Citation
Studies on the vegetation of Puerto Rico

Material Information

Title:
Studies on the vegetation of Puerto Rico
Series Title:
Special Publication, No. 1
Creator:
Dansereau, Pierre
Affiliation:
University of Puerto Rico -- Mayaguez -- Faculty of Arts and Sciences -- Institute of Caribbean Science
Place of Publication:
Mayagueze, PR
Publisher:
University of Puerto Rico, Institue of Caribbean Science
Publication Date:
Language:
English

Subjects

Subjects / Keywords:
Caribbean ( LCSH )
Puerto Rico ( LCSH )
Farming ( LCSH )
Agriculture ( LCSH )
Farm life ( LCSH )
Forests ( jstor )
Vegetation ( jstor )
Shrubs ( jstor )
Spatial Coverage:
North America -- Puerto Rico -- Mayaguez
North America -- Puerto Rico -- Mayaguez -- Roosevelt Roads Area
Caribbean

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
The University of Florida George A. Smathers Libraries respect the intellectual property rights of others and do not claim any copyright interest in this item. This item may be protected by copyright but is made available here under a claim of fair use (17 U.S.C. §107) for non-profit research and educational purposes. Users of this work have responsibility for determining copyright status prior to reusing, publishing or reproducing this item for purposes other than what is allowed by fair use or other copyright exemptions. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder. The Smathers Libraries would like to learn more about this item and invite individuals or organizations to contact Digital Services (UFDC@uflib.ufl.edu) with any additional information they can provide.

Downloads

This item has the following downloads:

00006.txt

00265.txt

00199.txt

00206.txt

00026.txt

00047.txt

00080.txt

00288.txt

00058.txt

00105.txt

00060.txt

00092.txt

00282.txt

00233.txt

00280.txt

00051.txt

00269.txt

00177.txt

00231.txt

00263.txt

00252.txt

00055.txt

00061.txt

00153.txt

00162.txt

00137.txt

00205.txt

00253.txt

00183.txt

00067.txt

00142.txt

00181.txt

00237.txt

00037.txt

00290.txt

00262.txt

00033.txt

00215.txt

00100.txt

00224.txt

00291.txt

00096.txt

00145.txt

00108.txt

00174.txt

00062.txt

00002.txt

00112.txt

00146.txt

00243.txt

00076.txt

00057.txt

00293.txt

00148.txt

00182.txt

00158.txt

00087.txt

00066.txt

00186.txt

00073.txt

00075.txt

00267.txt

00279.txt

00194.txt

00007.txt

00127.txt

00235.txt

00027.txt

00063.txt

00270.txt

00114.txt

00221.txt

00091.txt

00071.txt

00120.txt

00059.txt

00223.txt

00136.txt

00259.txt

00284.txt

00150.txt

00042.txt

00012.txt

00201.txt

00156.txt

00125.txt

00023.txt

00167.txt

00039.txt

00218.txt

00122.txt

00258.txt

00163.txt

00255.txt

00256.txt

00133.txt

00210.txt

00072.txt

00081.txt

00020.txt

00274.txt

00038.txt

00268.txt

00213.txt

00250.txt

00188.txt

00179.txt

00193.txt

00151.txt

00101.txt

00011.txt

00238.txt

00277.txt

00190.txt

00285.txt

00160.txt

00034.txt

00010.txt

00083.txt

00157.txt

00143.txt

00024.txt

00110.txt

00093.txt

00117.txt

00247.txt

00234.txt

00152.txt

00184.txt

00022.txt

00204.txt

00119.txt

00189.txt

00168.txt

00111.txt

00154.txt

00248.txt

00207.txt

00019.txt

00289.txt

00203.txt

00251.txt

00126.txt

00135.txt

00283.txt

00172.txt

00191.txt

00170.txt

00220.txt

00246.txt

00169.txt

00070.txt

00032.txt

00138.txt

00068.txt

00241.txt

00107.txt

00217.txt

00128.txt

00140.txt

00212.txt

00064.txt

00008.txt

00035.txt

00095.txt

00200.txt

00264.txt

00271.txt

00090.txt

00196.txt

00016.txt

00222.txt

00116.txt

00118.txt

00005.txt

00103.txt

00208.txt

00166.txt

00197.txt

00017.txt

00139.txt

00178.txt

00097.txt

00050.txt

00121.txt

00085.txt

00195.txt

00018.txt

00227.txt

00098.txt

00209.txt

00113.txt

00052.txt

00144.txt

00084.txt

00069.txt

00245.txt

00134.txt

00239.txt

00004.txt

00088.txt

00187.txt

00240.txt

00292.txt

00286.txt

00287.txt

00029.txt

00257.txt

00175.txt

00226.txt

00272.txt

00074.txt

00254.txt

00249.txt

00132.txt

00077.txt

00219.txt

00041.txt

00236.txt

00164.txt

00198.txt

00229.txt

00104.txt

00185.txt

00115.txt

00078.txt

00149.txt

00141.txt

00131.txt

00021.txt

00028.txt

00216.txt

00275.txt

00031.txt

00009.txt

00230.txt

00276.txt

00281.txt

00046.txt

00278.txt

00266.txt

00147.txt

00044.txt

00013.txt

00228.txt

00001.txt

00109.txt

00225.txt

00099.txt

00102.txt

00180.txt

00040.txt

00129.txt

00094.txt

00159.txt

00014.txt

00086.txt

00242.txt

00232.txt

00130.txt

00049.txt

00079.txt

00048.txt

00165.txt

00211.txt

00123.txt

00065.txt

00261.txt

00106.txt

00214.txt

00015.txt

00056.txt

00192.txt

00045.txt

00161.txt

00171.txt

00176.txt

00173.txt

00202.txt

00030.txt

00244.txt

00089.txt

00082.txt

00155.txt

00273.txt

00036.txt

00124.txt

00260.txt

00043.txt

00025.txt

00003.txt


Full Text




UNIVERSITY OF PUERTO RICO

Focully of Arts and Sciences

INSTITUTE OF CARIBBEAN SCIENCE


0



0 0


0 o


0


STUDIES ON THE VEGETATION OF PUERTO RICO
I. DESCRIPTION AND INTEGRATION OF THE 0
PLANT-COMMUNITIES by Pierre Donsereu
II. ANALYSIS AND MAPPING OF;,THE ROOSEVELT 0
ROADS AREA by Peter F. Buell and Pierre Dansereou


IJ


SPECIAL PUBLICATION No.I

NOVEMBER 1966

MAYAGUEZ, PUERTO RICO

























STUDIES ON THE VEGETATION OF PUERTO RICO


I. Description and integration of the
plant communities.
by Pierre Dansereau


II. Analysis and mapping of the
Roads area.


Roosevelt


b Peter F. Buell and Pierre Dansereau










University of Puerto Rico Faculty of Arts and Sciences INSTITUTE OF CARIBBEAN SCIENCE


SPECIAL PUBLICATION No. 1 November 1966
Mayag'iez, Puerto Rico






DORADO
BEACH
/ SAN JUAN .
-PTA MALDONADO
-PTA VACIA TALEGA EYNevare,,unue
SHaystack P'oe
~Sierra-de

Rio de la Plato YunqueA L u q uillo
Rio d to latoMte Britton

a nde oan Cristobal Rio Blanco


I0 miles










STUDIES ON THE VEGETATION OF PUERTO RICO


Explanation of Contents





Map of Puerto Rico (with place names). . Frontispiece

Studies on the vegetation of Puerto Rico. I. Description and
integration of the plant-communities. By Pierre Dansereau. .3

Introduction . . . 3
Ia. Littoral Subzone. .12
I. Lowland Rainforest Zone. 20
II. Seasonal Evergreen Forest Zone.26 IIa. Hill Scrub Subzone. 30
III. Semi-deciduous Forest. 31
IV. Lower Montane Rainforest Zone. 35
V. Montane Forest Zone. 40
VI. Montane Scrub Zone. 44
Conclusion. .45

(Figure 1 presents four transects across Map 1 (zonal
division of the Island). Figures 2 to 7 are linedrawings indicating relative positions in typical
regional (zonal) landscapes of the vegetation-types
and/or plant-communities described in the text and
enumerated in the Tables.)

Studies on the vegetation of Puerto Rico. II. Analysis and
mapping of the Roosevelt Roads area. By Peter F. Buell and
Pierre Dansereau. 6

Literature Cited in Parts I and II. 56

Appendix A: Terminology. .63

Appendix B: Map errata. 64

Naster-TablesA to J. 65

(Keys to the units and symbols used in the Text,
Diagrams, Naps, Figures, Photos, and Tables I-XIV. )

Tables ItoVII. 8?

(Regional analyses of vegetation-types and their corresponding site-features: landformn, habitat,
ecosystem. )





2


Page


TableVIII. 100

(All of the species of vascular plants mentioned
in the Text and found in the Relev6s (Tables IX
to XIV) are graded according to the criteria in the
Master-Tables.)

TablesIX to XIV. 125

(Phytosociological charts which record the Releves
used in the construction of Diagrams.)

Photographs 1 toll8. 180

(In most instances the legend includes (in parentheses) the number of the plant-conmmnity or vegetation-type listed in Master-Table D. In some examples the exact quadrat where the phytosociological
Relev6 was made is shown (Tables IX-XIV), and the
number of the corresponding Diagram is given.)

Diagrams1to73.212

(Each one was plotted from a Relev6 bearing the
same number and which is given in full in Tables
IX to XIV. They are drawn according to Dansereau's (1958) method as modified by Dansereau, Buell, and Dagon (1966) and shown in Master-Table A. The name of the community, its corresponding number, and the
related photo references are listed in Master-Table D.)

Maps 2 to 7 (in envelope) show various features and interpretations
of the vegetation of the Roosevelt Roads area.




3


STUDIES ON THE VEGETATION OF PUERTO RICO

I. DESCRIPTION AND INTEGRTION OF

THE PLANT-CONIJITIES


Pierre Dansereau1/

The New York Botanical Garden




INTRODUCTIONS

The plant-cover of the Island of Puerto Rico is now entirely controlled by man. There is no such thing as virgin vegetation, whether forest, savana, scrub, or grassland. The various "forest reserves" (some of which are not really or mostly forested/), however, contain stands that have never been actually clear-cut and/or have been free of serious interference for a long enough time to convey to the contemporary observer some idea of their original composition and quite possibly of their primeval structure as well.

The present series of papers is offered in part as an experiment in description and interpretation. I had devised, in 195l, a method of recording vegetation which has proven to be applicable very widely and which I have been encouraged to develop and amplify. The Waterways Experiment Station (Vicksburg, Mississippi) of the United States Army Corps of Engineers had given me a first contract (DA-22-079-eng-208) in 1957-58 which resulted in the publication, in 1958, of a revised scheme. A second contract (DA-22079-eng-332) extending from 1962 to 1965 was focused on Puerto Rico. Meanwhile, a number of field teams, some of them directed by the Waterways Experiment Station itself, and some of them under contract, applied and variously modified my 1958 system. A meeting held in Vicksburg in April 1963 brought these various groups together and a summary of the proceedings was published (U. S. Array Engineer Waterways Experiment Station 1963).

In the years 1958-1965, on the other hand, I have given my best efforts to the development of a more broadly based system of functional analysis of vegetation. A preliminary application of this multidimensional characterization was published in 1961 and a fuller account will soon be
released.


Head of the Department of Ecology at The New York BtnclGarden,
and Adjunct Professor of Botany and of Geography at Columbia University, New York.
2 See definition of Forest in Master-Table C, page 67.










My 1l958 system (as shown in Master-Table A) is adhered to in the present study for the purpose of drawing the diagrams. However, in a separate paper', my associates and I have tried out a number of modifications
(Dansereau, Buell, & Dagon 1966).*

I am much indebted to the chief of the Area Evaluation Branch,
Dr. Warren Grabau, for innumerable suggestions and enlightening discussion at all stages of this work. kty assistant, Peter F. Buell, did much of the field work and all of the airphoto interpretation. He and Ronald Dagon proposed many valuable alterations. My permanent research assistant, Virginia A. Weadock) provided much help in the compiling of data and editing the manuscript, John Marks and Ronald Dagon drew the maps and the diagrams, and helped in other ways. Charles C. Glare drew the figures. Linda MKarschner assisted in the compilation and preparation of the tables. Bob
Benn, chief of the San Juan area field team of the Waterways Experilment Station,, and the crews under his direction, provided transportation and othe%--r facilities. They collected a large body of field data which, however., are not used in these first two contributions. Roy Woodbury, of the Puerto Rico Agricultural Experiment Station, accompanied me on most of my field expeditions and I could have achieved very little without his identification of species. Brother Alain Liogier also named a number of plants for us and revisited several of our sites. I am also grateful to Dr. Gustavo Candelas, chairman of the Biology Department at the University of Puerto Rico in Rio Piedras, who invited me to give a course on tropical plant-formations and ecology, and who provided office space. He and his colleagues offered many stimullating discussions of vegetation problems.

I must also acknowledge the support given me by the National Science Foundation (under Grant GB-2101), since the present memoir is a contribution to a long-range project on functional analysis of vegetation.

My attempt at reconstruction of natural vegetation is based on
extensive travel to all parts of the island. A sharper focus was obtained by making a number of precise relev's, which are transcribed in Tables IX to XIV, are analyzed in the text, and are summarized in diagrams (based on Master-Talble A). I was also able to make useful observations on the ecological amplitude of many of the indicator species, and feel that I can suggest correlations of physiographic and edaphic features with vegetation structures (see Figures 2 to 7). I have been influenced by Gleason and Cook's (1927) earlier study and by my own experience of analogous situations elsewhere (southern Florida, eastern Mexico, Hawaii, the Canaries and Azores, WJest Africa, coastal Brazil). My interpretation of the tendencies detectable in vegetation change rests upon the following assumptions: 1) Quantitatively fluctuating assemblages of species having a common migzra-











3) Man's interference very frequently results in the confrontation of
individual species (free or associated) with a historically unprecedented constellation of environmental features.

I have defined elsewhere and in much more detail (1956a, 1956b,
1957a, 1961a, 1962, 1965) a number of ecological laws, principles, and proeesaes which I believe applicable to the origin, composition, structure, and dynamics of vegetation everywhere.

The following descriptive account is given within the framework of six principal vegetation zones which are conditioned primarily by climate and secondarily by either physiography or parent-material. Within each one of them, different ecosystems are distinguished which present a quantitatively uniform combination of resources. It is within these ecosystems that vegetation units are plotted and defined by their composition and structure and related (at least tentatively) to each other in a dynamic scheme.

Such an analysis of the landscape proceeds by singling out areas
of decreasing magnitude. The largest unit is the zone, which is bioclimatic. Following a precedent set by Egler (1950, p. 229), I have used zone for geographic units and belt for ecological units (see Dansereau 1957a). Map 1 shows the relative location of the six geographic zones (I to VI) and the two subzones that are given recognition herewith. The lines are largely extrapolated from physiograph-ic and climatic data, not from actual observation of vegetation, as will be made amply evident below. A more detailed mapping on a larger scale of a few selected areas is offered in subsequent contributions in this series. Table I aligns the principal characteristics of each zone, upon which a few comments will be made. They seem to deserve recognition because they occupy sizable portions of space on the island and are readily definable as physiographic and climatic units. It need not be assumed, however, that they are biogeographically equivalent. In fact, the littoral subzone, the hill scrub subzone, and quite possibly the montane scrub zone owe their distinctness to widespread but clearly edaphic conditions. In other words, the climatic climax below 200 meters is lowland rainforest, whereas a "littoral complex" is determined by edaphic conditions; in the
Limestone Hills the seasonal-evergreen forest is climax and the hill scrub occupies emerging spurs. It is not equally clear, however, that the montane scrub is edaphic, and it may be well to conserve both the montane forest and the montane scrub as separate bioclimatic zones.

Figure 1 offers four transects across Map 1 that show the main
features of altitude and topography and the contacts between zones, as defined. Subsequent figures will illustrate (on a much larger scale) the more immediate relation of vegetation to topography.
In view of the generalized disturbance, and in view of the exclusive occurrence of some landforms in each of these eight areal units, it has seemed more useful to maintain the subzones separate in devising a framework for the present descriptive account.

Therefore, in the following chapters (numbered to fit the units of Map 1, Figurel, and Table I), I will attempt to encompass the principal eco-








D C B A O


C' B' 40 miles

50 kilometers


Ia. Litforal


I. Lowland rainforest Ij . Seasonal-evergreen
forest

Eab. Hill scrub


M. Semi-deciduous
forest

.ower monlane
rainforest

-7 . Montane forest


V1. Monfane scrub


MAP1
Vegefafion zones of Puerlo Rico, generalized and extrapolated from physiographic and climatic features (see Table I). Transecfs across the island correspond fo profiles in Figure 1.


B


D


C


O,






7


systems within each landform and to give a preliminary description (or at least an enumeration) of the principal communities of plants that have been encountered (Master-Table D).

The landform is the major physiographic unit (lagoon, reef, dune) which is definable in geomorphological terms, a combination of relief, geological features, and drainage pattern. It is sometimes sufficiently differentiated (when observed at closer range) to allow several habitats (or minor topographic units) to appear. The habitat (pond, shallows, flat), in turn, generally favors the establishment of a definite ecosystem, although it may allow more than one to emerge. The ecosystem is the conjunction of living and non-living elements which, on a given site, operate the conversion cycles of environmental resources. It 0ften comprises several belts (not zones ) each one of which is occupied by a distinct community. Communities, in turn, are definable in both structural and floristic terms and only incompletely so in the one or the other.3/ Moreover, they are usually
related in a dynamic fashion.

Each ecosystem (and therefore all of the communities which it
harbors) is characterized by a regime which is defined according to the kind of physiological stress that prevails. This has been expressed in a variety of ways and has led to a rather confused vocabulary. I shall adhere to a scheme originally formulated by Huguet del Villar (1929) before the emergence of the theory of the ecosystem, and which I have redefined (1952, 1956b, 1957a, 1961b, 1964). Master-Table B offers a key and an explanation of this classification and also some examples of typical corresponding ecosystems. A given regime may extend to several habitats, whether they are contiguous or not. The regime itself, therefore, is conditioned by a predominant physiological aptitude which is required of the living members of the ecosystem and which marks their ability to utilize its resources by meeting the particular stress that it offers (for instance, excessive acidity, cold, hardness or looseness of substratum, etc.).

Finally, the plant-community (whatever the method whereby it is defined) is a more or less cohesive group of species, mutually interacting and characterized by various features of composition, structure, and dynamics. There are often several plant-communities in a single ecosystem; some may occur in several ecosystems. In fact, the very object of phytosociological research greatly concerns the power shown by the community (or by communities that have compositional, structural, or dynamic features in common) to transgress the barriers of: ecosystem, habitat, landform, zone!

In the present account it is not always possible to refer to actual
plant associations (sensu Braun-Blanquet 1932 and/or sensu Dansereau 1957a, 1959), which are definable by a large sampling of stands. They would form the basic unit, of course, but such an inventory is not available and often the broader and less-defined vegetation-type will be described, in



3This has been quite thoroughly discussed elsewhere (Dansereau l957a, 1961a). It is of special importance in the present context where structure is going to be emphasized.










8


which one or more associations may be involved. These units are recorded in Master-Table D and in the various tables given for each zone. They are also
sketched in on Figures 2 to 7.

In order to insure the clarity and legibility of the present texts and of the rather considerable illustrative materials, a liminary Explanation of Contents and Appendix A lend themselves to ready consultation for crossreference and definition of terms.

Regional patterns of vegetation, or vegetation zones, are determined primarily by climatic conditions, which in turn are due to the various interferences of airmasses and physiography. Lobeck's map and study (1922) and subsequent interpretations by Pic6 (1954, 1962, 1963) and Young (1955) provide us with an adequate description of the relief and of the landforms. Likewise, Roberts (1942), Thorp (1941), and Pic6 (1954, 1962) have mapped
and summarized the climatic features. Climatic data and diagrams will not be presented in this report.

The interactions of airmasses (climatic regimes), of physiography, and Tertiary migrations of flora have been studied by Beard (1944) who has presented a unified scheme for tropical America. Asprey and Robbins (1953) have applied this to Jamaica. It can be said that Puerto Rico repeats the Jamaican patterns (see their Figures 38 and 40) in a somewhat depauperate form. On the other hand, Stehl6 (1945, 1946) has emphasized the Lesser Antilles and made many useful edaphic and climatic correlations which it will be worth testing further by means of functional analyses such as are provided by the data offered in the present contribution.

Map 1 gives a much simplified account of the six zones and two subzones of vegetation whose principal characteristics are listed in Table I. Extrapolated from physiographic and climatic data, the zones correspond, on the whole, fairly well with most of Pic6's (1962) natural areas, and with Thornthwaite's climatic types (Thorp 1941). According to this latter definition, there is almost no arid land and not very much semi-arid either. The greater part of the driest zone is termed subhumid.

Earlier attempts at mapping of vegetation (Murphy 1916, Pic6 1954) provide very little help, for they are even sketchier than the present one. I am afraid I cannot make much, either of Kumme and Briscoe's (1963) application of the Holdridge system to the "forests of Puerto Rico."4/ On the other hand, Little and Wadsworth's (1964) map is based on the same concepts that have guided me here, and it will be seen that many of our boundaries between zones are nearly coincident. The "Common trees of Puerto Rico" is based on
many years of field experience and incorporates a full knowledge of forest types as revealed by foresters' studies, a few of which have been published in the "Caribbean Forester."

The present contribution is a survey and establishes the framework within which more detailed studies can be integrated. The following figures, photographs, and tables will set the stage. The vegetation of each zone will

4I should have much to say in both praise and criticism of Holdridge's definitions and mapping. Sawyer and Lindsey's (1964) recent application of these norms to Eastern North America certainly point to a lack of equivalence


between bioclimatic zones and vegetation zones. It is reassuring that Holdridge (1965) now calls his areas "life-zones."






A


o1000 TERS

750 500 250

0
U o


1000 750 500

250

0


1000 ME TERS 750 boo 250

0


50 KILOMETERS


FIGUR i1. Four transects across the Island as plotted on NAP)1 (which see for explanation of vegetation symbols


NO


0.


A 1000 METERS




6 00 -250



|1 0


B


C A C


D










therefore be described in the order in which it appears in Table I. The description of the vegetation-types and/or plant-communities will follow the order indicated in the table which lists the site-charanteristics.

The landscape of each zone will be represented, in a figure, by several transects that show the usual contiguities of communities. The numbers used are those listed in Master-Table D which carries references to photos and structure-diagrams. Similarly, each one of the regional (or zonal) tables (II to VII) will list all of the communities (by their names and numbers) to be found in the landscape under specified landform, regime (as in Plaster-Table B), habitat, and local ecosystem.

The scientific names of plants, with very few exceptions, are
those of Britton and Wilson (1923-30). It has not been possible to incorporate all of the numerous changes recently proposed by Liogier (1965).

Table VIII lists all the species of Puerto Rico that are mentioned in the text and tables, and it grades them according to the various criteria which serve the purpose of functional analysis and especially of structural representation.

