestria,
Floribunda, and Denticulata) are here combined under the name Rupestria.
The characters relied on in separating them are often uncertain, and the
presence or absence of regular dentation in the leaves is not even of specific
value.
The most interesting feature of the morphology of Diplostephium is
found in the variation exhibited by the styles of the disk flowers. In the
earlier revision three principal types of styles were distinguished and the
representative species listed. Many of the species not seen at that time have
since been examined, and a new list can now be presented. The style
characters in D. jelskii, D. cochense, D. obtusum, and D. ochraceum are still
unknown or insufficiently described, as well as those in authentic material
of D. meyenii, but the last species is not likely to differ from D. tacorense.
The specific names in quotation marks in the following lists are included on
the basis of Weddell's descriptions.
i. Style more or less clavate, merely bifid or even subentire, the branches very short,
ovate, usually obtuse, merely papillose: D. micradenium (Series Lavandulifolia); D.
revolutum, D. baccharideum, D. rosmarinifolium (Series Rosmarinifolia); D. rupestre, D.
eriophorum, D. schultzii, D. costaricense, D. lehmannianum, D. rhododendroides (approaching
type 2), D. pleistogynum, D. floribundum (Series Rupestria).
2. Style branches of medium length, oblong, linear-oblong, or lanceolate, acute, merely
papillose: D. anactinotum, "D. parvifolium" (Series Lavandulifolia); "D. cyparisias"
(Series Rosmarinifolia). A style intermediate between type 2 and type i is shown by D.
phylicoides (Series Rupestria), in which the branches are linear, acute, and hispidulous
dorsally, but often connivent nearly to apex; and styles more or less intermediate between
types 2 and 3 are found in D. weddellii, D. tacorense, D. leiocladum, and D. denticulatum (Series
Rupestria).
3. Style branches elongate, linear-subulate, acuminate, hispidulous or hispid dorsally,
the appendage sometimes thrice as long as the stigmatic region: D. empetrifolium, D.
glandulosum, D. oblanceolatum, D. macrocephalum, D. pearcei, D. callilepis, D. hippophae, D.
adenachaenium, D. gnidioides, D. lavandulifolium, D. hartwegii (Series Lavandulifolia); D.
carabayense (Series Rosmarinifolia); D. lechleri, D. pulchrum, D. haenkei, D. bicolor (Series
Rupestria). In D. spinulosum and D. antisanense (Series Lavandulifolia) the form of the
style branches is much the same, but they are merely papillose, and in D. spinulosum
appear to lack stigmatic lines.
In my earlier paper it was suggested that the primitive type of style in

Diplostephiunz is probably type 2, which approximates iaost naerly the
average or mean type of the tribe, and that types i and 3 have been derived
from it. It is equally possible that type 3, a form by no means uncommon
in the Astereae, is primitive, and that type i and type 2 have been derived
from it. In either case, the evolutionary course of the genus is apparently
directed toward the development of two groups, one with subentire or
slightly bifid styles and sterile disk flowers, the other with long-appendaged
styles and fertile disk flowers. This evolutionary tendency does not align
with the division into three series here adopted on the basis of inflorescence
and foliage-a division which, taking all the characters of the plants into

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Jan., I9281 BLAKE - DIPLOSTEPHIUM 45
consideration, seems to be a highly natural one -but cuts across it. Most
of the species of the Series Lavandulifolia fall into type 3, but D. micradenium
is typical of type I, and one or two other species enter type 2. Three of the
five species of Series Rosmarinifolia are of type i, one is of type 3, and one,
from description, appears to belong to type 2. Eight species of the Series
Rupestria belong to type I, four are of type 3, and five are variously inter-
mediate. That is, each of the two groups which may eventually attain
generic distinction by the development of differences in style and con-
comitant fertility or sterility of disk flowers is composed of species from each
of the three apparently very natural series into which the genus can now be
divided.
In connection with the evolutionary possibilities here discussed, the
variation shown by Diplostephium schultzii is of special significance. This
species is one of those with barely bifid style and a tendency to sterility of the
disk. In two collections examined, including the type number, the heads
were heterogamous, with IO to 44 rays and 4 to 32 disk flowers, a variation in
itself far beyond that known in any other species. Four heads dissected
from another collection, certainly referable to the same species, were
homogamous, with 27 to 38 ligules and no hermaphrodite flowers whatever.
The individual plant or plants from which these specimens were collected
apparently represent a remarkable mutation, of a sort I have never before
met with in wild Asteraceae, in which the head is composed wholly of
pistillate ray flowers. Whether a corresponding mutation resulting in th
production of discoid heads composed of staminate (infertile hermaphrodite)
flowers has taken place is uncertain, and probably unlikely, but it is evident
that a careful field study of this species would be of the highest interest.
The abortion of the ovules, in the achenes examined, would indicate that no
pollen was available in the vicinity of the plant or plants from which these
specimens were obtained.
The following abbreviations are used in this paper to refer to herbaria in
which specimens are deposited: F = Field Museum; G = Gray Herbarium;
K = Kew Herbarium; N = U. S. National Herbarium; Par. = Herbarium
of the Museum d'Histoire Naturelle, Paris. Except in a few cases, where
corrections in identification have been made, specimens already cited in my
previous paper have not been listed. My thanks are due the curators of the
herbaria mentioned for the opportunity to study and borrow the material
under their charge.
KEY TO SERIES
Heads solitary at tips of stem and branches . ........................ I. Lavandulifolia.

Heads cymose or cymose-panicled.
Leaves linear or essentially so, 2 mm. wide or less ................. 2. Rosmarinifolia.
Leaves linear-lanceolate to oblong or obovate, very rarely linear, 3-30 mm. wide.
3. Rupestria.

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KEY TO SPECIES
i. Lavandulifolia
Leaves densely and conspicuously impressed-glandular over whole upper surface.

Leaves finely impressed-glandular above, linear, about 25 mm. long, 1.5-2 mm. wide,
densely tomentose-pilose beneath and when young also sparsely pilose on upper
surface with comparatively long, loose, not crisped hairs; branches densely pilose-
tomentose with reflexed hairs ............................... I. D. micradenium.
Leaves coarsely impressed-glandular above, their tomentum dense, without long loose
hairs; branches closely tomentose, in one species glabrate or glabrescent.
Leaves 4-7 mm. long, essentially linear, not spinulose above; involucre 7-9 mm.
high ......................................... 2. D. empetrifolium.
Leaves usually IO-22 mm. long, if smaller then spinulose above; involucre Io-I3 mm.
high.
Larger leaves narrowly oblanceolate or linear-spatulate to nearly linear, 3 mm.
wide or less, closely ochraceous- or cinereous-tomentose beneath, the costa
glabrate and evident at least toward base; branches closely cinereous- or
ochroleucous-tomentose, usually glabrate or glabrescent; heads short-
peduncled ....................................... 3. D. glandulosum.
Larger leaves oblanceolate or spatulate-oblanceolate, I5-22 mm. long, 3.5-4.5
mm. wide, densely ochraceous- or fuscous-lanate beneath, the costa hidden;
branches densely brownish-lanate-tomentose; heads sessile or subsessile.
4. D. oblanceolatum.
Leaves not at all impressed-glandular above or only obscurely so (in D. tacorense and D.
meyenii), sometimes spinulose, glandular-tuberculate, or glandular.
Involucre io-IS mm. high.
Leaves narrowly spatulate or oblanceolate-spatulate, obtuse, fuscescently lanate-
tomentose beneath, glandular-tuberculate above ........... 5. D. macrocephalum.
Leaves strictly linear, acute or acutish, not glandular-tuberculate above.
Branches densely tomentose, not setulose.
Leaves spinulose along margin above; achenes glabrous or sparsely glandular.
6. D. pearcei.
Leaves glabrous above; achenes hirsutulous and glandular.... 7. D. callilepis.
Branches closely whitish-tomentose and densely setulose with thick yellowish
hairs; leaves spinulose along costa above, often also along margin; achenes
sparsely hirsutulous and toward apex glandular ........... 8. D. hippophae.
Involucre 4-9 mm. high.
Leaves densely glandular or resinous-coated but not pubescent above.
Leaf blades elliptic or oval, broadly rounded, definitely petioled.
9. D. anactinoturm.
Leaf blades triangular-oblong, acutish or obtuse, sessile . 1.0. . IO. D. parvifolium
Leaves not densely glandular or resinous-coated above, either glabrous, pubescent,
glandular-tomentose, or spinulose.
Leaf blades oblong or ovate-oblong, about 8 mm. long, 2.5 mm. wide, blackish-
lanate-tomentose beneath, appearing sessile, the angled petioles about I.5
mm. long, closely appressed to the stem and appearing adnate to it;
involucre black-lanate-tomentose ....................... ii. D. spinulosum.
Leaf blades linear to elliptic, or if tending toward oblong then with distinctly
spreading petioles; involucre not black-tomentose.
Leaf blades elliptic to linear-oblong, 7-I I mm. long, 1.5-3 mm. wide, soon
glabrous and shining above, not spinulose ....... . I2. D. adenachaenium.
Leaf blades linear to narrowly elliptic, not over 2 mm. wide.

