Biol. J. Linn. SOC.,1, pp. 19-30.

With 5 figures
April 1969

Patterns of communities in the tropics

ROBERT H. MAcARTHUR
Department of Biology, Princeton University, Princeton, New Jersey, U.S.A.

Tropical countries have many times more species of most taxa than temperate ones, and small
areas in the tropics have a smaller multiple of the number of species of small temperate areas.
Where many species are present, abundances tend to be more equal and geographic distribu-
tions more spotty. Most tropical environments are less seasonal and more productive, and the dry
areas and mountains which are relatively more seasonal and less productive have fewer species.
The species which have reached offshore islands are often much commoner there and occupy
expanded habitats.
T o account for these relations, the following general hypothesis seems necessary : species
interactions are important and the tropics have a head start on speciation. The head start,
or greater rate, allows extra species to pile up in the tropics, but because of the importanee of
competition, no single area becomes as greatly enriched. Rather, faunal differences between
areas increase. T h e lesser excess of tropical species in small areas is largely due to greater
productivity and reduced seasonality which make marginal ways of life profitable. With more
overlap in resources, the closely packed tropical species have more uniform abundances and
the coexistence of these species is more precarious, causing the spotty geographic distributions.
Neither the species diversity of the food supply nor the longer breeding season (supposedly
allowing staggered nesting seasons with an early shift and a later shift) is relevant to bird
species diversity.

CONTENTS
PACE
Introduction . . . . . . . . . . .
. . 19
The tropical environment . . . . . . .
. . . . 20
Community properties in the tropics .
An hypothesis to account for these properties.
. . . . .. ..
. . ..
. 21
26
. .
Acknowledgements
References . .. . . . . . . . . .
.
.
29
29

INTRODUCTION
The wet tropical lowland environment is different from that in temperate regions,
and in this tropical environment plant and animal communities exhibit various extreme
patterns, As is well known, there are more species of most taxa in the tropics. Coupled
with this, there are, in the tropics, lower birth rates, greater infant mortality (at least
in birds), more uniform relative abundances (i.e., less ‘dominance’), more spotty
geographic distributions and, probably, greater annual productivity. Any hypothesis
which can simultaneously relate all of these features of tropical biogeography must
be treated with respect.
At least four hypotheses have been seriouslyput forth to account for the great species
diversity in the tropics. (See Pianka, 1966, for an interesting but different review.)
(1) The tropics have offered more opportunities for speciation in the past, and these
species have been slow in adapting to climates with a winter (Fischer, 1960). This can
be completed in two ways :
20 H. MACARTHun
ROBERT
(a) Time is still short and the numbers of species are still increasing.
(b) There is a balance between new species entering and old ones vanishing.
(2) The tropics have more predation, parasitism and disease, putting a lower ceiling
on the abundance of any given species and thus allowing more species to fit in. A sort
of obverse of this theory says that moderate predation intensity not only allows more
species but is necessary for their persistence, possibly because some species can be
resource limited and other new ones predator limited. Paine (1966) has impressive
empirical evidence for intertidal communities.
(3) In the more uniform tropical climates, competitors can be packed closer together
(MacArthur & Levins, 1967) and, forthermore, the competition reduces the intrinsic
rate of natural increase I* and colonizing species will have a decreased chance of suc-
ceeding (MacArthur & Wilson, 1967).
(4) In the more productive tropical environments, what were marginally scarce
resources become adequate and new species will occupy these new ‘niches’. This is
presumably what Odum, Cantlon & Kornicker (1960) had in mind, and Cornell&
Orias (1964) have added that there may be a feedback with more species causing more
stability which in turn allows yet more species.
Now elementary events often have single clearcut explanations, but statistical events
like species diversity often have complex and multiple explanations. Hence there is no
reason that the validity of any of the explanations need preclude any of the others.
In fact,I believe all of the proposals are partly right, if incomplete. The first explanation,
which is essentially historical, allowing a future increase in the numbers of species,
seems inconsistent with the others which predict a saturation with species, but even
these can be mixed. Not only could some taxa have historical explanations while others
have ecological, but also, the total fauna of a country could continue to grow as in
hypothesis (lo), while the local biota of any subdivision might have reached saturation.
In what follows, I examine these ideas more carefully.

