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PUBLICATIONS

OF

FIELD MUSEUM OF NATURAL HISTORY

BOTANICAL SERIES VOLUME XVIII

THE LIBRARY OF THE

MAR 151939

UNIVERSITY OF ILLINOIS

CHICAGO, U.S.A.

1937-1938^

PRINTED IN THE UNITED STATES OF AMERICA BY FIELD MUSEUM PRESS

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

FOUNDED BY MARSHALL FIELD, 1893

VOLUME XVIII THE LIBRARY OF THE

MOV 5 -1937

UNIVERSITY OF ILLINOIS

FLORA OF COSTA RICA

PART I

BY

PAUL C. STANDLEY

CURATOR OF THE HERBARIUM, DEPARTMENT OP BOTANY

B. E. DAHLGREN

CHIEF CURATOR, DEPARTMENT OF BOTANY EDITOR

PUBLICATION 391

CHICAGO, U.S.A. OCTOBER 12, 1937

PRINTED IN THK UNITED STATES OF AMERICA BY FIELD MUSEUM PRESS

560.5

CONTENTS

PAGE

Introduction 5

Plant geography 6

Atlantic tierra caliente 10

Pacific tierra caliente 16

Temperate region (tierra templada) 21

Cold region (tierra fria) 29

Relationships of the Costa Rican flora 43

Botanical exploration in Costa Rica ... 45

Plan of the flora 57

Acknowledgments 58

Bibliography 62

Systematic list of families, genera, and species 63

FLORA OF COSTA RICA

PAUL C. STANDLEY

INTRODUCTION

Preparation of manuscript for the Flora of Costa Rica has been the most agreeable botanical work that the writer ever has under- taken, for two reasons. No other area of equal size anywhere in America possesses so rich and varied a flora, and none in North America is at all comparable in these respects. It is improbable that in any part of the earth there can be found an equal area of greater botanical interest. In the second place, work upon the flora has enabled the writer to relive many happy days spent in Costa Rica in 1924 and 1925-26, while making collections and becoming acquainted with several separated and representative regions: the Meseta Central; the Atlantic and Pacific coasts; the Province of Guanacaste, so unlike and yet in some respects so similar to central Costa Rica; the volcanoes; the Canton de Dota; and even the alpine paramos of Dota, the only ones, even if small and insignificant in comparison with those of the Andes, that exist in North America.

Working over these collections revived a host of memories of inspiring days spent in mountain and lowland forests, memories of the most varied kinds, all happy and pleasant ones. It is truly remarkable that in traveling so many miles, afoot, on horseback, and by other, often primitive means of transportation, there should have occurred no unpleasant incidents, nothing more embarrassing than minor failures of modern rather than primitive machines of transport. In few countries of the world, I believe, would it be possible to travel so much and find only pleasant and ever varied scenes, and be received everywhere with simple and sincere hospi- tality. Elsewhere in Central America the writer has always received most sympathetic treatment and most kindly hospitality from rich and poor, but in Costa Rica even the customary and expected courtesies have been exceeded.

It is unnecessary to expand this theme, for the writer has always been extremely enthusiastic in speaking of Costa Rica, and it would be difficult to develop the subject adequately. Suffice it to say that the country possesses a peculiar charm, in part based upon its great

6 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

natural beauty, with scenes that vary from densest tropical rain forest and jungle to semi-desert, the wildest mountain scenery, with steep scarps, majestic volcanoes, often with smoke issuing from their summits, lovely lakes, swift streams of clear, cold water; and alder- encircled pastures that recall the hillsides of New England. Costa Rica never has seemed to me a foreign country at all. Its atmosphere is homelike, and one feels immediately at home in any part of it. Costa Rica's greatest resource is its people. Ask any Central American outside Costa Rica what is the best part of Central America, and the answer is always the same: Costa Rica. This is not only because of the great beauty of the land, but on account of its inhabitants, who are celebrated for their good schools and stable and truly democratic government. It is a land where no one is very rich and no one hungry. The term that best describes the Costa Ricans is the Spanish word humildes. In its best sense this expresses their naturalness, their dignity without affectation, their contentment, and their happiness. One can not fail to be impressed with the intelligence of the mass of Costa Ricans, and the high learning of many of them, which always is combined with a lack of affectation that compels the respect that well-based educa- tion always deserves and receives.

PLANT GEOGRAPHY

Of the independent countries of the earth Costa Rica is one of the smallest. In flora it is one of the richest. Its area is approxi- mately 18,400 square miles, about that of the State of West Virginia, which it somewhat resembles in rugged terrain, but upon an exag- gerated scale. Bounded on either side by the Pacific and Atlantic (really the Caribbean Sea) oceans, it is scarcely more than a hundred miles in width, its longest axis running from northwest to southeast. Somewhere near this axis, but rather closer to the Pacific, it is transversed by the great cordillera that extends near the Pacific Ocean for almost the whole length of the two American continents.

Such, in brief, is the geography of Costa Rica. Land at either of its ports, but especially the Atlantic one, and look about you, and the geography seems far from simple. After spending a little time in the country, and trying to travel over it, even to the more accessible and more densely populated regions, you will agree that practically the geography is highly complicated.

So difficult is travel in Costa Rica, in spite of the railroad extend- ing from coast to coast, and so many are the regions never visited

FLORA OF COSTA RICA 7

by a botanist, even for the most desultory collecting, that it seems presumptuous to attempt at this time to offer a flora of a region so rich floristically as this. It is for that reason, principally, that the present flora appears as an annotated list rather than a more elaborate work with keys and full descriptions, for the time has not arrived when an adequate flora of the country can be written. There is no doubt that the great majority of Costa Rican plants have been collected, and that most of the important ones are described, but to judge from what is known of the flora, it is certain that the number of species will be greatly increased, far beyond even the astonishing number enumerated on the following pages.

Why should so small an area possess such a vast number of plants, a number much greater than exists in any of the other Central American countries, most of which are much more extensive in area? Why should Costa Rica have such a stupendous number of ferns and orchids, groups in which few other tropical countries, no matter how great their area, can rival it? Why is it that in America such wealth of plant life can be found in no other area of equal size, unless it be in portions of Colombia, Ecuador, or Peru? These questions may be answered satisfactorily by an under- standing of Costa Rican geography and climate.

In few tropical regions is it possible to find within such small compass so varied physical conditions. The surface of Costa Rica ascends from sea level to about 3,900 meters' elevation, an altitude above which, at this latitude, little plant life could be expected. Much of the provinces of Limon, Alajuela, Guanacaste, and Pun- tarenas consists of plains that rise gradually toward the mountains. About a third of the country is formed by high hills and mountains, of the most rugged character and the most involved structure. While upon the map one sees a neat chain crossing the country obliquely, in actuality this chain is much interrupted and bewilderingly complex in geography. All or most of the mountains are volcanic in origin; several volcanoes are almost constantly or intermittently active. Like most volcanoes, their slopes are steep and exceedingly rough, making travel over them tedious or even impossible. So difficult are conditions of travel that probably no one person has ever seen all the country, even superficially, despite the fact that so many Costa Ricans make long and frequent excursions on foot or horseback over their country. So difficult are means of communication that some of the richest agricultural lands have never been occupied. What motive can induce settlement in the productive valleys of

8 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

El General, since all produce, to reach a market, must be transported by a long and difficult route to the coast, then by boat to a port, and by railroad to the interior, or else carried on men's backs, or driven on the hoof, across the highest mountains of the country to market? Perhaps the airplane will solve this economic problem, as it has already done to a slight extent. The expense of building roads over such difficult territory is so great as to make them impractical for many years to come.

The writer has spent two winters in Costa Rica, making innumer- able trips by various means of transport, and visiting as many regions as could be reached during the months spent there. As he looks back at the collecting thus accomplished, and the list of localities visited, it seems that a great deal of traveling was done, but a glance at the map shows the routes as only thin lines, almost lost upon its surface. Few other botanists have seen so much of the country, and, in spite of all this, we know that botanically Costa Rica has received much more than its fair share of exploration, as compared with most other parts of tropical America! There is much still to be learned about the flora of the tropics.

The plains of Costa Rica, in the departments of Limon, Alajuela, Guanacaste, and Puntarenas, have a sparse population. On the Atlantic coast the chief and almost the only industries are the pro- duction of bananas and cacao, both of diminishing importance in late years, although flourishing formerly. It was in Costa Rica that the export banana trade, now of such huge importance, had its birth. The Pacific plains, in Guanacaste and Puntarenas, are devoted to stock-raising, although in limited regions agriculture is followed.

The great majority of Costa Rica's population is concentrated in the upland regions, chiefly within a small portion of the depart- ments of San Jose, Cartago, Heredia, and Alajuela. This elevated region, coinciding roughly with what is here called the tierra templada, is known commonly as the Meseta Central, although at times the latter term is restricted to the great valley between the two chief mountain ranges.

The high mountains of Costa Rica are arranged in two great chains running almost parallel. The northern and better known is formed primarily by the four great volcanoes, Turrialba, Irazu, Barba, and Poas, that overlook the Meseta Central, from any point of which at least one of them is visible. Northwestward the chain is continued by the low Sierra de Tilaran and the volcanoes of

FLORA OF COSTA RICA 9

Tenerio, Rincon de La Vieja, and Orosi, all of which except the first are quite unknown botanically.

South of this chain, and separated from it by valleys and low mountains, extends the shorter cordillera of Dota. This includes such high peaks as Cerro de La Muerte, Cerro de Las Vueltas, and Chirripo, the highest mountain of Costa Rica, and continues with little interruption to the Volcano of Chiriqui, in Panama, whose flora is naturally similar to that of the Costa Rican mountains. The flora of the more southern range, although it is little separated from the northern one, seems to possess a much closer affinity with that of the Colombian Andes.

The dominant factor in plant distribution in Central America is rainfall, moisture conditions being more or less uniform through- out corresponding parts of the region. Upon the Atlantic coast rainfall is heavy, in some places reaching 180 inches per year. Of greater importance is its continuity throughout the year, for while in some months rainfall is scant, there always is enough for luxuriant growth of vegetation.

On the Pacific coast rainfall is scarcely half as great, and periodic in distribution. During the invierno, May to September or October, the whole rainfall of the year is received. During the much warmer verano, coinciding with winter months of the North, there is little rain or more often none at all. For only half the year is there suffi- cient moisture for free growth of vegetation. During the dry months many plants are dormant, and many trees and shrubs shed their leaves. Crops can not be grown here during the dry season, but on the Atlantic coast they may be produced at all times of the year.

The mountains constituting the continental divide are the factor governing distribution of rainfall. During winter months they are an effective barrier against rain clouds driving inland from the Caribbean. These clouds are halted at the summits, but drift across upon the Pacific slope for a short distance. Ascending the Pacific slope of one of the central volcanoes during the winter, especially in early morning, it is easy to see how the moisture is distributed. On the lower slopes in March the fields are dry and brown, but at a certain level the dust in the road disappears, and the ground becomes progressively wetter and wetter. Immediately one notices that every tree is laden with orchids, ferns, and other epiphytes. It is to this line that clouds and mists descend at night.

Other factors affecting plant distribution are temperature and wind, the former dependent principally upon elevation, but partly

10 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

also upon rainfall. The mean temperature, in general, decreases as one ascends the mountains. The lowlands are hot, although seldom if ever so disagreeable as a hot summer day in central and northern United States. At middle elevations the temperature is delightful through all or most of the year. At San Jos£ the days may become rather hot during the dry season, but the nights are cool or cold. At Cartago, which has a considerably greater elevation, only the trite term of perpetual spring can properly describe the climate. There can scarcely be a more perfect climate, except that the nights are rather too chilly for one not born to the climate.

At 1,800 meters or higher the climate is really cold. The writer has suffered more from cold in Costa Rica than in all his life in the United States, and it is hard to recall a comfortable night spent in the uplands, even in a room almost hermetically sealed, which, unfortunately, is usually far from the case. Upon the Cerro de La Muerte and Cerro de Las Vueltas thin ice often forms at night. Although the temperature is sufficiently low for snow, this phenom- enon seems to be unknown in Central America, although a few possible exceptions are recorded.

The only places where wind is an important factor are the tops of the highest mountains, principally those of Dota, and there it is questionable whether wind is so important as temperature. In the high mountains, especially at night, winds are all too frequent, often reaching the force of gales. Even at such a low elevation as Tilaran, in Guanacaste, the wind howls all night long, and it is disagreeable to be on the road at night almost anywhere in the mountains, especially if rain is falling.

Most celebrated for wind and cold is the Cerro de La Muerte, whose name indicates its somber reputation. A rather important trail, crossing it from El General to Santa Maria de Dota, has long been an important thoroughfare. Because of its dangers, the govern- ment has built shelters where travelers may take refuge from cold and wind if overtaken by night, or sometimes even in daytime. Horses seldom are taken across the paramo, for it is said that they are almost certain to perish. It is claimed the Indians, when they had to cross the Cerro de La Muerte, used to carry bundles of nettles, with which they lashed their bodies to increase circulation and enable them to bear the cold.

ATLANTIC TIERRA CALIENTE

The Spanish term tierra caliente, generally used to designate the lowlands of Central America and other Spanish-American coun-

FLORA OF COSTA RICA 11

tries, signifies merely "hot land." As a term it is unsatisfactory, for it implies that the lowlands are regions of intense heat, which is not generally true. Some parts of them are uncomfortably hot, it must be admitted, especially desert areas, or those whose atmos- phere is saturated with water in association with high temperature, but in general the lowlands are less uncomfortable than most regions of the United States in midsummer.

Pittier places the upper limit of the tierra caliente in Costa Rica at 1,000 meters, while Werckl£ locates it at 800 meters. The former author gives 21°-28° C. as the mean temperature.

Entering the harbor of Limon, one has a good view of the Atlantic tierra caliente of Costa Rica. The scene is most lovely and impressive in early morning — in front a lofty wall of deep green, half veiled by low-hanging, fleecy clouds, swaying lazily like curtains stirred by light breezes. One does not realize the presence in the foreground of forested plains that extend for many miles inland from the coast, because only their edge is visible. The eye is held by the steep, green slopes of the high mountains, that appear to rise just beyond the port, but are actually many miles away.

The whole Atlantic slope of Costa Rica, from the sea to the tops of the mountains, except where cleared for cultivation, is occupied by dense rain forest of the type prevalent from southern Mexico almost continuously along the Central American coast, and far southward, at least to the mouth of the Amazon. If viewed from an airplane, nothing would be seen but a monotonous expanse of green, everywhere almost uniform in height, broken only by isolated peaks, and varied but slightly by lower mountains and hills.

When one is riding by railroad from Limon to San Jose", the land- scape of the tierra caliente is monotonous, even to a botanist. To others uninterested in vegetation it must seem even less varied, for the view consists of little but forest, interrupted by an occasional glimpse of a swift stream, especially the roaring waters of the Rio Reventazon. Here and there are plantations of bananas and cacao, but most of these are remote from the main line of the railway.

Along the Atlantic beaches is the usual type of tropical vegeta- tion so widely dispersed in America and even on shores of Old World tropics. The sand is carpeted with mats of rope-like stems of goat- foot morning glory (Ipomoea Pes-caprae) and Canavalia maritima. In salt flats shallowly flooded at high tide are sparse colonies of halophilous grasses and sedges, with Cakile, Sesuvium, Batis, Philox- erus, and other plants. Just back of the strand, in places seldom

12 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

reached by the waves, are compact thickets of shrubs and small trees, their outer edges usually banked with low shrubs of Caesalpinia Crista, Coccoloba Uvifera, and Chrysobalanus Icaco. Behind the shrubs, or frequently at the very edge of the water and projecting far into it, are often large areas of mangrove swamp, with their customary association of Rhizophora, Avicennia, Conocarpus, and Laguncularia. Usually the coast is fringed with graceful coconut palms, which add the necessary touch of picturesqueness to every tropical shore. Beyond the coastal thickets and mangrove swamps stretch miles of unbroken forest, uniform in appearance to the casual observer, but to the botanist observing its elements infinitely varied. Rainfall on the wide Atlantic plains of Costa Rica is so heavy that even the poorest soil can support a luxuriant vegetation. Some parts of these plains, only a fraction of which ever has been planted with bananas, are said to have very rich and productive soil, but much of the land in the wetter parts of Central America has the appearance of being sterile and probably useless for agriculture. It often consists of sticky red clay that collects heavily upon one's feet.

Where the forest has been cut, the ground cultivated for a time, and then abandoned, there often is a fine showing of bright-colored flowers, supplied by a large variety of shrubs and bushy herbs. Most plants that grow in these clearings are never seen in virgin forest, and one wonders what their habitat may have been before man altered the original disposition of the vegetation. If cleared land is neglected, it is soon covered with a lavish growth of herbaceous weeds, frequently six feet or more in height. By the second year there have sprung up coarse shrubs and seedlings of soft-wooded trees, which grow rapidly and soon form a low secondary forest. Growth of such plants is appallingly rapid, and the lowland settle- ments wage a constant fight to protect themselves from being overwhelmed with vegetation, which is growing every day in the year.

Weedy plants that repopulate clearings include such trees and shrubs as Cecropia, Luehea, Apeiba, Trium/etta, Trema, Ochroma, Spondias, various melastomes, Belotia, and dozens of others. The weedy plants are numerous in species and not confined to any special group. Away from the influence of tide water are many open or partly wooded swamps. The open swamps or marshes, often of great extent and affording homes for flocks of noisy aquatic birds, are rather uniform in vegetation. Dominant plants are the stiff- leaved Calathea lutea and Thalia, Canna, and Cyperus giganteus.

FLORA OF COSTA RICA 13

Other almost ubiquitous aquatic plants are Sagittaria, Pistia, Nym- phaea ampla, Pontederia rotundifolia, Eichhornia, and Limnanthemum.

In another type of swamp in which the water is shallow, some- times almost disappearing by the end of the spring months, and often influenced by the tides, there is a good growth of trees of such genera as Mora and Pterocarpus, and vast thickets of palms of the genera Raphia, Corozo, and Manicaria. A giant aroid with handsome inflorescences like callas, Montrichardia arborescens, grows abundantly in such places. These wooded swamps generally have a varied and interesting herbaceous vegetation, and numerous woody vines. One of their most beautiful flowers is Passiflora vitifolia, whose large blossoms are fire-red.

It is the upland forest that covers the major part of the lowland plains, the great plains of Santa Clara and San Carlos, the latter stretching far northwestward, almost to the Pacific coast. It consists of a dense stand of huge trees, often from 30 to 50 meters in height, occupying every available bit of land.

Entering this forest afoot or on horseback, the first feeling is one of bewilderment. One recalls the old remark about being unable to see the forest for the trees, a statement that here, especially for the botanist, is all too literally true. You are in a forest, with trees on every hand, but all you can discern in any direction is tree trunks and more trunks. You can not even guess at what they may be. For most botanists, unfortunately, tree trunks have little significance.

All the branches of the trees are so high overhead that one can form no idea of their foliage, especially because the branches of adjoining trees are interlaced, and even when leaves float down from their branches one never can be quite certain of the tree to which they belong. The only means of identifying these tall trees is to see them cut. Then it is revealed that they are astonishingly diverse as to species, and that pure stands of one species never, or very rarely, occur. Certain species often are especially abundant in a locality, but it is seldom that one dominates any limited forest area.

The most impressive feature of these forests is the vast height of the trunks and their gigantic diameter, especially when, as often happens, this is exaggerated by buttresses. A famous American naturalist a few years ago in a published account of Costa Rica remarked that he had never seen there any trees larger than those of the river valleys of Illinois. He wrote in his late years, long after he had seen Costa Rica, and his memory must have been at fault. I have seen many of the best existing forests of the eastern and

14 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

central United States, and none of their trees are comparable in size with those of the Atlantic forests of Costa Rica.

Because of the difficulty in obtaining material of them, it will be long before all the trees of the Atlantic plains are catalogued, but their principal components are well known. Among the more common ones are Luehea Seemannii, Pentaclethra, a great variety of Lauraceae in several genera, particularly Nectandra, Ocotea, and Phoebe; Ficus and Coussapoa, which often or usually begin life as epiphytes; species of Brosimum, Ogcodeia, Perebea, Poulsenia, Sym- phonia, Hippomane, Minquartia, Virola, Compsoneura, Dialyanthera, Prioria, Dialium, Dipteryx, Zanthoxylum, Bursera Simaruba, Protium, Swietenia macrophylla, Vochysia, Terminalia, Manilkara; the purple- flowered Jacaranda, probably the most brilliant tree of the Atlantic coast; Castillo,, Chrysophyllum, and hundreds of others. While most species are well represented and some abundantly, others are rare. Examples are Lecythis costaricensis, and that odd tree, Theobroma simiarum, whose tall trunk is decorated with sausage-like fruits.

The rain forest has three or four different levels of foliage. Al- though the dense crowns of the tallest trees intercept most of the sunlight, usually there is a definite understory of small trees— small here, yet of respectable size if in a temperate zone. Prominent among them is a variety of palms of minor or medium size, notably in such genera as Socratea, Welfia, Astrocaryum, Euterpe, Geonoma, Iriartea, Reinhardtia, and Chamaedorea. Tree ferns are plentiful in some localities. Zamia often occurs in wide colonies. Of small and medium-sized trees of higher groups may be mentioned Didymopanax, Pourouma, Carica dolichaula, Inga and Pithecolobium, Dracaena, Ravenia, Guatteria, Theobroma, Guarea, Posoqueria, Carapa, and Olmedia.

Shrubs are numerous in species, although seldom closely spaced. They include species of Piper, Heisteria, Siparuna, Swartzia, Quassia, Neea, Cupania, Pentagonia, Cephaelis, Rudgea, and Psychotria. There are many woody vines — call them lianas if the word seems more tropical; the words are synonymous — some of them reaching the tops of the highest trees. Marcgravias are often plentiful, and such water vines as the wild grape (Vitis tiliifolia) and certain Dillenia- ceae; several species of Carludovica, palm-like in foliage; Bigno- niaceae of several genera, with bright-colored flowers that are seldom seen; Allamanda; Rourea; Entada gigas, with gigantic, bean-like pods; species of Strychnos and Maripa; and the prickly sarsaparilla (Smtiax).

FLORA OF COSTA RICA 15

The light is so scant on the forest floor that herbaceous plants are typically sparse in growth. There are few grasses, and those vine-like in habit, \\keLasiacis, or of certain genera with broad leaves, especially adapted to forest conditions, species of Olyra, Streptachne, and Lithachne. The herbs thriving best are certain coarse Monoco- tyledoneae, of such genera as Heliconia, Calathea, Dieffenbachia, Renealmia, Costus, Xiphidium, and their relatives, which often grow in the greatest luxuriance. These plants are perhaps responsible for the greater part of the scant color found in these gloomy forests, whose atmosphere is as sober as that of a great church, and almost as silent, at least for most of the day. One who expects displays of brilliant color in a tropical forest will be disappointed in the Atlantic rain forest, where usually no other color than dark green is visible from a given point.

The Heliconias often grow so luxuriantly as to form, in the case of the larger plants, like H. Mariae, dense and for all practical purposes impenetrable thickets. Their stems are almost as thick and solid as those of banana plants, and so closely set that one can scarcely pass between them.

An important element in these forests consists of epiphytes. From the ground you can see the coarser ones, festooning the trunks of almost every tree with rope-like stems and cloaks of huge leaves. On a recently fallen tree you will discover a host of others that are invisible from the ground. Some of the more freely branched trees support dozens of epiphytic species, ranging in size from minute lichens and hepatics to the largest aroids.

The aroids are probably the most showy and abundant of these epiphytes, especially species of Philodendron and Anthurium. The Monsteras attract attention because of their ample leaves perforated with large holes or "windows." Lush bromeliads are almost as conspicuous as aroids, particularly species of Aechmea and Tillandsia, all of which usually grow high on branches rather than on trunks. Many kinds of ferns are common, but mostly of relatively unin- teresting and widely distributed species. There are a good many orchids, but no such abundance of species as at higher elevations. It is in this belt, however, that there is found the most celebrated of Costa Rican orchids, the guaria de Turrialba, Cattleya Dowiana, its lip crimson veined with gold. Vanilla likewise is a lowland orchid that thrives in dark, wet forest.