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Jan., I928] BLAKE - DIPLOSTEPHIUM 47
Leaf blades glandular-tomentose above at least when young, narrowly

linear or linear-oblanceolate, 8-30 mm. long ...........1 3. D. meyen
I4. D. tacorense.
Leaf blades sometimes tomentose but not glandular above, 3.5-II mm.
long.
Leaves acutely callous-pointed, the tomentum of the lower surface
usually concealed by the strongly revolute margin.
Leaves glabrous above; achenes densely hirsutulous.
15. D. jelskii.
Leaves spinulose above; achenes very sparsely hirsutulous.
i6. D. gnidioides.
Leaves obtuse, or if callous-pointed (D. hartwegii) then with
tomentum of lower leaf surface conspicuous.
Involucre 6-7 mm. high.
Stem and leaves beneath densely appressed-tomentose with
whitish or subcanescent hairs ........ I7. D. lavandulifolium.
Stem and leaves beneath densely and loosely tomentoge with
ochroleucous or reddish hairs .............. I8. D. hartwegii.
Involucre 4-5 mm. high ..................... I9. D. antisanense.
2. Rosmarinifolia
Branchlets glabrous, more or less viscid ............................. 20. D. revolutum.

Branchlets densely tomentose, sometimes glabrescent.
Heads comparatively large, the involucre about 9-IO mm. high; leaves 8-15 mm. long,
1.5-2 mm. wide ........................................... 21. D. carabayense.
Heads smaller, the involucre 5-7 mm. high; leaves (o.9) 1.5-4.2 cm. long.
Leaves slightly revolute-margined, pale green and persistently viscid-tomentellous
above, strictly sessile; heads subsessile ..................... 22. D. cyparissias.
Leaves strongly revolute-margined, dark green and glabrate or glabrescent above,
shortly but definitely petioled; heads pedicellate (pedicels usually 2-8 mm.
long).
Leaves I-2.5 cm. long; disk achenes sparsely hispid ......... 23. D. baccharideum.
Leaves 2.7-4.2 cm. long; disk achenes glabrous... . 24. D. rosmarinifolium.
3. Rupestria
Heads large, mostly on monocephalous peduncles in a usually simple or subsimple leafy-
bracted cyme; involucre (8) 9-I2 mm. high.
Leaves sessile by a comparatively broad base, impressed-punctate above (sometimes
obscurely so in D. weddellii).
Branchlets closely ochraceous-tomentose; heads subsessile ......... 25. D. weddellii.
Branchlets densely lanate-tomentose or pilose with spreading hairs; heads distinctly
peduncled.
Branchlets pilose or pilose-tomentose; leaves strongly viscid above, mostly
linear or linear-elliptic (by revolution of the margin), 2.5-5 cm. long, 3-7 mm.
wide ........... ........................... 26. D. rupestre.
Branchlets densely lanate-tomentose; leaves scarcely viscid above, elliptic or
ovate-elliptic, I.5-2.5 cm. long, 4-IO mm. wide ........... 27. D. eriophorum.
Leaves distinctly petioled or narrowed into a petioliform base, not impressed-punctate
above.
Leaves white-sericeous or -subsericeous beneath with a very close and comparatively
thin but persistent tomentum.
Leaves glabrous above, except on costa; branches closely and persistently
ochraceous-tomentose .................................... 28. D. lechleri.

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Leaves hispidulous on surface above; branches densely hirsute-pilose with yellowish

hairs as well as tomentose, the tomentum more or less deciduous.
29. D. pulchrum.
Leaves usually ochroleucous- to ferruginous-tomentose beneath, the tomentum dull,
thick, and comparatively loose ............................... 30. D. haenkei.
Heads smaller, usually in a cymose panicle; involucre 4.5-7 (8) mm. high.
Leaves very small, elliptic or oblong, thick-coriaceous, 5-II mm. long, I.8-3 mm. wide,
abruptly very short-petioled, not narrowed toward base of blade; heads in a close,
essentially simple, umbelliform cyme ........ .................. 31. D. phylicoides.
Leaves larger, mostly obovate or oblanceolate, narrowed to a petiole or petioliform base;
heads usually cymose-panicled.
Leaves persistently gray-tomentose or -tomentulose above, sometimes glabrate at
length (in D. obtusum).
Lateral veins 3-4 pairs, spreading at an angle of 45-6o', practically concealed
beneath by the tomentum.
Heads 27-50-flowered; larger leaves 2-3.3 cm. long.. 32. D. schultzii.
Heads I2-15-flowered; larger leaves 2 cm. long.. . 33. D. cochense.
Lateral veins 6-io pairs, spreading at nearly a right angle, usually evident
beneath.
Leaves acute or acutish, acutely callous-tipped; heads 22-57-flowered.
34. D. costaricense.
Leaves rounded or emarginate, not callous-apiculate; heads i8-2i-flowered.
35. D. obtusum.
Leaves green and glabrous or quickly glabrate above (except sometimes on costa).
Larger leaves 1.5-2.7 cm. long.
Involucre 4.5-5 mm. high; leaves 1.5 cm. long or less, obovate.
36. D. lehmannianum.
Involucre 7-8 mm. high; leaves 1.3-2.7 cm. long.
Leaves obovate, 2-2.5 cm. long, 8-io mm. wide; rays 6-8, disk flowers
about 20 ................................. 37. D. rhododendroides.
Leaves oblanceolate, I.3-2.7 cm. long, 4-7 mm. wide; rays about 27, disk
flowers 7 ..................................... 38. D. pleistogynum.
Larger leaves 4-IO cm. long.
Lateral veins slender, straightish, spreading from the costa at nearly a right
angle; petioles narrowly margined essentially to base.
Branchlets and peduncles persistently tomentose ..... 39. D. floribundum.
Branchlets and peduncles quickly nearly or quite glabrate.
40. D. leiocladum.
Lateral veins stout, curved, spreading at an angle of about 45-6o0; petioles
naked.
Leaf blades pergamentaceous, the secondary veins conspicuous and
reticulate beneath; achenes pubescent ........... 41. D. denticulatum.
Leaf blades more or less coriaceous, not conspicuously reticulate beneath,
the secondary veins mostly concealed by the tomentum; achenes
glabrous or sparsely glandular.
Leaf blades elliptic-oblong, rounded or bluntly cuneate at base.
42. D. ochraceum.
Leaf blades obovate or oblong-obovate, acutely cuneate at base.
43. D. bicolor.
SERIES I. LAVANDULIFOLIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

Heads solitary at tips of stem and branches or branchlets, the latter
often numerous and crowded.

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i. Diplostephium micradenium Blake, sp. nov.

Branches densely ochroleucous-pilose-tomentose with straightish reflexed
hairs; leaves linear, about 2.5 cm. long, obtuse, sessile, above green,
impressed-punctate, when young sparsely and loosely pilose, beneath
densely ochroleucous-pilose-tomentose with comparatively straight, antrorse
hairs, strongly revolute-margined; heads solitary, large; involucre I-I.2 cm.
high.
Shrub; branches few, erect, dichotomous, smoothly denudate in age, 2-4
mm. thick, terete, their pubescence grayish in age; internodes 1.5-3 mm.
long; leaves erectish, 2-3.2 cm. long, I.5-2.2 mm. wide, obtusely callous-
tipped, with broad costa; peduncle about I cm. long; disk about I2 mm.
high; involucre strongly graduate, about 5-seriate, the phyllaries lance-
ovate (outer) to linear, acute to subacuminate, whitish and indurate, with
purplish-brown tips and the costa thickened and darkened above, loosely and
sparsely pilose and ciliate, sessile-glandular; rays "white," the tube
hirsutulous above, 3.5 mm. long, the lamina linear-elliptic, 4-nerved, about
IO mm. long, I.8 mm. wide; disk corollas "brownish yellow," numerous,
hirsutulous on throat, slender, about 6 mm. long (tube I.5 mm., throat 3.5
mm., teeth I-I.3 mm.); achenes 4-5-nerved, rather densely hirsutulous and
toward apex sessile-glandular, those of the ray compressed, 4 mm. long, 2
mm. wide, of the disk linear; infertile, 5-6 mm. long; pappus bristles
brownish, 7 mm. long, slightly enlarged above, the outer pappus rather
obscure, setulose, 0.5-1.2 mm. long; style of disk flowers finely hirsutulous
above, minutely bilobate.
COLOMBIA: Shrub zone, below paramo, Cerro Tatama, Cordillera
Occidental, Dept. Caldas, alt. 3300-3500 m., 8-io Sept. I922, Pennell
I0533 (type no. I, I4I, 3I8, U. S. Nat. Herb.).
A very distinct species, remarkable for the apparently complete sterility
of the disk.
2. DIPLOSTEPHIUM EMPETRIFOLIUM Blake, Contr. U. S. Nat. Herb. 24: 73.