T H E TROPICAL ENVIRONMENT

At first I planned a careful statistical analysis of the stability of tropical climates but
this is more ambiguous than I had imagined and I now prefer a sort of bioassay of it.
The trees in temperate regions show marked annual growth rings because growth
nearly ceases in the winter, but trees at least in wet tropical forestsshow less conspicuous
growth rings, doubtless because the trees regard the climate as much less seasonal.
Tropical birds of one species or another nest nearly the whole year round and individual
species have longer nesting seasons indicating that their insect food regards the climate
as much less seasonal, and no bird breeding in Colombia (de Schauensee, 1964) or
Surinam (Haverschmidt, 1968), for instance, leaves to avoid an off season, so the birds
seem to regard the climate as less seasonal.
For these reasons, and in spite of the importance and variability of tropical dry
seasons, I believe the tropical environment must be regarded as more uniform than the
temperate one.
1 dN
y is the intrinsic rate of natural increase defined by - -where N is the population size.
N dt
 IN THE TROPICS
COMMUNITIES 21
That the lowland tropics are more productive is harder to demonstrate but equally
important. Rosenzweig (1968) has shown that actual evapotranspiration (the total
evaporation plus transpiration which equals the precipitation minus the percolation
and run-off) is closely correlated with net production, and evapotranspiration is surely
greater in the tropical lowlands (Holdridge, 1967). Winter, which for many organisms
is critical, is surely less productive in the temperate zone. The upper Amazon, where
Patrick (1966) found no increase over temperate rivers in diatoms or invertebrates,
was not more productive and has great seasonal variability in depth so the greater
productivity is not perfectly correlated with low latitude.
Other organisms of course form an important part of the tropical environment which
we can therefore conclude is more diverse (at least in numbers of species). The tropical
environment is also more diverse in other biological ways, For instance bird food not
only comes in a large range of sizes (Schoener & Janzen, 1968) but also in more
perennially available forms (e.g. fruit at all seasons).
It is often claimed that the tropical environments are actually structurally more
complex. This is surely true from the viewpoint of some animals, and epiphytes in
particular offer many new opportunities, but it is not so clear that the environment is
more structured for trees, or for the animals in savanna country.

COMMUNITY PROPERTIES IN THE TROPICS

As mentioned in the introduction, several correlated changes occur in the tropics.

The first is increase in numbers of species. Fischer’s article (1960) documents many
cases of this and I add another as shown in Fig. 1. Only freshwater invertebrates and
diatoms (Patrick, 1966) seem to fail to increase in diversity in the tropics. (See Table 1.)

Table 1. Comparison of species numbers of major groups of organisms found in hard water
with numbers found in temperate zone rivers and streams in the U.S.A. (After Patrick, 1966)

Quebrada de Ottawa River

Rio Tulumayo Puente Perez 1955-1956 Potomac River

Algae 73* 62 69 103

Protozoa 33 40 47 68
Lower invertebratest 5 6 8-1 5 27
Insects 104 78 51-64 104
Fish 26 22 17-28 28

These numbers represent established numbers of taxa and do not include taxa represented by less
than six specimens.
t Excluding Rotifers.

These patterns describe the numbers of species in large areas of many amalgamated
habitats. In many ways the local pattern is more interesting, if harder to study. Consider
these two hypothetical alternatives: in the first, tropical country A has ten times the
number of species of temperate country B (of equal area), and each component 10
acres of A also has ten times as many species. In this case, the ‘explanation’ of the
22 ROBERTH. MACARTHUR
tropical diversity is local-if we can explain the increase in 10 acres, there is nothing
further to account for. In the second alternative, country A still has ten times the number
of species B has but now the component 10 acres of A and B have equal numbers of
species. Here the explanation is global, not local-the tropical increase has nothing to
do with events within the 10-acre plots and instead has to do with the magnitude of

I I I I I I I I I I I
FIGURE1. The numbers of breeding land bird species in different parts of North America,
from various sources. (After MacArthur & Wilson, 1967.)

faunal differencesbetween habitats. To detect the difference between these alternatives,

plotting logarithms of the numbers of species is useful because than a constant vertical
distance between curves indicates the top one is a constant multiple of the bottom one,
In Fig. 2, I attempt to construct careful species-area curves for breeding land birds of
various tropical and temperate new world areas (‘land’ birds are those which appear in
the lists from ‘pigeons’onwards). In each case I have tried to use nested lists-the birds
of the smaller area are a subset of those of the larger area. Notice first that 5 acres of
woodland in southern Vermont support about the same fraction of the birds in 6 square
miles as do 5 acres on Barro Colorado Island in Panama of the total 6 square miles.
 COMMUNITIES IN THE TROPICS 23
Thus no new causes of tropical diversity appear in that area increase. Rather, even small
Panama censuses have the same multiple (about 2.5) of Vermont ones as do 6 square
miles. Second, notice that the tropical curve rises faster at the far right, for areas greater
than 6 square miles. This surely means that tropical bird species have subdivided the
topography of the country into more parts, each species occupying a smaller range.
Notice that because the graph is logarithmic, this accounts for most of the extra tropical
diversity. Thus, Ecuador has seven times as many species as New England while Barro
Colorado has only two and a half times as many as a similar area of southern Vermont.

