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From the U.S. C. & S. J. XXV.3:1953
Copiapoa cinerea (Philippi) Britton &
Rose is a little-known endemic restricted to a narrow coastal belt in
northern Chile bordering the Atacama desert in the southern part of the
Province of Antofagasta. C. cinerea is the most conspicuous and
widespread echinocactoid species from this area. It is also highly
polymorphic and appears to integrate with an undescribed, caespitose species
in the southern part of its range, and with Copiapoa marginata
(Salm-Dyck) Br. & R. in the north. In herbaria these integrades have
been variously determined. At least one new species (Copiapoa gigantea
Backbg) has been based on a plant from one of the outlying
populations in the northern range of C. cinerea. Another name (Copiapoa
haseltoniana Backeberg n.n.) has been proposed for a yellow-spined
variant of C. cinerea. These two names will be discussed later. The
purpose of the present paper is to clarify typification of C. cinerea
and to present data on natural variation in a population in Quebrada
Taltal and adjacent hills, an area which presumably includes the
locality of collection of the type. Field studies were made in January 1952
during the University of California Botanical Garden Expedition to South
America, 1951-52.
Philippi’s original description as
Echinocactus
cinerea reads:
“E. omino
cinereus diametri 4-pollicaris; costis numerosis; verrucis vix 2 lin.
inter se distantibus, diametri 21/2 lin., vetustioribus immersis, planis;
aculeis 5-6, nigris, teretibus, supremis duobus parvis, circa 2 lin.
longis, inferioribus circa 6-8 lin. longis, centrali 9-10 lin. longo;
apice lana alba densissima, 9 lin. longa tecto, flores plures vix e lana
emergentes, 9 lin. longos, flavos emittente. In litorali a valle Taltal a
25o 24’ lat. m. usque ad Cobre 24o 25’ lat m. frequens, inter
majores recensendus, valde ramosus, massas interdum diametri 1½ ped.
formans”.
No type plant has been designated in the
literature. Judging from the confusing array of annotation labels on the
various specimens of the type collection which I have seen, some elaboration
of the history of the dried material which Philippi collected is necessary,
since my choice of a lectotype differs from that of Rose. Rose’s choice was
later reiterated by Dr. Carlos Munoz P., who annotated Rose’s choice as
“Typus” and photographed this sheet at Santiago (SGO No. 041282) for
distribution to other herbaria. My citation of the Philippi collection
preserved at Santiago follows, and an explanation for it and the complex
history of the sheets examined is given immediately thereafter.
Chile, Prov.
Antofagasta, Dept. Taltal, Quebrada Taltal, “Hueso parado”, R.A. Philippi,
Dec. 14-15, 1854 (Herb. of F. Philippi No. 791) (SGO No. 502667 —
lectotype; SGO No. 041282 — isotype, US fragment).
Rose examined the material which Philippi
collected and deposited in the Herbarium of the Museo Nacional de Historia
Natural, Santiago, Chile. At the time Rose saw it, this material was
unmounted and there were three labels for it, each with slightly different
data. These labels read respectively, “Hueso parado. Decembri 18 4 Ph.”,
“Des Atacama”, and “Dr. R.A. Philippi leg. Taltal in des. Atacama.
Herb. F. Philippi 791.” Rose apparently assumed that these labels
referred to different collections and that the material, which showed
considerable variation in spine morphology, represented more than one
species. He separated from this material several areoles which showed
spination which agreed most closely with Schumann’s plate of this species
and took a fragment from his separation which is now mounted at the National
Herbarium, Washington, D.C. (US).
The remainder at Santiago was mounted later as SGO 041282, and Dr. Carlos Munoz P. annotated it as the “Typus” for
Echinocactus cinereus Philippi. I examined SGO 041282 at Santiago in
1952 and the fragment from it at the US Nat. Herb. in 1951 and this material
is definitely taken from a single plant which was a typical of Copiapoa
cinerea in a population at the type locality (see notes below) and
further, does not fit Philippi’s description. It may be described briefly as
follows:
areoles with
two central spines 12mm long, no radials; flower ca. 17 mm long, limb 13
mm wide. The label chosen (by whom?) for this material and mounted on SGO
041282 reads, “Dr. R.A. Philippi leg. Taltal in des Atacama. Herb. F.