pI. 2I. I922.
RANGE: Ecuador, Peru (?).
ADDITIONAL SPECIMENS EXAMINED: ECUADOR: Between San Lucas and
Ofia, Prov. Loja, alt. 2200-3IO0 m., 7 Sept. 1923, Hitchcock 2I526 (N).
This species is not closely allied to any other. It is readily recognized
by its very small, uniform, acutely callous-pointed leaves, green and densely
impressed-glandular above, and closely tomentose beneath. The rays, in
Hitchcock's comparatively fresh specimens, were evidently white; the
disk corollas, at least in age, are purple.
3. DIPLOSTEPHIUM GLANDULOSUM Hieron. Bot. Jahrb. Engler 2 1: 339.

I895.
Diplostephium spinulosum Hieron. Bot. Jahrb. Engler 29: 22. I900;
Blake, Contr. U. S. Nat. Herb. 24: 76. 1922. Not D. spinulosum
Wedd. (err. ident.).
Diplostephium cicatricosum Blake, Contr. U. S. Nat. Herb. 24: 76.
pI. 23. I922.

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50 AMERICAN JOURNAL OF BOTANY [Vol. IS,
TYPE LOCALITY: PAramo de la Boca del Mundo Nuevo, VolcAn de

Cumbal, Colombia.
RANGE: Colombia, Ecuador.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Paramo de la Boca del
Mundo Nuevo, VolcAn de Cumbal, Jan.-Feb. I870, Stuebel 445 b (type of
Diplostephium glandulosum; fragm. N); same locality, Karsten (herb. Sch.
Bip., fragm. N). Tuquerres, Prov. Tuquerres, alt. 3200 m., Triana 1257
(Par.). Mt. Azufral, Cordillera Meridional, i8 May I876, Andre (G). Ecu-
ADOR: Pilzhun, alt. 3965 m., Jameson (G). Arid hills near Riobamba, Sept.
I893, Sodiro 64/3 (fragm. N). Mirador, July I902, Rivet I83 (Par.), Aug.
1902, Rivet I94 (Par.). Naes, Feb. I903, Rivet 358 (Par.). El Pelado, Jan.
1903, Rivet 312, 315, 333 (Par.). Quinoaloma, Mar. 1904, Rivet 590 (Par.).
In my 1922 revision, Diplostephium glandulosum Hieron. was doubtfully
referred to D. spinulosum Wedd., the latter being interpreted by Hieronymus'
ample description 2 based on Sodiro 64/3. Examination of the type col-
lection of D. spinulosum, of fragments of two collections (including the type)
of D. glandulosum, and of fragments of Sodiro 64/3 has now shown that D.
spinulosum is a very distinct species and that D. glandulosum Hieron., D.
cicatricosum Blake, and "D. spinulosum" of Hieronymus and of Blake, not
of Weddell, are to be combined under the earliest name, D. glandulosum.
The specimens cited under D. spinulosum in my previous paper belong to D.
glandulosum. Karsten's plant in the Schultz Bipontinus herbarium bears
unpublished specific and varietal names of Schultz.
Rivet's labels describe the flowers as rose, violet, white, or yellowish.
The last can scarcely refer to the color of the rays. The plant is a shrub up
to about 45 cm. high, usually fasciculately branched above.
4. DIPLOSTEPHIUM OBLANCEOLATUM Blake, Contr. U. S. Nat. Herb. 24: 76.

PI. 24. 1922.
RANGE: Ecuador.
ADDITIONAL SPECIMEN EXAMINED: ECUADOR: Minas, Nov. I904, alt.
4000 m., Nov. 1904, Rivet 735 (Par., fragm. N).
This species is close to D. glandulosum, differing chiefly in its densely and
persistently tomentose branchlets, usually larger leaves with the costa
concealed beneath, and sessile or subsessile heads, those of D. glandulosum
being normally borne on short but evident peduncles 5-I5 mm. long. Rivet
notes the color of the flowers as violet.
5. DIPLOSTEPHIUM MACROCEPHALUM Blake, Contr. U. S. Nat. Herb. 24: 75.

PI. 22. 1922.
RANGE: Ecuador.
ADDITIONAL SPECIMENS EXAMINED: ECUADOR: Niarihuinia, alt. 3900 m.,
Dec. I904, Rivet 738, 743 (Par.).
Closely related to D. oblanceolatum, and distinguished principally by the
2 Bot. Jahrb. Engler 29: 22. 1900.

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entire lack of impressed glands on the leaves, which are conspicuously

tuberculate with blunt papillae above toward margin and tip. The flowers
are described as violet by Rivet. The plant is a low shrub under 30 cm.
high.
6. Diplostephium pearcei Sprague & Blake, sp. nov.

Branches densely griseous-tomentose; leaves linear, 7-II mm. long,
I-I.5 mm. wide, acute, green above, spinulose along margin and at apex,
densely griseous-tomentose or in age fuscescent-tomentose beneath,
strongly revolute-margined; heads solitary, large; involucre I2-15 mm.
high; achenes glabrous or sparsely glandular.
Shrub o.6-I.3 m. high, fastigiate-branched, very leafy; stem glabrate,
terete, 6 mm. thick, the extreme bases of the leaves persistent, appressed;
tomentum of the branches becoming fuscous; leaves spreading, obscurely
short-petioled, not glandular-punctate, the costa impressed above, concealed
beneath; heads sessile at tips of branches, 2.5-3 cm. wide; involucre
hemispheric, I.2-I.5 cm. high, about 6-seriate, strongly graduate, the
phyllaries lance-ovate (outermost) to linear, acute or subacuminate, pale and
indurate with narrow thin lacerate-ciliate margin, the outer persistently
fuscescent-tomentose dorsally, the inner glabrous or glabrate dorsally;
rays "white," 36 or more, the tube I.5 mm. long, sparsely hirsutulous, the
lamina linear, 3-denticulate, I3.5 mm. long, I.5 mm. wide, 4-nerved; disk
corollas numerous, sparsely hirsutulous toward base of throat, 5.3 mm. long
(tube I.5 mm., throat 3 mm., teeth o.8 mm.); ray achenes (immature) I.5
mm. long, obscurely glandular above or essentially glabrous, their pappus
brownish white, the inner bristles 5.5 mm. long, the outer 3.5 mm.; disk
achenes (immature) sparsely glandular, I.5 mm. long, their pappus brownish
white, the inner of bristles 5.5 mm. long, the outer of aristiform squamellae
about I mm. long, sometimes aristate-tipped; style appendages subulate-
acuminate, evenly hispidulous, 1.3 mm. long.
PERU: Boggy places in hills, Puitac (near Huanta), alt. 3660-3965
m. February I867, R. Pearce (types in Kew Herbarium and Brit. Mus.;
photog. and fragm., U. S. Nat. Herb.).
Allied to the Ecuadorian Diplostephium macrocephalum Blake, which is
readily distinguished by its narrowly oblanceolate or spatulate-oblanceolate,
obtuse, glandular-tuberculate leaves, i.5-3 mm. wide.
7. Diplostephium callilepis Blake, sp. nov.