The following descriptions, in most instances, will not be very detailed so far as floristic notations are concerned. However, since our field work consisted principally of phytosociological releves made according to the method defined in my "Biogeographyl an ecological perspective" (1957a) and exemplified in the Appendix of that book, these data have been assembled
in Tables IX to XIV. They have served as the base for the diagrams that bear corresponding numbers. They have been grouped under the following floristic complexes:
A. Mangrove complex (Table IX): principally Rhizophora, Avicennia, and
Laguncularia, but also many saltmarsh species. These are tropical
Atlantic species, very widely distributed.
B. Tropical beach-and-strand complex (Table X). Most of the species in
this group are pan-tropical, whether truly indigenous throughout the Tropics of Pacific and Indian as well as Atlantic shores, or secondarily distributed. Such are Ipomoea pes-caprae, Cocos nucifera.
C. Lowland marsh complex (Table XI). This contains a majority of pantropical species.
D. Hygrophytic complex (Table XII). Many of these are strictly American
tropical plants, some of them endemic to the Caribbean, to the Antilles, or even to Puerto Rico. The species range from the shade-loving, highrequirement species of the original rainforest to the ecologically unstable second-growth and jungle species.
B. Subxerophytic complex (Table XIII). The woodlands, savanas, and scrubs
of the Caribbean have possibly developed an even more characteristic
flora than the rainforest: Bursera simaruba, Plumiera alba are good examples. There is rather a broad range, in fact, from the seasonal evergreens characteristic of Zone II to the semi-deciduous plants of
Zone III, all of which are part of the same floristic group.






11


F. Montane complex (Table XIV). The isolated Caribbean highlands harbor
a flora of their own, ranging from genera of tropical affinity (Eugenia, Tabebuia) to widespread temperate-rainforest types (Ilex, Rapanea, Podocarpus, and possibly Euterpe), but no boreal ones.!
G. Tropical exotic complex. This comprises a number of invaders from other
lands that show various degrees of naturalization, from the planted Eucalyptus globulus (from Tasmania) to the subspontaneous Spathodea campanulata (from Central and South Africa) and the forest-invading
Erythrina poeppigiana (from Peru).

Some further discussion of these "complexes" will be given in the course of the description of communities. But it has been found useful to group the releves together under the above headings, whereas the individual species have all been graded (in Table VIII) according to floristic element (Master-Table H).
Throughout the following account, therefore, cross-references will constantly be made to the following: community: as listed in Master-Table D where essential facts appear concerning composition and structure (formation-type as defined in MasterTable C);
floristic composition (in some instances) is given for one or more stands
in Tables IX to XIV; and these field data, in turn, have served to
determine the
structure, as recorded in the diagrams (constructed on the basis of MasterTable A);
photos of most of the communities (or vegetation-types) show their particular position in the landscape which, for each zone, is schematized
in the
figures (2 to 7) that present typical profiles in each one of the zones that
appear on
Map 1 (and on Figure 1); each community bears the number given to it in
Master-Table D.







12


Ia. LITTORAL SUBZONE

Map 1; Figures 1, 2, 5; Tables I, II, IX, X;

Photos 1-h9.


This area, which lies near the sea and is variously prolonged
inland in the flatter areas, is under the influence of saltwater, saltspray, and sea-winds. It comprises the climatically driest portion of the Island (near Guanica, in the southwest) but also extends to very rainy territory. Not too many littoral communities seem to be affected by this difference in climate, although some variants are to be observed and also a difference in rate of development and replacement. Therefore, it is in contact, on its upper limits (see Fig. 1), mostly with Zone I, but also with Zone III, and occasionally with Zone II (see Photo 81). Figure 2 and Table II show the following major divisions: open sea, lagoon, reef, intertidal and supratidal belts, dunes, and estuaries. These are in turn under the impact of six ecosystematic regimes (seven if one considers man's influence). I propose to use these as the framework within which to describe the plant-communities.

Saltwater ecosystems (halohydrophytia)
These consist of the open sea (Atlantic or Caribbean), of partly submerged coral reefs, either isolated or forming fringes that enclose shallows, and of lagoons (embraced or closed by strands). The vegetation of
the sea, of the reef talus (or scarp), and of the permanently submerged part of the lagoon will not be considered here: it consists almost exclusively of algae. Its communities can rarely be accounted for without considerable attention to the animal components, especially on rocky shores (see Dansereau 1947a).

I have no knowledge at all of this marine vegetation5/ and will
make no attempt to include it in the present account, except to indicate its points of contact with palustrine formations. The kelp tangle and the sealettuce meadow do not have such contact. But the turtle-grass meadows that occupy the shallows do, The somewhat succulent linear-leaved, grasslike Thalassia testudinum grows in dense stands especially where a layer of sand covers the underlying rock. These plants have vigorous vegetative propagation. They are periodically uprooted by the sea currents or exceptionally high tides and deposited in great masses on the beach, on an otherwise sterile strip. Here they can offer conditions of germination and ephemeral growth to such halophytes as Cakile lanceolata which, however, does not become established. Moreover, these matted debris dry up and are blown
inward to the upper beach where they increase the organic content of the soil. This increment of organic (and mineral) elements can be of considerable importance and it would seem to operate in a variety of conditions (e.g., in the Arctic: see Dansereau 1954).



5This is being studied excellently by Diaz Piferrer and his collaborators.




ZONE La
SEA INTERTIDAL BELT SUPRATIDAL BELT UPLAND 24

REEF (ROC KY) (ROC KY)
LAGOON 14.7 159 137 16.4
6.4 i

~ ~ ~ ~ ~~ 0 .6 V v W V V, ,vvv vv v V V V vvv v v V v v
.5 4.0 2. 46
' I " i , v, v v, v v v v vV V v v V / v V, v v v v .1 , v v v ' v 4 v v v
v v
IlrvvVV V,~
vV VV v v'V

halo- I halo-- halo- heiohalophytia I petrohalophytia I psammohalophytia paranth
IIII
hydro-ophyia hydro- V phy
phytia phti
SHOAL (SANDY) (SANDY) DUNE UPLAND
20.4 43.1 e (
19.6 21.9 1818.7 20.4
18.7 17.6,\I : o 22.4 26.4_haiohydrophytia helohalophytia , psammohalophytia meso- ,h
,ydr h t axero- ,
' f phytia '
RE EF (SILTY) (SILTY) ESTUARY



5.1 6.1__ 7._ 6.1 214 34.5"53 3. 65
4.0 "7.4 4.0 3.6 9.6 1 2.5 7.4 12.5 2 37.4 3. 3. 3.5 31




,-~- . -- 0. . . . r o
halohydrophytia ,helohalophytia , helophy ia - limno-- -h eophyti phy"ia.h0 'p


corresponding to MASTE00-TABLE D and to TABLE TI. (See also FIGURE 6.)


ropo



12.1









ygrohytia




39.1 hy a



IV
V
-- _ v- _







14


Tidal saltwater ecosystems (helohalophytia)

I cannot offer a full account of the complex zonation of the tidal area, at this time, but will outline the principal formations' response to diverse substrata.

Whereas the sandy beaches are quite sterile in the intertidal zone, the rocky shores, especially where the surf beats strongly, are covered, at that level, with Fucus and especially with Turbinaria turbinata. These fucoid beds occupy the lower part of the intertidal belt (Photos 1 and 2), being exposed only a few hours, whereas the algal crust dominated by Enterom ha more often thrives on surf-spattered rocks higher up. Rawitscher (19447 described an analogous ecosystem in southeastern Brazil.

The reef itself, where it barely emerges above the high-tide level, often harbors bits and pieces of mangrove. These are isolated individuals of
Rhizophora mangle, well anchored in the brittle crevices of the crumbling coral. Where the reef platform is somewhat higher and inundated only by storm tides, very large, old Avicennia can be found (e.g., on La Gata, Photo 15). Colonies of Rhizophora are known to pioneer also on a sandy shore very much exposed to wind action (Photo 5).

The mangrove complex occupies various positions in the tropical
world (see map and bibliography in West 1956). In Indonesia, the Philippines, and Colombia it reaches great spatial development. Even more interesting, however, in a longer perspective, is the great floristic richness of the Indo-Pacific area to which Cuatrecasas (1958b) ascribed its origin. This same hypothesis was revived by van Steenis (1962), who listed 17 genera and 43 species in Indonesia, whereas Atlantic America has only 5 genera and 10 species, and Atlantic Africa, the same 5 genera with only 7 species. This migration radiation suggests to me an application of E. L. Braun's (1935) association-segregate mechanism. Both the lag in migration of genera and
species well suited to the mangrove and their relative inability to cross certain climatic barriers are indicated.

In Puerto Rico mangrove covers a good deal of territory on fine sand and silt and forms a broken ring around the island. It exhibits a number of patterns that are determined by the following features: 1) Rhizophora mangle, Avicennia nitida,/ Laguncularia racemosa, Conocarpus
erecta can withstand flooding by saltwater, in the order mentioned.
Therefore, here as elsewhere (in Brazil, for instance: see Dansereau 1947a), it can be expected that Rhizophora will be nearest the sea and Laguncularia
and Conocarpus farthest inland.
2) Avicennia is more tolerant of stagnant water than either Rhizophora or
Laguncularia. This partly explains the position of the latter two at the
opposite ends of the soil-moisture gradient (where water movement is
greater) and the exclusive presence of Avicennia over large tracts of the
flatte st land.


6 This species should now be called A. germinans (L.) L., according to Compare (1963) and Liogier (196 ).












3) There are many variations of structure in the mangrove as shown in
Diagrams 1 to 12: height to 20 meters (Photos 6 and 7), and canopy
coverage of 85 per cent, to height of 8 meters and canopy coverage of 50 per cent (Photo 9). It does not involve any other species but the four trees (and usually not Conocarpus at that). Coconuts are regularly washed in and are able to germinate in certain stands, but cannot
make much growth, even less maintain themselves. Where the mangrove
has been exploited (especially Avicennia), it tends to branch very low,
and many shrubby stands can be observed (Photo 11). (See Wadsworth 1959.)
4) In some areas, the back-waters favor a megaphorbia of Acrostichum aureum
rather than a forest, and in other areas, on the contrary, the occasionally flooded higher ground tends to concentrate salt, by evaporation,
and allows the development of herbaceous flats (with Sesuviura portulacastrum mats (Photo 19) and/or low Batis maritima scr). Many of these
lands are reinvaded by Avicennia and a savana may prevail (Photo 17).
5) Nowhere have I seen a scrub or forest of Hibiscus tiliaceus or Thespesia
populnea, although both of these species are present in Puerto Rico.

Some mangroves lie behind a strand. TWhere the vegetation of the
latter is dense (a coconut or an Australian pine screen (Photo 46) or a wellconsolidated sea-grape scrub), very little windblown sand is deposited on the emerging pnem'atophores of the black-mangrove or the white-mangrove. But where the levee is very open, the sand drifts in. It does so rather spectacularly where the levees are exploited by man for building purposes. If the sand ridge is low, in fact, the high tides will wash in a sheet of beach sand, eventually a foot or more in depth and the mangrove trees will be killed. This has been observed on a large scale at Punta Maldonado.

The literature on mangroves is extremely abundant, and the physiology of mangrove plants as well as their ecology have received a good deal of attention. Walter (1962) has brought these data together, on a world basis, and re-focused it in a very comprehensive way. In Puerto Rico itself, Holdridge estimated in 1940 some 16,000 acres and reported on changes induced by man. From the silvicultural point of view, Wadsworth (1959) has compiled the growth rates of white and black mangroves under various treatments. Golley, Odum, and Wilson (1962) have produced a pioneering study of the metabolism of a red mangrove forest.

Supratidal belts (halophytia)

This area is under the influence of salt spray and in unprotected exposures it can extend inland. This is much in evidence on the hard limestone slabs of the Guanica coast where salt-wind shearing affects the
scrubby vegetation many hundreds of yards from the shoreline.
Rocky shore (petrohalophytia)

Along this gradient, the occasionally surf-splashed rock has smaUl pockets of sand, some of it formed in situ by the weathering of the rock (San Juan sandstone in the North, Ponce limestone in the South) or blown in from neighboring beaches. Here, individual tufts of Euphorbia buxifolia and






16


Fimbristylis spadicea (Photos 2h and 25) are to be seen, as well as seedlings of Suriana maritima, Borrichia arborescens (Photo 22), Gundlachia corymbosa, in the form of a steppe or desert. Where more sand is available, Spartina patens is also found (Photo 38). The community that attains the highest development here, however, is a tight, low meadow of Fimbristylis spadicea (Photo 23).

Higher up, on the hard rock, firmly anchored in the fissures, a low, often succulent scrub develops, dominated by Gundlachia, Borrichia, or Suriana (Photo 21), almost always accompanied by Conocarpus erecta. The latter here is matted so densely as to exclude invaders (Photos 23 and 28). It is apparently very long-lived (Photo 29).

On the south shore, above this often disturbed area, subject to equinoctial and storm tides, the jointed and pitted slabs of limestone harbor a desert-like spiny and succulent vegetation, beginning with such low plants as Cactus intortus (Photo 26), and then the taller Opuntia rubescens, Cephalocereus royeni (Photos 27 and 30), Lemaireocereus hystrix (Photo 31), and Anthacanthus spinosus (Photo 32). The shearing effect becomes more visible where the scrub has somewhat consolidated, especially where the button-mangrove (Conocarpus erecta) is still involved. This plant, however, gives way to various xerophytic shrubs, mostly to the sea-grape. Photo 23 illustrates the salt-sedge to Conocarpus to Coccoloba gradation.

At the same level on the north shore, there is most likely to be a strong overlapping of sand, and the beach communities take over.

Sandy shore (psammophytia)

This sector comprises a beach and a levee habitat. The outer beach is sterile, as a rule, under the combined pacts of equinoctial and other very high tides, storms, and winds (Photos 33, 3h, hO, and h6). The inner beach commonly exhibits two belts. Nearest the ocean, the pioneering streamers of Sporobolus virginicus, Paspalum vaginatum, or Spartina patens extend precariously, sometimes accormnipanied by tufts of the annual Cakile lanceolata (Photos 3h and 46), or else the latter alone is present. The less often disturbed upper part also frequently receives and holds vast quantities of driftwood (Photo 39). Here the three grasses and the crucifer mentioned above may form a lush meadow or prairie which is reinforced by two vines, Ipomoea pes-caprae (Photo 34) and Canavalia maritima. A further step is the establishment of the succulent-leaved shrubs, Scaevola plumieri, Suriana maritima, Borrichia arborescens (Photos 35 and 36), and an eventual invasion by sea-grape (Photos 39 and hO).
The levee is almost everywhere planted to coconuts (Cocos nucifera) and also to Australian pines (Casuarina equisetifolia) (Photos "adh). But the dominant of the primeval scrub, Coccoloba uvifera, is rarely absent. This extremely aggressive plant is most obviously adapted to a variety of coastal habitats, ranging from the hard surfaces of sparsely-fissured schist or limestone (Photo 23) to the loose sand of dunes (Photo hO) where it responds to burying and to uncovering alike. It also takes the pruning of windshear (Photos 39 and hl). As a result, it may branch low or high, and








17


the size and even the texture of its leaves will vary. On the typical levee, it is low-branching and forms a closed canopy; it grows alone or is overwhelmingly more abundant than any other woody species. Table X has nine releves taken in this community (Releves and Diagrams 29 to 37). The floristic and structural variations of the prevalent type of coastal scrub show more persistence of beach plants gradually shaded out and invasion of more sciophilous species of the snake-bark scrub as the total mass and stratification increase. Inland, it is frequently replaced by maray-maray
(Dalbergia ecastophyllum) scrub.

In the lee of the levee, or in otherwise protected spots, a much taller, structurally complex, and floristically rich xerophytic scrub develops. This is Beard's (1944) "littoral woodland." Snake-bark (Colubrina ferruginosa) is most conspicuous. Many lianas are present and there can be a fair development of lower strata too. For instance, some stands have a great deal of Bromelia pinguin. Four releves and diagrams (39 to 42) represent this association. The formation of this thicket opens the way to an invasion by the members of what I have called above the subxerophytic floristic complex (Table XIII) which plays a major role in the consolidation of vegetation in the semi-deciduous forest (Zone III) and the seasonal-evergreen forest and scrub (Zone II).

The dune ecosystem (psammophytia)

Wind rather than salt is the influential factor on the dunes, which, however, do not occur inland in Puerto Rico. They largely share the flora of the levee (there are some high levees and some low dunes.) and some of the communities are quite identical if somewhat less stable. Thus the sharp, shoreward, steep, eroding face of the dune (top-heavy sand flowing down, wavecutting undermining) commonly has nothing but hanging shreds of one or more of the three beach grasses (mostly Spartina as in Photo 37) and vines (mostly Ipomoea), whereas it is capped by a tenacious Coccoloba scrub.

The interdunal areas (which are not very highly developed) will harbor a snake-bark scrub or sometimes a Malabar-almond tall scrub which looks spontaneous even though Terminalia catappa appears to have been introduced. Beach-grass (Sporobolus virginicus) forms low, irregular stands on both sides of the dune and can be very lush in the intervale.

The influence of man has deeply modified the littoral region,. The mangrove has been drained in many places and reclaimed for sugarcane (background of Photo 47); the strands are nearly all planted to coconut, under which are occasionally grown sugarcane (Mayaguez area) or pineapple (Jobos area)(Photo 45). Moreover, the ablation on beaches, levees, and dunes of
sand for construction purposes has caused mangroves to be covered by several feet of sand and thereby destroyed (as has been mentioned above).

Estur (limnophytia and helophytia)

Several rivers discharge into lagoons, mangroves, or breaks in the strand. Some of them are not too badly polluted, in that ultimate sector, to harbor several aquatic (limnophytic) and marsh (helophytic) communities that display characteristic zonation (Photos h7 and 48).







18


Floating mats, sometimes precariously secured to a shallow bottom or a protruding shore, respond to slightly different conditions: the waterhyacinth (Eichhornia crassipes, Photo h9) is very mobile, spreads rapidly, and is not carried away by running water so readily as the water-lettuce (Pistia stratiotes) would be. In fact, the latter does better in backwaters and in ponds than in rivers. The para-carib-grass mat (Photo h8)with its rhizomatous grasses (Panicum purpurascens, Eriochloa polystachya) has to grow outward from a bank, and waterlily pads are anchored in an organic bottom ooze in slow-flowing water.

It is a question of how brackish the substratum of the estuarine and other coastal marshes can be. Solid stands of cattail (Typha domingensis) and of reed-grass (Phragmites communis) here as elsewhere can certainly endure a small amount of salt. I know very little in this respect concerning the spikerushes (Eleocharis mutata, E. interstincta) of Puerto Rico. The marshes dominated by them, which are analyzed in Table XI and Diagrams h and h6, are (or were) well developed, forming rather a firm substratum, and tending to evolve to a fen structure by the invasion of Chrysobalanus icaco (and sometimes Drepanocarpus lunatus), associated with sedges (Cladium, Scirpus, Carex) and ferns (Blechnum occidentale). (The sawgrass, Cladium jamaicense, so important in the Florida Everglades, does not seem to dominate to any extent here.) The icaco scrub can be very dense and extensive. It is interrupted here and there by emergent pond-apples (Annona glabra) and button-mangrove (Conocarpus erecta). In fact, the latter form a swamp-woodland of rather irregular structure, riddled with pools of different depths, harboring true aquatics (such as Pistia, Nymphaea, Utricularia) and large sedge tussocks.

Three other helophytic communities seem to lie beyond brackish influence. The riparian trompetilla-grass marsh dominated by Panicum aquaticul and Hymenachne amplexicaulis is very similar in structure to the para-caribgrass mat, but is probably more firmly rooted in the mud, below constantly flowing although fluctuating water.

The rivercane brake is a gigantic grassy screen perched on the upper edge of the floodplain: Gynerium sagittatum (a relative of the pampas-grass)
grows in very dense, canelike formations and marks off the edge of sugarcane plantations, often alternating with planted bamboo (Photo 68).

The Pterocarpus officinalis swamp forest must once have occupied the upper plain of many of the estuaries, at least on the N, E, and SE of Puerto
Rico. Mere may be no more than two stands left today. The one at Dorado is probably the best example of its kind, with its streaming and twisting buttresses. This association, a well-developed forest of high dimensions, typically lies behind the mangrove on salt-free soil. Stehl6 (1945, Fig. 7) offers a good profile-diagram that also shows contact with the Acrostichum and Drepanocarpus communities. It was very well illustrated by Gleason and Cook (1927, Plate 17), whose study are yielded no living stand of Pterocarpus in 1963. It is interesting to compare the structure of this swamp-forest with one in Tahiti (Papy 195%) where the dominant Inocarpus edulis is physiognomically identical.

The littoral subzone is, of course, not distinct, climatically,
from the lowland rainforest zone. It is the influence of the sea that serves to define it: salt in air, water, and soil; offshore winds; tidal effect







19


(salt- and freshwater). In fact, some of its plant-communities (especially the marshes) extend considerably inland. Many of the upland littoral vegetation units also show a contact with one or more of the lowland rainforest communities. Thus Table X reveals the presence in coastal scrubs of some of the species of both the subxerophytic complex (Table XIII) and the hygrophytic complex (Table XII). This may well indicate a successional sequence as already suggested by Gleason and Cook (1927).

Transformation by man of the littoral subzone (as here defined) consists of:
1) sugarcane plantations on mangrove or marsh sites (Photos 67 to 69); 2) pineapple plantations on rich upland (Photos 63 to 65); 3) exploitation of sand and rock which are removed from strands and puntas; 4) port and marina installations, on various topographies;
5) salinas on mangrove or saltmarsh s ites; 6) pasturing and horticulture;
7) stripping of riparian zonation (Photo 51); 8) construction of wharves, buildings, roads, airports, gardens, cities.







20


I. LWL.kND RAINFOREST ZONE

Map 1; Figures 1 and 3; Tables I, III, XI;

Photos 51, 62 to 69


Even at the time (1926) of Gleason and Cook's investigation, extremely little was left of the primeval lowland rainforest. Nowadays such stands have been depleted even more drastically. The guesses which can be made concerning the composition and structure of the original plant-cover of the well-drained uplands are based on ecological laws and principles (which are themselves none too securely established: see Dansereau 1962,
196-5) and on analogues in other areas.

The geological nature of the substratum, the topography, relief and natural drainage patterns, the known levels and fluctuations of temperature and amounts of rainfall all point to the existence, at an earlier period, of some kind of tropical rainforest. The structure of this vegetation is most likelyV to have conformed to a widespread pattern of stratification such as has
been recorded in climatically similar places elsewhere and as summarized by Richards (1952). It would have consisted-of a number-of very stout and tall trees, unbranched in the lower two-thirds, some of them possibly overtopping the general canopy. Their shade would preclude the development of intermediate layers of any great density; lianas would reach high into the canopy and remain fairly close to the larger trees (not forming tangles in the lower strata). Epiphytes would be abundant-on the higher branches of trees, less so lower down. There would be no palms or tree-ferns. The very thin ground layer would consist primarily of woody plants,-many of-them-seedlings of the tree species.

Such a classicallyl structure is hardly to be observed anywhere in Puerto Rico at the present time.* At least, I have not seen any stand that nearly approximates the above description. The fairly mature stands I have seen-(at-Dorado and on the lower slopes of the-Luquillo Mountains) have-very few of the really big trees and the density of their lower layers suggests a state of unbalance or progress or "healing" which places the stand very much on this side of climax structure.

The clues that are to be gleaned from composition provide further insight. There is no atlas of the distribution of species in Puerto Rico. But Britton and Wilsonts (1923-30) flora, and additional unpublished herbarium and sight records made by Roy Woodbury and Alain Liogier, might allow us to line up a number of species that are never found (in Puerto Rico) above
aproiatl O eer.Mn.y of these, of course, be1-'lon n to.he littoral







21


quite show the zonal profile). Such are Manilkara bidentata, Lucuma multiflora, Diospyros ebenaster, Stenostemum obtusifolium, Ixora ferrea, Cassipourea elliptica, Faramea occidentalis, Hernandia sonora, Petitia domingensis. It is, of course, not excluded that some species (such as Zanthoxylum martinicense and Bucida buceras) which have a broader range of ecological and/or climatic valences (including shade tolerance) could have been authentic members of some "mixed mesophytic" complex.

