BOTANY OF THE MAYA:
BOTANY OF THE MAYA AREA: MISCELLANEOUS PAPERS
A METHOD OF PROCEDURE FOR FIELD WORK IN
TROPICAL AMERICAN PHYTOGEOGRAPHY BASED
UPON A BOTANICAL RECONNAISSANCE IN PARTS
OF BRITISH HONDURAS AND THE PETEN
FOREST OF GUATEMALA
BY HARLEY HARRIS BARTLETT
With fourteen plates
[Issued July 10, 19351
A METHOD OF PROCEDURE FOR FIELD WORK IN
TROPICAL AMERICAN PHYTOGEOGRAPHY BASED
UPON A BOTANICAL RECONNAISSANCE IN PARTS
OF BRITISH HONDURAS AND THE PETEN
FOREST OF GUATEMALA'
The modern point of view in botanical survey has hardly been applied in
the tropics. There is very little literature dealing with the definition and
analysis of tropical plant associations except such relatively simple ones as
the strand of the mangrove belt, and there are no ecological or phyto-
geographical publications for any restricted tropical area which give us such
definite information as we ha,4 for many temperate regions. Scottsberg
"As far as I know, not one description in a modern sense of a rich tropical
association exists. Possibly such detailed descriptions are not necessary; we
may be entitled to leave out many of the components and still be able to
separate the associations from each other. This is perhaps what we ought to
find out first of all."
As a matter of fact some observers have practically denied that rich
tropical floras can be divided into definable plant associations. Gleason
has said that the definition of an association in terms of character species is
impracticable or even impossible. He states that in a tropical forest a single
hectare may contain a hundred species of trees, not one of which can be found
in an adjoining hectare. Nichols 4 has questioned this statement and quotes
Whitford to the effect that he did not know of a single instance where quanti-
tative measurements have been made of the tree population in the tropics and
where this statement of Gleason's would hold true unless for plots which
presented radically different environmental conditions. Nichols also quoted
Shantz, who has had broad experience in Africa, to similar effect.
'Paper No. 406 from the Department of Botany and the Herbarium of the University
of Michigan. Based upon field work of the Maya Expedition of 1931, under auspices of
the Department of Historical Research. Carnegie Institution of Washington, the
Herbarium and the Museum of Zoology, University of Michigan.
'Carl Scottsberg, Plant communities of the Juan Fernandez Islands, Proc. Internat.
Cong. Plant Sci., Ithaca, N. Y., vol. 1, 565-574, 1929. (See p. 573.)
'H. A. Gleason, The individualistic concept of the plant alssociitions, Bull. Torr. Bot.
Club, vol. 53, 7-26, 1926.
*G. E. Nichols, Plant associations and their classification, Proc. Internat. Cong. Plant
Sci., Ithaca, N. Y., vol. 1, 629-641, 1929.
BOTANY OF THE MAYA AREA
The most informative studies that have been made in rich tropical forests
seem to be those of Whitford,1 Brown and Matthews,2 and Brown 3 in the
Philippines. Since Whitford has had especially long experience in tropical
forest ecology and had made quantitive studies of the tree population in
many places in the tropics, the following opinion of his, quoted by Nichols,
is particularly significant:
"I recall one instance in the Philippines where strip surveys were made in
a forest covering thirty square miles and where practically every hectare
measured showed a repetition of the same species. They were mixed, to be
sure, with a large number of other species which occurred much more rarely;
but even such species, upon careful examination, were found to be repre-
sented by small specimens in the adjacent hectares. To the average botan-
ist entering a tropical forest all seems confusion, and yet, with a knowledge
of the species and quantitative studies of the composition, tropical forests
can be classified by associations and can be designated by the generic and
sometimes the specific names of the predominating trees."
The writer has made no quantitative studies, but as a collector his experi-
ence is wholly in accord with Whitford's. Nevertheless there is every reason
for believing that Gleason's statement might be found entirely true at
many places in the tropics where cleared land has been abandoned and
covered by a relatively uniform secondary forest of a few species which are
so rare as to be easily overlooked even if they occur in adjoining primary
forest. In Sumatra, for instance, old secondary forest ("beloekar toewa")
only very slowly attains the climax composition of virgin forest ("rimba
raja"), and certain it is that a small patch of the latter surrounded by
second growth might actually have a hundred species that had not entered
the secondary forest. The latter, even if merely a successional stage, is a
long-enduring and definite plant association and has to be treated as such.
Ecologists who have worked in the Malayan region have indeed done so.
Brown, for instance, divides the vegetation of Mount Maquiling in Luzon
into (1) parang vegetation (secondary forest and abandoned clearings);
(2) dipterocarp forest (subdivided into (a) virgin forest, and (b) culled
forest; (3) midmountain forest, with two subdivisions (a) the Quercus-
Neolitsea association, and (b) the Astronia rolfei association; (4) the mossy
forest. Here we have a very serviceable classification of the vegetation
which is none the less useful because of the obvious fact that some of the
associations are successional phases and that there must be much
II. N. Whitford, The regetation of the Lamao Forest Reserve, Philip. Jour. Sci., vol. 1,
373-431, 637-682, 1906.
Studies in the vegetation of the Philippines. The Composition and volume of the
dipterocarp forests of the Philippines, Philip. Jour. Sci., Bot.. vol. 4, 600-723, 1909.
The forests of the Philippines. Philip Bur. Forestry. Bull. 10.' 94; 10,' 113, 1911.
SW. H. Brown, and D. M. Matthews, Philippine dipterocarp forests, Philip. Jour. Sci.
9 Par. A: 413-561, 1914.
'W. H. Brown, Vegetation of Philippine mountains, Manila, 1919.
A POINT OF VIEW AND A METHOD OF PROCEDURE FOR
RAPID FIELD WORK IN TROPICAL PHYTOGEOGRAPHY
The experience of tropical botanists with the highly complex flora of the
Philippines indicates that definite plant associations do exist in the tropics
and may (with certain practical limitations) be defined. Success in the
rapid study of any new area by a non-resident botanist will depend upon
getting a proper systematic acquaintance with the flora while in the field,
and such an acquaintance, the writer believes, is generally most ad-
vantageously obtained from the natives of the country. Their knowledge
is often found to be surprisingly extensive when scientifically checked. Folk
science takes account not only of a great number of plant species, but also
of plant associations. Its utilization by a botanist requires, in the first place,
that he must for a time subordinate his naturally greater interest in species
found in flower or fruit to an interest in prevalent plants for which he can
learn the vernacular names, regardless of whether or not they can be found
in determinable condition. Eventually they will be found fertile, but until
they are it will be necessary for the botanist to have names for them and
to recognize them in a sterile state as the natives do. Otherwise he can not
take notes. Nothing is easier, as the writer can affirm from his own experi-
ence, than to glean from a tropical forest a few species, possibly including
rare and interesting ones without being able to make a single useful note
about the plant association as a whole. Of course every isolated fact that
can be acquired is valuable, and therefore I do not wish in the slightest de-
gree to detract from the value of collecting specimens of whatever is deter-
minable, even one or two species whenever it is not possible to work more
thoroughly. I do, however, wish to emphasize the importance, in a genuine
biological survey, of recognizing the plant associations.
A botanist working in a new tropical area is seldom so fortunate as to
recognize in the field more than a small proportion of the species. If the
flora is complex he is confronted with a multitude of species which are not
only new to him, but which flower and fruit only at some other season than
that of his visit, or perhaps so sporadically that he can hardly hope to find
them fertile. Furthermore, just such plants are likely to be character plants
of the associations. They are often gigantic trees of which flowers are not
easily detected from the ground; or lianas, whose leaves, even, are in-
accessible in the tops of the trees; or palms, extremely difficult to collect
adequately; or bamboos, very seldom found in flower. When confronted
with such a situation, the botanist will find that his difficulties vanish as
if by magic if lie undertake to learn the flora as the natives know it, using
their plant names, their criteria for identification (which frequently neglect
the fruiting parts entirely), and their terms for habitats and types of land.