Branchlets closely subargenteous-tomentose; leaves linear, about I.2 cm.
long, glabrous above, densely and closely subargenteous-tomentose beneath,
short-petioled, subacutely callous-tipped; heads solitary, subsessile, large;
involucre io mm. high, the outer phyllaries densely whitish-tomentose, the
inner linear, obtuse, slightly ampliate at apex, with deeply lacerate-ciliate
scarious margin; achenes sparsely hirsute and glandular.
Low shrub, fastigiately branched, glabrescent in age, the petiole bases
persistent; petioles i mm. long; blades more or less spreading, 8-I5 mm.
long, I-I.5 mm. wide, shining above, sometimes tomentulose at base of
costa, rather strongly revolute, the costa concealed beneath; heads solitary
at tips of branchlets, about 2.5 cm. wide; disk 8-io mm. high; involucre
strongly graduate, about 6-seriate, the outer phyllaries triangular, acute,
lacerate-ciliate toward apex, the inner linear, thinly pilose or glabrate, the

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52 AMERICAN JOURNAL OF BOTANY [VO]. I5,
body o.6-o.8 mm. wide, purplish-brown, its deeply lacerate-ciliate, brownish-

white, scarious margin about as wide; receptacle alveolate; rays about 35,
"pale hortense violet," the tube sparsely puberulous, 1.5 mm. long, the
lamina linear, bidenticulate, 1.3 cm. long, I.5 mm. wide; disk corollas
numerous, purple above, sparsely pilosulous, 5.8 mm. long (tube I mm.,
throat cylindric-funnelform, 4 mm., teeth o.8 mm. long); achenes sparsely
hirsutulous and especially above glandular, 2 mm. long; pappus barely
purplish-tinged, 4.8 mm. long, some of the inner bristles slightly thickened
above, the outer pappus setulose-squamellate, 1.2 mm. long; stigmatic
region o.5 mm. long, the style appendages subulate, acuminate, I.2 mm.
long, short-hirsute.
PERU: Rocky banks on p'aramo, Paso de Tres Cruces, Cerro de Cusil-
luyoc, Dept. Cuzco, alt. 3800-3900 m., 3 May I925, Pennell I3858 (type no.
558I44, Field Mus.; photog. and fragm., U. S. Nat. Herb.).
Allied to D. pearcei and D. hippophae; distinguished from the latter by
the lack of setulae on the branches, and from both by the glabrous upper
surface of the leaves and the apically somewhat ampliate, elegantly lacerate-
ciliate inner phyllaries.
8. Diplostephium hippophae Blake, sp. nov.

Branches closely whitish-tomentose and densely setulose with short,
thick, spreading, yellowish hairs, these long-persistent; leaves linear, about
I.5 cm. long, acutely callous-pointed, short-petioled, above green, antrorse-
spinulose along costa and often along margin, beneath densely silvery-
tomentose with a very closely matted tomentum, narrowly revolute-
margined; heads solitary, large; involucre I3 mm. high.
Shrub i m. high, fastigiate-branched above, the branches simple; inter-
nodes I-3 mm. long; petioles i mm. long; leaves spreading, I.2-I.8 cm.
long, I-2 mm. wide, the costa rather narrow, usually naked beneath; the
upper leaves often bearing axillary fascicles of reduced leaves; head ap-
parently sessile, about 2.5 cm. wide; disk I.4 cm. high; involucre strongly
graduate, 7-8-seriate, the outer phyllaries (about 4 rows) triangular-ovate,
acute, closely whitish-tomentose, glabrescent, the inner linear, acute or
subacuminate, lacerate-ciliate at apex, on back glabrate or sparsely pilo-
sulous, all indurate and brownish-white; rays 20 or more, " purplish,"
hirsutulous on tube above, the tube 2-2.5 mm. long, the lamina linear,
tridenticulate, 4-nerved, 14 mm. long, I.5 mm. wide; disk corollas rather
numerous, purple, hirsutulous chiefly at base of throat, 7.7 mm. long (tube 2
mm., throat 4.5 mm., teeth I.2 mm.); achenes sparsely hirsutulous and
toward apex sparsely glandular, 2-2.8 mm. long, those of the disk at least
sometimes fertile; pappus purple-tinged, 6-7 mm. long, somewhat graduate,
the longer bristles somewhat thickened at apex, the outer pappus sparse,
setulose, about 0.5 mm. long; style branches linear or linear-subulate,
hirsutulous above.
PERU: Grassy slope, Villcabamba, an hacienda on Rio Chinchao, Dept.
Huanuco, alt. I830 m., I7-26 July I923, Macbride 5136 (type no. 536184,
Field Mus.; photog. and fragm., U. S. Nat. Herb.).
9. DIPLOSTEPHIUM ANACTINOTUM Wedd. Chlor. And. I: 20I. pI. 35, f. B.

I857; Blake, Contr. U. S. Nat. Herb. 24: 72. 1922.
RANGE: Sierra Nevada of Santa Marta, Colombia.

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ADDITIONAL SPECIMEN EXAMINED: COLOMBIA: Piramo, San Miguel,

Sierra Nevada de Santa Marta, alt. 3000 m., Karsten (herb. Sch. Bip.,
fragm. N).
Easily recognized by its small, elliptic or oval, thick leaves, 5-IO mm.
long, 2-4 mm. wide (including the petiole, this I-2.5 mm. long), covered
with a viscid coat above, and its very short ligules surpassed by the styles.
The labels of the specimens in Schultz's herbarium bear an unpublished
generic name referring to the fragrant quality of the leaves.
IO. DIPLOSTEPHIUM PARVIFOLIUM Blake, Contr. U. S. Nat. Herb. 24: 74.

I922.
Diplostephium microphyllum Wedd. Chlor. And. I: 20I. I857. Not D.

microphyllum Nees, I832.
Linochilus microphyllus Sch. Bip.; Wedd. Chlor. And. T: 201. I857, as
synonym.
SPECIMENS EXAMINED: COLOMBIA: Santa Marta, alt. 3200 m., Jan.
I843, Funck 477 (herb. Sch. Bip., fragm. N); same locality, Galeotti 388
(herb. Sch. Bip., ex herb. Turcz.; presumably of the type collection).
The affinity of this species is with D. anactinotum, as is indicated by its
short rays (the lamina 3 mm. long, according to Weddell), and the densely
glandular upper surface of its closely imbricate, sessile leaves. The im-
mature achenes are strongly compressed and rather sparsely appressed-
pubescent, and the outer pappus is composed of slender setae I-2 mm. long.
II. DIPLOSTEPHIUM SPINULOSUM Wedd. Chlor. And. I: 200. I857.

SPECIMENS EXAMINED: COLOMBIA: Shrub zone (p6ramillo), Mt. Pan de
Azucar, Cordillera Central, Dept. Cauca, alt. 3300-3600 m., i6 June I922,
Pennell 7041 (N). ECUADOR: Alpine pastures, Andes of Quito, alt. 4270 m.,
Jameson 406 (type, Par.; photog. and fragm. N).
The confusion of this species with D. glandulosum by Hieronymus and
the writer has been discussed under species no. 3 of this revision. Diplo-
stephium spinulosum is easily recognized by its blackish tomentum and its
small, oblong or oblong-ovate leaves borne on closely appressed petioles.
The spinules of the upper leaf surface are nearly absent in Pennell's plant.
The rays in this collection are described as white, the disk as yellow. The
achenes are sparsely glandular and hispidulous, and the outer pappus
consists of setiform paleae 0.5-I mm. long.
12. DIPLOSTEPHIUM ADENACHAENIUM Blake, Contr. U. S. Nat. Herb. 24: 72.

I922.

ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Shrub zone (paramillo),
Mt. Pan de Azucar, Cordillera Central, Dept. Cauca, alt. 3300-3600 m., I6

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54 AMERICAN JOURNAL OF BOTANY [Vol. IS,
June I922, Pennell 7035 (N). Mt. Azufral, Tuquerres, alt. 3800 m., Karsten
(herb. Sch. Bip., fragm. N). ECUADOR: Mirador, July I902, Rivet I82
(Par.). El Angel to La Posta (?), alt. 3500 m., Feb. 1903, Rivet 408 (Par.).
Readily recognized by its small, shining, elliptic to linear-oblong,
distinctly petioled leaves, and densely glandular achenes. Pennell describes
the rays as purple-violet; Rivet gives the flower color in one case as white,
in the other as blue. Karsten's plant bears an unpublished specific name of
Schultz Bipontinus.
13. DIPLOSTEPHIUM MEYENII Wedd. Chlor. And. I: 20I. I857; Blake,

Contr. U. S. Nat. Herb. 24: 72. I922.
Linochilus meyenii Sch. Bip.; Wedd. Chlor. And. I: 20I. I857, as
synonym.
?Aster (?) trachyticus F. Phil.; Phil. Anal. Mus. Nac. Chile Bot. I89I: 37.
I89I.
RANGE: Definitely known only from the type locality, Cordillera of
Tacora, altitude 4000-4500 m., Department of Tacna, Peru [now Chile?].
This species and the following one are closely allied if not identical.
Weddell's description is not sufficiently detailed to enable the two to be
distinguished satisfactorily. The following specimens, all from the De-
partment of Arequipa, Peru, alt. 2600-3700 m., belong to D. tacorense or D.
meyenii: Rose I8954 (N), Pennell I3202 (F), I32I3 (F), I326I (F), 14272
(F). A closely related but possibly distinct plant, with oblanceolate leaves
about I.5 cm. long and 3 mm. wide, and involucre IO mm. high, is represented
by Pennell I3333 (F), from the same region.
I4. DIPLOSTEPHIUM TACORENSE Hieron. Bot. Jahrb. Engler 2I: 337.