Philippi 791.”
The balance of the material left after Rose
separated the material of SGO 041282 (and the fragment of it at US) was
eventually mounted at Santiago as SGO 052667 and bears the label “Hueso
parado. Decembri 1854 Ph.” Rose’s annotation label “a mixture here” is now
mounted on this sheet. This material may be briefly described as follows:
rib section of
8 areoles, central spines two, 19 to 22 mm long, radial spines 3 to 6, 3
lower radials 12 to 15 mm long, the central lower one usually 1 to 3 mm
longer than the two lateral ones, 3 upper radials much shorter (up to 4 mm
long) but occasionally 1 to all of the upper radials missing, flowers as
in SGO 041272.
Rose’s concept of this species, which he did
not study in its habitat, was apparently largely based on Schumann’s figure
of a plant with usually one or two short central spines and no radials.
Philippi, however, describes his Echinocactus cinereus as having 5 to
6 spines, so apparently his concept was based primarily on the material now
mounted at Santiago as SGO 052667, which Rose, and later, Munoz, considered
to be of another species, or at least atypical of Copiapoa cinerea.
As is shown below, all of Philippi’s material falls within the range of
variation in a population of C. cinerea at the type locality, but it
was apparently taken from two different plants. Rose correctly separated the
material from these two different plants, but was in error in assuming that
what was left represented a different species. In fact the material left is
the most typical of C. cinerea and since it fits Philippi’s
description more closely, it should be designated the lectotype (i.e., SGO
052667).
As to the proper locality for the type
collection, and the status of the three available labels, it is my opinion
that F. Philippi, (R.A. Philippi’s son) wrote out additional labels for this
material for his herbarium and casually altered the data each time. The data
on all labels could equally well apply to one locality, indeed there is a
hierachy of specificity with regard to that locality as is readily seen by
the following information on the localities concerned.
Johnston (Contr. Gray Herb. 4 (85): 1-138,
1929) reports that Philippi collected in the Quebrada Taltal on Dec.
14th & 15th 1854. He labelled all specimens taken on these dates as “Hueso
Parado”.
His campsite was recorded as “Agua del Clerigo” which is
probably either one of the two localities now known as Agua de Perales
(a spring on the south base of Cerro de Hueso Parado) or Agua de
la Lora (a spring in a Quebrada of the southeast flank of the same
mountain, now known as Cerro Perales). While at this station he
collected from there to the shore now called Caleta de Hueso Parado.
His collection of C. cinerea must have been taken inland near his
camp, however, since shoreline populations of C. cinerea have
generally longer, thinner spines which are brownish or yellowish (compare
P.C. Hutchinson 401, UC) rather than black or grey as is typical of inland
populations of this species, as well as of Philippi’s herbarium material.
Since one of the two plants which Philippi preserved has no radial spines,
it is likely that it was taken in one of the plateau populations on the
flanks of Cerro Perales in Quebrada Taltal, for as is mentioned below, forms
with no radial spines were found by the author only in plateau populations
and those on the slopes above were always more spinose and always bore
radial spines. Forms with radial spines, conversely, do occur in the plateau
populations, and probably Philippi took his material of this type of plant
(designated above as a lectotype) from a plant nearby the one from which he
took the material with no radials (SGO No. 041282).