The ground covering of small, herbaceous plants seldom is very ample, but it is sufficiently varied. One might expect in such low-

16 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

lands to find only uninteresting and widely distributed species of herbs and such species are frequent enough; but it is here that one finds a good many endemic Costa Rican plants. I have often been impressed by their local distribution. In one place a certain rare herbaceous plant may be abundant, while in another forest area a few miles away, exactly similar in general appearance, perhaps not a single individual of the same species can be discovered. On this account it seems probable that many more endemic lowland species will be described from Costa Rica when such areas as the now almost unknown plains of San Carlos have been explored.

Acanthaceae, in wide variety as to color and habit, are a feature of the rain forest, also Commelinaceae. Several Selaginellas form lacy carpets over the soil. Terrestrial Araceae are not rare, and there are many dwarf plants of Cyclanthus, and huge clumps of Carludovica palmata, so much like a palm, except for its odd inflores- cence. Low-growing Rubiaceae are rather numerous. The Cucur- bitaceae are represented by large-leaved vines of the genera Gurania and Anguria, with small but handsome, orange and red flowers.

The Atlantic forests of Costa Rica have been neglected by most botanical collectors, who may have been eager to reach the more attractive regions of the mountains, more probably because they expected to find little of interest, or perhaps because of the lurking menace of malaria. It has seemed to me that in a land of so many interesting regions, this was one of the most profitable for explora- tion. Nowhere is it possible to find a greater number of plants that are likely to prove new.

PACIFIC TIERRA CALIENTE

In practice, it usually is a long distance from the tierra caliente of the Atlantic coast to the corresponding belt of the Pacific. By air line the distance is not great, but in Costa Rica one does not travel that way. Between the two is usually an interval spent in the Meseta Central, and after that the Atlantic rain forest seems very far away. Then it is not surprising to find on the Pacific coast, at the same elevations — 800 meters or less- — altogether different con- ditions prevailing. So far as general appearance of scenery and vegetation is concerned, the two regions might be a thousand mi-les apart. Equally unlike are the species on the two coasts: most of them are different.

In the Pacific lowlands there is rarely much suggestion of the conventional type of tropical vegetation. The general appearance

FLORA OF COSTA RICA 17

is not so unlike western or southern Texas. For half the year, at least, there is less of green than brown and yellow. In the wet season the general hue of the landscape is not the deep, dreary green of the rain forest, but a livelier green, brightened by abundant sun, more like the vivid green of temperate lands.

The principal factor differentiating the vegetation of the Pacific tierra caliente is rainfall. The fact that a continental divide inter- venes is probably of great importance, but not an insuperable barrier, since there are plenty of low gaps by which plants could cross it if the environment were favorable. As stated previously, rainfall on the Pacific coast is limited to half the year. The Atlantic forests are evergreen, those of the Pacific, such as they are, mostly deciduous, many of the trees and shrubs being leafless during much of the dry season, and many of the herbs dying if annual, or remain- ing dormant if perennial. A botanist can always find on the Pacific coast plenty of plants in flower or fruit, even at the driest season, and it is during the dry season, when travel is easiest, that most botanical exploration has been carried on.

The major part of the Pacific tierra caliente is or has been covered with either forest or thickets of more or less deciduous trees and shrubs. In general, the vegetation may be described as xerophytic. In gross aspect as well as in component species it is most unlike the Atlantic rain forest.

Most of the trees are smaller than those of the wet region, but there are exceptions, the ceibas, sandbox trees (Hura), and guana- caste (Enterolobium) having few rivals in size among Central American trees. These very large examples, however, usually grow as isolated individuals, or at least not densely crowded, as in the Atlantic forest. The trees of the Pacific forest usually are not crowded but generally rather widely spaced, and their tops are often broad and spreading, there being ample sunlight to induce free branching. Much of the soil in the Pacific tierra caliente is rather poor, and the rainfall is not heavy enough to compensate for soil deficiencies.

Descending by train from the Meseta Central toward the Pacific coast, one is impressed by the sparseness of vegetation, especially if the time is the dry season. Many hillsides and plains support but a scanty growth of plants, in which shrubs often are more con- spicuous than trees. The land is so open that it affords natural grazing areas, and in some places trees and bushes have been cut to improve pasturage. Much of the Pacific tierra caliente is devoted to the cattle industry, which is the principal livelihood of this area.

18 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

There are cultivated fields of considerable extent in some parts, but crops are grown only in the rainy season or invierno.

In the lowest parts of the coast, toward the seashore, there often are dense thickets of low and tall shrubs, above which rise scattered trees. Here, as in the Atlantic tierra caliente, it is unusual to find a pure stand of any tree, but normally a large number of species grow in association. Sometimes, it is true, one species of tree domi- nates a locality, as the guanacaste and sandbox trees in much of the Province of Guanacaste.

In large areas of the Pacific tierra caliente, possibly because the land is too poor for the growth of many shrubs or trees, perhaps partly on account of seasonal fires, and probably because of still other conditions, there are no shrubs or trees, or only scattered ones, and the land is a grassy savanna. This type of vegetation does not extend north of Costa Rica except in insignificant areas, but it is well developed in Costa Rica, in Guanacaste and farther southeast- ward in TeYraba and Boruca, toward the Golfo Dulce. Savannas are characteristic of Pacific Panama, and of many regions of north- ern South America, notably Venezuela and the Guianas.

These savannas are beautiful after the rains begin, when they are covered with fresh, bright green grass, and capable of supporting large herds of cattle and horses. The grasses, which often form a dense sward, represent numerous species, mostly plants less than a meter in height and often much lower. Among them grow many other plants, particularly a large number of Cyperaceae, and repre- sentatives of such genera as Cipura, Polygala, Curtia, Melochia, Hyptis, Centrosema, Sauvagesia, Crotalaria, Eriosema, Stylosanthes, Zornia, Evolvulus, Buchnera, Ruellia, and Borreria. There often is an abundance of small but brightly colored flowers. Water collects in shallow pools all over the savannas, and about these are borders of aquatic or hygrophilous plants, such as species of Schultesia, Bacopa, Limnanthemum, and Nymphaea. During the dry season the savannas are parched and brown.

Along the Pacific coast, as along the Atlantic, are mangrove swamps, with their peculiar association of species. The strand vege- tation is nearly, or quite identical with that of the Caribbean shore.

Among the principal large trees of dry Pacific forests are Anacar- dium excelsum, Pseudolmedia, Licania platypus, Sterculia apetala; various species oiFicus; guanacaste or ear tree, Enterolobium, one of the giants of Central American forests, and abundant in many regions, where it is an important source of lumber; Platymiscium;

FLORA OF COSTA RICA 19

Pithecolobium Saman; Pterocarpus Hayesii; Sweetia; Lauraceae in various genera although fewer than in wetter and more elevated regions; several Sapotaceae; Tabebuia pentaphylla, with gorgeous pink flowers; the almost equally showy Triplaris; the ant-infested Cor dia alliodora; Cassia grandis; Ceiba, Bombax, and Bombacopsis; Terminalia; Cedrela; Hura crepitans; Rheedia edulis; Gyrocarpus; Calycophyllum, with sheets of white "flowers" and distinctive, pale, peeling bark; Andira inermis; Dalbergias, the cocobolo trees; and Genipa Caruto. Especially in evidence are trees of the family Leguminosae, and even more numerous are shrubs and herbs of the same family, the group of plants most abundantly represented in the Pacific tierra caliente, as in many other lands of similar climate in remote parts of the earth.

Of characteristic smaller trees of the Pacific tierra caliente one may mention Dipterodendron, with handsome, fern-like foliage; Byrsonima, which often forms extensive groves of distinctive aspect; several species of Coccoloba; Tabebuia chrysantha, with bunches of golden blossoms; Hymenaea Courbaril; Crataeva; a few species of Caesalpinia; Cochlospermum, often only a shrub, with flowers like yellow roses; a rubber tree, Castilla nicoyana; Gecropias, but much fewer than on the Atlantic coast; Chlorophora, the fustic or mora tree that furnishes dyewood; Guazuma; Pourouma;Lacistema; Trophis racemosa; Plumeria acutifolia with beautiful white flowers, produced when the tree is leafless; Anona pur pur ea; Rollinia; Anacardium occidental with curious, edible fruits; Psidium Guajaba, which often forms groves or thickets, as does also the calabash tree, Crescentia Cujete; Sloanea quadrivalvis ; Diphysa robinioides with racemes of yellow blossoms ;Esenbeckia; Simaruba glauca, with edible, olive-like fruits; Muntingia Calabura, also with edible, intensely sweet fruits; Gliricidia;Erythrina rubrinervia, with pale-red, sword-shaped flowers; Spondias purpurea, with edible, plum-like fruits; Pereskia; Trema micrantha; Licania arbor ea; various species of Lonchocarpus ; and Bursera Simaruba.

The shrubs of the Pacific thickets are legion in both individuals and species. While some, like the trees, reach the Atlantic tierra caliente, many, and probably the majority, are restricted to the Pacific coast. They include species of Casearia and Gouania; Ouratea with bright yellow flowers and leathery leaves; Rauwolfia; the abundant Hamelia patens; Chiococca alba, with pure white, disk- like fruits; numerous species of Psychotria (these are more plentiful in wetter regions); Coutarea; Ruprechtia; Curatella americana, the

20 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

sandpaper tree, distinctive in its rough leaves, often forming thickets of characteristic aspect; Cassia biflora and C. nicaraguensis ; various Anonaceae, particularly the carrion-flowered Sapranthus; Helicteres; Melastomaceae, but of fewer species than in wetter regions ;Erythro- chiton; Acacia Farnesiana; Psidium guineense; Triumfetta, with bur- like fruits; several species of Capparis; Heisteria concinna; several Solanums with prickly branches; Ximenia; Mimosa pigra, mostly in marshy places; bullhorn Acacias; Prosopis, chiefly on seashores; Erythroxylon; various species of Acalypha, especially A. diver sifolia. There are many Pipers, but fewer than in wet regions. In the brushy thickets often are extensive areas so closely covered with Bromelias that they are impenetrable by any large animal.

In the Pacific tierra caliente grows the only tree cactus of Costa Rica, Cereus Aragoni, which may not be a native plant, according to statements of some authorities. Here abound palms of the genus Bactris, often producing an impenetrable undergrowth of stiff, spiny stems in places where water stands during the wetter months. An outstanding feature of the landscape in most of the Pacific tierra caliente is the coyol palm, Acrocomia vinifera. In thickets and forests are all too many plants of the sole climbing palm of the region, Desmoncus, with dangerously armed leaves. Other common palms are species of Pyrenoglyphis and Scheelea.

Epiphytes are scarce in most of the Pacific coast, but a few exist, especially Bromeliaceae, principally Tillandsia species, and, less frequently, hardy aroids. Orchids are scarce, but some showy ones grow upon the trees, species of Laelia and Epidendrum, and even the superb guaria morada, Cattleya Skinneri.

Woody vines thrive in the dry thickets and even in the forest, often in dense tangles over shrubbery. Many of them are Bignonia- ceae with bright-colored flowers, produced at the end of the dry season. Likewise plentiful are Malpighiaceae, displaying masses of golden blossoms. Other common vines are Vitis tiliifolia; species of Cissus, often with long, pendent, aerial roots; the blue-flowered Petrea; Trigonia; the pink Securidaca; species ofBauhinia, the stems of some of them ribbon-like and perforated with holes, to suggest the common name escalera de mono, "monkey ladder" ; species of Mucuna, a few with painfully irritant hairs on their large pods; and one or two species of Combretum, with showy, red and yellow flowers attrac- tive to bees and hummingbirds.

The area designated as tierra caliente is, naturally, not sharply separated from the belt next above it, the two gradually merging

FLORA OF COSTA RICA 21

except where there is abrupt transition in climate, as may happen at the crest of a range of hills or mountains. Certain lower hills of the Pacific slope do not rise above the tierra caliente, while others bear on their summits small areas of different vegetation. The height to which the typical tierra caliente vegetation ascends depends largely upon rainfall, and in places where there is plenty of rain, especially if long continued, the tierra templada vegetation descends to a lower level, as in the region of Tilaran. Where the climate is markedly dry, the tierra caliente vegetation rises to higher levels. Its upper belts often remain green nearly or quite throughout the year.

While the vegetation of the Pacific tierra caliente is varied enough, and composed of a large number of species, botanically this is the least interesting part of Costa Rica. Certainly it is the least agree- able in which to work, for the climate is hot, the forest and thickets particularly so, and full of tangled vines and spiny branches, not to mention the ticks that thrive better than elsewhere. On this account, and for lack of good means of transportation, partly also because of the sparsely settled country, the Pacific tierra caliente has been relatively little investigated by botanists. Its exploration involves long rides on horseback on obscure trails, where there are few and often uncomfortable lodging places. It must not be forgotten that some localities on the Pacific coast are noted for a virulent type of malaria.

TEMPERATE REGION (TIERRA TEMPLADA)

In relative botanical interest, the tierra templada ranks just above the Atlantic tierra caliente. Its flora includes large numbers of rare or unusual species, many of them endemic. It is the region in which most of the Costa Rican people live, the one in which all or nearly all the coffee is produced. Wherever in Central America coffee is grown commercially, the climate is agreeable.

Pittier locates the tierra templada between 1,000 and 2,600 meters in altitude and gives its mean temperature as 14°-21° C. These wide limits include practically all the uplands, excluding only the cold regions about mountain summits. It seems to me that Werckle does better in placing the limits of the temperate region (the term is only a relative one) at 800-1,500 meters. That, as remarked, includes all the coffee region, and even extends somewhat higher.

This belt is one of transition, and not sharply marked anywhere, as so often is true of plant belts. It is no easy matter to indicate

22 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

species most distinctive of the area, because so many extend higher or lower, but the same may be said of most of the other regions more or less arbitrarily defined here. On the Pacific slope the so-called tierra templada is much better marked than on the Atlantic, for in the former it may be taken to include all mountain slopes at middle elevation that have a well-defined dry season. The upper limit is recognized readily when, in climbing the slopes of the volcanoes, one meets the line at which trees begin to be heavily infested with orchids, mosses, and other epiphytes.

On the Atlantic slope, where there is almost everywhere plenty of rain throughout the year, it is not easy to recognize any vegetation belts at all, except those marked by different types of agriculture. There the tierra templada may be roughly indicated as beginning in the region of Turrialba and Pejivalle, approximately where coffee cultivation begins, and extending up the volcano slopes to the fields where potatoes are cultivated, and to the pastures for dairy cattle.

Climate in some parts of Costa Rica plays strange pranks. If altitude alone were considered, the mountains of Tilaran in Guana- caste would fall wholly inside the tierra caliente, but as a matter of fact at only 750 meters they display almost the same vegetation seen in central Costa Rica at twice that elevation or more.

The explanation of this phenomenon is that here the continental divide is extremely low, only about 700 meters, and that rain-laden winds from the Caribbean coast are not wholly stopped by the summits of the mountains. The rain clouds drift westward across their tops, and throughout the year considerable rain falls on the west slope of the Tilaran range, which should be arid and parched during the dry season. The climate at Tilaran is the most curious I have encountered in Central America. Although the elevation is only 500 meters, in the dry months there falls almost all day long a fine mist, the pelo de gato, "cat fur," a very descriptive term. Although the sun is shining, in half an hour one's clothes become quite damp. There is a rainbow nearly always in sight. It is said that the name Tilaran, of Indian origin, signifies "the place where rain is always falling." In the town the fields are beautifully green, while only two or three miles away, toward the Gulf of Nicoya, everything is as dry and brown as a desert.

The tierra templada, as stated, contains most of Costa Rica's population, all the area mentioned on the following pages as the Meseta Central, and such important regions as Tilaran, Heredia, San Jose", Cartago, and Santa Maria de Dota. Because of the high

FLORA OF COSTA RICA 28

percentage of cultivated land in the tierra templada, its vegetation has been modified more than that of any other part of Costa Rica, and less remains of the former forest. Originally all or practically all the land was covered by heavy forest, centuries ago, for the large aboriginal population (attested by the abundance of pottery buried everywhere in the soil), must have cleared a good deal of land, and their Spanish successors have almost completed the work.

This is the best farming area of the country, nearly all of it, except where too steep, being suitable for some sort of agriculture. Costa Rica is fortunate in having available for its increasing popula- tion large areas yet unsettled, chiefly because of present lack of transportation. Such lands exist in the mountains of Guanacaste, on the more remote slopes of the central volcanoes, and in the difficultly accessible mountains of the Province of Puntarenas, as well as in the less distant region of Dota, and the plains of San Carlos.

Clearing of land is progressing constantly in some of those regions, and there probably are being destroyed a good many orchids and other epiphytes that never will be known to science. Many species must have passed out of existence thus in the 'central region. There is, for instance, a handsome ornamental shrub, Bouvardia glabra, common in Costa Rican gardens but unknown elsewhere, and not found recently in Costa Rica in a wild state. Probably it grew formerly about San Jos£ or Cartago, but has been wholly destroyed.

Originally the tierra templada must have been covered (except in Guanacaste) by a dense, moist or wet forest in which oaks (Quercus} predominated. In the more heavily populated valleys all these trees have been cut, except upon slopes too steep for cultivation, and even there most usable trees have been removed. It is possible to judge of the primitive forest about San Jose", Heredia, and Cartago only by fragments remaining in quebradas or on slopes above the culti- vated fields, in such places as the beautiful Cerro de La Carpintera, near Cartago, and El Tablazo, near San Jose". About Santa Maria de Dota there are larger forests little disturbed, although even there the better forest lies in what is here called the tierra fria.

Other large trees associated with the oaks are great numbers of Lauraceae, especially species of Nectranda, Ocotea, and Persea, espe- cially P. Schiedeana and P. caerulea; Cedrela or Spanish cedar; Sapium; Inga in several species; Chaetoptelea ; Talauma;Zanthoxylum; Podocarpus; Engelhardtia ; the endemic genus Alfaroa of the Jug- landaceae; and Ladenbergia, a relative of the true quinine trees. Among smaller trees and large shrubs are numerous species of

24 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

Croton; Citharexylum; several woody Compositae, especially the white-flowered Montanoa; Myrtaceae, chiefly species of Myrcia, Calyptranthes, and Eugenia; Urera and Myriocarpa; three species of Hedyosmum; species of Geonoma and several other genera of palms; Malvaviscus and Robinsonella ; Myricas; Panopsis; Litsea; Arctostaphylos ; Conostegia xalapensis; Hauya; Trichilia and Guarea; Roupala; Symplocos; Eurya; Oestrum aurantiacum; and several white- or pink-flowered Rondeletias.

Among smaller shrubs Melastomaceae, Rubiaceae, and Piper are prominent, the last in greater numbers than at any other level. There are many shrubby Compositae, including species of Calea, Vernonia, Eupatorium, and Verbesina. In wetter parts of the Atlantic slope are innumerable red-flowered Gesneriaceae and Ericaceae, mostly epiphytic. Further groups well represented by species or individuals are Solanum, Rubus, Clematis, Paullinia and Serjania, Calliandra, Palicourea, Smilax, Myrsinaceae, Siparuna, and Mollinedia.

Immediately about Cartago and San Jose most of the native shrubs are to be sought in the dense roadside hedges so characteristic of all cultivated parts of Central America. Certain common shrubs are almost confined to hedges. Among the most characteristic shrubs and small trees of such places are Xylosma, Randia Karstenii, Zanthoxylum Limoncello, Iresine Calea, Trichilia havanensis, Mauria, Acnistus, Picramnia, Oestrum aurantiacum, and Erythrina rubrinervia, the last one of the most noticeable small trees of the tierra templada, although by no means confined to it. In the Meseta Central a delightful hedge or roadside plant is the rosa de Castilla, a rose with clusters of small, pink, double flowers that has become naturalized.

Herbaceous plants, both weedy and endemic or rare species, are an important element of the temperate forests. Among them are species of Passi flora, Begonia, Lamourouxia, Salvia, Desmodium, Leucocarpus, Ipomoea, Ischnosiphon (in Guanacaste),Gerarmm, ferns in great variety, including some tree ferns and numerous epiphytes; Cuphea, Spigelia, Gynandropsis, Xanthosoma; Heliconias; wide fields ofEleocharis, and many other Cyperaceae;Canavalia;Loasa;Lopezia; Tibouchina, Centradenia, and other small melastomes; Gynandropsis ; Zebrina; species of Juncus and Phaseolus; Valeriana; Coccocypselum. Interesting among aquatic plants are the several Podostemonaceae that grow submerged on rocks in swift streams.

The pasture lands encircling San Jose and Cartago, especially the former, become dry during winter months, but in the rainy

FLORA OF COSTA RICA 25

season, and at Cartago for most of the year, they are beautiful with fresh, green grass and myriads of small plants with prettily colored flowers. Nothing is more pleasant during the wet season than a walk through the lanes about those places, with high hedges on either side, the road bordered with closely cropped grass from which spring many conspicuous flowers. Common plants of pastures are Lamourouxia and Russelia, Lobelia laxiflora, and Kohleria, all with red flowers; Mimosa albida; Hypoxis; Desmodium; Parosela; Poly- galas; Oenothera rosea; Salvias; Vernonias and Caleas; Sisyrinchium; Centaurium;Evolvulus; Dichondra; Lobelias; EryngiumCarlinae; and several species of Tagetes. Lovely beyond description are the pastures of Cartago, on the lower slopes of Irazu, unbroken sheets of the beautiful hazy blue of Santa Lucia (Alomia microcarpa), which looks exactly like the Ageratum of gardens. Around Cartago and Santa Maria the cornfields often are invaded by tall plants of Santa Catalina, a dahlia with single or double, white or pink flowers, which behaves like sunflowers (Helianthus annuus) in the middle western states.

Epiphytes are plentiful in the temperate region, especially on the Atlantic slope. They are never absent anywhere, and are more varied in species than in the tierra caliente, although much less diversified than in the tierra fria. Orchids are sufficiently common, and many of the species are so ubiquitous as to rank almost as weeds. Here grows the beautiful guaria morada (Cattleya Skinneri). The country people often take numbers of large plants to their homes, and throw them upon tile roofs or on the tile covering of the adobe walls. There the plants thrive, and produce gorgeous masses of flowers in springtime. Another plant of similar habit is the endemic Echeveria australis, which is most at home on tile or dirt roofs.

Other common epiphytes are aroids, generally species of Philo- dendron, Anthurium, and Monstera. Epiphytic ferns are numerous, and there are plenty of mosses, hepatics, and lichens. The Cactaceae are represented by a few species of Cereus and Rhipsalis.

There are many fascinating collecting grounds for plants in the tierra templada, some of which have been celebrated among local and visiting botanists since the days of Oersted. Around San Jos4 so much land is cultivated that little representative native vegetation remains, but only a short distance away by bus are the hills of Escasu and Aserri, where there are tantalizing bits of forest, with large numbers of rare plants. El Tablazo also is easily reached, the type locality for dozens of Costa Rican species. During the rainy season there is a fine showing of beautiful flowers in hedges and

26 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

pastures, but one is not likely to find there species of great rarity. With a horse it is possible to visit in a day the lower slopes of Irazu and Barba, although those regions, at least the ones conspicuously different from the environs of the city of San Jose", lie above the tierra templada.

At Cartago botanical conditions are much more favorable for a collector. It is an easy stroll up the lower slopes of Irazu, through grassy lanes bounded by weathered stone walls, to quebradas where there are many interesting plants, including the rarer endemic species. In such stream beds are scattered individuals of high mountain plants, borne by the current far below their normal habitat. Every roadside tree, large or small, bears a varied burden of epiphytes, orchids and the more conspicuous, brilliant-flowered bromeliads. The mountain air is so cool and invigorating that walking is a pleasure, especially where on every side there is nothing but beauty, wide meadows banked with forest, extensive views in every direction, upward to the summit of Irazu, southward across Cartago and the Reventazon to the mountains beyond Navarro and Orosi. Cartago charms every one who visits it.

South of Cartago, one or two hours by horseback, are some of the richest collecting fields of Costa Rica. The Rio Navarro and Santa Clara compare favorably even with the forests of the upper mountain slopes, and it will be long indeed before their flora is completely known.