The names most often used will in general be those of the species of
greatest utility, or of the greatest importance in land classification. The
latter are the ones most important for the definition of associations. As
BOTANY OF THE MAYA ARRE
many names as possible should be listed, and the ones coming up most
frequently in conversation should be associated as soon as possible with the
species to which they apply. There is always a residue of names for rare
species with which no plants are associated, and, conversely, many plants
are found which are not specifically talked about by natives, and therefore
have no specific vernacular names. The result of applying the native-name
method in learning the flora, is to concentrate attention on the dominant and
most useful species and upon the associations which they characterize, rather
than upon the few and possibly non-characteristic species which are merely
convenient to collect, or which are accidentally found in readily determin-
By learning native plant names the botanist is able at once to make
serviceable notes upon plant associations in which the characteristic species
may never be seen except in sterile condition, or not found fertile until long
after their occurrence should have been recorded many times. It is of course
possible, and very frequently necessary, to use arbitrary designations for
plants in tropical field work, or numbers, which are not easily remembered,
but a vernacular name if one can be learned is infinitely preferable because
it makes botanical inquiry possible. Many interesting plants and facts
about them come to light only if one makes a habit of talking about botani-
cal matters with natives. If one is interested in vernacular names he will
at first hear them faster than he can find the plants to which they belong.
Unchecked names in his list will indicate gaps in his information about
important plants which he will be constantly on the look-out to fill. Native
companions are general more interested in hunting specific plants, or
habitats, than they are in helping with routine collecting. Insistence upon
the necessity of finding dominant plants in flower or fruit will generally
result in some sort of determinable material being collected. Nothing would
be easier than for a casual collector in Central America to leave the charac-
teristic species of most of the associations uncollected and unnoted-to have
nothing to prove the presence of the mahogany, for instance, in the ma-
hogany forest, or the chicle tree in the great zapote forest. The writer, in
spite of as close adherence as practicable to his scheme of following up
unassociated names, had to leave the field at the close of the 1931 season
with several of the more important plants quite unaccounted for. The same
was even more true at the close of field work in a limited region of Sumatra
in 1927; there remained on the list of unidentified plants several of such
importance that place names were derived from their dominance at par-
The preceding paragraphs will indicate something of the point of view
with which the writer made his first contact with Central American vege-
tation. After experience in the East Indies, and Mexico, part of which
had been mere collecting, the writer felt when he started work in Central
America that he had developed a ready and serviceable method of procedure
for ascertaining folk knowledge of plant associations, which would suffice
for the rapid reconnaissance that is undertaken in a general biological
It speedily became obvious that in British Honduras and Guatemala the
people knew an unusually large number of plants, had names for them,
which might be "creole," Spanish or Maya, and, furthermore, that they had
a perfectly definite land and vegetation classification, with appropriate
nomenclature in English, Spanish and Maya. Such knowledge furnished a
perfect basis to build upon, and as a matter of fact the local classification
and nomenclature of vegetational types is so serviceable and natural that
it needs only to be taken over and scientifically formulated to make avail-
able to science information that is already folk knowledge. Thus we shall
have an incomparably more adequate knowledge than we shall ever possess
if we wait for the application of ideal quantitative methods. Many natives
have a knowledge of local natural history far superior to that which a
scientific visitor could possibly glean without their assistance in any reason-
able length of time. It must be remembered that in the region under con-
sideration, as in many other forested tropical regions, much time is spent in
felling forest to make clearings, and that the natives have a lively curiosity
about what they find. Furthermore many of them have spent a large part of
their lives in scouting for mahogany, tapping the chicle tree, and in the
exploitation of other forest products, so that they inevitably acquire a great
deal of astonishingly precise knowledge about plants.
It is only forest associations that are more difficult to analyze in the
tropics than the simpler associations with which we deal in temperate
regions. Grass-lands, etc., present no especial difficulties and are not
markedly rich in species. The difficulty concerns the old forest, with its
great diversity of species which are inaccessible from the ground, which
can not be collected without laborious climbing or chopping, and which
are too often found to be sterile when felled for identification.
The technique of assimilating folk knowledge of tropical plant associa-
tions into conventional plant geography is in theory simple enough, but in
practice it is found to present its share of difficulties. Instead of making a
quantitative vegetational analysis by felling strips and quadrats, which is
rarely or never practicable for a non-resident botanist with limited time and
labor resources, the procedure is to utilize the experience which the natives
have gained in seeking wild products and in locating new land for their
primitive agricultural utilization of forest clearings (milpas). Obviously
the best possible training in the method is to be present during the making
of clearings, and to study and secure a name for everything that falls,
whether sterile or fertile; to accompany natives in their quests for specific
forest products; to inquire minutely into their basis for selecting land for
different kinds of utilization. The information obtained must be put to the
BOTANY OF THE MAYA AREA
test in the forest, with natives as critics. Interest in native names is the
almost indispensable prerequisite in a non-resident botanist who is to be
a reasonably competent plytogeographer as well as a collector.
The method involves the assumption that natives who have occupied an
area for generations will have names for the dominant species in the flora,
and that persistence in collecting the names and finding out how they are
applied will result in accounting for most of the species which should be
taken into consideration if plant associations are to be recognized in the
phytogeographic reconnaissance of botanically new country.
The utilization of folk science as a method in phytogeographic work is
commended to botanists who have found it difficult (as the writer con-
fessedly has) to give proper attention to the vegetation because of too great
concern over collectible flowers and fruits. The rules should be (1) to
learn each species for which the natives know a name, (2) to be able to use
their criteria for recognizing each species, even if sterile, (3) to be ever-
lastingly persistent in inquiring about and finding each named species, and
(4) to be equally persistent in securing, eventually, good determinable
botanical material of each. The last rule is the hardest to observe, but
essential to a successful survey. The writer believes that sterile material is
better than none, and is exceedingly grateful to those systematists, among
whom Dr. Standley is an outstanding example, who have been willing to
attempt the identification of imperfect specimens.
MAJOR TYPES OF VEGETATION IN CENTRAL
BRITISH HONDURAS AND THE PETEN
The northern part of British Honduras and the Department of Peten in
Guatemala are phytogeographically as well as physiographically a part of
the Yucatan Peninsula. Although having a well-defined dry season, they
lie south of the region of dry thorn forest in country which, with less dis-
turbance and greater rainfall, supports tall rain forest. As one goes west-
ward, there is a well-marked change in the vegetation which corresponds
with the absence of permanent streams, and roughly coincides, at least
near El Cayo, with the Guatemalan boundary. This change is chiefly
noticed in the dropping out of the cohune palm, except in restricted areas,
when the higher underdrained limestone country is entered. The region to
the east of the international boundary, below a line of hills located to the
westward of the international boundary, is Lundell's "Eastern Coast." The
underdrained region without permanent streams is "Northern Peten" of his
Since in British Honduras the language is prevailing English (or what
passes for English, although a stranger may at first imagine that the Creole
dialect is Carib or some other equally unknown language) it will be useful
Cyrus Longworth Lundell, Preliminary sketch of the phytogeographv of the Yucatan
Peninsula, Carnegie Inst. Wash., Contrib. Amer. Archaeology, No. 12, Pub. No. 436,
to outline first the classification of land and vegetation that is current among
the "Creoles" in British Honduras and then the more elaborate but basically
similar one of the Spanish-speaking inhabitants.
In the central and northern parts of British Honduras the halophytic plant
associations (strand and mangrove swamps) are succeeded inland by three
major plant associations which conform to soil conditions. These are
(1) Cohune ridge (Subclimax cohune-mahogany forest).
(2) Broken ridge (Climax mahogany-sapodilla forest and sequelar).
(3) Pine ridge (Climax pine forest).
It must be explained at the outset that "ridge" has nothing to do in local
parlance with elevation. However low and flat the land may be, even
though water stands upon it, it is nevertheless "ridge."
The lowest land near the coast is underlain by sandy Quaternary alluvium;
inland there are somewhat more elevated areas of Tertiary limestone,
known as the Rio Dulce limestones and marls, which according to Owcr
are of Oligocene age, but have been found by C. W. Cooke (unpublished in-
formation) to be partly Eocene. The lower Tertiary limestone areas are in
part covered with a thin mantle of sand, the distribution of which has been
modified by erosion. Cohune ridge and broken ridge occupy residual
calcareous soils or alluvial soils of mixed origin. Pine ridge occupies sour
sedgy sandy land which (as in the high "Mountain Pine Ridge" of the
central third of the colony) overlies granitic rocks, or other formations
deficient in lime, or land where superficial sand deposits or Quaternary
alluvial deposits are thick enough to minimize the effect of lime underneath
(as in more northern and coastal parts of the colony). In a land where the
soil is thoroughly leached during much of the year by heavy tropical rain-
fall and not reached by flood water from the rivers, the presence of lime-
stone beneath the soil does not necessarily greatly influence the vegetation.
Cohune ridge occupies the banks of the rivers and low level lands not far
from running streams. The soil is rich loam and the vegetation is locally
reputed to be "perhaps the richest in the world" (Dillon,' p. 18). The
plant association, or soil type which is coextensive with it, takes its name
from the great abundance of the cohune palm, Orbignya cohune (Mart.)