This notion of a floristic cohort of species that have very closely overlapping ecological ranges, first formulated by E. Lucy Braun in 1935, was later elaborated by her (1950). I made an initial application of it to tropical vegetation in Brazil (1947b) and later attempted to re-formulate it (1956a, 1965) and to extend its application to the origin and growth of plantcommunities in general (1961a). I am convinced that this approach, if critically followed, will yield the most valuable clues. Our improving knowledge (Berlioz 1965) of geographic relationships in the Antilles will soon allow us to discern the affinities of what I have called (above) the hygrophytic complex from those of the subxerophytic complex. Aubreville (19h9) and others have gone to some length in exploring a similar relationship in Africa.

Our criteria for delimiting this geographical zone at present are therefore largely floristic. Its boundaries will coincide with the upper limits of a certain number of species (mostly the trees mentioned above); with the lower limits of others (for instance Dacryodes excelsa), and this is roughly 350 meters. To be sure, ravines, crests, scarps, nature of parentmaterial make such a line waver upwards and downwards.

At all events, such a zonal delimitation based upon the presumed
extension (on average upland topography) of a climatic climax association has more theoretical than practical value and surely no descriptive value at all, at this time, in Puerto Rico. Other forms of vegetation now occupy virtually all of the surface and must be considered in some detail. Figure 2 and Table III give us our bearings in this respect. Table XII, on the other hand, refers exclusively to Zone IV, the lower montane.

Lake and river ecosystems (limnophytia)

Rivers and lakes prolong the estuarine conditions described above. They have floating mats of Eichhornia crassipes which are sometimes dense enough to seal in the water surface altogether. In ponds or very still water the duckweed (Lemna perpusilla) performs this function; also, the water-lettuce (Pistia stratiotes).7Panicum purpurascens often comes next in the shoreward zonation, forming a mat which is attached, however, to the
bank by a rooted hinge. This grass grows to a height of about 60 centimeters. Behind this, marsh conditions prevail, rather than truly aquatic ones.

Riparian ecosystems (helophytia)

A marsh ecosystem of varying complexity is determined by the
drainage pattern of the stream or lake border. Cattail communities (Typh~a domingensis, dominant) alternate with the taller reeds (Phragxnites communis).






LAKE limnophytia


FLOOD-PLAIN '
helophytia 33.5 I


54.5


CULTIVATED LAND
paranfhropophyfia


ZONE I


v3Jj JiYI OC(
PASTURE
paranthropophyti-a
49.6 52.6:I
51.6 , , .3L , 67.5


The principal structural variations of vegetation in the Lowland Rainforest Zone (I) at present and


FIGURE 3.







23


They commonly consist of pure populations of these species, with only a scattering of other plants, such as Eleocharis spp., Cladium *amaicense, Sagittaria lancifolia. On rather steep banks, the river-cane (yer agittatum) may rise as high as 15 meters. At present it is usually seen to occupy rather a narrow strip. However, it is evident that with only a slight improvement in drainage the rich muck of the lowlands has been converted to the cultivation of sugarcane.

The torrential streams, especially in the southwest, have periodically emersed river-gravels and flats (Photo 51) that mostly carry ephemeral vegetation. The spider-flower (Cleome spinosa), an annual, is often the most conspicuous member of such a community, where it grows in clumps.

Upland vegetation (hygrophytia)

Spontaneous and subspontaneous vegetation occupy a relatively small portion of the well-drained lands below 350 meters. Figure 3 shows the principal communities that are to be found on former tropical rainforest sites. I have already mentioned the difficulty of describing the lowland rainforest and even the hazards of identifying its principal floristic components, in the absence of extensive and well-conserved relicts. Lists of species given by Stehl& (1945, 1946), Beard (1946, 1949), Asprey and Robbins (1953) elsewhere in the Caribbean indicate the persistence of many second-growth species in otherwise well-structured stands. It follows that it is also difficult to distinguish it from the more mature rainforests with which it is in contact, namely the seasonal-evergreen and the lowland montane, for both of which I have more data (see Table XII and Relev's 50-53 in Table XIII). The more mature second-growth has also been recorded better in Zones II and IV (which see). Virtually all forest stands in the lowlands are second-growth, where the presence of such species as trumpet-wood (Cecropia peltata), Didymopanax morototoni, Casearia spp., Andira inermis, testify to an opening-up of the canopy without which these species are not likely to have germinated and to have gone through their early growth stages. Some of the stands, of course, may never have been clear-cut and they testify to the "healing" of a disturbed rainforest. There are certainly many types of forest in this category, resulting from various kinds of interference, all the way from the natural, and more or less cyclical, effect of hurricanes to selective cutting and planned silvicultural practices.

Jungle (mesoxerophytia)

Jungle, in the proper sense of the term, is the result of either degradation or recovery, probably more often of the former than the latter. It is vegetation of difficult access_7/ because of its encumbrances in the lower layers. This is mostly caused by woody and/or herbaceous vines. Much jungle is of shrub height or consists of scattered trees overtopping tangled shrubs and themselves draped in vines, forming a savana or eventually a wood7It is quite remarkable that the etymology and folklore of vegetation
terms repeatedly lead to this anthropocentric and peregrine origin of the meaning of words. Tundra, for instance, means hostile" land, as does also desert.







24


land. Very rarely, if a true forest structure develops, does the lianescent element retain much prominence.

There are not too many examples of this vegetation type in the
Puertorican lowlands: for the most part, either the forest is exploited as such or is cleared for agricultural purposes. The most conspicuous jungles lie at the foot of the limestone hills, in the north, in valleys and ravines on shallow soils. (These areas will be described below: see Zone II.)


Controlledvegetation (paranthropophytia)

It seems useful to distinguish three levels of human interference:
1) managed land, which involves old fields in various stages of subspontaneous or spontaneous revegetation on the one hand and pastures on the other; 2) cultivated land, which is geared more or less intensively to production of crops;
3 occupied land, which is not strictly productive but is used for gardens, dwellings, transport, recreation, commerce, etc./

1) Managed land. The scrubs and savanas are mostly derived from
agricultural land. These communities are better developed in the lower montane zone than in the lowlands and will be considered in a later section. It may be worth noting, however, that the guava scrub is probably more aggressive and successful in this (warmer) zone.

As for the pastures, it is not impossible that some of the successional grasslands near sea-level were developed subsequently to the downward migration of certain species such as Andropogon bicornis. Garcia Molinari (1952) has given an account of many of the introductions that have been attempted and of their relative success. As Figure 3 illustrates very schematically, a fairly dense low sward of St. Augustine-grass (Stenotaphrum secundatum) occupies deep soils or rolling topography, with rather a high water table, whereas the carpet-grass (Axonopus compressus) meadow is in shallow depressions. The wirybeardgrass (Andropogon gracilis) dominates on dry limestone ridges and crests. The cerillo-grass (Sporobolus indicus) is a very coarse plant of low requirements and is quick to invade even somewhat'salt soils.'

2) Cultivated land. This occupies by far most of the surface. The
principal crops are sugarcane (Photo 68), pineapple (Photos 63-65), and coconuts (Photo hh); there are also orchards (mango, citrus, bananas, plantains). Pic6
(1963), recently reviewing his earlier land-use and crop distribution maps, shows us the extent of these principal crops, and their regional correlations.

The vegetation structures of croplands undergo a number of important
variations. Whereas the orchards maintain their savanna structure over long periods of time and the coconut groves their woodland or forest formation-type, sugarcane and pineapple do not, Photo 69 shows the conspicuous differences at



8 There is, of course, a larger gamut of human interference (see Dansereau 19%7a, Chapter V, and Dansereau 196 ), but this is not involved in the present discussion. Obviously these criteria are paramount in land-use classification and have been fully applied by Pic6 (195h, 1962, 1963).












harvest time between a tall prairie of cane in the alluvial fields of the northeast, and Photo 70 the low prairie structure on the drier calcareous hills of the southwest. Likewise, pineapple starts with a steppe-like structure, which it maintains for some time, and ends up as a low prairie. Photo 45 shows pineapple grown under coconut, and thereby forming a herb layer in a forest.

3) Occupied land. Roadside and garden vegetation mostly involve exotics. Some trees are regularly planted in rows along the highways and streets. The most common are: Spathodea campanulata, Albizzia lebbeck and A.procera, Poinciana regia, Gliricidia sepium. Others are rather scattered or even quite local, such as Artocarpus incisa, Ochroma pyramidale, Montezuma speciosissima, Adenanthera pavonina, Bucida buceras, Ceiba pentandra, Eucalyptus globulus, Cordia sulcata, Ficus nitida.

Wolcott (1945), considering the various species that do well in
this habitat, warned against some of the entomological perils involved, such as the pink bollworm of cotton (Pectinophora gossypiella) which breeds on Montezuma speciosissima, a Puertorican endemic.






26


II. SEASONAL-EVERGREEN FOREST ZONE

Map 1; Figures 1 and 4; Tables I, IV, XIII;

Photos 54, 555, 7-59, 72, 74, 78



The range of limestone hills that extends from the extreme west almost to the extreme east of the Island, in the north, is in part immersed in the preceding Zone I (Map 1) or else it comes in contact with the foothills of the Central Range and even with Zone IV to the south. It also extends, in places, all the way to the coast and contacts Zone Ia, as shown in Figure 1. Rivers cut through the range and are bordered by alluvial plains (Photo 66) and by gentle slopes that accumulate the eroding materials. The altitudes of the hilltops (Zone IIa, which see) are not very great (at most, and quite exceptionally, 600 meters), so that there are no appreciably low temperatures. The precipitation throughout runs quite high, but decreases somewhat to the southwest.

The differences in vegetation (at the same altitudes) seem principally due to topographic and edaphic features. There are two principal types
of topography depending upon the extent of the ridge-tops: the sugarloaf type (or "'mogote") is compact, has a domed plateau, few spurs and exposed surfaces; the haystack type (or "pepino"), on the contrary (Photos 54, 80), offers many exposures. The lesser the asperities, the more dense is the vegetation and-the higher it rises on the hills. However, some coastal areas (Photos 81, 82) and a few of the interior uplands harbor quite a bit of scrub.

This geographical region is characterized by the periodicity of its vegetation. It may seem that the terms "seasonal-evergreen" (used for Zone II) and "semi-deciduous" (used for Zone III in the southwestern region) are six of one and half-a-dozen of the other. They correspond respectively to Beard's (1944) "evergreen-seasonal forest" and to his "semi-evergreen seasonal forest"; and also probably to Asprey and Robbins' (1953) "wet limestone forest" and "dry limestone scrub forest." Inasmuch as some of the most important tree species are common to both (for instance, Bursera simaruba and Bucida buceras) such an impression is reinforced, and the same "subxerophytic complex" can be assumed to provide the raw materials for these closely related vegetations. Actually, a difference in emphasis is implied which is supported by other
characteristics of the vegetation.

An assignment of Puertorican tree species to the three categories "deciduous," "semi-deciduous," and "evergreen" (Dansereau 1951, 1958, and Master-Table A) is not very satisfactory. It seems, for instance(fide Frank Wadsworth and Roy Woodbury), that dormancy and leaf-shedding in a number of woody species responds to short-day periods. Some species, such as Plumiera alba, Bursera simaruba, and Poinciana regia, lose their leaves whether or not preipitto falls below a critical level. Others, such as Bucida buceras,
shed them sooner or later and more or less completely (or even not at all') according to the intensity of precipitation during the short-day (winter)






27


period. Beard's (1942) study in Trinidad makes these essential physiological distinctions and uses them to characterize his evergreen semi-monsoon forest which, maybe, is similar to the gateado forest of Puerto Rico. Needless to say, we do not have systematic records for many species of this potential vs. actual periodicity. Neither Britton and Wilson (1923-30) nor Little and Wadsworth (1964) provide information beyond the statement of whether the tree is deciduous or evergreen.

Whatever the cause of deciduousness, I have assigned a category to each species listed in Table VIII. It remains that the seasonal-evergreen forest of the northern limestone hills (Zone II) is in full leaf almost all year and does not suffer, at any time, as drastic a depletion of foliage as can be witnessed every year in the southwest (Zone III). Table IV and Figure 4 show the correlations of plant-communities.

Intervales and lower slopes (hygrophytia)

The lowland rainforest reaches various altitudes on the flanks of the hills and is particularly well-developed in the ravines where they are not too steep; it must earlier have occupied much of the intervale surface. At this time, however, the intervales and lower slopes are cultivated to sugarcane (Photo 66) or are occupied by pasture. The latter have been described in the previous section.

Trumpet-wood forest (listed here under No. 43.1, and shown in Photos 54 and 57) certainly includes many plant-communities belonging to the subclimax of lowland rainforest, of seasonal-evergreen forest, and of lower montane forest. Cecropia peltata (Photo 56) plays an important role in all of them. These forests contain trees (ecologically like our boreal birches and pines) that do not readily regenerate in the shade. Acrocomia aculeata (Photo 58), Ochroma midale (Photo 55), and Montezuma speciosissima are
among them. But they also contain some of the truly sciophilous species. Moreover, their multistoried structure is not unlike that of the climax and does not usually involve the heavy masses in the lower layers that characterize jungle.

The latter is nowhere better represented than in this zone and in its various structural aspects, from forest (44.1) to scrub (44.4). Persisting banana plants and large Zingiberaceous forbs, a great tangle of many lianas, woody and herbaceous, occasional clumps of bamboo, make this vegetation very difficult to penetrate and quite useless for grazing.

Lower slopes (mesoxerophytia)
On slightly more drained topography, where soil is less mature and the surface more eroded, the lowland rainforest has not fully developed, and a somewhat more xerophytic vegetation, the gateado forest, takes over the whole middle portion of the hills. Where they are more or less flat-topped and of the "sugarloaf" type, it can extend to the top. The largest individuals of Bucida buceras are to be encountered there. Other abundant trees











ZONE II






60.1




42.1 r-'-,


60.4


'PLATEAU
mesoxeroI. phytia


CLIFF chasmophyfia


SLOPE
mesoxerophytia


hygrophyftia


SLOPE
mesoxerophytia


paranfhropophytia


44.1


48.2


48.3


47.4


INTERV AL E AND SLOPE
pa ran t h r op op h y f i a


FIGURE h. Vegetation of the Limestone-hill Seasonal-evergreen Forest Zone (I) (and the Hill-scrub'.


N)
Co


43.1


CLIFF chasmophytia


44.2


44.3


.46.4


52.6







29


are Coccoloba laurifolia and C. diversifolia, Quararibaea turbinata, Torrubia fragrans, Zanthoxylum martinicense (Photo 72). There are no tree-ferns, but quite a few terrestrial and epiphytic ones: Blechnum occidentale, Tectaria heracleifolia, Cyclopeltis semicordata, Adiantum tenerum. Lianas and epiphytes are not at all conspicuous, and play a very minor role. The trees are straight-trunked, very infrequently buttressed (Ficus laevigata). The leaflitter is not so rapidly decomposed as in the lowland rainforest and can form a more or less persistent mulch.

This forest, of course, has been very heavily damaged and the disproportion in diameter between some very large surviving individuals of Bucida buceras and Hernandia sonora and the associated trees testify to their
relict status. 9/

Abandoned fields (paranthropophytia)

Cut-over land, once pastured and/or plowed, has often reverted to firebrush (Photo 62) and more often to pepper scrub which is mostly Piper aduncum, an extremely aggressive steeple-shaped shrub (Photo 60) with slightly drooping large leaves that catch the light, and is most conspicuous. This serves as a nurse-shrub for the development of many taller woody plants, indigenous (Randia aculeata, Didymopanax morototoni) or introduced (Spathodea campanulata), that eventually form a savana. This particular line of succession does not inevitably involve masses of lianas (and therefore does not
necessarily lead to jungle formation). With the advent of R quena, Acrocomia aculeata (Photos 57, 58), Ochroma pyramidale (Photo 55), and Cecropia peltata (Photo 56), the much more hygrophytic trumpet-wood forest takes over.


9 This reminded me of a New Caledonian hill forest where Hernandia, Garcinia, Kentia, Calophyllum, and Manilkara are all represented.






30


lla. HILL SCRUB SUBZONE

Map 1; Figures 1 and 4; Tables I, XV, XII;

Photos 77, 79P-82



In the upper part of the hills (and yet sometimes extending to low altitudes, see Figure 1), the xerophytic character is more accentuated.

Cliffs (chasmophytia)

The somewhat shaded walls in the forest are typically hung with the stilt-roots of potential stranglers such as the balsamfig (Clusia rosea) and Ficus laevigata. They send downwards considerable lengths of aerial roots that fasten themselves laterally with great solidity. Many shrubs (such as Gesneria alba) grow in the minor ledges and pockets and sometimes a great abundance of Anthurium acaule and/or Pitcairnia angustifolia (Figure 4 and Photos 77, 78 Y( r

Plateaus and gentle slopes (mesoxerophytia)

The crabwood scrub (never quite a forest in height, but often quite dense) is the highest type of vegetation to develop and shows little evidence of being displaced. Gymanthes lucida is often dominant. It is frequently
branched at mid-height or slightlyEabove. Ground layers are very poorly developed. Epiphytes are not abundant.

Spurs (subxerophytia)

Wherever the tops of the limestone hills are narrow enough to receive light and be exposed to wind on all sides, and where the original forest cover has been stripped well beyond immediate recovery, the sciophilous species, without all being eliminated, no longer have any advantage over the heliophilous ones. Moreover, the rapidly eroding, karstic limestone allows a complete runoff, and drought-resistant species are favored (Photos 79, 80). Some rather typical southwestern (Zone III) species are here: Bursera simaruba, for instance, is a dominant of scrubs and savanas. Some indeed are only here in the northern part of the island, such as Plumiera alba (Photo 82). Others are on the hilltops and also in the coastal scrub (Anthacanthus spinosus). Yet others are fairly widespread but particularly prominent in this position (Comocladia glabra). Finally, a small group are strictly confined to this habitats the palm Gaussia attenuata (Photo 80) offers the most noteworthy example. It can reach a height of 10 meters, its plumose rosette overtopping the canopy at the end of a very slender trunk.

This subxerophytic formation approximately six meters in height,
varying from scrub to savana, is fairly open. It sometimes consists of as many as five layers, is rich in epiphytes. This is not unlike Asprey and Robbins' (1953) "dry limestone scrub forest."

The roble-prieto scrub (Photos 81, 82) has a greater development of


herbs, mostly grasses, and is possibly best developed in areas of contact with the littoral subzone.






31


III. SEMI-DECIDUOUS FOREST

Map 1; Figures 1 and 5; Tables I, V, XIII;

Photos 70, 71, 73, 75, 76, 83-88, 100



The southwestern part of the island is very much the driest. The prevailing vegetation has often been called a deciduous forest. It seems to me that it more closely corresponds to Beard's (1944) "semi-evergreen sea-sonal forest" than to his "deciduous seasonal forest."

At this time, in fact, there is rather little forest of any kind to be seen, if forest is to be defined (as in Master-Table C) as taller than eight meters with a canopy of more than 60 per cent coverage. As for deciduousness, many patterns are to be observed (see Section II, above). The determinants are: light-period, vegetative-sexual antagonism, drought. The responses are: A) leaf-shed just before flowering and rapid leafing during and after flowering (this usually coincides with the short-day period); B) leaf-shed with or without pronounced drought during the short-day period, with very rapid recovery of leafing activity; C) as in B but with one to three months dormancy; D) leaf-shed never complete, but very pronounced depletion of leaf-mass during dry period; E) complete evergreenness, but new growth somewhat in evidence at the beginning of the rainy season. No one
has provided us, so far, with a rating of the wood species of Puerto Rico in this respect. Britton and Wilson's flora (1923-30) offers no such information. Nor do we have a record of systematic observations through the years that would permit a partial unraveling of the internal-external forces responsible for leaf-shedding. Inhabitants of the island, however, do remember the typical Guanica area as remaining green throughout some of the wetter years and being virtually bare in the driest. In 1963 the contrast between the rather green late-February and the much grayer late-March aspects was quite notable. Table V and Figure 5 show the principal features of vegetation in this zone.

Gravelly slopes and plateaus (mesoxerophytia and subxerophytia)

Bucaro forest seems to be the regional climax where a relatively
deep soil has had time to develop. It can be fairly dense (coverage to 75 per cent in the canopy). The species associated with Bucida buceras are
Savia sessiliflora, Coccoloba laurifolia, Gymnanthes lucida, Schaefferia frutescens, Thyana portoricensis, and occasionally Bursera simaruba. Lianas and epiphytes are not conspicuous or abundant.
This forest shows many floristic affinities with the limestone-hill vegetation, especially with the communities of the higher and drier places. It also has a good deal in common with the hammocks of the Florida Everglades and Keys.

The bucaro does not often reach the girth or height that it attains in the gateado forest in spite of the fact that it often forms a woodland






67.5 68.5 69.6
ZONE Ia ZONE II

65.7 SRE

415.4 a 'Iv -64.2, vvNeV% 01S HIS
14.6
14.7 - vv v v rvvv v vV v V V V parardhropophylia
14.8 IME ST0N E PAVEM ENT / .
V, Vv v- VV v, v64.1
.;. . . .,.'.-
V V VV v v .V .hyperxerophy,ia

petro-halophyfia,,,,
~WELL- DRAINE



16.4 � v


65.7' V ,,,,
, vvv Vsvhy ,v
v V , v v / Iwv V/ vv v V V , ,Ie o


v V VPv Vq V v v v v
VV V' /VV V vv ' v v
VV Vv v v
hyperxerophyia 67.5 51.6 70.7
ZONE" 66.


I. .-.- .- . . . . * .** .

vparanhropophytiah

psammo-halto-; phy6ia750


FIGURE . Hillside semi-deciduous ve-etation in the Guanica re-ion 'Zone III" and art of th. L"


V







33


rather than a forest. The more open the canopy the greater the role played by gumbolimbo (Bursera simaruba). Such thinning-out of the upper layer and increased xerophytism occurs on ridges where the soil is thinner and on slopes where the rubble is coarser, and in areas where the original bucaro forest has been disturbed.

Gumbolimbo savana is commonly found at mid-elevations, beyond the coastal (or at least the marine) influence, but possibly at a level of greater condensation of humidity, judging by the increased abundance of epiphytes (Photos 83, 84). Here the trees can be quite tall (to 20 meters), but are widely spaced. The cactus Cephalocereus royeni probably reaches its optimum under these conditions (Photo 85). A number of spiny or otherwise xeromorphic shrubs are present: Plumiera alba, Croton lucidus, Pictetia aculeata, Comocladia dodonaea. This savana appears to be stable, although it is possible that it would be replaced, in time, by the Bucida forest. In some of the valleys where a windy corridor allows even greater condensation of moisture, it becomes a woodland and carries a heavy investment of epiphytes, mostly Tillandsia spp.

Limestone pavement (hyperxerophytia)

Hard, pitted, and fractured limestone (see Littoral Subzone Ia, above) shows alternating areas of bare rock with sprigs of Krameria ixina, Comocladia dodonaea, Croton discolor, or Cactus intortus emerging from the small pits (Photo 26) or mats of Aristida adscensionis or of Bouteloua heteroste aand of very dense thickets of sebucan-tachuelo (Cephalocereus royeni-Pictetia aculeata) (Photos 27, 30, 31). These thickets accumulate a good deal of leaf litter and harbor virtually no undergrowth at all. It is most likely in this biotope that Bursera simaruba (which eventually emerges) gets its start. A mosaic involving those several communities occupies a good deal of the south-facing territory (Photo 27). This can be compared with the "thorn-woodland to cactus-scrub" complex outlined by Curtis (1947) in Haiti.