Unique among collecting grounds of the central region is the Cerro de La Carpintera, near Tres Rios and Cartago, an elevated and isolated mountain that rises above the fields of Ochomogo, where was fought a famous and almost the only Costa Rican battle of the past hundred years. I have a special liking for La Carpintera, because it can be reached easily on foot from either Cartago or Tres Rios, and because it has an extraordinarily diverse flora, which has yielded dozens of new species. This flora is, however, referable for the most part rather to the tierra fria than to the tierra templada.

The lower slopes of the mountain are cleared for grazing, and there are easy trails to the edge of the forest that covers the upper, painfully steep sides about the summit. Wood is cut at the edge of the forest, so that every year the trees are reduced in number. It would be a patriotic act if this last remaining bit of fine natural vegetation were permanently reserved as a national monument, that future generations might see how beautiful their country was in its primeval state.

FLORA OF COSTA RICA 27

The upper forest of La Carpintera is dense and wet, and dif- ficult to penetrate. Nowhere in Costa Rica have I seen more varied and luxuriant vegetation. There is a profusion of rare trees, many tree ferns, a luxuriant growth of innumerable kinds of epiphytes, Ericaceae and Gesneriaceae, endless ferns, and a great variety of beautiful flowers. Along the ridges are tall plants of Marattia, one of the most primitive ferns, and of great rarity in Costa Rica.

The most memorable feature of a climb up La Carpintera is the fact that all day you are entertained by the roaring of a few naulingos that somehow manage to survive in their isolated and restricted home. Nowhere else so close to the cities of Costa Rica is it possible to hear the magnificent voices of these great black howler monkeys, the largest monkeys of Middle America.

One of the regions in the temperate belt best deserving the attention of a botanist is that of Santa Maria de Dota, south of Cartago and some sixty miles by road from San Jose". Situated on the Pacific watershed, its climate is relatively dry, decidedly so during winter months. Its most famous locality botanically is El Copey, appearing on even the smallest maps of Costa Rica, and consisting of half a dozen houses! There still is plenty of forest about Santa Maria, but little close to the village, and even this remnant is shrinking rapidly. From the settlement it is possible in a day's trip with a horse to reach many rich localities lying at greater elevation.

The vegetation around Santa Maria, while similar in many respects to that of San Jose and Cartago, shows manifest differences or at least is unlike anything now remaining at the latter places. On slopes near the town is a rather open oak forest, the trees often widely spaced and with spreading crowns, although even under such conditions exhibiting little resemblance to northern oak forests. Between them is a comparatively scant growth of shrubbery, with many characteristic species lacking elsewhere in forest. One of the most remarkable is Dodonaea, a shrub more usually found at sea level and most often on seashores.

Perhaps the most distinctive and profitable region in which the writer has ever worked in Central America is that of Tilaran where, in company with Professor Juvenal Valerio, he spent approximately the month of January, 1926. Reference already has been made to the unique climate and physiography of Tilaran. Botanical interest arose primarily from the fact that it was virgin land for exploration, and even more from its surprisingly rich and varied flora, which

28 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

abundantly justified our hopes. An unexpectedly large number of new species was discovered.

Not the least important element contributing to the success of our excursion to Tilaran was the hospitality of its people, who showed an intelligent interest in our work. The land about Tilaran has been settled in recent years by people from the Meseta Central, who form an oasis in the otherwise almost Nicaraguan culture of the Province of Guanacaste. The somewhat primitive but pros- perous and comfortable conditions prevalent at Tilaran suggest pioneer days in the United States, which likewise were noted for their hospitality.

The most vivid impression of Guanacaste — the small part of it that I have seen — is of its abundant animal life. Along streams running into the landlocked Gulf of Nicoya are countless flocks of aquatic and other birds. Every tree along the rivers supports colonies of them, noisily flapping about their nests. Evidently they are little molested, for they pay slight heed to passing boats.

More astonishing than the birds are the mammals, and above all the monkeys. Contrary to popular and generally erroneous pictures of the tropics, monkeys seldom figure in a tropical land- scape; in such places as Panama and Salvador, or even Honduras, you may spend a long time in the forest and see not a single one. But if I described literally the abundance and tameness of Guana- caste monkeys, particularly the big black howlers elsewhere so shy, along the country roads or even on the very edge of the village of Tilaran, I should not be believed. They are so tame that they seem quite indifferent to man, and their numbers are beyond belief. Traveling the roads one is accompanied by their roaring, that suggests nothing so much as the rumbling of a gargantuan coffee mill, and is one of the loudest sounds made by any animal.

It is quite defensible to assign to the temperate region the west- ward slopes of the Sierra de Tilaran, for their flora is strangely like that of the woods about San Jose" and Cartago, where the elevation is twice as great. The Atlantic slopes of the same mountains, being dripping wet, support a flora definitely referable to the Atlantic tierra caliente.

Although the Tilaran flora is so like that of the Meseta Central, it differs in important respects. Oaks (Quercus) are said not to grow in Guanacaste, and I saw none; neither are there members of the genus Rubus, so well represented in the central regions. Never- theless, the flora is sufficiently similar to remind the settlers of their

FLORA OF COSTA RICA 29

former homes in the Meseta Central. It is also, I judge, much like that of the incredibly rich region of San Ramon, where Professor Alberto M. Brenes has made such comprehensive collections.

The Tilaran forests are infinitely varied in composition, with much the same groups of trees that have been listed for the tierra templada. The undergrowth is unusually dense, so tangled that often it is impossible to enter it without a machete. Rich pastures of tall, rank Guinea grass exist where forest has been cleared, and the climate in some places is excellent for coffee production. As previously stated, a short distance often makes a great difference in agricultural potentialities of Guanacaste land. One landowner whom we visited remarked in all seriousness that on his finca of only moderate extent there were practically three distinct climates.

Another outstanding and well-defined temperate region I have visited is the lower slopes of the Volcan de Poas, above Alajuela. They have few inhabitants, chiefly, it seems, because the land is unproductive. Much of it is open, and perhaps formerly under cultivation, at least by the Indians, who must have been numerous there, and wide stretches are covered with bracken (Pteridium), whose presence is usually considered an indication that land has little agricultural value. There are numerous fincas in some parts of the slopes, and especially at lowest elevations most of the natural vegetation has disappeared. In the quebradas are dense growths of rather small trees and many shrubs, and some of the densest thickets of tall bamboo to be found anywhere in Costa Rica. Bamboo thickets are a customary feature of the highest mountain slopes, but they are not a usual type of vegetation on the lower flanks.

COLD REGION (TIERRA FRIA)

The fame of Costa Rica's flora for bewildering variety and exquisite beauty is based primarily upon the vegetation of the tierra fria, and when one comes to discuss this portion of the plant life, one is appalled by the complexity of the task. Only the vocab- ulary and the descriptive genius of a Reginald Farrar could do justice to the subject. I am sure that the plants of the Chinese mountains upon which he lavished his incomparable vocabulary hold no superiority in beauty and variety to those of the high mountains of Costa Rica.

If a botanist has only a short time to spend in Costa Rica, he should hasten to the upper slopes. He will never forget what he sees there of profuse vegetation, and he will have seen, no matter

30 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

how long he remains, only a small fraction of the whole. For many years botanists have been visiting La Palma de San Jose", the place at which this type of vegetation is mostly easily reached from the capital, but late visitors find there many plants never seen or at least never collected by their predecessors. It still seems possible to discover almost as many new species as when La Palma was first visited by a botanist. After seeing the region, it is easy to understand why this is true.

The variety of vegetation in the wet tierra caliente is well known, but it is monotonous uniformity in comparison with what exists in the high mountains. In all North America no region can compete in variety of vegetation and number of species with the higher mountains of Costa Rica, except the similar adjoining Province of Chiriqui in Panama. In all the Americas it is improbable that any region is at all comparable, except the mountains of Colombia, with their related flora, or certain regions of the eastern slopes of the Ecuadorian and Peruvian Andes.

Pittier, as before stated, limits the tierra fria to regions above 2,600 meters, with a mean temperature of 5°-15° C., and there is no doubt that the small area so circumscribed does possess a dis- tinctive flora. It seems to the present writer preferable to follow Werckle, who applies the term tierra fria to all slopes above approxi- mately 1,500 meters. If this is done, it is necessary to subdivide the cold region into two belts, lower and upper, the upper representing the tierra fria of Pittier.

The lower belt of the cold region, then, is an area of dense forests (except where modified by man), the forests composed of small or only medium-sized and densely crowded trees. Their tops are drenched every night and sometimes all day with drizzling rain or heavy downpours, and much of the time they are half hidden by drifting clouds and fog. If you touch a tree or bush you are showered with cold water. Every branch and trunk is swathed in epiphytic vegetation, which is dripping wet. The bunches of mosses, hepatics, and lichens are like saturated sponges. One's feet never touch dry soil .

Agriculturally the cold belt is the region of potatoes and the dairy industry. Wherever you go, you will eat potatoes and eat (clabbered) or drink milk, very likely potatoes boiled in milk, or, at higher elevations, the potatoes may be lacking. We once spent a night at one of the high ranches where so far as could be seen the people (we were well supplied with food ourselves) had nothing to eat except milk, and what game they could kill.

FLORA OF COSTA RICA . 31

The term tierra fria is an accurately descriptive one, for the climate is really cold. There is nothing in the temperature to remind one of the tropics, no matter how tropical the vegetation may appear. I have spent a good many nights in the tierra fria of Costa Rica, as well as in the corresponding belt in other countries of Central America, and I can remember only two when I was comfortably warm while sleeping — one of those was spent on an open veranda! The native people have to be accustomed to the cold and wet, but the children, at least, suffer severely, and often have hacking coughs that bode no long or happy future.

The effect of the cold is intensified by the dampness, for in these regions it is impossible to keep any object dry. Outside the house one's footwear is always wet. Only the tightest houses, and they are few indeed, are impervious to the cold winds and rain that bluster throughout the night.

By day rains are less persistent, but they are often followed by dense fogs, dangerous for the stranger or even the residents them- selves. In regions where fogs prevail, and elsewhere in the uplands for that matter, it is unsafe for a stranger to venture far without a guide; it is a careless host who permits him to do so. I have had few more painful half hours than once in the high mountains of Costa Rica when, after sending back to the house a guide who had proved himself a hindrance to the work I was doing, I suddenly realized that I did not know just where the house was from which we had started.

Another danger that may deserve mention in this place is that of falling branches. Most of the taller trees bear such a heavy load of epiphytes that a wind sends branches crashing noisily to the ground. Especially in the higher mountains it is unsafe to wander through a forest when much wind is blowing. After the end of a gale, it is possible to make a rare haul of epiphytes from fresh branches strewn upon the ground. It is even more fortunate to reach a place where tall trees have been cut recently, but the majority of the epiphytic plants wither on fallen trees in an astonishingly short time.

The reason why these wet mountain forests still yield so many new plants to even the most casual collector is that one person can never see more than mere fragments of them. No collector, unless giving his whole attention to some limited group, will travel far through them in one day, for he will be unable to carry away speci- mens of the plants he finds. Almost anywhere in the region, if a collector experienced in the tropics will follow a trail along a forest,

32 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

at the margin of a meadow, and collect all the plants he sees, none or scarcely any of which will be common ones, in two hundred yards he will gather more specimens than he alone can transport. One wholly unfamiliar with such places will be so bewildered that he can only pick here and there at the most brilliant or curious bits of vegetation, and will overlook the great majority of the plants, many of which, particularly among orchids and ferns, are small and unobtrusive.

Originally all these mountain slopes, except in the highest parts where there are paramos or paramillos, were densely covered with forest. It must have been discovered in comparatively recent years that the upper slopes, when cleared, made excellent pastures if seeded with European grasses, and thus many fine collecting grounds have become accessible to the botanist.

Formerly, as now to some extent, the populated Meseta Central was isolated from the Atlantic coast by mountains that were every- where densely forested. These forests were penetrated only by a narrow, paved cartroad that crossed the divide between Barba and Poas, and descended to the San Juan River. The roads were so long and steep that people who could afford the cost, as I have heard very old people relate, were carried along the trail in chairs on the backs of professional cargadores.

At present it is easy to reach hundreds of places on the upper slopes where dairy farms exist, by roads which, if often painfully steep and boggy, can at least be traversed on horseback. The places thus accessible are so numerous that no one person ever has seen them all, although all lie within a region of no great extent, as the earth's surface is measured.

Each of these many isolated localities yields plants seen nowhere else. It is a remarkable fact that one may go to any secluded nook on the slopes of Barba, Irazu, and Turrialba, whose floras seem to be richer than that of Poas, and find there an always surprising number of showy and conspicuous plants. At another spot in the next quebrada, or in one three or four miles away, there may be an altogether different or at least conspicuously different association of species, in which many of the most conspicuous plants of the first locality are absent. On this account, when visiting a certain area it never is safe to pass a desirable plant, for it may never be seen again.

Many of the species seem to be extremely local in distribution, and this is true of some of the showiest ones. Good examples are

33

the two species of Wercklea (Malvaceae), trees with huge, bright- colored flowers that can not be overlooked. Each species is abundant in certain limited localities on the slopes of Irazu, but they do not grow together, so far as known, and each is known from a single station in the central mountains. Both do grow in other parts of Costa Rica and, I have no doubt, at other places on Irazu and Barba. Some of the most showy plants, naturally, are rather general in distribution, and may be seen almost anywhere one goes.

A catalogue of characteristic trees of the forests of the cold region would include almost the whole mountain forest flora of Costa Rica, but there are certain species, genera, or families that are particularly well represented. Large areas, in Dota up to the edge of the paramo region, are covered with Quercus in almost pure stands. It may be remarked that often it is difficult to determine what a particular tree may be, for it bears so many woody or herbaceous epiphytes that their foliage is much more prominent than that of the host itself. Conversely, often it is hard to decide whether a certain branch represents the tree or an epiphyte.

Podocarpus is another genus typical of the higher mountains. P. oleifolius is not a conspicuous tree, because its branches are so high above the ground, but P, montanus, on account of its fir-like foliage, with leaves white beneath, is more easily discerned. Laura- ceae are everywhere numerous, with several genera and a fair number of species. On many slopes the Weinmannias, most fre- quently W. pinnata, are more plentiful than any other trees, and small trees of Melastomaceae, especially of the genera Blakea and Topobea, are often dominant. In many open places are groves or thickets of Alnus and Vismia. Other large and small trees occurring in some abundance, generally or locally, are Araliaceae, especially Oreopanax, in great variety, some with strikingly handsome and well-differentiated foliage; Prunus; Morus insignis, with greatly elongate but inedible fruit; Magnolia; endless tree ferns, including the largest individuals of the whole country; various Myrtaceae,. especially species of Eugenia and Myrcia; Hedyosinum; Gaiadendron, with showy, bright yellow flowers; Drimys Winteri, a relative of the magnolias, with glaucous leaves, pungent bark, and small, white flowers; slender Phyllonomas, whose minute flowers issue from the upper leaf surface; Brunellia; Pithecolobium species of the subgenus Cojoba; Fuchsia arbor escens, which becomes a round-topped tree of some size, especially if isolated in pastures; the two species of the endemic genus Wercklea, one with mauve, the other with pumpkin-

34 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

yellow flowers as large as those of hollyhocks ;Eupatoriumfistulosum, with huge leaves and panicles of vividly purple flowers; Sambucus; Sty rax, with dead-white flowers; Symplocos; Myrica; Sapium; very numerous Rubiaceae, mostly species of Psychotria, Palicourea, and Faramea; numerous Myrsinaceae; the lovely pink-flowered Hydran- gea Oerstedii, which usually is a high-climbing vine; Bocconia; numerous Guttiferae, especially species of Clusia, with waxy, white flowers, fragrant like orange blossoms; Solandra, with trumpet- shaped, white corollas similar to those of Datura; several species of Ilex; the Icacinaceous tree Calatola, which sometimes grows at lower elevations, as in the region of Dota; and the flaming red- flowered Billia, related to the buckeye (Aesculus) of the North.

Among small shrubs Pipers are plentiful, exhibiting here as much diversity as almost everywhere else in Costa Rica. There are numerous species of Rubus, usually forming thickets in or at the borders of clearings. The one with the best fruit is Rubus glaucus, whose handsome berries are deliciously flavored. Several Senecios with yellow or white heads are much in evidence in some localities. One of the showiest shrubs of the slopes of Irazu is Solenophora calycosa (Gesneriaceae), with trumpet-shaped, dull yellow flowers dotted with purple and as large as a small coffee cup.

On the upper -slopes of most of the higher mountains are inter- laced thickets of tall bamboos, growing beneath the forest trees. These tangles are so dense that it is necessary to cut a trail in order to pass, and scarcely any other plants are found among them. The ground usually is covered with a deep mulch of their fallen leaves.

As might be expected, palms are far less numerous than at lower levels, but there are a good many species of such genera as Euterpe, Geonoma, and Chamaedorea, especially the last, whose representa- tives are low and graceful, slender palms of handsome appearance because of their often brightly colored fruiting spadices. Woody vines are conspicuously few on the upper slopes; mention already has been made of the native Hydrangea. On the. slopes of Irazu a famous plant is the flor del volcan, Solanum Wendlandii, with large clusters of vivid blue blossoms, which have made the vine a favorite in cultivation. Muhlenbeckia drapes old stumps and trunks. Several species of Carludovica with biparted, palm-like leaves cover the tree trunks, but these distinctive plants are fully as much at-home in the tierra caliente, perhaps more so.

The tierra fria is the region par excellence of epiphytes. Condi- tions are ideal for their growth — moderate shade and constant,

FLORA OF COSTA RICA 35

abundant moisture. First in interest are the orchids for which Costa Rica is famous. In the tierra caliente, if moisture is sufficient, there are plenty of orchids, but few are seen because they are so high overhead on branches of tall trees that they can be reached only when a tree is felled. In the tierra templada orchids are much more plentiful than in hotter regions, and some localities, like those south of Cartago and the environs of San Ramon, have yielded great numbers, but even there many of the orchids grow on the taller trees, where it is impossible to collect them except when trees have been felled. But when, ascending the slopes of the volcanoes that overlook the Meseta Central, one comes to the cloud line previously described, one realizes that there at last is the paradise for orchid hunters.

The trunk of every low tree beside the road bears several large clumps of orchid plants. Dismount from your horse, inspect the tree closely, and you will discover dozens of small plants invisible from the middle of the road, species of Pleurothallis, Stelis, Lepanthes, etc. Lift your eyes to the branches, and you will see that every one has small and large clumps of orchids perched along it. The trees are so low, with heavy, broad tops like apple trees, that it is easy to climb into them, and upon any random tree one interested in orchids can find enough to keep him busy for some time. In any other country of Central America one such tree would be a boon, but here are countless thousands, and on every tree one may expect something quite new, not only of orchids but of other groups of plants. It is because of this great diversity and the immensity of the field that it may be stated confidently that the number of orchids known from Costa Rica will be vastly increased with time.

Let no one suppose that the abundance of orchid plants involves a profusion of orchid flowers such as may be seen in orchid houses of the North, for that is far from the case. Orchid species with large and showy flowers are few, and their blooming season seems to be usually short. Yet on every trip one may have the thrill of discovering a few clumps of perfectly flowered Miltonias, varied Epidendrums, Oncidiums, and hosts of others. There are few more delicately beautiful flowers than a thrifty clump of the lovely Epidendrum Endresii (certainly to be envied is the orchid collector for whom it was named), with its panicles of small, white and purple blossoms.

It is not true that small-flowered orchids lack beauty — they merely are not showy. Some of the almost minute flowers, if ex-

36 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

amined closely, prove quite as beautiful and even more intricate than the large ones, but they never will be popular in foreign lands — fortunately, since so many of the showiest orchids of tropical America face extinction by commercial collectors.

Well known to botanists are the orchids — -some of them — of La Palma, a paradise indeed; but dozens of other localities are equally productive or better, and in the Costa Rican mountains are hundreds of other promising places never discovered by a col- lector. Locate a clearing anywhere in the higher mountains, cattle pastures or small openings made apparently by nature, perhaps relics of former small clearings made by man, and you are certain to find all the orchids you will be able to care for. They are abundant as to both species and individuals. From any one of these favored spots it would be possible to carry away wagonloads of orchid plants, if one were inclined to such vandalism.

Popularly orchids are associated with hot, dense forests, but they do not develop best in either heat or deep shade, the great majority of them at least. They thrive where the climate is cool or even chilly, and orchids, except the smallest and most delicate, require plenty of light. The richest collecting grounds are not inside the dark forest but at its outer edge, where the branches are exposed to abundant sunlight.

Orchids are but one element of the epiphytic vegetation of the tierra fria, and not the most voluminous one. Probably the greatest mass of epiphytes consists of mosses, hepatics, and lichens, the last least in evidence. Mosses and hepatics serve a useful purpose as sponges, to maintain abundant moisture at the bases of orchids and other larger epiphytes. Every limb is shrouded in them, so that usually nothing of the bark is visible. From every branch dangle feathery streamers of mosses and hepatics. These same plants exist also on the ground, but in lesser quantity.

For one interested in ferns there is probably no richer field on the whole earth, for the number of ferns in these dripping forests is beyond belief. The best region for them I have seen is the upper slopes of Turrialba, where there seems really no end to them. They cover every tree, and almost as many others grow profusely over the ground. Many hundreds of species have been collected in Costa Rica, many of them still without names, and possibly almost as many more await discovery. They represent most of the known genera, but are particularly numerous in such groups as Polypodium, Asplenium, and Elaphoglossum. It is here that the delicate filmy

FLORA OF COSTA RICA 37

ferns ( Hymenophyllum and Trichomanes) find a congenial habitat, their fronds always bathed in water. Lycopodiums, likewise mostly epiphytic plants, are almost confined to this belt.

Aroids, especially Anthurium species, are very abundant but not particularly showy. They attract notice because of the varied shapes of their leaves, and some have curiously colored spathes or fruit spikes. Bromeliads are exceedingly conspicuous and often confusingly abundant. They continue so up to the very limit of woody vegetation, some of the largest and showiest of all Costa Rican bromeliads being confined to the highest thickets. Well represented there are the genera Thecophyllum, Vriesia, and Guz- mannia, not to mention the Tillandsias, which range from the coast almost to the limit of vegetation. Many bromeliads are far showier than any orchids, with clean clusters of leaves and brilliantly colored inflorescences, the bracts often intensely red or pink, and the petals blue or white.

Important in numbers and diverse in foliage, but possessing few claims to beauty, are the many Peperomias that grow on the upper slopes, but these epiphytes are found in equal or greater profusion at middle or lower elevations. Everywhere, too, are Pileas, which may be either epiphytic or terrestrial. Among the strangest of the epiphytes are several species of Utricularia that often are mistaken for orchids. U. Endresii, with large, rose-purple flowers, really is quite orchid-like in appearance.

The most prominent flowers of the uplands are produced not by orchids but rather by groups which, if represented at all in the North, are not noted for showy blossoms. The profusion of bright flowers in the upper mountains of Costa Rica reminds one of the wealth of color in alpine parks of the Rocky Mountains, the upper slopes of the White Mountains of New Hampshire, or meadows of the mountains of North Carolina and Tennessee. Nature is all too prodigal of yellow, a quite practical hue that wears well, but becomes excessively tedious when overdone, as in most parts of North America. In the Costa Rican uplands one is for once freed of the dominance of yellow. While there are no wide meadows filled with color, there are reckless displays of the most lively hues, and not yellow but reds and pinks, mixed with some white and blue, predominate.

It is probably epiphytic shrubs of the tribe Thibaudieae of the family Ericaceae that contribute most to this riot of color. They are small or large shrubs that abound at almost every forest edge.

38 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

They are not confined to such high elevations, for they are familiar enough in the lower hills beyond Cartago, but wherever they are, no matter how great their abundance, they always seem fresh and new, and one is never sated with them. Each individual flower is so perfect, so fresh, so beautifully and variedly tinted, that each plant seems a new discovery. These fine shrubs have decorative foliage, often bronzed or painted with red and pink, the bracts are as intensely colored as the flowers, and the corollas present many shades of red, pink, and white. Beautiful, too, are the scarcely less abundant Gaultherias of the same family, very unlike the northern ones in habit of growth; and graceful and dainty are some of the epiphytic species of Vaccinium.