Dahlgren (Attalea cohune Mart.), called by the Spanish corozo, of which
Standley 2 says (p. 111):
"This is by far the finest and most imposing of all Central American
palms, its huge leaves greatly exceeding those of every other species. Not
withstanding their size, they are very graceful, and the corozo palm is a
tree of great beauty."
The Spanish name of the plant association characterized by this noble
palm is corozal. An alternative name for cohunee ridge," found on some
'A. Barrow Dillon, Geography of British Honduras, 1923, London. (Published by the
Crown Agents for the Colonies.)
'P. C. Standley, Flora of the Lancetilla Valley, Honduras, Field Mus. Nat. Hist..
Bot. Ser., vol. X, 1931, Chicago.
BOTANY OF THE MAYA AREA
of the government maps is "mahogany ridge," since mahogany is the chief
economic product of the vegetation type. When the mahogany is culled out,
mahogany ridge becomes cohunee ridge." In British Honduras and in the
Peten there are areas of rich forest without cohune, appropriately enough
known as "mahogany ridge" or sapodillaa ridge," to which the term cohunee
ridge" can not, of course, be applied at all.
Pine ridge occupies the areas between the rivers where the soil is sandy,
and also certain interior areas at higher elevations where the soil is not
alluvial or water-worked but locally derived from underlying granitic or
other crystalline rock. On this basis it is possible to make a distinction
between (1) lowland pine ridge, and (2) mountain pine ridge, which, as far
as studied by the writer, comprises the "Mountain Pine Ridge," called on
some maps "The Great Southern Pine Ridge." The lowland pine ridge is
so flat that it may be submerged more or less constantly in places, but there
is no great opportunity for variation in vegetation caused by relief, except
by alternation with wet savanna. In the Mountain Pine Ridge, however,
there are (1) deep ravines with a well-marked ravine flora, (2) the pine
land proper, and (3) swampy meadow, often with mat vegetation of
Cyperacewe and Sphagnum, at the edge of streams. Pine ridge soils are
seldom or never utilized for agriculture. The great areas in the lowland
extend back from the mangrove belt as tongues between the rivers, some-
times practically touching the rivers. The best-known pine ridges lie be-
tween the New River and the Hondo (August Pine Ridge and large areas
not named on maps), between the New River and the Belize northwest of
Big Falls, between the Belize and Sibun Rivers (Young's Pine Ridge), and
along the coast from Belize southward to Deep River. The canal which
extends southward from Belize, crosses the Sibun River, connects with
Northern Lagoon and Manatee Lagoon, and gives access to savanna
country forming the transition from the mangrove association to pine ridge.
Along the river banks, mangrove grades, by an interesting intermediate as-
sociation of Bactris, Oreodoxa, Pachira, Diffenbachia and other plants,
occupying low muddy alluvial sandy clays, into cohune ridge. Near Gales
Point and elsewhere on the Manatee Lagoon, the pine ridge country
extends to high-tide level and practically reaches the sea.
A beautiful and typical pine ridge may be seen within a short distance
up the Belize River from Belize, at the aviation field prepared for (but not
at present used by) Pan-American Airways. (See Plates 1 and 2.)
Pine ridge reminds the botanist from the United States of the South-
eastern Coastal Plain. The flora is extremely interesting but not at all
more complex than we customarily find in our coastal plain pine barrens.
The character tree is Pinus caribma, and the live oak, Quercus oleoides C. &
S. var. australis Trel. which often dominates the association, is closely re-
lated to our own live-oak, Quercus virginiana Mill. (See Plate 1, fig. 2.)
Broken ridge is the name applied locally to areas between cohune ridge
and pine ridge. When the belt of cohune palms and transitional broad-
leaved species is narrow between the river and pine ridge, the description
given by Morris (see below) is admirable. However, as Lundell states,
there are areas in northern British Honduras where the forest between the
cohune ridge and the pine ridge may cover a strip as much as ten miles
in diameter. In such instances, proceeding from the cohune ridge which
is subclimax, we next encounter the climax sapodilla-mahogany (zapotal-
caobal) forest which except for the absence of the cohune palms is as
luxuriant as the forest of the cohune ridge. This climax forest occupies
black shallow clays where outcropping limestone is common. Proceeding
from the climax sapodilla-mahogany forest toward the pine ridge, a zone
of marginal scrubby growth is encountered as the pine land is approached.
This latter zone is sometimes designated as the sequelar. Broken ridge
therefore includes climax and marginal scrub forest when the belt between
the river and pine ridge is wide, but only scrubby marginal growth when
the belt is narrow.
These three primary areas were described many years ago by Morris.1
As far as the major types of vegetation are concerned, his descriptions are
graphic, although he did not mention many species. He was vague, more-
over, in his conception of the geological factors concerned in plant distribu-
tion, since he thought that the subsoil of the colony was prevailingly non-
calcareous. Thus he said (p. 59):
"Geologically speaking, a cohune ridge has been formed by a river
valley, or depression in the quartzy ground-floor of the country, being, in
process of time, filled up by large deposits of fine alluvium and vegetable
As a matter of fact, it is more likely that the thin sandy mantle which
covers the flat country, overlying limestone in all the northern parts of the
colony, was eroded away in the stream valleys, exposing limestone on which
the calciphile vegetation, reaching its subclimax in cohune ridge and climax
in mahogany-sapodilla forest, established itself. Subsequently some of the
stream banks have been built up higher than the land back of them, by
silting, and arc possibly maintained in a calcareous or neutral condition by
being periodically overflowed by calcareous river water.
Morris (1.c., p. 63) particularly well described the vegetation types en-
countered in going across country from one stream to another within the
calcareous region. He may be quoted at some length in this connection:
"In addition to pine-ridge and colune-ridge there is sometimes known a
district possessing a vegetation of its own, to which the colonists apply the
term broken-ridge. This broken-ridge country generally appears to lie on
the outside, and generally parallel to and continuous with the cohune-ridge;
and, in fact, is an intermediate belt of vegetation coming between it and the
'Daniel Morris. The Colony of British Honduras, 1883. London
BOTANY OF THE MAYA AREA
pine-ridge country. The trees in this belt are smaller than in the cohune-
ridge; the undergrowth is denser and more scrubby in character; and, gen-
erally, the conditions indicate a poorer and less luxuriant phase of plant life,
toning down more and more until it merges into the scant sparse vegetation
of the pine ridge country... The broken ridge is no doubt due to a difference
in the character of the soil, which, having a slight depth only of humus and
alluvium, is able to support a less luxuriant vegetation than the cohune-
ridge but a little more so than the pine-ridge, which is almost devoid of
these elements of plant food."
What Morris describes is true only where the broken ridge belt is narrow.
"Starting from a river bed and traversing the country at right angles to
its course, there first comes the cohune-country, then the broken-ridge and
lastly the pine-ridge. The latter generally acts as a watershed between the
several river basins, and the order in which the ridges come may be shown
Cohune- Cohune- Broken- Pine- Broken- Cohune-
ridge River ridge ridge ridge ridge ridge River
"The vegetation of the cohune-ridge comprises tall-towering timber trees,
the lordly mahogany and luxuriant palms; while the ground below is cov-
ered with shade-loving ferns, selaginellas, and aroids. The broken-ridge has
fewer, less luxuriant, and somewhat stunted timber-trees, such as the cock-
spur, abundant in prickles and thorn; the supa, or gru-gru palm (Acrocomia
sclerocarpa) and small-leaved spiny shrubs of Randia, Capparis, etc. In
the more open spaces, coarse bromeliads, rank grasses, and prickly creep-
ers impede one's movements until at last the open pine-ridge country is
reached. Here, as already mentioned, the tall (Scotch-looking) firs, or
pines, are the more striking objects, surrounded, when in clumps, by the
shrubby pimento palms,1 with the craboo2 and the haha8 trees dotted
here and there. Amongst the hard, coarse grass of the pine-ridge, small low-
spreading shrubs are found, such as Pithecolobium ligustrum and Cassia
diphylla; a few ground orchids (Habenaria and Stenorrhynchus) and small,
From the economic standpoint the calcareous or semicalcareous alluvial
lands have been of preeminent importance to British Honduras for the rea-
son that they are the habitat of the mahogany tree. To one who is con-
vinced, as the writer is, of the paramount importance of the distribution of
limestone in determining the major ecological subdivisions of the flora, it
seems impossible that there should not be two fundamentally distinct climax
floras (namely broad-leaved and coniferous) on the higher lands, one for
each primary soil type, and that between them (occupying what has been
See the paper following in this volume for a description of the pimento palm as
Acelorraphe pinetorum Bartlett.
