Camp 1949b

Cinchona at High Altitudes in Ecuador
Author(s): W. H. Camp
Source: Brittonia , Feb. 21, 1949, Vol. 6, No. 4 (Feb. 21, 1949), pp. 394-430
Published by: Springer on behalf of the New York Botanical Garden Press
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VOL. 6, No. 4, pp. 394-430 BRITTONIA FEBRUARY 21, 1949
CINCHONA AT HIGH ALTITUDES IN ECUADOR
W. H. CAMP
TABLE OF CONTENTS
I. Introduction. . . 394
II. Source of the Data . ... ..... ..... . . .... . 394
III. The Alkaloids of Cinchona Bark ............. . .398
IV. The Pitaya-Serrana-Pata de gallinazo complex ...398
V. The Alkaloids of the Pitaya-Serrana-Pata de gallinazo complex . . . 402
VI. A Transect of the Western Escarpment ..... 405
VII. An Explanation of the Aberrancies Met With in the Rio Saloya-Corazon Pass
Transect ... ....1......2.......................... .......................... .. 412
VIII. The Possible Origin of the Serrana Type ........................... .415
IX. The Possible Origin of the Pata de gallinazo Type . . . 417
X. On the History of the Group and the Nature of the Cline in Alkaloids in the
Pitaya-Serrana-Pata de gallinazo Complex . .. . ..... .. 421
Summary .............. . ............. 428
Literature cited . . . . ... .... .. .....430
I. INTRODUCTION
Those who have made critical field studies of the members of the genus Cin-
chona on the cloud-swept and often precipitous upper slopes of the northern
Andean escarpments-where, at elevations between 8,000 and 11,500 feet, the
rainfall may be as much as, or exceed, 200 inches per year-will agree that
in certain of its phases the genus can be complex. Yet, in spite of the often con-
fusing appearance of local stands on some particular slope, or in a single valley
head, field study over a period which is sufficiently long and covers enough ter-
ritory (both in area and altitude) generally yields clarifying results, for out of
this seeming welter of variabilities a pattern eventually begins to emerge. Even
so, the over-all complexity will remain, for this is the character of the Cinchona-
flora as it exists today, especially in the southern part of Ecuador. But it is, in
my opinion, an orderly complexity rather than chaos. It is a pattern in vari-
ability which, when eventually understood, ought to lead to a satisfactory nomen-
clatural treatment. This is to be hoped for because the systematics of the genus
at present is in a sorry condition.
This paper, however, is not one wherein the problems of nomenclature will
be given more than passing mention; instead it is designed only as an exposition
of several of the lesser patterns of variability running through a few segments
of the Ecuadorean Cinchona-flora, together with certain speculations on the pos-
sible ways in which these portions of the general pattern might have developed.
Furthermore, to a considerable extent, the discussion will be based on the kinds
and amounts of crystallizable alkaloids present in the different types of Cinchona
considered. The obviously correlated study of the morphological variabilities
within these same populations leads directly to the problem of nomenclature;
since that phase of the study must be taken up separately (and it is expected by
others), the morphological characters of the individuals of these populations will
receive mention only where they seem to be pertinent to the present discussion.
II. SOURCE OF THE DATA
The general background of the activities of the group employed by the United
394

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1948] Camp: Cinchona in Ecuador 395
States Foreign Economic Administration' to explore for and locate stands of

exploitable Cinchona in Ecuador has already been outlined by both Rainiey (10)
and Steere (11) and need not be recounted in any detail except for certain items
which have a partieular bearing on the methods whereby the data presented
here were obtained.
In the early years of World War II, I had been occupied with various prob-
lems related to the war emergency in other parts of Latin America. Therefore
I did not join the group working in the Ecuadorean Cinchona forests until April,
1944; my connection with that organization continued until the cessation of its
exploratory work in April of the followinig year. I then rejoined the staff of
the New York Botanical Garden (having been on leave for three years) but re-
mained in Ecuador to continue for an additional period of about six months ob-
taining collections and making observations on the general flora of the region.
Since the basic purpose of the Cinchona-group of the Foreign Economic Ad-
ministration was to obtain the greatest amount of quinine-yielding barks in the
shortest possible period, emphasis necessarily was placed upon those areas wherein
the trees were thought to contain the largest quantities of this particular alka-
loid. Consequently, during that critiial period it was not possible to allot more
than hurried reconnaissance surveys to the areas wherein the low-yielding stands
were known to predominate. During the latter part of my stay in Ecuador it
therefore was possible to spend some time in certain areas which were not only
of general botanical interest but which also contained stands of low-yielding
Cinchona. I was thus able to fill in certain blank places in our general informa-
tion and observations which otherwise would not have been possible had my work
in Ecuador terminated with the cessation of concentrated exploratory work for
the alkaloidally better types of the genus. It was during this latter period-
when the over-all picture of these complex populations could be obtained-that
the data previously assembled by the group of workers with which I had been
associated began to fall into what, to me, appeared to be a more logical pattern.
In practice, in the field, small parties of the group of explorers penetrated
either those areas where stands of Cinchona were reported to exist (but about
the composition of which little was known), or into areas where they might be
expected to occur. There, a working estimate of the available trees in the stand
was made; at the same time representative samples of the bark of a varying num-
ber of individual trees were collected, together (in general) with a certain num-
ber of authenticating herbarium specimens.2 The bark samples then were ana-
lyzed in the central laboratory in Quito. From these combined data a rather
accurate estimate could be made, not only of the tonnage of bark, but also of the
anmount and kinds of alkaloids available in a given area.
In the early part of the work it seemed evident that those areas wherein the
populations were most nearly homomorphic were likely to yield bark whose
analyses were quite similar. However, as the work extended farther afield we
were more and more forced to the conclusion that what at first glance appeared
to be very similar trees (even those growing in the same stand) might, on occasion,
I The pertinent phases of the work here treated were carried out mainly under the Foreign
Economic Administration successor to both the Board of Economic Warfare and Office of
Economic Warfare; they were concluded under the U. S. Commercial Company.
2 It is to be regretted that more authenticating herbarium specimens at times were not
collected in certain areas. Failure to obtain them in reasonably large series did not in any
way hamper or retard the exploitation of quinine-yielding barks; the lack of them is most
keenly felt now that the emergency period is passed and some attempt is being made to in-
terpret the various available data in terms of systematics and bio-dynamics.

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396 Brittonia [VOL. 6
contain not only different amounts but also different kinds of alkaloids. This
led to the necessity of making very careful observations on characters which
either had been overlooked or deemed unworthy of consideration by some of the
earlier students of the genus. The problem confronting us in the field was not
one of making broad specific interpretations for ease in the filing of herbarium
specimens; our problem was that of finding field characters which would permit
the recognition of populations of high-yielding types, even ahead of the results
of the necessary analyses made in the central laboratory in Quito, and so facili-
tate our explorations.
As has been noted, it is unfortunate that an authenticating herbarium speci-
men was not always collected with each bark-sample.8 However, it should be
added that after a short period of field experience the "on-the-spot" identifica-
tions by the majority of the group were usually fairly accurate. It also should
be made clear that, since various members of the exploratory group were profes-
sional botanists, these did make every effort to obtain authenticating herbarium
materials where possible, especially if any of the trees seemed to be at all dif-
ferent from the general population in which the work was being carried on at
that particular time. Notes of such items were then usually appended to the
laboratory reports and kept in the files of the central office in Quito.
Because of various circumstances, I was the last of the exploratory group to
be retained in Ecuador. Therefore the task of closing up certain activities of
that organization fell to me. In leafing through the files of the group in Quito,
it became apparent that they contained considerable amounts of data which likely
would be of more than passing interest to any systematist attempting to work
over the taxonomy of the genus in the future and certainly much of exceeding
importance to one interested in the dynamics of its species. Furthermore, events
which took place at that time made it seem unlikely that more would be done in
the immediate future with the data on chemical analyses of the various field
samples than to ship them to the central office of a presumably temporary Govern-
mental agency. There would be the chance that there they might be filed in such
a manner as scarcely to be easily available in the more distant future, or possibly
lost, or even discarded by those who did not understand their importainee, much
as seems to have been the fate of a similar set of analyses made of materials col-
lected from the same genus a quarter-century previously by Rusby and Pennell.
The time available to me was short-much too short to make a complete abstract
of the information-and so I was forced to the alternative of taking such por-
tions of the data as seemed most likely to yield immediately pertinent infornia-
tion. Also, I knew that already parts had been extracted for use in a paper
which was then being prepared and which has since appeared.4 The general
scope and method of approach of that paper was known to me, therefore it was
thought best to select those parts of the data as might be useful in a study of
the major population types within certain more restricted areas and especially
from the standpoint of the variation in individual trees within these stands.
A preliminary sorting of the data seemed to indicate that, within the limited
time available, the most significant results would be obtained by a consideration
3 It will be understood that in the emergency period of the work, and where rapid recon-
naissance surveys were made usually under rather trying conditions, it was not always feasible
to spend the length of time necessary to fell a large forest tree just for the sake of several
herbariumn specimens, much as the various members of the group might have llked to obtain
them; instead a very satisfactory bark sample could be made from the tree in situ merely by
a few blows with a machete and with scarcely a pause in the exploratory transect.
4 The paper by Martin and Gandara (8).

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of the problems revolving about certain forms which occur at fairly high eleva-
tions and also in outlining the situation regarding the alkaloids present in the
forms met with in a geographically related altitudinal transect. In thus limiting
the scope of the study it was necessary to use the data mainly obtained from the
field samples made by my former associates who had worked primarily in the
central and northern provinces. Certain data taken from the studies of my own
field goup in the southern provinces have also been inieluded. The reason for
choosing a high-altitude population for the first part of the study was that, of
the forms met with in any abundance in the northern provinces, it yielded by
far the highest amounts of crystallizable alkaloids and therefore had received the
most attention from the field group. As will be seen, the data on similar popu-
lations in the southern provinces are not so detailed; there, our preliminary sur-
veys indicated that the cognate high-altitude populations were so poor in alka-
loids that no extensive investigations of them were made. Nevertheless, the data
on them appear to be representative.
The region chosen for an altitudinal transect also was in the north, in the
Province of Pichincha, specifically in the Rio Saloya-Corazon Pass area. Al-
though technically these are different localities they constitute a definitive region
of rather small compass and are within the same drainage system; it was there-
fore thought best to combine them since no observable differences were noted in
the general populations at the same elevations. This unit area is at present
traversed by a road leading approximately west from Quito (the Quito-Sanlto
Domingo de los Colorados road via Chillogallo). The upper levels of this area
also are penetrated by a spur road which diverges toward the west from the
Quito-Latacunga road (the present Pan-American Highway) about 25 kilometers
south of Quito, and then swings in a northwesterly direction before it stops; as
no more than a poor trail at present, this continues to lower levels where it
ultimately connects with the Quito-Santo Domingo de los Colorados road men-
tioned above. The available tranisportation system in this area favored the ob-
taining of a more complete and representative group of samples than from cer-
tain other regions.5
The data on the chemical composition of certain Ecuadorean barks presented
in this paper, therefore, were taken from individual-tree analyses of materials
collected by the exploratory group of the "Mision de Cinchona del Ecuador"
during the course of its routine work in 1943, 1944, and briefly in 1945, together
with several other pertinent items, the sources of which will be noted where they
appear in the text. The preliminary manuscript of this paper was submitted to
several of my former colleagues before publication since they were more familiar
than I with the northern provinces of Ecuador, whence the data largely were
obtained. Therefore, it is a pleasure to thank Doctors William B. Drew, Gerald
W. Prescott, William C. Steere, and Ira L. Wiggins for their criticisms and sug-
gestions. It is to be understood, however, that the interpretations presented
here are my own and do not necessarily imply agreement on their parts.
It will be noted that all altitudes given are in feet. During our work in
5 In the foregoing there is no intimation that my associates working in the northern

provinces tended mainly to stay close to graded roads in their explorations. It will, I trust,
be understood at all times that, while the task of our group was one of exploration, it also was
necessary to take into account the factor of transportation. It is obvious that, faced with
the necessity of obtaining the bark in the shortest time possible, it was at times advisable to
concentrate on those areas closest to existing means of transportation rather than-as was
sometimes necessary-to construct trails through uncleared wildernesses over a most difficult
terrain.

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Ecuador, altimeters calibrated in meters were not generally available to the

field parties, hence the readings from those we had were recorded directly on
the field sheets rather 'than chance the possibility of introducing errors by hastily
calculated, conversions. Since this is true of nearly all the records of the group,
the present paper follows this same system.
III. THE ALKALOIDS OF CINCHONA BARK
Although the alkaloid quinine is extracted from Cinchona bark it does not
follow that all Cinchona barks contain quinine. It is well known that many
kinds contain no quinine; in fact, several types of bark from within the genus
apparently contain no more than mere traces of any alkaloid.
Reference to any standard work on the subject will reveal that a series of
alkaloids may be present in Cinchona barks; of these various kinds, the crystal-
lizable alkaloids are the ones dealt with by those primarily interested in a study
of antirmalarial substances. Four crystallizable alkaloids are generally recog-
nized. Three of them, cinchonine, quinine, and cinchonidine-in order of the
prevalence of their presence in the general group of Ecuadorean barks-may be
classed as antimalarials. The fourth, quinidine, important because of its use
in the treatment of auricular fibrillation, is found in small quantities in certain
types of bark. Where used in this paper, the term "total crystallizable alka-
loids" (abbreviated as TCA) refers to the sum of the anhydrous forms of these
four alkaloids expressed in terms of the per cent weight of the dry bark.
IV. THE PITAYA-SERRANA-PATA DE GALLINAZO COMPLEX
As has been set forth by Steere (12), that portion of the Cinchona population
occurring primarily between the elevations of 9,000 and 10,000 feet in the nortlr-
ernmost provinces of Ecuador is relatively homogeneous. In that paper he
refers to this material as C. pitayemsis, a species first described from Colombia,
although he writes (12, p. 476), "In Ecuador, the trees are smaller. . . . Fur-
thermore, the leaves are more pubescent, especially when young, and the capsules
are less strongly ribbed. The Ecuadorean form was proposed as a new species,
Cinchona corymbosa, by Karsten (1858), on the basis of specimens from southern-
most Colombia. Although this name has generally been treated as a synonym of
C. pitayensis (Standley 1930), there appears to me to be reason to keep them
separate, perhaps with varietal rank, especially after having seen both forms in
the field. "
The differences between the Colombian and northern Ecuadorean stands
of C. pitayensis as pointed out by Steere seem to be significant and will be further
noted in another part of the paper; yet, with him, I feel that the differences are
not sufficient to warrant the recognition of two separate species. Because of
their great similarities they are collectively referred to in this paper as the
"Pitaya" type.
Somewhat southward, in the Province of Pichincha, another form begins to
be evident in the high-altitude population; this is the type of material called
"Serrana" by the local casearilleros (Cinchona bark gatherers). By direct im-
plication, Steere separates this material from C. pitayensis for he refers to it (12,
p. 471) as ". . . the 'Serrana' variety of Cintchona pubescens. . ." This same
viewpoint is also taken by Martin and Gandara in their paper on the composition
of Ecuadorean and other barks. However, they admit (8, p. 188) that "This
bark [Serrana] has been considered a type of C. pubescens, and is so listed here
because of similarity of [alkaloid] composition. In reality, the tree in leaf, fruit,