Another group of epiphytic plants with gorgeously colored flowers is the Gesneriaceae, which are legion in Costa Rica, and infinitely diversified in habit, form, and color. Pre-eminent are the Columneas, rather coarse in habit, and ordinary in foliage, but with the most gaudy, bright red or orange-red flowers. They are scarcely less abundant or conspicuous than the Ericaceae. Strange are the scandent Campaneas, with bell-shaped flowers suggestive of Gloxinias, luridly colored in green and brown, and dangling on cord-like peduncles far below the supporting branches. Not all Gesneriaceae of the tierra fria are epiphytes. There are many herbaceous or half- woody terrestrial plants, especially in the genus Besleria, but most of these have far less vivid flowers.

A further group of red-flowered plants is the Lobeliaceae, one of the families best represented and most conspicuous in these upper regions. They are either epiphytic or terrestrial, the latter more numerous. Their flowers are prevailingly red, often with some admixture of yellow, or they may be green or green and purple. The genera Burmeistera and Centropogon have many species in both middle and upper regions of Costa Rica, and, while their foliage is mediocre and their habit often ungainly, their flowers, if not so beautiful as those of Ericaceae and Gesneriaceae, yield nothing to them in brilliance of color.

Among the epiphytes are a few species of Eupatorium with white, pink, or purple heads. The prettiest is E. eximium, with neat, succulent leaves and vivid pink flower heads. Species of Senecio and Liabum supply all the yellow needed to vary the color picture. The white clusters of Metternichia cover many half-decayed stumps, often in company with festoons of Marcgravia and Ruyschia. The Marcgravias, while represented even in the tierra caliente, attain

FLORA OF COSTA RICA 39

greatest luxuriance in the cold regions. Their foliage is most decora- tive, the young leaves usually tinged with bronze and red. Further tints of the same colors are exhibited by the bizarre flowers, or rather bracts. Epiphytes are rare among the Rubiaceae, but in these forests two genera have adopted this habit, Ravnia, with bright red, tubular corollas little suggestive of the family to which they belong, and Hillia, with pure white blossoms.

In the tierra fria, as at lower elevations, herbaceous plants play only a minor role in the forest, but there are more of them in open places, or at the margin of the forest. Among the characteristic groups are Acanthaceae, some with handsome blue, white, or red corollas; Hydrocotyle; Hemichaena, with bright yellow flowers, in Dota; Cardamine; Monnina; Dysopsis; Symbolanthus ; Sibthorpia; and Nertera, with small, deep green leaves and bright red berries, reminiscent of its near relative, the partridge berry (Mitchella) of the United States. In wet forests high in the mountains of Dota grows a species of Greigia, the single North American member of this otherwise South American genus of bromeliads. On the slopes of the central volcanoes clumps of large herbs often are tangled with graceful foliage of a slender vine, Cobaea, decorated with bell-shaped flowers of delicate pink. Near it grows Loasa speciosa, a tall, coarse plant with large, pale yellow, bell-shaped flowers. Like most Loasas, it is well protected by a dense covering of long hairs that inflict more painful stings than any nettle.

In the densest forests of Dota one, if fortunate, may find three strangely familiar and welcome northern plants, Conopholis, Mono- tropa, and Chimaphila. In the same canton grows an umbellifer, Myrrhidendron, that is most unlike its northern relatives in attaining the size of a large shrub, with foliage of corresponding dimensions. Two other conspicuous plants of upland forests are species of Symbo- lanthus, a gentian, and a Gynandropsis that sometimes bears brilliant red flowers, but more often only dirty pink ones.

The Costa Rican plant of most individuality, certainly one of those best known in the country, is the higuera (Gunnera), which suggests distantly by habit the garden rhubarb (Rheum'). Its incon- spicuous, greenish inflorescences are hidden beneath the foliage. The leaves are gigantic, the rounded and deeply cordate blades supported on stout stalks and sometimes almost two meters in breadth. The stiff higuera leaves serve well as umbrellas if one is surprised on some mountain trail by a rainstorm. I have seen half a dozen people trudging one after the other stolidly through pouring rain along a

40 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

slippery, muddy trail, each protected by one of the great leaves held by its stalk above his head.

It is not alone among the epiphytes that bright red flowers are found in the high mountains, for some of the terrestrial plants rival them in color. Most beautiful of them are the Bomareas, slender vines with profuse umbels of bell-shaped flowers, deep red, yellow, and green. Two other decorative plants with confusingly similar flowers of exactly the same intense shade of red are Impatiens and Tropaeolum.

One of the most delightful mountain regions of Costa Rica consists of the meadows in the upper, wettest belt that have been cleared for pasture. These are of wide extent, and are being enlarged yearly. They support large herds of dairy cattle, this being the only Central American country in which dairying is an industry of im- portance. From these high, isolated ranches a plentiful supply of milk is carried in metal cans on horses' backs every morning to San Jos4 and Cartago, and on the most remote ranches the milk is made into butter or cheese, the cheese of the Volcan de Turrialba being justly famous.

These meadows, except where unusually well drained, are almost bogs, their soil always muddy, and the close vegetation like a dripping sponge. In tracks made by cattle live myriads of little frogs, some of which have a call suggesting a silver hammer upon a tiny anvil. The grass is vividly green, and it is said there is no change in the appearance of the vegetation from one end of the year to the other.

After the trees have been felled, the meadows usually are planted with grass seed imported from Europe. Mixed with it as impurities are seeds of common European weeds, some of which have become firmly established here, as nowhere else in Central America. Many of them are familiar weeds of the United States, but others are rare or absent in most of temperate North America. Particularly pleasing are the mats of English daisy (Bellis perennis), which finds a moist, cool, uniform climate exactly suited to its growth. After attempting the difficult feat of growing daisies in the United States, it is a joy to see how they prosper throughout these pastures.

With them many other small plants abound, Cerastium viscosum, Silene gallica, Ranunculus, Veronica, Lotus, Rumex Acetosella, two or three species of Trifolium with white or yellow flowers, Medicago lupulina, Taraxacum, and a good many others. On the slopes of Irazu are colonies of foxglove (Digitalis), and in marshy spots along the walled lanes on the slopes of the same volcano are wild colonies

FLORA OF COSTA RICA 41

of callas (Zantedeschia). Elsewhere there have been planted long hedges of Datura Candida with huge, pendent, trumpet-shaped, white corollas that exhale a heavy fragrance. Small groves of alder (Alnus) and isolated trees of Fuchsia arbor escens and showy-flowered Oestrums often add picturesqueness to the pasture landscape.

Introduced plants, principally grasses, often dominate the meadow vegetation, but there is a large element of small native plants, few of them distinguished by showy flowers. Common among them are Sisiyrinchium with either blue or yellow flowers; Alchemillas; Tri- folium amabile, the only native clover; Oenothera cuprea; Halenias; Calceolarias; species of Hypericum; Hydrocotyle with prostrate, rooting stems; and several species of Valeriana. Nertera often forms close mats over shaded banks or mossy fallen logs. In dense tangles in drier parts of the meadows grow two ferns, Pteridium and Dicran- opteris, the latter often scandent.

A very specialized flora reigns about the summits of the volcanoes, or at least about the limit of vegetation, which often is lower than the summit. On the summits of Irazu and Turrialba are extensive fields of volcanic rock, and lava flows often extend far down the slopes. In the mountains of Dota are wide expanses of natural meadow, covering high plains or gentle slopes, constituting the only paramos of North America, perhaps better called paramillos because of their meager extent. They are similar in every respect to the paramos of Colombia, Ecuador, and Peru, as indicated by the typically Andine plants growing in them.

Just below or at the edge of these alpine meadows or paramos are usually interlacing thickets of shrubs and low trees of localized species. Typical are Escallonia, Ribes, Berberis, Mahonia, Hes- peromeles, Holodiscus, Myrtus, Pernettia, Buddleia alpina, Arcyto- phyllum, and a few Senecios. All or most of these are confined to such places. Most of the larger shrubs and small trees are densely invested with epiphytes, such as certain large bromeliads, mosses, and lichens, from which water is dripping constantly. Two con- spicuous herbaceous plants of these places are Myrrhidendron and a giant Rumex sometimes four meters high or more, truly tree-like in habit, although its hollow stems are herbaceous and succulent.

The paramos of the mountains of Dota undoubtedly constitute one of the most interesting floristic regions of Costa Rica, if not the most interesting of all. They are best developed upon Chirripo, the highest peak of Costa Rica, Cerro de La Muerte, and the near-by Cerro de Las Vueltas.

42 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Among their distinctive plants are several large mosses, which often carpet the ground to the exclusion of most other vegetation. There are extensive beds of Sphagnum, of several species, crowded into soft cushions saturated with water. Arising conspicuously from the sphagnum are several species of Lycopodium, stout, stocky, stiffly erect plants, often differentiated from their neighbors by a tinge of dark red. Here, too, are colonies of the narrow, stiff fronds of the Jamesonias, a genus of ferns restricted to the higher Andes, and in North America to these mountains.

Of small herbs, many of which form dense and elevated cushions, few of them with showy flowers, there are a great many, and since so many of these paramo plants are tiny, it seems likely that a great many of them are still to be collected. It must be remembered that the weather here is so uniformly disagreeable, with fogs, mists, and even drenching rains, and a cold wind blowing almost constantly, that only the most enthusiastic collector will linger long to hunt diminutive plants while his feet and knees are wet and sinking deeper every moment into the moss and ice-cold water.

L/upinus and Castilleja are two bright-flowered plants of the drier spots of the paramos and volcanic meadows. Other alpine plants are the almost minute Gentiana sedifolia, several species of Carex, Juncus, and Luzula; Xyris; Eriocaulon and Paepalanthus; a few low and some taller grasses; Acaena cylindrostachya; several species of Alchemilla, one of a definitely Andean type; Halenias; and Eryngium.

It is here in the paramos of Dota that the definitely Andean genus Puya (Bromeliaceae) finds its northernmost outpost, isolated many hundreds of miles from its nearest occurrence in the mountains of Colombia. Except a few scattered shrubs and an occasional stunted tree, it is the tallest and stateliest paramo plant, its pole-like stems scattered through the meadows so as to resemble perplexingly the mullein stalks of some New England pasture.

Some of the best displays of certain of these paramo plants are near Santa Maria de Dota at a comparatively low elevation, probably not more than 2,400 meters, around small or large sphagnum bogs enclosed by dense but not very high forest. These bogs are gradually filling small ponds, and are not greatly unlike similar sphagnum bogs in the mountains of northwestern Montana. It is suspected that they represent remains of former more extensive paramos that have been gradually occupied by forest.

It gives one an uncanny feeling to push for an hour through the densest sort of wet oak forest, with the light so scant that a lantern

FLORA OF COSTA RICA 43

would be useful, then to emerge without warning upon one of these openings flooded with sunlight. The water is bordered with a broad belt of sphagnum, into which the feet sink so deeply that plants only a few feet from the bank, no matter how desirable, can not be reached. Eriocaulon, Paepalanthus, Xyris, and a low, shrubby Hypericum luxuriate in such situations, and it was in such a place that I first saw the Costa Rican Puya. The particular bog in which it grew, a rather large one, was bordered by a plant so strange that the spot seemed almost supernatural. Could an unsuspected cycad possibly occur at such an elevation? The trunks, three or four feet high and very thick, were surely cycad trunks, and the leaves, thick and hard to the touch, were surely those of a cycad. Very slowly it dawned that the plant was a fern, probably Lomaria Wercklei Christ, the only North American representative of another Andean group of plants.

RELATIONSHIPS OF THE COSTA RICAN FLORA

In order to get some idea of the relationships of the Costa Rican flora with those of other regions it is necessary to consider the vegeta- tion according to the belts discussed on preceding pages. A mere glance at the map affords no reason for supposing that the flora of Costa Rica should be particularly notable, or markedly different from that of other parts of Central America, at least Panama and Nicaragua or, more distantly, Guatemala. Yet the fact remains that after composition of the vegetation is studied, Costa Rica is found to have a flora which, taken as a whole, is distinctive, and quite dissimilar from that of other parts of Central America. The Volcano of Chiriqui in Panama is essentially Costa Rican in flora, but other regions of Panama are not.

Of all Central American countries Costa Rica possesses by far the richest flora. Nowhere is it even approached except in the Coban district of Guatemala, which is limited in area as well as in number of species.

The variety of the Costa Rican flora is best illustrated in such groups as orchids, Piperaceae, and ferns, in all of which it is eminent. The percentage of endemism is extraordinarily high, nor is it believed that further exploration of adjoining countries will greatly reduce it. The reason for this is that the high mountains, where the greatest variety of plants exists, are isolated by either elevation or climate, or both, from all neighboring regions. The mountains of Nicaragua, to the north, are low and comparatively or absolutely dry. Those

44 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

of Panama, to the south, except the adjacent Volcano of Chiriqui, are too low to invite comparison. In order to find comparable areas it is necessary to look to such distant countries as Colombia and Ecuador, where, indeed, a similar flora exists. The montane flora of Costa Rica is definitely allied with that of Colombia and Ecuador, and shows little affinity with that of Guatemala, which is essentially Mexican.

The least peculiar region of Costa Rica is the tierra caliente, the majority of the species here having a relatively wide range. The rain forest of the Atlantic tierra caliente is in every respect similar to that prevailing from Panama to at least Guatemala, and probably even to the State of Veracruz in Mexico. Many of the trees have a still wider range, southward as far as the Amazon Basin. A good many endemic species of trees and shrubs are reported from the Atlantic tierra caliente, but these are to be expected in Panama and Nicaragua, or even in regions more remote. The herbaceous plants include a large number that are endemic so far as present knowledge indicates, but herbs are of relatively little significance in rain forest, and even the present endemics may appear in other countries.

The flora of the Pacific tierra caliente of Costa Rica is continuous with that occupying the Pacific coast from Sinaloa in Mexico south to Panama. Essentially the same flora reappears on the coast of Ecuador and, rather strangely, on the north coast of Colombia and Venezuela. It is characterized by a high number of Leguminosae, as usually is the case in tropical regions of limited or periodic rainfall. There are many apparently endemic Costa Rican species in this division, and probably many of them really are confined to the country, since endemism is not unusual in this coastal strip. But not a great deal is known about the Pacific coastal floras of Nica- ragua and Panama, consequently many of these supposed endemics may later be discovered in those countries, if not even farther away.

The flora of the Costa Rican tierra templada is largely endemic. Its great variety of trees and shrubs is unequaled elsewhere except on Chiriqui. Many of them belong to South America genera, but one does not expect mountain species of trees and shrubs to con- tinue over so wide a region as separates Costa Rica from the nearest similar regions, in Colombia. It is only the tierra templada plants growing about settlements or in places altered by man that have, as a rule, a wide range, extending in either direction, but more often toward the south. There are, I believe, a great many more species having in Costa Rica a northern than a southern limit.

FLORA OF COSTA RICA 45

Perhaps the most distinctive feature of the tierra templada flora is the abundance of oaks. Oaks are few in South America, and more numerous in Mexico than anywhere else in the world, but this does not imply any close relationship with the Mexican flora. More significant is the fact that pines fail to reach Costa Rica, but find their southern limit in central Nicaragua.

Mention should be made of the plants of grasslands, not only in the tierra templada but in the Pacific tierra caliente. Most savanna species have a wide range. Their affinities are obviously South American. North of Costa Rica are no savannas worthy of con- sideration, but farther south, near by in Panama and far away in Venezuela and the Guianas, are vast tracts of savanna land, of which the Costa Rican ones are distant outliers. The savanna flora of Costa Rica is essentially South American, not only in genera but often in the very species.

It is the flora of the tierra fria in Costa Rica that probably shows the highest percentage of endemism, and it certainly is the one most clearly South American in taxonomic affinities. This relationship is impressive in such groups as orchids, Melastomaceae, Rubiaceae, Gesneriaceae, the tribe Thibaudieae of the Ericaceae, Cunoniaceae, Araceae, Bromeliaceae, and many more that might be mentioned.

The closest link with the South American, or rather the Andean flora is exemplified in the vestigial paramos of southern Costa Rica, on the mountains of Dota. These represent a purely Andean type of vegetation, which vanishes even before the center of the country is reached.

BOTANICAL EXPLORATION IN COSTA RICA

Of the Central American countries Costa Rica has received most attention from botanical collectors. This is the result of its fame for exuberant and varied vegetation, and also of its delightful climate and scenery, and the fact that in the uplands at least one is safe from malaria and other plagues that menace health in so many parts of Middle America.

Most other Central American countries were explored to some extent by Spanish botanists in colonial days or by other Europeans more than a century ago, but Costa Rica seems almost wholly to have escaped their attention. Plants were collected in Guatemala and Panama 150 years ago or more, but we find no mention of Costa Rican vegetation in the works of early writers, except for casual refer- ences in Oviedo's History of the Indies (1541).

46 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

The botanist who was privileged to reveal to science the first glimpse of the riches of the Costa Rican flora was a Dane, Anders Sandoe Oersted, who landed at Puntarenas in 1864. Fortunate indeed was the man who first with the critical eye of a botanist had the opportunity of exploring such a flora as this. Except for the common weeds of roadside and field, almost every plant he saw after reaching the uplands was new to science. By accident he landed at the strategic point for beginning his exploration, the point at which the least distinct flora was to be found. As he journeyed inland, up the Pacific slope, to the Meseta Central, and later the wet Atlantic slope, and the steep sides of the volcanoes, one climax after another was unfolded before him.

Oersted was an enthusiastic and capable botanist, who fully appreciated the opportunity offered him. From the coast he traveled slowly toward the interior, collecting at various stations along the way. Especially valuable were the collections made at the mines of Aguacate, a locality not revisited by later botanists, so far as I know. Reaching San Jose", he remained there some months, making excursions in different directions. He visited Pacaca and the valley of Jaris, then inhabited only by Indians. His pen has left a vivid picture of the forests of the mountains of Candelaria, that rise above San Jose", as they existed in his day. At present nothing remains of those forests where he collected so many of his species except a few scattered groves of scant extent, much of their remaining vegeta- tion now damaged by pasturing cattle.

In Cartago Oersted passed several months. He made the acquaint- ance of Don Francisco Maria Oreamuno and Don Francisco Gutierrez, who greatly faciliated his explorations. To the former he dedicated the genus Oreamunoa of the Juglandaceae, to the latter two beautiful plants of the Cartago region, Lamourouxia Gutierrezii and Sipho- campylus Gutierrezii.

From Cartago Oersted must have made many excursions in search of new plants, but probably the regions accessible then were much less distant than those that now can be reached easily. It is probable that in the immediate vicinity of Cartago there was then a great deal more forest than exists today. He visited the slopes of Irazu that rise steeply above the town, and reached the summit. It was thus that so many of the high mountain plants of Costa Rica have their type localities on the slopes of Irazu.

Descending toward the coast, Oersted stopped at El Naranjo, now Juan Vinas, where he collected many more new plants, and also

FLORA OF COSTA RICA 47

at the settlement of Turrialba. Thence he made his way by most difficult trails down the Rio Reventazon to Moin, and left the country by the Sarapiqui Valley. In Nicaragua he continued his botanical explorations, and also had an opportunity to visit Guanacaste in Costa Rica, where he made what were until recently the only collec- tions known from the major part of that isolated province.

After three years in Central America, Oersted returned to Copenhagen, to devote the remaining twenty-five years of his life to study of his collections. Unfortunately he never completed a report upon his plants, although the herbarium shows that he named all or most of them, but he did publish many papers discussing the larger families, and describing many of the new species he had discovered. Groups that he treated in some detail included the palms, Lobeliaceae, Acanthaceae, Begoniaceae, Gesneriaceae, and Compositae.

Oersted's original specimens, preserved at Copenhagen, represent the cream of the endemic species of the Costa Rican mountains. Few of his specimens, apparently, were distributed to European herbaria, but in the United States a good many of his duplicates now are found in Field Museum of Natural History and the United States National Museum.

In 1848 Costa Rica was visited by a Polish gardener, Warscewicz, who had been a member of a Belgian commission sent to Central America with the idea of establishing a colony for settlement by Belgian emigrants. He visited several countries of Central America and northern South America, devoting much of his attention to orchids and hummingbirds. Little or nothing is known of his routes in Central America, for as a rule no locality data appear on his labels, and there is often doubt as to whether the species based upon his specimens were collected in Costa Rica or Panama, or even in Colom- bia or Ecuador. He did, however, gather original material of many Costa Rican orchids.

Later naturalists to explore the country were Moritz Wagner and Carl Scherzer, the latter commemorated in the handsome aroid, Anthurium Scherzerianum, now grown in hothouses all over the earth, but neither of these was a botanist. In 1853 or 1854 there came to Costa Rica a German, Carl Hoffmann, who spent the few remaining years of his life, until 1859, in study of the flora of his adopted land. He worked principally about the capital, and pub- lished accounts of his ascents of the volcanoes of Irazu and Barba. His collections are preserved at Berlin, where they have served as

48 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

basis for description of many of the most handsome ornamental plants of the mountains.

In 1856 and 1857 Hermann Wendland, gardener of the court of Hanover, visited Costa Rica and made a small but highly impor- tant collection of plants. He gave special attention to palms, orchids, and aroids, plants difficult to make into specimens, and consequently neglected by most collectors. He entered the country by the Sarapiqui Valley, and explored especially the mountain chain from Barba to Turrialba.

The German gardener Julian Carmiol, who died in San Jos£ in 1885, assembled a large number of the finer ornamental plants of Costa Rica in his garden. He was responsible for introducing to Europe many native species that are still favorites in cultivation.

Between 1870 and 1880 Costa Rica was visited by several Euro- pean collectors, among them F. C. Lehmann, who seems to have made here only a small collection. He devoted many years to work in the mountains of Ecuador and western Colombia. Endres is best known for the orchids he discovered, and while nothing is available regarding his routes in Costa Rica, he must have found some of the best regions for these plants, because many species were described by Reichenbach from his collections. In the Natural History Mu- seum of Vienna, where his specimens are deposited, there is also much material in other families, but little has ever been published concerning it. Otto Kuntze crossed the country from Limon to Puntarenas, collecting at various localities, especially in the Atlantic tierra caliente, and he mentions many of the plants in his account of his voyage around the earth.

In 1875 Dr. Helmut Polakowsky came to Costa Rica, in company with other teachers imported by the government to reorganize the secondary schools, and for two years he was instructor in natural history in the Institute Nacional. He botanized particularly in the vicinity of Cartago and San Jose", and many of his specimens came from the Cerro de La Carpintera. He was author of several important papers upon the flora of Costa Rica, and described a number of new species. His specimens, at least in part, are in the Berlin herbarium.

Costa Rica has always been famous for its own distinguished men of letters and sciences, and it is not surprising to learn that the major part of its flora is the result of work by native botanists or others who adopted it as their home. Modern increase in knowledge of Costa Rican vegetation began with Professor Anastasio Alfaro, for many years Director of the National Museum of Costa Rica,

FLORA OF COSTA RICA 49

who in 1888 published a list of the known plants of the country, compiled from Hemsley's recently completed Botany of the Biologia Centr ali- Americana. It is interesting to note that previous explora- tion had resulted in the report of 1,218 species of phanerogams and Pteridophyta from Costa Rica. Probably three-fourths of these were discovered by Oersted.

Professor Alfaro began, with the publication of this list, an interest in botany that has continued for many years. He has given most attention to orchids, ferns, mosses, and Cactaceae, in all of which he has made notable discoveries, in spite of the fact that his major work has been in other branches of natural history. His contribu- tions to our present knowledge of the Costa Rican flora are very extensive indeed.

He was instrumental in arousing to interest in botanical collecting Juan J. Cooper of Cartago, already an enthusiastic naturalist, who made a large collection of specimens, mostly in the mountains south of Cartago. These were sent for study to Captain John Donnell Smith, who described from them a large number of new species.

Henry Pittier came to Costa Rica in 1887 from Switzerland, one of several Swiss educators invited to the country about this time to reorganize the educational system, He remained in Costa Rica until 1903, and during those years he and the several botanists or naturalists associated with him executed a systematic exploration which has not been equaled in any other country of tropical America. Pittier's interests were catholic, as indicated by the long list of his publications, covering almost every branch of natural history in its broadest sense, and other subjects as well, for Costa Rica and other regions of Central America and northern South America.