Wire-grass steppe (Uniola virgata) (Photo 86) is rather an unusual type of grassland (Garcia-Molinari 1952)7 It is a bunch-grass or tussockgrass type, high and coarse, and tends to be invaded by the surrounding spiny scrub, forming a grassy maquis.

Screes and gravels (paranthropophytia)

Three grassland communities in this area that have structural
analogies show definite soil preferences. According to Garcia-Molinari (1952) the lanilla meadow is restricted to the lateritic plateau and the rabo-degato prairie occurs on serpentine. As for the Guinea-grass prairie, it is compatible with a number of substrata and has a wide range in Puerto Rico, as it has in many other tropical areas.

Flats, sandy and silty (hyperxerophytia)

The lowermost block in Figure 5 shows the contact between the littoral zone (Ia) and the extension inland of flats. At the extreme left, the











salt-grass steppe is in contact with grama-grass steppe and with other plantcommunities that could either be quite spontaneous or maintained by man.

This flat area is the domain of mesquite savana which is, however, quite restricted. It does not extend east of Salinas and is really important only in the Lajas Valley (Photo 88) and in the vicinity of Coqui. It requires fine-textured soils. At least, it does not occur on the limestone pavement or on the gravelly rubble. A study made at Jauca (Photo 87) suggests considerable stability. Prosopis juliflora of all age-classes is observed, and the grass layer (mostly of Chloris inflata) alternates with patches of semiwoody Achyranthes indica. Other woody plants are Parkinsonia aculeata and Capparis flexuosa. Other grasses are Andropogon annulatus and Eragrostis plumosa. Pasturing probably maintains the present structure. In sandier areas where woody growth could be more favored, the stands do not have such big trees and the other thorny species, mostly Cephalocereus royeni and Anthacanthus spinosus, tend to take over, much as they do in Haiti (Curtis
1947)0

On the contrary, where the pressure of grazing is not so great,
bucaro (Bucida buceras) comes in, but I can find no indication that it would consolidate to forest, although woodland might develop.

Pastures (paranthropophytia)

The complex lithology of this area, mentioned above, makes all early stages of vegetation development potentially very different. Quite apart from the halophytic influence (considered in Zone Ia), the limestone-schistserpentine-laterite-sand alternatives allow more than ten grassland communities to develop. Prairie, steppe, and meadow are the three structures involved: the steppes' angleton-grass, grama-grass, and wire-grass are the driest and are highly susceptible to invasion by woody plants. The meadows may or may not be distinct from the herbaceous layer that normally forms part of the mesquite savana. This is not so of the Guinea-grass but most likely is of the
paraguiito and of the rabo-de-gato, and maybe of the lanilla.

As for the cerrillo-grass, it belongs on low-fertility, depleted sites, and it also often forms a transition from the halophytic zone.











IV. LOWER MONTANE RAINFOREST ZONE


Map 1; Figures 1 and 6; Tables I, VI, XII;

Photos 50, 52, 53, 56, 60, 61, 89-95, 99, 101



There is a great deal of forest at present in this zone, which is very extensive (see Map 1). Most of it, of course, is second-growth or planted. The relief varies a good deal, consisting in foothills of the Luquillo Mountains, of the Guavate or the Maricao Highlands, and of the Cordillera Central. (The latter is the principal coffee area: see Photos 90, 91.) The east-central region also offers a particular topography: the "llanos y honduras" consist of very deep canyons cut into plateaus that
are level or gently rolling. (This is the tobacco country: see Photos 99, 101.) Figure 6 provides a sketch of the topographic variation and of the corresponding vegetation.

Rainfore st (hygrophytia)

Tabonuco forest differs quite substantially, in structure, from the lowland rainforest which is the "rainforest" of Beard (1944), whereas this corresponds to his "lower montane forest." The presence of tree-ferns, for one thing, is quite important, since they are absent in the lowlands (and indeed not to be expected in typical tropical rainforest anywhere). There are also rather more herbs, in greater abundance, than are found lower down. This is Wadsworth's (1952a,b) "tabonuco type."

The releves in Table XII are not of virgin stands, which would hardly contain such taxa as Cecropia peltata and the two species of Inga. However, these stands have a comparatively mature structure and probably contain most of the climax characteristics.

The climax (what we can know of it) and some of the subclimax
forests of this zone in some respects do not differ very markedly from those of the lowland zone. However, quite a few of the lowland species (Mammea americana, Diospyros ebenaster, for instance) do not extend above 350 meters and are therefore absent from the lower montane rainforest; some of the others take on rather a more important role at this level: Zanthoxylum martinicense, Sloanea berteriana, Lucuina multiflcra, Cordia sulcata; and finally, some species make their appearance here that are not to be found lower down, for instance, Dacryodes excelsa (Photo 89), Magnolia splendens, Buchenavia capitata, and Cyathea arborea. A host of others could be mentioned also that properly belong to the montane forest and that can be found as stragglers or in "privileged" positions within the lower montane zone:
no doubt the sierra palm (Euterpe globosa) is the most notable of these. It is quite possible that altitudinal limits in other Antillean islands are different, as witness the composition of tabonuco forest in Guadeloupe (Stehl6 19L16, Tables 89-111).

Thus the development of an association-segregate (Braun 1935) out of a rich "undifferentiated" matrix seems to follow the same pattern as in







36


ZO N E IV


74.1


43.1


UPLAND (Ilanos)
hygrophytia ."
CLIFFF chasmophyfia



lithophyfia 77.4

ch


73.5


61.4


UPLAND (lanos) paranthropophylia


CANYON (hondu ras)


61.4


CLIFF chasmophytia


79.5


77.3


78.5


RIVER-EDGE helophyfia


POND limnophytia


PASTURE SLOPES
parant hropophyftia


75.1


59.2


55.3


81.5


58.3


80.5


59.7


CULTIVATED LAND


OCCUPIED


LAND


pa rant h r opophy t i a


FIGURE 6. Vegetation and relief in the Lower Montane Zone (IV) and its
characteristic vegetation. Numbers of communities correlated to
MASTER-TABLE D and to TABLE VI.


79.1


44.1


44.4







37


Appalachia (Braun 1950) and, to reach for a nearer analogy, in southeastern Brazil (Dansereau 1947b). Asprey and Robbins (1953, Fig. 38) indicate just such a derivation, in Jamaica, of both "lower montane rain forest" and "wet limestone forest" from "rain forest."

This phenomenon extends to subclimax and/or second-growth vegetation as well. For instance, the trumpet-wood (Photos 52, 56) forest shows an admixture of the endemic royal palm (Roystonea borinquena) and of Buchenavia capitata. It is most abundant in ravines and on the more shaded slopes (Photo93).

Managed land (paranthropophytia)

However, most of the forest is managed for the cultivation of coffee (Photos 90, 91). It was found that Inga vera and Erythrina poeppigiana (which were introduced from Peru for the purpose)provided excellent shade. These two trees are now very abundant since they have freely naturalized. Wadsworth (1945) reports that several other species were tried, but with a good deal less success; he also points out the abusive exploitation of many of the steeper slopes and suggests means of restoring the forest cover. The indigenous Inga laurina is also frequently found on coffee lands. And, since many of these plots were never clear-cut, most of the primeval forest trees are present as scattered individuals, and occasionally in good numbers: Lucuma, Dacryodes, Sloanea, Zanthoxylum, Cordia, etc. The tree-ferns Cyathea arborea and emiteliahorrida are occasiona---ore frequent on the forest edge). It is to be expected that in spite of a fairly unmodified structure the coffee forest will have been managed mostly in its shrub layer where the coffee plants are substituted for both native shrubs and transgressing trees.

Modifications in structure and composition of forest, here as in
other zones, can be brought about by hurricanes. Wadsworth and Englerth (1959) have given us tables that show the susceptibility to windthrow and to breakage of various species. In fact, they point out that depth of soil is not nearly so important as the density of the stand itself. Other silvical characteristics had been established earlier by Wadsworth (1952a,b), such as: maximum size, insect and disease resistance, decay resistance, strength of wood, tolerance, growth-rate, and ease of reproduction. These features, as estimated, were
then cumulated in the form of an index which resulted in very high economic values for such species as Andira inermis, Ocotea leucoxylon, Byrsonima coriacea, Linociera domingensis, Inga laurina, Cupania americana, and much
lower ones for Didymopanax morototoni, Nectandra coriacea, Alchorneopsis portoricensis.

Abandoned land and rough pasture (paranthropophytia)

The Ing-coffee forest shows the effects of many forms of management, overmanagement, mismanagement, neglect, and abandonment. Therefore, it ranges from very orderly forest or woodland all the way to liana-infested jungle.

Tree-ferns, especially Cyathea arborea and Hemitelia horrida, are the prime indicators of lower montane conditions. Released by lumbering and clearing from their subordinate position in the forest stratification, they cominate two rather distinct communities. The first is a tall scrub which is






38


quite closed at the top by their contacting crowns (Photo 96). It is a prominent feature of open ravines and hillsides and appears able to maintain itself for many years. The second is more frequent on very steep slopes that have been stripped of forest: there the tree-ferns are isolated or in small patches, forming a savana whose lower layers are a continuous and cascading carpet of Dicranopteris nervosa, D. bifida, sometimes containing an admixture of Odontosoria aculeata and Lycopodium cernuum (Photos 97, 98). Where even this has been destroyed, the African molasses-grass (Melinis minutiflora) often invades. Big rosettes of Furcraea gigantea are also present.

Orderly pasture is not common, but some meadows of carpet-grass (Axonopus compressus) are seen.

A bluestem (Andropogon bicorne) prairie can maintain itself under light grazing (Photo 94), but will revert to scrub, involving roble-prieto (Tabebuia heterophylla, Photo 95) or pepper (Piper aduncum, Photo 61),if overgrazed or abandoned.

The latter, in turn, are soon invaded by Didymopanax morototoni,
Dendropanax arboreum, Casearia spp., Randia mitis, and other heliophilous tall shrubs and small treesi(Phot7o6O). But the introduced rose-apple or pomarrosa (Eu enia jambos) is even more aggressive, to the point of dominance in many places (Photos 60 and 93). As it has been used for posts and fuel, and has a tremendous growth-rate, Wadsworth (1943) has recommended its discriminate use under careful control.

Cultivated and occupied land (paranthropophytia)

Plantain and banana plantations are in optimum conditions at this level and can be seen to occupy small and large areas. They probably do best on northeast slopes, even steep ones. Individual plants or groups are seen to persist under all kinds of conditions.

This is the tobacco zone as well, mostly in the eastern sector where
it does best on the plateaus (llanos). This high-yielding crop is managed with much care (Photos, 99, 100, 101).

Very few if any of the species grown in orchards, gardens, along
roadsides, etc., at lower altitudes are eliminated at the lower montane level.

Honduras

The "llanos y honduras" landscape marks a very sharp contrast: the
deep canyons (see Figures 1 and 6) that abruptly break these hills and plateaus have vertical cliffs (with waterfalls and ponded waters).

Watercourses (limnophytia and helophytia)

The pools (lixnnophytia) may harbor waterlilies, but more often contain a thick growth of green algae (sometimes sealed in by Lemna) and are bordered by luxuriant Colocasia antiquorum (Photo 50). There is little development of marginal marsh�, ajvthough a uyperus iS fairly abundant and stands of cattail are well established. The riverbed vegetation is not very well






39


developed: such characteristic plants as Jussiaea are only occasional. It must be added that there is virtually no unpolluted free water anywhere in the lowland and lower-montane zones and that this alone accounts for most of the destruction of spontaneous aquatic and palustrine communities.

The cliffs are clothed with algae and mosses under the waterfals or where the seepage is fairly constant. On the dry faces, thinner crusts of algae, mosses, and lichens are formed; if the surface is somewhat pitted and where small ledges are available the balsainfig screen (Clusia-Anthurium) described above takes over. Clusia rosea roots can dangle down some 60 mtersi

The scree that forms at the foot of the canyon cliff consists of coarse gravel and is more or less stable due to periodical fall of new materials and rise of water level. Where it is best consolidated the tree-fern scrub will tend to take hold and straggling woody plants, such as the roseapple and several of the savana shrubs and trees, will be mixed in.







4o


V. MONTANE FOREST ZONE

Map 1; Figures 1 and 7; Tables I, VII, XIV;

Photos 96-98, 102-116



This tier is characterized by the montane floristic complex of which the sierra palm (Euterpe globosa) is typical. It contains a number of species not found at lower levels (such as: Cyathea portoricensis, Cyathea pubescens, Podocarpus coriaceus, Didymopanax gleasoni, Clusia minor, Ocotea portoricensis, Icacorea luquillensis), many of them endemic, and some of them very narrowly so. This is Beard's (1944) "montane rain forest" which is quite widespread in Caribbean lands. There are four areas (above 75O meters) where montane vegetation is developed, from east to west: Luquillo, Guavate, Toro Negro, and Maricao (see Map 1). Table VII and Figure 7 show typical features and relationships of the vegetation.

Watercourses (limnophytia, helophytia)

The topography being rather steep, there is not much aquatic, even less palustrine, vegetation. A few pools (some of them artificially dammed) hold waterlily or pondweed communities. There is a more promising development of algal and mossy growth on seeping rocks and in waterfalls.

Bog (oxyphytia)

The temperate character of this zone being very much in question, on
the grounds of cli-mate, flora, and vegetation, one does not expect true bog to develop, but rather fen, marsh, or swamp. The presence of Sphagnum is not enough, of course. And there do not seem to be any species that are restricted to this habitat. The melastome Nepsera aquatica is also in non-acid wetlands. Altogether I have seen only one fragmentary stand of this vegetation and must refer to it as a very dubious bog.

S s(subhygrophytia

I have named four variants of the montane rainforest: they most likely form a continuum that involves the entire subhygrophytic montane complex, which extends to the scrub as well. Table XIV has nine releves that fairly indicate this.
Stehl6 (19h6) has drawn attention to the prevalence of the sclerophyll and microphyll character of these communities.* White (1963) has measured the changes in structure and composition that take place from the lower montane to the summit in the Luquillo sector: thus many wide-ranging species emerge as dominants between much narrower limits.* Such are, at the lower altitudes: Calycogonium squaxnulosum, Micropholis garcinifolia; and at higher altitudes: Tabebuia rigida and Ocotea spathulata.






















88.1


88.4b


88.4a


88.1c


UPLAND SLOPES
subhygrophytia


DOME
mesoxerophyfia

ZONE VI


88.1b


88.1a


ZONE


83.5


87.4a


86.4


V


29.6


84.0


85.5


WATERFALL


UPLAND SLOPES subxerophytia


BOG
oxyphytia


POND
limnophyfia


MARSH helophyfia


Vegetation at the Montane Forest level (Zone V) and at the Montane Scrub level (Zone VI).
Numbers of communities correlated to MASTER-TABLE D and to TABLE VII.


FIGURE 7.


H











Palm forest undergoes several variations (Photos 107, l12, 113, ll4, ll5). In its optimum climatic conditions (in the Luquillo massif, where the rainfall is highest), it forms pure stands (Beard's (194h) "palm brake") and shows evidence of enough regeneration to maintain its effectives. However, it would not seem able to keep out the more tolerant broadleaved evergreens (Clusia minor, Calycogonium squamulosum, Tabebuia rigida) (Photos 112, 115). It is quite possible that-not unlike Pinus strobus in New England-it owes its ecological prevalence to hurricanes. On El Yunque the palm dominates in a canopy that rises to 15 meters. Its very straight trunk contrasts (especially on the steep mountain sides) with the arcuating stems of its associates. In the Cordillera Central it is more abundant in ravines (Photo 107); in Guavate (which is the lowest of the montane areas) it is scattered and marginal (Photo 89).

The broadleaved trees gain a decided advantage in the drier montane areas of the west. This is essentially Wadsworth's (1952a,b) "Colorado type." In Maricao, for instance, the sierra palm hardly ever attains dominance, whereas Clusia minor usually does, whether forming a scrub (Photo 104) or a true forestC(Photos104, 105). The presence of Podocarpus coriaceus at both ends of the island is rather a special situation. Although this species shows perfect vitality, reproducing abundantly wherever it grows, it does not actually dominate and most often does not quite reach canopy size.l0/

The areas of dense cloud condensation show another variant-the
"mist forest" of Asprey and Robbins (1953)-in which Tabebuia rii is very prominent, but where mosses play a conspicuous role. Like the moss-forests of the Pacific (Japan, Philippines, Borneo), the low-growing trees exhibit a definite "moss-line" consisting of a continuous sheath of bryophytes enveloping branches and trunk completely, and thinning out both downward and upward.

Wadsworth and Bonnet (1951) have analyzed the soil of such a "montane thicket" and found it to differ sharply from that of the lower montane forest: greater accumulation of organic matter near the surface and consequent fuller swing of the podzolization regime account for both floristic and structural differences. In other words, the climatically-induced edaphic development is more important than the direct effect of increased rainfall and exposure to wind of the woody plants themselves.






10 had observed the sane relatively uneasy and incomplete adaptation of
Podcarusin the cloudforest of the Sierra Madre oriental in Mexico. Both the Northeast-Mexican and the Puertorican podocarps are hardly to be compared with the great trees that arise in the "bosque andino" of Colombia (see Cuatrecasas 1958b, Plate XIV-2), a more fully developed example of this type of vegetation. The speciation and behavior of Poo u is of great plantgeographical interest since this genus holds the largest common denominator for all temperate rainforest development in the world today (see map in Dansereau 1957b).








43


Maquis (subxerophytia)

There may not be a considerable array of xerophytic and/or heliophilous shrubs at this level. The ever-abundant moisture and low evaporation allow rapid healing of the forest where it has been disturbed and the forest species mentioned above, especially ferns (Photo 98) and palms, probably
reinvade very readily.

At least this seems to be the usual pattern in the Luquillo massif. The Cordillera Central, however, and even more the Maricao area are more exposed to drought effects, for three reasons: 1) the mass of the montane zone is lesser; 2) the rainfall is reduced; and 3) serpentines and laterites are present in the westernmost sector.

I have ventured to distinguish two scrub communities. The montane maquis (Photo 102) is quite sclerophyll (Didyopanax gleasoni) and contains some spiny plants (not succulents). It does appear to be limited to serpentine rock and possibly it can be invaded by the montane broadleaf scrub (Photos 103, 104). The latter in turn differs structurally but not floristically from the sierra broadleaf forest of which it could be a variant on a poorer site.












VI. MONTANE SCRUB ZONE


Map 1; Figures 1 and 7; Tables I, VII, XIV;

Photos 117, 118



At the highest levels, below the outcropping and isolated rocks, the palm drops out almost completely and so-called elfin forest develops (Photo 117). This formation is less than eight meters high-therefore a scrub by my present standard (Master-Table C). Like the elfin forest-or the "elfin woodland" of Beard (1944)-it may have a moss-line at mid-height. The
almost sarmentose trees are twisted into a very complex network.

It is tempting to consider this montane vegetation as temperate instead of tropical. The simplicity of its composition, the well-marked dominance of a few species, the importance of mosses, all liken it to woody formations in Madeira or New Zealand. Several of the genera are widespread temperate rainforest characteristics: Podocarpus, Ardisia, Ocotea, Alsophila, Cyathea, Hemitelia, M ca, Magnolia, Ilex, Symplocos (see Dansereau 1957b Another
feature is the accumulation of humus. One of these stands (near Cerro de Punta) had a great depth (35 cm. or so) of fibrous organic matter, a dry peat of sorts.

The rock outcrops at the tops (where they have not been stripped of their natural vegetation for military or other purposes) have a low, windcropped, even-topped scrub, consisting of some (but not all) of the montane scrub evergreen woody species: Micropholis arcinifolia, enia bornquensis,
Tabebuia rigida, Daphnopsis philippiana (Photo11.












CONCLUSION


The foregoing description is based upon a concept of vegetation dynamics that gives recognition to composition and structure in plant-communities and ascribes their relative position to variously weighted determinants in the non-living environment. There is a resultant ordination of vegetation units according to their magnitude: the zone (Map 1, Figure 1, Table I) is under the influence of physiography and climate and is characterized by climatic formation-classes (structurally defined). The mosaic of vegetation-types or communities (for which a sampling of releves made in selected localities is available in Tables IX to XIV) is assembled, within each of the six zones, according to its fitness to the physiographic, edaphic, and historical determinants that have created the present ecosystematic matrix.

A coordinated series of figures, tables, photos, diagrams, and
releves is commented upon in the text, which is intended as a unified description of Puertorican vegetation.

As for interpretation, the master-tables offer a series of graded criteria which have been applied to all the species involved in the sampling. Table VIII fully displays this grading, upon which the diagrams also have been based. It will remain to make a functional analysis of the releves in Tables IX to XIV by applying an already-tested procedure (Dansereau 1961a) which has brought out significant differences between stands (and, by extension, between whole communities) that mere floristic differences, as such, do not reveal.

In spite of much care for detail, there no doubt are some errors and misapprehensions in the present account. It is hoped that they will be pointed out and corrected. It is also hoped that the present framework will prove useful for more detailed ecological research and experiment, such as phytosociological surveys, and population studies of both plant and animal
taxa.

A comparison also remains to be made with the vegetation of other Caribbean areas. The excellent work of Cuatrecasas (1934), of Beard (1942, 1944, 1946, 1949, 1953), of Stehle (1945, 1946, 1947), of Asprey and Robbins (1953) provide abundant materials. There are also valuable indications in Verdoorn's (1945) compendium, and in numerous other papers, such as Shreve (1914), Marie-Victorin and Leon (1942, 1944, 1956), Seifriz (1943), Howard
(1952), Lindeman (1953), Lindeman and Moolenaar (1959), Taylor (1959), Heyligers (1963), Espinal and Montenegro (1963), Espinal (1964), Harris (1965), etc.

Continued floristic exploration (Liogier 1965), especially where it is geared to ecological surveys (Duke 1965), will provide new insights in the next few years. Also, comprehensive agricultural surveys (Koenig 1953) and silvicultural reports and plans (as published throughout the span of the Caribbean Forester's existence (1939-1965) by Frank Wadsworth and his collaborators) iIcertainly allow a better understanding of both natural and induced vegetation.










46


STUDIES ON THE VEGETATION OF PUERTO RICO.

II. ANALYSIS AND MAPPING OF THE ROOSEVELT

ROADS AREA


Peter F. Buell and Pierre Dansereau/




The present study of a very small area is offered principally as an experiment. It consists in airphoto readings which are coded and transferred to a map at a scale of 1:20,000 (Map 2). A part of this has then been enlarged to 1:6,667 (Map 3); it is this map which is further subjected to various interpretations that concern the ecological features of the landscape.

The area mapped is approximately eighteen square miles (7L x21
miles) and is noteworthy principally for the extent and diversity of its mangrove swamps (about 20 per cent of the map area). It is situated at the eastern end of Puerto Rico in the townships of Ceiba and Naguabo, and most of it lies within the boundaries of Roosevelt Roads Naval Base (U. S. Navy). Aside from the extensive tidal lands dominated by the mangroves there are level, alluvial valleys which are predominantly under sugarcane cultivation, and low hills (less than 150 meters above sea level) which are vegetated with grassland (uninproved and abandoned pasture), various scrubs, and successional stages between the two. Areas of considerable extent are taken up by military installations and residential complexes.



AIRPHOTO-,TYPES

The airphotos which were available to us in 1962-64 were a series
made by the U. S. Air Force in January 1951 at a scale of approximately l:15,000. We knew of some instances where there had been change in the vegetation and in other aspects of the landscape, of course, but could use only the documentation at our disposal at the time. Ultimately (in 1965) we obtained more recent airphotos (January 1964, scale approximately 1:20,OOO) and were able to record these changes (compare Maps 3 and 7).