In January 1952, the author found Copiapoa
cinerea abundant on level rocky plateaux, slightly raised above the
quebrada bottom-lands, 10 km east of Taltal in the Quebrada Taltal, and
again on similar plateaux 5 to 10 km inland in the Sierra Esmeralda 30 to 50
km to the south. It occurs in less dense stands and often as isolated
individuals on the hills above Quebrada Taltal and extends from there
continuously over the coastal range, reaching the coast about 5 km north of
Taltal, then receding into the hills about 15 or 20 km north of Taltal. In
the Sierra Esmeralda its distribution, as far as known, is almost
exclusively on small inland plateaux, rarely on hillsides, and apparently
not at all on the coast. I was unable to confirm its distribution further
north than Quebrada Guanillo above Paposo although, as stated
above, Philippi records it as far as Cobre and Johnston reports it in
“the region north of Paposo”. In its densest stands on the plateaux, it is
rarely associated with any other cactus but on hillsides a common associate
is Eulychnia iquiquensis (Schum.) Br. & R.
Most plants are columnar, to 1.3 meters tall
usually single or with 1 to 5 branches commonly from the base. There is
little or no variation in stem shape. One extremely large plant examined in
the hills 2 km west of Breas (P.C. Hutchison No. 394) had 19 branches
from the base, each about a meter long, with 29 smaller branches from the
apex of these stems. In this same area I found the only examples of
fasciated stems which were observed in any population of Copiapoa -
about ten large plants each with three to eight stems ca. 1 meter tall, and
with one to several stems fasciated. These are apparently the first example
of fasciation to be reported in the genus Copiapoa. Several of the
fasciated stems were in flower. It was interesting to find that these
fasciated plants were all growing together at the top of a narrow, steep
gorge cutting the sides of the Quebrada Taltal, and apparently occurred
nowhere else. I could find no evidence that an environmental factor was
responsible for the fasciated stems. They were not isolated from normal
plants.
On the slightly elevated, flat plateaux of
lower altitudes, where C. cinerea is most abundant, the plants are
usually smaller, seldom more and generally less than 0.7 meters tall. A
single main stem is most common, with subsidiary stems from the base or, in
most cases, from the lower half of the plant, being half as long or less:
Occasionally a cluster is found in which all stems are of uniform length and
form a mound. The smaller size of the plants in plateaux populations is
probably attributable to a scarcer water supply. The plateaux were below the
limit of the winter fogs (called “camanchaca” apparently the primary source
of water in most years, for rain is rare in this area), whereas all of the
largest specimens examined on the sides of the quebradas grew in the fog
zone.
Britton and Rose in The Cactaceae 3: 85-90;
1922 used the number of ribs as a key character for distinguishing species
of Copiapoa. Subsequent authors have considered this character to be
diagnostic for some species of this genus. In C. cinerea the
variation in number is at least from 14 to 30. Number of ribs is probably
correlated with age and size of the plant as influenced by climatic
conditions, since the oldest plants in the most favourable habitats had the
largest number. The various stems of a single plant show little variation in
number when they are of equal size (usually a variation of 0 to 3 ribs), but
when different stems of one plant are of various sizes (and, presumably,
ages) the variation is larger. Areoles may be nearly approximate or up to 2
cm apart. The epidermis in all specimens examined was always ash-grey.
Central spines are either one or two, most
often the latter, with the uppermost spine appearing first and often
attaining greater length; length varies from 1.3 to 3.3 cm. Radial spines
may be lacking (rare) or from 1 to 7, but commonly 2,4, or 6; length varies
from 0.5 to 2 cm, but radials are usually shorter than the centrals. On most
specimens there is fluctuation in spine pattern on various parts of a single
stem, the only notable exception being plants with a single central and no
radial spines. When radial spines occur the number of centrals may vary at
random between one and two on a single stem, two being more common. In
general, plants at higher altitudes on the quebrada slopes within the fog
belt are more heavily armed with stouter and longer spines and bear at least
two centrals and 4 or more radial spines (commonly 4, 5, or 6). Nearer the
coast and at lower altitudes in the plateau populations, spines are much
shorter and plants with
1 or 2 centrals and no radials occur. There is considerable variation,
however, within the general limits described above, but not exceeding these
limits i.e. 1 or 2 centrals, with 0 to 7 (usually 2, 4, or 6) radial spines. There is no consistency in arrangement of
spines in any particular area other than as pointed out above and the
dominance by numbers of plants with radial spines occurring in pairs.
