The North American Species of Cactus, Anhalonium, and Lophophora
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John M. Coulter >> The North American Species of Cactus, Anhalonium, and Lophophora
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A Preliminary Revision of the North American Species of Cactus,
Anhalonium, and Lophophora by John M. Coulter.
U. S. Department of Agriculture
Division of Botany
CONTRIBUTIONS FROM THE U. S. NATIONAL HERBARIUM
Vol. III--No. 2
Issued June 10, 1894
Preliminary Revision of the North American Species of Cactus,
Anhalonium, and Lophophora.
by
John M. Coulter.
Published by Authority of the Secretary of Agriculture
Washington
Government Printing Office
1894
LETTER OF TRANSMITTAL
U. S. Department of Agriculture
Division of Botany
Washington, D. C., March 21, 1894
SIR: I have the honor to transmit herewith, for publication as
Vol. III, No. 2, of Contributions from the U. S. National
Herbarium, a Preliminary Revision of the North American species
of Cactus, Anhalonium, and Lophophora, by President John M.
Coulter.
Respectfully,
Frederick V. Coville,
Chief of the Division of Botany.
Hon. J. Sterling Morton,
Secretary of Agriculture.
PRELIMINARY REVISION OF THE NORTH AMERICAN SPECIES
OF CACTUS, ANHALONIUM, AND LOPHOPHORA.
Prefatory Note.
In the fall of 1890 Dr. George Vasey, then Botanist of the
Department of Agriculture, arranged with me to prepare a revision
of North American Cactaceae. Owing to the peculiar difficulty of
preserving material the family was poorly represented, even in
our leading herbaria. To secure a large amount of additional
material in the way of specimens and field notes the Department
authorized me to visit the region of the Mexican boundary during
the summer of 1891. Preliminary to this exploration it was
necessary to examine the Engelmann collection of Cactaceae, in
the possession of the Missouri Botanical Garden. This
collection, supplemented by the continual additions made at the
garden, is by far the largest collection of skeletons and living
specimens in this country, and also contains the large majority
of our types.
In March, 1891, I visited this collection and made such notes as
seemed necessary for use in the field, and in June, accompanied
by Mr. W. H. Evans and Mr. G. C. Nealley, I began field work in
the neighborhood of El Paso, Tex. After ten days of exploration
it was necessary for me to leave the field work in charge of Mr.
Evans, who, with Mr. Nealley, continued work westward, during
July and a part of August, to southern California, along the
Southern Pacific Railway. As a result a large number of complete
plant bodies was secured, but very few of them were in flower and
the field notes indicated little besides collection stations.
During the following fall and winter preliminary determinations
of this material were made by Mr. Evans.
In the fall of 1892 critical study of this and other collections
was begun in connection with my assistants, Dr. Elmon M. Fisher
and Mr. Edwin B. Uline, who have ever since rendered constant and
most import assistance in the examination of material and
bibliography, which alone has made the work possible in the midst
of other pressing duties.
In the spring of 1893 these two gentlemen spent several weeks at
the Missouri Botanical Garden in the critical study of its rich
material, and during the latter part of their stay I assisted in
the work. Dr. William Trelease, the director of the garden, had
hastened the arrangement of the Engelmann material, and had
mounted in convenient form the large mass of notes left by Dr.
Engelmann. These notes contained not only critical remarks upon
known species, but also the diagnoses of many unpublished species
which had come into his hands, notably those collected by Mr.
William Gabb in 1867 in Lower California. The collections that
have thus far been studied are:
(1) Those of the Missouri Botanical Garden; and thanks are
especially due to Dr. Trelease for his generous cooperation in
the use of this material, without which the work would have been
impossible.
(2) Those of the Department of Agriculture, including the results
of several recent explorations, for the use of which I am
indebted to Mr. Frederick V. Coville.
(3) Those of the Gray Herbarium at Harvard University, which Dr.
B. L. Robinson kindly placed at my disposal.
(4) Those of the California Academy of Sciences, notably rich in
forms from Lower California and the adjacent islands, kindly
loaned by Mr. T. S. Brandegee.
(5) Those of Dr. Louis Eschanzier, of San Luis Potosi, Mexico,
who send a large series of Mexican forms collected in 1891.