SByrsonima pulchra DC. is, according to to Mr. C. V. Morton's determinations, the
common "craboo" of British Honduras.
SCuratella americana is a small tree or shrub with broad very rough leaves, known
by the Spanish-speaking inhabitants as chaparro and by the "Creoles" as ahha.
well called a tension zone) there should be a transitional flora (sequelar of
the broken-ridge) of extremely variable composition. It is only fair, how-
ever, to point out that not all observers come to this point of view.
Mr. N. S. Stevenson (Deputy Conservator in the Forest Department,
British Honduras)' has expressed the opinion that-
"the mahogany forests of British Honduras have originated in two ways.
(1) The primary or natural way by invasion of the pine-lands by groups of
broad-leaved trees, producing patches of forest containing mahogany and
sapodilla (Achras sapota L.) locally known as 'broken-ridge.' Coalescence
of these groups follows, to form the more advanced type known as 'high-
ridge,' with final advancement, where conditions permit, to the climax,
cohune-ridge, with Attalea cohune Mart. Spontaneous reproduction of the
succession, even from the pine stage, may take place after fire or other for-
est destruction. (2) The secondary or artificial way, chiefly seen in the hill
region, takes place by recolonization of the abandoned clearings made by
the scanty and shifting population. The forest phase preceding these clear-
ings has been, in all probability, climax forest of the natural type. Such
secondary growth is known as huamil. Spontaneous regeneration of trees
is most prolific at the broken ridge stage and decreases progressively to-
wards the climax; regeneration in the cohune ridge being almost suppressed
by the dense shade of the Attalea."
Stevenson considers the cohune ridge to be climax, whereas in all proba-
bility it is subclimax, the climax being the mahogany-sapodilla forest of
the "high ridge."
Stevenson's conception, like that of Morris, disregards completely the
view that the calcareous (neutral) or non-calcareous (acid) nature of the
soil fundamentally controls plant distribution and the nature of the climax
vegetation. His idea that pine ridge is supplanted by broken ridge and
finally by cohune ridge could only be true within the zone of silting and
supply of calcareous water by stream overflow. Given geological time
within which streams could base-level the country and meander far from
their present beds, his idea might be correct, since the substratum of the
pine ridges would be, by the geological process of silting, modified into some-
thing essentially different. As far as succession on the geologically unmodi-
fed soil of the typical pine ridge is concerned, however, it is perfectly safe
to say that the pine climax would never become a cohune or mahogany-
This point of view, i.e., that there is a rigorous control of the vegetation
by the distribution of lime, found an energetic local advocate in L. H.
Ower,2 Government Geologist of British Honduras, whose conclusions did
not agree with those of the Conservator of Forests.
'Quoted from Abstract in Jour. Ecol. XVII Supplement 4. British Empire Vegetation
Abstracts, p. 67. Neil A. Stevenson, Hilvicultural treatment of Mahogany Forests in
Britih Honduras, Empire Forestry Jour.. vol. 6; 219-227, 1927.
'Leslie H. Ower, The Geology of British Honduras. Printed by the Clarion, Limited,
Belize, British Honduras. (Pamphlet ofi 24 pages; unpaged, no title page, no date, with
over title as follows: "Map of British Honduras, Compiled in Surveyor General's Depart-
ment, by G. A. Elliott, C. L. S.. from Notes and Surveys of L. H. Ower, Government
Geologist. Scale 1/1.000.000. Published by permission of Royal Geographical Society.")
BOTANY OF THE MAYA AREA
Ower holds that the extent of Maya settlements (i.e. those that were
based upon agriculture that left vestiges of any sort) coincided strictly
with the extent of calcareous soils. He states that all calcareous soils in
the colony (except trifling areas) have at some remote time been in cultiva-
tion, and that they are therefore now occupied not by primary but by sec-
ondary forest. On the contrary, he concluded that the non-calcareous areas
have a primary plant cover which has never been extensively modified by
man, since they are not now and never have been cultivated. In the course
of his geological survey he found traces of terrace cultivation in practically
all of the limestone upland. He concludes that the area of mahogany and
sapodilla coincides with the area of old cultivation, and therefore with the
limestone area. He says:
[The Maya] "lived almost entirely upon the calcareous area. On the
limestone plateau abandoned agricultural terraces are very frequent wher-
ever a few yards of soil were obtainable . The whole of the limestone,
with the exception of that near the source of the Rio Grande, contains the
relics of terrace cultivation. This old cultivation has been of the greatest
importance to British Honduras, for where it is absent mahogany is seldom
seen . The whole of the Colony, with the exception of the pine belts,
the slates, porphyries, and some small limestone areas, has probably been
under cultivation at some time, so much of the present vegetation is of sec-
ondary growth. . [Cohunes] seldom extend more than a mile back from
the streams but will be found in the limestone depressions and on rich
soil. . Reports by agricultural and forestry experts have been published,
but the geological factors influencing these resources have hitherto been
unrecognized. The geological map to a large extent serves as a vegetation
or soil map of British Honduras. . Mahogany is nearly always found on
the land that was once cultivated, so is seldom seen in the slate country."
Except for the dubious implication that former human occupation of the
calcareous areas extended the range or abundance of mahogany (which
may be quite true in spite of lack of evidence) the conclusions of Ower coin-
cide closely with those of the writer.
It would seem almost beyond question that there are two chief upland
climax vegetations, one calciphile and one calcifuge. They have few species
in common. The influence of the lime is, of course, in part secondary,
through preventing soil acidity, and the plants which appear to avoid lime,
in some measure, but no means entirely, are those that are tolerant of acid
soils and are not successful competitors in the vigorous flora of limy soils.
It is thus possible to account for the sequelar flora of the broken ridge.
LOCAL DESIGNATIONS OF PLANT ASSOCIATIONS:
THE SPANISH TERMINOLOGY.
In the Peten upland forest the plant associations are not very different
from those of the adjoining but lower calcareous districts of British Hon-
duras. There is indeed some change, as would be expected, corresponding
SOwer here implies but does not definitely say that he believes mahogany to prevail
in the calcareous region because of former human occupation of the land. He writes:
"The Conservator of Forest is not in agreement with the statement ...
to the transition from the adequately watered coastal districts to the under-
drained limestone region of the Peten, and the change is not altogether
or even primarily caused by water supply, since a large part of the flora of
Mid soils found in the pine ridges of British Honduras, and straying at least
into the edge of the plant habitats underlain by limestone, drops out as the
Peten is entered. Nevertheless some of the pine ridge plants also grow in
the drier parts of wooded swamps (bajos) in the calcareous Peten upland,
and thus are important components of two physiographically very dissimi-
lar habitats. As an example the live oak, Quercus oleoides var. australis,
may be instanced. It is characteristic of oak "islands" in the pine ridges,
where it is a dominant plant. Apparently the same oak varies in abundance
from a minor to a common constituent of the vegetation in the drier parts
of the logwood swamps. Its associates are for the most part an entirely dif-
ferent group of species from those found in the pine ridges. The bajos
(wooded swamps) do not exclude plants usually found in sour soils, even
though they occupy depressions in a limestone country. They are doubt-
fan acid habitats. On the adjacent limestone of the "high bush," however,
I did not see a single oak.
I have chosen to list the plant associations (corresponding to habitats)
of the whole region of central British Honduras and the adjoining Peten
forest and to indicate particularly those associations which are encountered
only in a particular part. Incidentally as helping in an approach to com-
pleteness, I have listed some associations which the natives refer to, but
which I have not seen.
.i we omit the halophytic plant communities, the others are as follows:
(1) Zapotal (Peten high bush). The writer is in agreement with Lun-
de1's conclusion that Achras sapota is a more general constituent of the
"high bush" of the calcareous lands of northern British Honduras and the
Peten than any other one of the major forest trees, and that therefore the
term sapotal is the best of several eligible terms by which to designate inolu-
&ively the several plant associations that differ from one another in present-
ing with different local dominants much the same assemblage of species.
The writer had used "ramonal" as the inclusive term for the "high bush"
ieociations, but Lundell's observations have been more extensive than the
writer's on this point. Not only has he studied the distribution of zapote
in the "Eastern Coast" area of British Honduras but he has also traversed
Oh entire western and central parts of the Peten "high bush." The writer's
Eterrations of the zapotal have been made in the vicinity of El Cayo and in
the stretch of country from El Cayo northwestward to Uaxactun, Peten,
that is, all in the eastern part of the Peten forest.