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and flower characteristics is much closer to C. pitayensis, which occurs in northern

Ecuador in the same elevation zone."
This same Serrana type continues southward along the western escarpment of
the Cordillera through the central provinces and at least into the central part of
the Province of Cafiar where I found it to be locally abundant. There, it is
scarcely to be distinguished in morphological characters from the trees which
occur in the Province of Pichincha. Concerning one of the central provinces,
Steere writes (12, p. 467): "Cinuchona pitayensis was . . . found to be abundant
in the Province of Leon [the Province of Cotopaxi on the most recent maps];
north of Pilalo by G. W. Prescott, and in the region of Sigehos by M. Acosta-
Solis. A curious feature of this species in Leon [Cotopaxi] Province is that al-
though its morphological character and its habitat preference are quite typical,
its bark is not particularly rich in alkaloids, especially quinine. The cause of
this unusual condition will probably be found in some soil factor, although the
species itself may be more variable at the edges of its geographic range. "
In considering this material later, I came to the conclusion that it more prop-
erly belonged with the Serrana type than with the Pitaya type. It is my opinion
that when the notes on which Steere 's statement was based were taken, the mor-
phological characters of this material from the Province of Cotopaxi (Leon) led
him toward the conclusion that it was a "curious" low-alkaloid form of Pitaya;
conversely, my study of the alkaloid types in the same population, in conjunction
with its morphological characteristics, influenced me to the belief that it was
more properly the Serrana type. This difference of opinion indicates something
of the tenuous nature of the morphological characters which separate the Serrana
and Pitaya types; and the quotation from the Martin and Gandara paper serves
to give weight to my contention that they are morphologically so close that they
deserve to be treated as parts of the same complex. Certainly, if the Serrana
type is not to be considered as specifically distinct from the other Cinchonas, but
as part of a species-complex, it belongs with C. pitayensis rather than with C.
pubescens, a species which, in the more important characters, is morphologically
quite different.
In summary, then, the Serrana type, being quite similar to the Pitaya in ap-
pearance but differing considerably in its alkaloid picture, apparently occurs at
least from the Province of Pichincha southward along the western edge of the
escarpment into the Province of Cafiar.
In the work of my own group at comparably high altitudes along the crest
of the eastern escarpment, in the Provinces of Cafiar, Azuay, and Loja, another
"type" of bark was located; this was called "Pata de gallinazo" (or sometimes
"Pata de gallinazo blanco" or, more rarely, "Hoja de luema redonda") by the
local casearilleros. It is usually a smaller tree than those at comparable eleva-
tions northward, but it must not be forgotten that these southern provinces are
the classical Casearilla (Cinchona-bark) areas and consequently have been greatly
over-exploited, so much so that it is indeed rare in certain localities to find a
fully grown speci,men of any species. It is a well known fact that for at least
two centuries even the essentially quinine-free barks have been harvested to be
used in the preparation of "tonicy" wines, sparkling waters and "bitters," and
also as adulterants for the better kinds.6 Conversely, as he pointed out, it was
not until Steere rediscovered the stands of Pitaya in northern Ecuador that any
6 Having bought quite a few commercial lots of bark while in the southern provinces, I am
fully aware of the many tricks which the casearilleros of the region can play; they are past-
masters at the gentle occupation of "doctoring" a lot, having brought it almost to the state
of a fine art, so that even an experienced dealer is sometimes fooled.

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attempt was made to exploit them; therefore except for incidental clearing along
with other forest trees, in the north the stands of Pitaya were in an essentially
virgin condition.
My work took me into the southern provinces without opportunity to see
stands of either Pitaya or Serrana. Thus the Pata de gallinazo type struck no
note of recognition when I first saw it. Months later, when an opportunity pre-
sented itself to examine those areas in the northern provinces where both Pitaya
and Serrana grow, it became immediately evident that this material of the south-
ern provinces was so exceedingly similar in general appearance to those of the
northern provinces that I was at a loss to find any easily definable (or stable)
morphological character, or group of characters, which would be infallible iil
separating them. However, although morphologically similar, it was easily de-
termined that Pata de gallinazo was neither Pitaya nor Serrana by the simple
expedient of chewing a piece of the bark. It was bitter but, being almost devoi(d
of alkaloids, to me the bark of Pata de gallinazo produced neither the sharp,
tingling sensation in the mouth such as is caused by the einchonine of the Serrana
bark, nor this same tingling coupled with the accumulative, lingering bitterness
of the additional alkaloids (such as quinine) in the Pitaya type; it had the flat,
puckery bitterness closely akin to that of certain tannins.-
Adequately large series of herbarium specimens taken throughout the com-
plete ranges of these three types-Pitaya, Serrana, and Pata de gallinazo-are
not available to me as this is written, but from my observations of them in the
field it is my opinion that they form a sufficiently compact group so that, for the
purposes of the present study, they might better be treated as parts of the same
morpho-taxonomic unit. Their chemo-taxonomy is quite another item and I
think has been more responsible for the many attempts over a long period to
keep them separate than have any differences in their morphology.
Therefore, looking at the problem from the standpoint of the community of
morphological characters which they share (rather than attempting to over-em-
phasize their admitted differences), it would seem that, among the many and
widely diverse types of Cinchona in Ecuador, these three form a reasonably com-
pact unit. If we are to consider them as parts of a single complex which perhaps
has developed out of a basically common ancestry we are then confronted with a
series of items needing explanation, among the more important of these being a
consideration of the sources of the variant morphological characters and chemo-
synthetic abilities which are present. Are these three a peculiarly assorted group
of sub-populations derived by a series of gene-mutations from within a single
species? Again-assuming a basically common origin-is there any evidence
that the differences in their chemo-synthetic abilities and such morphological
differences as are present have been the result of genic infiltrations from several
other species with which the putative basic population might have come into con-
tact, thereby giving rise to the segments of the present "Pitaya-Serrana-Pata de
gallinazo" complex? Or, has the general morphological similarity of these three
populations led me astray in my basic assumptions?:-are the chemo-synthetic
abilities of these populations of equal or even greater significance in the delimita-
tion of the specific units than their morphological manifestations? An attempt
7Considerable variance of opinion exists concerning the taste of these various alkaloids
as they occur in the fresh and dried bark. Recent studies indicate considerable differences
between individuals in their sensitivity to bitter substances. This probably explains why some
of the local cascarilleros are much more successful than others; since this ability to distinguish
bitter substances apparently is genetically controlled it also probably explains why the pr o-
fession "runs in certain families."

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will be m-ade in succeeding passages to explore these-and additional-possibili-

ties and thus perhaps ultimately arrive at a working basis for a functional taxon-
omy of the group.
V. THE ALKALOIDS OF THE PITAYA-SERRANA-PATA DE GALLINAZO COMPLEX
As is indicated by figure 1, when one goes in a general north-south direction

in Ecuador between the elevations of 9,000 and 10,000 feet there is considerable
difference in the kinds and amounts of alkaloids in the group of forms here re-
ferred to as the Pitaya-Serrana-Pata de gallinazo complex. In the two northern
provinces-Carchi and Imbabura-the percentage of the crystallizable alkaloids
is not only highest, but individual trees often contain all four alkaloids. This
is the region wherein the essentially pure stands of Pitaya are to be found.
Along the western escarpment in the middle provinces-Cotopaxi (Leon of
the older maps), Bolivar, Chimborazo, and Caniar-although considerable
amounts of alkaloid are present, it is a rare tree which yields anything but ein-
chonine. Where departures are found, these consist only of small amounts-
never more than 0.2 per cent-of quinidine. This is the region in which the
Serrana population is most typically developed.
If we now go to both sides of the crest of the eastern cordillera (and still
within the same elevation range as before) in the two southernmost provinces,
Azuay and Loja, we find that the alkaloids, have dwindled to the point where,
in some cases, no traces were even reported in the analyses.8 -This is the region
of Pata de gallinazo.
Two areas have not been mentioned in the preceding paragraphs. These are
the Province of Pichincha and the eastern part of the Province of Caiiar. It is
obvious that, in the Pichincha population, three types of individuals are present:
(1) those which most closely resemble the Serrana type of analysis, (2) those
which to a certain extent approach the Pitaya type, and (3) those which seem
to be intermediate.9 This Pichincha population, therefore, is one which is tran-
sitional between the typical Pitaya and Serrana types. Likewise, although con-
sisting of only a relatively few examples, the sample of the population in eastern
Caniar is clearly transitional between the Serrana and Pata de gallinazo types.
Although there is a marked difference, not only in the amounts but also in
the kinds of alkaloids present in the Pitaya, Serrana, and Pata de gallinazo types,
a casual glance at figure 1 would seem to indicate that there also is a rather coi-
sistent decline in the amount of total alkaloids as one goes from north to south.
If we assume that this is a single complex-aand for the moment disregard the
different kinds of alkaloids present-it would seem apparent that a definite
North > South "dline" in total crystallizable alkaloids (TCA) exists.
Since the northern end of the Pitaya complex extends well into Colombia, it
night be pertinent to ascertain if there is evidence of the continuation of a dline-
8 Additional analyses of trees with ''0.0 per cent" alkaloids were available from the
southernmost provinces, but there seemed to be little point in attempting to find space for ad-
ditional "blank analyses" on an already over-crowded chart.
9 Attention should be called to the fact that in this and other places in the figures there
are instances where cinchonidine and quinine are not differentiated. Because of critical short-
ages of certain laboratory reagents-temporary shortages brought about by the conditions of
the period-where the sum of these two alkaloids fell below 1 per cent there were times when
it was not expedient to split the tartrates and so carry the analyses to completion. Where
necessary iii the charts, these combined alkaloids are indicated by cross-hatching. At this
point I also wish to pay tribute to the excellent work and spirit of complete cooperation of
Doctors A. W. Bastress and J. A. Gandara as well as their various assistants who formerly
were in the analytical laboratory in Quito.

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IL4/{yU!mUe4~(/ -1-6
-o
The Andearn provinces (2) Prov. Imnbabur- -

NEGuaCfCor. I6.
(%)
(3) Prov Pich in ch- a6
Pd rov 6,ozolwe (5),.BaixvSd (6),i_%il

n 5t -470 010 -1 % __111 0 Qu- in
West -3 ;C1?chon
if] f]((8)RAzv y g ECiconidu
(7)ProvCca&d ? (9)PpLOV d
FIG. 1. Individual-tree analyses of the crystallizable alkaloids in representative members
of the Pitaya-Serrana-Pata de gallinazo complex occurring between the elevations of +9,000
and 10,000 feet in Andean Ecuador.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
effect in that region. On this- item I quote from Steere (12, p. 470): "In Ec-
uador, the alkaloid content of the bark [of C. pitayensis] does not seem to be so
high as in the northern parts of its geographic range, in Colombia, and it ap-
parently becomes progressively less rich in alkaloids, toward the south." In his
remarks he notes a tree from Colombia with 8.1 per cent TCA.
In addition to the single analysis listed by Steere, the officials of a similar
group working in Colombia kindly forwarded to me the analyses of 19 other
trees which had been authenticated as to species by the botanists of that group
and taken in series from the Colombian-Ecuador border northward essentially
to the known limits of C. pitayensis at about 3? North Latitude.10 While show-
ing a somewhat greater spread of variability (due primarily to the single speci-
men cited by Steere), when averaged this series of 20 trees did not seem to be
significantly different from the "stands" of Pitaya across the border in Ecuador.
In order to test the entire series in somewhat more graphic form, the popu-
lations as represented by individual-tree analyses from the various areas noted
in figure 1 were averaged and these placed as nearly as possible in relation to
their proper latitudes north and south of the Equator, together with the 20
Colombian trees (these being taken as a unit). The result of this is shown in
figure 2.
Pit yvd" 'Pdtd deyd//uzdzo"
$ t . . . . ~~~~~~Contdce ihree &f,es /s4wd/@e 4ret

Q 8- Nz/uzdxm 7CA
7 AzIhP s1nt7 of /Aese

> I / tc tn WofJd4' 9,o -O, ooofte/evleiIon,
/ ~~~~~~~~~~~~~~~~~~~WdL/Z OCc451,0nd/ Z'/-ee5 dt~
6 6- 6 (LimIts Ovfldd/trt) 8)SOO -/0jS700 /t e/ev
7b,rov,lic e a- reye 4I.