I895; Blake, Contr. U. S. Nat. Herb. 24: 72. I922.
RANGE: Known only from the type locality in northern Chile (Tacora).
SPECIMEN EXAMINED: CHILE: Tacora, alt. 4270-5i85 m., April I831,
Meyen (type, fragm. N).
i5. DIPLOSTEPHIUM JELSKII Hieron. Bot. Jahrb. Engler 36: 376. I905;
Blake, Contr. U. S. Nat. Herb. 24: 73. I922.
RANGE: Known only from the type locality, Cutervo, Dept. Caxamarca,
Peru.
SPECIMEN EXAMINED: PERU: Cutervo, May I879, de Jelski 622 (type,
fragm. N).
Distinguished by its small, rather acutely callous-pointed, linear or
"linear-lanceolate" leaves, glabrous and shining above, and very strongly
revolute-margined.
i6. Diplostephium gnidioides Blake, sp. nov.

Branches loosely cinereous- or in age fuscous-tomentose and yellowish-
setulose; leaves linear to narrowly lance-linear, about 8 mm. long, acutely

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callous-pointed, short-petioled, spinulose above, usually so strongly revolute-

margined as to conceal the loose grayish tomentum beneath; heads solitary,
medium-sized, subsessile; involucre about 8 mm. high; achenes sparsely
hirsutulous and toward apex sessile-glandular.
Shrub 30-60 cm. high, fastigiately branched, the petiole bases persistent,
the branches glabrescent; internodes I-3 mm. long; petioles about I mm.
long; blades spreading or erectish, 6-ii mm. long, 0.6-2 mm. wide, above
green, sparsely or usually densely spinulose, often with axillary fascicles;
disk 8 mm. high; involucre strongly graduate, about 5-seriate, the phyllaries
lance-ovate to linear, acute or subacuminate, indurate, whitish brown,
rather densely and subpersistently cinereous-pilose-tomentose and ciliate;
rays essentially glabrous, the tube 2 mm. long, the lamina linear, bidenticu-
late, 3-4-nerved, I3 mm. long, 1.3 mm. wide; disk corollas purple above,
puberulous on throat and teeth, 6.3 mm. long (tube 2.5 mm., throat thick-
cylindric, 3 mm., teeth o.8 mm.); achenes I.8 mm. long, those of disk
5-nerved; pappus brownish white or purple-tinged, 5 mm. long, the inner
bristles slightly thickened apically, the outer pappus setulose, o.5-i mm.
long; stigmatic region 0.5 mm. long, the linear-subulate, acuminate,
hispidulous appendages i.6 mm. long.
PERU: Grassy uplands, 9.6 km. south of Mito, Dept. Hu6nuco, alt.
3050 m., i-5 Aug. 1922, Macbride & Featherstone I858 (type no. 5I8353.
Field Mus.; duplicate no. II2I760, U. S. Nat. Herb.). On open slope,
Villcabamba, on Rio Chinchao, Dept. Huanuco, alt. I830 m., I7-26 July
I923, Macbride 5I84 (F, N).
17. DIPLOSTEPHIUM LAVANDULIFOLIUM H. B. K. Nov. Gen. & Sp. 4: 97.

pl. 335. I820; Wedd. Chlor. And. I: I99. pl. 36, f. A. I857.
Diplopappus lavandulifolius Cass. Dict. Sci. Nat. 25: 96. I822.
Aster lavandulaceus Willd.; Nees, Gen. & Sp. Ast. I89. I832, as
synonym.
Linochilus lavandulifolius Sch. Bip.; Wedd. Chlor. And. I: 200. I857,
as synonym.
RANGE: Ecuador.
SPECIMEN EXAMINED: ECUADOR: [Near Mulalo, alt. 2930 m., near base
of Mt. Illinissa and Mt. Cotopaxi,j Bonpland 3064 (type, herb. Humb. &
Bonpl.).
I8. DIPLOSTEPHIUM HARTWEGII Hieron. Bot. Jahrb. Engler 21: 337.

ADDITIONAL SPECIMENS EXAMINED: ECUADOR: Saraguru Mountains,
Hartweg 763 (type, fragm. N). Andine region, Mt. Pichincha, 4000 m.,
I9I7, Mille (N).
Described by Father Mille as a shrub 2-3 meters high.
I9. DIPLOSTEPHIUM ANTISANENSE Hieron. Bot. Jahrb. Engler 21: 338.

Diplostephium pycnophyllum Blake, Contr. U. S. Nat. Herb. 24: 75.
I922.
RANGE: Ecuador.

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SPECIMENS EXAMINED: ECUADOR: Paramos of Cerro Antisana, near Las

Cimarronas, alt. 4000 m., Oct. I871, Stuebel 235a (type, fragm. N). Paramo
near Cafiar, i6 Sept. I9I8, Rose & Rose 22750 (type of D. pycnophyllum,
N). PAramo between Cuenca and Huigra, Sept. I923, Hitchcock 2I66I (N).
Distinguished from D. lavandulifolium and D. hartwegii by its smaller
heads and shorter involucre. Hitchcock describes the plant as a shrub 0.3-I
meter high, with white rays.
SERIES 2. ROSMARINIFOLIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

Heads several to many at tips of stem and branches, cymose or cymose-
panicled; leaves linear or slightly spatulate-linear, 2 mm. wide or less
(rarely slightly wider in no. 2I).
20. DIPLOSTEPHIUM REVOLUTUM Blake, Contr. U. S. Nat. Herb. 24: 78.

1922.
RANGE: Vicinity of Bogota, Colombia.

ADDITIONAL SPECIMEN EXAMINED: COLOMBIA: Moist p'aramo near
Laguna Verjon, near BogotA, alt. 3200-3400 m., 27 Sept. I9I7, Pennell 2263
(N).
Unique in its group in its essentially glabrous branchlets and pedicels.
The rays are described as white. Pennell's plant was distributed as a
Baccharis, which it much resembles in appearance.
2I. DIPLOSTEPHIUM CARABAYENSE Wedd. Chlor. And. I: 202. I857;

RANGE: Known only from the type locality, in alpine region of south-
eastern Peru.
SPECIMEN EXAMINED: PERU: Near San Juan del Oro, Prov. Carabaya,
June-July I847, Weddell 4648 (type, Par.; photog. and fragm. N).
Distinguished in its group by its large heads and comparatively small,
linear leaves (8-I5 mm. long, I.5-2 mm. wide), green and glabrous above,
densely lanate-tomentose beneath with ochroleucous or whitish hairs.
Weddell's label gives the color of the flowers as pale violet or lilac.
22. DIPLOSTEPHIUM CYPARISSIAS Wedd. Chlor. And. I: 203. I857; Blake,

Contr. U. S. Nat. Herb. 24: 77. 1922.
SPECIMEN EXAMINED: COLOMBIA: Sierra Nevada of Santa Marta, alt.
2745 m., Funck 387 (type, Par.; photog. and fragm., N).
This species, known only by the original collection, has leaves 1.5-2 cm.
long, 1.5-2 mm. wide. The flowers are described as white.
23. DIPLOSTEPHIUM BACCHARIDEUM Blake, Contr. U. S. Nat. Herb. 24: 77.

PI. 25. I922.
RANGE: Colombia.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Without definite

186.11.18.239 on Fri, 13 May 2022 16:51:05 UTC
locality, Triana I263 (Brit. Mus.). Paramo de Choachi, near Bogota, alt.
3700 in., 8 Aug. I922, Killip & Ariste-Joseph II927 (N). Shrub zone below
pAramo, Cerro Tatama, Cordillera Occidental, Dept. Caldas, alt. 3300-
3500 m., Sept. I922, Pennell I0535 (N).
The flowers are described as white (no. II927). Pennell gives the
vernacular name "romero" for this shrub.
24. DIPLOSTEPHIUM ROSMARINIFOLIUM (Benth.) Wedd. Chlor. And. I: 202.

I857; Blake, Contr. U. S. Nat. Herb..24: 77. 1922.
Linochilus rosmarinifolius Benth. P1. Hartw. I97. I845.
RANGE: Known only from the type collection from near BogotA,
Colombia.
Closely similar to D. baccharideum, but with considerably long and
somewhat broader leaves.
SERIES 3. RUPESTRIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

Heads several to many at tips of stem and branches, cymose or cymose-
panicled; leaves linear-lanceolate to oblong or obovate, rarely linear, 3-30
mm. wide.-Including Series Floribunda and Denticulata Blake, Contr. U. S.
Nat. Herb. 24: 69. 1922.
25. DIPLOSTEPHIUM WEDDELLII Blake, Contr. U. S. Nat. Herb. 24: 79.