The general prevalence of zapote chico (Achras sapota) throughout so
great an area, and the apparent fact that the vast majority of the trees
seem to belong to a very old age class suggest that the factor of human
:_upation of the land was all important in determining this major plant
aseoeiation. It is possible that the usefulness of the very sweet fruit as
food in a sugarless diet may have led the Maya to spare the zapote when
other trees were destroyed in the clearing of land for planting annual crops.
BOTANY OF THE MAYA ABBA
It is almost certain that during the period of densest population, numerous
islands of trees would have been left for shade at house and village sites,
and that as wood was needed for various purposes there would have been
an elimination of species that bore no useful fruit or other product. We can
imagine the Maya of old sparing useful wild trees just as they do today, and
just as primitive man does in other parts of the world.
The writer has observed the extreme reluctance of the lowland Batak of
the East Coast of Sumatra to destroy certain useful plants of the jungle,
and believes that the extent to which the forest is "improved" by these people
through setting out seedlings has never been properly appreciated. Very
often one may find near a native clearing an appropriate site in the edge of
the forest occupied by a seed bed of the more useful species of wild rotan,
which are dug when big enough to transplant, done up into bundles and
planted in favorable sites throughout the forest, in replacement of mature
plants removed for sale. The chiefs require that this be done, and one of the
frequent complaints of the natives when they are moved out of lands granted
to foreigners as plantation concessions is that their claim to private owner-
ship of apparently wild plants is made light of. The highland Batak both
spare and propagate useful woody plants about their villages, so that the
landscape resembles a sea of rice fields with numerous wooded islands in
which the villages are concealed. If their land were abandoned, the presence
of these wooded islands would certainly influence the flora for centuries,
both through the actual persistence of individual trees and through the fact
that certain selected species would have the advantage of propinquity in
seeding the rice terraces and would thus reestablish new forest of a particu-
lar type. This would be especially true if the cultivated and semicultivated
species were themselves natives of the land. The dry-land milpa of the
Maya would never have required the clean clearing of the wet rice terrace,
but would have resembled more the Malay ladang or Batak djoema of
Sumatran primitive agriculture, in which we may be sure that a useful fruit
tree would have been left. The sugar-palm groves of lowland Sumatra,
which are one well-marked plant association of second-growth forest are
often if not always of artificial origin. To argue about what the Maya may
have done from what a tropical race on the other side of the world actually
do may not seem very convincing, but the line of thought is one which
should be productive in the endeavor to determine the ecological succession
from cleared field to forest and the effect of ancient human occupation upon
the present flora of the Maya area.
The zapotal is a highly variable plant association which grows on the
high well-drained calcareous areas. It is characterized from Uaxactun east-
ward to the Guatemalan boundary by an abundance of at least part of the
A. Exogenous trees, arranged alphabetically, starred if sometimes among the six or eight
dominant trees of the association:
Acacia collUnsl Safford *(fwalpini violaces (Miller) Standl.
Ansona (Bartlett 18474) COealpini a .atanesmi Oreenm.
*Aspidosperma megaltocrpon Muell. Arg. *Cedrels msican~ Roem.
Aspidosperms sanguineum Bartlett 0*etba peatandra (L.) Gaertn.
Astrocasia phyltantheodes Robins. & Millsp. *Celt hotftet Standl.
*Burerire oephylla Stand). Olusia (2 species) (See Plates 3 and 4)
*Brofimum alicastrum Sw. *Oupanla belisesus Standl.
*Bumelis mayana Standl. *Dipholite oaliefolia A. DC.
*Burera aimaruba (L.) Sarg. Drypetes brownat Standl.
ktp~er laterjifera (Sw.) Krug. & Urban
Dnthe paniesuato (Guss.) Radlk.
*eorchammerts trifoliata Radlk.
squaress eseelas H. B. K.
*Brtlll americana L.
Ifrugodendron ferreum (Vahl) Urban
eea iftsamnta Sw.
sowsloearpur coatiloi Standl.
*Utems eampechiana H. B. K.
tLuWana darlandii Standl.
Mfalma depreua (Bail.) Fr.
*JM ta ofsteaUit Lindl.
S'roism eopal (S. & C.) Engl.
*Pamsdolmedia spura (Sw.) Griseb.
3. Xxogenous shrubs or treelets:
Ptper coobanes Trel.
Piper villipetiolatum Trel.
Piper yaloehanum Trel.
C. Bndogenous woody trees or shrubs:
Stat morrNaisna Bartlett (See Plates 3
Orpooephlta argentea Bartlett'
Asthurium &eulum Schott
Moatesr guatemalenais Bartlett
PlWodendron mithit Engl.
Deasonve ferom Bartlett
Daonscus uamactunensis Bartlett
Quararibea fleldii Millsp.?
Sapindus saponaria L.
Sickingia salvadorensis (Standl.) Standl.
*Siderooylon amygdalinum Standl.
*Sideroaylon meyeri Standl.
*Simaruba glauca DC.
'Stemmadenia donnell-mithii (Rose) Wood-
*Swietenia macrophylla King (See Plates 3
*Tabebuia pentaphylla (L.) Hemal.
*Talisia floresii Standl.
*Trichilia minutiflora Standl.
*Vites gaumeri Greenm.
Wimmeria concolor S. & C.
Psychotris limonenais Krause
Ruellia stemonaeanthoides ((Erst.) Hemsl
Opsiandra maya Cook (See Plates 10 and
Chamadorea (several species)
Dracana americana Donn. Sm.
Dioscorea bartlettii Morton (See Plate
Smilax medical S. & C.
Adenoealymna fissum Loes.
Dalbergia glabra (Mill.) Standl.
Petrea arbores H. B. K.
Vitis tiirefolia H. B. K.
Phases of the zapotal to which separate names are locally applied are
characterized by fluctuations in abundance of the dominant species. The
chief ones are:
(la) Ramonal. The dominant species is the ramon, Brosimum alicastrum
8w. This species is one of the economically very important trees of the
region, since it is cut for fodder, and is essential in the "high bush" for feed-
ing mules, which provide the only transportation of the region. Lundell 2
has noticed a high correlation between the abundance of ramon and the
presence of Maya ruins. It is very probable that the ramon was left stand-
ing or may even have been planted by the Maya, as a resource in the event
BSee following paper in this volume, and also Plate 7.
'He writes: "I have found groves of the ramon tree covering every Southern Culture
frua which I have visited, and it is no mere coincidence that this species is so abundant
there. Of the other trees, cacao and mamey were doubtless of the greatest importance
as additional tree crops, for they also bear fruit in the dry season. It is probable that
these trees and many others which are now found in the region also grew in some of
the plazas and streets, providing both shade and food." C. L. Lundell, The Agriculture
of the Maya, Southwest Review, vol. 19, 65-77, 1933. (See pp. 71-72.)
BOTANY OF THE MAYA AREA
of crop failures. Its fruit may be used as a human food, as indicated by the
Creole name, bread-nut. If this supposition regarding ramon should be sup-
ported by future investigations it will afford additional support for the sup-
position that the plant associations of the Peten forest were determined
largely by human agency centuries ago. The ramon is presumed to have
been given its original dominance by preservation when less useful trees were
destroyed or to have been actually planted, and never to have been forced
into a secondary r8le as the forest reclaimed land deserted by human occu-
(lb) Caobal. The important, even if not dominant, species is mahog-
any (caoba), Swietenia macrophylla King. (Plates 3, 4.) It may be assumed
to have spread quickly to abandoned lands because of its winged seeds.
(Ic) Uacutal. This phase is very rare and characterized by the gigantic
tree called uacut, Bernoullia flammea Oliver. (See Plates 4 and 5.) I have
seen it only amid the ruins of Tikal, where the flora was not fully collected
in the brief time at my disposal, but contained the following species:
Bernoullia flammea Oliver Cryosophila argentea Bartlett
Achras sapota L. Chamwdorea (several species)
Brosimum alicastrum Sw. Bactris sp.
Bumelia mayana Standley Dracena americana Donn. Sm.
Jacquinia aurantiaca Ait. Piper tikalense Trel.
Miconia impetiolaris (Sw.) Don Psychotria flava (Erst.
Ouratea pyramidalis Riley Dryopteris melanosticta (Kuntze) Kuntze
Sabal morrisiana Bartlett
(ld) Cedral. Characterized by the dominance of Cedrela mexicana Roem.
(le) Manaxal. So called if "cherry," Pseudolmedia, (manax), is
(If) Higueral de las ruinas. The undisturbed vegetation of the ruins is
similar to that of the ramona or the caobal, but has two characteristics
of its own, namely, the abundance of strangling figs, which, enveloping the
stones of the masonry and eventually falling, are responsible for much of
the destruction of the old buildings, and the presence of certain plants that
may possibly represent vestiges of an ancient ruderal and semicultivated
plant association. On the uncleared ruins of Uaxactun one finds a random
assortment of the "high bush" (caobal) flora with at least the following
species more in evidence than usual, and some of them with the aspect of
being weeds or of weedy propensity:
Ficus lapathifolium (Liebm.) Miq. Celosia nitida Vahl
Bumelia mayana Standl. Iresine celosia L.