0 -Averciye TCA or v//of
the -cao-eub. i '( ,, , j g . ,, 1f:,' ~~~~~~~~~lVd-ti 'I

Co/ombid~~ ~~~~~
S5 nCotc wddd'o Peru/
~~~~~~~~uPe,- ?~
comle along th ars of th Ande fro aboutl 3? N.Lt,inClmi,o5.a.na
0~~~~~~~~~~~~~~~~~~~~~
JON. 20 /0 00 702030 0 40 Lf05 5Ld t.
Co/ombid Ecada'or Peru
FiG. 2. The dline in total crystallizable alkaloids in the Pitaya-Serrana-Pata de gallinazo

complex along the crest of the Andes from about 30 N. Lat., in Colombia, to 50 S. Lat. near
the Ecuador-Peru border.
10 Unfortunately, only the figures on the TCA of these Colombian samples were made
available to me although I particularly requested complete analyses; hence there is no op-
portunity to make a comparison.of the proportiolns of the individual-alkaloids in the Colombian
and. Ecuadorean Pitaya populations. However, it is my understanding that they are fairly
similar, the Colombian Pitaya trees apparently producing slightly larger percentages of quini-
dine than those of Ecuador.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
On the basis of this chart it would seem that, while a generalized North >
South dline in TCA does exist, it is not a simple one; instead, it appears to con-
sist of three rather distinct and fairly homogeneous populations with rather
sharp sub-clines descending through intermediate forms. Therefore, it is not
a continuous dline but, at least as indicated by the existing data, an example of
a modified or "step dline," with sharp transitional populations.
One additional point should, I think, be noted at this place. The question
might legitimately be raised whether these individual-tree samples are at all
representative of the general populations of these types as they exist in Ecuador.
Turning to the recent study of Ecuadorean barks by Martin and Gandara I took
their figures for the average TCA of C. pitayensis (Pitaya of this paper) and
Serrana and placed them on my owni chart. Although the basic data for the
Martin and Gandara study and this one are in part common to both, the methods
of sampling these data were somewhat different."1 While some of the data in-
cluded here were not used in the Martin and Gandara paper (having been ob-
tained after theirs was assembled), they had an additional series of samples
which I did not include because the basic field-data-exact locality, elevation,
etc.-were not present on the analysis slips in Quito.12 It is therefore interesting
to note (fig. 2: circles) that the averages for Ecuador as a whole as listed by
Martin and Gandara for these two barks (8, p. 186) -although taken from a
much larger group of individuals and to a considerable extent from those not
used here-are quite similar to the averages of single-tree samples of the same
types of bark from the various provinces (fig. 2: dots).13 For this reason-al-
though regretting that my data are not so extensive as one might wish them-
I dQ feel that, in general, a sufficiently large sample was taken so that the results
may be relied upon to a reasonable extent. Thus, feeling more secure about the
adequacy of the data, we are left with the question of their interpretation.
The peculiar nature of the step-dline present in the alkaloids of the Pitaya-
Serrana-Pata de gallinazo complex might easily lead one to the conclusion that
here we are dealing with three essentially different populations. Certainly,
throughout the greater parts of their individual ranges they appear to be rea-
sonably homogenous.14 The fact that these populations apparently have zones
of intergradation would not necessarily bar them from consideration as groups
worthy of recognition under some sort of separate taxonomic treatment. Such
transition populations are of regular occurrence between subspecies; and they
11 In selecting the data of individual-tree analyses for this paper, only those were chosen
which had complete field-data and which had been made from samples primarily collected
either by the professional botanical members of the exploratory group or under their personal
supervision. Since the botanical members of the group were more likely to make certain that
authenticating herbarium specimens were collected along with their bark samples, it was
thought that by so selecting the data there would be a better chance of ultimately correlating
the chemosynthetic and morphological characters of these populations.
12 For the greater part, this represents a large series of special analyses made of material
collected personally by Martin, then of the Office of Foreign Agricultural Relations of the
U. S. Department of Agriculture. The field-data on these specimens of Martin were not avail-
able to me during the time I was checking through the files of the Misi6n de Cinchona.
13 Pata de gallinazo was not listed by Martin and Gandara because it is not considered
to bo a " commercial bark," although it frequently appeared as an adulterant of such things as
Costrona fina and Urituzinga of the C. officinalis-complex.
14As noted previously, it would seem that the northern limit of the Pitaya type is at
about 30 N. Lat., in Colombia. The Pata de gallinazo type is known to extend southward to
the Ecuador-Peru border; whether it extends into Peru-and if so, how far-is not known
to me. However, it should be noted that occasional specimens from northern Peru seem to
belong here.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
are by no means unknown where regularly accepted and quite distinct species
(or at least populations usually considered to be different species) make contact.
On the other hand, as mentioned in earlier parts of this paper, in spite of the
wide differences in chemosynthetic abilities of these three types, their morpho-
logical characters are such that, to me at least, they seem to be parts of a basically
common complex. Thus, when considered against the background of the rela-
tively wide differences which are to be found between other species of the genus
as they exist in Ecuador, these three have so much in common that I do not feel
it quite proper either arbitrarily to give them different specific names (as has
been traditional in many of the previous works on the genus), or to assign the
Serrana segment to what, morphologically, is asdmittedly a quite different group
of forms-namely, the C. pubescens complex-as has recently been done. How-
ever, before we attempt any conclusion on the possible solution of the questioll,
it might be well to approach the problem from an entirely different angle.
VI. A TRANSECT OF THE WESTERN ESCARPMENT
As noted in a previous section of this paper a region called the Rio Saloya-
Corazon Pass area, located in the Province of Pichincha, was selected for a fran-
sect-study of the western escarpment of the Cordillera. Here the genus Cin-
chona exists in three rather well-marked zones. As is shown in figure 3, the
-6 P4, P4rp c/awed enecUv l//ied4i'e: o0ooa(.

_ _o/e1un7, Trc44eo/am, 'd i l
------ 0,~~~~~6tdfi-S
-4Xs d/zd VVyOqOOAOy6
pdStue CV-9,Ov 0,tl
V-iSerr4ncd A-ineer,~eidg4e O-Piz4Ya,

The Serrand -PitzV complex. x'6,-000ft1
SJ]Quinidine 7 OOOf t. Zef generd7 770

@ Cinchonidine ,Rv6ture.
| (RrbodEeso{Q3w/-)3R 4,ooDy. -3 VW i-4

Cincoie Nco&_e
Soinge-tree an4j/y
an d/ittIdin4/ trdseCt.- , )f dherrd BoJac RFR 'i
Lk Id/oy4- CordPdsscdorec. The 'bubescens co

FIG. 3. A transect of the frontal esearpment of the western Cordillera in the Province
of Pichincha, Ecuador, indicating the altitudinal zonation of the various major Cinchona
populations.
uppermost zone consists of a mixture of materials very closely approaching Pitaya

and Serrana, together with what appear to be intermediates between the two.
As has also been noted, this Pitaya-Serrana zone is best developed between the
elevations of 9,000 and 10,000 feet, with a few trees at slightly lower levels.
Below this, on essentially the middle slopes of the escarpment and primarily
between the elevations of 6,000 and 8,000 feet, the "Bofuda" type appears; as
might be expected, occasional outliers of the basic population of this type may

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
be found at somewhat lower and higher elevations. The Bofuda type is part
of the Cinchona pubescens complex."5
The lowermost of these Cinchona-zones occurs at about 4,000 feet elevation
and is characterized by the type called "Roja-roja" (for brevity in this and other
studies usually shortened to "Roja").16 The Roja type originally was given
the name of Cinchona succirubra and so appears in the majority of the older
works, although in recent years it has become the custom with certain authors to
unite it with C. pubescens. Since the present study does not include a general
consideration of C. pubescens and its close relatives (a complex of forms which,
without doubt, geographically is the most widespread and probably taxonomically
as vexing as any in the genus) there is little point in discussing the nomenclatural
position of this material. In passing, however, it is to be suggested that the
morphological characters of Roja are such that it would seem to merit rather
critical study in connection with the other segments of the pubescens-complex.
In this particular part of the transect samples from but four trees were available,
although there is considerable evidence that the Roja zone once extended over
a wider range of altitude in this particular area [Martin and Gandara (8, p. 185)
give a general range in Ecuador of from 2,500 to 4,000 feet]. It is quite obvious
that over-exploitation and clearing in this particular region have, in general,
been so extensive that it exists today only as isolated trees or at best in small
residual stands.17
On the basis of the data presented, except for individual trees, these three
populations would appear to be reasonably disjunct. H-owever, in this region
there is little opportunity of ascertaining whether this is a natural or artificial
condition for today the zones of apparent disjunction to a certain extent are
cleared and cultivated, or in some way disturbed. In the upper zone of disjune-
tion-that between the Bofuda and Pitaya-Serrana zones-the people of the
region carry on a typical high-altitude type of Andean cultivation. Among the
more characteristic of the crops are various native tuberous plants such as the
"papa" or potato (Solanum sp., several species including S. tuberosum being
involved), the "mashua" (Tropaeolum tuberosum), and the "oea" (Oxalis
tuberosa); the "quinoa" (Chenopodium quinoa) is sometimes encountered; and
secondary pastures following cultivation are also to be found. The lower of
these disjunct zones-that between the Roja and Bofuda populations-is also
considerably cleared and pastured; where the terrain permits, maize (Zea) is
grown along with other general crops.
Since but little can be gained in ascertaining the cause of these two disjune-
tions by further consideration of the data available to me from this particular
transect, it would seem pertinent to extend the study, if possible, to another
point on the same escarpment where less clearing and relatively little cultivation
exists. Fortunately, such a transect was made by Steere near the Colombian
border in the Province of Carchi. Although the original of the report on this
15 For all practical purposes, Bofuda of the northern provinees, Rosada of the central
provinces, and Zapallo, Hoja de Zambo, and to a lesser extent Rosada of the southern provinces
are names applied to essentially the same kind of tree; in my opinion these are all variant
local names for Cinchona pubescens in a restricted taxonomic sense.
16 It is well known that among those peoples with not overly extensive vocabularies-such
as the local casearilleros, the bark gatherers-it is- a common practice to designate an object
with special qualities by repetition of the name solely for the sake of emphasis. Thus, to the
casearilleros, the usually excellent "Redbark" of commerce-for which a premium was paid-
became "'Roja-roja. "
17 The Roja type is also found in small plantations along the western escarpment; analyses
of wild trees only have been included in this study.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
transect was in Quito when I went over the files of the exploratory group, there
was no opportunity to have a copy made; more recently Steere made a personal
copy available to me. With his permission portions of the data have been ex-
tracted. Before we consider this transect, it should be recalled that, in the
Province of Carchi, no Serrana type was found, and that apparently only the
Pitaya type exists between the elevations of 9,000 and 10,000 feet (see fig. 1).
The pertinent details of this report are as follows.
Extract of a report on an exploratory transect (Expedicion Q-5) in the
western Cordillera of the Province of Carei, Ecuador, by Dr. Wm. C. Steere and
E. M. Ferdon, Jr. (August 20-29, 1943).
Route: From Tulean westward (approximately) via the Volean de Chiles
paramo (14,500 ft. elev.)'and the Rio de la Plata valley to the Rio San Juan
valley in the region of the town of Maldonado (5,070 ft. elev.), and return.
The striking feature of the transect of the western slope of the Cordillera in
this region is the clear stratification 'of the arboreseent rubiaceous vegetation,
there being three zones characterized by species of Cinchona separated by two
zones of Ladenbergia. In descending order these are as follows:
Cinchona pitayensis . 10,000-9,000 ft.

Ladenbergia macrocarpa 9,000-8,500 ft.
Cinchona putbescens 8,500-7,000 ft.
Ladenbergia sp? 7,000-6,000 ft.
Cinchona sp? 5,500-4,500 ft.
The material of the uppermost layer-beginning just below the paramo zone
is apparently the same as that found earlier in southern Colombia and referred
to Cinchona pitayensis. Here for a distalnce of about 3.5 kilometers along the
Tufinio-Maldonado trail (and between the 10,000 and 9,000 ft. levels) the trees
of this well-marked species were fairly abundant, although scattered and rarely
in "manchas" (groups) as is sometimes the case.18 It appears not to have been
previously exploited in.this region.
In the Cinchona pubescens zone, herbarium specimens and a bark sample were
taken at Tambo Bella Vista. The material from this zone is morphologically
very similar (if not identical) to the Bofuda form of this species which occurs
at comparable elevations in the western Cordillera to the southward; further-
more, having 2 per cent TCA in the bark-and this almost entirely einchonine-
it also is chemically akin to the Bofuda of other regions.
The lowest of these three Cinchona zones is at present poorly defined because
of the scarcity of trees, the result of excessive exploitation during the last 100
years. It is locally called "Roja-roja" (or "Roja de Maldonado"). The leaves
are leathery but thin, ovate to round in outline, and become bright red with age;
they are somewhat similar to the leaves of C. pitayensis but have the margins
markedly revolute, a character not present in that species. Analyses of three
bark samples from trees still existing from what must formerly have been ex-
tensive stands and expressed in the per cent weight of the dry bark are as fol-
lows (the WCS specimens from the vicinity of Maldonado; the Janouch speci-
men from the nearby Cerro Golondrines):
WCS No. 7 WCS No. 8 Janouch-SC14

Quinidine trace trace trace
Cinchonidine 1.64 1.44 1.0
Quinine 1.22 0.97 0.6
Cinchonine 2.23 3.84 4.3
T.C.A. 5.09 6.25 5.9
1s Analyses of material coll

in -figure 1 of this paper.-W.H.C.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
408 Brittonia [VOL. a
For the present, this material is not taxonomically placed and may represent
an undescribed species [end of extract] .19
In discussing further the implications of the data presented in his report, in
a personal communication Steere pointed out that in the region of the western
escarpment under discussion there are three rather well defined cloud belts, one
at the low elevations, one at the middle elevations and one at the high elevations,
and that the primary populations of Cinchona appear to be limited to these three
cloud belts. On the other hand, the stands of Ladenbergita are to be found in the
intercalated zones where the cloud layers are less likely to be well developed
and, because of less "drip rain" (the result of condensation of moisture on the
arboreseent vegetation), are inclined to be somewhat drier sites. Thus, in a
relatively undisturbed area on the western slope of the esearpment in the Province
of Carchi, there are three very moist zones in which populations of Cinchona
occur and two intercalated drier zones wherein may be found species of Laden-
bergia, the latter a genus of the Rubiaceae closely related to Cinchona but ap-
parently with less exacting ecological requirements.
Returning now to the transect in the Province of Pichincha (fig. 3), it is
obvious that the three Cinchona zones have the same fundamental relationship
as they do in Carchi. Such differences in their absolute elevation as are present
may well be due to relatively minor changes in cloud levels as a result of slight
differences in the angle of slope of the escarpment and a resultant shift in air
currents, or to some other local factor, such as nearness to the moist lowlands of
coastal Esmeraldas. At least, it is obvious that three major Cinchona zones do
exist in Pichincha and those of us who have visited this area know that the pre-
cipitation in these zones is such that they hardly can be said to be suitable for
agricultural pursuits. Conversely, the intercalated zones, being somewhat drier
and therefore more suitable for agricultural crops, would be the zones most likely
to be cleared first by those seeking home sites.20 Therefore, the disjunctions of
19 Although this was then thought by Dr. Steere to be taxonomically different from the
material treated as ''Roja'' in figure 3 of this paper, the general similarity in alkaloid content
may be noted. Further study and comparison doubtless will reveal important points of rnor-
phological similarity between the Roja of Maldonado and vicinity and the Roja (C. succirubra)
of the Rio Saloya area.-W.H.C.
20 The clearing operations mentioned here are post-Colonial and for the most part quite