We read our airphoto-types by concentrating on the structural features which are defined in Master-Tables A and C. The structural characteristics recorded in Master-Table E, therefore, concern: 1) the habit-form (as


1Respectively, research assistant and head of the Department of Ecology, The New York Botanical Garden.












in Master-Table A) of the dominant plants, with their coverage and height; 2) the formation-type (Master-Table C) of the vegetation as a whole. Only ten such formation-types are recognized here (see Dansereau 1958).

Identification of plants to species or even to genus was not possible in all instances. Therefore, floristic data are very unevenly recorded. For this reason, and because we do not have a complete ground-check on all areas mapped, our correlation of airphoto-types with the vegetation-types and/or plant-communities is not always secure and many question marks must be retained.

The site features can usually be read fairly well by combining
airphoto and topographic-map information. The substratum, therefore, can be correlated to the landforms listed in Table II. As for the ecosystematic regimes, they are defined in Master-Table B and have been discussed in Part I.



MAPPING PROCEDURE

In the initial mapping of the area, boundaries were drawn wherever there appeared to be vegetation differences in the photoimage. At the same time as these boundaries were delimited, whatever structural and floristic
characteristics of the vegetation-types and their environments could be interpreted from the photos (in the light of limited ground information) were noted. Both during the progress of the mapping and after the mapping was essentially completed, it was apparent from the initial type-descriptions that some of the types were quite similar. The photoimages of these types were re-examined and carefully compared, and in many instances the types were equated. However, since this procedure was applied very conservatively, there are probably several instances where what is essentially one type may be mapped as two or more because variation in such factors as wind velocity and angle of lighting may result in variations in the photoimage which are independent of any variation in the vegetation. It is hoped that the converse situations (two or more vegetation-types mapped as one because of their similar photoimages) have been eliminated, but this possible error, unfortunately, is inherent to airphoto interpretation.l2/

Mapping was started on acetate overlays of the airphotos. This
method was later abandoned in favor of mapping on Xerox copies of the photos. The procedure for this method is as follows (examples drawn from the Roosevelt
Roads mapping):





12 The basic principles and techniques of airphoto interpretation are
discussed in the American Society of Photogrammetry's Manual of Photographic Interpretation (1960). The Society's Manual of Photograxumetry (1966) Presents a comprehensive treatment of the subject of photogrammetry.







48


1) Make a Xerox copy of each of the airphotos to be used in the mapping. (Xerox seemed to be the most satisfactory for this purpose of any of the photo-copying processes experimented with.)

2) Under a mirror stereoscope map the central region of each Xerox copy using the copy and an adjacent photo as a stereopair.

Assume a sequence of nine-inch by nine-inch airphotos, with 60 per cent overlap between photos and 30 per cent overlap between adjacent flight lines, covering a portion of the map area, is numbered 149, 150, 151, etc. (from right to left or east to west). The eastern edge of the map area is assumed to fall approximately in the center of photo 149. Place the Xerox
coyof photo 349 under the right mirror of the stereoscope and phot7T50
under the left mirror. (This gives a readable, though somewhat coarse and blurry, stereoimage.) Map Xerox 149 from the center to about 21 inches from the left (west) edge and to about 11 inches from the top (north) and bottom (south). If at any point greater clarity of detail is needed for interpretation, photo 149 can be placed temporarily over Xerox 2J49, without disturbing the position of the Xerox, to give a normal stereoimage.

Replace Xerox 149 with photo 2J49 and photo 150 with Xerox 150, and map the right half of Xerox 150 over to the line where mapping was stopped on Xerox 149.

Replace photo 149 with Xerox 150, place photo 151 under the left mirror,, and map the left half of Xerox 150 (see Figure 8).

Continue in this manner until the sequence has been mapped. Follow
the same procedure to map adjacent series (to the north and/or south) of photos.

Using a red pencil for the mapping helps considerably in avoiding
confusion of vegetation boundary lines with linear features on the photos and, at a later stage in this procedure, with contours, roads, streams, etc., on
the topographic map.,

3) Make photostatic enlargements (or reductions) of a topographic map of the area to a sequence of scales which spans the range of scales found on the airphotos.

As an example: for the Roosevelt Roads area, a topographic map at scale of 1:20,000 was used (U. S. Geological SurVey 4 iuetoorpi series). The scale on the photos ranged from about 1:14,500 to 1:17.,500. Enlargements were made to the scales of 1:.4,500, 1:15,000, 1:15.,500, 1:16,'100, 1:16,80 n 11,0. A greater number of enlargements made at smaller
intervalstrogd terag1o:caevritonwulav5ivngrae
accurcy. Aso,1tachi vagvndgeofacrythnubrftp-


































FIGURE 8. Mapping with Xerox copy and adjacent photo used as a stereopair.
1) photo l1; 2) Xerox copy of photo 150; 3) photo 1O;
4) 25-watt spotlight,








































Tracing vegetation boundaries from Xerox copy to topographic-map enlargement. photo 151; 2) cardboard divider; 3) Xerox 150; 4) portion of a topographic-map enlargement (1:17,500); 5) 25-watt spotlight.


0ln
0


FIGURE 9.










now the Xerox copy is illuminated only by the light transmitted through it from the light table, whereas the adjacent photo is illuminated by a small (e.g., 25 watt) spotlight. (An ordinary desk lamp may be used if a vertical cardboard divider is placed between the photo and the Xerox copy.) (See Figure 9.)

5) Place one of the topographic-map enlargements over the Xerox copy, and, within a small area of reasonably uniform scale, attempt to match the contours, streams, roads, buildings, etc., on the enlargement with the ridges, hilltops, drainageways, streams, roads, buildings, etc., on the stereoimage. Unless a nearly perfect match can be made, replace the enlargement with the next smaller or larger (depending on whether the scale of the stereoimage seems smaller or larger than the scale of the first-tried enlargement), and again attempt to match. Repeat until that enlargement which best fits the stereoimage in the small area under consideration is determined, and, with this enlargement in place over the Xerox, trace onto the enlargement any vegetation boundaries which lie in this area of fit. Select another small area, and repeat the above procedure until all the boundaries on all the Xerox copies have been transferred to one or another of the topographic-map enlargements.

Bear in mind throughout this operation that the scale at any point
on a vertical airphoto varies from the datum-plane scale of the photo depending on the difference in elevation between that point and the datum plane, on the degree of slope of the ground at that point, and on the aspect (with respect to the principal point) of the slope at that point.

Any direct lighting on the map enlargement (overlying the Xerox copy) tends to decrease the readability of the Xerox copy--hence the spotlight and/or the cardboard divider. The readability of the Xerox copy is increased if the photostats of the topographic maps are made lighter (with less contrast) than is customary in photostating. Also, the photostats should be made on matte
or semi-matte (non-glossy) paper.

6) Reduce (or enlarge) photostatically all of the topographic-map enlargements to the scale of the original map, thereby reducing all the vegetation boundaries to a common scale.

7) The original topographic map is now used as a base map on which to draw the vegetation map. Superimpose the topographic map successively on each of the enlarged-and-reduced topographic-vegetation maps, and trace (using the light table) the vegetation boundaries onto the base map.
At this point it will be obvious that, while the enlargements of the
topographic map need not have been made exactly to the predetermined scales,
knowing exactly how much they were enlarged facilitates their reduction to exactly the original scale.

The resulting map is likely to have a confusing abundance of information on it and must be given some further cartographic treatment (such as deleting some or all of the topographic and cultural markings) in order to produce a useful map. (See Map 2.)












The map made from the 1964 photos (Map 7) was done in much the same manner, except that the final version at 1:20,000 scale was enlarged to 1:6,667 for ease of comparison with Map 3.

It is felt that, in the absence of sophisticated photogrammetric equipment, this system is remarkably accurage, rapid, and inexpensive. It would probably not be so satisfactory in areas of greater relief or in areas where good topographic maps at a scale close to that of the airphotos were
not available.



NAP SYMBOLOGY

The code-numbers for airphoto-types, which appear in Master-Table B., have the following meaning:
Firstiit: formation-type (as in Master-Table C): 0 - 9 Second digit: strength of the prevailing ph siological stress of the ecosystematic regime (see Master-Table B): 1 - 9j3/
ThidAdiit: vegetation zone or subzone (see Map 1): even numbers (0, 2, hP
~6578)= lowland rainforest zone (I); odd numbers (1, 3., 5p 7P 9)= littoral
subzone (Ia)
These code-numbers appear on Maps 2, 3,, and 7, whereas they are correlated with other information on Master-Table E.

On Maps 2, 3, and 7, heavy lines indicate a land-water boundary.
Broken lines show the approximate location of a boundary where the transition from one vegetation-type to another takes place over an appreciable distance and the exact point of change cannot be defined.

Maps 4, 5, and 6, on the other hand, present a visual analysis of the airphoto-types in Map 3, according to several criteria. The mosaic Will consist, generally, of fewer pieces and of correspondingly larger areal units. The procedure adopted here, actually, follows the model proposed earlier (see Dansereau 1961b) in a schematic form. In that example,, the basic units (plantassociations) had not been presented in any actual spatial situation, whereas in the present instance they are.

Map 4 shows the distribution of vegetation-types or associations as described in Part I and as consigned in Master-Table D. A comparison with












Map 3 shows that several airphoto-types can sometimes be referred to a single vegetation-type. It also points to the uncertainty of cross-reference already
in evidence on Master-Table E.

Map 5 is even more simplified than the preceding one since there are only ten possibilities (see Master-Table C) and since, moreover, two of the formation-types (tundra and crust) do not happen to materialize in this particular area. This map, being exclusively structural, gives an excellent view of differential biomasses: the vast expanse and almost unbroken continuity of scrub is remarkable, as is the continuity of forest. Woodland and meadow, possibly because of their instability, are very patchy.

Map 6 shows the distribution of the controlling ecosystematic regimes. The 25 categories listed in Master-Table B follow Huguet del Villar's (1929) scheme. The symbols assigned to them on this map permit a number of
combinations, such as halohydrophytia, halohelophytia, psammohalophytia, psamohelophytia, chasmohalophytia. The other regimes represented in this map are: subxerophytia, mesoxerophytia, paranthropophytia, helophytia. This makes a total of only nine units.



INTERPRETIVE PROJECTIONS

General trends in vegetation changes in the Roosevelt Roads area,
based on a comparison of the 1951 photos (Map 3) with the 196h photos (Map 7), are as follows. The structural shifts are easily read by direct reference to the first digit: thus (prairie) is frequently replaced by h(scrub) and even by 3 (savana), indicating a progressive succession. Area in scrub is rapidly expanding at the expense of grassland. This generally occurs by way of a savana-like stage and is probably due to a relatively recent discontinuance of pasturing on the hilly land. The most rapidly expanding upland scrub community seems to be the thicket dominated by Leucaena glauca.

This leguminous plant is a relatively recent invader. In other places throughout the Tropics it has been deliberately introduced and its spread encouraged (Fosberg 1960). Whether or not this is so in the Roosevelt Roads area we do not know, but it does seem to be favored, at least initially, over the other scrub types. It generally starts in a savana-like distribution
of individuals and small clumps, usually associated with a dense growth of vines. As the clumps consolidate into a thicket the vines greatly decrease in importance. There are some indications that the Leucaena thickets are later replaced by other scrub communities, but as this evidence is meager the possibility that the thickets are stable for a long period of time and give way slowly to a forest community cannot be discounted.

Mangrove seems to be expanding slowly seaward and consolidating its outliers. This process is occurring more rapidly in the shallow, sheltered lagoons and at the mouth of the principal river course. In the inner mangrove, where the trees have been cut for firewood and poles in the past, scrubs and savanas are rapidly becoming forest with the cessation of cutting (see type











323 on Map 3, and type 325 on Map 7, west edge). Some mangrove scrubs in the outer mangrove belt (e.g., the extensive area in the northeast corner of Maps
3 and 7) seem to be relatively stable as such even though their edges are expanding into the tidal flats. It may be that the less firm substrate of
the outer mangrove is the cause of this contrast.

More specific changes are as follows. In the northwest corner of Map 3, airphoto-type 34 (Avicennia-Batis savana) shows a sharp boundary with the surrounding inner mangrove forest. On the basis of photo-interpretation and of field examination of types similar to this elsewhere in Puerto Rico, it was assumed that this type had developed on an old beach ridge which, although now surrounded by silt deposits, remained slightly higher and sandier than the adjacent swamp. Furthermore, that as long as the difference in substrate remained, the site would continue to support this quite different vegetation-type. However, 13 years later (Map 7), the area of type 34 has diminished considerably. Part of the original 34 area is now indistinguishable from the inner mangrove forest, whereas other parts of it are woodland (type 231) which, it may be assumed, is in the process of consolidating into a forest.

In another place (north-central part of Maps 3 and 7), a strip of type hh3 (Batis scrub) has become type 3l. This gives us a logical successional series: Batis scrub (4h3) invaded by mangrove to become AvicenniaBatis savana (3hl7 ,-Tavana becoming woodland (231) by consolidation of the mangroves and decrease in the importance of Batis, and finally, woodland becoming forest (121) by further consolidation of the mangroves and disappearance of Batis. There are at least two possible explanations for this succession. One is that these areas were formerly in mangrove which was cut. The harder, drier substrate has retarded the natural reforestation rate in comparison to that on the surrounding, siltier substrate more favorable to mangrove. The other possibility is that these areas have never before been mangrove-covered because of the unfavorable substrate; but because of gradual silting-over of the sandy beach ridge the substrate is becoming soft enough and wet enough to support mangroves. Considering the rate at which the change has taken place, the former explanation seems more likely, except that in the sites examined on the ground the lack of old mangrove stumps is somewhat perplexing.

The most striking changes in the character of the vegetation are those resulting from mants activities. Prominent among these is the alteration of the predominantly scrubby area in the southeast corner of Map 3 to a large expanse of meadow on Map 7. This is the result of the erection of a housing development here, and it can be assumed that the "meadow" (lawns) will remain essentially unchanged for an indefinite period of time. On the other hand, the topography was considerably smoothed off by bulldozers before
construction started and some of the rubble from this operation was dumped into two small mangrove swamps (airphoto-type 121) to the south and east. In the southern one the formation-type has been changed from forest to steppe (or perhaps desert inuediately after the filling operation), the ecosystem h~s been changed from helohalophytia to paranthropophytia and is probably approaching meso- or subxerophytia, and the community has been changed from a relatively stable one to a rapidly changing late pioneer or consolidation











stage. Barring further disturbance this will become a scrub (probably Leucaena), later a forest, and eventually a lowland rainforest, but it will never revert to mangrove. This "reclamation" process is occurring in other places in the map area.

The fill-material for some of these reclaimed areas seems to be coming from two rather extensive quarries or "borrow pits, only parts of which are in the northeast corner of Maps 3 and 7. There has been considerable expansion of these pits between 1951 and 1964, but evidence that in some parts they have been worked more recently than in others is evident from the vegetation patterns. In the current and most recent workings the ground appears virtually bare. Slightly older workings have a desert or sparse steppe cover (type 750). In the oldest, a Leucaena savana (354) and eventually a Leucaena thicket (352 or 354) develops. Whether or not
this will eventually give way to some other scrub type cannot be told from the photos. (See discussion of Leucaena glauca, page 53.)












LITERATURE CITED IN PARTS I AND II


AMERICAN SOCIETY OF PHOTOGRAMETRY, 1960. Manual of photographic interpretation. Amer. Soc. Photogrammetry, Washington, D.C., xv + 868 pp.
AMERICAN SOCIETY OF PHOTOGRAMNETRY, 1966. Manual of photogrammetry (third
edition). Amer. Soc. Photogrammetry, Falls Church, Virginia, 2 Volumes,
xx + 1192 pp.
ASPREY, G. F., and R. G. ROBBINS, 1953. The vegetation of Jamaica. Ecol.
Monogr., 23(4) :359-412.
AUBRtVILLE, A., 1949. Climiats, forts et desertification de l'Afrique tropicale. Soc. Edit. Geogr., Maritimes et Colon., Paris, 351 pp.
BEARD, J. S., 1942. The use of the term "deciduous" as applied to forest
types in Trinidad, B.W.I. Empire Forestry Jour., 21(l):12-17.
BEARD, J. S., 1944. Climax vegetation in tropical America. Ecology, 25(2):
127-158.
BEARD, J. S., 1946. The Mora forests of Trinidad, British West Indies.
Jour. Ecol., 33(2):173-192.

BEARD, J. s., 1949. The natural vegetation of the Windward and Leeward
Islands. Clarendon Press, Oxford, 192 pp.
BEARD, J. S., 1953. The savanna vegetation of northern tropical America.
Ecol. Monogr., 23(2):149-215.
BERLIOZ, J. (ed.), 1965. Contributions A la biog6ographie des Antilles.
C. R. Soc. Biog6ogr., N' Special (Nos. 308-313, 317-319, 323-325):1-64.

BRAUN, E. Lucy, 1935. The undifferentiated deciduous forest climax and the
association-segregate. Ecology, 16:514-519.

BRAUN, E. Lucy, 1950. Deciduous forests of eastern North America. The
Blakiston Co., Philadelphia, xiv + 596 pp.
BRAUN-BLANQUET, J., 1932. Plant sociology (translated by H. S. Conard and
G. D. Fuller). McGraw-Hill Book Co., Inc., New York, xviii + 439 PP.

BRITTON, N. L., and Percy WILSON, 1923-30. Botany of Porto Rico and the
Virgin Islands. Spermatophyta and Pteridophyta. New York Acad. Sci.,
Scientific Survey of Porto Rico and the Virgin Islands, 5(1-4):1-626;
6(1-4) :1-663.
COMP RE, P., 1963. The correct name of the Afro-American black mangrove.
Taxon, 12(4):l50-l52.










CUATRECASAS, Jose, 1934. Observaciones geobotanicas en Colombia. Trab.
Museo Nac. Cienc. Nat., Madrid, Serie Bot., 27:1-144.
CUATRECASAS, Joso, 1958a. Introduccion al estudio de los manglares. Bol.
Soc. Bot. Mexico, 23:84-98.
CUATRECASAS, Jose, 1958b. Aspectos de la vegetacion natural de Colombia.
Rev. Acad. Colomb. Cienc. Exactas, Fsicas y Nat., 10(40):221-268.
CURTIS, John T., 1947. The palo verde forest type near Gonaives, Haiti,
and its relation to the surrounding vegetation. Carib. Forester, 8(1) :l-12.
CURTIS, John T., 1959. The vegetation of Wisconsin. Univ. Wis. Press,
Madison, xi + 657 pp.

DANSEREAU, Pierre, 1945. Essai de correlation sociologique entre les plantes
superieures et les poissons de la beine du Lac St-Louis. Rev. Canad.
Biol., h(3):369-h17; Contrib. Inst. Biol. Univ. Montreal, 16:369-h17.

DANSEREAU, Pierre, 1947a. Zonation et succession sur la restinga de Rio de
Janeiro. I. Halosere. Rev. Canad. Biol., 6(3):448-477; Contrib.
Inst. Biol. Gen. et Zool. Univ. Montreal, 20:448-477.
DANSEREAU, Pierre, 1947b. Notas sobre a biogeografia de urea parte da Serra
do Mar. Rev. Brasil. Geogr., 9(4):497-520.
DANSEREAU, Pierre, 1951. Description and recording of vegetation upon a
structural basis. Ecology, 32(2):172-229; Bull. Serv. Biogeogr.,
8:172-229 (1953).
DANSEREAU, Pierre, 1952. The varieties of evolutionary opportunity.
Rev. Canad. Biol., 11(4):305-388.

DANSEREAU, Pierre, 1954. Studies on Central Baffin vegetation. I. Bray
Island. Vegetatio, 5-6:329-339.

DANSEREAU, Pierre, 1956a. Le coincement, un processus ecologique. Acta
Biotheoretica, I1(3-4) :157-178.

DANSEREAU, Pierre, 1956b. Le regime climatique regional de la vegetation et
les contrles 6daphiques. Rev. Canad. Biol., 15(l):1-71.
DANSEREAU, Pierre, 1957a. Biogeography: an ecological perspective. The
Ronald Press Co., New York, xiii + 394 pp.
DANSEREAU, Pierre, 1957b. A preliminary note on the structure variations of
temperate rainforest. Proc. 8th Pac. Sdi. Congr., IV(Botany):4O7-436.

DANSEREAU, Pierre, 1958. A universal system for recording vegetation.
Contrib. Inst. Bot. Univ. Montreal, 72:1-58.











DANSEREAU, Pierre, 1959. Phytogeographia laurentiana. II. The principal
plant associations of the Saint Lawrence Valley. Contrib. Inst. Bot.
Univ. Montreal, 75:1-147.

DANSEREAU, Pierre, 1961a. The origin and growth of plant conmunities. In:
"Growth in Living Systems," Proc. Syrup. on Growth, Purdue Univ., June1960;
ed. by M. X. Zarrow; Basic Books, New.York, pp. 567,-603.

DANSEREAU, Pierre, 1961b. Essais de representation cartographique des e6lments
structuraux de la vegetation. In: "Methodes de la Cartographie de la Vegetation," Colloques Internationaux du Centre National de la Recherche
Scientifique, Paris, 97:233-255.

DANSEREAU, Pierre, 1962. An application of ecological laws to woodlots.
Proc. Lockwood Conf. on Suburban Forest and Ecology, March 1962, New Haven;
Conn. Agric. Exp. Sta., Bull. 642, pp. 45-56; Sarracenia No. 7, 14 pp.

DANSEREAU, Pierre, 1964. The future of ecology. BioScience (July), 14(7):20-23.

DANSEREAU, Pierre, 1965. Ecological impact and human ecology. In: Symposium
on "The Future Environments of North America," Conservation Foundation,
New York, mimeogr. version, 77 pp.

DANSEREAU, Pierre, Peter F. BUELL, and Ronald DAGON, 1966. A universal system
for recording vegetation II. A methodological critique and an experiment.
Sarracenia No. 10, 64 pp.

DANSEREAU, Pierre, and Kornelius LEMS, 1957. The grading of dispersal types
in plant communities and their ecological significance. Contrib. Inst.
Bot. Univ. Montreal, 71:1-52.

DUKE, James A., 1965. Keys for the identification of seedlings of some prominent woody species in eight forest types in Puerto Rico. Ann. Missouri
Bot. Guard , 52(3):314-350.

EGLER, Frank E., 1950. Southeast saline Everglades vegetation, Florida, and
its management. Vegetatio, 3(4-5.) :213-265.

ESPINAL T., Luis Sigifredo, 1964. Formaciones vegetales del Departamento de
Antioquia. Rev. Fac. Nac. Agron., 24(60):1-84.

ESPINAL T., Luis Sigifredo, y Elmo MONTENEGRO Mo., 1963. Formaciones vegetales
de Colombia. Republica de Colombia, Inst. Geogr. "Agustin Codazzi,"
Dept. Agrologico, 201 pp. + mapa ecologico.
FOSBERG, F. R., 1960. The vegetation of Micronesia. 1. General descriptions,
the vegetation of the Narianas Islands, and a detailed consideration of the
vegetation of Guam. Bull. Amer. Nus. Nat. Hist., 119(l):l-76.

GARCTA-NCLINARI, Ovidio, 1952. Grasslands and grasses of Puerto Rico. Univ.
Puerto Rico, Agrico Exp. Sta., Bull. 102, 167 pp.










GLFASON, H. A., and Mel T. COOK, 1927. Plant ecology of Porto Rico. New
York Acad. Sci., Scientific Survey of Porto Rico and the Virgin Islands,
7(1-2) :1-173.