Amyris sylvatica Jacq. Dioscorea bernoulliana Prain & Burkill
Bauhinia divaricata L. Passiflora biflora Lam.
Hybanthus yucatanensis Millap. Smilax mollis Willd.
Bernardia interrupt (Schlecht.) Muell.-Arg. Tournefortia hirsutissima L.
Piper cobanense Trel. Antharium tetragonum Hook. var. yuea-
Carica papaya L. tanense Engl.
Solanum lanceifolium Jacq. Agave sp.
Rolanum nuaum L. Euphorbia graminea Jacq.
Borreria levis (Lam.) Griseb. Neurolwna lobata (L.) R. Br.
Psychotria pubescena Sw. Rhoeo discolor (L'Her.) Hance
Capsicum macrophyllum (H. B. K. ) Standl. Adiantum tricholepis Fee
Hibiscus Bp. Asplenium dentatum L.
Russelia campechiana Standl. Dryopteris patens (Sw.) Kuntze
Otopappus scaber Blake Pteris biaurita L.
Bchistocarpha oppositifolia (Kuntze) Rydb. Pteris grandifolia L.
Amaranthus dubiau Mart. Pteris quadriaurita Retz
(1g) Guarumal. If the "high bush" is cleared and allowed to revert to
forest, a very conspicuous phase in the succession is that in which Cecropia
(guarumo) springs up as a dense growth. At Uaxactun it is accompanied
by a wild papaya (Carica papaya L.) with very small practically worth-
less fruit. The guarumo is a very rapid grower and with the papaya makes
a dense thicket in which the soft young trees can be as easily chopped with
a machete as stems of herbaceous plants. The hollow stems of guarumo are
inhabited by ants that bite viciously. The flora is exceedingly weedy and
is probably seeded in large part from the more scattered and less conspicu-
ous weedy plants of the ruins. At Uaxactun were listed:
Cecropia obtusa Tricul
Carica papaya L.
elitis trinervia Lam.
GOaltunm ulmsfolia Lam.
Frema floridana Britton
Hamelia patens Jacq.
Psychotria pubescens Sw.
Buhinia divaricata L.
Bypbnthue yucatanensis Millsp.
Piper auritum H. B. K. var. amplifolium
Piper cobanense Trel.
Piper villipetiolatum Trel.
COisanpelos tomentosa DC.
Dioscorea bernoulliana Prain & Burkhill
Passiflora adenopoda DC.
Pa8siflora biflora Lam.
Calonyction aculeatum (L.) House
lposeoa polyanthes R. & S.
Amaranthus dubius Mart.
Iresine celosia L.
Celosia nitida Vahl
Desmodium frutescens (Jacq.) Schindl.
Borreria larvis (Lam.) Griseb.
Rivina humilis L.
Acalypha (Bartlett 12464)
Sida glutinosa Commers.
Hibiscus clypeatus L.
Capraria biflora L.
Russelia campechiana Standl.
Capsicum macrophyllum (H. B. K.) Standl.
Capsicum viscidum Standl.
Cestrum panamense Standl.
Solanum nudum L.
Tournefortia hirsutissima L.
Cirsium mexieanum DC.
Schistocarpha oppositifolia (Kuntz) Rydb.
Neurolaena lobata (L.) R. Br.
Leptochloa virgata (L.) Beauv.
Dryopteris patens (Sw.) Kuntze
Pityrogramma ealomelwna (L.) Link
(lg) Botanal: guanal. In the borders of the "high bush" along the
bajos the flora is characterized by a predominance of palms, the species
being also found on the higher ground with the plant associations already
enumerated. I have called the transition flora botanal if Sabal morrisiana
dominates and if the soil is moist enough so that there are filmy ferns on
the bases of the botan trunks. (See Plate 6.) The botanal passes into the
escobal, a true bajo association, in which Cryosophila dominates, often
accompanied by Desmoncus.
(2) Corozal. This is also one of the vegetational units classified in Brit-
ish Honduras as "high bush" or cohunee ridge." It extends through Peten,
but is not found near Uaxactun. It occupies alluvial stretches, overlying
limestone, and takes its names from the dominance of the corozo, or cohune,
palm. It is found along the rivers and in some favorable areas far from
running water. Here are found dominantly:
poroso (Orbignya cohune (Mart.) Dahlgren.) jobo (Spondias purpurea L.)
ewob (Swietenia macrophylla King) zapotillo (Lucuma durlandii Standl.)
oerdoncillo (Piper, various shrubby species) cherry: (Pseudolmedia spuria (Sw.) Griseb.)
zapote macho (Achras chicle Pittier)
(2a) Guamil: matorral. If an area in "high bush" (ramonal or corozal)
is cleared and used for pasture, or otherwise kept open until it becomes
grassy, it is quickly occupied, when abandoned, by coarse weeds, shrubby
plants and quick-growing second-growth trees. The association is very
diverse. A phase which can hardly escape particular notice is:
(6a) Encinal. Oak "islands" or groves in the pine ridge. If the oaks
(mostly Quercus oleoides var. australis Trel.) are dispersed uniformly
among the pines, this phase can not be distinguished, but the oak islands
are generally well defined and are sometimes groves of considerable size,
containing large trees. (See Plate 1.)
From the standpoint of the shrubby and herbaceous vegetation, which is
greatly influenced by the nature of the shade and by minor variations in
level, it would be possible to distinguish a considerable number of sub-
associations of the pine ridge, some of which would have exact or very nearly
exact counterparts in the sabana. One which is perhaps named for practi-
cal reasons (just as we distinguish a huckleberry swamp from other
unclassified but very similar swamps because it has huckleberries in
-bundance, or as we designate a poison sumac swamp as such, ignoring
the species which do not immediately concern us) is:
(5b) Nanzal. An area in which Byrsonima pulchra DC. (nanze) is pre-
dominant. This subassociation is not coordinate in importance with the
esninal, since the nanze is but one of several shrubs of similar habit of
growth which fluctuate in abundance. It is always associated, for instance,
with Cwatella americana L. (haha; chaparro) which is often dominant.
(See Plate 2.) This association does not intrude upon the encinal, but may
~b a part of either the pinar or the sabana. If the pines of the pinar are
destroyed and reproduction is prevented by fires, the pinar becomes:
(6) Sabana. This is a more or less open plain with herbaceous vegeta-
tion, isolated trees of Crescentia (see Plate 2), and scattered shrubs, with
or without clumps of palmetto and oak. The sabana which is seen along
the coast of British Honduras may be entirely natural, the growth of pine
being perhaps prevented by the presence of too much salt in the soil.
There may be other natural sabanas.
(1) Cipresal: barrancal. Within the Mountain Pine Ridge of British
HRnduras are ravines containing a very different vegetation from that of
the fiat pine land. The characteristic plants of the ravines will probably
be found to belong to the flora of the non-calcareous mountain area to the
suthward, which is as yet inadequately known. Here the relatively rich
flara includes Podocarpus pinetorum Bartlett 1 (cipres), Ormosia, Ilex,
Qreopanax, Clusia, the mountain cabbage palm, two tree ferns, and many
Vbar interesting plants. The very edge of the ravine, especially if it slopes
Olfsomewhat gently, has a very different association from that of the pine
rdge proper (pinar) or the cipresal. It is a dense bracken thicket called:
(7a) Cizal: helechal. The predominant plant, nearly a pure growth, is
itsilchican, Dicranopteris pectinata (Willd) Underw.
(8) Escobal. This is the type of wooded swamp which lies most nearly
at the "high bush" level and is therefore really transitional from the "high
.ush" associations to the bajos, or swamps. It surrounds the small per-
manent or subpermanent water holes (the aguadas) of the upland and may
'Padocarpus pinetorum sp. nov. a P. oleifera differt foliis supra hand sulcatis, a
P. lateaelense stature arborea, diametro 25 cm., et foliis duplo brevioribus, rectis nee
rlcatia-Specimen typicum legit H. H. Bartlett n. 13109, "Mountain Pine Ridge, El Cayo
District, British Honduras, 8 May, 1931." I have somewhat reluctantly separated this
frBtP. watemaleaeia Standley, since the type of the latter (Standley 25090, "wet thicket"
at Puerto Barrios, Dept. Izabal, Guatemala) was a shrub 6 feet tall with falcate leaves
twice as long as those of P. pinetorum (i.e. up to 15 cm. long, and 9-11.5 mm. wide).