recent-even current. The presence of ancient building sites at numerous places on the
western escarpment-mentioned by Steere (in litt.t) as being common in Carchi, and seen
by me in other provinces-indicate that it supported a much larger human population in pre-
Conquest times than at present. Whether this means that the population pressure on the
Plateaui (and in the inter-Andean valleys) was then so great that segments of these early peo-
ples were forced to seek home sites on the relatively steep slopes of the escarpmenft, 'or that
the climate on the Plateau was then less favorable for the development of agriculture than at
present must, for the time being, remain in the realm of speculation. The fact remains, how-
ever, that the relative abundance of ancient home and temple sites on the escarpment would
seem to indicate a former population sufficiently large that it certainly must have greatly dis-
turbed the then-existing natural vegetation. If such be the case, it would follow, therefore,
that the present forest type is one which has developed after such a disturbance. The possible
effect of clearing and the development of agricultural areas followed by a cultural regression
and the return to a natural vegetation-and these sometimes in recurrent cycles-in a region
which has been peopled for thousands of years would seem to merit consideration by students
of distribution, ecology, and speciation. It is by no means beyond the bounds of reasonable
speculation to suppose that such early vegetational disturbances have been more instrumental
in fostering imbalances in gene distributions within complex species, or the elimination of
certain- of their incipient biotypes and the chance preservation of others, than we sometimes
have thought. It is entirely possible that the destruction of segments of a somewhat poly-
morphic and wide-ranging species and the spread of the few remaining and possibly quite
different biotypes into ecologically acceptable habitats after a regression of the human popu-
lation of an area may have greatly accelerated speciation in those regions where human cultures
have been cyclically persistent for a considerable period, or where they once were present.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
1948] Camp: 'Cinchona in Ecuador 409
the Cinchona zones in Pichincha seem likely to be of the same general type as
in Carchi and not entirely the result of the present cycle of clearing. However,
it should be noted that, while Steere reported no individuals of Cinchona in the
intercalated zones in Carchi, there is evidence in the Pichincha transect that at
least occasional individuals of Cinchona are to be found there.
It would seem that but little might be accomplished by speculating on the
situation before the current cycle of clearing, yet the presence of these few in-
dividuals in the intercalated zones in the Pichincha transect during the investiga-
tions of the years 1943 and 1944 do indicate that, had the clearing not been so
great, others undoubtedly would have been found. Therefore there is consider-
able reason to believe that, in the Pichincha transect, these three major Cinchona
populations are not (or were not) so completely disjunct as they seem to be in
Carchi. Let us, therefore, consider this Pichincha transect again in some detail.
[Note: Since this was set in type I have come across an item in my field
notes on the economic uses of Andean plants by the inhabitants of the provinces
,of Azuay and Loja which might be of considerable importance here. In those
areas at comparable elevations in the southern provinces of Ecuador termites
are abundant and the inhabitants consider material of the genus Cinchona to
be among their better termite-resistant woods, seeking it especially for the main
supporting timbers of their houses. (It is well known that alkaloids are present
in the wood of Cinchona as well as in the bark, sometimes in considerable quan-
tity.) The wood also is used in the fabrication of household furniture. It is
most unlikely that the areas where habitations are most abundant on the western
escarpment are termite-free; it is quite as unlikely that knowledge of the ter-
mite-resistance of Cinchona wood is confined to the peoples of the southern prov-
inces. Therefore, it would seem safe to assume that selective cutting of this
material solely for local construction purposes has tended to make it consider-
ably more scarce today in the residual forest stands in the intercalated zones
favorable to agriculture in the Rio Saloya transect than it actually was under
natural conditions.]
Attention already has been called to the heterogeneous nature of the com-
bined Serrana-Pitaya populations at the uppermost level. This population
might easily be thought of merely as a mingling of the two types in a region where
their ranges overlap; yet (and I am anticipating here) it will be shown later that
neither the part of this population designated " Serrana" in figure 3 nor the part
called "Pitaya" is exactly typical of either.
As mentioned in an earlier passage (especially footnote 15) the Bofuda type
has very close counterparts which occur essentially at the same elevation in other
parts of Ecuador, these being such types as are locally called Rosada, Zapallo,
and Hoja de Zambo. Martin and Gandara (8, p. 186) list 36 samples of Bofuda
from Ecuador having an average TCA of 2.12 per cent and 165 samples of Rosada
averaging 2.72 per cent. In both Bofuda and Rosada the only alkaloid present
was einchonine. With these two types could be listed additional materials from
my own studies of the cognate Zapallo and Hoja de Zambo from the southern
provinces; in both of these the result would be the same-a bark with relatively
low TCA, and all of it einchonine. In other words, throughout considerable
areas in Ecuador at what might be called the middle elevations there exists a
Cinch6na population which morphologically is reasonably uniform, the bark of
which is low in TCA and (at least in more than 200 of the samples known to me)
one which contains no alkaloids except einchonine. This Bofuda-Rosada-ete.
group, therefore, in spite of its various local names, is one of the more stable and

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
widespread Cinchona populations in Ecuador; that is, except in certain local

areas-and one of these happens to be in the Rio Saloya-Corazon Pass transect
here under consideration.
Returning now to figure 3, it is immediately apparent that the population in
the middle elevations is made up of individuals of two types: those which might
be classed as rather typical Bofuda and those which, for want of a better name,
I have called " aberrant Bofuda. " Thus, if this apparently basic and widespread
type is so uniform in other places, we are confronted with the problem of at-
tempting to explain its aberrancy in this particular area.
To date so little actual published data are available concerning the ability
of a wide series of species of Cinchona to hybridize that one can do little more
than speculate.21 However, in reviewing the present situation, certain items be-
come evident:
1. At the middle elevations in Ecuador a wide-ranging form is present; this
is C. pubescens in a restricted taxonomic sense (i.e. Bofuda and its cognate forms
which, as has been pointed out, are not so much different types as different local
names applied to the same type of tree).
2. Throughout the greater part of its range this material is essentially homo-
geneous, having-because of its community of morphological characters and
chemosynthetic abilities-very much the appearance of a single, wide-ranging
biotype.
3. In the Pichincha transect at an altitude where we logically would expect
to find only this basic [eu-] pubescens type, we find additional material which,
chemically at least, is aberrant.22
4. Also, there is considerable evidence in this same transect that, although
the population at the middle elevations still tends strongly to be basically dis-
junct from the lower (Roja) and higher (Serrana-Pitaya) populations, the dis-
junctions are lot (or until recent clearing were not) complete. That is, there
is evidence that prior to the recent clearings at least a few individuals bridged
the gaps between the basic Roja and Bofuda populations and between the Bofuda
and Serrana-Pitaya populations.
5. The aberrant individuals of the middle-elevation population in the Pichin-
cha transect-in at least their chemical picture-have the appearance of being
basically Bofuda, to which have been added genetic potentialities for the syn-
thesis of alkaloids not possessed by the Bofuda ( [eu-] pubescens) segment of the
population.
From the foregoing array of observations and facts I can come to only one
conclusion: that the aberrancies present in the middle-elevation Bofuda popula-
21 Since this was written I have been in receipt of data concerning the amount of inter-
specific hybrids which recently have been produced under controlled conditions in Cinchona.
These involved the types classed in literature as C. succirubra, C. calisaya, C. ledgeriana, and a
series of spontaneous hybrids; in one year alone about 900,000 seed were produced, adequate
samples of which indicated a very high fertility. The confidential nature of these data does
not permit further discussion prior to its publication by those immediately concerned, except
to say that in the materials involved there appear to be few, if any, genetic barriers to inter-
specific crosses. This merely confirms experimentally what seemed to be apparent everywhere
I carefully examined wild stands: that, in Cinchona, the only real barrier between species is
distributional (primarily altitudinal) disjunction.
22 Since I did not personally explore this particular population in detail in the field, but
made only brief visits to the various elevations, its modulus of morphological variability is
not too well known to me, although certain differences in leaf shape and pubescence were
noted; however careful examination of such herbarium material as possibly can be assembled
in the future very likely will show statistical differences. This is a task for those who will
be -dealing with the nomenclature of the group.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
tion in the Rto Saloya-Corazon Pass transect in Pichincha are the result of the
action of genes which have migrated into it, either from the low-elevation Roja
population or from the Pitaya members of the high-elevation population, or pos-
sibly from both.
As may be seen in figure 3, the aberrant members of the Bofuda population
are rather evenly scattered between the elevations of 6,000 and 8,000 feet. Had
they been localized at the lower level, one logically might conclude that they were
the result of gene infiltration from the Roja type; conversely, had they been
primarily at the upper level, one might suppose that they were caused by genetic
elements which had drifted downward from the Pitaya segment of the Serrana-
Pitaya population. Since in figure 3 the block diagrams of the individual-tree
analyses are not arranged in altitudinal sequence but in order of their TCA
(this for ease in constructing the diagrams and also as an aid in more rapidly
visualizing the extent of variability in the segments of the several populations),
T/ze '[3oJcf I Reo/dtio- - Sd/oyd -CordZcW Rs5 ?dre.
E/evdtio"s i? feet.
FIG. 4. Individual-tree analyses of the alkaloids of the "Blofuda" population in the
iRio Saloya-Corazon Pass area arranged according to their altitudinal occurrence. The symbols
are the same as those in figure 3.
the individuals of the entire Bofuda population have been brought together in
a single diagram and arranged in order of their altitudinal occurrence.
An examination of this diagram (fig. 4) seems to yield little if anything be-
yond what we already know-that the aberrant members are scattered about
evenly through the Bofuda population. However, another line of attack is yet
open to us. This is a consideration of the morphological characters, of these dif-
ferent populations.
On1 the situation regarding the morphological characters I have this to offer.
One of the members of my own group working on the western escarpment in the
Province of Caniar encountered a few individuals at appropriate elevations wvhich
seemed to bridge the gap between Roja and Bofuda, not only in their morpho-
logical characters-and these admittedly are not overly strong-but also to some
extent in the type and amount of alkaloids. While I have- made no extensive
personal study of the Pichincha transect, I have seen representatives of it down
to about the 5,500 ft. level. What struck me in this hasty examination in the
field was the presence of " problematical " individuals. This observation is
borne out by certain field notes which appeared on the sheets from which these
analyses were taken. In addition, as I search further through my notes taken in
Quito, I find one analysis made from material collected in this same general region
at an elevation of 8,600 feet and which was of such a nature that the collector

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
-considered it impossible to identify. (Because of this the specimen was not

figured in the preceding diagrams, but does appear in figure 5, where I have taken
the liberty of calling it "Bofuda x Pitaya. ") Thus, until cytological and breed-
ing studies prove otherwise, on the basis of both the chemical and morphological
characters, I feel that it is permissible, for the present, to proceed upon the as-
sumption that the aberrant members of the Bofu da population in this Pichincha
transect are the result of genie infiltrations, almost certainly from the Pitaya
members of the Serrana-Pitaya population (or more probably from its pure
Pitaya precursor), and possibly also from the Roja population. It is to be em-
phasized here that, although a snmall amount of evidence points in that direction,
I am much less certain of the Roja x Bofuda combination than I am of the Bofuda
x Pitaya combination.
VII. AN EXPLANATION OF THE ABERRANCIES MET WITH IN THE

RIO SALOYA-CORAZON PASS TRANSECT
Since information is lacking on the cytogenetical situation in these various

populations, it is perhaps unwarranted to proceed too far solely on the basis of
the available data. Appearances in the field at times may be quite deceiving.
Yet, even after having taken part in advocating caution regarding a too hasty
conclusion that hybrid swarms are present when the possibility of a population
of segregative polyploids may be involved instead, I do feel that much can be
gained by proceeding with the hypothesis that the aberrancies in this transect
are of hybrid origin. At least this gives us a vehicle for discussion, although it
may be necessary to modify it should future cytological data prove otherwise.28
In this connection, however, the recently added footnote 21 might be consulted.
As mentioned in a preceding paragraph, I am not thoroughly convinced that
genlie exchanges have been active between the Roja and Bofuda populations, al-
though there is some evidence for it. Having had some experience with the
structure of polyploidally complex populations in other groups (Camp 1, 2) the
thought has come to me that, in view of their morphological similarities, Roja
and Bofuda might possibly be phyletically rather closely related but-at the
present time-genetically disjunct because of some form of polyploidy. How-
ever, this is merely a suggestion and certainly not a conclusion. The point is-
and this will become evident in succeeding paragraphs-it does not seem necessary
to include Roja in a discussion of the possible causes of the obvious aberrancies
present in the other members of this transeet.24
Thus, although Roja is included (even if there seems no need to do so), we
may proceed directly to figure 5, which will be used as a graphic basis for this
23 In a paper on the structure of species [Camp and Gilly (4, bottom of p. 363 and first
two paragraphs of p. 364)] it was pointed out that the incidence of a few autopolyploids in
the population of one species may give rise to simulative individuals which can hybridize with
those of another species on a different ploidal level and so make it appear as if the two basic
species have produced hybrids; also the nature of a segregative allopolyploid population is
such that it often has the appearance of a hybrid swarm.
24 It is interesting to note, however, that, although Martin and Gandara (8, p. 186) list
no quinidine as being present in their 31 samples of Roja, one of the four trees at the base
of the Rio Saloya transect (fig. 3) had an appreciable amount (0.5%) of this alkaloid. In
spite of the fact that no evidence (beyond an occasional "trace") of quinidine was noted in
tho "aberrant Bofuda" of this transect, one can but wonder whether the genetic ability to
produce this alkaloid might not have been transferred in some manner from the Pitaya popu-
lation-where quinidine usually is present-through the Bofuda population and thence to the
Roja population where, through some linkage effect, there was opportunity for the expression
of this potentiality.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
part of our discussion. In this diagram a division was made only between ein-
chonine and the other alkaloids (i.e. quinine, einchonidine, and quinidine).25
The block diagrams represent averages for various of the populations and popu-
lation segments dealt with in previous parts of this paper.
If, for the present, we should assume that hybridization between the Pitaya
and Bofuda types is possible and that the resulting hybrids are not completely
sterile-and the evidence for this in so many families of plants is now so ex-
tensive that we no longer need to cite examples; see again footnote 21-we would
expect to find intermediate hybrid types, or at least individuals exhibiting char-
acters derived from both parents. Also, it might be expected that such individ-
6-
5-
4 ~-
3-~~~~~~~~~I
09NO 2-?,',P
Reid Bo/udd Itrn1Ntd
6errd id Pit dyd """Y't
PicAh'cha Pic/u IW'h4 Pi/c/c p/ma l
JrX,00?ft. 6,000-8,o0oJ. 8,600/1. +9,000-70,000fi.

- Cinchonine -Other al/AM/oid (.e. Q^inne, Cinchondinee fivzdzie) |
