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Camp 1949 PDF
Camp 1949 PDF
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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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.
The general background of the activities of the group employed by the United
394
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.
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).
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
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.
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,
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."
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.
e ta 25gur~~4vdhal)_ \|
-o
(%)
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.
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.
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.
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.
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.
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
Soinge-tree an4j/y
an d/ittIdin4/ trdseCt.- , )f dherrd BoJac RFR 'i
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.
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):
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.
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
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.
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),
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
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.
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
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.
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.
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.
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
-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.
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.
X. ON THE HISTORY OF THE GROUP AND THE NATURE OF THE CLINE IN ALKALOIDS
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.
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).
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.
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
(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.
SUMMARY
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.
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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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.