(6) Numerous small sets from different correspondents, who have
given both time and material in aiding the work.
It is needless to say that Dr. George Engelmann, the great
pioneer student of this difficult family, has opened the paths in
which we must follow, and it was exceedingly unfortunate that he
was not able to complete the final revision that he had in mind.
The difficulties which beset the critical study of this group can
not be easily exaggerated. Such scanty material as has been
collected has been for the most part very incomplete, consisting
of plant bodies without flower or fruit, flower or fruit without
plant bodies, and bunches of spines without either. The species
are displayed also in the most inaccessible regions, and their
culmination is found in the still poorly known regions of Mexico.
On account of their singular forms and often brilliant flowers
they have long been extensively cultivated, especially in Europe.
These cultivated forms have formed the basis of original
descriptions in almost all of the European publications, and in
very rare cases have any types been preserved. As a result, the
bibliography of Cactaceae is appalling, and it is questionable
whether satisfactory conclusions can be reached in the case of
hundreds of published names. The earlier descriptions were not
only meager, but were based upon what are now regarded very
insufficient characters, and in the absence of types it is not
only unsafe, but impossible to venture an opinion concerning
their identity. In view of these facts, I have thought it
advisable to present a preliminary revision of the order, which
shall contain the results of the study of material confessedly
insufficient. With such knowledge as we possess brought
together, it is hoped that the study of this very interesting and
much neglected group will be stimulated, and that more critical
exploration of our southwestern territory and adjacent Mexico
will make a more satisfactory presentation possible. It would be
useless to notice the vast number of reputed species that are not
represented by actual specimens in our possession.
In the proposed preliminary account of the family, of which the
present paper is the first part, only those genera are considered
which form a part of the flora of the United States, and those
species which I have been able to examine and to identify with
reasonable certainty. All forms credited to the United States
have been studied, and the account of these species may be
considered fairly complete, but the far more numerous Mexican
species are but scantily represented. The Mexican boundary is so
unnatural a dividing line in the distribution of Cactaceae that
it has been disregarded, and all the species studied have been
arranged in a lineal series of uniform prominence. So far as
known the subject of geographical distribution is considered, but
it will be seen how meager is our knowledge of this subject. It
is to be hoped that this preliminary presentation will provoke
exploration and study, and that species will not only be
collected, but all the facts of their distribution noted. It is
more than probable that our present notion of species in this
group must be much modified, and doubtless many forms are at
present kept specifically distinct which will prove to be but
different phases of a single species.
In the matter of generic delimitation we are in still greater
uncertainty, and several generic lines at present recognized must
be regarded as purely arbitrary, a fact which must become still
more evident with additional material. The whole group is to be
regarded as made up of poorly differentiated forms and only long
observation under cultivation can determine the possibilities of
specific variation under the influence of environment, of age, of
inherent tendencies. For instance, that these plants change in
form and in spine characters with increasing age and after they
have begun to flower can not be doubted, but what described forms
have thus been separated in descriptions can only be guessed at.
John M. Coulter.
Lake Forest University,
Lake Forest, Ill., January, 1891.
CACTUS, ANHALONIUM, AND LOPHOPHORA.
1. CACTUS Linn. Sp. Pl. 466 (1753), restricted.
MAMILLARIA Haw. Synop. 177 (1812), not Stackh. (1809).
Usually globose to oblong plants (simple, branching or
cespitose), but sometimes slender-cylindrical, covered with
spine-bearing tubercles: flower-bearing areola axillary (with
reference to tubercles), entirely separate from the terminal
spine-bearing areola, although sometimes (Coryphantha) connected
with it by a woolly groove along the upper face of the tubercle:
ovary naked: seeds smooth or pitted: embryo usually straight,
with short cotyledons. Originally defined by Linnaeus in his
Systema, ed. l (1735).
The Linnaean genus Cactus of 1753 included 22 species and was
coextensive with the present order. In 1812 the species were
separated by Haworth into five genera, the original generic name
Cactus being discarded. Among these species C. mamillaris seems
to have stood as the type, not only of the Linnaean genus Cactus,
but also of Haworth's Mamillaria, and as such should retain the
original generic name. Besides, Mamillaria was used as the
generic name of an alga in 1809. Cactus mamillaris L. is the
West Indian Mamillaria simplex Haw.