FIG. 5. A comparison of the alkaloid types in various Cinchona populations in Ecuador.

These are averages, with the exception of one marked Bofuda x Pitaya, which is an individual
tree. For a complete explanation the text should be consulted.
uals would be most abundant where the parental types make (or have made)
coitact. Unfortunately, in this particular transect, this happens to be one of
the cultivated and disturbed zones (8,000-9,000 ft.; fig. 3). However, as in-
dicated previously (Section VI), one such collection was made at an elevation
of 8,600 feet in this region; the analysis of this specimen appears in figure 5,
where it is labeled "Bofuda x Pitaya." There is no intimation that this is an
F1 individual; in fact, there is considerable chance that it is some sort of segregate
or back-cross. With such individuals as this one at 8,000 feet (and presumably
others previous to the present cycle of clearing) bridging the gap between Pitaya
and Bofuda, it is not difficult to think of the genetic elements which influence
(or control) the synthesis of the alkaloids other than einchonine as having drifted
downward into the Bofuda population, there producing such individuals as I
have termed "aberrant Bofuda."
Now if, as hypothecated here, genetic elements characteristic of Pitaya have
25 This was made necessary because of the numerous instances where einchonidine and
qujinine were not differentiated-see footnote 9.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
entered the Bofuda po

gene-flow, a reciprocal exchange should have takeni place. lIn an earlier passage
of this paper (Section IV) an excerpt from Steere's paper was quoted, a portion
of which will be repeated here. In contrasting the Colombian and Ecuadorean
Pitaya types he says: ''In Ecuador . . . the leaves are more pubescent, especially
when young . . etc." Since Cincho'na pub escens (Bofuda) is very aptly named
it is my opinion that here, in Steere's remarks, we have at least an important
indication- that genetic elements of Bofuda (in this instance those influencing
the development of pubescence) have entered the Pitaya population. It w-ould
therefore seem evident that there has been a mutual exchange of genes between
the Bofuda and Pitaya populations. Lacking evidence to the contrary, it is my
opinion that these two are homoploid and, at least to a certain degree, mutually
interfertile. That they have not become a common (blended) populatioll mav,
I think, be laid to the fact that the intervening zone, in general, does not have
the requisite ecological conditionis favorable to the building up of an extensive
Cinchona population-i.e., that the "biotic bridge" between the two, while strong
enough to permit a certaini amounit of gene exchange, is not (or was not.) suffi-
ciently, well developed to permllit a free and continuous flow of hereditary nma-
terials between the two basic populations. Linkages also mav have been very
importanit, but it is obvious that those genes determining pubescence and the
t!ypes of alkaloids synthesized are not liniked with those determining, ecological
preferelnces.
The other aberrant portion of this transect is the Serrana-Pitay a population
between the elevations of 9,000 and 10,000 feet. As may be noted in figure 3,
I have split this high-elevation population into three segments-"'Serrana,"
"'intermiiediate,'" and "'Pitava.'" This was donie somewhat arbitrarily for, as
mav be seen in figure 5, the 'Serrana" of this Pichincha population, while havinig
onlv cinchonine, averages higher in TCA than is characteristic of this type in
the Provinces of Cotopaxi, Bolivar, and Chimborazo, where Serrana is most
typically developed. Like-wise, in Pichincha, the "Pitaya" members of the pop-
ulation, while closelY approaching the typical Pitaya of the neighboring Province
of Imbabura in TCA, have a much higher percentage of cinchonilne thani is to
be expected in this material. The range of variabilitv in the "intermediate"
members of this population is not shown in figure 5 but may be seen in figure 3.
Again, in figure 3, it may be noted that the "Serrana," ''intermediate,' and
"'Pitaa'a" indcividu Is are about evenly] scattered in the general hioh-elevation
population.
In examining(r these data, it therefore would appear that, in Pichinlcha, wve do
not lhav e a simple mixture of inidividuals of genetically pure Serrana anld Pitaya
types but, instead, a populationi exhibiting a pattern which wNould indicate at
least a partial hereditary blendling of these two. Ilere, the blending is of an
initerestinig type anid may indicate certaini genic reactions, for quite separate
factors seem to be operative. Onie of these, as shown by the so-called "'Serrana"
segment, is the tenidelncy toward the synthesis of a greater amount of alkaloids
(TCA) than is to be expected in this tvpe; in this character it shows the influenice
of the hiigh-vieldinig potentiality of typical Pitaya. Converselv, although higher
in TCA than one would expect for straight initermediates between pure Serrana
and Pitaya, the so-called "Pitaya" members of the Pichincha population conitain
proportionately much more cinehonine than we would expect if they, were lheredi-
tarily pure; in this thev tend toward the real Serrana type. The implicationi of
the possible nature of the reaction of these gene complexes xvill probably be of
solmie initerest to those concernied with the genetics of Cinchona.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
1948] Camp: Cinchonwa in Ecutador 415
When we consider the regionally compact nature of this population, it is ob-

vious that at least partially segregative factors have been involved in producing
the present pattern of the Serrana-Pitaya complex in Pichincha. The view-
point expressed here concerning the hereditary control of synthetic abilities is
at considerable variance with the great bulk of the literature on Cinchona-some
of it quite recent, even current-where stress has been laid on climatic, edaphic
and site factors. To be sure, such items can and do influence the vigor of a
plant and, secondarily, its ability to synthesize greater or less amounts of ma-
terials such as alkaloids; this has been demonstrated repeatedly in plantation
culture. But there is no evidence that these ecological factors influence the basic
ability-pattern of the plant to synthesize different kinds of alkaloids. To a very
large extent these would seem to be hereditarily controlled and it is my opinion
that the aberrant members of the several populations in this Pichincha transect
are indication of it. It also seems likely that the differences between the Colom-
bian and Ecuadorean Pitayas, noted by Steere, may be the result of the migra-
tions of a few Bofuda genes, via Serrana, into the Ecuadorean segment of the
Pitaya type.
VIII. THE POSSIBLE ORIGIN OF THE SERRANA TYPE
As pointed out in the preceding section, there seems to be considerable evi-

dence that hybridization is possible between the Bofuda and Pitaya types, ac-
companied by the production of both intermediate and segregate forms. So far
as the alkaloids are concerned, among the segregates we might logically expect
to find those which, like the Bofuda ancestor, produce only einchonine but which,
tending toward the high-yielding Pitaya ancestor, would have a higher amount
of this alkaloid than Bofuda. Likewise, on the morphological side, we would
expect to find a spread of variability in the segregating progeny. Furthermore,
"preference" as to habitat-or, rather, the ability to compete and survive in
particular habitats and under certain coniditions-also would be segregative.
Those who have worked with large populations containing hybrids, back-crosses,
segregates, and genic introgressions, both under experimental conditions and in
the wild, are aware of the potentialities for natural selection out of such hybrid
complexes; these are not so evident if one works only with material in the her-
barium. In the field, the variability of certain of these populations of Cinchona
seemed to be analogous to other complex ones derived from hybrids met with else-
where [Darrow and Camp (5) ; Camp (2) ].
Therefore, it would seem to be reasonable to envision, at some time in the
past, at least a small populatioha of hybrids between Bofuda -nd Pitaya, possibly
in the general region where they still seem to make slight contact, as for example
in what is now the Province of Pichincha (fig. 2). Among the segregates of
this population there undoubtedly would have been those which carried the po-
tentiality for synthesizing primarily einchonine, but in larger quantities than
the ancestral Bofuda type. Of these there would be some which, to a considerable
extent, had the life-form (morphological similarity) and ecological requirements
of Pitaya. These would be the ones most likely to persist in the uppermost
(9,000-10,000 ft.) Cinchona zone. And it is my opinion that just such individ-
uals were the nucleus out of which the present Serrana population developed
(fig. 5). Theoretically, at least some of the pubescence of the Bofuda ancestor
might be expected to appear in the Serrana population unless it happened to be
associated by linkage with some factor which was deleterious. Apparently pu-
bescence itself is not a limiting factor nor does it seem to be linked to any which
is, for, on the whole, the Serrana population is more pubescent than Pitaya, but
less pubescent than Bofuda.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
The question immedia*ly appears: If the Serrana population arose in what

is now the Pichincha region of the western escarpment, why did it spread south-
ward and not northward? In the first place it is not necessary to suppose that
the Serrana type did arise in this immediate area; it may well have had its in-
ception in another area somewhat to the south, possibly at a time when the Pitaya
population was more widespread than it is today. There is, of course, also a
possibility that the present Serrana population may be polytopic-that it de-
veloped in several places along the edge of the western escarpment between
what now are the Provinces of Pichincha and Bolivar; this would demand the
one-time presence of extensive stands of Pitaya in the middle provinces.
It certainly would be unwise to exclude the possibility of a polytopic origini
of the Serrana type from our considerations, but until better evidences are pre-
sented of extensive stands of Pitaya in the high-elevation populatikn along the
western escarpment in the Provinces of Cotopaxi, Bolivar, or Chimborazo, I must
assume that the southern limits of the effective breeding populations of Pitaya
are-and for some time have been-in the Pichincha region.26 Therefore if we
think of Serrana as having originated subsequent to Pitaya, the disseminules of
the original Serrana individuals would have found the more favorable upper-
level Cinchona-habitats to the northward already occupied by the Pitaya type.
Conversely, since no other high-elevation species of the genus is known to me to
the southward until we come to the Caniar-Azuay border, the Cinchona-habitats
would have been more open to colonization in that direction. As a consequence
the newly developed Serrana type would have been more successful in the central
provinces than in the already occupied Pitaya territory northward. Naturally,
other factors such as deposits of volcanic ash, or local glaciation, could-and as
we will later see did-have considerable influence on the migration, persistence,
and present occurrence of these types. However, regardless of the factors in its
spread along the western escarpment, Serrana is the arboreseent rubiaceous type
which today dominates the high-elevation Cinchona zone in the middle provinces
of Ecuador.
Before we leave this Serrana population an additional item should be noted.
Reference to figure 1 will show that, of the 35 specimens from Cotopaxi, Bolivar,
and Chimborazo, 5 contained minute amounts of quinidine. When averaged into
the, general population from these three provinces the amount of this alkaloid was
so small that it could not satisfactorily be shown in figure 5 and so was omitted
from that.diagram. However, it does seem significant that a few of these Serrana
trees did contain even small amounts of this alkaloid in addition to the charac-
teristic einchonine. Since, as may be seen in figure 1, the majority of the Pitaya
individuals contain small amounts of quinidine, the appearance of this alkaloid
in at least a few Serrana individuals would seem to indicate a distinct and sep-
arate set of genic controls for the synthesis of these and various other kinds of
alkaloids. Further examination of figure 1 also will show that certain of the
individuals of the Pichincha high-elevation population have significantly greater
amounts of quinidine than do the members of the typical Pitaya population in,
for example, the Province of Imbabura. Since quinidine is a ohemical isomer of
quinine (as einchonidine is of einchoniiie), one cannot help being a little curious
26 There are rumors of trees in the central provinces alkaloidally somewhat like Pitaya,
but I have no exact records of them. Furthermore, were they present in appreciable quantities,
I feel certain that there should be areas where populations similar to that of the 9,000-10,000
ft. elevation in the Province of Pichincha would have been built up. These also appear to be
absent. The presence of quinidine in the bark of a few trees in the central provinces will be
noted and discussed in the text.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
as to the exact nature of the genie balances which control the synthesis of these
two pairs of alkaloids. Lacking data on such items, all one can do is to reaffirm
a strong belief in the efficacy of hereditary predetermination in these matters.
Certainly an examination of the populations in and surrounding this Pichincha
complex gives considerable evidence that genetic factors rather than climatic,
edaphic, or site factors, are primarily responsible for their differences. The
least that can be said is that the presence of quinidine in various memnbers of the
Serrana population would seem to be another indication of its genetic link with
the Pitaya population. To me, the Serrana type appears to be no more than
Pitaya which has undergone slight morphological changes coupled with a modi-
fication of its synthetic abilities as a result of gene exchanges with Bofuda.
IX. THE POSSIBLE ORIGIN OF THE PATA DE GALLINAZO TYPE
As previously noted, another form which I have associated with the Pitaya-
Serrana complex occurs at comparably high elevations in the southern provinces
of Ecuador. In the Province of Azuay, this is called Pata de gallinazo (or some-
times Pata de gallinazo blanco) ; in Loja, the southernmost province of Ecuador,
it is usually so rare that many of the casearilleros have no name for it and those
who have one call it either Pata de gallinazo (if they have worked in the Azuay
forests), or "Hoja de Luema redonda," thus differentiating it from the rather
rare but more valuable Loja bark called "Hoja de Luema" (Cinchona lucumae-
folia), a tree of lower elevations. Because of the great similarity of these forms
as they appear in both Loja and Azuay (and for at least a small distance in ad-
jacent easternmost Cafiar)-and to avoid confusion in a multiplicity of local
names-they will here be referred, to collectively as Pata de gallinazo.
There are indications from several small "hand samples" of bark brought in
by the Cholo residents that both Serrana and Pata de gallinazo may yet be found
on the escarpment of the western Cordillera as well as on the "nudos" (trans-
verse inter-cordilleran ranges) in these southern provinces. However, there is
considerable evidence that they are not so well developed there as in other areas-
Serrana to the northward on the western escarpment and Pata de gallinazo south-
ward on the crest of the eastern Cordillera.
The clearing of the plateau and transverse inter-Andean ridges in this region
is so extensive that but few stands of Cinchona (or for that matter other forest
types) remain, yet it is easy to eilvision several former migration routes for the
Serrana type of the western Cordillera across the Plateau to the eastern Cordil-
lera in the region of the Provinces of Caniar and Azuay. Here the Plateau and
transverse ridges are at the requisite height for this type. Furthermore, al-
though it has often been stated that Cinchona is limited to steep slopes, this is
not necessarily so, for I have occasionally found small stands of Serrana on rolling
or even essentially level areas on the Plateau in central Cafiar; directly opposite
-and north of the Rio Paute-in eastern Cafiar, Serrana was found, together
with what appeared to be Pata de gallinazo, on what certainly is mountainous
terrain but, for the Andean region in Ecuador, not particularly steep slopes.
These great transverse ridges ("nudos") in Cafiar and Azuay would, therefore,
have given ample opportunity for a population of Serrana migrating southward
to cross from the escarpment of the western Cordillera to the eastern Cordillera.
At least this much is obvious: in the eastern Cordillera in the southernmost part
of the Province of Cafiar, what morphologically appears to be Serrana is found
mixed with aberrant Pata de gallinazo; the alkaloid picture of this population
(fig. 1, 2) bears this out. At this nnint-iust south of the Caniar-Azuay boundary

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
-the eastern Cordillera is cut by the Rio Paute, and on the heights south of
its great gorge in Azuay I made definite contact with abundant stands of the
Pata de gallinazo type.
During the first year of our work in the southern provinces, members of my
own exploratory group and I had encountered the Pata de gallinazo type in var-
ious parts of the eastern Cordillera, first in the Province of Loja and later in
Azuay, and each time it was ascertained to be so worthless-because of its ex-
ceedingly low alkaloid content, or even the complete absence of any crystallizable
alkaloids-that there was no point then in expending the time sufficient for a