Thre is a possibility that Standley's specimen represents a juvenile state of the same
ppeeies, but to reach any certainty about the matter it will be necessary to study
P. ustemalessi at the type locality.
BOTANY OF THE MAYA AREA
constitute much of the higher part of the bajo itself, the term bajo being
used in its broad sense as including all the wooded muddy swamps. (See
Plates 7 and 8.) In the escobal are found:
escoba (Cryosophila argentea Bartlett) Dioscorea bartlettii Morton
raoba (Sw'ieenia marrophylla King) Hirwa obovata (Kunth) Niedenzu
botan (Sabal morrisiana Bartlett) Paullinia fjuceseen H. B. K.
jobo (Spondian purpurca L.) Serjania scatena Radlk.
capote chiro (Aehras sapota L.) Passiflora (Bartlett I2si0)
milady blanco (Aspidosperma megalocarpon Petrea arborea H. B. K.
Muell. Arg.) Strychnos panamensi8 Seem.?
milady colorado (Aspidosperma sanguinale. Aretottonia sempervireM Trel.
Bartlett) Piper cobasense Trel.
grand betty (Cupania belisenai Standl.) Piper pailorrhaeke C. DC.
water wood (Cestrum panamense Standl.) Ardisia escaltonioides 8. & C.
mora (Chlorophora tinctoria (L.) Gaud.) Coccoloba refleeifora Standl.
Trichilia moschata Sw. Borreria oeimoides (Burm.) DC.
Ouratea jurgensenii (Planch.) Engl. Morinda yucatanensis Greenm.
Acacia collinsii Safford Psychotria fruticetorum Standl.
Lonchocarpus guatemalensis Benth. Psychotria undata Jacq.
Lonchocarpus (Bartlett 12573) Agave (Bartlett 12850)
Lonchocarpua hondurensis Benth. Blechum pyramidatam (Lam.) Urb.
Bucida buceras L. Vernonia aschenborniana Schauer
Nectandra glabrescens Benth. Zeemenia serrata Llave
Bauhinia dn. Euvatorium macrophjllum L.
tfits (Hemmatoxylum campechianum L.)
uayabilUo (Hasseltia mexicana (Gray)
.spote chico (Achras sapota L.)
black poison-wood (Metopium brownei
neiWo (Quercus oleoides H. B. K. var.
O lsia Bp.
Pithecolobium lanceolatum (H. & B.) Benth.
Mimosa hemiendyta Rose & Robins.
Ardisa escallonioidee S. & C.
Eugenia lundellii Standl.
Oryosophila argentea Bartlett
Berardia interrupt (Schl. Muell.-Arg.
Croton niveus Jacq.?
Oroto reflemifolius H. B. K.
Capparis cynophallophora L.
Gymnopodium antigonoides (Robins.) Blake
Ternatrwmia spherocarpa (Rose) Melch.
Miconia ambigua (Bonpl.) DC.
Morinda (Bartlett 12808)
Psychotria flava (Erst.
Hirea borealis Niedenzu
Passiflora palmeri Rose var. sublaceolata
Serjania scatens Radlk.
Berjania adiantoides Radlk.
Wedelia adhwrens Blake
Wedelia parviceps Blake
Borreria verticillata (L.) Meyer
Egletes viicosa (L.) Less.
Eupatorium odoratum L.
Perymenium peckii Robins.
Seleria melaleuca S. & C.
This association represents a late phase in the silting up and transforma-
tion into dry land of shallow lagoons (lagunas) which were their geological
predecessors. (See Cooke.1) With difficulty one may generally get a mule
through the tintal even in the wet season.
(10) Tembladeral. This is the wettest part of the bajo, and is avoided
at all seasons since mules are likely to be mired in it and lost. Its charac-
teristic plants vary from place to place. Two definitely different associa-
tions which are generally recognized (but were not studied by the writer)
are numbered 10a and 10b.
(10a) Julubal. Here grows a plant called julub (hulub), together with
vines and sedges.
(10b) Zapotebobal. Here the tree called in British Honduras "provision
tree" and in Peten zapotebobo is said to be dominant. Although character-
istie of quagmires near the coast above the mangrove belt, zapote bobo
(Pachira aquatica Aubl.) is said to grow in the wettest bajos within six
miles of Uaxactun. The plant is very easily recognized, and there is little
reason to doubt the reports of chicleros. At any rate they talk of zapote-
bobales, and of zapotebobo which grows in them almost to the exclusion of
everything else except vines (bejucos).
(11) Laguna: charco. At various places in British Honduras and Peten
there are long narrow shallow lakes in various stages of extinction through
silting and invasion by vegetation. They represent an early stage in the
development of bajos whose different plant associations have been defined.
The flora of the lagunas has not been investigated, but names were obtained
at Yal'och for two associations as follows:
(1la) Naabal. The waterlily association, characterized by utop naab,
Nymph a ampla (Salisb.) DC.
(11b) Carrizal. The reed association in the edge of the water. (Species
(12) Aguada. The water holes which are not parts of big bajos and lie
at a high enough level to be surrounded by some association other than the
tinta (either the escobal or botanal) constitute a distinct habitat. It
would be worth while to investigate as many of them as possible since they
vary greatly in floristic composition. The number of species found in each
S0. W. Cooke, Why the Mayan cities of the Peten District, Guatemala, were abandoned,
Jour. Wash. Acad. Sci., vol. 21, 1931.
BOTANY OP THE MAYA ARIA
is not great, and each one seems to have some two or three species which
may be missing at the next. Two types are as follows:
(12a) Lechugal. Pistia stratiotes L. (lechuga) floats in a dense mat on
the surface of the water, with Lemna and Wolffia. (See Plate 12.) Shrubs
and trees which grow in the water of an aguada of this type at Uaxactun are:
MsaShynomene deamni Robins. & Bartl. Loschooarpm guatemalensti Benth.
Alternanthera obovata (M. & G.) Millsp. Psyehotria arrtediana Standl.
(12b) Pital. The edge of the water is filled with a dense mass of saw-
toothed pita, Ananas magdalenm. The aguada at Tikal is of this type.
Here there are no free-floating species, but the following plants actually
grow in the water:
Anan.a magdalesn (Andre) Standl; Cagsa grand L. f.
Ardisia compresra H. B. K. Lonohocarpus guatemalen8ia Benth.
Trichilis havanenmis Jacq. Inoga purti H. & B.
(12c) Aguacatillal. Perhaps not practicably distinguished from the
escobal, but, appearing at the edge of aguadas, is an association in which
the Lauraces called aguacatiUo are conspicuous. There are probably
no species in this subassociation which are not also found along water
courses and in the escobal at the edge of the bajos, but the vegetation has a
different physiognomy. The species for a definite locality, Uaxactun, are as
Nectandra olabresaen Benth. (?) Chamr dorea (Bartlett 144t)
Bucida uceras L. Dioscorea mwtagalpenami Uline?
eebastiana longioupis Standl. Renealmis occdentalit (Sw.) Sweet (See
Lonohocarpus guatemalenss Benth. Plate 12.)
Trichilla motohte Sw. Typha angustifolio L.
Oalyptranthes ehytraouNli (L.) Sw. Oyperu (3 species)
Piper psilorrhache C. DC. Alternmanther obowata (M. & G.) Millsp.
Plsehotria aratediana Standl. Spilanthes americano (Mut.) Hieron. f.
Thevetoa sp. laniteota A. H. Moore.
Xyltoma flesuosa (H. B. K.) Hemal.
(13) Bambonal. A very characteristic association is that of the banks
of streams in which there is constantly or generally running water. The
character plant in many places is a viciously spiny bamboo, Guadua sp.
This association soon drops out after the Peten is entered, as the higher
limestone country is underdrained and the channels have no water during
the dry season. It was last seen at Holmul in the Peten. The river bank
associations have been described by Lundell (see p. 8).
(14) Arenal. One of the habitats which takes its name from the nature
of the soil rather than the plant association is the arenal-a sandy, but well-
drained, or at least not sour-soiled, place. The word is applied to sandy
beds of dry arroyos, and also to sand banks or sand bars exposed at low
water along streams. A characteristic plant is Xanthosoma, with which
grow a variety of herbs such as species of Ageratum, Acalypha, grasses,
sedges, etc. No arenal in British Honduras was carefully examined and
listed, but the slight representation (probably not typical) of this associa-
tion on sand bars in the dry Arroyo Uaxactun, at Uaxactun, had the follow-
ing species in identifiable condition:
Zotlosome lvuoatanense Engl.