I922.
Diplostephium sessiliflorum Wedd. Chlor. And. I: 204. I857. Not D.

sessiliflorum Spreng. I826.
SPECIMENS EXAMINED: COLOMBIA: Sierra Nevada, Prov. Rio Hacha,
alt. 366o-4I20 m., March I852, Schlim 8o6 (type, Par.; photog. and
fragm. N). San Miguel, Sierra Nevada of Santa Marta, Karsten (herb. Sch.
Bip., fragm. N).
In Karsten's plant the heads are apparently solitary in some cases.
26. DIPLOSTEPHIUM RUPESTRE (H. B. K.) Wedd. Chlor. And. I: 206.

Aster rupestris H. B. K. Nov. Gen. & Sp. 4: 94. pl. 334. I820.
Tetramolopium ? rupestre Nees, Gen. & Sp. Ast. 203. I832.
Aster pichinchensis Willd.; Nees, Gen. & Sp. Ast. 203. I832, as
synonym.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Shrub-forest on slope,
P'aramo del Quindio, Dept. Caldas, alt. 4200-4400 m., Aug. I922, Pennell &
Hazen 9907 (N); grassy pAramo, alt. 4IOO-4400 m., same locality, Pennell &
Hazen 9835 (N). ECUADOR: Ruici-Pichincha, Bonpland (type, Par.).
Mt. Pichincha, alt. 4000 m., Andre' 391 I (Field Mus.). Mt. Pichincha, alt.
4IOO-4500 m., Aug. I7, I923, Hitchcock 2IO87 (N). Mt. Rumifiahui, alt.
4300 m., Aug. I9I7, Mille (N).

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The rays in this species are white, the disk corollas "yellow brown" or
"purple," according to Pennell & Hazen's labels.
27. DIPLOSTEPHIUM ERIOPHORUM Wedd. Chlor. And. I: 206. pl. 36, f. C.

RANGE: Departments of Tolima and Caldas, Colombia.
ADDITIONAL SPECIMEN EXAMINED: COLOMBIA: Shrub forest on slope,
P4ramo del Quindio, Cordillera Central, Dept. Caldas, alt. 4200-4400 m.,
Aug. I922, Pennell & Hazen 9908 (N).
The label describes the rays as white and the disk as purple.
28. DIPLOSTEPHIUM LECHLERI (Sch. Bip.) Wedd. Chlor. And. I: 204.

I857; Blake, Contr. U. S. Nat. Herb. 24: 8i. I922.
Liabum (Oligactis) lechieri Sch. Bip. Bonplandia 3: 236. I855.
RANGE: Known only from the type collection from near Sachapata,
Andes of Carabaya, Peru.
ADDITIONAL SPECIMEN EXAMINED: PERU: Sachapata, Prov. Carabaya,
Aug. I854, Lechler 2517 (type coll.; Par., photog. and fragm. N).
29. Diplostephium pulchrum Blake, sp. nov.

Branches thinly and closely silvery-tomentose and densely hirsute-
pilose with many-celled, spreading, yellowish-white hairs; leaves lance-
linear, about 5 cm. long, finely and evenly hispidulous above, thinly and
closely argenteous-tomentose beneath; heads large, in a terminal leafy-
bracted cyme; involucre about 14 mm. high; pappus purple.
"Small tree, 3.6 m. high," or "shrub, I.3-2 m. high," the branchlets
fastigiate, at length glabrate or glabrescent, 2-3 mm. thick; internodes I-5
mm. long; petioles broad, 1-2 mm. long; blades lance-linear or elliptic-
linear, 4-6.5 cm. long, 5-8 mm. wide, acuminately callous-tipped, truncat
rounded at base, usually only slightly revolute, entire, thin-coriaceous, above
deep green, shining, impressed-veined, the costa prominent and nearly
glabrous beneath, the lateral veins I5-25 pairs, delicate, spreading nearly at
a right angle, forked and anastomosing; heads about 2.5 cm. wide, in nearly
or quite simple cymes of 9-14 toward apex of branches, the peduncles I-7
cm. long, pubescent like the branches, bracted with linear strongly revolute-
margined leaves about 2 cm. long; disk in fruit I-1.2 cm. high; involucre
graduate, about 6-seriate, the phyllaries linear-subulate, o.8-i mm. wide,
attenuate, thinly tomentose, scarcely ciliate, with loose subcirrhous yellowish
tips; receptacle alveolate, the alveolae lacerate-fringed; rays "white,"
about 54, the tube puberulous toward apex, 2 mm. long, the lamina linear,
bidenticulate, 4-nerved, I5 mm. long, 1.5 mm. wide; disk corollas very
numerous, purple above, puberulous, 6.5 mm. lQng (tube 2 mm., throat
cylindric, 3.5 mm., teeth I mm.); achenes sparsely hirsutulous and glandu-
lar, 5-nerved, 2 mm. long; pappus deep purplish except at base, the inner of
bristles 5 mm. long, scarcely thickened apically, the outer setulose-squamel-
late, whitish, I mm. long; style branches with subulate, acuminate, dorsally
hispidulous appendages nearly or quite as long as the stigmatic region.
PERU: Wet mossy rocky open uplands, Tambo de Vaca, Dept. Huinuco,
alt. 3965 m., 10-24 June I923, Macbride 4346 (type no. 53543I, Field Mus.;

186.11.18.239 on Fri, 13 May 2022 16:51:05 UTC
duplicate no. I,19I,499, U. S. Nat. Herb.). Mufia, Dept. Huanuco, alt.

2745-3050 m., May I863, R. Pearce (K, fragm. N).
In addition to the characters mentioned in the key, the species differs
from D. lechieri in its involucre. In that plant the involucre is only 9-io
mm. high, and there are 3 or 4 outer series of triangular-ovate phyllaries
4 mm. long or less, while in D. pulchrum even the outermost phyllaries are
linear-subulate and 6-8 mm. long.
30. DIPLOSTEPHIUM HAENKEI (DC.) Wedd. Chlor. And. I: 203. I857;

Simblocline haenkei DC. Prodr. 5: 297. I836.
Diplostephium afline Wedd. Chlor. And. I: 203. I857; Blake, Contr.
U. S. Nat. Herb. 24: 86. I922.
Diplostephium mandonii Sch. Bip. Linnaea 34: 534. I865-66, and
Bull. Soc. Bot. France I2: 8I. I865, nomen nudum.
Aster sejaense Kuntze, Rev. Gen. Pl. 32: I3I. I898.
Diplostephium mandoni Rusby, Bull. N. Y. Bot. Gard. 4: 383. I907.
Diplostephium liabioides Rusby, Bull. N. Y. Bot. Gard. 4: 384. I907.
Diplostephium atropurpureum Rusby, Bull. N. Y. Bot. Gard. 4: 384.
I907.
Diplostephium sejaense Blake, Contr. U. S. Nat. Herb. 24: 8o. I922.
RANGE: Peru, Bolivia.
ADDITIONAL SPECIMENS EXAMINED: PERU: Without definite locality,
Haenke (type of S. haenkei, Prodr. Herb.). Cordillera of Carabaya, alt.
3000-3500 m., i847, Weddell 4740 (type of D. affine, Par.; photog. and
fragm. N). Rocky thicket, shrub zone, alt. 3500-3800 m., Paso de Tres
Cruces, Cerro de Cusilluyoc, Dept. Cuzco, I925, Pennell I3899 (F). BOLIVIA:
Near Sorata, Mandon 215 (chirotype of D. mandonii Sch. Bip., herb. Sch.
Bip.; fragm. N.).
Comparison of the types of D. haenkei and D. affine with material of D.
sejaense by the writer in 1925 disclosed no differences of any moment.
Buchtien's labels (nos. 3030, 303I) describe the plant as a shrub 2 meters
high, with violet flowers. The heads are rarely solitary.
3I. DIPLOSTEPHIUM PHYLICOIDES (H. B. K.) Wedd. Chlor. And. I: 205.