GOLLEY, Frank, H. T. ODUM, and R. F. WILSON, 1962. The structure and metabolism of a Puerto Rican red mangrove forest in May. Ecology, 43(l):9-19.

HARRIS, David R., 1965. Plants, annals, and man in the Outer Leeward Islands, West Indies. An ecological study of Antigua, Barbuda, and
Anguilla. Univ. Calif. Publ. Geography, 18: ix + 164 pp.

HPEYLIGERS, P. C., 1963. Vegetation and soil of a white-sand savanna in
Suriname. Koninkl. Nederl. Akad. Wetenschappen, afd. Natuurkunde,
Tweede Reeks, 54(3):1-148.

HOLDRIDGE, Leslie R., 1940. Some notes on the mangrove swamps of Puerto Rico.
Carib. Forester, 1(4):19-29.

HOLDRIDGE, Leslie R., 1965. The tropics, a misunderstood ecosystem. In:
Symposium on "Human Populations in Relation to Food Supply, AIBS Ann.
Meeting, Urbana, Illinois (August 1965), 12 pp. mimeogr. + figures.

HOWARD, Richard A., 1952. The vegetation of the Grenadines, Windward Islands,
British West Indies. Contrib. Gray Herb. Harvard Univ., No. 174, 129 pp.

KOENIG, Nathan, 1953. A comprehensive agricultural program for Puerto Rico.
U. S. Dept. Agric., 299 pp.

KTE, K. W. 0., and C. B. BRISCOE, 1963. Forest formations of Puerto Rico.
Carib. Forester, 24(2) :57-66.

LMlEAN, J. C., l1953. The vegetation of the coastal region of Suriname.
Uitgevers-Maatschappij v/h Kemink en Zoon N.V., Utrecht, x + 135 pp. + map.

LINDEMAN, J. C., and S. P. MOOLENAAR, 1959. The Vegetation of Suriname,
Vol. I, Part 2: Preliminary survey of the vegetation types of northern
Suriname. Van Eedenfonds, Amsterdam, 45 pp. + maps.
LIOGIER, Alain, 1965. Nomenclatural changes and additions to Britton and
Wilson's "Flora of Porto Rico and the Virgin Islands." Rhodora, 67(772):
315-361.

LITTLE, E. L., and F. H. WADSWORTH, 1964. Common trees of Puerto Rico and
the Virgin Islands. U. S. Dept. Agric., Agric. Handbook 249, 548 pp. LOBECK, A. K., 1922. The physiography of Porto Rico. New York Acad. Sci.,
Scientific Survey of Porto Rico and the Virgin Islands, l(4):301-384.

MARIE-VICTORIN, Fr~re, et Fr~re LEON, 19h2. Itin~raires botaniques dans
l'Ie d Cua premiere serie). Contrib. Inst. Bot. Univ. Montreal, No.
Wl, h96 pP. + map.






60


MARIE-VICTORIN, Frere, et Frere LEON, 1944. Itineraires botaniques dans
l'Ile de Cuba (deuxime serie). Contrib. Inst. Bot. Univ. Montreal, No.
50, l410 pp.
14ARIE-VICTORIN, Frere, et Frere LEON, 1956. Itineraires botaniques dans
1'tle de Cuba (troisieme serie). Contrib. Inst. Bot. Univ. Montreal,
No. 68, 227 pp.
NJRPHY, Louis S., 1916. Forests of Porto Rico; past, present, and future,
and their physical and economic environment. U. S. Dept. Agric. Forest
Service Bull. 354:1-99.
PAPY, H. Ren6, 1955. Tahiti et les Iles voisines. La vog6tation des Iles de
la Socite et de Makatea (Oceanie Frangaise). 2e partie. Tray. Lab.
For. Toulouse, Tome V, IIe Sect., 1(3):163-386.
PIC6, Rafael, 1954. Geograffa de Puerto Rico. Parte I. Geografloa fisica.
Editorial Universitaria, Rio Piedras, Puerto Rico, xiii + 2)43 pp.
PIC6, Rafael, 1962. Puerto Rico: planificaci6n y acci6n. Banco Gubernan.
Fomento Puerto Rico, San Juan, xv + 312 pp.
PICO, Rafael, 1963. The commonwealth of Puerto Rico. Focus, 14(2):l-6,
RAUNKIAER, C., 1934. The life forms of plants and statistical plant geography.
Clarendon Press, Oxford, xvi + 632 pp.
RAWITSCHER, Felix K., 1944. Algumas nog6'es s~bre a vegetagao do litoral
brasileiro. Bol. Assoc. Ge6gr. Bras., No 5:13-28.
RICHARDS, P. W., 1952. The tropical rain forest. An ecological study.
Cambridge Univ. Press, xviii + 450 pp.
ROBERTS, R. C., 1942. Soil survey of Puerto Rico. U. S. Dept. Agric., Bur.
Plant Ind., Soil Survey Series 1936, No. 8, 503 pp.
SAWYER, John 0., Jr., and Alton A. LINDSEY, 1964. The Holdridge bioclimatic
formations of the Eastern and Central United States. Proc. Indiana Acad.
Sci., 72:105-112.
SEIFRIZ, William, 1943. The plant life of Cuba. Ecol. Monogr., 13:375-426.
SHREVE, Forrest, 1914. A montane rain-forest. A contribution to the physiological plant geography of Jamaica. Carnegie Inst. Washington, Publ.
199, 0ll pp.
STEENTS, C. G. G. J. van, 1962. The distribution of mangrove plant genera and
its significance for palaeogeography. Proc. Koninkl. Nederl. Akad.
Wetenschappen, Ser. C 65(2) :164-169.
STEHLE, Henri, 1945. Forest ty-pes of the Caribbean Islands. Carib. Forester,
6( Suppl. ) : 273-414.










STEHLE, Henri, 1946. Les types forestiers des Iles Cara bes. Carib.
Forester, 7(Suppl.) :337-709.

STEHLE, Henri, 1947. La v6getation sylvatique de l'Archipel caraibe. Etude
d' eco-phytosociologie. These, Universit6 de Montpellier, 548 pp.

TAYLOR, B. W., 1959. Estudios ecologicos para el aprovechamiento de la
tierra en Nicaragua. Min. Econ., Inst. Fomento Nac., y ONU FAO, Vol. I,
xv + 338 pp.

THORP, James, 1941. Climate and settlement in Puerto Rico and the Hawaiian
Islands. In: Climate and Man, Yearbook of Agriculture, 1941, pp. 217-226.

U. S. ARMY ENGINEER WATERWAYS EXPERIMENT STATION (ed.), 1963. Military evaluation of geographic areas, reports on activities to April 1963. Corps of
Engineers, Vicksburg, Mississippi, Misc. Paper No. 3-610, December 1963,
vi + 237 pp.

VERDOORN, Frans (ed.), 1945. Plants and plant science in Latin America.
Chronica Botanica Co., Waltham, Mass., xxxvii + 383 pp.
VILLAR, E. Huguet del, 1929. Geobotnica. Editorial Labor, BarcelonaBuenos Aires, 339 PP.

WADSWORTH, Frank H., 1943. Pomarrosa, Jambosa Jambos (L.) Millsp. and its
place in Puerto Rico. Carib. Forester,(TBY-19.

WADSWORTH, Frank H., 1945. Forestry in the coffee region of Puerto Rico.
Carib. Forester, 6(2):71-75.

WADSWORTH, Frank H., 1952a. Forest management in the Luquillo Mountains, II.
Carib. Forester, 13(2) :49-61.

WADSWORTH, Frank H., 1952b. Forest management in the Luquillo Mountains, III.
Carib. Forester, 13(3):93-119.
WADSWORTH, Frank H., 1959. Growth and regeneration of white mangrove in
Puerto Rico. Carib. Forester, 20(3-L):59-71.

WADSWORTH, Frank H., and Juan Amedee BONNET, 1951. Soil as a factor in the
occurrence of two types of montane forest in Puerto Rico. Carib. Forester,
12 (2) :67-70.

WADSWORTH, Frank H., and George H. ENGLERTH, 1959. Effects of the 1956 hurricane on forests in Puerto Rico. Carib. Forester, 20(1-2):38-51.
WALTER, Heirich, 1962. Die Vegetation der Erde in 1 kologischer Betrachtung.
Band I: Die tropischen und subtropischen Zonen. VEB Gustav Fischer
Verlag, Jena, AV + 538 pp.

NEST, Robert C., 1956. Mangrove swamps of the Pacific Coast of Colombia.
Ann. Assoc. Amer. Geogr., [46(l) :98-121.






62



WHITE, H. H., Jr., 1963. Variation of stand structure correlated with
altitude in the Luquillo Mountains, Carib. Forester, 24(1)Lj46-52.

WOLCOTT, George N., 1945. Trees for roadside planting in Puerto Rico.
Carib. Forester, 6(3):115-120.

YOUNG, Robert N., 1955. A geographic classification of the landforms of
Puerto Rico. Chapter II In: C. F. Jones and Rafael Pic6 (eds.),
"Symposium on the Geography-of Puerto Rico.," Univ. Puerto Rico Press,
Rlo Piedras, pp. 27-46.







63


APPENDIX A. -- Terminology


Most of the vocabulary used here is defined in the glossary (pages 317-336) of Dansereau's "Biogeography: an ecological perspective" (1957a).
Additional terms are to be found in Dansereau, Buell, and Dagon (1966).

Airphoto-type: a vegetation unit and its habitat as determined chiefly by
airphoto image interpretation and whose description often lacks some of the definitive elements, notably the floristic, included in vegetationtype descriptions (see Master-Table E). It is the basic unit used here
in large-scale mapping (see Maps 2, 3, 7).

Belt: a vegetation-type or a plant-community that occupies a definite position
within a habitat or an ecosystem. It is usually poised on a gradient, such
as a floodplain. (Not to be confused with zone, which see.)

Ecosystem: the conjunction of living and non-living elements which, on a given
site, operate the conversion cycles of environmental resources.

Ecosystematic regime: the prevalent physiological stress that dominates an
ecosystem and determines the behavior of its living members (see MasterTable B).

Formation-type: the purely structural definition of a stand of vegetation,
based on height and coverage of woody and/or herbaceous elements. In the
present context, only ten of these are recognized (Master-Table C).

Habitat: the physiographic and edaphic unit within which an ecosystem develops.

Plant-community: a phytosociological unit described in terms of at least
summary floristic composition and outstanding structural features.

Regime: regular and often rhythmic behavior, e.g., climatic regime (mediterranean, monsoon), soil regime (podzolization, laterization), ecosystematic
regime (helophytia, tropophytia).

Stand: an actual area of continuous and homogeneous vegetation, e.g., a
stand of cattails, a stand of mountain palms.

Vegetation-type: a purely descriptive, sometimes very inclusive, characterization of the aspect of vegetation; it may contain some floristic, some
structural, and some site notations. (Many examples in text and in MasterTable D.)
Zone: a geographical area that combines certain features of topography and
climate (see Map 1 and Table I).






64


APPENDIX B: Map Errata



Map 2. 1) Unlabeled area in the north-central part, bordered on the wetOt by
the head of a bay and on the northeast by type 341: should be
labeled 889.
2) Legend: for 1952 read 1951.

Map 5. Narrow strip in the southeast part-bordered on the north by Scrub,
on the west by Forest, and on the south and east by the sea, and
corresponding to type 781 on Map 3- should be Steppe.

Map 6. 1) Area in the southeast part-bordered on the north and west by
subxerophytia, on the east by chasmohalophytia, and on the south by the sea, and corresponding to type 121 on Map 3-is indicated
as "helophytia" but should be "helohalophytia."

2) Two areas in the northwest part-bordered on the north, east, and
south by subxerophytia, and on the west by helohalophytia, and corresponding to type 534' on Map 3-are indicated as "helohalophytia" but should be "helophytia."


Map 7. 1) Unlabeled area in north-central part-bordered on the east and
west by type 456, on the north by type 364, and on the south by
type 362-should be continuous with type 362 to the south.

2) Large, very irregularly shaped area just southeast of the center
is labeled 448, but should be 456.
3) Area in the southeast part-bordered on the north, east, and west
by type 644, and on the south by the sea--is labeled 450 but should
be 750.


4) Legend: for 1963 read 1964.












MASTER-TABLE A. Categories and symbols used to draw structure
diagrams (Dansereau 1958, Dfasereau, Buell, & Dagon 1966).


i. LIFE-Fa

W ( erect woody plants

L climbing or decumbent woody
plants
A epiphytes and non-rooted
aquatic plants

H herbs

M ( bryoids and crusts


2. STRATIFICATICN

7 more than 25 meters

6 10 - 25 meters

5 8 - 10 meters

4 2- 8 meters

3 0.5 - 2 meters

2 0.1- 0.5 meters

1 0.0 - 0.1 meters


3. COAGE

b barren or very sparse

i interrupted, discontinuous

p in patches, tufts, clumps

c continuous


4. SEASONALITY
d []deciduous or ephemeral

s semideciduous
e evergreen

evergreen-succulent or
evergreen-leaf less




. FA ShAP - AND SIZ

o leafless

n 0 needle, spine, scale, subulate

g 0 graminoid

a medium or small
h oroad

v ) conipound

q Q)thalloid

6. 1 mA XTmI

o leafless

f 0 filmy

z Elmembranous x * sclerophyll

k succulent or fungoid






AS.R-TAME B. Huguet del Villar' s (1929) ecological, non-geograplhical classification of the physiological regimes tat characterize ecosystems


ELATIV HARMONY


Harmony of factors


partly aquatic Dominant


discrepwany of one factor


NATRE OF CONTROL


Chemical Thermic


HABITAT


Biotic Mepritic accuiulations


Harmory of factors ISSUHYTIA


Dominant discrepancy of one factor


Water scarce: XEROFETIA Teperat-ure extreme


Reaction diverging


Reaction divergingq
from neutral

EDAPHOPHYTIA

Phy sical condition
unfavoriCble;
substrattnm
excessively:

Perturbing biotic
factor


BIOGEN0"HYIA


Texttre of


Harmon- of factorsIsiostratim


HarmoxV of factors


Texture of


substratum


Constant Subccnstaent Discon"inurous


Moderately Extreamely Excess Deficiency


Acal i iityAcidity


Loose
Dry

PETROPHYTIA Compact:

Putrescible
accrulations

General transformation
of the envirmrbient


Constant


Constant


SUBSTRATM


7. Hygrophytia . Subhygrophytia

9. Tropophytia '0. Mesoxerophytia i!. H~reopyi 12. Subxeroohytia 13. Psychrophytia


14. Halophytia 15. Omypytia


16. Psamophytia 17. Chersophytia lb. Chasmophti*a 19. Lithophytia "" (Pezosaprophytia)

20. Biogrenopht-ia (s.str.) 21. Paramthropophytia 22. *Hydrosaprophytia 23. *"Pezosaprophytia


24. Ectobiophytia 25. Endobiophytia


QUALITY OF CO1 0L Constant Subconstant Alcaiity Acidity Excess Deficiency


Tropical rainforest Subtropical and temperate
rainforest
Monsoon and deciduous forest Mediterranean forest Desert

Savana
Tundra


Seashore
Bogs and needle-leaf forest


Dunes


Dune s Shallow gravels Crevices


Bird cliffs Buildings, yards, rai!vWys Logs under vater Rotting logs Bark of trees, sheaths of
bromeliads
Intestines of animals,
living wood


Totally or


TYPICAL ECOSYSTEMS Lakes, ponds, streams Marshes, temporary ponds Sea, salt lakes Acid lakes Warm springs Arctic seas, ice, snow


geophysical: ECOPHYTIA


H'YROPHYTI A


emerged PEZOPHY. TIA


Aquatic Emerged


Supporting Harboring


organic:


SAPROPHYTIC (dead)
BIO
PRYTIA (living),


i.Limophytia 2. Helophytia

3. Halohydrophytia
4. Oqlhydrophytia

5. Hydrothermophytia
6. Cryophytia

(Hydrosaprophtia)


I


0%






67


MASTER-TABLE C. Formation-type. The height and coverage of the ten formation-types: the height, in meters, is average (not always maximum); the "class" refers to the stratification and coverage classes of Table III. Slightly modified from Dansereau 1958.


WOODY PL-ANT (W, L) 1M iS PLANTS (H, 9E,)
SYMBOL AION- Height Coverage Height Coverage
TYPE _ _ _ _ _ . .
meters class % class meters class % class

1 FOREST 8 7-5 +60 c var. var. var, var,

2 WOODL AND 16 7-5 25-60 i-c v.0r. var. var. var.

3 SkVANA 2-10 5-4 10-25 i-p 0-2 3-1 25-100 i-p-c

4 SCRUB 0.1-68 4-2 25-1OG i-c var. var. var. vr.

5 PREARIr 0.5-2 3 50-100 c

60M.0-0.5 2-1 50-100 c
STEPE 0.1-2 3-2 0-25 b-1 0.0-2 3-2 10-50 i-p


8 ESElT 0.0-10 5- 0-0 b 0.0-.5 3-1 0-10 b

9 TUNDA 0.0-0.25 2-1 10-60 b-c 0.0-0. 25 2-1 0-20 b-p

10 CRUST 0.0-0.1 1 50-100 p-c








68


NAST -TALE D. List of the vegetation-types and plant communities of Puerto Rico.
The last digit (at the left) refers to formation-type (as in Master-Table C).
Photo numbers show stands of vegetation; if underlined,they display a whole
landscape; if in parentheses, an individual dominant or characteristic plant.


Vegetation-type Zone (Map 1 and Table I) Composition Diaor Plant-comaunity gram Photo
. -- ,and. mmName Ia I 11a, III IV V VI Dominants Other species relev6 ber
ber ~mnber

0.0 vegetation X X X X X X X 34
33
1.5 kelp tangle X Laminaria sp.
2.6 algal meadow X Ulva lactuca

3,6 turtle-grass X 'Thalassia
3 bed r" testudinum 3

4.0 marine- algal X Enteromorpha sp.
araist

4.6 fucoid mat X Turbinaria 1
tuxbinta 2


51 forest A mangle nIi
i ( 1 2 )

3
5.4 red-mangrovesubX Rhi zophora Avicemnianiid 2 , 4!



black-mangrove Aviceymia Laguimx lar ia 10
6.1 forest nitida racemosa 4 13
black- mangrove Avi cennia 5 11
. scrub.Xnitida 1
Rhizophora mangle 6
7.1 mixed mang ove Avicennia nitida 7
forest Laguncularia
racemosa 9
7-3 mangrove savana X Rhizophora mangle Batis maritima 10 17
.-.--.Avicennia yitida
74 mixed nkgrove Lagunclariia 11 (l
scrub racemosa 12

9.6 glasswort mat X Salicornia
bigelovii
sea-purslane Sesuvium 18
10.6 mat X portulacastrum19
___ ___ (20)
11.4 saltwort scrub X Batis maritime Avicennia nitid a
. . . . X- . .i . .Acrostichum
12 salt-fern aureum
125 prairie Acrostichum
danaefolium






69


NameI III IV V V Dominants Oer species 1ia Photo
ber I I a gram

bay-cedar- Suriana maritima 21
13.4 horse-bush X 13 21
scrub , corymbosa .
bay -cedar x Suriana maxitima Borrichia (22)
13"7 steppe arb~trescens

14.6 salt-sedge Fimbristylis
meadow x spadicea 27
14-7 spurge-sedge x Euphorbia 14
rMat buxifolia

14.8 spurge desert x Fibristylis15

Opuntia rubescens
high button- Conocarpus erecta 17 28
15.4 mangrove- X Cephalocereus 17
cactus scrub royeni

low button- Conocapus erect 26
15.-a mangrove- x Cactus intortus
cactus scrub

15.9 button-mangrove x Conocaxpus erect 23
matted scrub
sOpuntia rubescens sebucan- Conocarpus erecta Leaireocereus
16.4 tachue1 o X Cephalocereus hystrix (31)
thornscrub royeni Plumiera alba 32

beach-grass- Ipomoea- Canaval'a 35
17.5 mornirg-glory X Sporobolus- arvitia 19 36
prairie Spartina

17.6 mxrnig-glory X Ipomoea pes-caprae 20
festoon 34

17.8 sea-rocket X Cakile lanceolata 46
ctrip

salt-grass x 21 37
prairie
Spartina patens 22
salt-grass 23
io.7 steppe X 24 (38)
25
26
b Sporobolus Paspalum
19.6 eahras X27 39
si.ad virginicus vaginaturn 28

29
30

20.4 sea-grape X Coccoloba uvifera 33
scrb 41
=, 42
36g 43
37







70


Num
ber


Name


sara--mara 21.4 scrub


22. snake-bark "4 scrub



Leucaena woodland

234 Leucaena
thicket

Coconut
24.2 . ant at ion


Australianpine screen

26.4 Ilbaralmond scrub
27.6 water-h cinth
mat


28.6 water-lettuce
mat

29.6 waterlily mat


para-carib30.5 grass marsh



trorupet il la3 1 -c5 grass "arsh


32.5 cattail marsh


Ia


X


x


x


x


X

x


X


x

x


x


x


x


X


x


x


x

x


x x


IaI


IV


x


x


x


33 reed-grass xK N
marsh


VI


Domirants


Dalbergia ecastophyllum


Colubrina ferruginosa


Leucaera glauca


Cocos nacifera


Casuarina equisetifolia

Terminalia catappa

E ichhornia crassipes


Nymphbaea amp la
Panic ca purpurascens Eriochloa polystachya
Panicum
aquat icuM Hynmeachne amplexicaulis

Typha
domingensis


Pbragmites
conmmnis


Other species


.Arris elen.ifera Anthacanthus
spir osus Comocladia dodoraea Reynosia uncinata


Eleocharis
34-. spike-rush x x interstincta Sagittaria 45
3 marsh Eleoch&aris lancifolia 46
mutata

Cladium
35.5 sedge marsh X X Carex AZ
Scirpus

rivercane X X Gyneritum
36.5 brake sagittatum


37.4 icaco scrub xChrysobalaus Blechnum indicu 47


82 pond-apple Annona glabra Acrostichum 48
8 swamps - Conocarpus erecta aureum


Photo


48


44
(45)


Diagram


39 40 41 42


43 44


1--l. 4- 0 4-m - - a, Im"le


-L--. - ---


(49


Imln 11 ":,l






71


Dia
NUraer Name Ia IIII IVI V i P omirants Other species Dram Photo
ber I I ~ I IDmnat te
391 Pterocaxpus Pterocarpus49
1 SWmp officinalis

40.0 duckweed crust X X Lemna perpusilla 50
41.7 dr-flowerX Cleome spinosa 51

Diospyros
ebenaster Nectandra
42.] wanrest X X Mameaamericana membranaces
ianilkara
nitida

Andira inermis
Acrocomia
;;A
x K x aculeata 5
43.1 trumpet-wood X X X Cecropia peltaa Erytrina 61A6
forest poeppigiana57
0chroma (58)
pyramidal0

Casearia
44.1 Andira
44.2 jungle X X X Piper
44410 Abrus
44.4 Ipomoea

45-4 guava scrub X Psidium guava
(59)
46.4 pepper scrub X X X Piper aduncum 66

47-4 firebrush scrub X X Croton lucidus 62
second-growth
48.2 woodland x x Randia aculeata
. -- .- Didymopanax cpao laes
43 second-gyow-th x x x Moro totoni ca6panulata _0
49.6 St. tigustine Stenotaphrum
grass sard x x (x) secundatum

50.6 carpet-grass xfmonopuz
sw rd Xcompressus

51.6 cerrillo-grass X x Sporobolus
s56 m rd indicus
52.6 wiry-beardgrass Andropogon
sward X Xgracilis

537 field X Ananas msativa 64
~(65)

66

54.5 canefield x x (x) iofficiarum 69
~70 ~71






72


Num-
ber Name Ia I I iii IV V Vi Dominants Other species Dia- Photo

55.3 orchard Mangifera irdica
Citrus limonium
58.3 gardens x (X) (X) x (ornamentals)

Casuarina
59.2 screens X (x) (x) X equisetifolia Terminalia
Poinciana regia catappa

Hibiscus
59.4 hedges X (X X llamandi
Ba~mbusa

Stach y tarpheta 73
59. 7 roadsides X (X) (X) X jamaicensis Ceiba pentanra 74
Sida carpinifolia 75

Zanthoxylum
martinicense
%amia
latifoliolata 50
61gateado Coccoloba, Torrubia fragrans 51 (72)
forest X laurifolia Faramea 52
occidentalis 53
Quararibaea
turbinata
Guazuma ulmifolia

60.i crabwood scrub X Gymnanthes
lucida Ehugeniia inonti cola 54

Pitcairnia
balsamfig Clusia rosea gstifolia577
61.4 screen Anthurium acaule Dipholis 55 (78)
� salicifolia
62.3 limbe-gumbo-X Gaussia Sideroxylon 56 79
limbo v attemata foetidi ssimum
Bursera Ilex nitida
62.4 limbo-subo X simaruba Conocladia glabra 57 80
limbo scrub

Randia aculeata
roble prieto Tabebuia Plumiera alba 38 81
" scrub X heterophylla Elaeodendron 58 (82)
xylocarpum
Savia
64.1 bucaro forest X Bucida sessiliflora
. .-buceras Krugiodendron
64.2 bucaro woodland X ferreum 59

gumbolirnboCephalocereus(8' 64.3 osamana X Bursera simaruba Tb .a 18 84

portoricensis(8
67 wire-grass XTnoavraa8
67 steppeXUilairaa8
S graca-grass Bouteloua
66.7 steppe heterostega

67.5 prii X X ! Panicum maximum







73


N um - II D ia- h o
ber Name Ia I II I IV V VI Dominants Other species am Photo

Arundinella
rabo-de-gato X confines
pasture Andropogon
semiberbis
Leptocoryphium
lanatum
69.6 lanilla pasture X aritum

portoricensis

angleton-grassxAndrpogon 7 sward amulatus
Chloris inflata
71.6 paragto X Pennisetum
meadow ciliare

Chloris inflata
72 osopis Leonotis 88
savant juliflora - nepetaefolia

Parkinsonia
Bucida buceras aculeaa
72.3a bucaro-mesquiteX Prosopis achyranthes 60 87
sa vana julifloradc a

Colocasia
73-5 aroid-sedge x antiquorum 50
belt Cyperus spp.