From one point of view the two sections of the genus
(Eumamillaria and Coryphantha) deserve generic separation, for
the character of grooveless and grooved tubercles seems to hold
without exception, and the sections are separated with more
certainty than are certain species of Coryphantha and
Echinocactus. If genera are simply groups of convenience the
separation should be made.
I. EUMAMILLARIA. Flowers from the axils of the older or
full-grown tubercles (hence usually appearing lateral), mostly
small, and generally from whitish to pink or red: tubercles never
grooved: fruit almost always clavate and scarlet.
A. Tubercles more or less quadrangular.
* Central spines not hooked.
+ More than one central spine.
1. Cactus alternatus, sp. nov.
Subglobose, 10 cm. in diameter, simple: tubercles long (15 to 20
mm.) and spreading, with woolly axils: radial spines 3, rigid and
recurved, 5 mm. long; central spines 3, very stout and much
recurved, 20 to 30 mm. long, alternating with the radials; all
ashy colored and often twisted: flower and fruit unknown.--Type
in Herb. Coulter.
The few spines, with the very short radials alternating with the
very long and stout centrals, furnish a striking character.
Occasionally one of the centrals is wanting.
2.Cactus acanthophlegmus (Lehm.) Kuntze, Rev. Gen. Pl. 260
(1891).
Mamillaria acanthophlegma Lehm. Delect. Sem. Hamb. (1833)
Subglobose with a deeply depressed vertex, or becoming
cylindrical, 3 to 8.5 cm. in diameter: tubercles sharply
quadrangular-conical, with densely woolly axils: radial spines 15
to 30, white, very slender (bristly) and radiant, sometimes
coarse capillary, 4 to 7 mm. long, interwoven with those of
neighboring tubercles and so covering the whole plant; central
spines 2 to 4, robust and straight, erect or divergent, whitish
or reddish, black-tipped, 5 to 6.5 mm. long: flowers reddish, 1
to 2 cm. broad: fruit unknown. Type unknown.
From Coahuila and San Luis Potosi to Oaxaca. Fl. May.
Specimens examined: Coahuila (Poselger of 1856; Pringle 3116 of
1890): San Luis Potosi (Eschanzier of 1891).
The central spines are quite variable in number and arrangement.
In case there are two they are vertically placed and are either
erect and parallel or widely divergent. Even three centrals may
occur in the same vertical plane; but more usually the three or
four centrals are arranged about a center and are widely
divergent. The tubercles are apt to persist and to become naked
and corky with age. The axillary wool and the capillary radials
are also apt to be more or less persistent, thus giving the whole
plant a woolly appearance.
3.Cactus brandegei, sp. nov.
Cylindrical: tubercles sharply quadrangular-conical, 6 to 8 mm.
long, with densely woolly axils: radial spines about 10, slender
and rigid, whitish with dusky tips, spreading but not radiant, 7
to 10 mm. long; central spines 3 or 4, stouter and slightly
longer, erect-spreading (sometimes slightly curved),
reddish-brown below, becoming blackish above: flowers small
(scarcely longer than the tubercle?): fruit unknown. Type in
Herb. Calif. Acad.
San Jorge, Lower California. Fl. April.
Specimens examined: Lower California (Brandegee of 1889, at San
Jorge).
The species has somewhat the spine characters of C. palmeri, but
the sharply quadrangular and longer tubercles with axillary wool
free from bristles suggest a very different affinity.
4.Cactus densispinus, sp. nov.
Globose, 7.5 cm. in diameter, simple: tubercles short, with
woolly axils: radial spines about 25, erect-spreading, slender
but rigid, yellow (brownish to black with age), unequal, 8 to 10
mm. long; central spines 6, a little longer (10 to 12 mm.) and
straight, more rigid and darker, black-tipped: seeds obovate,
reddish-brown, 1 mm. long. Type in Herb. Coulter.