Henry Pittier undoubtedly has gained a more intimate knowledge of the natural history and especially the botany of Central America and northwestern South America than has ever been possessed by any single person. His many papers upon plants of Central America are fundamentally essential for any botanical work that ever may be undertaken there. The writer had the good fortune to be associ- ated intimately with him for a number of years, and heard from his lips much about his work in Costa Rica. It was his sympathetic and fluent description that first gave the writer a desire to visit a country that seemed to possess such a distinctive charm, not only botanically but in many other respects.

The combined labors of Pittier and Adolfo Tonduz, Pablo Biolley, Carlos Werckle, Alberto M. Brenes, and the Brade brothers

50 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

resulted in the formation in the National Museum in San Jos£ of a national herbarium which in 1903 was unequaled below the Rio Grande del Norte. They visited almost every corner of the country, and while neither they nor many others could succeed in collecting all the plants of Costa Rica, they brought together a marvelous number of them, estimated by Pittier at 5,000 species. He began publication of the first comprehensive account of the Costa Rican flora, in the Primitiae Costarricenses, a work that most unfortunately was not brought to completion. In his account of the Plantas Usuaks de Costa Rica he presented more information regarding Costa Rican vegetation than had ever been published before.

The herbarium so formed consisted of more than 18,000 numbers, which were sent to specialists of North America and Europe for determination. It may be seen from the data adduced that Pittier and his collaborators added to the known flora of Costa Rica almost 4,000 species! Besides the specimens filed in San Jose", many thou- sands of duplicates were distributed to the leading herbaria of the world. In all the larger herbaria of Europe and the United States there are so many of them that one marvels at the patient labor expended in their collection.

Adolf o Tonduz is well known to the botanical world for his collec- tions of plants. He gave most of his life completely to them and spent more than 30 years in exploring the mountains and lowlands of Costa Rica. A man of somewhat eccentric nature, devoted to the flora of his adopted land, he built an enduring monument in the many Costa Rican plants that bear his name, and in his contributions toward advancement of botanical science. Carlos Werckl£ likewise was a man with rather eccentric traits — many botanists seem to be so — and at his unfortunate death he is believed to have possessed an unequaled store of information regarding the far corners of Costa Rica and their plant life. The writer had an opportunity of meeting him only once, and then but briefly, and was amazed at the fund of knowledge that he exhibited so fluently. Werckl£ collected less extensively than the other persons named but discriminately, giving his attention to special groups, particularly bromeliads and orchids. The majority of endemic bromeliads of Costa Rica were described from his collections. His specimens usually were disgraceful, for on his tramps over the country he used to cram any strange plants into his pockets or into a bag, and they reached the herbarium in a sad state, but still in good enough condition for study. After all, such a method of collection is not ill adapted to bromeliads. Werckle

FLORA OF COSTA RICA 51

published an important paper upon the phytogeography of Costa Rica, a work that is tantalizing because of its hints concerning rare plants that unfortunately still remain unknown to science, except as briefly mentioned by its author. His thorough knowledge of the country is well proved by this important paper, the only extensive one that he published.

Werckle"'s later years were devoted to the collection of orchids, chiefly at the instigation of Dona Amparo de Zeledon, under whose patronage there were collected so many of the species figuring in Schlechter's classic account of Costa Rican orchids. Much of the material upon which Schlechter worked was gathered also by the Brade brothers, whose prime interest was in plants having horti- cultural value.

The work of United States botanists in Costa Rica began with Captain John Donnell Smith of Baltimore, Maryland, who, after a business career, retired about 1884 to devote the rest of his life to the botanical work in which he had long been interested. These years were equal to many a lifetime, for he lived until 1928, and lacked but a few months of attaining a full hundred years.

Captain Smith's attention probably was first drawn to Costa Rica by the collections of Juan J. Cooper, which were sent him for determination. In 1896 he visited the country, where he was accom- panied on expeditions by Anastasio Alfaro. A charming gentleman, distinguished in appearance, highly cultured, with keen enthusiasm for his work, Captain Smith was a welcome visitor, and Professor Alfaro has spoken affectionately of his companionship upon their excursions. I have also heard Captain Smith recall with evident pleasure his memories of the Costa Rican mountains, where he saw so many plants that previously he had known only from mere dried specimens. Among the regions that he visited were the plains of Santa Clara, La Palma de San Jose", and the mountains south of Cartago.

For many years Captain Smith was the authority upon the flora of Central America. He financed collectors in several countries, and received large collections for study. He assembled a very exten- sive herbarium, chiefly of Central American plants, with hundreds of types of species he had described. These, with his large library, are now in the United States National Museum in Washington.

Reports upon Costa Rica by enthusiastic visitors have caused several United States botanists and collectors to travel to Costa Rica, where they have obtained large series of new and valuable

52 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

material. Among them have been Dr. William R. Maxon of the United States National Museum, who has visited Costa Rica twice, in search primarily of ferns. Dr. 0. F. Cook and C. B. Doyle of the United States Department of Agriculture made a substantial collection in different parts of the country, including many specimens of palms. Dr. J. M. Greenman of the Missouri Botanical Garden collected in the Costa Rican mountains, where he was especially interested in the genus Senecio. Prof. H. E. Stork of Carleton College, Minnesota, has made two admirable collections, largely in the region of Dota. Dr. C. W. Dodge of the Missouri Botanical Garden has visited Costa Rica twice, in search of cryptogamic plants, particularly lichens. There are also a number of other visitors from the United States who have made collections in Costa Rica.

A few years ago the well known mycologist, H. Sydow of Berlin, made an extensive collection of Costa Rican parasitic fungi about which he has issued various publications. Additional material has been collected for him by Professor Brenes. Specimens of some of the host plants are deposited in the Berlin herbarium, and others have been sent to the writer for determination.

Of special importance because of the publications based upon them are the collections made recently by the Austrian expedition under Dr. Porsch. Most of the botanical material was collected by Dr. Giorgi Cufodontis, who has published an admirable list of the plants collected, with critical notes and descriptions of new species.

Among the residents in the country who have made important contributions to knowledge of its flora is Mr. C. H. Lankester of Las Concavas. With wide interests in natural history, he has made intensive studies of the birds and butterflies. In botany his special field of interest has been the orchids, which he has hunted assiduously in the mountains and lowlands. With facilities for care of living plants, he has brought them into flower, and the specimens he has sent to the Kew Gardens and to the United States have served as types for dozens of new species. To most of the foreign botanists visiting Costa Rica in recent years Mr. Lankester has been a genial host and a tireless guide.

Of the Costa Ricans who have given serious attention to the flora of their native land one is preeminent for his work. Professor Alberto M. Brenes, botanist of the National Museum for many years, took up the work of exploration after Pittier left the country. With unequaled zeal and devotion he has continued his collections to the present time, and has accumulated a herbarium of more than

FLORA OF COSTA RICA 53

20,000 numbers. In volume and value of his collections, he has no rival in Central America. In fact, it is questionable whether they have been surpassed even in South America, unless by such botanists as Spruce, Glaziou, and Ducke in Brazil.

The writer is extremely fortunate in being able to include in this flora the unique collection made by Professor Brenes. Although representing several distinct regions, it comes in chief part from the mountains of San Ramon, a center of inexhaustible botanical variety. It well illustrates the floristic wealth of a region of the Costa Rican mountains. No such intensive study has been given to any part of Central America, and the San Ramon region is now better known in herbaria than any other area of equal extent in Middle America, unless it be the Panama Canal Zone, or Morelia, Mexico. How fortunate botanical science would be if only there were more collectors of equal industry and discrimination!

Other Costa Ricans besides those already mentioned have been active in expanding the available knowledge regarding the flora. For many years Oton Jimenez has worked both directly and in- directly to explore the forests, and it is much to be regretted that the demands of business affairs have precluded a greater amount of personal field work on the part of one who has such a keen perception of facts and the ability to discover them in strange places. As it is, he has acquired a vast amount of interesting data concerning Costa Rican plants, which it is hoped may be made available to the public. Many of his specimens are found in the United States National Museum.

Professor Rube"n Torres Rojas of Cartago in recent years has made extensive collections, especially of ferns, in the varied region of Cartago. In the Herbarium of Field Museum there is an extensive series of specimens of his collection that has proved unusually valu- able because of the numerous vernacular names that accompany it. Some of his earlier collections are in the United States National Museum.

The first collection made in the Tilaran Mountains, so far as I know, was one by Prof. Juvenal Valerio Rodriguez, now Director of the Museo Nacional. This material, from El Silencio de Tilaran, was fascinating because of the curious flora that it indicated, and aroused in the writer a desire, later gratified, to visit the mountains of Guanacaste. Mention should be made of Prof. Romulo Aguilar and the late Prof. Fidel Tristan, the latter a congenial companion and a wise guide to many foreign naturalists who visited Costa Rica.

54 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Prof. Manuel Valerio has forwarded to the writer more than a thousand numbers of plants from the most varied regions, some of the species new to the country and others new to science. He has also made extensive collections of mosses that have added many species to the known Costa Rican flora.

Very recently there have been received excellent and sometimes extensive collections made by promising new collectors resident in Costa Rica. Among them are Fernando Solis Rojas and Professor Manuel Quiros Calvo, and it is greatly hoped that the number of Costa Rican botanists may increase. There is no good reason why floras of Central American countries should not be written by the resident botanists who have a deeper understanding of their native lands. Only when this is done shall we ever have a fully satisfactory knowledge of the plant life of Middle America. The foreign botanist, no matter how sympathetic his attitude toward a strange country nor how well he may explore it, misses so much of the local atmos- phere and overlooks data that are common knowledge among the whole population. No matter how widely he may travel, he can never gain the intimate geographic and floristic knowledge that a person born in the country could acquire.

It is unfortunately true that local floras published in Latin America by resident scientists often have been superficial or worse, but that is equally true of many published in Europe and the United States. Every Central American country should and could possess a botanical library adequate for study of its flora, and a compre- hensive herbarium, formed by local collectors. Such a herbarium now exists in Costa Rica. By sending the local collections to special- ists in countries where facilities exist for their study, there can be assembled an authoritatively determined series of any country's flora, and with this as a basis, assisted by the proper literature, in which any Latin American institution can easily obtain aid from North American botanical libraries, the local botanists could begin original creative work that would form a basic contribution of the highest practical value to science.

The writer's personal experience in Costa Rica has consisted of two visits to the country, in the winters of 1923-24 and 1925-26. While the time spent there was sadly limited, special facilities graciously provided by friends in Costa Rica made it possible to visit a surprising number of localities, and obtain a large quantity of diversified herbarium material. Without such exceptionally generous assistance the work would have been far less successful.

FLORA OF COSTA RICA 55

During the first season attention was centered upon the central region, with many short trips about San Jose" and Cartago. Longer excursions were made to La Palma and La Hondura, on the old road to the Atlantic coast; Las Nubes, high on the slopes of Irazu; El Coyolar and the Rio Grande de Tarcoles, near the Pacific coast; the regions of Navarro, La Estrella, and Orosi south of Cartago; and the Cerro de La Carpintera, near Cartago. A memorable excur- sion was made to Alajuela, and an ascent to the summit of the Vol- cano of Poas, with its forest-walled lake and its awe-inspiring crater. Another had as its object the Volcano of Turrialba, as far as the famous finca situated well toward its summit. Very profitable visits were made to several parts of the Atlantic coast, especially La Colombiana and Guapiles, regions that have received little attention from collectors.

During the second visit to Costa Rica more distant regions were visited, although collections were made at some of the places explored during the preceding years, such as La Palma and La Hondura, Cartago, Navarro, and La Carpintera. Almost a month was spent in work about Santa Maria de Dota, an area little explored pre- viously. After the Dota expedition a month was devoted to Guana- caste, where, from headquarters at Tilaran, daily excursions were made in every direction. Our trips extended as far as the Lake of Arenal, on the Atlantic side of the Sierra de Tilaran.

Several excursions of a day each were made to the upper slopes of the Volcan de Barba, a volcano which, in spite of its accessibility, has attracted few collectors. An excursion to Fraijanes, on the slopes of Poas, was especially noteworthy for the new plants it revealed. At Pejivalle a rich collection was made of the plants of the border line between the tierra templada and the tierra caliente. Two days were passed in collecting in the rain forest along the banks of the Reventazon near El Cairo.

During these two expeditions to Costa Rica approximately 15,000 numbers of plants were obtained, representing all groups, with special emphasis upon orchids. Although the field covered was rather wide, and a good deal of effort expended to utilize it, it is realized that only a small part of Costa Rica was seen. Travel to the farther parts of the country, or even to some of those quite near the capital, as a bird flies, is so tedious that many months and years of time and effort are necessary if one will see most of Costa Rica.

As the map is studied, with a knowledge of the localities at which plants have been collected, it is plain that most of Costa Rica never

56 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

has been visited by a botanist, except most casually. Whole regions are botanically unknown, and until every mountain side has been explored, every region of the plains visited repeatedly, there can be no certainty that the flora is even approximately well known. Some places have had more than their share of exploration, the environs of San Jos£ and Cartago, and the southward slopes of Irazu, the volcano slopes most visited by a long line of botanists, and apparently the least interesting ones of the country. The region of San Ramon has been combed assiduously, to reveal an unparalleled flora. But even in the outskirts of San Jos4 the writer has found undescribed species, and many others doubtless can be found in even the best explored localities.

Regions conspicuously in need of exploration are so numerous that it is impractical to list them. Most alluring of all, perhaps, are the forests of El General and the slopes of Chirripo and the other high mountains of the Cordillera of Dota. Scarcely less promising are the volcanoes of Guanacaste, whose very summits have scarcely been seen even from afar by a botanist. There is no reason to suppose they are less interesting than the central volcanoes, and what sort of a flora they possess is a matter for speculation. Close as they are to the Meseta Central, the northward slopes of the central volcanoes are wholly unknown, nor are they easy to reach. The savannas and plains of the southeastern part of Costa Rica are almost virgin territory.

One of the most promising fields awaiting botanical collectors is the wide plains of San Carlos. Practically nothing is known of the whole region toward the Rio San Juan, and not a great deal about the rest of the Atlantic plains.

Most of the unexplored regions mentioned are fairly easy of access, so far as human transportation is concerned. I have often envied ornithological collectors, who carry in saddle-bags all the necessary equipment for collecting birds, and transport their finished collections with ease. Unfortunately a botanical collector needs bulky and heavy equipment, and the finished specimens are even more bulky, making transportation a serious problem. A botanist can easily visit the most remote and desirable parts of Costa Rica, but without almost unlimited funds for hire and care of pack animals, he can not make collections there, and even with such funds, pack animals can not always be obtained. It is on this account that for many decades to come Costa Rica will continue to attract botanists, for they will be able only to nibble from year to year at regions that

FLORA OF COSTA RICA 57

become gradually more accessible. Even progressive accessibility can not be depended upon, for many regions of the tropics, including some in Central America, are far less accessible now than they were decades or even centuries ago.

PLAN OF THE FLORA

On the following pages is presented a systematic, annotated list of the phanerogamic plants of Costa Rica. In ferns the country is one of the richest areas of the earth, and it is unfortunate that they can not be listed here, especially since their literature is even more widely scattered, perhaps, than that of the flowering plants. There have been published extensive lists of the mosses, hepatics, and lichens, but all these, like the fungi, are still known but imperfectly, the mosses and lichens being better known than the other groups.

In most details the plan of the following list is obvious. For each family there have been cited all Costa Rican species known to the writer, with indication of their distribution within and outside the country. For names based on Costa Rican material there is cited the place of publication, and for such species there are cited also their synonyms with place of publication. The first citation of material for a species published from Costa Rica is the type collec- tion, with the locality and the name of its collector. Type collec- tions of synonyms are cited in parentheses after the citation of the synonym. Species without citation of literature were described from other countries. The species believed to be endemic, that is, con- fined to Costa Rica, are so indicated (when only a single collection is known, this has not seemed necessary), but some of these doubtless will be found later in other countries.

Cultivated plants are included in the list, so far as they are known. For some obscure reason, cultivated species are despised by the aver- age systematic botanist as being something unworthy of his atten- tion, with the natural result that common garden plants are less well known, as regards nomenclature and taxonomy, than our native flora. While horticultural plants do not constitute a part of the native flora of a region, from a practical standpoint and as a matter of interest to residents and visitors, they are often more important than the native ones. It is regretted only that the list of cultivated plants of Costa Rica can not be presented more completely than it appears on the following pages.

In the present flora are listed all the species known or reported for Cocos Island, so far as they are familiar to the author.

58 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Unfortunately, but little is known of its flora which, from general reports, must be a rather rich one. Although lying far from Costa Rica, Cocos Island is a possession of that republic and, somewhat strangely, the Cocos Island flora seems to be as closely related to that of Costa Rica as to that of any other continental region.

At this place it may be worth while to insert a word of caution regarding Costa Rican place names. The names of saints, particu- larly, are all too lavishly employed to designate geographic localities in Central America, and within a given country, often indeed within the same province or department, the same saint's name may desig- nate two or more settlements. Locally the places usually are dis- tinguished by the addition of a reference to the province or to some near-by locality, as San Isidro de Coronado and San Isidro de Heredia. Frequently the distinguishing phrases do not appear upon collectors' labels, sometimes leading to the assumption that two plant localities are identical when in fact they may be many miles apart, in quite unlike phytogeographic regions. Especially to be noted in Costa Rica is the name La Palma. La Palma de San Jose" in the Depart- ment of San Jose* has long been a favorite collecting ground for local and visiting botanists, and is the type locality for many species. Almost equally famous for orchids, however, is another place of the same name, La Palma de San Ramon, at which Professor Brenes and others have found so many new species. Upon the following pages these and other similarly duplicated names have not always been distinguished.

ACKNOWLEDGMENTS

Were it possible to name here all persons who have given material assistance to the writer in the course of work with the plants of Costa Rica, the list would be a very long one, of many dozens of names. The preparation of a local flora well illustrates the necessity of cooperation by many people to achieve a definite and visible end. How many collectors, how many students in both field and her- barium, working together or separately for almost a century, have collaborated to make possible the presentation of facts that appears on the following pages! Unfortunately it is practical to mention only a few of the persons to whom the writer is indebted, but these few are important.

With his wonted hospitality toward visiting scientists, Professor Anastasio Alfaro, then Director of the Museo Nacional, generously provided convenient working quarters in the museum building during all the time the writer spent in Costa Rica. These were more than

FLORA OF COSTA RICA 59

ample, and an extremely pleasant place in which to be, even during the earthquake weeks of March, 1924! Professor Alfaro contributed freely of his time to the success of the work, and gave helpful advice as to its execution, besides collecting a large quantity of highly desirable specimens. He was a congenial guide upon a delightful excursion to the Volcano of Poas, of which the most interesting and vivid memories are preserved by the writer. With his simplicity, gentleness, and abundant store of accurate knowledge, Professor Alfaro represents the best type of scientist, a man whom I shall always remember with deepest affection.

On both his visits to Costa Rica very direct assistance of the most substantial kind was given the author by Don Oton Jimenez Luthmer of San Jose", who is the good counselor of North American botanists. His advice upon practical matters of procedure as well as upon regions best suited for exploration has been invaluable. The writer looks back with longing to the many pleasant evenings spent in his company, when the work of the past and future was discussed so sympathetically. Even more to be remembered are rides with him to the mountains, and it was, in fact, in his company that I made an excursion to La Palma, where I saw for the first time the wonderful diversity of the flora of the tierra fria. No petty detail of the writer's work was too small for his consideration; without his unstinted kindness the writer's memories of Costa Rica would be less perfect. His sympathetic and witty exposition of the history and culture of Costa Rica have contributed more than the reading of many books to an understanding of the principles that distinguish this country among all lands of tropical America.

To Mr. C. H. Lankester of Las Concavas the writer owes many favors and many of the brightest pictures in his gallery of Costa Rican recollections. One who has experienced the hospitality of Mr. and Mrs. Lankester at Las Concavas has something to remember. Nor does their hospitality end there. The writer has not forgotten that they sent a special messenger upon a two days' journey to bring a greeting at Christmas time. Such courtesies are not forgotten.

To Mr. Ferdinand Nevermann there are special obligations for a most pleasant and profitable visit to his fincas in the lowlands along the Reventazon River. Enviable is the botanist who receives a welcome from so considerate a host, or visits the forest with so competent a guide.

Another who will ever have a special place in the writer's affections is Professor Ruben Torres Rojas of Cartago, to whom he is indebted

60 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

for some of the happiest days of all his many memorable ones in Central America. I have been fortunate in having his company upon several important excursions to Navarro, to the upper slopes of Turrialba, to Orosi, and to the beautiful region of Frai janes on the slopes of Poas. His gentleness and consideration, his inflexible punctuality, and his joyous enthusiasm will ever be remembered with pleasure.

I have left until the last one person to whom I am most indebted for practical help and companionship through many days of work in the mountains and lowlands of Costa Rica — Professor Juvenal Valerio Rodriguez, Director of the Museo Nacional of Costa Rica. During my first visit to Costa Rica we made but one excursion to- gether, to La Hondura de San Jose". During the second expedition I was more fortunate. Professor Valerio accompanied me during part of the time spent in Santa Maria de Dota, and together we made two ascents to the paramos of Cerro de Las Vueltas, spending there the unforgettable New Year's Eve of 1926. For a month we were to- gether in Guanacaste, a province that without his aid I should scarcely have seen. The great success of that expedition and the warm welcome received in Guanacaste were due solely to his fore- sight and to the high esteem in which he is held in that alluring region. Later we visited the lowlands of Pejivalle and El Cairo, and made several memorable ascents of the slopes of the Volcan de Barba, setting out from Heredia in the madrugada or earlier, so that we saw the awesome spectacle of the sun's slow illumination of the Meseta Central, with a final blaze of light and color that no painter's brush could exaggerate. The whole day would be spent in the forests and meadows high on Barba' s slopes, and riding homeward after dark we would see the less spectacular but very peaceful and comforting view of the Meseta Central, dotted with myriad clusters of electric lights, each locating a city or a tiny village.

All these many days of excursions were very profitable ones, all took us to fascinating places where there was much beauty and entertainment. Sometimes there was aching weariness when the day was over, because of the great amount of work that had to be done. Such an opportunity and necessity for work seldom comes to a botanist. Throughout these weeks Professor Valerio was always the most kind and entertaining companion, conversing eloquently upon interesting subjects pertaining to science and many other fields of knowledge. He was a tireless worker, always patient and considerate, even when he had frequent reason for provocation to

FLORA OF COSTA RICA 61

quite other moods. His kindness and friendship will always be treasured by one who often has sorely tried both.

There are many other Costa Ricans who deserve mention here, rich and poor, who extended the hospitality of their homes to shelter a guest who was undoubtedly a pest — this is a presumption; there never was any visible sign— with his peculiarities. There were also many others who were generous in providing information of the most varied sorts, upon voyages or while collecting material in the field. Everyone seemed to have a genuine and intelligent interest in the work being done, and a desire to be of service, if even in the most modest way. I can not remember ever to have met with dis- courtesy from any Costa Rican. If the eyes of any of these persons should ever light on these pages, let them feel that their sympathy has helped to give the foreign world an idea of the flora of their country, and let them know that wherever in the world there is a person who knows something about Costa Rica, their country has a friend.

While it is impossible to mention all the people who gave direct assistance to the writer's exploration in Costa Rica, there must be mentioned a few others who aided in other ways than the making of collections. From Don Prospero Mena of Santa Maria de Dota many favors were received. To Dona Isabel Valverde of Santa Maria and her family the writer is indebted for the most homelike life, at least of the longest duration, that he has ever enjoyed in Central America. And especially remembered are the evenings spent in Tilaran with Don Federico Carmioland Don Daniel Esquivel, whose geniality had an important part in the success of our work in Guanacaste, and in the pleasant recollections that are retained of it.