Aster phylicoides H. B. K. Nov. Gen. & Sp. 4: 93. I920.
Tetramolopium ? phylicoides DC. Prodr. 5: 262. I836.
Linochilus phylicoides Sch. Bip.; Wedd. Chlor. And. I: 205. I857, as
synonym.
Diplostephium umbelliferum Blake, Contr. U. S. Nat. Herb. 24: 80.
PI. 26. I922.
RANGE: Vicinity of Bogotd, Colombia.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Piramo de Choachi,
near BogotA, alt. 3700 m., 8 Aug. I922, Killip & Ariste-Joseph II926 (N).
5

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6o AMERICAN JOURNAL OF BOTANY [Vol. I5,
Bogota, Karsten (herb. Sch. Bip., fragm. N). Without definite locality
(" Nova Hispania"), Bonpland (type of A. phylicoides, herb. Humb. &
Bonpl.).
This species is distinguished by its small, thick, elliptic or oblong leaves
and umbelliform clusters of small heads. The flowers are described as
white in no. II926; those of other collections have been described as
purple blue. The characters supposed to distinguish D. umbelliferum from
D. phylicoides are of no consequence. The style branches tend to adhere,
and the supposed difference in shape was evidently due principally to the
degree to which they were separated in dissecting.
32. DIPLOSTEPHIUM SCHULTZII Wedd. Chlor. And. I: 204. I857; Blake,

Contr. U. S. Nat. Herb. 24: 83. 1922.
Linochilus iodopcappus (jodopappus) Sch. Bip.; Wedd. Chlor. And. I:

204. I857, as synonym.
RANGE: Colombia.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Volcan de Tolima, alt.
4025 m., Jan. I843, Linden 90I (type collection; Par., herb. Sch. Bip.,
photog. and fragm. N). Along stream, Paramo del Quindio, Cordillera
Central, Dept. Caldas, alt. 4100-4400 m., Aug. I922, Pennell & Hazen 9825
(G, N.). Shrub-zone, below paramo, Cerro Tatama, Cordillera Occidental,
Dept. Caldas, alt. 3300-3500 m., Sept. I922, Pennell I0532 (G, N). Without
definite locality, Triana go (G).
This species and the next, which are very closely allied, are most closely
related to the group consisting of D. lehmannianum, D. rhododendroides, and
D. pleistogynum, and are distinguished principally by the persistent to-
mentum of the upper leaf surface. Linden's plant is described as purple-
flowered. In one of Pennell's plants the disk is described as red, and the
rays pink; in the other the rays are said to be blackish-violet, although they
appear to have been of the same color as in the first specimen. The pappus
and disk corollas become deep purple at maturity. In a head of no. 9825,
many of the alveolar margins of the receptacle were prolonged into slender
smooth setae up to I mm. long. Reexamination of Triana 90 shows that it
belongs to D. schultzii and not to D. cochense, to which I previously referred
it.
A head of the type collection dissected by the writer contained about 44
rays and 6 disk flowers. The two collections by Pennell, which are certainly
specifically identical with each other and with the type, differ remarkably
in the composition of the heads. In one (no. 9825) two heads dissected, from
different specimens, contained in one case 37 rays and 4 disk flowers, and in
the other IO rays and 32 disk flowers. Four heads of no. I0532, very
carefully dissected from sheets in two different herbaria, showed respecti
38, 32, 3I, and 27 rays, and no disk flowers whatever! The heads appeared
normal, the rays were normally formed, although often shorter than the

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Jan., I928] BLAKE - DIPLOSTEPHIUM 6I
styles, and none showed any sign of stamens; the mature achenes were
normal in appearance, but their embryos were abortive. This collection
apparently represents a remarkable mutation, of a sort I have never before
met with in Compositae. Its significance is discussed in the introduction to
this paper.
33. DIPLOSTEPHIUM COCHENSE Hieron. Bot. Jahrb. Engler 21: 341. I895;
RANGE: Colombia.
SPECIMENS EXAMINED: COLOMBIA: Llafiura del Rio Cocha, Aug. I869,
Stuebel 353 (type, fragm. N.).
Triana go, doubtfully referred to this species in my former paper, prov
on reexamination to belong to D. schultzii.
34. DIPLOSTEPHIUM COSTARICENSE Blake, Contr. U. S. Nat. Herb. 24: 82.

PI. 27. I922.
RANGE: Mountains and paramos of Costa Rica, altitude I500-3100
meters.
ADDITIONAL SPECIMENS EXAMINED: COSTA RICA: Common, wet paramo
thicket, Cerro de las Vueltas, Prov. San Jose. alt. 2700-3000 m., Dec. I925-
Jan. 1926, Standley & Valerio 43722, 43809 (N).
Described as a dense gray shrub 3-4.5 m. high, with white rays and dark
purple-red disk. The heads in this species are 22-57-flowered, with i6-27
rays and 6-36 disk flowers. In no. 43722 the ray corollas are normally
formed but very short (the lamina I.7-2 mm. long), equaling or shorter than
the styles.
35. DIPLOSTEPHIUM OBTUSUM Blake, Contr. U. S. Nat. Herb. 24: 84.

I922.
RANGE: Paramos of the State of Trujillo, Venezuela, alt. 2800-3200
meters.
36. DIPLOSTEPHIUM LEHMANNIANUM Hieron. Bot. Jahrb. Engler 21: 340.

Diplostephium schultzii var. lehmanniana Hieron. Bot. Jahrb. Engler I9:
48. I 894.
SPECIMENS EXAMINED: COLOMBIA: Highest shrub-zone, Paramo de
Guanacas, Prov. Popayan, alt. 3000-3500 m., Jan., Lehmann 4893 (type,
fragm. N). ECUADOR: Eastern andine region, Pifo, near Paluguillo, Prov.
Pichincha, alt. 3800 m., I898, Mille (N).
Distinguished, among the species with small leaves glabrous above, by its
small heads and involucre. Mille's plant is described as a very handsome
shrub i.5 meters high.

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62 AMERICAN JOURNAL OF BOTANY [VO1. I 5,
37. DIPLOSTEPHIUM RHODODENDROIDES Hieron. Bot. Jahrb. Engler 2I: 340.

I895; Blake, Contr. U. S. Nat. Herb. 24: 8i. 1922.
RANGE: Known only from the type collection, from Azufral de Tuquerres,
Colombia.
SPECIMEN EXAMINED: COLOMBIA: Laguna Verde, near Azufral de
Tuquerres, Jan. I870, Stuebel 429 (type, fragm. N).
38. DIPLOSTEPHIUM PLEISTOGYNuM Blake, Contr. U. S. Nat. Herb. 24: 82.

1922.
RANGE: Known only from the type collection, from PAramo de Buena
Vista, Huila group, Cordillera Central, Dept. Cauca, Colombia.
39. DIPLOSTEPHIUM FLORIBUNDUM (Benth.) Wedd. Chlor. And. I: 205. pl.

36,f.B. I857.
Linochilusfloribundus Benth. P1. Hartw. 203. I845.
Linochilus ochraceus Sch. Bip.; Wedd. Chlor. And. I: 205. I857, as
synonym.
Diplostephium ochroleucum Klatt, Bot. Jahrb. Engler 8: 37. I886.
Aster ochroleucus Kuntze, Rev. Gen. P1. 3 2: 131. I898.
RANGE: Departments of Popayin and Cauca, Colombia.
ADDITIONAL SPECIMENS EXAMINED: COLOMBIA: Moist thicket in ptramo
("llano"), PaletarA, Cordillera Central, Dept. Cauca, alt. 2950-3IO0 m.,
June I922, Pennell 6969 (N). Shrub-zone ("paramillo"), Mt. El Derrumbo,
Cordillera Occidental, Dept. Cauca, alt. 2700-3000 m., June I922, Pennell
7489 (N).
40. Diplostephium leiocladum Blake, sp. nov.
Branches, peduncles, and pedicels quickly glabrate; leaves elliptic or
lance-elliptic, about 6 cm. long, repand-toothed or entire, glabrous above,
whitish-tomentose beneath, slender-petioled; heads rather small, numerous,
cymose-panicled; involucre about 7 mm. high.
Branches stout, 3-5 mm. thick, ridged by lines decurrent from the leaf
bases, at first thinly but closely whitish-tomentose, soon nearly or quite
glabrate, very leafy; internodes 2-4 mm. long; petioles slender, glabrous or
quickly glabrate, very narrowly margined essentially to base, 8-io mm.
long; blades 4.5-7.5 cm. long, I.2-1.8 cm. wide, acute, at base acuminate,
repand-serrate (teeth acutish, about 3 mm. apart, 0.5-I mm. high) or entire,
coriaceous, narrowly revolute-margined, above deep green, shining, glabrous,
beneath densely and closely whitish- or slightly ochroleucous-tomentose, the
costa glabrous and prominent beneath, impressed above, the principal lateral
veins 7-I2 pairs, spreading at nearly a right angle, slender, prominulous,
forked and anastomosing, usually becoming glabrate beneath, impressed
above; heads about I3-I5-flowered, in small pedunculate cymes in the axils
of the uppermost leaves, forming a dense rounded panicle about 7 cm. wide,
about equalled by the leaves, the peduncles pubescent like the stem and
similarly glabrate, about i.5 cm. long, the pedicels 3-7 mm. long; involucre
narrowly campanulate, strongly graduate, about 5-seriate, the phyllaries
ovate (outer) to linear-oblong or lance-linear, acute (outer) to obtuse,