AMiipkA vwillos Jacq.
triwUtylidium pittieri Hack.
'Pflisthanu fereo Standl.
yerppteris patens (Sw.) Kuntze
gopteris aubtetragona (Link) Maxon
PfWijpdium phvllitJdi L. (as a terrestrial
bouelis humili (Sw.) Urb.
Priva lappulacea (L.) Pers.
Lobelia berlandieri DC.(?)
Eupatorium pyonooephalum Less.
Bomeria ulmifolia Wedd.
Stenandrium aubcordatum Standl.
Opismenus hirtellus (L.) Beauv.
Panioum virgwltorum Hack.
Selaginella (Bartlett 12545)
This paper has been lying about two years unpublished, but has been
jgessible to Lundell, who has cited the manuscript. His more extensive
;tadies exemplify the use of the field method and contain much more
detailed lists for some of the same plant associations that I have described.
He has studied them at other places and has been able to present a general
.mccount of the plant formations and associations of the entire Yucatan
Peninsula. One of his longer contributions has been cited (see page 8)
and a still more extensive one is now in preparation and will be published
ma "Botanical Studies in the Department of Peten, Guatemala." I have had
the advantage of consulting Mr. Lundell's unpublished work, as he has had
of seeing mine, and many of our ideas have developed in partnership. The
Writer still plans to write a somewhat more extended account of the Moun-
tain Pine Ridge of British Honduras, based upon observations made in
1931, but Mr. Lundell's more extensive experience makes it desirable to
arve further and more detailed publication on the phytogeography and
Scology of the Peten forest to him.
DESCIPnON or PLATz 1
Fle. 1-Pine ridge near aviation field, Belize, with low foreground occupied by stretch
of sedge savanna (sabana), at edge of which, on left, one of filets of pimento
palm (Acelorrmphe pisetoru) just shows; pines (Pinas caribos) and oaks
(Quereos oleoides var. australi) in background.
FlP. 2-View of vegetational islet in pine ridge at Cornhouse Creek, Manatee River,
Belize District. Plant association is the eneinal, composed primarily of Quere
oleoides var, ausaralis bordered by pimenta palm, Acelorraphe pinetorum. Picture
was taken from stretch of sedge saba~a.
CARNEGIE INAT. WVA91IrN'r)N P(~ B. (161 BriI ILT
I)ESCRIPTION OF PLATE 2
Fi(. 1- View in pine ridge north of aviation lield. Belize. showing alternation of pine
association proper (piniar) in Iiackgroind with areas of sedlge abahnan which are
dotted here andi there with sniall pimento islets. The latter contain as chief
woody diot.vyledlon the halh (('uraltela americana) which is the little irregular
tree at Ihft. Sometimes tie haha and the nanze, lByrsonima pulchra, form
iSlets withulit tlle pimlentlot palmn. the association being then called chaparral
ifrolni happIaro, an e(rone(lros popular name of the hhaha) or nan:al.
Fl]. 2- ('rerm'ltiat rujete L., a liii harteristic tree of coastal pine ridges, which often
grows iln .ahbaa areas as isolated individuals. Pine ridge north of aviation field,
CARNEOIE INT. WASHIIIN(TlN 1'vii. 461 (D)-BIrtlcl Plm 2
I t;. 2
DESCRIPTION OF PLATE 3
Flu. 1- Edge of clearing at I'axactun. P. Ptn looking down into low flat th t was occupied
by escobal and botanal prior to clearing. Land rises at edge of the clearing
to climax forest. caoobal. The gigantic mahogany tree, Nwrietenlia marrophylla
King (note man at base) is locally known as caoba from whic-h name the
association gets its designation. At right ae two botan palms (Nabal morrisiana
Bartlett), almost as tall as the mahogany.
FIG. 2--Tle base of mahogany tree (croha) in climax forest casuall noear I'axactun,
Peten. At right of trunk is a ('lusia, one of species called chuinuip by the .Maya.
Below. its adventitious roots form larcuate stilts: above, they cling tightly to
trunk of the cooba, surrounding it in large numbers. increasing in girth. under-
going lateral fusion by spontaneous grafting. and ultimately forming a tight collar
or vest around doomed caoha. (See Plate 4, fig. 1.)
CA8m1E INST. WASIHINGTON PIB. 161 (PLA:ll ctI 1'r.TE 3
DESCRIPTION OF PLATE 4
Fia. 1-Saime tree as shown in Plate 3, figure 2. photographed with camera pointing up
almost vertically. Near top of picture adventitious roots of chunup are forming
collar around the cuoba.
Flu. 2-Base of a iuacut, Bernoullia flammen. Oliver. among ruins of Tikal, Peten. It is
easy to imagine that this species. with its flaming orange flowers. may have had
some esthetic or ceremonial significance to the ancient Maya that would account
for its persistence amid the ruins. with such obviously valuable species as ranmon
(Brosimum alicastrum) and chiro zapote (.Ahras sapota). Left. Mercedes
Chanek; right. Percy Gentle.
CARNEGIE INST. WA.Il~~11MN P(I. -161 (D11;01t t01 PLAri, 4
DESCRIPTION OF PLATE 5
FiG. I I P1i(ot(og glph ( fron1 olle of til p' iii, ds ) of tI oll of a MII III, 1., Iollia flam inn
n hiich iS loclllY ahMIndlalit at Tikal aijl char acterizes tile 110CHNIfl, a JiAlCse oft
,iu,,ionaiil ii ich the raprron is not dominant. In background is a tirrrple-crowne ur
py Iam Lid covered with forest (higueroal dei laxt iasrub ).
FIG. 2-1 ap a ruins convered I withI v-eget atiron. I(Term ple 1. C rounp .A. 1'a xa etinn. Peten.)
Larger trees oil bruins a re generally Ficus in prputhi lirum (Liebrn ) AIihq.
CA! ET;I 1%'T. P[~II,1~ B' iIu (I M~iI P
DESCRIPTION OF PLATE 6
FIG. 1-View in hotanal near Tikal. Peten. Guatemala. This association is characterized
by the botan palm. Sabal morrisiana Bartlett. of which a young stemless indi-
vidual is shown. Note that petioles of gigantic leaves are four tinms the height
of man at base of plant, i.r. about 7 meters tall. Blades of larger leaves have
spread of abolt 2 meters.
FiG. 2--Picture taken in same botanal at Tikal as figure 1. At a somewhat open place in
forest the camera was pointed up vertically in order to photograph silhouette of
crown of a boton, Ss'nba morrisinia Bartlett. Branched inflorescence exceeds
leaves in length.
CARNEGIE INST. WASHING'TON PIH. 461 (I)--B:rtlhtt
DESCRIPTION OF PLATE 7
FlI. 1-Enrobal at I iaxactlun. Peten. characterized by e srioa or "give-and-take," Cryo-
sophila argentca. Slender thorny trunk is obscured by petioles of collapsed dead
lower leaves. At right is Mercedes Chanek of El ('ayo. the writer's Maya
assistant. He was the principal informant (although there were others) with
regard to the local classification of plant associations in Peten and bordering
British lHondurnas. This paper may be looked upon as very largely a statement
in botanical terms of what this illiterate old Indian could tell about plant life
of his environment. Much of whatever success the writer's work miay have had
was the result of his kindliness and untutored wisdom.
Flo. 2-T'angle of clambering bamboo (A.rthrostylidiim piittieri Hack.) and climbing palm
(IrsInsmone n iiaactuinensis Bartlett) at edge of an escobal, characterized by the
sroba or "give-and take." 'ryosophila argentea Bartlett. Photograph taken
from natural opening provided by an arroyo lbhl.
CARNEGIE INST. XX S'IuNHM;'N PIuh. 461 (I) -l1VartioPL E
DaeCRIPTION OF PIATE 8
FIG. 1-View near Tikal. 1'eten showing, especially. climbing palm Desmoncus ferox
(actual type specimen. Bartlett 12584 in upper center), at border between the
pital (edge of aguada, characterized by Annaas magdalenew) and the escobal
(characterized by give-and-take palm. (ryosophila argentea). The give-and-take
(fan-like leaves in right foreground) gets its name from the dense investiture
of the trunk with spine-like branched roots. Whoever brushes against this palm
gives some tatters and takes some brittle. loose spines.
FIG. 2-Thi'ket of climbing palm. Deumonircus uaxaralunenisii Bartlett. photographed from
a natllal opening (bed of arroyo) in the esrobaol association. characterized by
the escoba or "give-and-take." Cryosophila argenica Bartlett. Uaxactun, Peten.