To Professor Oakes Ames of Harvard University the writer is indebted for the long list of Costa Rican orchids that appears on the following pages, and also for substantial aid in the success of the two expeditions to Costa Rica. To Dr. William R. Maxon of the United States National Museum special thanks are given for the loan of a large amount of material studied in preparation of the Flora. Last, but not least, special acknowledgment should be made of the many favors received from the United Fruit Company, which has so often aided the writer's work in Central America. That company's cus- tomarily generous assistance toward scientific work contributed greatly toward the success of the two visits to Costa Rica. Its employees in the Atlantic lowlands were courteous in affording facilities for work in the tierra caliente, and to some of them, par-

62 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

ticularly Dr. Paul V. Siggers, then of La Colombiana, the writer is indebted for many profitable and pleasant days spent in the luxuriant forests that border the great banana plantations.

BIBLIOGRAPHY

The publications cited below are those most important or useful for study of the Costa Rican flora. The list is brief, but if there were included all papers containing references to Costa Rican plants, it would fill many pages. A more complete bibliography may be found in Pittier's Plantas Usuales. Unfortunately, the works practically useful for study of Central American plants, at least for nonprofes- sional botanists, are few. Those that treat isolated groups or contain descriptions of new species are scattered through serials and other volumes, many of which are available in only the larger libraries. In the systematic list forming the principal part of this volume there will be found references to monographs of families and genera that are useful in study of the Costa Rican species concerned.

Alfaro, Anastasio. Lista de las plantas encontradas hasta ahora en Costa Rica y en los territories limitroifes, extractada de la "Biologia Centrali- Americana." Anales del Museo Nacional de Costa Rica 1: 1-101. 1888. A list of 1,218 species, cited in Hemsley's classic account of the Central American flora.

Bartram, Edwin B. Costa Rican mosses collected by Paul C. Standley in 1924- 26. Contr. U. S. Nat. Herb. 26: 51-114. /. 1-39. 1928. An enumeration of a large number of species occurring in Costa Rica, with references to previous literature upon mosses of the region.

Calvert, Amelia Smith, and Galvert, Philip Powell. A year of Costa Rican natural history. 577 pages, numerous plates, map. The Macmillan Company, New York, 1917. A good description of Costa Rica, with extensive notes regarding both animals and plants.

Dodge, Carroll W. The foliose and fruticose lichens of Costa Rica. I. Ann. Mo. Bot. Gard. 20: 373-467. 1933. Includes a map and a general account of geography, vegetation, and exploration.

Durand, Th., and Pittier, H. Primitiae florae costaricensis. 2 vols., Brussels and San Jose, 1891-1901. The only proper flora ever published for Costa Rica and very useful, but unfortunately it does not cover all the families.

Hemsley, W. B. Botany, in Godman and Salvin, Biologia Centrali-Americana, 5 vols., ill. London, 1879-88. Lists all Costa Rican species of phanerogams and pteridophyta known at the time of publication.

Pittier, Henri. Ensayo sobre las plantas usuales de Costa Rica. 176 pages, 31 plates. Washington, 1908. The most useful published paper for gaining a general idea of Costa Rican vegetation, and equally useful for almost all other parts of Central America.

Record, Samuel J., and Mell, Clayton D. Timbers of tropical America. 610 pages, frontispiece, 50 plates. New Haven, 1924. A descriptive account of the principal woods of tropical America, with much information regarding trees of Central America and their economic uses.

Standley, Paul C. Trees and shrubs of Mexico. Contr. U. S. Nat. Herb., vol. 23. 1,721 pages. Washington, 1920-26. Contains descriptions of many Costu Rican species.

FLORA OF COSTA RICA 63

Standley, Paul G. Flora of the Panama Canal Zone. Contr. U. S. Nat. Herb., vol. 27. 416 pages, 51 plates. Washington, 1928. Useful for the coast regions of Costa Rica.

Flora of the Lancetilla Valley, Honduras. Field Mus. Nat. Hist., Bot. Ser., vol. 10. 418 pages, 68 plates. Chicago, 1931. Contains descriptions and illustrations of many Costa Rican plants.

— and Calderon, Salvador. Lista preliminar de las plantas de El Salvador. 274 pages. San Salvador, 1925. Includes notes and lists of vernacular names that are somewhat useful also in Costa Rica.

Werckle, Carlos. La subregion fitogeografica costarricense. 55 pages. San

Jose, 1909. An excellent account of the phytogeography of Costa Rica, by

one who knew it well. It is unfortunate that one who knew so much of the

>•' plants of Costa Rica, through many years of keen observation, should have

published so little.

SYSTEMATIC LIST OF FAMILIES, GENERA, AND SPECIES

CYCADACEAE. Cycad Family

One of the most ancient groups of higher plants, represented in the native flora of Central America only by the genus Zamia.

CYCAS L.

Cycas revoluta Thunb. Cultivated at Desamparados, and doubtless in other regions. Native of the East Indies. A handsome, palm-like plant with a short, thick trunk, the very stiff leaves with very numerous narrowly linear leaflets.

ZAMIA L.

Zamia Skinneri Warscewicz. Wet forests of the Atlantic coast, ascending to Pejivalle (900 meters), and in the mountains of Guana- caste (up to 700 meters) ; abundant in many places. Also in Panama. A large plant of palm-like appearance, the thick, rough trunk as much as a meter high, or the leaves often rising directly from the ground; leaves pinnate, the segments few or numerous, rigid, lanceo- late, with conspicuous, elevated nerves; fruit cone-like, the scales covered with a brown or rusty wool. A very handsome plant, which has been introduced to the greenhouses of Europe. Specimens of Zamia from Costa Rica have been referred to Z. pseudoparasitica Yates, which is doubtfully distinct from Z. Skinneri.

In the coast of Honduras there is a species of this group (Z. fur- furacea L. f.), which is known by the name Camotillo. Its root is highly poisonous, and has been employed at times for criminal poisoning, as well as for poisoning noxious animals. There is a popular belief that the root, if out of the ground two days, kills its human victim in two days; if dug a week before, it kills in a week, and so on. I do not know whether the Costa Rican species is

64 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

used in the same manner, or whether the same belief is held there regarding it. It goes without saying that, although the root is a dangerous poison, it does not kill in accordance with the popular schedule.

TAXACEAE. Yew Family Reference: R. Pilger, Pflanzenreich IV. 5. 1903.

PODOCARPUS L'Her.

A genus of wide dispersal in tropical regions of almost the whole earth. Besides the species enumerated here, another Central Ameri- can one grows in Guatemala and British Honduras. The wood is yellowish or brownish, of fine and uniform texture, and easy to work. In the regions where it is found it is employed for the same purposes as pine (Pinus). Pittier states that in Costa Rica it is favored for the manufacture of certain parts of ox carts.

Podocarpus montanus (Willd.) Lodd. Ciprecillo, Cobola. Volcan de Poas; mountains of the Canton de Dota; at 2,000-3,000 meters. A South American species that does not extend north of Costa Rica. A tall tree, 25 meters high or more; leaves 1-2 cm. long, linear, white on the lower surface; fruit juicy, purplish black, with a single seed. The branches look almost exactly like those of fir (Abies'), and I well remember the first time that I ever saw some of them, at the Lecheria on Volcan de Poas thrown on the ground where they had been used as the basis for a bed, just as fir branches are sometimes used in the United States. I could not believe that they were not fir branches, but at the same time could not guess how they might have reached Costa Rica! The tree is plentiful on the slopes of Poas from the Lecheria to the crater.

Podocarpus oleifolius Don. Ciprecillo, Cobola. Wet forest of Volcan de Poas and mountains of the Canton de Dota; El Muneco, Prov. Cartago; region of San Ramon; at 1,400-2,700 meters. A South American species that does not extend north of Costa Rica. A tall tree, up to 20 meters or more in height; leaves linear-lanceolate, 3-8 cm. long, about 1 cm. wide, green. All parts of the plant exhale an unpleasant odor. The trees are abundant in the mountains, especially in Dota, but are so tall that a person on foot ordinarily sees nothing of their foliage, except the young seedlings, and is unaware of the adult trees.

Pilger reports for Costa Rica (Poas) P. macrostachyus Parl., another South American species. The specimens I have seen were

FLORA OF COSTA RICA 65

all sterile, and it has been impossible to recognize more than a single species, because the differences are based upon the inflorescences. However, I consider it highly improbable that there exist on Poas two (this and P. oleifolius) closely related species of the genus. Consequently, it seems more nearly in accordance with the probable facts to refer all the specimens with broad leaves to a single species.

CONIFERAE. Pine Family ARAUCARIA Juss.

Several species of this genus are rather frequent or common as ornamental trees in Central America, often attaining a great size. They are natives of South America, Australasia, and the Pacific islands.

Araucaria excelsa R. Br. Planted frequently in the temperate region. Native of Norfolk Island. A handsome tree, at first pyrami- dal in form, with horizontal or drooping branches, the rigid, linear leaves somewhat curved, not closely imbricated. Doubtless other species are planted in Costa Rica, especially A. Rulei Muell., with appressed, densely imbricated leaves.

CRYPTOMERIA D. Don

Cryptomeria japonica D. Don. Planted as an ornamental tree in the temperate region. Native of Japan and China. A tall tree with reddish brown bark peeling off in shreds; leaves linear-subulate, laterally compressed, somewhat incurved, 6-8 mm. long.

CUPRESSUS L. Cypress

Cupressus Benthamii Endl. Cipres. Cultivated commonly in the Meseta Central and elsewhere as an ornamental tree. Native of the mountains of Guatemala and Mexico. The species has been reported for Costa Rica as Juniperus flaccida Schlecht. The branches are employed commonly as decorations in churches and houses.

PINUS L. Pine

In spite of the fact that ill-informed writers, with little regard for accuracy, have reported the existence in Costa Rica of pine forests, these exist only in their imagination, for pines in a wild state do not exist in the country. The genus has its southern limit in America in central or northern Nicaragua.

Pinus oocarpa Schiede. Pino. Planted occasionally in small numbers as an ornamental tree in the region of San Jose", and espe-

66 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

cially about fincas on the middle slopes of the mountains. Ranging from Mexico to Honduras and probably to Nicaragua.

THUJA L.

Thuja orientalis L. A shrub or small tree, native of eastern Asia, sometimes planted for ornament in parks and fincas in Costa Rica. In general appearance it suggests Cupressus, but is distin- guished by its evidently compressed branchlets.

TYPHACEAE. Cat-tail Family TYPHA L. Cat-tail

Typha angustifolia L. Espadana, Tule balsa. T. domingensis Pers. A plant of open swamps, forming wide colonies in some parts of the Atlantic coast, and probably in other regions of the country. Widely distributed in both hemispheres. In other parts of Central America the plant is called Tule and Enea. It is a coarse herb, 1-2 meters high, with spongy, flat, long, linear leaves; the inflorescence is a thick, chestnut-colored spike 10-40 cm. long. In some parts of the Central American coast the fluffy "wool" from the spikes is utilized for stuffing pillows and cushions.

In gardens there are cultivated sometimes as ornamental plants species of Pandanus (P. tectorius Soland. and P. dubius Spreng.; Pandanaceae). They are tall plants, somewhat palm-like in habit, the narrow leaves sword-shaped, their margins armed with fine, spine-like teeth. They are natives of the East Indies.

POTAMOGETONACEAE. Pondweed Family

POTAMOGETON L. Pondweed

Potamogeton foliosus Raf. Meseta Central and probably in other regions. A species widely distributed in America. An aquatic plant with long, slender stems, floating in water, the leaves linear, the minute flowers green, in small spikes. The species of Potamogeton grow in lakes or streams, attached to soil or stones, with their leaves all submerged or some of them floating on the surface of the water. It is probable that other species grow in Costa Rica, although these plants are not plentiful in the tropics.

It is probable that the genus Naias (Naiadaceae) is represented in Costa Rica. Two or more species occur elsewhere in Central America.

FLORA OF COSTA RICA 67

ALISMAGEAE. Arrowhead Family ECHINODORUS L. Rich.

Echinodorus tenellus (Mart.) Buchenau. Open swamps in the coasts or in regions of scant elevation. Widely distributed in America. A small herb, the leaves linear to elliptic; flowers small, white; fruit a small head of minute achenes. Undoubtedly other species of Echino- dorus are to be found along the Atlantic coast.

SAGITTARIA L.

Sagittaria lancifolia L. Atlantic coast to the Meseta Central; Guanacaste; in swamps or in the edges of streams and lakes. Widely dispersed in tropical America. An aquatic plant, the leaves lance- linear to elliptic, 20-50 cm. long; flowers large, white, in long racemes; fruit a head of many achenes.

Sagittaria latifolia Willd. Swampy places in the coasts, usually growing in shallow water. Widely distributed in America. Leaves arrow-shaped.

BUTOMACEAE HYDROCLEIS L. Rich.

Hydrocleis parviflora Seub. Collected in Guanacaste by Oersted. An aquatic plant, otherwise South American in distribu- tion, unknown elsewhere in Central America.

There is to be found almost certainly in Costa Rica, at least on the Pacific slope, Limnocharis flava (L.) Buchenau, another aquatic plant of this family. It is known from both Nicaragua and Panama.

Also to be expected in Costa Rica are species of Triuris and Scia- phila, of the family Triuridaceae. They are delicate saprophytes of dense, wet forests of the tierra caliente. Species of Sciaphila are known from Honduras and Panama.

GRAMINEAE. Grass Family

Reference: A. S. Hitchcock, The grasses of Central America, Contr. U. S. Nat. Herb. 24: 557-762. 1930.

One of the largest families of plants, and the one containing the plants of greatest importance to man. Grasses are most abundant in temperate regions, but they are well represented in Costa Rica.

AEGOPOGON Humb. & Bonpl.

Aegopogon cenchroides HBK. Meseta Central. Mexico to Bolivia.

68 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Aegopogon tenellus (Cav.) Trin. Meseta Central, and prob- ably in other regions. Arizona to northern South America.

AGROSTIS L.

Agrostis bacillata Hack. Oesterr. Bot. Zeitschr. 52: 59. 1902. Type from Cerro de La Muerte, 3,100 meters; paramos of that peak, and of Cerro de Las Vueltas. Endemic.

Agrostis Hoffmann! Mez, Repert. Sp. Nov. 18: 3. 1922. Irazu, Hoffmann; also Cerro de La Muerte, 3,100 meters. Guatemala to Chile.

Agrostis Pittieri Hack. Oesterr. Bot. Zeitschr. 52: 60. 1902. Known only from Volcan de Poas, 2,500 meters, growing in sphagnum bogs. Endemic.

Agrostis stolonif era L. Abundant in meadows of the volcanoes, where it has been sown for pasture. Imported from Europe.

Agrostis tolucensis HBK. Paramos of the high peaks, 2,700- 3,100 meters. Mexico to Chile.

Agrostis turrialbae Mez, Repert. Sp. Nov. 18: 4. 1922. Volcan de Turrialba, Pittier. Turrialba and Cerro de Las Vueltas, in pastures and paramos, 2,000-2,600 meters. Endemic.

ANDROPOGON L.

Andropogon bicornis L. Cola de venado. Meseta Central, and regions of less elevation, growing in savannas and abandoned fields; region of San Ramon. Widely distributed in America.

Andropogon brevifolius Swartz. Meseta Central and regions of less elevation, in pastures and on open banks; region of San Ramon. Tropical regions of both hemispheres.

Andropogon condensatus HBK. Meseta Central to the coasts. Mexico and Lesser Antilles to Argentina.

Andropogon glomeratus (Walt.) BSP. A common grass of the Meseta Central, and in regions of less elevation. Southeastern United States to Argentina.

Andropogon hirtiflorus (Nees) Kunth. Canton de Dota; Meseta Central, extending to the coasts. Widely distributed in America.

Andropogon leucostachyus HBK. Meseta Central, extending to the coasts. Widely distributed in America.

FLORA OF COSTA RICA 69

Andropogon semiberbis (Nees) Kunth. Meseta Central, and in the tierra caliente. Florida and Mexico to Argentina.

Andropogon virgatus Desv. Savannas of the tierra caliente. Extending to the West Indies and Brazil.

Andropogon virginicus L. On banks and in sterile fields at low elevations. Panama to the West Indies and United States.

ANTHEPHORA Schreb.

Anthephora hermaphrodita (L.) Kuntze. A common plant of the coasts. Generally distributed in tropical America.

ANTHOXANTHUM L.

Anthoxanthum odoratum L. Frequent in the pastures of the volcanoes, 1,500-2,500 meters, naturalized from Europe. The plant has a sweet, agreeable odor.

ARISTIDA L.

Reference: A. S. Hitchcock, The North American species of Aristida, Contr. U. S. Nat. Herb. 22: 517-586. 1924.

Aristida capillacea Lam. Boruca, Buenos Aires, and other regions of little elevation, usually in savannas; region of San Ramon at 1,100 meters. Mexico to Bolivia and Brazil.

Aristida jorullensis Kunth. Pacific coast, in dry or sterile fields. Mexico to Panama.

Aristida ternipes Cav. Pacific coast, in dry or sterile places. Arizona to Colombia and Cuba.

Aristida torta (Nees) Kunth. A. breviglumis Mez, Repert. Sp. Nov. 17: 152. 1921 (Buenos Aires, Tonduz). Region of Buenos Aires. Extending to Brazil.

ARTHROSTYLIDIUM Rupr.

Arthrostylidium Maxonii Hitchc. Proc. Biol. Soc. Wash. 40: 80. 1927. Wet forests of Santa Clara de Cartago, 1,950 meters, Maxon 8154; region south of Cartago, and on the slopes of Barba, 1,400-2,000 meters. Endemic. A bamboo with long, slender stems as much as 4 meters in length.

Arthrostylidium Pittieri Hack. Oesterr. Bot. Zeitschr. 53: 75. 1904. Wet forests in the region of the Meseta Central, the type collected near San Jose". Also in Guatemala.

70 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Arthrostylidium racemiflorum Steud. Carrizo. Mountains of Guanacaste, and probably in other places on the Pacific slope; region of San Ramon. A slender bamboo with stems as much as 5 meters long.

ARUNDINARIA Michx.

Arundinaria Standleyi Hitchc. Proc. Biol. Soc. Wash. 40: 79. 1927. Wet forest, El Muneco, Prov. Cartago, 1,400-1,500 meters, Standley & Torres 51050. Endemic; known only from the type locality. An erect bamboo, or sometimes almost scandent, the stems as much as 3 meters long.

Arundinaria viscosa Hitchc. Proc. Biol. Soc. Wash. 40: 79. 1927. Bosques de Velirla, Copey, 1,800-2,700 meters, Tonduz 11729. Canton de Dota, abundant in many places and forming a dense undergrowth in forest. Also in Venezuela. An erect bamboo, up to 4 meters in height.

ARUNDINELLA Raddi

Arundinella Berteroniana (Schult.) Hitchc. & Chase. Meseta Central, slopes of the volcanoes, and in places of less elevation; common in many localities; region of San Ramon. Mexico to Brazil.

Arundinella confinis (Schult.) Hitchc. & Chase. A grass of pastures and open fields. Mexico to the West Indies and Paraguay.

Arundinella Deppeana Nees. Cola de venado. Common in the Canton de Dota, Pejivalle, and other regions, ascending to 2,000 meters; region of San Ramon. Mexico and the West Indies to Brazil.

AVENA L.

Avena sterilis L. San Jose". A European plant, introduced but probably not naturalized.

Avena sativa L. Avena. The common oats, so important as a grain crop in temperate regions, has been planted experimentally in Costa Rica, but does not thrive in Central America, even in regions where potatoes are grown.

AXONOPUS Beauv.

Axonopus aureus Beauv. A grass of pastures and savannas, at low elevations. Central America to Brazil and Bolivia.

Axonopus capillaris (Lam.) Chase. In fields and savannas. Central America to Brazil.

FLORA OF COSTA RICA 71

Axonopus chrysoblepharis (Lag.) Chase. Savannas of the Pacific coast, at 500 meters or less. Ranging to Paraguay.

Axonopus compressus (Swartz) Beauv. Zacate amargo. Abundant in waste ground, forests, and thickets, especially in the tierra caliente; Meseta Central, and sometimes in pastures of the volcanoes, ascending to 1,800 meters or higher; Guanacaste; Cocos Island. Generally distributed in tropical America.

Axonopus Purpusii (Mez) Chase. Zacate amargo. Paspalum Purpusii Mez. Savannas and pastures at low elevations; collected also on the slopes of Barba, 1,800 meters. Mexico to Argentina.

Axonopus scoparius (Fluegge) Hitchc. Pie de paloma. Savan- nas and pastures, region of Cartago and elsewhere. Planted in some regions as a pasture grass. Salvador to Brazil and Bolivia.

BAMBUSA Retz. Bamboo

Bambusa vulgaris Schrad. Bambu. Planted almost everywhere for ornament, and also for its tall, thick stems, which are utilized in various ways. An Asiatic plant, naturalized in all tropical regions.

BOUTELOUA Lag.

Reference: David Griffiths, The grama grasses, Contr. U. S. Nat. Herb. 14: 343-428. 1912.

Bouteloua alamosana Vasey. Pacific coast. Ranging to northern Mexico.

Bouteloua americana (L.) Scribn. Savannas and pastures, at low elevations. Honduras to Venezuela and British Guiana.

Bouteloua chondrosioides (HBK.) Benth. Nicoya, 150 meters. Unknown in other parts of Central America. Mexico and south- western United States.

Bouteloua pilosa (Hook, f.) Benth. Savannas of the Pacific coast. Guatemala to Peru.

Bouteloua repens (HBK.) Scribn. & Merr. Pacific coast. Mexico to Panama.

BRAGHIARIA Griseb.

Brachiaria plantaginea (Link) Hitchc. Wet fields and waste ground, Meseta Central to the coasts, at 1,300 meters or lower. Southern United States to Brazil and Bolivia.

72 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

BRACHYPODIUM Beauv.

Brachypodium mexicanum Link. Forests and thickets of the higher mountains. Mexico to Bolivia.

BRIZA L.

Briza minor L. Pastures near Cartago, 1,500 meters. An annual, naturalized from Europe.

BROMUS L.

Bromus laciniatus Beal. Triguillo. B. proximus Shear. Meseta Central and slopes of the volcanoes. Guatemala and Mexico.

CALAMAGROSTIS Adans.

Calamagrostis intermedia (Presl) Steud. Paramos of Cerro de Buena Vista, 3,100 meters. A South American species, unknown north of Costa Rica.

Calamagrostis Pittieri Hack. Oesterr. Bot. Zeitschr. 52: 108. 1902. Wet paramos of Cerro de Buena Vista, 3,100 meters, the type; also Cerro de La Muerte. Endemic.

CENCHRUS L. Sandbur

The fruit in this genus is a bur, covered with very sharp spines that penetrate the skin easily or adhere to clothing. The plants are troublesome weeds in cultivated fields.

Cenchrus echinatus L. Abundant from the Meseta Central to the coasts. Generally dispersed in tropical America.

Cenchrus viridis Spreng. Abundant in waste ground of the coasts, often invading cultivated ground. Generally distributed in tropical America.

Cenchrus pauciflorus Benth. Pacific coast, usually on beaches. Argentina to the United States.

CHAETIUM Nees

Chaetium bromoides (Presl) Benth. Setilla. Pastures and fields at low elevations; abundant in the Meseta Central and Canton de Dota. Extending to Mexico.

CHLORIS Swartz

Chloris orthonoton Doell. Pastures and fields, Meseta Central and probably in other regions. Mexico to Brazil.

FLORA OF COSTA RICA 73

Chloris petraea Swartz. Atlantic coast. Panama to southern United States.

Chloris radiata (L.) Swartz. Meseta Central to the coasts, often abundant in cultivated or abandoned fields, at 1,300 meters or less. Generally distributed in tropical America.

CHUSQUEA Kunth

The plants of this genus are tall bamboos, with thick or slender stems that are used in many ways in the regions where they grow. For Costa Rican species of this genus there have been reported the following names: Bejuco de canasta, Cafiuela; Uka (Bribri); Petara (Guatuso); Krugro, Uirba (TeYraba).

Chusquea Lehmannii Pilger. Forests of the volcanoes (Cascajal and Poas), 1,500-3,150 meters. Plants 3-6 meters high, forming dense thickets in forests. Also in Colombia.

Chusquea Meyeriana Rupr. Cascajal, at 1,650 meters, Lankes- ter 105. Also in Brazil and Bolivia.

Chusquea Pittieri Hack. Oesterr. Bot. Zeitschr. 53: 153. 1903. Cana brava. Cuesta de los Arrepentidos, between San Marcos and Santa Maria de Dota, 1,400 meters; known only from Canton de Dota. Endemic. The stems are 3-6 meters long.