73.5a yerbade- X Jussiaea repens
clavo strip


Dacryodes berteriana
tabonuco excelsa Philodendron 62
74.1 forest Lucuma krebsii 63 89
multiflora Ruellia cocciea
Hemitelia horrida

Inga-coffeexInga vera Erythrina 61
75.1 fIore atfx Inga lauria
Coffea arabica poeppigiana

rose-applexEugenia m 92
764 Ithicket
Cyathea arborea (97)
Dicranopteris spp. 98
77.4 fern scrub x Cyathea arborea
Hemitelia horrida
molasses-grass Me linis
7o.5 field X minutiflora

Ardropogon 61
bluestem X bicornis 6
b l-5 e sr ai rX S p o r o b o l u s r o9 4
prarieindicus (95)
99
80 5 t b c o f e &XN icotiana 100
805tbcofedtabacu (101)


81.5 banana X IMusa sp.
815 plantation










Num- II Dia-II IhontM .
ber Name Ia I a iI IV V VDomits Other species gram Photo

2 cliff algal
02 0 crust x


83.5 ponlweed Potamogeton sp.?
tangle


84.0 algal carpet


85.5 sedge marsh

86.4 bog scrub


87.4 montane maquis


montane
87.4a broadleaf
scrub



sierra
88.1 broaAleaf
forest




sierra
88.1a broadleaf-palm
forest



88.lb sierra-palm
forest



8.lc sierra mossforest


88.4a elfin forest


x



x


X





x




x




x


x


Cyperus sp.

Nepsera aquatica Sphagnum spp.

Di dymopanix
gleasoni

Clusia minor calycogonium cquamulosum



Clusia Minor Calycogonium sGuamulosum



Euterpe globosa Calycogonium

squamulosum Clusia minor


Euterpe globosa


TabebuLa rigida Podocarpus coia~eus Muscinae

Mi cropholis garcinifolia Clusia krugiana


Cyathea pbescens Clusia krugiana


Ocotea leucoxylon





Podocarpus coriaceus


Tabebuia haeantha Rapanea ferruginea


66
67 68




69


70
71


Thibaudia krugii
Eugenia Ilex sintenisii 72
68.4b high montane X borinquenss Ocotea spathulata 118
" scrub Tabebuia rigid--I-a lcorea 73
luquil learns,


102


103


104


106
log



110
illl
112
115 118





117






FIASTER-TAULE. Airphoto-types of the Roosevelt Roads area shown on Maps 2, 3, and 7 (n.a. = non-applicable; em. emulsionn or mixture of two format


(5) (6) (7)


(8)


(9)


(10)


(11)


(12)


Structural character


Formationt-pe


Dominant habitform


Height
(MI.)


Cover
(CA)


eristics


Other structural
features


Floristic characteristics Composition Abundance


Site features


Substratum


Regime


1 895 0.0 desert n.a. n.a. n.a. barren n.a. wave-ished, sandy beach
-" 1 Ieset na. l~d I ________ ________________ ph~ti


desert


may have occasional algal growth


tidal flats; silty


high(> 2 in.), in- out
(8) 7- terlocking stilts ; Rhizoa mangle do. mae
3 151 5.1 forest trees 12(15) 70-80 conmon stem om.
diamter:�~ c. -along tidal channels .00 diameter: 15 cm.

lowm (( ".), inscrub 2-10 terlockirg stilts;
desert) shrubs 2- (50-80 small (ca. 5 m. hizo. (occ tidal flats or very
4e r ) wi h i n d i a m t e r)- ) w i d e ly . .do. ) s h l l o w w t e r
clumps) scattered clumps
of vegetation

low( 8scrub 20-5 terlocking stilts;
sr5 (7080 small (ca. 5 M. Rhizphora mangle dom. (occ.) tid
431 5.4 .desert) shrubs within diameter) clumps .ra. .
em.)chimps) spaced 5-15 in.
apart

scrub shrubs interlocking l
5.4-5.1 (forest) (trees) 6-8(10) 7C-90 stilts tidal channel margins

6 49 Rhizohramangle outer grove;
5.4 scrub shrubs 2-3(5) 5-80 low (< 1 m.) in- generally not so firm
terlocking stilts as in 141

outer mangrove;
7 47 5.4 scrub shrubs 2-3(5) 50-80 low (< 1 m.) in- hizohagenrly sfI
terlocdng stilts . a.g.eneas ll n sf


EL '~IA
belobaloA)


Key num
ber


Map Airphototype code

ber


Vegetationtype, or plantCommunity (MsterTable D)


889


4.0


HELOALOP14YTIA


df


;ion-types).






(.) (2) (3) (4)


(5) (6) (7)


(8)


(9)


(10)


(11)


iRhizophora mangle dom.?
6.low(<,m.) in. inner side of outer
8 423 5.4 scrub shrubs 3-4? .65-80 terlocking stilts -race r 0Cc.-OMM nrove;
7.4? - cmdier tan 14
Avicennia nitida? rare
Rhz2hra mangle dom.; ab. stilts abundant L lcularia conn.
9 141 7.1 forest trees 15(18) 6o-8o and usually raceMosa (dom.; ab.) outer nove
interlocking . . . . . .
Avicennia nitida occ.
woodland 3065 clumps and groves; RhLo hora e dom.

10 225 5.1-4.0 (scrub) trees (5)8- 25-5(y/o nn- L .u.cularia
(m) (shrubs) 10 within vegetated(?)
clumps) tidal flat racemosa occ.

shrubs 3-4(8) 6c-85 lRhzera le dom.?
Lvr aculaia recently out-over;
11 323 3 sva r avery in areas nrmlly 141
323 7.3 scrub) trees 8-15 5-15 height " 0" 1 occ. or 121
Avicennia nitida
'RhizqSIhra mange

Avicennia nitida dom.
12 121 6.1 forest trees 10-15 65-85 (reproduction) Lagunclaria intr mangrove
slight racemose
L_.cularia
r-cems rare-occ.

trees 8-20 2Avicennia nitida? rare-occ.

Upland tree spp. occ.
. - - . . . . . .inner mangrove;

13 319 7-3 savana shrubs 4-8 10 Upland tree spp. occ. probably cut-over
axi rare-occ.
stems dense . nd racemose
shrubs 2-i 20 low-branching Avicennia nitid4 rare-occ.

Upland tree spp.? occ.

herbs ( 2 75-90 Similar to 528 similar
to 528


IMOHALCPLT IA





(1) (2) (3) (4)


(5) (6) (7)


undergrowth (reproduction 0.5-1.0
m) locally dense


Avicennia nitida dom.


inner mangrove; estuary head


filled


inner mangrove; slight15 443 11.4 scrub shrubs 0.5 40-80 dense and tangled Baimitima doma. ly raised, slightly
Salicornia ? areas
inner mangr ove; sIi ght ly
shrubs 1-4(5) 20-50 Avicennia nitida occ. raised, slightly sandy
i6 341 7.3 savana . ._areas but slightly
shrubs 0.5 30-70 dense and tangled Batis maritime dom. lower and less sandy
tihn 443


Avicennia nitida ab.


____________________ +


Bati s maritima


abb


similar to 341


IELOHALOPHYTIA


shrub I(2? 2-5Probably haloshrubs 1(2?) 2-5 phytic; Batis? occr
I.7 savara,- -restitdtliiiy HM L
18 355 11 (meadow) Beach grasses? of 831 (q.v.); along (Helop
(scrub) herbs 0.5 60-70 Ilomhyi abs sl
Walophytic
succulents?

Shrubs probably restricted to vicinity
stems dense and halophytic; of 831 (g.v.) ; low, HAOY
15 ?relatively slen- haophtic h
15.4a? one or two spe- flat areas; may be
19 4 13.4? scrub shrubs 3.4(6) 65-85 der, possibly
13 ?abundant multiple cies probably subett casoa ao
stems and low overwhelmingly tidal (perhaps fresh Subxe
sranohing dominant water) in tons

1-2 dense canopy; Halophytic ,dom.
4 80-90 height may vary shrubs;
considerably with Coccoloba?
scrub shrubs topographic ir- lower slopes facing sea
20 451 20.4 sregularities and (Halo
<0.1- 0-15 exposure to salt- Opuntia dillenii patches
0.5 bearing winds velaff.f- c
SBeach grasses? dore o general
meadow herbs? (0.1?) 60-100 very low and sedges? vines? vi t o 3 v
(prairie) .5(?) quite dense;
21 535 17.5 savana) evenly spaced; . generally adjacent to P
scrub) shrubs 1-20) < i even height rare ocean


0PHYTIA hytia) erophytia)



MrIA
ialophytia) cerophytia) cwopytia)









[TIA
noal ophytia) 6halophytia) srophytia)


119


forest


trees


(8)


12


70-90


(9)


(10)


(11)


(12)


17


433


11.4


scrub (woo land)


shrubs to small trees


shrubs


(5)6-8


0.5


4o-60


30-60







(1) (2) (3) (4) (5) (6) (7)


relatively dense
(o.0)2 .50-70 canopy; stems sclerophyllous dom. Wi oa neve
2 471 2 scrub (5) probably closely halophytes
4?.shrubs- paced and often
13.4? oeta 0 ees H'
0.1- 015 multiple; ind Opuntia dillenii
0.5 and/or salt pruned Ve ffpatches-fromshore

(0.5?) all shrubs and
steppe shrubs ?-2 0-5 most herbs growirE
(2esert)1herbsd<0e1 2)10 out of crevices occ. coastal cliffs (Per
1 1.7dherbs < 0.1 2-10 a on small
() ledges
brush solidly
dense to dense probably preshrubs 1-3(8) 50-80 in elongate dominantly dom.
24 445 13.4- scrub strips parallel sclerophyllous b
(24.2) (savana) to old shoreline
(8) i0-Cocos nucifera or

trees (8)10- <2 -f- r- - s oaCC.

shrubs 1-2(3) <1 halophytes? occ.-rare
10.6? steppe halophytic? of 831 (.v.) ;
25 765 17.8? scrub) graminoids; patches slightly higher ground (Haic
17.5? meadow) herbs 4(0.5 30-50 halophytic and occ. than most of 831 (Hale
succulents?
halophytic; restricted to vicinity PSbMI
26 775 10.6? scrub) b < O. 30-40 ga
ima ow shrubs?) a nd/or and occ. along shall w Eelc
succulents drainage-ways Hac
consisting principally
halophytic? of one fairly extensive
herbs (< O'l?) graminoids? area (with inclusions)
05 5 an idsor occ.-rare at inner edge of a man17.8 succulents? grove swamp; level to s MC
27 831 10.6? . . . . . . . very gently sloping; (P s
11.4? either say or dry
shrubs 1-2 (1 halophytes rare saline flats; ay be
infrequently inundated
by storm tides


fTIA




HALOPHYTIA ohalophytia)




ALOPHYTIA





)HALOPHYTIA xeroph3tia)


HALMPfYTI A DhaIOPT ia) )pltia)




MALOPHYTIA mopbytia) a,rxerophytia?)


(8)


(9)


(10)


(D4


(12)


(11)







jubescens 7 oc.
grasses ma be Paicum Maximum filed estuary heads-28 528 67.5 prairie herbs 2 85-100 planted by man vel aff. dorm. usually adjacent to IEL0PH
for cattle halophytes? 119, or
forage or hay .y.121.
I~omoa spp. patches
non-mangrove o e
(5)8 dom.extree innrtedg ofpeciesE
29 117 5.1 forest treesA(5)8-) 75-90 relatively free ieoexrmange; e ry (Melo
10(15) of undergrowbh Laguncularias
racemosa ccs
simiar t 117level to gently sloping I .I? (7)8-l0 ~ relatively free similar to 117floor of smll valley
30 116, forest trees75-90 of undergrowth out lacking dora. (cove) adjacent to
4E: 2? 15) mangrove inner m ove

considerable variation in tree Prosops
trees 6-8 20-50 density; similar julf lora? or dora. wind-sheltered coastal
31 342 72.3? savana (shrubs) to 444; quite free PitheelTobium? cove; only one oc- ITS0IE
. . of undergrowth; currency in map area (Subx
herbs 0.5-1 60-80 graminoids may be graminoids dora.
bunch grasses

gently to moderately
sloping hillsides;
not exposed to wind- IMSSC
32 232 48.2? woodland trees 15-20 40-60 dorm. blown salt; open canopy (Subx
may result from selective cutting of forest

sedge dom. lower slopes ( ca. 25
33 534 35.5 prairie herbs 0.5"1 70-90 tangled, coarse- - grade) ; mi be M SGX1
occ.- ephemeral seep area
bunch sedge Ipomoea spp. patches


beach grasses? restricted to one area;
0b herbs 0. 5-1I 80-9 5 halophytes? dm
34 5a e18.5?orairieband between 560 and NESGC
347. 3: . 151; lw veygently (Helo

shrubs 1-3 2-5 mangrove ooc. ( <57 grade) sloping Helo

may be some saltsprVy damage to at least two spe- reSUB(R
465 13.4 s shrubs 25 80-90 the extent of in- cies of probably rdm
creasing4twiggi-5sclerophyllous ward slopes facing c (Hype
creasing twiggi- halophytes (Halo
hess of shrubs


R0PHYTIA
balophytia) phytia)



ROPHYTIA




ROPHYTI A erophyia)




,RFHYTIA erophytia)



RCPHYTI A



RCPHYTIA haloph'ria) rphytia)

MYTIA rxerophytia) phytia


(1) ) 9 4


(8)


(9)


( o)


(11)


(]. )







(1) (2) (3) (4) (5) (6) (7)(80


(12)


PHYTI A




PHYTI A bytia)


(9)


(io)


(11)


Considerable diSscrub shrubs 5-10 65-95 undergrowth very versity of usually steep hillsides SUEEa
(forest) (trees) dense species and ravines

near edges of
area, resembles
37 44 72.3? scrub shrubs 342; widely spa- ro low level area between
23.4? sru (trees) 4-5(8?) p6b-80lced stem julflora? or m.52 d (eop
Probably quite ee~obium om4 lone (
free of under- and Leucaena occurrence in map
growth


shrubs (trees)


5-8(10) 160-90


resembles 465 in density


Sclerophyllous species?; at least two species


dom.


leeward (generally Wfacing) sides of broad, shallow ravines on steep hillsides; within 465 areas; possibly slightly influenced by
salt spray


commonly on, but not
39 452 23.4 scrub shrubs 3-4(5) 75-90 Leucaena uca dom. restricted to, hillsides and crests

low-branching; may occur on any well40 454 23.4 scrub shrubs 2-3(4) 75-90 small stems Leucaena glauca dom. drained site except
close spcedimmediately adjacent closly sacedto salt waiter


shrubs


1-2


>6o


dense stem spacing; most common stem diameter ca. 3 cm.


Croton spp.?


dom.


dense stem spac42 456 44.4 scrub shrubs 2-5 50-70 ing; multiple
stems and lowbranching common

shrubs 132-5 10-70 occ.-dom.
43 360 44.3 savana rough pasturecomlo
herbs (O.i) 40-80 grasses & forbs dom. conhillsides

(.5-1 and crests
bunch grasses occ. ab.


SIT OPHYTIA


38


462


13.4


scrub (forest)


458


47.4


scrub


co
0


(8)




(1) (;') (3)


23.3


(4) (5) (6) (7)


saval a (scrub) (em.)


shrubs


( 2) 4-5


20-50


vines ;(2)4-5 5-50
(li anas)
0.5-1 5-70


herbs


0.5-1


20-70


small clumps of
Leucan
more or less overrun with morningglory; area between clumps also vinecovered or, less comnonly, grassy


(8)


Leucaena 1g


(9)A


(10)


(1)2- . !
shrubs 5(8) 1-10occ

savana rough pasture
44 362 48.3 (prairie) (0.I) gasses & forbs cr.
herbs 0.5-i 75-90

bunch grasses occ.-ab.


45


46


564


354


--1. 4- L 4 4 4.


47


346


48.3? 44.3?


savana


trees


shrubs


10-20


2-3(5)


5-10


8c-9o


rairie savan/a)


shrubs


herbs


(1i)2;5(8)


LI


(0.1)
C.5-1


<1


80-95


rare-oct.


rough pasture grasses & forbs bunch grasses


domn. cc. -ab.


occ.-dom.


______________ F -


others


000.


climbing
I~ppooea
'0' hes occ.-dom.
creeping m - spp.?


rough pasture grasses & forbs


occ.-dom.


occ.

dom.


(11)


commonly on hillsides and crests


hillsides and crests; commonly peripheral to or in association
with 452


sites similar and adjacent to 232; probably results from selective cutting


(12)


shrubs <2 5-20 stem spacing and
48 770_13-7?_Steppe_ overall density oco. restricted to 831 area SUB3NO
. . . .level, probably sandy (Psam
4 7.1- intermediate be- occ.-dom. in and dry
herbs 0.5(1?) 10-30 tween 831 & 466 patches

shrubs locally taller, especially
along drainage low, level to gently
wa~s; dense stems; slooing,?slightly
49 466 47.4? scrub shrubs 1(2) 70-80 most common stem ro.hg

diameter 3 cm.; adjacent to, 831
apparently similar to .45


t + + 4 F F 4 +


WHYTI A Lophfbi) IYTIA





H









grasses dom.

50 560 67.5? prairie herbs 0.5-1 75-95 forbs occ. co rly s
. . . . . . . . . . . . .p h y t i a
bunch grasses occ. -ab.

very low an
evenly distribu- restricted to vicinity
ted (except for beach grasses?; of 831 (i.v.)" gentle S
51 572 17.5 ~meadow herbs (0.1) 75-85 faint, fine sedges?; dom. (2-5 grade) slopes;
(prairie) < 0.5 linear distribu- creeping vines? sandy?; very shallow,
tion paallel fine bullying
to slope)
in and on excavations
pioneer -r oc.
shrub 2(3r5re-oioncr. ,and construction sites SEJBXERC
52 75C 51.6? steppe shrubs 2() 515 and early ra1.ocnor
herbs 0.1-0.5 10-40 successional occ. (ab. soil generally very (Parar
52.6? herbs .1-05 IO-40(weed) species in patches) poor (topsoil removed)
ground between
desert shrubs apparently SUBXER
53 786 51.6? steppe shrubs (0.5) 2-10 bare ( mZy hIve - r.-re-occ.
savaa 1(2?) sparse graminoid arar

cover)
A 1.0"312 4 spaced (usually

trees I01 40_60 evenly) in grid Cocos nucifera dom.
54 245 24.2 woodland pattern. . . . . . casionally other level P
undergrowth sady areas near coast (Psamm
(45)1- usually kept occ.
s 3(4? open by cutting


(24,2)51.6


savara


trees herbs


10-15

0.5-1


(2)5-10 75-95


trees widely spaced (>25 m. apart); similar to 560 with addition of trees


Cocos nucifera;
others grasses


f orbs


Occ. dom.


Occ.


low hillsides near coast


trees widely trees 10-15 (2)5-10 spaced ( > 25 m. Cocos nucifera oc.
ap art)
56 332 (24.2)- savana Saccharum valley bottoms and low,
herbs (<0.1) 0-95 similar to 532of dom. ge
2-4
forbs occ.


IPHTI A thropoTMYTIA ophytia)




PHYTIA )hytia) thropophytia)


EFHTI rA aophyrtia) itropophytia)




IRPCPHYTIA aohalophytia)


PARAMTHROPO?YTI A


55


3.28


i


(1co


( 2)( 1) ( )


(8)


(9)


(lo)


(11)





(1) (2) (3) (4) (5) (6) (7)


-r I1 I____I__I___I__________I___I


359


49. 6- 50. 651.6-55.35Q.3-59.259.4-59.7


350 58.3-59.259.4-59.7


532


savaria


savana


trees shrubs


57


trees and shrubs


herbs


various various


i f


various various


various various


various I various


-4-- +


various various


includes small ( < 1 acre) fields planted to garden crops; occasionl fruit trees


similar to 352, exclusive of
fields in garden crops


shade and ornamentals


occ.


ornamentals occ.


garden crops,
"lawn" grasses


shade and fruit trees; ornamental shrubs and trees


"lawn" grasses


occ. dora. patches


ccc.


occ.-dom.; patches


m i ii i i 4i i4


54.5


prairie


herbs


(< 0.1)
2-4


0-95


height and cover vary annually and regularly


S.accharum
of f I crarum


Ipomoea and other "weeds"


dom.


occ.-ab.


around dwellings


around dwellings


valley bottoms; occasional hillsides


PARA0nTOPM YI A


Saccbham
uneven height and or forage and/or valley bottoms only;
PARANP
60 536 54.5 prairie herbs 2-4 85-95 spacing;mayfbe haygrass or doam, probably porly (HeIo
poor ctXOP of some otherdrandgod
sugar car tall coarse
graminoid.
pasturelow, gentle slopes;
49.6-50.6 meadow < even turf pdo. commonly situated be61 622451.6 (prairie) herbs (1? 90-100grasses teen 532 and 560

in general vicinity of
62 644 49.6-50.6-<0Idelns te cu
62 644 51.6 meadow herbs (O.1 75-100 dense, even turf 'lawn" grasses d lm.lings, her
nationn dumps, roadsides 51.6 <0.1*

63 648 49.6-50.6 meadow herbs < .I 6085 irregularly dense 'lawn" grasses dam.-;n ccpe
51.6 (0.5) turf patches dw1 nrs
around non-resident ial

64 862 52.6? desert herbs <0.1 <10? ground apparently 'weeds" rare-occ. buildings and on
(0.5)bare recent excavations


65 894 0.0 desert n.a. n.a. n.a. ground bare n.a. n.a. astad v


vpytia)












TM0PMDYTI A


herbs


0


. - i


(8)


(9)


(lo)


(12)


(11)











MASaR-TABLE E( 1). Guide to symbols used in Master-Table E and on Maps 2, 3, ad7.