careful study of its distribution. It was not until my last six months in Ecuador
that an opportunity presented itself to study this material in any detail in the
field. During that period I made a point of returning to the eastern Cordillera
in the Province of Azuay and for a period of about forty days was in a region
where various segments of the local Cinchona population could be studied.27
During that period of study it became obvious that Pata de gallinazo was a
form which, among the galaxy of Cinchona types in the southern provinces, de-
served special taxonomic attention. Also, as detailed in an earlier passage, it
appeared, on the basis of its. morphological manifestations, that this material
merited serious consideration as a segment of the high-elevation population better
developed in the central and northern provinces and here collectively called the
Pitaya-Serrana-Pata de gallinazo complex. Again, I do not mean to imply that
a statistical study will not reveal points of morphological difference between the
Pata de gallinazo and Serrana types, any more than I have said that there are
no differences between Serrana and Pitaya. The point which I have tried to
make is this: looking at the pattern of variabilities in the general Cinchona popu-
lations in Ecuador-and I have seen examples of the majority of them in the field
-it is my opinion that the three here under consideration form a reasonably
well-defined unit. If the Pitaya, Serrana, and Pata de gallinazo types are not
to be considered as different specific entities among all the other species of Cin-
chona, certainly it would seem that the Serrana and Pata de gallinazo types
deserve in some manner to be treated as being more closely related to Pitaya
than to any of the others. Although these three differ greatly in the kinds and
amounts of alkaloids which they produce, their morphological manifestations are
such that they might easily fall within the limits of a single and not overly com-
plex species. Furthermore, such morphological differences as these forms exhibit
seem to be the result of not overly large genie infiltrations from other species; and
what appear to be rather large chemtcal differences may very well be the result of
a difference in relatively few genes.
As has been noted, the Pata de gallinazo type appears to be best developed
along the eastern Cordillera in the two southernmost provinces. There it does
not occur in a characteristically broad band as do the Serrana and Pitaya types
in the northern provinces; instead, the population is rather limited and some-
times attenuated, usually being little more than a thin line of trees or small
27 The period of study in this particular area extended from the latter part of July to
the first part of September, 1945, during the height of the rainy season in that region. Those
who have traversed the eastern Cordillera of the Andes in Ecuador at that season will under-
stand the kinship I felt with Noah who, it seems, also had some slight experience with forty
days and nights of rain. Also, much like the famous Ark, the little mudwalled Cholo cabin
in which I set up headquarters on the slopes above the villages of El Pan and Sevilla de Oro
had its quota of pigs, ciickens, guinea-pigs, sheep, cows, horses, mules, cats, dogs, fleas, and
other more or less domesticated Animalia which wanted to-and often did- 'come aboard"
to get out of the rain.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
1948] Camp: Cinchona tn Ecuador 419
"maniehas" (colonies) and these often scattered and with large gaps. It was
very difficult to come to any conclusion concerning this type in the Loja region
because of the extreme over-exploitation of all types of bark-even the worthless
ones such as Pata de gallinazo. However, in the eastern cordillera of Azuay,
where the extermination of individuals had niot been so great and also where
there was more opportunity for detailed field study, it became evident that the
Pata de gallinazo type had an apparently serious competitor within the genus
Cinchona for at least the upper part of its normal altitudinal range. A similar
competitor was present in the Loja region.
In the eastern Cordillera of Azuay, between the elevations of 10,500 and
9,500 feet, or sometimes even lower, we encountered large stands of a type called
"Crespilla." This well-marked form is quite different from Pata de gallinazo
and is to be found illustrated in Howard's classical study of the genus (6) under
the name of Cinchona rugosa or "Casearilla crespilla de Cuenca. "28 The illus-
tration in Howard's work is reasonably representative of this type, although the
leaves are not always so elongate as the figure would indicate.
This material of C. rugosa and its variant forms must be discussed at greater
length in another place. It is sufficient here to state that while making a special
study of the transition zone where Crespilla de Cuenca and Pata de gallinazo
make contact I came to the conclusion that no sharp line of demarcation could be
drawn between them. This was not because of any great similarity between them
for, basically, they differ by a whole series of excellent characters. Above 10,000
feet, Crespilla was reasonably clear-cut and with little variability. The same
might be said for Pata de gallinazo at about the 9,300-9,500 ft. elevations in cer-
tain areas.29 But in elevations between 9,500 and 10,000 feet there were areas
where both Pata de gallinazo and Crespilla in apparently "typical" form were
found together with such complete series of intergradent individuals-and these
in all manner of character combinations-that I could come to no other conclusion
except that these two had hybridized freely. It is of further interest to note at
this point that at no time did we get specimens of the true Crespilla de Cuenca
type which gave any indication of alkaloids beyond the merest traces.
Let us now, for a moment, return to the Pichincha transect discussed earlier
and recall the presumed effect of Bofuda genes on Pitaya. There the result ap-
parently was the Serrana type, a form rather much like the Pitaya type in ap-
28 Cuenca, the capital of the Province of Azuay, and my general headquarters for over a
year, was once an important center for the exploitation of bark in this region.
29 It will perhaps be noted that I have placed the lower part of the stable portion of the
Pata de gallinazo population in certain areas somewhat above the usual lower limits for mem-
bers of this complex for, theoretically, it should go down to 9,000 feet. Here in the eastern
Cordillera, another species-" Costrona fina," a member of the " Cinchona officinalis complex"
-is present just below the Pata de gallinazo zone. From what data I have, this type seems
to have contrib'uted aberrant members (via hybrids and genie introgressions) to at least the
lower portions of the Pata de gallinazo population. Actually, this narrow belt wherein may
be found rather typical members of the Pata de gallinazo population is not a true zone, fox
it varies considerably as a result of what, in the field, appeared to be both genic interference
and population dominance from both the Crespilla and Costrona fina types. In certain areas,
because of the configuration of the terrain, and possibly as a result of previous clearing and a
re-establishment of the forest, the Crespilla population sometimes drifts downward through the
Pata de galliiiazo population, almost to the Costrona -fina zone; in other local areas the reverse
appears to have taken place, with the Costrona finla approaching the base of the normal Crespilla
zone. The result has been a "pinching out" of segments of the Pata de gallinazo zone, ac-
companied by the production of locally abundant, mixed (in part hybrid) populations. It is
quite likely that the specimen on which the Howard plate of Cinchona macrocalyx (6) was
based came from this hybrid complex.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
420 Brittonia LVOL. 6
pearance (excepting a slight increase in pubescence and a few other minor

aberrant characters), the most notable change being the inability of the new type
to synthesize any alkaloids except einchonine (again, except for a few individuals
able to produce very small amounts of quinidine). It is, therefore, possible to
suppose that much the same thing might have happened in Caniar where, so far
as the data indicate, the southward (and at that point also eastward) migrating
Serrana met the alkaloidally barren Crespilla type. In this instance it would
appear that the genie introgression-which, from the presence of apparent hy-
brids and back-crosses, I must suppose took place-resulted in the almost complete
loss, in the Serrana type, of ability to produce crystallizable alkaloids, the gross
result of this exchange being the Pata de gallinazo type of individual. At least,
as shown by figures 1 and 2, there is a marked break in the synthetic ability of
this Serrana population in eastern Caiiar in the region where contact is made
with the edge of the Crespilla population. In fact, as one looks at these two
diagrams, one can but wonder whether this alkaloid-depressing effect of the Cres-
pilla genes-if it be such-might not have migrated backward (by gene-flow)
across the transverse ridges through the bridgihg population (which once must
have existed) and left its mark on the Serrana population of western Cainar.
Certainly, although the specimens are fewer than one would wish, they do average
less TCA in western Cafnar than the basic Serrana population to the nlorthward
(see fig. 2).
Southward, in the Province of Loja, Pata de gallinazo (locally called lloja
de luema redonda) is also found. There, because of the extreme over-exploita-
tion in former years of all types it is no longer abundant. Yet, there, even as in
Azuay, it has (or did have) a potent competitor in the high-elevation Cinchona
population. This is the "Crespilla de Loja " of the modern casearilleros, which
is the same as the "Casearilla Crespilla con Hojas de Roble" or Cinchona micro-
phylla of the Howard plate (6).30 Also, it is to be noted that-just as in the
Crespilla de Cuenca-at no time were we able to obtain specimens of this Crespilla
de Loja which yielded more than mere traces of any crystallizable alkaloid.
In conclusion then, it is my opinion that these high-altitude Crespilla forms
of the southern Provinces have acted as a "brake" on the Pata de gallinazo pop-
ulation, both of them being adapted to conditions somewhat above and within
the upper levels of the normal Pata de gallinazo zone, thus restricting its
development into an effective unit-population, although not completely excluding
it from the general Cinchona-flora of the region. Furthermore, it would seem
that these-through hybridization and subsequent gene exchanges-have had a
depressing effect on the ability of the basic type to synthe'size alkaloids, in most
individuals almost completely eliminating it. Morphologically, the Pata de gal-
linazo type is still very much like Serrana, yet it has a certain cast which is all its
own. In my opinion this may be laid to the genes which it has acquired from its
neighbors which, as outlined in the general text of this section, are the Crespilla
de Cuenca on one hand and (as briefly mentioned in footnote 29) Costrona finia
on the other. As noted earlier, there is some slight evidence that Pata de gal-
linazo extends southward along the eastern Cordillera into Peru, although I have
no personal knowledge of it beyond the present Ecuador-Peru border.
30 C. microphylla and C. rugosa appear on the same unnumbered plate in this work. As
seen in the field, the populations of these are quite distinct from each other;' they apparently
make contact with a zone of intergrades in the eastern Cordillera opposite Onia, a small -village
approximately midway between the cities of Cuenca and Loja. It is my opinion that the
'common name I listed by Howard for C. microphylla 'is a spurious one given it by Europeans;
it is not used today by the casearilleros and, for that matter, I never even heard them use the
word "Roble" and think that this Spanish name for oak is probably not in their vocabularies.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTCC
Before we turn to other items, another observation should be recorded. While

yet in the field it occurred to me that, morphologically, Pata de gallinazo and
Crespilla de Cuenca are serially rather intermediate between the widely different
Crespilla de Loja and Serrana, at least with respect to leaf shape, pubescence,
and the presence of a glossy sheen on the foliar structures. Therefore it does
not strain one's imagination too greatly to think of Pata de gallinazo possibly as
the Serrana-like segregate type (which also has much of its ecological prefer-
ences), and the Crespilla de Cuenca as the reciprocal segregate type morpho-
logically and, especially ecologically, more like its Crespilla de Loja ancestor.
X. ON THE HISTORY OF THE GROUP AND THE NATURE OF THE CLINE IN ALKALOIDS
IN THE PITAYA-SERRANA-PATA DE GALLINAZO COMPLEX
Although it is often thought that a clilie or gradient-series of characters should

be continuous within a population, H-uxley (who gave us the term) notes that
they sometimes may be in the form of a "staircase or stepped ramp" (7, p. 211,
etc.). As pointed out in section V of the present paper, if we are to consider
the group of forms called Pitaya, Serrana, and Pata de gallinazo as being parts
of a single-species complex (and I see no other simple solution to our problem)
then the North > South dline in total crystallizable alkaloids (TCA) present in
this series is of the "step" or "staircase" type. On the basis of the alkaloids,
other descriptive N > S dlines could be recognized, such as the number of dif-
ferent alkaloids synthesized, or the percentage of individuals which contain
quinidine (see fig. 1). In either of these, the result would be somewhat similar
to the TCA dline demonstrated in figure 2.
In sections VIII and IX an attempt was made to account for the presence of
the Serrana and Pata de gallinazo types in the central and southern provinces.
There the hypothesis was proposed that the Serrana type was the result of genic
introgressions from Bofuda into Pitaya following a series of primary hybridiza-
tions. - It would seem apparent, since Serrana exists under conditions quite com-
parable to those under which Pitaya flourishes, that the different abilities in
alkaloid synthesis are of no selective importance.
It was further suggested that the Serrana type then migrated southward
along the upper edge of the escarpment of the western Cordillera, at least into
what is now the Province of Caniar. There it could have migrated from the
western to the eastern escarpment. Once on the eastern escarpment it would
have met the alkaloidally barren Crespilla type, another high-elevation species.
At the present time there is every evidence that the Serralna and Crespilla types
can exchange genes by hybridization. Thus it might appear that the present
Pata de gallinazo population is the result of the infiltration of Crespilla genes into
the southward migrating Serrana population.
If the events of the foregoing accoulnt of the Pitaya-Serrana-Pata de gal-
linazo complex are at all correct (and the hypothesis is to be further tested in
this section), it is to be concluded that the step-dline in total crystallizable alka-
loids present in this series (fig. 2) is the result of a successive contamination of
the basic Pitaya genotype by genic elements derived from two other specifically
distinct types, namely Bofuda and Crespilla.
However, before this subject can be closed, another possibility should be ex-
plored. In considering the natural distribution of the genus Cinchona it is im-
mediately obvious that its present center of complexity is in northern Peru and
adjacent southern Ecuador. I have no personal knowledge of the conditions in
Peru, but in southern Ecuador, in the Province of Loja (as on the slopes of the

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
Cordillera de Zamora) we found five-and sometimes additional-morphQlog-

ically quite distinct types in a single transect. The same can be said for a tran-
sect in that part of the escarpment lying eastward of the Province of Azuay in
the Oriente (the Province of Santiago-Zamora). There, in descending order
from about 10,500 feet to somewhat less than 4,000 feet, in a single transect along
the Rio Negro3' were found zones of Crespilla de Cuenca (C. rugosa), Pata de
gallinazo (of the C. pitayensis-complex of this paper), Costrona fina (a member
of the C. officinalis-complex), Zapallo (a cognate of Bofuda in the C. pubescens-
complex), Hoja de capuli (distinct from the others but imperfectly known bo-
tanically), and an additional unnamed type which occurred in altitudinally
somewhat disjunct and morphologically different forms toward the base of the
transect (both of these, morphologically and on the basis of their alkaloid pat-
terns, appear to be northward-extending members of the primarily Peruvian C.
calisaya-complex, known to be quite variable there). This certainly is a much
more complicated situation than that described by Steere on the western esearp-
melit in the northernmost part of the Province of Carchi (Section VI), where
only three types of Cinchona were found.
If, for the present, we were to assume that the center of complexity in a
genus in a general way can be taken as its center of origin-and I am fully aware
of the various arguments against- and the many known exceptions to such a view-
point-we might think of northern Peru and southern Ecuador as a region in
which the basic types of the group perhaps developed and from which they radi-
ated. Such a viewpoint would possibly necessitate a revision of the history of the
Pitaya-Serrana-Pata de gallinazo series as previously outlined.
In our conversations while yet in Ecuador, and more recently through cor-
respondence, Steere (who had considerable experience with the Cinchona-popu-
lation in Colombia before transferring his activities to Ecuador) pointed out
that, in Colombia, the Pitaya type did not extend northward beyond the zone of