Chusquea serrulata Pilger. Cana brava. Cerro de Las Vueltas and Volcan de Poas, 1,300-3,000 meters. Also in Panama and Colombia. A bamboo 3-6 meters high, often forming dense thickets. The stems, which are 2-5 cm. thick, are used locally for making rockets and a kind of firecracker.

Chusquea simpliciflora Munro. Rio Naranjo. Ranging from Guatemala to Panama.

Chusquea subtessellata Hitchc. Proc. Biol. Soc. Wash. 40: 81. 1927. Batamba. Paramos of Cerro de La Muerte, Tonduz3367; Cerro de Las Vueltas, 2,700-3,000 meters. Panama (?). Plants only 1-3 meters high, forming dense thickets in the paramos; stems thick and flexible, almost as hard as iron. The plant is said to be an important source of forage in the elevated region where it is found.

Chusquea Tonduzii Hack. Oesterr. Bot. Zeitschr. 53: 155. 1903. Known only from the summit of Volcan de Poas. Endemic.

Chusquea virgata Hack. Oesterr. Bot. Zeitschr. 53: 156. 1903. San Marcos, 1,355 meters, type; and El Copey. Endemic.

74 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XVIII

CINNA L.

Cinna poaeformis (HBK.) Scribn. & Merr. Slopes of the higher mountains, 2,000-3,000 meters. Mexico to Peru.

COIX L. Job's tears

Coix Lacryma- Jobi L. Lagrimas de San Pedro. Waste ground, Meseta Central and the coasts, preferring wet soil. Imported from the tropics of the Orient. The handsome, smooth and shining, gray seeds are used for making necklaces, rosaries, and other articles.

CORTADERIA Stapf

Cortaderia nitida (HBK.) Pilger. Bogs in the high mountains, often growing in sphagnum, 2,000-3,000 meters, frequent in Canton de Dota. Ranging to Peru. The genus Cortaderia, typically Andean, does not extend north of Costa Rica.

CYMBOPOGON Spreng.

Cymbopogon citratus (DC.) Stapf. Zacate de limon, Sontol. Planted commonly in gardens and often naturalized; native of India. All parts of the plant have a strong and agreeable odor of lemon. The rhizomes are employed for flavoring tobacco, and an infusion, te de limdn, is a popular domestic remedy.

CYNODON L. Rich. Bermuda grass

Cynodon Dactylon (L.) Pers. Capriola Dactylon Kuntze. Common in many places, Meseta Central to the coasts. A species of pantropic distribution. Perhaps the best lawn grass, certainly the one most used, for the tropics. It forms a very dense, close sod that thrives even during long dry seasons. When, however, Bermuda grass invades cultivated ground, it is a bad pest, extremely difficult of eradication.

DACTYLIS L. Orchard grass

Dactylis glomerata L. Naturalized in pastures in the region of El Copey; imported from Europe. Perhaps planted in some places as a pasture grass, a purpose for which it is much used in temperate regions.

DACTYLOCTENIUM Willd.

Dactyloctenium aegyptium (L.) Richt. A common grass of the coasts, growing in waste places, especially about the ports; naturalized from the Old World tropics.

FLORA OF COSTA RICA 75

DIECTOMIS HBK.

Diectomis fastigiata (Swartz) HBK. Fields and savannas at low elevations. Widely distributed in the tropics of both hemispheres.

DIGIT ARIA Heist. Crab grass

Digitaria argillacea (Hitchc. & Chase) Fernald. Atlantic coast. Guatemala to Panama; Greater Antilles.

Digitaria horizontalis Willd. Meseta Central to the coasts; common in many places. Widely distributed in tropical regions of both hemispheres.

Digitaria panicea (Swartz) Urban. Carrillos de Poas, Brenes 19303. Mexico to Panama and West Indies.

Digitaria sanguinalis (L.) Scop. Syntherisma sanguinalis Dulac. Abundant in waste or cultivated ground, Meseta Central to the coasts. Widely dispersed in both hemispheres.

Digitaria villosa (Walt.) Pers. Meseta Central. Ranging to the West Indies and southern United States.

ECHINOCHLOA Beauv. Barnyard grass

Echinochloa colonum (L.) Link. Panicum colonum L. Wet or swampy places, Meseta Central to the coasts; introduced from the tropics of the Old World.

Echinochloa Crus-galli (L.) Beauv. var. Crus-pavonis (HBK.) Hitchc. Cola de gallo. Wet or swampy places, Meseta Central to the coasts. Mexico and West Indies to Argentina.

ELEUSINE Gaertn.

Eleusine indica (L.) Gaertn. Meseta Central to the coasts; a common species of waste and cultivated ground. Introduced from the tropics of the Old World.

EPICAMPES Presl

Epicampes Emersleyi (Vasey) Hitchc. Meseta Central and in regions of less elevation; region of San Ramon. Panama to south- western United States. A coarse grass, as much as 1.5 meters high.

ERAGROSTIS Host

Eragrostis amabilis (L.) Wight & Arn. A delicate annual, growing near the coasts; introduced from the Old World.

76 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Eragrostis cilianensis (All.) Link. E. major Host; E. mega- stachya Link. Meseta Central to the coasts; a common species of waste and cultivated ground. Naturalized from the Old World.

Eragrostis ciliaris (L.) Link. An abundant grass of lowlands, growing in waste ground; probably introduced from the Old World. One of the most common weedy grasses almost throughout tropical America.

Eragrostis hypnoides (Lam.) BSP. Wet soil near the coasts. Brazil to the United States.

Eragrostis limbata Fourn. Frequent at low and middle eleva- tions; region of San Ramon. Mexico to Bolivia.

Eragrostis lugens Nees. Common at middle elevations. Southwestern United States to Argentina.

Eragrostis maypurensis (HBK.) Steud. Common in waste ground in the lowlands, and sometimes at higher elevations; region of San Ramon. Mexico to Brazil and Bolivia.

Eragrostis mexicana (Lag.) Link. Las Concavas, Prov. Car- tago. Southwestern United States to Chile. Unknown elsewhere in Central America.

Eragrostis prolif era (Swartz) Steud. Pacific coast ; a halophilous plant, growing on beaches. Central America to West Indies and Brazil.

Eragrostis secundiflora Presl. San Francisco de Guadalupe; the only Central American locality known for the species. Mexico and southwestern United States.

Eragrostis simpliciflora (Presl) Steud. Fields and savannas of the Pacific slope. Mexico to Panama.

Eragrostis tephrosanthos Schult. Fields and waste ground, at low or middle elevations. Southern United States to Brazil.

ERIOCHLOA HBK.

Eriochloa distachya HBK. In savannas at 1,000 meters or less. Guatemala to Paraguay.

Eriochloa punctata (L.) Desv. A perennial plant, growing in wet or boggy places, at 1,000 meters or less. Southern United States to Argentina.

FLORA OF COSTA RICA 77

ERIOCHRYSIS Beauv.

Eriochrysis cayennensis Beauv. A perennial grass, growing in swamps at low elevations. Mexico and West Indies to Uruguay.

FESTUCA L.

Festuca elatior L. Meadows of the volcanoes, probably planted as a pasture grass; introduced from Europe.

Festuca rubra L. Pastures of Volcan de Turrialba, introduced from Europe.

Festuca tolucensis HBK. Pastures of Volcan de Irazu. A Mexican species, unknown elsewhere in Central America.

It is probable that there grows on the Atlantic coast Guadua aculeata Rupr., a native bamboo as large asBambusa vulgaris, armed with hooked spines. The species is abundant in many parts of the coast from Guatemala to Panama, but there are extensive areas where it does not occur.

GYMNOPOGON Beauv. Gymnopogon f astigiatus Nees. Boruca. Extending to Brazil.

GYNERIUM Willd.

Gynerium sagittatum (Aubl.) Beauv. Cana blanca, Cana de Castillo,. G. saccharoides Humb. & Bonpl. Abundant in open swamps of the coasts, especially along the Reventazon, where it occupies large areas to the exclusion of most other plants. Southern Mexico to Paraguay. A giant grass, almost as large as a bamboo. Stems as thick as those of sugar cane, employed commonly for the construction of huts in the tierra caliente. In the larger towns the stems are employed in the better houses in place of laths. The stems have many other uses, in their utility being scarcely inferior to bam- boo. Pittier states that in Talamanca this grass is so important that the Indians have adopted its flowering season to mark the beginning of their summer. Among the indigenous names reported for the species are: Ukakul, Ukakur (Bribri); Kagru, Uka (Cabe"cara); Bak-kra (Brunka); Soro (Te"rraba).

HACKELOGHLOA Kuntze

Hackelochloa granularis (L.) Kuntze. Manisuris granularis Sw.; Rytilix granularis Skeels. Chiefly on the coasts, growing in waste places; San Ramon, 1,050 meters. Naturalized from the Old World.

78 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

HOLCUS L.

llolcus lanatus L. Mielilla. Notholcus lanatus Nash. Rather common in pastures of the volcanoes and in Santa Maria de Dota, 1,400-2,800 meters. Perhaps sown for pasture, or introduced by accident with seeds of other pasture grasses. Native of Europe. All parts of the plant are covered with a fine, soft pubescence. In Santa Maria I was given for this species the name of Raigra, which is employed ordinarily for Lolium perenne.

HOMOLEPIS Chase

Homolepis aturensis (HBK.) Chase. Atlantic coast, in wet places or swamps. Mexico to Brazil and Bolivia.

HYMENACHNE Beauv.

Hymenachne amplexicaulis (Rudge) Nees. Atlantic coast, in wet ground or swamps. Tropics of both hemispheres.

HYPARRHENIA Fourn.

Hyparrhenia bracteata (Humb. & Bonpl.) Stapf. Fields and savannas at low elevations. Mexico to Paraguay. A coarse perennial , 1-2 meters high.

ICHNANTHUS Beauv.

Ichnanthus axillaris (Nees) Hitchc. & Chase. Panicum axillare Nees. In forests of the lowlands. Central America and West Indies to Brazil and Ecuador.

Ichnanthus nemorosus (Swartz) Doell. Wet forests and thickets, abundant in many places, 1,000-2,000 meters; sometimes in localities of less elevation; Guanacaste. Mexico to Panama and West Indies.

Ichnanthus pallens (Swartz) Munro. Panicum pattern Swartz. Wet forests, region of Cartago to the coasts; region of San Ramon; Guanacaste; at 1,400 meters or less. Generally distributed in tropical America.

Ichnanthus tenuis Presl. Wet forests at low and middle eleva- tions; region of San Ramon. Ranging to Trinidad and Colombia.

IMPERATA Cyrillo

Imperata contracta (HBK.) Hitchc. In savannas at low elevations; Guanacaste, common in the region of Tilaran. Mexico and West Indies to Brazil and Chile.

FLORA OF COSTA RICA 79

ISACHNE R. Br.

Isachne arundinacea (Swartz) Griseb. A vine or a more or less clambering plant, its stems as much as 6 meters long, growing in wet forests and thickets, 500-2,000 meters; region of San Ramon. Mexico to Jamaica, Venezuela, and Bolivia.

Isachne polygonoides (Lam.) Doell. Wet savannas and swampy places, at low and middle elevations. Guatemala to Brazil.

ISCHAEMUM L.

Ischaemum latifolium (Spreng.) Kunth. Canton de Dota, and in other regions. Southern Mexico to West Indies, Brazil, and Ecuador.

IXOPHORUS Schlecht.

Ixophorus unisetus (Presl) Schlecht. Zacate de Honduras. Swampy places in the lowlands; abundant in Guanacaste, where it is sometimes planted for pasture. Mexico to Colombia.

LASIACIS (Griseb.) Hitchc.

Except for L. procerrima, the plants of this genus are vines or more or less clambering shrubs with elongate and branched stems.

Lasiacis divaricata (L.) Hitchc. Panicum divaricatum L. Abundant in forests and thickets of the lowlands; region of San Ramon. Florida and Mexico to West Indies and Argentina.

Lasiacis oaxacensis (Steud.) Hitchc. Tierra caliente of both coasts; abundant in Guanacaste. Mexico to Ecuador.

Lasiacis procerrima (Hack.) Hitchc. Proc. Biol. Soc. Wash. 24: 145. 1911. Panicum procerrimum Hack. Oesterr. Bot. Zeitschr. 51:431. 1901. Type collected near San Jose". Meseta Central to the coasts. Mexico to Venezuela. A coarse, erect, herbaceous plant, as much as 4 meters high.

Lasiacis rhizophora (Fourn.) Hitchc. Panicum rhizophorum Fourn. Wet forests, Meseta Central and on both slopes at lower elevations; mountains, at 500-2,000 meters. Extending to Mexico.

Lasiacis ruscifolia (HBK.) Hitchc. Panicum ruscifolium HBK. Wet thickets and forests of the coasts, ascending to 900 meters. Mexico and West Indies to Peru.

Lasiacis scabrior Hitchc. Meseta Central and other regions, at 1,300 meters or less. Guatemala to Panama.

80 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Lasiacis Sloanei (Griseb.) Hitchc. Panicum Sloanei Griseb. Turrialba. Mexico and West Indies to Colombia.

Lasiacis sorghoidea (Desv.) Hitchc. & Chase. Carricillo, Carri- cillo trepador. Panicum sorghoideum Desv. Common in the Meseta Central and other regions of middle elevation; region of San Ramon. Mexico and West Indies to Argentina and Bolivia. It is probable that the name Carricillo is given to all the Costa Rican species.

Lasiacis Standleyi Hitchc. Proc. Biol. Soc. Wash. 40: 86. 1927. La Tejona, north of Tilaran, Guanacaste, 800 meters, Standley & Valeria 45839. Wet forests of the mountains of Guancaste; region of San Ramon; El General. Endemic.

LEERSIA Swartz

Leersia grandiflora (Doell) Prodoehl. Meseta Central and regions of less elevation. Southern United States and Mexico to Brazil.

Leersia hexandra Swartz. Tepalon. Homalocenchrus hexandrus Kuntze. Swamps or wet soil at low and middle elevations, usually near the coasts. Southern United States and Mexico to Brazil.

LEPTOCHLOA Beauv.

Leptochloa filiformis (Lam.) Beauv. A common grass of the coasts. Generally distributed in tropical America.

Leptochloa scabra Nees. Wet soil of the coasts. Widely dispersed in tropical America.

Leptochloa virgata (L.) Beauv. Abundant in moist ground at low and middle elevations; region of San Ramon. Generally distributed in tropical America.

LEPTOCORYPHIUM Nees

Leptocoryphium lanatum (HBK.) Nees. Dry or sterile places, Meseta Central and in other regions of slight or middle eleva- tion. Mexico and West Indies to Argentina.

LITHACHNE Beauv.

Lithachne pauciflora (Swartz) Beauv. Wet forests of the coasts, ascending to 2,000 meters. Mexico and West Indies to Argentina. A low plant with broad leaves. Referred by many authors to the genus Olyra.

FLORA OF COSTA RICA 81

LOLIUM L.

Lolium perenne L. Raigras, Raigrd. Commonly sown for pasture in almost all the meadows of the volcanoes. Imported from Europe. The local name is, of course, a modification of the English "rye grass."

MELINIS Beauv.

Melinis minutiflora Beauv. Zacate gordura. A grass of African origin, sometimes sown for pasture, and completely naturalized in some places, as in Santa Maria de Dota; region of San Ramon. This plant has a strong odor, and possesses properties that seem to repel insects. It is claimed that cattle pasturing in fields of it are free from ticks.

MEROSTACHYS Spreng.

Merostachys multiramea Hack. Carrizo. La Palma de San Ramon, 1,050 meters, Brenes 51$3. Also in Brazil. A slender bamboo, growing in forest clearings. The genus is unknown else- where in North America.

MUHLENBERGIA Schreb.

Muhlenbergia ciliata (HBK.) Kunth. In shaded places, 700- 2,000 meters. Mexico to Panama.

Muhlenbergia diversiglumis Trin. Meseta Central, forests and on open banks, 1,000-1,800 meters. Mexico to Peru.

Muhlenbergia implicata (HBK.) Kunth. Meseta Central and other regions of middle elevation, 1,000-2,000 meters. Mexico to Venezuela.

Muhlenbergia quadridentata (HBK.) Kunth. M. flabellata Mez, Repert. Sp. Nov. 17: 213. 1921 (Cerro de Buena Vista, Pittier}. Known in Central America only from Cerro de Buena Vista, at 3,100 meters. Also in Mexico.

Muhlenbergia setarioides Fourn. In forest or other shaded places, Meseta Central and slopes of the volcanoes, 1,000-2,000 meters. Mexico to Panama.

Muhlenbergia tenella (HBK.) Trin. Zacate de seda. A common species of the Meseta Central, in shaded places. Mexico to Panama.

OLYRA L.

Olyra caudata Trin. 0. Pittieri Hack. Oesterr. Bot. Zeitschr. 51: 461. 1901. Known in North America only from forests of El

82 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Cordoncillal, 300-500 meters, type locality of 0. Pittieri. British Guiana to Brazil and Peru.

Olyra lateralis (Presl) Chase. In forests at 700-1,100 meters. A South American species, known in Central America only from Costa Rica. It has been collected in El General and elsewhere.

Olyra latifolia L. Gamalote. Common in wet forests of the coasts; Guanacaste. A coarse grass resembling a bamboo, sometimes as much as 5 meters high but usually lower. Mexico and West Indies to Brazil and Bolivia.

Olyra Standleyi Hitchc. Proc. Biol. Soc. Wash. 40: 86. 1927. Wet forest, El Murleco, Prov. Cartago, 1,400-1,500 meters, Standley & Torres 50932. Endemic ; known only from the region of El Muneco.

OPLISMENUS Beauv.

Reference: A. S. Hitchcock, The North American species of Oplismenus, Contr. U. S. Nat. Herb. 22: 123-132. 1920.

Oplismenus Burmannii (Retz.) Beauv. Zacate de raton. 0. Humboldtianus Nees. Meseta Central to the coasts; abundant in moist places of the coasts, especially in banana plantations. Pan- tropic in distribution. Said to be a native of the Old World, but to one who has seen its present abundance in America, this seems improbable. It is one of the most common weeds of Central America.

Oplismenus hirtellus (L.) Beauv. Meseta Central to the coasts, an abundant grass of forests and wet places. Generally distributed in tropical America.

ORTHOCLADA Beauv.

Orthoclada laxa (L. Rich.) Beauv. Common in forests of the Atlantic coast. Mexico to Brazil and Peru.

ORYZA L.

Oryza latifolia Desv. Wet or swampy places of the Atlantic coast, perhaps also on the Pacific slope. A native plant that it is almost impossible to distinguish from cultivated rice. It is said that in the Amazon region its seeds have been employed for food. Guatemala and West Indies to Brazil.

Oryza sativa L. Arroz. Rice. A plant native in the Orient. Rice of good quality, but insufficient for local consumption, is grown in all inhabited portions of the Pacific slope and in the plains of San

FLORA OF COSTA RICA 83

Carlos and Sarapiqui. Only the upland variety is grown, this needing no irrigation in its cultivation. Indigenous names reported for rice are: lok-koro (TeYraba); Sunoji-ku (Guatuso).

PANICUM L.

Reference: A. S. Hitchcock and A. Chase, Tropical North Ameri- can species of Panicum, Contr. U. S. Nat. Herb. 17: 459-539. 1915.

The largest genus of grasses, composed of about 500 species. From Central America 65 species are known.

Panicum altum Hitchc. & Chase. Buenos Aires. British Honduras to Panama; Trinidad and Tobago.

Panicum boliviense Hack. Reported as collected in El General by Skutch. Guatemala to Paraguay.

Panicum cayennense Lam. Buenos Aires. Central America and West Indies to Brazil.

Panicum cordovense Fourn. Piedra del Convento, Pittier. Mexico to Brazil and Bolivia.

Panicum fasciculatum Swartz. P. fuscum Swartz. On the coasts and sometimes in regions of middle elevation; common in waste places and often in cultivated ground. Generally distributed in tropical America.

Panicum frondescens Mey. Swampy places along the Atlantic coast. Mexico to Brazil.

Panicum geminatum Forsk. Wet or swampy places, Guana- caste (Bebedero) and probably in other regions. Tropics of both hemispheres.

Panicum Ghiesbreghtii Fourn. Meseta Central to the Pacific coast. Mexico to Brazil and Bolivia.

Panicum glutinosum Swartz. Wet forests at middle eleva- tions; region of San Ramon. Mexico and West Indies to Paraguay.

Panicum Haenkeanum Presl. P. costaricense Hack. Oesterr. Bot. Zeitschr. 51: 428. 1901 (between Buenos Aires and TeYraba, Pittier 3636). Reported also from Boruca and El Cordoncillal. Mexico to Venezuela.

Panicum helobium Mez. Swampy places in forests, region of Santa Maria de Dota, 1,500-1,800 meters. A South American species, unknown elsewhere in Central America.

84 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Panicum hirsu turn Swartz. Wet places of both coasts. Mexico and West Indies to Brazil and Ecuador.

Panicum hirticaule Presl. Pacific coast, and probably in other regions. Southwestern United States to Bolivia.

Panicum laxum Swartz. Tepaldn. Abundant on the coasts, and sometimes in regions of middle elevation; region of San Ramon. Mexico and West Indies to Paraguay.

Panicum maximum Jacq. Guinea, Zacate de Guinea. Probably native of Africa, planted almost everywhere, but especially in the lowlands, as pasture for stock; Cocos Island. Pittier states that the seed was first brought to Costa Rica about 1885 by Don San- tiago Millet. This is considered the best pasture grass everywhere in the tierra caliente of Central America.

Panicum megiston Schult. Guanacaste, and probably in swamps of other regions. Mexico and Cuba to Paraguay.

Panicum molle Swartz. Pacific coast (Puntarenas), and prob- ably in other regions. Mexico and Cuba to Argentina.

Panicum olivaceum Hitchc. & Chase. Meseta Central and slopes of the volcanoes; Canton de Dota; common in some regions, 1,200-1,800 meters. Mexico to Colombia and Venezuela.

Panicum orbiculatum Poir. Meseta Central and Atlantic slope. Southern Mexico to West Indies and Paraguay.

Panicum parvifolium Lam. Savannas of Buenos Aires. Not known elsewhere in Central America. West Indies to Paraguay.

Panicum parviglume Hack. Oesterr. Bot. Zeitschr. 51: 429. 1901. Rio Torres, near San Jose", Pittier 9080. Meseta Central. Also in Mexico.

Panicum pilosum Swartz. Common in the lowlands, in wet or swampy places. Mexico and West Indies to Paraguay.

Panicum polygonatum Schrad. Common in wet places of the tierra caliente; region of San Ramon; Cocos Island. Mexico to Paraguay.

Panicum pulchellum Raddi. Meseta Central and in the low- lands; a plant of shaded places; region of San Ramon. Mexico and West Indies to Brazil.

Panicum purpurascens Raddi. Para, Zacate de Para. P. barbinode Trin.; P. molle of some authors, not of Swartz. Planted generally for forage on the Atlantic slope, up to 1,400 meters; also

FLORA OF COSTA RICA 85

on the Pacific slope. Perhaps introduced to Costa Rica from Brazil or elsewhere. Tropics of both hemispheres.

Panicum Rudgei Roem. & Schult. Savannas of El General, Buenos Aires, and other localities of that general region. Central America and West Indies to Brazil.

Panicum Schiffneri Hack. Meseta Central. Mexico and West Indies to Brazil.

Panicum Sellowii Nees. El General. Mexico and West Indies to Paraguay.

Panicum sphaerocarpon Ell. Meseta Central and Canton de Dota, 1,200-1,800 meters. Eastern United States to Venezuela.

Panicum stenodes Griseb. Buenos Aires. Guatemala and West Indies to Brazil.

Panicum stenodoides Hubbard. Buenos Aires. British Hon- duras, Panama, and Trinidad.

Panicum strigosum Muhl. Meseta Central. Colombia to West Indies and United States.

Panicum trichanthum Nees. Common in wet forests of the Atlantic coast. Mexico and West Indies to Paraguay.