Airphoto-tye Key numbers on
Master-Table E

1. FORESTS


117 118
119 121 141 151


225 231 232
245


319 323
332 341
346 350
352 354 355 360
362


423 427 429 433
441 443 444 445 44
451 452


29 30 14 12 9
3
2. WOODLANs


3. SAVANAS


4. SORUBS


10
5
32 54


13 11 55
56
16 31
47 58 57 18
43 44


8
7
6
17
4 15 37

20
39


Airphoto-type


Key numbers on Yaster-Table E


454 40
456 42
458 41
462 3
465 35
466 49
471 22
475 19
5. PRAIRIES


528 529 532
534 535

564
572


622




750
765 770 7 5
7 1
786


831
862 889 894 895


6. MiAImmS


7. STEFP S


8. IESERTS


28
34 59 33
21 6o 50
45 51


61
62 63


52
26
23
53


27
64
2
65
1








85


M.STER-TALE F. Life-forms according to Braun-Blanquet (1932), to
Raunkiaer (1934), and to Dansereau (1945, 1957a, 1959). Two examples
from Eastern North America are given in each case: the first one a
native plant, the second a cultivated one; the third example is
from Puerto Rico.






P PHAERCPHYTES: trees, or woody plants, with regenerating buds high (at least
50 cm.) above ground

Pg meahaneroyhgtes: tall trees, more than 25 m. high
Ex.:-Ulmus amercana, Picea abies, Dacrodes excelsa

Pm mes haeraohytes: small trees, 10-25 m.
Ex.: Ostrya v iniana, Morus alba, Avicennia nitida

Pp micrtes: tall woody plants, 2-10 m.
Ex.: Crlsont, ~ ~ g riAnnonaLabra

Pn nano 1aeryh tes: low shrubs, 0.5-2 m.
Ex.: S raea7laifolia, Berberis vulgaris, Croton poecilanthus Ps P. Ps clirubing_ phaerqkh ts
YCeTilT5t~rfui scandens, Wisteria sinensis, Rhabdadenia bliflora


Ch CH4XAFPYTES: plants with regenerating buds not much above soil level
(less than 50 cm.)
Ex.: Lcoodiunannotinum, Vinca minor, Cactus intortus

Chin moss chama2Ie_ es: bryophytes and lichens


H M1CRYITOPTY TE S: plants that die down to soil level during the unfavorable
(cold or drj) season

Hspose hemio to hytes: with a well-developed stem, branching quite
prof- e .y
Ex.: Euatorium ru.osum, Capanula media, Pilea k-.Wii

Hr rosette he itop es: with a crown of leaves at soil level
. . M __ _ . . 10- " " i -ui
Ex,-, SaXifraaw e ,Primula floribunda, Bronelia nR _y

Hc cespitose heicr'tophytes: with many tufts or short branches arising
at ground lev
Ex.: Danthonia s~cata, Poa2ratensis Bouteloua a zrica

Hg decumbent or climbin- hemcrt yhes: with a weak plagiotropic stem
Ex.: P onumci-__ed ome, acoerulea, Comnelina elegant







86


NASIP-T f HlE F (continued)




G GEOEYS: plants with regenerating buds well under soil level
Gb bulb-. p tes
Ex.: Alliur-tricoccum, Tuli __snerina, Hymenocallis declinata

Gr rhizome.o2 te es
Ex.: Smilacin raceraosa, Iris gerrianica, Canna Rlauca

Gg root _eo hy e s
Ex.: A-sc aassriac, Dahlia variabilis, Alpinia antillarum


Thl TMOPHEs (a=nuals)
Ex.: IFitiens ca ensis, sum saxatile, Cakile lanceolata


Th2 THEPOPHYTES (bie nals)
Ex.: Oenothera biermis, Althaea rosea, Leorurus sibiricus


E EPIPHYTES. plants that rest upon the boughs or stems of other plants
Ex.: Usnea !orissimoa and mnwn other lichens, C tl abiata,
Tillandsia recurv-i



HH HYDROPHYTES: true aquatics (subdivisions in this category are according to
Dansereau 1945)

HE'S free-liing, submerged, or emered plants
Ex.: Ceratophyllu -demersum, Salvinra"natans, Pistia stratiotes

HHf emerged aquatics with broad leaves
Ex.: Pontederia cordata, Nelumbo lutea, Sagittaxia lancifolia

HHj emerged aquatics with thin (reed-like) stems and/or leaves
Lx.: sci- - sacutus, -e-papis ,--X:s _-s

HHn floating-leave& rooted aquatics
Ex.-. N iH d-s-D, aema-ohea z nzibarica, N yTTmLw e a

HHv submerged aqyutics with ribbon-like leaves and/or stems
Ex.: Vallisneria arnerT n, Potaoeton crisus7aJass ia testudinum

H~r submersed rosette Rlants
Ex.: Lobelia dortmanna, I oetes sris, Codocea rmmatorum

HHt submersed an s or he_tes
Ex.: Najas flexilis, Najas a l uensis, Najas marina

HHa ephy-tic or epilithic aquatics
Ex.: Font-riais spp., Podostermon spp., Helianthium tenelluum







87




MASTER-TABLE G. Leaf-size classes (.aunkiaer 1934).



SmoArea Cat~o

1 1 less than 25 mm.2 LTpYLL

2 n 25 - 225 mra.2 NANO L

3 m 225 - 2,025 mm.2 1RCPHYLL

4 NM2,025- 18,225 . 2 ISOPHYLL

5 W 18,225 - 164,025 mm.2 1AAR OPY L

6 IM more than 164,025 mm.2 EGAPHYLL













MASTER-TABLE H. Floristic elements in Puerto Rico, based on geographical distribution.


Symbol

E Endemic: only in Puerto Rico (IR~pstonea borinquena)

A Antillean: also found in other parts of the West Indies (Gundlachia cormbosa)

C Caribbean: Antilles and ContinentalCentral America, Florida (Bursera siraruba)

G Northern Neoroical: Caribbean and Northern South America (Casearia guianensis)

B Neotropical: tropical America (Acacia westiana)

P Pantrojical: tropics of old -and new world (Albizzia procera)

W Cosmoolitan: (e )
N Naturalized: (Albizzia lebbeck)

0 Unk nown
X Planted: (Castilla elastica)







88






MASTER-T.AHLE J. Diaspore types, as proposed by Dansereau & Lems (1957).

(D" means diaspore.)




D. does not disarticulate from the parent plant before being deposited at the site of further development. (Parent plInt may be dead or alive at
that time.).e. . OCHCRE

D. disarticulates from the parent plant before the dispersal phase.

D. veri voluminous in relation to the actual reproductive part, consisting
of a loose spherical framework. . . . . .CYCLCHaE 2

D. consisting for the main part of the reproductive organ.

D. with scarious or stiff appendages.

Appendages thin, light, often flexible.

Appendages scarious, wing-like, or saccate. . PTEROCHORE 3

Appendages long, hairlike, or plumose . .POGONOCHRE 4

Appendages short, stiff, spiny, or glandular, adhering to rough surfaces. . DESIC1 5

D. without appendL3es (except for arils).

D. with juicy or fleshy outer layers.S OCHCo 6

D. with hard outer leer.

D. small or light enough to be carried by a breeze.
.*. SPOROCHORE 7

D. too heavy to be c~.rried by breeze.

Parent plant without mechanism of expulsion.

D. light enough to be carried by wind.
. SCLOCOE 8

D. very heavy. . . . . . .B ROCOHRE 9

Parent plant with mechanism of expulsion. . . BALLCCFORE 10








TARE I. The vegetation zones of Puerto Rizo and their prirnipal correlations and characteristics.


VEGETATION
ZONE
(Ma 1)


GEOGRAPHICAL REGIONS (Pico, 1962)


EAN ALTITUIES


PHYSTOGR_HY


VI MONTANE SCRUB 9(p) 10(P) 1000 + peaks


V MONTANE FOREST


IV LOWER MONTANE
RNFCREST


!III SEIIDECTX.OUS
FOREST


Ha FILL SCITOB
subzone


II SEASONAL
EVERGREEN
FOREST


7b( )
I.( ) 1 o'p)


79(p)
9(r)


39(p)




5a(p) 5b, c, d, e 6(p)


lb(p)

7a(p) 7b ,c


75o-.Ioo


200- 750


50- 350


50- 200


50- 200


up-er slopes


foothills, valleys, slopes, ravines canyons


foothilIs, valleys


tops of ills, cliffs, sharply eroded


hills, of var able slope


PARENT -M.CR I AL


grani te


mostly granite and shale, serpentine


mostly granite and shale


eroded limestone, granite


limestone, very sharply eroded,
exposed


limestone, very permeable


vIll
(Thorntlwaite) CQoo, very moist, AB' r, BB' r


warm to cool, very moist AB'r


warm, moist BB'r BA'r


ECOSYSTEMAT IC PEGIXIES
(Master-Table B)


subhygrophytia


- A


hygrophytia lithophytia


hygrophytia lithophytia
mesoxerophytia paranthropophytia


VE TATION STRUCTURES
(Master-Table C)


scrub


forest, savaa,
scrub, grassland


forest, savanna, scrub, grassland


4. __________


very warm with dry seas on CA'r CA'd Die d. EA'd


very warm, moist


BA'r BA'w


BB'


subxerophytia lithophytia chasmophtia

subxerophytia lithophytia chasmophytia


hygrophytia
mesomerophytia lithophytia
chasm phytia


forest, savana, scrub, desert


scrub


forest, scrub


hygrophytia
plains hills, very warm, helophytia
aL(WLA25-ssediments, moist subxerophytia
RATkNFREST cla I5(p) 25- 200 0 u. orcrops BA' r paranthropophytia scrub, prairie
2a 2b(p) lithophytia
2c 2d . .-. . .__ - - -_ _ _ -
2b )g elvey warm, helophytia

Ia LITTORAL 4d ) flat, shoeline rock, san moist to dry halohydrophytia desert, meadow
subzone 0- 50 fandbays; sltI BA'r CA'd psam hia steppe, scrub
neClisi, ffs CA'r DAI'd lithophytia savane, forest
som cliffs EA' d chasuophytia





TABU II. Charx-cteristics and plant comtrn.un-ties of the littoral subzone (Ia).


ECOSTSM


LAMICEE


SEA


LAGMONS


FM ATIONTYPE (aster Table C)


BI1DIE
(MasterTable B)



HALCIYMO-


VEGETATI CN-TYE or PLAIO-CCU INITY (Milaster-Table D)


1.5 kelp tangle


DaaINANT SPECIES


Lamiriaria sp.


POND saltwater pond 2.6 algal meadow Ulva lactuca
meadow . .
SHALLOi' saltwater shallows 3.& turtle-grass bed JTnalassia testudium


coral cliff


salt-washed rock




sandy salt-flat


mangrove


saltersh


crust


scrab


meadow


scrub


desert


scrub


safna forest


eadow


4.0 marine algal crust


o.4 black-mangrove scrub

4.6 fucoid bed

5.4 red-mangrove scrub


Enteromorpha sp.


Avicermia nitida Fucus sp.
Turbinaria turbirata ,Rhizophora mangle


b.4 black-manrove scrub Avicennia nitida

00 e( ne) (none)

54 red.mangrove scrub Rhizophora mangle


6.4 black-mangrove scrub 7.4 mnid-angrove scrub


7.3 mangrove savana

5.1 red-mangrove forest


Avicenria nitida Rhizophora mangle Avicennia ritida Laguncularia racemose Rhizophora mangle


.I b lack-mangrove foresw Avicennia nitida


7. mixed-mangrove forest


9.6 glasswort atl 10.6 sea-purslane mat


Rhi zophora mangle Avicennia nitidca Laguncularia racemosa Salicornia sp. Sesuvium portulacastrum


sBatis .maritima
scrub 11.4 saltwort scrub Avicernia nitida


ai 12.5 salt-fern prairie


Acrostichum aureui Acrostic-hur danaefolium


HABITAT


OPO WATR


marine benthos


SEEFS


prairie


PHYTIA


HALOPHYTIA


SCAP


PLATR


TIDAL FLAT


rocky


or


s ilty


TIDAL


BELTS


prairie











littoral platform


SUPRATIDAL BELTS


CLIFF


FETROHALOPHYTIA


BE17H


TLVEE


salt-spra cliff


liiestone pavement


outer beach


inmer beach




marine saiy


ridge


scrub steppe


caesert eadow tundra






scrub


desert meadow


prairie steppe


13.4 bay.-cedar -~


13.4 bay-cedar -horse-bush scrub 3.7 ba -cedar steppe 14.7 spurge-sedge mat


14.8 spurge desert 14.6 salt-sedge meadow

Icrr button-mangrove
1" - r-tted scrub
low; but tocn-mangrove15.4 cactus scrab 15ahigh button-mangrove15.4a cactus scrub


16.4 sebiucan-tacielo
thornscrUD


17.8 sea-rocket strip


lO~ sea-.uurslane mat


17.6 moring-glory festoon


b each- gr ass - -morni'ngglory prairie

il.5 salt-grass prairie 18.7 salt-grass steppe


Suriana maritima Gundlachia corymbosa Borr chia arborescens Spartina patens

Euphorbia buxifolia Fimbristylis spadicea

Fimbristylis spadicea


4 ----~---


Conocarpus
erecta


Caielanceola


--~ Iporroea pes-caprae


Cactus intortus

Cephalocereus royeni

Lemaireocereus hvstrix Plumiera alba


Sesuvium portulacastrum


Ip omoea-S por obo lus-Spar t3ina Canavalia Spartina patens


meadow 19.b beach-grass sward Sporobolus virginicus

20.4 sea-grape scrub Coccoloba uvifera

scrub 21.4 meray-nmaray scrub Dalbergia ecastophyllum

22.4 snake-bark scrub Colubrira ferruginosa


woodland


24.2 coconu-t grove


25.2 Australian-pine screen


Cocos rxucif era


Casuarira equisetifolia


ROCKY SLOPE


strand


PSAM1OHALOPHYTIA PARMTZ OPOPHYTIA


-I I


-- - .r








Ir I. -- - , Ir


PS IOPHYTi A


31.5 trompetlla-gras rab.


32.5 cattail marsh


Coiubrin ferruginosa Leucaena glauca Termialia catappa Eichhorrda crassipes Pistia stratiotes NYprnhaea a. pla Paniarum purpurascens Ericchloa polystachya


Panicam aquaticum Rjyrenachme armplexicaulis T,'ha domingensis


33.5 reed-grass marsh Phragmites communis

34.5 spike-rush marsh Eieocharis spp.


35.5 sedge marsh 3c. 5 rivercane brake 37.4 icaco scrub


Cladiun, Scirpus, Carex

Gyneriu msagittatum Chrysobalanus icaco


Anmona glabra
woodland 3 .2 pond-apple amp oncarpu erecta

forst 39.d teoapu wmpPooarpus off cnali


DM1S


ESTUARIES


22.4 sra.,e-bark scrub 23.4 Leucaena thicket M.4 Xalabar almond scrab 27.6 "ater-hyacinth mat 2.o wter.oettuce ,at 29.6 w.teriir omat



30.5 pra-cari-gracss mat


PHYTI A


EELOPI{YTI A


RISE


Sid ATE


RIVER


LAE


4-OODPLAIN


eow prairie steppe Meadow


prairie


prairie scrub


terdune open mter


marsh


river-


17.o morring-glory festoon Ipomoea pes-aorae

16.5 sal-grass prairie Spartina paten
Iw.7 salt-r:,ss steppe �.6 beach-grass sAxrd Sporobolus virginicus

20.4 sea-grape scub Coccoioba uvifera


f or e st


39.-1 Pterocarpuns swamp


Pterocarpus officinalis








TABLE III. Chact.aristics and counties of the lowland r'dnforest zone (I).


(masterTable B) LIMNOPHYTIA


marsh


FOIRMATIONTYPE (M as ter Table C)





Meadow


ECOSYSTEM





lake


or


pond


VEET ATION- TYPE
or PLANT-CMAJITY
(Master-Table D) 27.6 water-hyacinth mat 28.6 -ter-lettuace mat 29. vaterlily mat 30-.5 para-carib-grass mat


crust 40. duclkweed crust


prairie


swamp forest


31.5 troupetilla-grass marsh


D INANT SPECIES Eichhornia crassipes Pistia stratiotes Nymphaea ampla


Pai cure purpurascens Eriochloa polystachya


Lemna perpusilla


Panicum aquaticum Hymenachne amplexicaulis


32.5 cattail marsh Typha domingensis


33-5 reed-grass marsh Phragmites communis


34-5 spike-rush marsh Eleocharis spp.


36.5 rivercane brake


39-1 Pterocarpus swamp


IGynerium sagittatum


Pterocarpus officials


river-flat steppe 41.7 spider-flower flat Cleon~ spinosa


LANDFCOM


LAKES







and RIVERS


prairie


HELO










PHYTIA


HABITAT


LAME AND


POND


FLOODPLAIN


river-f lat


steppe


41.7 spider-f Iwer f lat


Cleome spinosa






RYGRO- vmlPHYTIA DRAINED


PHYTIA


PAANTHRcPOPRYTI A


UPLAIND


LANDM
L ZD


CTLTIV AD

L AND


O COPIED LAND


forest


abandoned


pasture


tilled & cUltivated land




stabilized







landscaping


forest


woodland





scrub


42.1 lowland rainforest 43.1 trunpet-wood forest


AA4.2
44.3__jungle 44.4
45-4 guava scr I


A6.4 pepper scrub 47.4 firebush scrub 23.4 Leucaeoa scrab


23.2 Leucaena moci Land woodland
46.2 second-growth woodland


savana


saow



steppe

prairie wood land!


savaa


scrub


woodlanl


steppe


46.3 second-gromth savana


49.0 St. Augustine grass sward 50.6 carpet-grass smrard


Di ospyros ebenaster Mammea americant I-lmi!kara ni tidla

Cecropia peltata


Casearia arborea Andira inermis
Piper aduncum Abrus praecatorius Ipomoea spp.

Psidium guava


Piper adncum Croton lucidus

Leucaena glaica Spathodea campanulata Randia aculeata Didymopanax morototoni


Stenotaprmum secndatni Axomopus compressus


51.6 cerrillo-grass swaxr Sporobolus indices

52.6 dr-beardgrass sward Andropogon gracilis
53.7 pineapple fiel& Anaras sativa

54.5 caufiel& Sacchax-am officinarum
24.2 coconut plantation Cocos nacifera


25.2 Australian nine screen 55.3 orchard 5b.3 gardens & parks


59.4 hedges 59.2 screens


59.7 roadsides


Casuariia equisetifolia Citrus limonium 1angifera in-ica (ornaentals)


Hibiscus schizopetalus Allamnnda cathartica Bambuisa valgaris

Casuarina equisetifolia Poinciana regia

Ceiba pentandra
Albizzia julibrissin Poinciana regia


UPLANDS


J 414. 'L -


--- --- I






T�BUE IV. Characteristics nd co, runities of the seasonal-evergreen forest zone (Ii), and the I'l-scrub subzone ( Ha).


ILANDPQGRM


INTERVATIS



and SLOPES


SLOPES

and

LATE ALTS


ISYMGI-(MasterTable B)


HYGRO-


PHYTT A MH-S0XCERQPYTI PARANPHYTIA


HABITAT


DRAINED


lol-SOUEROYTI UPLAND


ECOSYSTEM


forest


FORMATIONTYPE (aster Table C)l


forest


VEGETATION-TYPE or PLANT-CaMUNITY (Master-Table D)


42.1 lowland rainforest


DIXDNANT SPWISS


Diospyros ebenaster Mamxea americana Manilkara nitida


43.1 tnuipet-wood forest Cecropia peltata

44.1 jule Andira inermiis
Roystonea borinquena


60.1 glateado forest


Coccoloba laurifolia


49.6 St. Augustine grass sward Stenotaphrum secundatum 50.6 carpet-grass suard Axonopus compressus
pasture Ymeaow 51.6 cerrillo-grass suard Sporobolus indicus
52.6 wiry-beardgrass swrd Andropogon gracilis


cultivated land


abandoned


fields plateau scrub


scr-ab


woodland


savana


54.5 canefield


Saccharum officinarum


pepper scrub Piper adncum


47.4 firebrush scrub


4b.2 second-g rowth woodland


4b.3 second-growth savaxa


___________ + I


scrub


0.4


craowmod scrub


Croton lucid"us


Randia aculeata Diaywopanax moratotoni


Gymnanthes lucida


Clusia rosea
CLIFFS CHAS4OPHYTIA STEEP ROCK shaded cliff face scrub 61.4 balsanfig screen nthuium acaule

savana 62.3 llume-gumbolimbo savana Gaussia attenuata
SUB 0 - NARROW 624 me-gubolimbo scrub ursera simaruba
SPURS LhmeSTONE limestone64scb
crest Tabebuja heterophylla
PHYTIA'RIDGE63.4 roble prieto scrub Plumiera alba


\0t


-.-- - - -L-







TrBIE V. Characteristics and communities of the se- -deci0uous forest zone (III).


rRIIT -FM':'ION - VEETAT ION-TYII
LADFCR,'i(Mster- HA B!TAT ECVIM SS M =I F (ivfs- or PL IN - -"SDIMMN NT PE IE
Tal )Iter Table C) (laster-Table D)
ortCJI


I.S0XER0PHYTIA SLBXEROPTI A


HYPZRXER0P YTI A



PAAN





THROPOPEYTI A









HYPEXEROP YTIA


IFAIED LIiESTONE
GRAVEL & LLSTC1

LIUSSTONE PARENT




RAVEL, SMST

NETINE
SCHIST

LATiTIC PLATEAU




OPEN







FLATS


forest


forest


woodland


64.


84.2


bucao forest


buc3ro woodland


--I oucica


buceras


open savan savara b4.3 gumbolimbo savana Bursera sinaxuba

rub . seb ucan- tachuelo Cephalocereus royeni
thosrscrub Pitetia aleaa
Saqir s -65.7 wire-grass steppe Uniola virgata
steppe
66.7 grama-grass steppe Boutelaua heterostega

meaiow 51.6 cerrillo-grass sward Sporobolus indicus

7.5 Guirea-grass prairie Paxnium maximum
prairierArundinella confinis &5 r Andropogon semiberbis

Leptocoryphium laratum
PastureMeadow nAristida portoricensis

steppe 6-.7 grama-grass steppe Bouteloua heterostega

prairie o7.5 Guinea-grass prairie Panium maximum

meadow 51. cerrillo-grass sward Sporobolus indices

steppe 70.7 angleton-grass sward Andropogon armulatus

Chloris inflata
mradow 71.6 ptaragto meadow Pennisetirn ciliare


savant


savana


72.3 e suite savam. 72.3a bucaro-mesquite savana


Prosopis julflora


Bucida buceras Prosopis juliflora


ROCKY










HILLS


SAND


and SILT FLATS


13. --* -, -


- --- lv


I