Very easily distinguished by its dense, erect spines, which so
completely cover the plant as to give it the appearance of a
large chestnut bur. Another much smaller form, which seems to be
a variety, has stouter and longer ashy-white spines, the centrals
darker-tipped, and the lower centrals slightly curved.
++ One short central spine (rarely two or none): ovaries
immersed: seeds small, yellow and rugulose: simple.
5.Cactus heyderi (Muhlenpf.) Kuntze, Rev. Gen. Pl. 260 (1891).
Mamillaria heyderi Muhlenpf. Allg. Gart. Zeit. xvi. 20(1848).
Mamillaria declivis Dietr. Allg. Gart. Zeit. xviii. 235
(1850).
Mamillaria applanata Engelm. Pl. Lindh 198 (1850).
Mamillaria texensis Labouret, Monogr. Cact. 89 (1858).
Depressed, globose, usually with depressed vertex, 8 to 12 cm.
broad, 2.5 to 5 cm. high: tubercles elongated: radial spines 10
to 22, whitish, 5 to 12 mm. long, the lower usually the longer,
stouter, and often darker; central spine 4 to 8 mm. long, light
yellowish-brown, stout, straight, and porrect: flowers 2 to 2.5
cm. long, reddish-white: fruit incurved, 1.5 to 3 cm. long. (Ill.
Cact. Mex. Bound. t. 9. figs. 4-14). Type unknown.
From the Guadalupe River, Texas, to the mouth of the Rio Grande,
and westward to Arizona and Sonora. Fl. April, May.
Specimens examined: Texas (Lindheimer of 1845, 1847, 1853; Wright
226, also collections of 1849, 1852, 1853, 1855, 1856; Bigelow of
1853; Trelease of 1892; Nealley of 1892): New Mexico (Wright 311;
Bigelow of 1853, Evans of 1891): Arizona (Pringle of 1881): also
growing in Mo. Bot. Gard. 1893; and in the World's Fair
collection of Mrs. Nickels.
The radial spines are somewhat variable in relative length, often
becoming almost equal, while sometimes the upper radials are very
much reduced. The figure referred to in Cact. Mex. Bound. is not
satisfactory as to the general habit of the plant, which is
flat-topped rather than hemispherical.
6.Cactus heyderi hemisphaericus (Engelm.).
Mamillaria hemisphaerica Engelm. Pl. Lindh. 198 (1850).
Differs in being hemispherical instead of flat-topped, in its
fewer (9 to 12) and shorter (4 to 8 mm.) radial spines, and much
smaller less rough and lighter-colored seeds. (Ill. Cact. Mex.
Bound. t. 9. figs. 15-17) Type, the "Goebel's Garden" plants in
Herb. Mo. Bot. Gard.
Throughout southern Texas and southern New Mexico, and southward;
not extending so far north or west as the species, and apparently
not so abundant within the United States. Fl. May.
Specimens examined: Texas(Schott 322, 614): New Mexico (Evans of
1891): also specimens cultivated in the Goebel Garden, St. Louis,
in 1847, brought from "below Matamoras on the Rio Grande" by the
St. Louis Volunteers, in 1816.
On account of its convex top the variety becomes somewhat higher
than the species (5 to 7.5 cm.), and the flowers are sometimes
slightly longer (2 to 3 cm.).
7.Cactus meiacanthus (Engelm.) Kuntze, Rev. Gen. Pl. 260
(1891).
Mamillaria meiacantha Engelm. Syn. Cact. 263 (1856)
Hemispherical or with depressed vertex, 7.5 to 12.5 cm. in
diameter, with a broad top-shaped base: tubercles compressed, 14
to 18 mm. long: radial spines 5 to 9 (usually about 6), stout
and strongly subulate, 6 to 10 mm. long, straight or somewhat
curved, whitish or yellowish, the lower mostly a little longer,
the upper one sometimes wanting; central spine shorter and stout,
darker, straight, and porrect, turned upwards among the radials,
or rarely wanting: flowers 2.5 to 3 cm. long, reddish-white:
fruit incurved, 2 to 3 cm. long. (Ill. Cact. Mex. Bound. t.
9, figs. 1-3). Type specimens are those of the collections of
1847, 1851, 1852, and 1853, from which the original description
was drawn and all of which are in Herb. Mo. Bot. Gard.