At all localities spines are extremely brittle,
especially the older (lower) ones. The lower half of most old stems is
usually completely naked due to the loss of these brittle old spines. Spines
are usually black, fading to grey-black or grey, but at one locality ca. 20
km north of Taltal on the coast road to Paposo some plants bore brownish or
yellow-brown spines which became under cultivation, more markedly yellowish
(compare P.C. Hutchison 401 UC, also cultivated at UC Botanical garden No.
52.585 and No. 52.585S). This form has been called Copiapoa
haseltoniana Backeberg n.n. (C. & S. J. U.S. 23.4:1951).
Flowers of C. cinerea are 1.7 to 3.5 cm
long and 2.0 to 3.5 cm wide across the spread limb; generally length and
width are about equal. Colour ranges from yellow to yellow shaded pink to
dark red on the outer surface of the petals. Flowers of a single plant are
usually of the same size and colour, but variation occurs between flowers of
different individuals. No significant variations in morphology, other than
size, were noted. Both flowers and fruit have a mass of white or
cream-coloured hairs attached to the base below the ovary. In extracting
flowers from the stem apex I found that the hair is often not attached, as
it is apparently more strongly affixed to the areolar tissue than to the
flower tissue. The individual strands of hair in C. cinerea are more
or less 2 cm long.
The fruit is cream-coloured tinted pink, to
almost pure carmine, naked or more usually with 1 to 5 short, erect, deltoid
to lanceolate bractlets at the truncated apex and occasionally with a few
smaller bractlets subapically. These bractlets may be lost during
development. The ovary is at first totally immersed in the densely matted
hairs of the stem apex. Shortly after fertilisation it begins to colour. As
it matures it swells upwards until finally the apex is almost flush with or
a mm or so above the surface of the apical hairs. The perianth dries and is
at first persistent on a circular disc at the apex of the ovary. Dehiscence
is usually accomplished by the drying of this disc prior to the drying of
the ovary walls. As the disc shrivels it breaks from the sides of the walls
of the ovary; later it is attached by a very narrow section to one side and
may be lifted at an angle above the ovary. The withering-persistent corolla
usually breaks away in the wind at this time. Attachment of the flower to
the disc is always weaker than that of the disc to the ovary; if the
withered flower is grasped, it usually pulls away and the disc remains. As
the disc first ruptures, seeds are extruded through the narrow opening and,
in most cases, these seeds are still attached by the funicles to the
placentae, or at least, the broken funiculus remains attached to the hilum.
In the sun the fleshy remnants of the funiculus dry rapidly and the seed
becomes scattered over the apical wool, to be blown away or eventually to
drop to the base of the plant. As dehiscence nears the halfway point, the
ovary begins to dry up and shrink back into the wool; the disc and attached
flower usually break away at this time. As the capsule shrivels, all seed is
extruded or some may remain, so that a dried capsule may be empty or contain
seed. The dried capsule is usually again totally immersed in the matted
hairs of the stem apex.
Occasionally the disc may not rupture and the
capsule dries about the seed without dehiscing. The capsule is later
disgorged and lies on top of the apical hairs of the stem. This dried
capsule is round or oval, very brittle and tough, and solidly filled with
seed. The withering-persistent flower, in such cases, becomes lost before
the capsule is disgorged. Capsules which have lost their discs sometimes
continue to grow and are usually distorted in shape and fissured by pressure
against surrounding structures.
Plants in the plateau population 10 km east of
Taltal were infested with a scale insect (Diaspis sp.). The
infestation was most severe on young plants or on young basal offsets of
mature plants. No infestation by scale insects was noted at any other
locality. There was no evidence of infestation or damage to C. cinerea
by any Buprestidae, as has been demonstrated for C. humilis
(Phil) Hutchison and C. taltalensis (Werd) Looser.
In summary, it has been shown that variation in
certain characters, previously presumed to be diagnostic and static, of the
Chilean endemic species Copiapoa cinerea (Phil) Br. & R., is
extremely wide. This is particularly true with reference to habit, ribbing,
areole placement, spine number, size, arrangement and colour, flower colour
and size, and finally, fruit colour and morphology. Conversely, there was
little variation in general shape of stems, epidermal colour, details of
floral morphology other than size, and seed morphology.