relatively recent volcanic activity. It was his suggestion that fairly new vol-
canic ash in the soil-by supplying certain mineral nutrients-might be an im-
portant factor in the distribution of this type. While it may be a potent factor,
the fact remains that the Pitaya type, in its strict interpretation, apparently
stops rather abruptly in the Province of Pichincha in its southerly extension into
Ecuador. It therefore is interesting to note that, in Ecuador, with the exception
of two volcanic cones, Reventador (which is in the north at some distance east
of.the Cordillera in the Oriente and quite active today) and Pichincha, the others
which have been active within historical times-Antisana, Cotopaxi, Tungurahua,
and Sangay-are in the same latitude as the Serrana type.32
31 According to several personal traverses of the region and considerable work on the
backbone ridge, the Rio Negro rises in a series of boggy areas at about 13,500 ft. elevation
in the Paramo del Castillo at the edge of the eastern escarpment in the Oriente, roughly op-
posite and across the range from the villages of El Pan and Sevilla de Oro in easternmost
Azuay; the union of the Rio Negro with the Rio El Cruzado forms the Rio Chupiangas, which
empties into the Rio Paute southeast of Mendez near the confluence of that stream with the
Rio Upano. The position of all these streams is only approximate on current published maps.
32 Of the numerous volcanic cones in Ecuador, only two-Reventador and El Sangay-are
today explosively active. However, Wolf (13) in his monumental work on Ecuador, published
in 1892, records eruptions within historical times of the following (I do not have available
records of eruptions since the time of Wolf): Pichincha-1566 to 1582, 1660 (very heavy)*
Antisana-1590, 1728?, 1801; Cotopaxi-1534, 1743 to 1768 (with heavy devastation over this
period in the Prov. of "Le6n," now Cotopaxi), 1803, 1845, 1851, 1853, 1855, 1856, 1877,
1880; Tungurahua-1641?, 1771, 1881, 1886 (this last period very-violent and the one which
dammed the valley of the Rio Pastaza); Sangay-Wolf (p. 366) notes a count made by S.
Wisse in December, 1849, of 267 strong explosions per hour. In May, 1944, Dr. Steere and

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
Before we discuss this point further, let us look at the southern Provinces.
Oppenheim's recent geological map of South America (9), although necessarily
generalized, divides the Andean region of southern Ecuador into two parts, the
eastern being classed as Precambrian sediments, metamorphics, and igneous rocks,
and the western as Paleozoic and Mesozoic igneous rocks. Yet in spite of the
age of the basic strata, there were pockets of material-and little more than mere
remnants at times-along the fresh workings where the Pan-American Highway
was being cut through, which I would interpret as being quite definitely volcanic
ash of a later age. Fresh exposures along the road-cuts on the shoulders of Cerro
Villanaeo just west of the citv of Loja also would lead me to think that this is
a not too ancient volcanic cone. In brief, it is my opinion, although the major
volcanies of the southern provinces may be quite old-probably no younger than
the Mesozoic-that there was considerable eruptive activity at least locally in this
region which lingered well into the Cenozoic, and therefore into the time during
which the basic members of the genera of the present flora were evolving and
deploying.
It may well be, then, that the ancestral Pitaya did evolve much to the south
of where it occurs today, and there became established as a type which succeeds
best on soils which have been enriched or in some way modified with volcanic ash.
If so, it might well have arisen near what is now the Ecuador-Peru border and
moved north in normal ecological succession following the volcanic cycle into
what now is northern Ecuador and southern Colombia. Should the foregoing
be the case, we then are left with alternative explanations for the present distribu-
tions of the cognate and apparently derived Serrana and Pata de gallinazo types.
1. Having originated in the south and later, by migration, reached the north-
ern limit of the soil type to which it is ecologically adjusted (and where other
limiting factors probably also might become effective), the Pitaya type would
then become static in its migration. However, with a possible rejuvenation of
volcanic activities to the south in what now are the central provinces (and re-
gional activities of vuleanism often do reappear in long-term cycles), there would
be opportunity for Pitaya, or a population with much the ecological requirements
of Pitaya, to reoccupy this territory. It is therefore interesting to note that the
derived Serrana type-which, in its ecological requirements, appears to be almost
a duplicate of Pitaya-does occupy the territory which is so placed, because of
the prevailing winds, that it is within the "ash-shadow " of the more recently
active volcanoes.33 Furthermore, as a dominant member of the high-elevation
Cinchona-population, Serrana does not stray for any considerable distance south
of the region in which the ash-dust from Sangay would be likely to fall. South
of that zone, its place is taken by the Pata de gallinazo type and, as has been out-
lined, basically this type appears to be Serrana to which have been transferred
certain genie elements of the Crespilla type. It is further interesting to note
that, so far as my data indicate, the Crespilla type does not extend northward
into the zone of the present or more recent volcanic activities. In fact, both the
I were north of Sangay and noted approximately two strong explosions per hour, accompanied
by heavy ash fall. In December of the same year, Henning Jorgengen and I were in the valley
of the Rio Upano about 80 kilometers S-SE of Sangay and on one clear evening we could
distinctly see the fire column given out at each major explosion; at that time the explosions
lbad increased to as many as twelve per hour.
33 The ash-dust of the more northerly Reventador is commonly detected throughout the
northern provinces and is much in evideliee in Quito, 95 kilometers distant; and I have been
informed that the ash of the centrally located Sangay (at 20 S. Lat.) is often noted in Guaya-
quil, 125 kilometers away on the opposite side of the Andean ranges.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
424 Brittonia L[VOL. 6
Crespilla of the Azuay region and that of the Loja region appear to be limited
to the ancient, primarily sedimentary and metamorphic rocks on the crest of the
eastern Cordillera in the southern provinces where, if it once was present, any
surficial ash-mantle has long since been removed by erosion. One therefore can
but wonder whether, in the exchange of genes where the Serrana and Crespilla
types met, the resultant Pata de gallinazo type-although morphologically still
to all general appearances a member of the Pitaya-Serrana phylad-might not
have acquired the ability to break its old hereditary ecological pattern and so
venture with some success onto soils which, today and in the recent past, have
not had a constant renewal of certain necessary-and possibly rare-minerals
through the medium of ash-dust from at least intermittently active volcanoes.
2. The other alternative-if we assume that the original Pitaya type arose
in the south and migrated northward-is that, during its northward migration,
it left behind residual stands which, siice, have become modified by certain gene
exchanges with other species which perhaps, as the Crespilla type, may have mi-
grated into the region more recently.
The latter of these two explanations certainly is the simplest and perhaps most
obvious one and I personally would be inclined to accept it were there not another
item to be considered-that of Pleistocene high-altitude glaciations in the Ecua-
dorean Andes.
Today, from the border between the Provinces of Caniar and Chimborazo,
northward through Ecuador, there are numerous quiescent or active volcanic
cones capped with snow. The permanent snow-line varies from 15,000 to 16,000
feet depending on the size of the accumulating area, direction of slope, the
seasonal, regional, and climatically cyclic precipitation. Snow is, common on
the paramo zone below the snow-fields; and crops are often frosted down to
11,000 ft., and occasionally even lower than 10,000 ft. in certain areas (e.g. in the
inter-Andean valley near Quito where according to Steere (in litt.) " granizo "
hail-like snow-has fallen to a reported depth of two feet).
Throughout much of Andean Ecuador there are numerous small lakes. A
few of these are the result of the damming of streams by eruptives and seismic
disturbances; the majority, however, are glacial in origin and owe their presence
to terminal moraines of former valley glaciers. It is not surprising to find many
of these small lakes clustered around the higher mountain masses which -still
support permanent snow fields and which must have been nuclei for former
(Pleistocene) glacial activities. But they also are a common feature in the
valleys surrounding the great paramo area of the western part of the province of
Azuay, in the southern part of the country. Here these lakes are usually between
10,000 and 11,500 feet elevation.34 Where I have examined them they obviously
are the result of damming by terminal moraines of former valley glaciers, just as
are the similar lakes northward where the mountains are higher.
An interesting feature of this paramo area in western Azuay, an area approxi-
mately 40 miles long and averaging about 10 miles wide, is that its surface is
rolling, is but little dissected, and lies mainly between 11,500 and 13,000 feet
elevation, its highest point known to me being only about 13,650 feet. Yet from
this rather low area (that is, low for Andean Ecuador), there once radiated a
considerable system of valley glaciers. The only conclusion which can be reached
34 Photographs of one of these U-shaped, glacially scoured valleys (the Surueucho), with
its terminal valley-moraine of the last cycle of glaciation and typical high-Andean lake, to-
gether with a non-technical account of the surrounding flora, appeared in a recent publication
(Camp, 3).

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
is that, in the most recent cycle of Andean glaciation this area of comparatively
low.relief was covered with an extensive ice field. From the configuration of the
terrain and the position of the terminal moraines of the valley glaciers which were
spawned in this area, while studying it in the field I came to the further con-
clusion that the zone of permanent ice-the snow-line-must then have been as
low as 12,000 feet, or possibly even 11,000 feet in local areas. This would have
forced the zone of occasional frosts to 8,000 or perhaps as low as 7,000 feet.
Physiographic features comparable to the foregoing in rather continuoun series
northward indicate that much the same conditions prevailed throughout Andean
Ecuador and probably southern Colombia at comparable elevations during the
maximum of the last major glacial cycle.35
While I have no actual data on the item of frost resistance in Cinchona, it
would seem apparent that the upper altitudinal limit of the Pitaya-Serrana-Pata
de gallinazo complex might well be determined by the lower limit of killing frost.
In fact, the only Cinchonas with which I am familiar that enter the upper alti-
tudinal zone where, on the sole basis of elevation, frost theoretically might be
expected, are the two Crespillas (Cr. de Cuenca and Cr. de Loja). And the
highest stands of these types found were in passes in the Cordillera Oriental at
the heads of valleys facing toward the east and the Amazonian basin, and there-
fore so placed that the warm and essentially continuous easterly winds of the
equatorial lowlands, sweeping up these valleys, would locally preclude the possi-
bility of frost.
Briefly, then, during the last glacial cycle the entire montane flora of Ecuador
requiring frost-free conditions-and this includes all species of Cinchona-must
have been at considerably lower levels than at present.
It is most unlikely that the present tendency for species of Cinchona to be
altitudinally stratified on the Andean slopes is a newly acquired trait-this habit
must go back into the primary evolution of the group during the early or middle
Tertiary, with its further elaboration during the late Tertiary and the (pre-
sumed) repeated cycles of the Pleistocene. At least it, would be safe to assume
that, at the approach of the last glacial cycle, various species of Cinchona already
were present in the Ecuadorean Andes and that those which were so close phyleti-
cally as not to have yet achieved potential genetic disjunction maintained their
integrity as specific units because of latitudinal or altitudinal disjunction. For
those separated by altitudinal disjunctions the approach of the last glacial cycle
would have been a critical time, for to have continued existence would mean that
it would have been necessary to migrate to lower levels. Where a series of these
were stratified on the slopes, a "jamming together" of the various Cinchona-
zones would have been inevitable.
In the previous parts of this paper we have seen what happens when various
species of Cinchona make distributional contact. The constant reference to such
things as the "C. pubescens-complex," the "C. officinalis-complex," etc., may
have additional meaning in the light of this part of our discussion, for it is my
35 It is here suggested that the development of natural paramo areas in the northern Andes
may be as much related to edaphic conditions as to elevation and related factors, for the
paramos (at least those of Ecuador) seem to be fairly well correlated in extent with those
areas which apparently were covered and therefore probably scoured and planed by the ice of
the last glacial cycle. Characteristically, the p'aramos are rather poorly drained areas of low
relief. It is true that the paramos have been enlarged by clearing and pasturing-in pre-
Conquest times probably in certain areas by vast herds of domesticated llama and since then
obviously by introduced stock-but it also is equally true that the paramos are a natural
phenomenon and have been an Andean feature for a very long time, else there would not be
so many characteristic genera and species endemic to them.

143.106.201.2 on Thu, 16 Jul 2020 15:25:18 UTC
opinion that much of the complexity of the Cinchona-flora can be laid to the events
of the Pleistocene as they affected its history and development. The conclusion
to which I came as I studied these groups in the field was that sometimes they
were widespread species whose obvious regional variants had been the result
primarily of genie introgressions from adjacent species. Or, again, on occasion
it appeared that these introgressions, while not always disorganizing too greatly
the morphological characters which the taxonomist uses in his determinations, had
yielded segregate biotypes capable of entering new territories. Such events
certainly have added greatly to the complexity of the Cinchona-flora in Ecuador.
It is possible to suppose that Cinchona pitayensis, in a taxonomically restricted
sense (as the Pitaya type), was distributed from southern Colombia southward
through Ecuador before the last glaeial cycle; that its present Serrana and Pata
de gallinazo cognates are the result of genic introgressions, respectively, from C.
pubescens and C. rugosa (or, mnore probably, C. microphyllU) over a wide front
where these today are regionally associated, but during the last glacial cycle when
it is certain that they would have been forced down slope and into closer contact.
It is an intriguing theory, but I have difficulty in accepting it.
The transect along the Rio Negro (briefly outlined in a previous paragraph of
this section) is perhaps enlightening. There C. pub escens (masquerading under
the local name of Zapallo) occurs at the elevations where we would expect it, yet
there is no evidence of gene exchange between it and either a hypothetically
present Pitaya or Pata de gallinazo (the local variant of the C. pitayensis-complex
as interpreted in this paper). The wide distribution of C. pubescens-it is on
the flanks of both the western and eastern escarpments at least fromr Peru into
Colombia-would seem to indicate its presence in those areas before the last
of the glacial cycles. It therefore is concluded that any member of the C. pitay-
ensis complex present on the eastern Cordillera of the southern provinces has
migrated into the area since the last of the Andean glaciations. In fact, this
transect of the Cinchona-flora along the Rio Negro has much the appearance of
a group of altitudinally intercalated species, some of which migrated into the
area from the north and some from the south; and still others appear to be more
or less stable local derivatives of former hybrid complexes.
Furthermore, as previously noted, Crespilla de Cuenca (C. rugosa) has much
the appearance of a segregate form developed from a combination of genes de-
rived out of Serrana and Crespilla de Loja (C. microphylla). Therefore, it may
well be that both Pata de gallinazo and Crespilla de Cuenca have been derived, as
segregate biotypes, from hybrids between Serrana and Crespilla de Loja, with one,
of these (Crespilla de Cuenca) taking over the ecological niche of one parental
type (Crespilla de Loja) and the other (Pato de gallinazo) that of the other
parent (Serrana). The morphological -"break over' (apparently with a zone of
intergrades) between the two Crespilla types appears to be east of the village
of Onia near the Azuay-Loja border. Since time is always a factor to be reckoned
with, it is possible that the' segregate Pata de gallinazo biotype, in the southern
portion of its distribution along the eastern border of the Province of Loja on the
Cordillera de Zamora, is just now in the process of migrating southward as a form
intercalated between the uppermost species of the genus (Crespilla de Loja) and
the next one below (Hoja de luema) in the normal, eastern Province of Loja
transect.6
36 In the Loja region, transects by my own group from the crest of the eastern Cordillera