From the Guadalupe River, Texas, to the "Great Bend" of the Rio
Grande, westward through western Texas and New Mexico; also
northern Mexico (Hemsley); Fl. May, June.
Specimens examined: Texas (Wright of 1851, 1852; Bigelow of
1853): New Mexico ("Missouri Volunteers" of 1847; unknown
collector in 1880); also specimens cultivated in St. Louis in
1853, and others growing in Mo. Bot. Gard. 1893.
Dr. Engelmann regarded this species as possibly only a variety
of C. heyderi, to which it is certainly very closely allied
through var. hemisphaerica, but the different tubercles and
fewer stouter spines serve so well to distinguish it that it
seems best to retain its specific rank.
In reference to the citation of the original description an
explanation seems necessary, which will apply to numerous similar
cases. The Pacif. R. Rep. iv. 27 (1856), Syn. Cact. 263
(1858), and Cact. Mex. Bound. 9 (1859), have each been cited
as the original publication. The confusion has arisen from the
fact that in both the publications of 1856 the description in the
Rep. Mex. Bound. is referred to, and in that report the plant
is fully described as "sp. nov." However, the publication of
the Boundary Report was long delayed on account of the
preparation of the plates, and in the meantime both the
publications of 1856 had appeared, in each one of which the
species is distinctly characterized and reference made to the
description in the forthcoming Boundary Report. As between the
two publications of 1856 the Syn. Cact. (Proc. Amer. Acad.
iii. 259) was evidently distributed first.
8.Cactus gummiferus (Engelm.) Kuntze. Rev. Gen. Pl. 260
(1891).
Mamillaria gummifera Engelm. Wisliz. Rep. 21 (1848).
Hemispherical, 7.5 to 12.5 cm. broad and 6 to 10 cm. high:
tubercles 12 to 15 mm. long: radial spines 10 to 12, the lower
stout, with dusky apex, 12 to 15 mm. long, twice or thrice as
long as the whitish setaceous upper ones; central spine
(sometimes two) shorter (about 4 mm.), stout, dusky and porrect:
flowers 3 cm. long, reddish-white, brownish-red outside: fruit
unknown. (Ill. Cact. Mex. Bound. t. 9. figs. 18-20) Type
probably lost, as no specimens could be found in the Engelmann
Herbarium.
Chihuahua, near Cosihuiriachi.
So far as can be discovered, this species has not been collected
since the original Wislizenus collection of 1846-47. The plants
were cultivated by Dr. Engelmann and made to bloom, showing the
flowers to be larger and darker colored than in the rest of the
group, from which the species also differs in its more robust
habit, its very unequal radial spines, and the occasional
occurrence of two centrals.
** Central spine hooked.
9.Cactus uncinatus (Zucc) Kuntze, Rev. Gen. Pl. 261 (1591).
Mamillaria uncinata Zucc. in Pfeiff. Enum. 34 (1837).
Mamillaria bihamata Pfeiff. in Otto and Deitr. Gart. vi. 274 (1840)
Mamillaria adunca Scheidw. (1845-1849?).
Mamillaria depressa Scheidw. (1845-1849?).
Usually globose (occasionally depressed or even subcolumnar), 5
to 6 cm. in diameter (doubtless becoming larger): tubercles 8
to 10 mm. long, woolly in the upper axils: radial spines 4 to
6, rigid, 4 to 6 mm. long, the upper one stouter than the rest
and sometimes shorter, reddish-brown and horny, straight or
slightly curved, the remainder straight and white with dusky
tips; central spine stout and horny, reddish-brown, 7 to 10 mm.
long: flowers greenish-white or tinged with red: fruit unknown
Type unknown.
Entirely Mexican, reported from Chihuahua to Saint Luis Potosi.
Specimens examined: San Luis Potosi (Gregg of 1848; Parry 268;
Eschanzier of 1891): Chihuahua (Wislizenus of 1846-47; also
Chihuahua specimens cultivated in the Jacoby Garden in 1856 and
1857).