Comments on Copiapoa
cinerea
from H.
Middleditch
In the course of searching the literature for
information dealing with the Copiapoa cinerea species-complex, I have
had occasion to read material written by Backeberg, Buining, Knize and
Ritter and I must admit that the above article by Hutchison - although
perhaps lengthy - would appear to be not only the most thorough treatment
but also a model of clear descriptive writing. The account of the Philippi
herbarium specimens is so explicit that one almost feels, in the course of
reading it, that one has actually been examining the herbarium specimens
oneself.
Quite apart from the valuable observations made
by Hutchison there would seem to be several interesting points unresolved.
For example, Hutchison offers his opinion that F. Philippi “casually altered
the data” on the herbarium specimens. Now I would not be unduly surprised at
such goings-on in a commercial nursery, but it is hardly what one expects of
a professional botanist. However, this begs two questions: was R.A.
Philippi’s son a trained botanist? And were other Natural History Museums
equally casual at that day and age? On either point one may perhaps feel
that Hutchison’s opinion could be of value.
This species is described as
intergrading with
an undescribed caespitose species to the south of its distribution; when one
considers the relative ease of travel through this dry northern territory of
Chile and the very thorough collecting activity of Ritter there, not to
mention other collectors in that field, it is somewhat remarkable that any
undescribed species could be left, even in 1953. With a geographical plot of
the various species reported from Chile, including those found by later
collectors, it is still not at all clear whether Hutchison was referring to
plants which were given a species name by another collector between that
time and the present date. In his account of his collecting work in this
vicinity (Chileans No. 16) Knize refers to the richness of the cactus flora
between Taltal and Paposo, thereby implying that it is more sparse to the
south of Taltal. About 25 km to the south of Taltal is Cifunchos, the
place reported by Lau as the location for our cover plant; Knize reports
collecting C. cinerea KZ 77 “South of Taltal” but this does not tell
how far to the south the collection was made. Hutchison states that C.
cinerea is found as far south as the Cerro Esmeralda (which lies
about 40 km south of Taltal) but indicates that plants were found as far as
50 km south of Taltal. This is barely 10 km from Latitude 26o S
which Ritter gives as the habitat location for Copiapoa columna-alba.
These plants, however, are described by Ritter as solitary and so can hardly
be the “caespitose species in the southern part of the range of cinerea”
with which cinerea intergrades, according to Hutchison.
The ground to the north of Paposo was not
covered by Hutchison, but it was travelled by Knize, who refers (Chileans
No. 16) to the absence of Copiapoa cinerea in that area, although it
was reported from there by Philippi. Knize poses the question whether a
marginata-like plant was taken for a northward extension of cinerea,
whilst Hutchison suggests that C. cinerea intergrades with C.
marginata to the north. It would be nice to have some clear idea of what
was each writer’s conception of “marginata”, for then it would be
less difficult to know what each one meant when comparing other species with
C. marginata.
There are some excellent illustrations
accompanying the article by Hutchison; some of these depict grooves between
the ribs which are slightly wavy on younger growth in the crown, but none
show the very wavy groove to be seen on our cover illustration and on
Ritter’s photographs of C. dealbata. There is a passing
reference to white or cream wool in the crown, but no reference to the
orange coloured wool on C. haseltoniana. Some of these illustrations
would suggest that sunken areoles, such as those described by Ritter for
C. dealbata, do occur on plants of C. cinerea which we observed
by Hutchison, but this receives no mention in the text. Either the
photographs are misleading, or perhaps Hutchison considered it a matter of
no importance.
The barrel-shaped fruit is described by
Hutchison as a “dehisced capsule”, presumably a normal type, since he then
describes the tapering tri-cornered fruit as abnormal. Of those Copiapoa
which have set fruit for me (not cinerea sp.) this latter fruit
shape has certainly not been abnormal.
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