westward into the interior valley of the Rio Catamayo and adjacent areas characteristically
yielded the following in descending order: Crespilla de Loja (C. microphylla), Pata de gallinazo

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1948] Camp: Ci,nchona in Ecuador 427
In summary, then, it is my opinion that, before th

only the Pitaya portion of the Pitaya-Serrana-Pata de gallinazo high-altitude
complex was in existence, and that its distribution was in, or concentrated in,
what now is northern Ecuador and southern Colombia. During this last glacial
cycle it necessarily was foreed to lower elevations where, in one or perhaps several
local areas, it made distributional contact with the wide-spread C. pubescens
which occurred below it on the slopes of the escarpment. Out of the derived
hybrid complex (much as one now finds in the Rio Saloya-Corazon Pass area)
there appears to have arisen a. high-altitude biotype, the Serrana, capable-in
early post-Pleistocene time-of migrating southward along the upper -part of the
esearpmient, then unoccupied by any competing species of Cinchona, until it
apparently was blocked from further migration by the series of broad and deep
canyons which cut into the western esearpmnent at about 30 S. Latitude. There,
because of transverse ridges, there would have been opportunity to migrate east-
ward. Arriving on the eastern escarpment, it then would have had opportunity
to migrate southward. However, at the elevations necessary for the free migra-
tion of the Serrana type it would have met another-the more southerly distrib-
uted Crespilla, then probably migrating northward. There also may have been
ecological (edaphic) barriers to a further southward migration of the genetically
stabilized Serrana.7 At least, in those areas in the region of the Rio Paute,
there is evidence that Serrana and a form of Crespilla have exchanged genic ele-
ments. Out of this exchange there then apparently developed another biotype,
the Pata de gallinazo, which, with its newly acquired genetic balance with
the changed ecological conditions of the region, has since been able to migrate
further southward along the escarpment, at least into the Cordillera de Zamora
east of Loja, and perhaps via the transverse Nudo de Sabanilla for a short
distance into northern Peru.
Yet, regardless of how the problem of the migration of these forms ultimately
is to be decided, the fact remains that, today, these three morphologically com-
parable members of the Pitaya-Serrana-Pata de gallinazo complex do exhibit a
modified or "stepy' dlinie in their alkaloids. Furthermore, it would seem obvious
that the items of variability exhibited by them have not arisen by mutation or
some change from within the complex itself, but are the result of modifications
induced by genetic elements which have been received by means of, genie intro-
gressions from quite distinct species outside the complex. Therefore, since the
variabilities seen in this complex are exogenous and not endogenous, the exhibited
(of the C. pitayensis-complex and locally called Hoja de luema redonda), Hoja de luema (C.
Iucumaefolia), Urituzinga (C. officinatis, cognate with Costrona fina of the Cuenca area), and
Hoja de Zambo (C. putbescens, cognate with Bofuda, Rosada, and Zapallo). Westward, at still
lower elevations, in the Zaruma Basin of adjacent El Oro, Roja (C. succirubra) also was found
in normal sequence below the putbescens zone (see fig. 3).
37 It would be legitimate at this point to ask why Serrana did not turn here and migrate
northward along the eastern escarpment. I do not know, but those who have traversed the
area tell me that from the valley of the RRio Paute, northward for well over a hundred miles,
there are large areas on the escarpment where, although the general flora is rich, members of
the Rubiaceae (the family- to which Cinchona belongs) are conspicuously absent, or at best
represented only by the more ubiq4iitous Andean "weedy" species of the family. In my own
explorations in the Rio Zamora-Rio Sabanilla region in the Oriente east of Loja a similar
zone, almost devoid of the usually rich rubiaceous flora, was encountered. Such "'barren
zones,'' wherein essentially a whole family of plants suddenly drops out of the general floristic
pattern, are positive indication that some ecological factor (probably edaphic, assoeiated with
the underlying rock type) is operative; such things will easily block migratioii and it very
probably was just such a barrier which did not permit the Serrana type of Cinchona to migrate
northward once it made the crossing from the western to the eastern Cordillera.

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dline, if we wish to classify it, might be called an exogenous cline. In addition to

being an exogenous dline, it also is a modified or "step" dline.38
SUMMARY
On the basis of their community of morphological characters, it is concluded

that three of the major types which occur at the upper (9,000-10,000 ft.) eleva-
tions of the altitudinal distribution of Cinchona in Andean Ecuador-namely
Pitaya, Serrana, and Pata de gallinazo-are parts of a single complex. The
Pitaya type, which occurs in the northern provinoes (and also in adjacent Colom-
bia), is thought to be the basic member of the complex as it exists today.
A transect of the frontal escarpment of the western Cordillera in northern-
most Ecuador in the Province of Carchi revealed three distinct zones in which
Cinchona is present. The low-elevation zone is characterized by a type called
. Roja" (C. "succirubra") ; the middle-elevation zone by "Bofuda" (C. ptbes-
cens, sens. str.) ; and the high-elevation zone by "Pitaya" (C. pitayensis).
Somewhat to the south, a transect of the same escarpment in the Rio Saloya-
Corazon Pass area in the Province of Pichincha revealed the same general zona-
tion, but here there are indications that the three populations are not (odr in the
past were not) completely disjunct. Here the middle-elevation Bofuda type
contained a high percentage of "aberrant" individuals. On the basis of morpho-
logical appearances and evidences from their different abilities to produce various
of the alkaloids present in the genus, it is concluded that these aberrant indi-
viduals of the Bofuda population are the result of the infiltration of genie ma-
terials from one or both of the altitudinally adjacent types. While there is a
possibility that Roja has contributed to the aberrancies of Bofuda, an analysis
of the situation would seem to indicate that Pitaya may have been primarily con-
cerned.
Since there is evidence that the specifieally quite distinct Bofuda and Pitaya
are able to hybridize and exchange genes, one would expect that out of such a
nuclear hybrid population (and presumably in the general region of contact)
a segregate type could arise which, because of its life-form and ecological pattern,
would potentially be able to build up a successful population in the high-elevation
Cinchona-zone. It is also obvious that non-selective characters, if segregative,
could also appear in this new population. Thus, where the Bofuda and Pitaya
types make contact, there is present a local population wherein the expected
variant individuals are found. Extending southward from this region is found
the Serrana population which, in its morphological manifestations, gives clear
evidence of its close affinity with and primary derivation from the Pitaya type,
but which to a slight extent exhibits characters of a type patently derived from
Bofuda; the occurrence of Serrana at the same elevation and on much the same
soil type indicates that its ecological requirements also are mainly those of its
Pitaya ancestor. So far as the alkaloid picture of Serrana is concerned, it par-
takes of both ancestral forms for in synthesizing primarily einchonine it is much
like Bofuda, but in its ability to produce greater amounts than this type it is more
like its alkaloidally high-yielding Pitaya counterpart.
In the southern provinces of Ecuador, Serrana makes distributional contact
with material belonging to quite a different group-the Crespilla-and there is
incontrovertible evidence in the field that here these two have hybridized to a
38It will be obvious that both normal and ''step' dlines may be either endogenous or
exogenous; in fact, there is no reason why, in some of the more complicated populations, both
endogenous and exogenous dlines might not be present at the same time in a single species.

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certain extent and exchanged genie materials. From this point southward, the
third member of the complex under scrutiny-Pata de gallinazo-is to be found.
Again, while morphologically well within the limits of variability of this complex,
and showing its close affinities with Serrana and thence to Pitaya, this Pata de
gallinazo population exhibits certain items which would indicate the influence
of the former gene-exchange of some member of the complex with Crespilla.
Throughout the known range of Crespilla-and two Crespilla types are involved
-this high-altitude associate of Pata de gallinazo is barren of alkaloids; and
perhaps the outstanding point of difference between Serrana and Pata de gal-
linazo is the lack, in the latter, of ability to synthesize alkaloids. It is concluded
that this is one of the results of genic introgression of Crespilla elements. An-
other point is that, while at elevations comparable with the other members of its
complex, Pata de gallinazo flourishes on soil types different from those in which
Pitaya and Serrana are found. Again, it is concluded that this apparently new
potentiality in the complex may have been derived through the medium of gene-
exchange with Crespilla, edaphically adjusted to the region.
The dline in total crystallizable alkaloids exhibited by this complex-being
highest in Pitaya and least in Pata de gallinazo-is a modified or "step" dline.
Since the differences in ability to synthesize these alkaloids appear to be the result
of genic introgressions from species outside the complex, this would be classed
as an exogenous dline.
It is indeed unfortunate that data on the cytogeneties of the members of the
genus primarily dealt with in this study are not available for, should natural
polyploidy be found among them, it might necessitate a recasting of the present
hypothesis. However, if we proceed on the basis of the available data, taken
fromn the morphological manifestations and the evidences obtained from the chem-
ical analyses of single-tree samples in these various populations, three items stand
out in bold relief:
1. While large populations composed apparently of a single biotype do exist
in Cinchona, there is abundant experimental and field evidence that specifically
different members of the genus can exchange genes. This results in the presence,
today, of locally complicated populations seemingly of hybrid origin (e.g. the
series of individuals from 8,000 ft. to 10,000 ft. in the Pichincha transect in-
volving Bofuda, Pitaya, and Serrana).
2. If hybridization is possible at the present time, there is no reason to sup-
pose that it could not have happened in the past, yielding population segments
which, today, are atypical as a result of active gene-flow (e.g. the "aberrant
Bofuda" individuals in the Pichineha transect).
3. The last item (and perhaps the most important from the standpoint of an
interpretation of the group) is the apparent segregation of forms ecologically
attuned to particular habitats-forms which, presumably, have had opportunity
to build up reasonably stable populations over fairly extensive areas (e.g. the
Serrana and Pata de gallinazo types).
It is therefore apparent that future students of the systematics of the genus
Cinchona face no simple problem. They may, if they so desire, "split" the
species so finely that an understanding of the group becomes lost in a multiplicity
of binominals, the delimitation of each covering only a local population, as has
been the tendency of some writers; or they may "lump" the species to the point
where all semblance of a phyletic interpretation is lost, as also has been done.
Nor will the mere a priori adoption of a "middle ground" viewpoint suffice.
The problem which students of the systeinaties of the genus Cinchona face is

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the rationalization of our present system of nomenclature with the actualities of

a dynamic evolution which exhibits itself in a reticulate phyletic pattern.
If the present paper-dealing, as it does, with only a few forms-appears to
be over-long, it should be considered as a "case history" examined in some detail
so as to clear the way for a study of the structure of natural populations in
Cinchona in those regions in which the genus is even more complex, but where
the same naitural phenomena appear to have been operative.
As may be evident from the foregoing discussion, it is my opinion that the
Pitaya, Serrana, and Pata de gallinazo populations should be treated as members
of the same specific complex-probably as subspecies-rather than parcelled out
among other species with which they have only minor morphological affinities.
By agreement with others, I have not pursued this study to its logical nomencla-
tural conelusion, having deferred the decisions in these matters to those now
primarily concerned with the systematics of the Rubiaceae.
Literature Cited
1. Camp, W. H. A preliminary consideration of the biosystematy of Oxycocctus. Bull. Torrey
Club 71: 426-437, 1944.
2. . The North American blueberries with notes on other groups of Vaceiniaceae.
Brittonia 5: 203-275. 1945.
3. . The Surueucho. Jour. N. Y. Bot. Gard. 47: 25-31. 1946.
4. and Gilly, C. L. The structure and origin of species. Brittonia 4: 323-385.
1943.
5. Darrow, G. M. & Camp, W. H. Vaccinium hybrids and the development of new horticultural
material. Bull. Torrey Club 72: 1-21. 1945.
6. Howard,T J. E. Illustrations of the Nueva Quinologia of Pavon. London. 1862.
7. Huxley, Julian. Evolution: the modern synthesis. New York and London. 1942.
8. Martin, W. E. & Gandara, J. A. Alkaloid content of Ecuadorian and other American
Cinchona barks. Bot. Gaz. 107: 184-199. 1945.
9. Oppenheim, Victor. First generalized geologic map of South America. Tech. Paper No. 2.
Pan Am. Inst. Mining. Eng. & Geol. New York. 1945.
10. Rainey, Froleich. Quinine hunters in Ecuador. Nat. Geogr. Mag. 89: 341-363. 1946.
11. Steere, Wm. C. The Cinchona bark industry in South America. Sci. Monthly 61: 114-126.
1945.
12. . The discovery and distribution of Cinchona pitayensis in Ecuador. Bull.
Torrey Club. 72: 464-471. 1945.
13. Wolf, Teodoro. Geografia y geologia del Ecuador. Leipzig. 1892.
NOTE: My original copy of the most recent general work on the einchonas of Ecuador
[Acosta Solis, M. Cinchonas del Ecuador. 271 pp. Quito. 1946.] was not available, because
of mis-filing or loss, during the preparation of manuscript for the foregoing paper. A sub-
stitute copy was obtained only after it was set in type. This will explain the absence of this
work from the bibliography. It might be noted, however, that such differences as exist would
be mainly those of nomenclatural interpretation and these, as mentioned in the text, are outside
the scope of the present discussion.-W. H. C.

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Cinchona at High Altitudes in Ecuador

Stable URL: http://www.jstor.com/stable/2804925

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CINCHONA AT HIGH ALTITUDES IN ECUADOR

II. SOURCE OF THE DATA

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States Foreign Economic Administration' to explore for and locate stands of

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5 In the foregoing there is no intimation that my associates working in the northern

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Ecuador, altimeters calibrated in meters were not generally available to the

III. THE ALKALOIDS OF CINCHONA BARK

IV. THE PITAYA-SERRANA-PATA DE GALLINAZO COMPLEX

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and flower characteristics is much closer to C. pitayensis, which occurs in northern

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will be m-ade in succeeding passages to explore these-and additional-possibili-

V. THE ALKALOIDS OF THE PITAYA-SERRANA-PATA DE GALLINAZO COMPLEX

As is indicated by figure 1, when one goes in a general north-south direction

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val &4y)4/ 5 6, o t 7)r Cd,- chz

The Andearn provinces (2) Prov. Imnbabur- -

(3) Prov Pich in ch- a6

Pd rov 6,ozolwe (5),.BaixvSd (6),i_%il

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Co/ombid Ecada'or Peru

FiG. 2. The dline in total crystallizable alkaloids in the Pitaya-Serrana-Pata de gallinazo

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VI. A TRANSECT OF THE WESTERN ESCARPMENT

-6 P4, P4rp c/awed enecUv l//ied4i'e: o0ooa(.

V-iSerr4ncd A-ineer,~eidg4e O-Piz4Ya,

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| (RrbodEeso{Q3w/-)3R 4,ooDy. -3 VW i-4

Lk Id/oy4- CordPdsscdorec. The 'bubescens co

uppermost zone consists of a mixture of materials very closely approaching Pitaya

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Cinchona pitayensis . 10,000-9,000 ft.

WCS No. 7 WCS No. 8 Janouch-SC14

1s Analyses of material coll

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widespread Cinchona populations in Ecuador; that is, except in certain local

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T/ze '[3oJcf I Reo/dtio- - Sd/oyd -CordZcW Rs5 ?dre.

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-considered it impossible to identify. (Because of this the specimen was not

VII. AN EXPLANATION OF THE ABERRANCIES MET WITH IN THE

Since information is lacking on the cytogenetical situation in these various

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JrX,00?ft. 6,000-8,o0oJ. 8,600/1. +9,000-70,000fi.

FIG. 5. A comparison of the alkaloid types in various Cinchona populations in Ecuador.

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entered the Bofuda po