186.11.18.239 on Fri, 13 May 2022 16:51:05 UTC
indurate, obscurely greenish toward apex, scarious-margined, ciliolate and

toward apex persistently tomentose; receptacle shallowly alveolate; rays
about 5-8, "white," the tube sparsely puberulous, 4.5 mm. long, the lamina
spreading, elliptic-linear, bidenticulate, 3-3.5 mm. long, o.6 mm. wide; disk
corollas about 8-io, whitish, densely puberulous on slender part of throat,
the tube and greater part of throat isodiametric, 4 mm. long, the upper part
of throat abruptly dilated for 0.5 mm., the lance-ovate teeth I.5 mm. long;
achenes glabrous or sparsely sessile-glandular above, 5-nerved, I.8-2.5 mm.
long, those of disk apparently infertile; pappus brownish white, 5 mm. long,
the longer bristles slightly dilated at apex, the outer pappus sparse, setulose,
I mm. long or less; style branches linear-oblong, hispidulous throughout,
with short acute or subacuminate appendages.
COLOMBIA: Shrub zone, below paramo, Cerro Tatama, Cordillera
Occidental, Dept. Caldas, alt. 3300-3500 m., 8-io Sept. 1922, Pennell
I053I (type no. 1,I41,3I6, U. S. Nat. Herb.); also Pennell 10530, on edge of
paramo, same data.
Closely related to D. floribundum, but distinguished by its quickly
glabrate peduncles and branches, as well as by its larger leaves. The leaves
in no. I053I are conspicuously toothed, in no. I0530 entire, but the two
collections are clearly forms of one species.
41. DIPLOSTEPHIUM DENTICULATUM Blake, Contr. Gray Herb. 53: 25.

I9I8; Contr. U. S. Nat. Herb. 24: 85. I922.
RANGE: Known only from the type locality, Guadalupe, Colombia,
altitude 3000 meters.
Distinguished from the two following species by the conspicuous
reticulation of the lower leaf surface and the pubescent achenes.
42. DIPLOSTEPHIUM OCHRACEUM (H. B. K.) Nees, Gen. & Sp. Ast. 201.
Aster ochraceus H. B. K. Nov. Gen. & Sp. 4: 85. I820.
Tetramolopium ? ochraceum DC. Prodr. 5: 262. I836.
RANGE: Known only from the type, collected "in montibus Quitensi-
bus? "
SPECIMEN EXAMINED: ECUADOR (?): Without definite locality, Bonpland
(type, herb. Humb. & Bonpl.; photog. N).
43. DIPLOSTEPHIUM BICOLOR Blake, Contr. U. S. Nat. Herb. 24: 85.

pI. 28. I922.
RANGE: Known only from the type locality, headwaters of the Rio
Lopez, Rio Palo Basin, Tierra Adentro, Cauca, Colombia, altitude 2500-
3000 meters.
Closely allied to D. ochraceum, and apparently distinguished principally
by its more or less obovate, rather thicker leaves, with denser tomentum.

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64 AMERICAN JOURNAL OF BOTANY LVo1. I5,
EXCLUDED SPECIES 3
DIPLOSTEPHIUM CORYMBOSUM Donn. Smith, Bot. Gaz. 23: 8. 1897;

This is ARCHIBACCHARIS CORYMBOSA (Donn. Smith) Blake, Journ.
Washington Acad. Sci. I7: 6o. I927 (Hemibaccharis corymbosa Blake,
Contr. U. S. Nat. Herb. 20: 553. I924).
DIPLOSTEPHIUM INCANUM Hieron. Bot. Jahrb. Engler 2I: 340. I895;

This is a species of the tribe Senecioneae, belonging to Senecio or a
related genus. The fragments of the type examined are insufficient to
enable the genus to be determined.
DIPLOSTEPHIUM PANICULATUM Donn. Smith, Bot. Gaz. 23: 8. I897;

Blake, Contr. U. S. Nat. Herb. 24: 86. 1922.
This is Archibaccharis mucronata var. paniculata (Donn. Smith) Blake
(Hemibaccharis mucronata paniculato Blake, Contr. U. S. Nat. Herb. 20: 551.
1924; Archibaccharis mucronata paniculata Blake in Standl. Contr. U. S.
Nat. Herb. 23: I509. I926).
8 Including only species not disposed of in my I922 revision.

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Review of The Genus Diplostephium

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Review of the Genus Diplostephium

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(Received for publication May 28, I927)

In 1922 the writer' published a revision of Diplostephium, a genus

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In my earlier paper it was suggested that the primitive type of style in

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Heads solitary at tips of stem and branches . ........................ I. Lavandulifolia.

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Leaves densely and conspicuously impressed-glandular over whole upper surface.

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Leaf blades glandular-tomentose above at least when young, narrowly

Branchlets glabrous, more or less viscid ............................. 20. D. revolutum.

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Leaves hispidulous on surface above; branches densely hirsute-pilose with yellowish

SERIES I. LAVANDULIFOLIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

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i. Diplostephium micradenium Blake, sp. nov.

2. DIPLOSTEPHIUM EMPETRIFOLIUM Blake, Contr. U. S. Nat. Herb. 24: 73.

3. DIPLOSTEPHIUM GLANDULOSUM Hieron. Bot. Jahrb. Engler 2 1: 339.

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TYPE LOCALITY: PAramo de la Boca del Mundo Nuevo, VolcAn de

4. DIPLOSTEPHIUM OBLANCEOLATUM Blake, Contr. U. S. Nat. Herb. 24: 76.

5. DIPLOSTEPHIUM MACROCEPHALUM Blake, Contr. U. S. Nat. Herb. 24: 75.

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entire lack of impressed glands on the leaves, which are conspicuously

6. Diplostephium pearcei Sprague & Blake, sp. nov.

7. Diplostephium callilepis Blake, sp. nov.

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body o.6-o.8 mm. wide, purplish-brown, its deeply lacerate-ciliate, brownish-

8. Diplostephium hippophae Blake, sp. nov.

9. DIPLOSTEPHIUM ANACTINOTUM Wedd. Chlor. And. I: 20I. pI. 35, f. B.

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ADDITIONAL SPECIMEN EXAMINED: COLOMBIA: Piramo, San Miguel,

IO. DIPLOSTEPHIUM PARVIFOLIUM Blake, Contr. U. S. Nat. Herb. 24: 74.

Diplostephium microphyllum Wedd. Chlor. And. I: 20I. I857. Not D.

II. DIPLOSTEPHIUM SPINULOSUM Wedd. Chlor. And. I: 200. I857.

12. DIPLOSTEPHIUM ADENACHAENIUM Blake, Contr. U. S. Nat. Herb. 24: 72.

RANGE: Colombia, Ecuador.

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13. DIPLOSTEPHIUM MEYENII Wedd. Chlor. And. I: 20I. I857; Blake,

I4. DIPLOSTEPHIUM TACORENSE Hieron. Bot. Jahrb. Engler 2I: 337.

i6. Diplostephium gnidioides Blake, sp. nov.

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callous-pointed, short-petioled, spinulose above, usually so strongly revolute-

17. DIPLOSTEPHIUM LAVANDULIFOLIUM H. B. K. Nov. Gen. & Sp. 4: 97.

I8. DIPLOSTEPHIUM HARTWEGII Hieron. Bot. Jahrb. Engler 21: 337.

I9. DIPLOSTEPHIUM ANTISANENSE Hieron. Bot. Jahrb. Engler 21: 338.

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SPECIMENS EXAMINED: ECUADOR: Paramos of Cerro Antisana, near Las

SERIES 2. ROSMARINIFOLIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

20. DIPLOSTEPHIUM REVOLUTUM Blake, Contr. U. S. Nat. Herb. 24: 78.

RANGE: Vicinity of Bogota, Colombia.

2I. DIPLOSTEPHIUM CARABAYENSE Wedd. Chlor. And. I: 202. I857;

22. DIPLOSTEPHIUM CYPARISSIAS Wedd. Chlor. And. I: 203. I857; Blake,

23. DIPLOSTEPHIUM BACCHARIDEUM Blake, Contr. U. S. Nat. Herb. 24: 77.

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24. DIPLOSTEPHIUM ROSMARINIFOLIUM (Benth.) Wedd. Chlor. And. I: 202.

SERIES 3. RUPESTRIA Blake, Contr. U. S. Nat. Herb. 24: 69. I922.

25. DIPLOSTEPHIUM WEDDELLII Blake, Contr. U. S. Nat. Herb. 24: 79.

Diplostephium sessiliflorum Wedd. Chlor. And. I: 204. I857. Not D.

26. DIPLOSTEPHIUM RUPESTRE (H. B. K.) Wedd. Chlor. And. I: 206.