Panicum trichoides Swartz. Abundant in the lowlands, ascending to at least 1,100 meters. One of the most common weedy grasses of all the tierra caliente of Central America. Dispersed throughout tropical America.

Panicum virgultorum Hack. Oesterr. Bot. Zeitschr. 51: 369. 1901. La Verbena, near Alajuelita, Tonduz 8829. A common species of the Meseta Central; region of San Ramon. Mexico to Panama.

Panicum viscidellum Scribn. Meseta Central; El Muneco; El General. Mexico and Cuba to Colombia.

Panicum xalapense HBK. Regions of Cartago and Dota. Extending to the United States; Hispaniola.

Panicum zizanioides HBK. Wet or swampy places of the Atlantic coast. Mexico and West Indies to Paraguay.

PARIANA Aubl.

Pariana zingiberina Doell. Wet forests of the Atlantic coast. Ranging to Brazil. The genus, otherwise South American, does not extend north of Costa Rica.

86 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

PASPALUM L.

Reference: Agnes Chase, The North American species of Pas- palum, Contr. U. S. Nat. Herb. 28: 1-310. 1929.

One of the largest genera of grasses, with 380 species, of which at least 67 are known in Central America.

Pa spa 1 urn candidum (Humb. & Bonpl.) Kunth. Common in the Meseta Central and slopes of the volcanoes; region of San Ramon; 1,100-1,800 meters. Southern Mexico to Chile.

Paspalum centrale Chase. Pacific Coast (Puntarenas and Atenas). Salvador to Panama.

Paspalum clavuliferum C. Wright. P. Pittieri Hack, ex Beal, Grasses N. Amer. 2: 88. 1896. Llanos de Turrucares, Pittier. Southern Mexico and Cuba to Brazil.

Paspalum conjugatum Berg. Turvara. Meseta Central to the coasts, abundant in wet or waste places, often in cultivated ground. Pantropic in distribution.

Paspalum convexum Humb. & Bonpl. Meseta Central to the Pacific coast. Mexico to Brazil.

Paspalum costaricense Mez, Repert. Sp. Nov. 15: 72. 1917. Zacate de caballo. San Jose", Tonduz. Meseta Central and Canton de Dota. Also in Guatemala and Salvador.

Paspalum decumbens Swartz. Common in the tierra caliente, ascending to 900 meters; Cocos Island. Guatemala and West Indies to Brazil and Bolivia.

Paspalum dilatatum Poir. Meseta Central; meadows of Volcan de Turrialba. Believed to be a native of South America, but now widely introduced elsewhere. It is said to have considerable value as a pasture grass.

Paspalum distichum L. Meseta Central and at middle elevations on the Atlantic slope. Widely distributed in America.

Paspalum fasciculatum Willd. Gamalote. Meseta Central and on the coasts, abundant in many places. The Guatuso name for the species is reported by Lehmann as Tonun.

Paspalum Humboldtianum Fluegge. Meseta Central. Mex- ico to Argentina.

Paspalum Jimenezii Chase, Contr. U. S. Nat. Herb. 28: 159. /. 101. 1929. Rio Bebedero, Las Playitas, Guanacaste, Otdn Jimenez 742. Endemic, and known only from the original collection.

FLORA OF COSTA RICA 87

Paspalum Langei (Fourn.) Nash. Reported for Costa Rica by Hitchcock and Chase, without indication of the locality. Southern United States to Venezuela.

Paspalum lineare Trin. Known on the North American conti- nent only from Cabagra, where it was collected by Tonduz. Cuba to Argentina.

Paspalum microstachyum Presl. On the coasts. Guatemala to Ecuador and Brazil.

Paspalum minus Fourn. A grass of the coasts; collected also at San Jose". Mexico and West Indies to Paraguay.

Paspalum multicaule Poir. Grecia; Buenos Aires; Boruca; region of San Ramon, ascending to 1,050 meters. Southern Mexico and West Indies to Brazil.

Paspalum notatum Fluegge. Gengibrillo. Meseta Central to the coasts. Mexico and West Indies to Argentina.

Paspalum nutans Lam. Pejivalle; San Jose"; El General. Central America and West Indies to Brazil.

Paspalum paniculatum L. Abundant in many places, Meseta Central to the coasts. Tropics of both hemispheres.

Paspalum pectinatum Nees. Guanacaste;Puntarenas. Hon- duras to Brazil.

Paspalum pictum Ekman. P. maculatum Nash, N. Amer. Fl. 17: 186. 1912 (savannas of Boruca, Pittier U?4). Ranging to Brazil and Bolivia. Known in North America only from Costa Rica.

Paspalum pilosum Lam. Meseta Central; Turrialba; Pacific slope. Ranging to Brazil and Bolivia.

Paspalum plenum Chase. Nuestro Amo; Agua Caliente. Southern Mexico to Colombia.

Paspalum plicatulum Michx. Meseta Central to the coasts. Southern United States to Argentina.

Paspalum propinquum Nash. Puntarenas. Florida and Mexico to Panama.

Paspalum repens Berg. Guanacaste and Puntarenas; an aquatic plant. Southern United States to Paraguay.

Paspalum saccharoides Nees. Atlantic coast; region of San Ramon. Extending to West Indies and Bolivia.

88 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Paspalum squamulatum Fourn. Meseta Central and slopes of the volcanoes; Canton de Dota; in pastures and shady places. Ranging to Mexico.

Paspalum stellatum Humb. & Bonpl. Boruca. Southern Mexico to Argentina. A species easy of recognition because of the broad, yellow wings of the axis of the flower spike.

Paspalum Tonduzii Mez, Repert. Sp. Nov. 15: 72. 1917. In cornfields, Santa Rosa del Copey, 1,800 meters, Tonduz 11767. Endemic.

Paspalum vaginatum Swartz. Atlantic coast. Southern United States to Argentina and Chile.

Paspalum variabile (Fourn.) Nash. Guanacaste. A Mexican species, unknown elsewhere in Central America.

Paspalum virgatum L. Meseta Central to the coasts; Cocos Island; abundant in many places, growing in moist or swampy ground. Texas to West Indies and Brazil.

PENNISETUM L. Rich.

Reference: Agnes Chase, The North American species of Penni- setum, Contr. U. S. Nat. Herb. 22: 209-234. 1921.

Pennisetum bambusiforme (Fourn.) Hemsl. Guapiles, El Copey, and probably in other regions, at 1,800 meters or less; region of San Ramon. Mexico to Peru. Stems branched, as much as 5 meters high; flowers arranged in dense spikes 5-12 cm. long.

Pennisetum distachyum (Fourn.) Rupr. Chiefly in the Meseta Central, in thickets and on stream banks, also at San Ramon. Mexico and Central America. Stems 1-4 meters high.

Pennisetum purpureum Schumacher. Yerba elefante. Native of Africa, planted in some regions (Dota, etc.) for forage. Spikes conspicuously tinged with purple.

Pennisetum setosum (Swartz) L. Rich. Pacific slope, at low elevations, in fields and savannas, forming dense clumps 1-2 meters high. Widely distributed in tropical America.

Pennisetum vulcanicum Chase. Nuestro Amo. Also in Salvador.

FLORA OF COSTA RICA 89

PEREILEMA Presl

Pereilema Beyrichianum (Kunth) Hitchc. Llano Grande de Puriscal. A South American species, known in North America only from Costa Rica.

Pereilema crinitum Presl. Meseta Central and regions of middle elevation, growing on moist banks and in shaded places. Mexico to Ecuador.

PHALARIS L.

Phalaris arundinacea L. var. picta L. Zacate de listdn. A European grass, grown for ornament in gardens. The green leaves are striped with yellowish white. The plant rarely blooms in cultivation.

PHARUS L.

Pharus cornutus Hack. Oesterr. Bot. Zeitschr. 52: 9. 1902. An endemic grass, growing in moist or wet places at low elevations, the type collected at Tsaki; at 200-500 meters.

Pharus glaber L. P. Mezii Prodoehl, Bot. Archiv Mez 1 : 250. 1922 (type from Costa Rica). Abundant in wet forests of the tierra caliente, ascending to 1,100 meters. Mexico and Guatemala to Brazil. The large leaves are lanceolate and 3-5 cm. wide. The Bribri name is reported as Tsubuk-uo.

Pharus latifolius L. Yerba de hierro (Cufodontis). Common in wet forests of the coast regions. Guatemala and West Indies to Brazil.

Pharus parvifolius Nash. Wet forests of the tierra caliente; common in Guanacaste. Mexico and West Indies to Brazil.

PHRAGMITES Adans.

Phragmites communis Trin. Abundant in swamps of coast regions, especially on river banks, sometimes forming colonies of large extent. In temperate regions almost throughout the earth, and extending into the tropics. A coarse plant, 2-3 meters high, its stems as thick as a finger, the leaves ashy green. The stems are employed for many purposes by the people of the coastal regions of Costa Rica.

POA L.

Poa annua L. Zacate de raton. Abundant in almost all meadows of the higher mountains; growing on Poas Volcano in sphagnum

90 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

bogs. An annual grass, here doubtless imported from Europe. Widely dispersed in temperate regions of both hemispheres.

Poa pratensis L. Pastures of Volcan de Turrialba, and prob- ably in those of other volcanoes. Imported from Europe, and widely naturalized in temperate regions of America. A favorite pasture and lawn grass (bluegrass) in many parts of the earth.

POLYPOGON Desf.

Polypogon elongatus HBK. Common in the Meseta Central and pastures of the volcanoes; region of Dota; growing in swamps or wet meadows. Mexico to Argentina.

PSEUDECHINOLAENA Stapf

Pseudechinolaena polystachya (HBK.) Stapf. Meseta Cen- tral; Pejivalle; on shaded banks or sometimes in cultivated fields, 900-1,500 meters. Mexico to Bolivia and Uruguay; tropical Africa.

RADDIA Bertol.

Raddia concinna (Hook, f.) Chase, Proc. Biol. Soc. Wash. 21: 185. 1908. Olyra concinna Hook. f. in Curtis' Bot. Mag. III. 52: pi. 7469. 1896. Wet forests of the Atlantic coast (Hamburg Finca). Known only from Costa Rica; described from plants believed to have grown from Costa Rican seeds in greenhouses of the Kew Gardens, London. A low perennial with short, broad leaves, forming small, dense tufts somewhat suggestive of a tiny bamboo.

Raddia costaricensis Hitchc. Proc. Biol. Soc. Wash. 40: 87. 1927. Forests of Rio Hondo, near Madre de Dios, 200 meters, Pittier 10352. Endemic.

ROTTBOELLIA L. f.

Rottboellia aurita Steud. Savannas of the tierra caliente. Extending to Argentina and Bolivia.

SACGHARUM L. Sugar cane

Saccharum officinarum L. Cana, Cana de azucar. A native of the Orient, sugar cane is one of the most important economic plants of Costa Rica. It is cultivated chiefly in the temperate regions but also in almost every part of the tierra caliente. Two main varieties are grown locally, yellow and purple, the former chiefly as a source of sugar, the latter for forage for stock. Among indigenous names cited for this plant are the following: Ipacru,

FLORA OF COSTA RICA 91

Ipakur (Bribri); Pashtu (Cabe"cara); Soror-bo (T&raba); Budi (Boruca); Afo-fora (Guatuso).

SETARIA Beauv.

Reference: A. S. Hitchcock, The North American species of Chaetochloa, Contr. U. S. Nat. Herb. 22: 155-208. 1920. (The generic name Chaetochloa is a synonym of Setaria.}

Setaria geniculata (Lam.) Beauv. Abundant in almost all inhabited regions, Meseta Central to the coasts; often abundant in cultivated ground. Generally distributed in tropical America, but without economic value, since it is not useful for pasture or forage. A low, perennial grass with dense, yellowish flower spikes.

Setaria magna Griseb. Pacific coast. Ranging to the West Indies and United States. An annual as much as 4 meters high.

Setaria paniculifera (Steud.) Fourn. Zacate de mula. Meseta Central to the coasts, in wet forests or swamps. Mexico to West Indies and Colombia. A perennial grass, sometimes 4 meters high, the leaves as much as 10 cm. wide.

Setaria scandens Schrad. Meseta Central. Guatemala to West Indies and Paraguay.

Setaria tenacissima Schrad. Meseta Central; Canas Gordas. Guatemala to West Indies and Brazil.

Setaria viridis (L.) Beauv. Pastures near Cartago; the only Central American locality known for the species, which is European in origin. In the United States this grass is one of the worst pests of cultivated ground.

Setaria vulpiseta (Lam.) Roem. & Schult. Atlantic coast and probably in other regions; region of San Ramon. Southern Mexico to West Indies, Argentina, and Peru. A coarse plant, as much as 2 meters high.

SORGHASTRUM Nash

Sorghastrum incompletum (Presl) Nash. Dry fields and savannas at low elevations. Mexico to Venezuela; tropical Africa. It is probable that there grows in Costa Rica also S. nutans (L.) Nash, which is known from Honduras and Panama.

SORGHUM Moench

Sorghum halepense Pers. Holcus halepensis L. Waste or cultivated ground. Naturalized from Europe, but not common in

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Costa Rica. In the United States this (Johnson grass) is considered one of the worst weeds of cultivated land.

Sorghum vulgare Pers. Maicillo, Maiz de millo. Holcus Sor- ghum L. A plant of the Old World, cultivated in some parts of Costa Rica as forage and for its seeds. A tall plant, much like maize in habit, its spikes of fruit forming a large, rounded inflorescence. A form of the species with large, open panicles is cultivated for use in manufacture of brooms.

SPARTINA Schreb.

Spartina Spartinae (Trin.) Merrill. S. Pittieri Hack. Oesterr. Bot. Zeitschr. 52: 237. 1902 (type from Costa Rica). A grass of sea beaches, known in Central America only from Limon. Ranging to Mexico and southern United States.

SPOROBOLUS R. Br.

Sporobolus cilia tus Presl. Savannas of the Pacific slope. Honduras to Brazil.

Sporobolus cubensis Hitchc. Savannas of Cafias Gordas. Extending to the West Indies, Venezuela, and British Guiana.

Sporobolus elongatus R. Br. Meseta Central and slopes of the volcanoes, ascending to 2,000 meters; also in the lowlands. Widely distributed in tropical America, perhaps naturalized from Asia.

Sporobolus indicus (L.) R. Br. Pitilla. Savannas and dry fields at low elevations. Generally distributed in tropical America.

Sporobolus littoralis (Lam.) Kunth. A halophilous plant of sea beaches. Florida to Brazil.

Sporobolus minutissimus (Steud.) Hitchc. S. confusus Vasey. Meseta Central and regions of less elevation. Extending to western United States.

Sporobolus purpurascens (Swartz) Hamilt. In fields at low or middle elevations. Mexico and West Indies to Brazil and Bolivia.

Sporobolus ramulosus (HBK.) Kunth. Pastures of Irazu. Extending to southwestern United States.

STENOTAPHRUM Trin.

Stenotaphrum secundatum (Walt.) Kuntze. Common in the coasts. Southern United States to Argentina.

FLORA OF COSTA RICA 93

STIPA L.

Stipa Ichu (Ruiz & Pavon) Kunth. Cerros de Escasu; abundant on grassy slopes, the long, slender, and very tough leaves forming dense clumps. Mexico to Chile. This is the celebrated ichu grass of the Andean paramos, where it is often the dominant species, and is used extensively for thatch and fuel.

STREPTOCHAETA Schrad.

Streptochaeta Sodiroana Hack. Wet forests of the Atlantic coast. Honduras to Ecuador. A coarse and showy grass with broad leaves as much as 7 cm. wide.

STREPTOGYNE Beauv.

Streptogyne crinita Beauv. Wet forests of the Atlantic coast. Mexico to Trinidad and Brazil. A perennial grass, 1-1.5 meters high, its leaves 1-1.5 cm. wide.

THRASYA HBK.

Thrasya campylostachya (Hack.) Chase, Proc. Biol. Soc. Wash. 24: 115. 1911. Panicum campylostachyum Hack. Oesterr. Bot. Zeitschr. 51: 367. 1901. Type collected at Canas Gordas. Ascending to 1,200 meters. Guatemala to Bolivia.

TRACK YPOGON Nees

Trachypogon Montufari (HBK.) Nees. Dry fields and savan- nas at low elevations. Southwestern United States to Uruguay i

Trachypogon Montufari var. mollis (Nees) Anderss. In savannas. A form of the species with densely villous foliage.

TRICHACHNE Nees

Trichachne insular is (L.) Nees. Valota insularis Chase. A common grass of fields and pastures, Meseta Central to the coasts. Generally distributed in tropical America. The soft, silky flower panicles are used for decorating altars and nacimientos.

Trichachne Pittieri (Hack.) Hitchc. Proc. Biol. Soc. Wash. 40: 83. 1927. Panicum Pittieri Hack. Oesterr. Bot. Zeitschr. 51: 367. 1901. Valota Pittieri Chase, Proc. Biol. Soc. Wash. 19: 188. 1906. Meseta Central, 1,000-2,000 meters, the type from Rio Tiribi. Endemic.

TRICHOLAENA Schrad.

Tricholaena rosea Nees. Ilusion, Zacate de seda. An African grass, thoroughly naturalized in the Meseta Central, Pacific coast,

94 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

and probably other regions, abundant in many places. It is com- mon in many parts of Central America. The silky panicles are handsomely colored with pink or purple.

TRINIOCHLOA Hitchc.

Triniochloa stipoides (HBK.) Hitchc. Meadows of the higher mountains, 2,000-3,000 meters. Mexico to Bolivia.

TRIPSAGUM L.

Tripsacum laxum Nash. Maicillo, Cana de la India. Planted on the coasts and in regions of middle elevation for forage; natural- ized in some places. Mexico to Colombia. A tall grass as much as 5 meters high, the stems 2.5 cm. thick.

TRISETUM Pers.

Trisetum irazuense (Kuntze) Hitchc. Proc. Biol. Soc. Wash. 40: 82. 1927. Calamagrostis irazuensis Kuntze, Rev. Gen. 763. 1891 (Volcan de Irazu, Kuntze). Meadows of Irazu and Turrialba, 2,000-3,000 meters. Ranging to Ecuador.

Trisetum Pringlei (Scribn.) Hitchc. Paramos of Cerro de Las Vueltas, 3,000 meters. Panama to Mexico.

Trisetum viride (HBK.) Kunth. Meadows of the high moun- tains, 2,000-2,500 meters. A Mexican species, known in Central America only from Costa Rica.

TRITICUM L.

Triticum aestivum L. Trigo. Wheat is said to grow well in the temperate regions of Costa Rica, and has been planted on a small scale, but it is grown only experimentally.

UNIOLA L.

Uniola Pittieri Hack. Oesterr. Bot. Zeitschr. 52: 309. 1902. Type collected at the Bay of Salinas; Pacific coast, on sea beaches. Mexico to Ecuador. A tall, coarse grass, with large panicles of handsome, flat spikelets 1-2 cm. long.

ZEA L. Maize

Zea Mays L. Maiz. Maize and beans are the two important Costa Rican products for local consumption. Maize is cultivated from sea level to the uppermost limit of cultivation on the higher mountains, three crops a year being obtained sometimes in the tierra

FLORA OF COSTA RICA 95

caliente, one only in the cold regions. All the corn grown is of the hard-kerneled or flint type. Believed to be of Mexican origin, maize has never been found in a wild state. Its only close relative that does grow wild is teosinte (Euchlaena), of Guatemala and Mexico. It is believed by some authorities that maize is a hybrid between teosinte and some other unknown grass. It has been grown in Costa Rica doubtless since the very beginning of agriculture, and reached Peru long before the Spanish conquest. Indigenous Costa Rican names reported for maize are: Ik (Cabe"cara, Bribri); Ip (Te"rraba), Ai (Guatuso); Rukra (Brunka); Hoie" (Talamanca); Ko-ep, Cup (Boruca).

ZEUGITES P. Br.

Zeugites mexicana (Kunth) Trin. Senites mexicana Hitchc. Meseta Central, Canton de Dota, and probably elsewhere; shaded places, often abundant, 1,000-2,000 meters. Mexico to Bolivia.

Zeugites Pittieri Hack. Oesterr. Bot. Zeitschr. 52: 373. 1902. Wet forest, 500-1,100 meters. Type from Alto del Rodeo. Endemic.

CYPERAGEAE. Sedge Family

References: C. B. Clarke, The Cyperaceae of Costa Rica, Contr. U. S. Nat. Herb. 10: 443-471. 1902; Paul C. Standley, The Cyper- aceae of Central America, Field Mus. Bot. 8: 239-292. 1931.

Although so similar to grasses in general appearance, the Cyper- aceae have few or none of the important economic uses of that family.

CALYPTROCARYA Nees

Calyptrocarya glomerulata (Brongn.) Urban. C. fragifera Kunth. Atlantic coast; Cocos Island. British Honduras to South America.

CAREX L.

The largest genus of the family, with numerous species in most temperate and arctic regions, but with very few in the tropics, and then not in the hotter regions.

Carex albolutescens Schwein. Slopes of the volcanoes, 1,500- 1,800 meters, wet places, often in sphagnum bogs. United States to northern South America.

Carex Bonplandii Kunth. C. heptastachya Boeckl. Linnaea 39: 114. 1875 (based upon Costa Rican material); C. Durandii Boeckl. Allgem. Bot. Zeitschr. 2: 189. 1896 (type from Costa Rica).

96 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XVIII

Higher mountains, 2,400-3,100 meters. A South American species, unknown north of Costa Rica.

Carex Donnell-Smithii Bailey. C. viridis Boeckl. Linnaea 40: 330. 1876, ex parte, non C. viridis Jungh.; C. Pittieri Boeckl. Allgem. Bot. Zeitschr. 2: 190. 1896 (type from Costa Rica). High mountains, 2,500-3,300 meters, sometimes in sphagnum bogs. Listed by Clarke under the names C. Jamesonii Boott and C. pichinchensis HBK.

Carex Humboldtiana Steud. Forests of the temperate region, 1,200-1,800 meters. Mexico and West Indies to northern South America.

Carex Jovis Clarke, Contr. U. S. Nat. Herb. 10: 470. 1908. Without locality, Pittier & Tonduz 10757.

Carex Lemanniana Boott. Higher mountains, at 2,000 meters or more. Southward to Ecuador. Reported from Costa Rica under the names C. pichinchensis HBK. and C. Jamesonii Boott.

Carex polystachya Swartz. C. cladostachya WahL; C. acrolepis Liebm. Dansk. Vid. Selsk. Skrivt. V. 2: 270. 1851 (type from Costa Rica); C. Oerstedii Liebm. op. cit. 272. 1851 (type from Costa Rica). Common in forests of the Meseta Central and on the higher moun- tains, 1,000-2,500 meters; region of San Ramon. Mexico and West Indies to northern South America. The most common species of Carex in Central America.

Carex Purdiei Boott. Forests of the higher mountains, 2,000- 3,000 meters. Southward to Ecuador. A characteristic species of the paramos of Cerro de Las Vueltas, often growing with sphagnum.

CYPERUS L.

The largest genus of Cyperaceae in tropical regions, well repre- sented in Costa Rica as well as elsewhere in Central America.

Cyperus alternifolius L. An aquatic plant, originally from Madagascar, cultivated in gardens, especially in fountains. It is so similar to the native C. canus Presl that it is difficult to distinguish the two species in herbarium specimens.

Cyperus amabilis Vahl. Delta of Tsuritkub, Tonduz 8606 (fide Kiikenthal). Widely distributed in tropical and subtropical America and Africa. The Costa Rican collection is referred by Kiikenthal to var. macrostachyus (Boeckl.) Kiikenth.

Cyperus articulatus L. Common in the tierra caliente and perhaps in regions of greater elevation; growing in shallow water

FLORA OF COSTA RICA 97

or wet soil. Generally distributed in tropical regions of the earth. Easily recognized by its leafless, terete, spongy stems with numerous cross partitions.

Cyperus canus Presl. Tierra caliente. Extending to Mexico. In Salvador this species is cultivated commonly for its tough, some- what spongy stems which are used for making the mats generally used as mattresses in Central America.

Cyperus caracasanus Kunth. Mariscus flabelliformis HBK. ; C. breviradiatus Liebm. Dansk. Vid. Selsk. Skrivt. V. 2: 26. 1851.