The variations observed in this species do not seem sufficient
for the establishment of varieties. The type form seems to have
been globose, with 4 radial spines and a stout central one. The
depressed forms with 6 radials and a more slender central
represent var. spinosior Lem. (M. depressa Scheidw.); and the
subcolumnar forms with 6 radials (the upper one of which is
somewhat curved) and a stout strongly hooked central represent
var biuncinata Lem. (M. bihamata Pfeiff.) Such combinations of
characters, however, do not hold, as any one of the plant body
forms may display any one of the spine characters referred to.
B. Tubercles terete.
* Central spines none: mostly simple globose plants, with very
numerous straight whitish setaceous radials.
10. Cactus lasiacanthus (Engelm.) Kuntze, Rev. Gen. Pl. 259
(1891).
Mamillaria lasiacantha Engelm. Syn. Cact. 261 (1856).
Globose or ovate globose, 2 to 2.5 cm. high and 1 to 2 cm. broad:
tubercles 4 mm. long, about 2 mm. in diameter, with naked axils:
spines 40 to 60, in many series, very unequal, 2 to 4 mm, long,
white and pilose, the upper exterior usually longer than the
rest, the innermost usually much shorter: flowers 12 mm. long,
whitish or pinkish (petals with red median band): fruit 1 to 2
cm. long: seeds about 1 mm. long, blackish and conspicuously
pitted. (Ill. Cact. Mex. Bound. t. 3). Type, the specimens of
Wright in Herb. Mo. Bot. Gard.
From western Texas ("west of time Pecos, on low limestone hills,
among herbage") to Arizona and Chihuahua. Fl. April, May.
Specimens examined: Texas (Wright 121, also of 1852; Parry of
1852): Arizona (Miller of 1881): Chihuahua (Pringle 213,
250,258): also specimens cultivated in St. Louis in 1852 and
1855.
11. Cactus lasiacanthus denudatus (Engelm.).
Mamillaria lasiacantha denudata Engelm. Cact. Mex. Bound. 5
(1859).
Larger, 2.5 to 3.5 cm. in diameter, with longer tubercles (5 to 6
mm.), and more numerous (50 to 80) longer (3 to 5 mum.) spines
which are naked or nearly so. (Ill. Cact. Mex. Bound. t. 4)
Type, Wright specimen in Herb. Mo. Bot. Gard.
From western Texas (with the species) to Coahuila.
Specimens examined: Texas (Wright of 1852): Coahuila (Palmer of
1880).
In the Syn. Cact. Dr. Engelmann merges this variety with the
species, and has been followed in this by subsequent writers, but
the characters seem so (distinctive that its varietal rank has
been restored.
12. Cactus micromeris (Engelm.) Kuntze, Rev. Gen. Pl. 260 (1891).
Mamillaria micromeris Engelm. Syn. Cact. 260 (1856).
With depressed top and very rarely branching, 1 to 3.5 cm. in
diameter: tubercles very small (about 1 mm. long) and wart-like,
crowded, shedding the spines with age and giving the base of the
plant a tuberculated appearance: spines from white to ashy-gray,
1 to 3 mm. long; in young plants and on lower tubercles of adult
plants about 20, equal and radiant; on flower-bearing tubercles
30 to 40, stellate-porrect in every direction, the 6 to 8 upper
ones two to four times longer than the rest (4 to 8 mm.), clavate
toward the apex and acute (the clavate top at length deciduous),
intermixed with loose wool of about the same length and forming a
small tuft on the top of the plant which includes and partly
hides flowers and fruit: flowers whitish to light pink, almost
central, very small (6 mm. in diameter), much reduced (3 to 5
sepals, 5 petals, 10 to 15 stamens, 3 stigmas): fruit 8 to 12 mm.
long: seeds 1.5 mm. long, black and shining. (Ill. Cact. Mex.
Bound. t. 1 and 2. figs. 1-4) Type, the specimens of Wright in
Herb. Mo. Bot. Gard.
On naked mountain tops and sides, extreme southwestern Texas (Val
Verde County to El Paso) and southward into Coahuila and
Chihuahua.
Specimens examined: Texas (Wright 227 of 1849, also of 1852;
Nealley of 1892): Coahuila (Bigelow of 1853): Chihuahua (Pringle
212): also growing in Mo. Bot. Gard. 1893.