I I I I I I 1 1 I
6

I :0.

.-
v1)
0

Q
v)

I I I I I I I I I
10-2 1 102 lo4 I6
Sq. miles

FIGURE 2. The number of breeding land bird species plotted against the area in square miles
for tropical and temperate areas. 1, Five-acre census on Barro Colorado Is. (MacArthur,
Recher & Cody, 1966). 2, Barro Colorado Island (six sq. miles) (Eisenmann, 1952). 3, Panama
Canal Zone (Eisenmann& Loftin, 1967). 4, Republic of Panama (Eisenmann, 1955). 5, Ecuador
(de Schauensee, 1966). 6, Columbia plus Ecuador plus Peru (de Schauensee, 1966). 7, Five-
acre Vermont census (MacArthur& MacArthur, 1961). 8, Six sq. miles in southern Vermont
(unpublished data). 9, Southern Vermont. 10, New England. 11, Northeastern U.S. and
adjacent Canada. 12, Eastern U.S. and Canada. 13, Texas. 14, Western U.S.15, North America
(North of Mexico).
The points for western United States indicate the effect of greater topographic diversity.

This topographic subdivision is fairly clearcut, but the local increase is more puzzling :
how do even 5 acres of Panama forest support two and a half times more species than 5
acres of Vermont forest ? One more bit of evidence is relevant here :this local increase
is largely in number of genera. Thus Barro Colorado has 175 species and 121 genera
while the 6 square miles in Vermont have 80 species and 65 genera. The ratio of species
is 175 :80, i.e. 2.18 while the ratio of genera is 121:65, i.e. 1.86 which is nearly as great.
Although genera are slightly arbitrary, I think this clearly means there is an increase
in bird structural diversity in the tropics. Orians (in press) has some very interesting
tables showing just which new ways of harvesting foods actually appear in the tropics.
I shall discuss this later. The density of species is of course not uniform in the tropics.
24 H. MACARTHUR
ROBERT
Highlands and dry areas generally have fewer species than low wet areas. Holdridge
(1967) in his figure 3 shows how tree diversity (designated by number of species, tree
height, basal area and stem numbers) varies, generally decreasing with lowered tem-
peratures (i.e. altitude in the tropics) and reduced rainfall. Although the data are less
complete, numbers of bird species show the same general features. Seasonality is much
greater in dry areas and slightly greater in high ones (Atlas Estadistico de Costa Rica,
1953) and the productivity as indicated by annual evapotranspiration is greater in low
than high and in wet than dry areas. I t is an informative exercise to explain why montane
areas in the tropics should have temperate species diversities (although tropical clutch
sizes).
One more feature of tropical species diversity needs to be pointed out here. It is not
simply related to the species diversity of food. This is obvious for plants, whose food
does not come in species. For insects, which eat the trees, there may be a correlation,
but probably the number of insect species is more closely correlated with the number of
plant genera than the number of plant species. Arthur Shapiro (pers. comm.) has
found a correlation between plant genera and butterfly species, but I am not aware
of an analysis for other insects. Particular species are sometimes associated with par-
ticular plant species, but the number of bird species is not usually associated with
the number of plant species. In fact, forests of a single tree species will have as many
bird species as others if they have a many-layer aspect due to trees of varied ages present.
(MacArthur & MacArthur (1961) for temperate U S . ;Orians (pers. comm.) for tropics.)
Tropical naturalists have long been aware that nesting mortality rates were high in
the tropics. The only case I know of a single species whose nesting mortality rates in
tropics and temperate zone are known is the red-winged blackbird (Agelaiusphoenicmus)
which nests from Costa Rica to Canada. Dr Gordon Orians has kindly supplied me
with data on its mortality in Costa Rica (his own data) and the State of Washington
much nearer its northern terminus (data of Celia R. Haigh). These appear in the
following table set up for 2' x 2 X 2 test:

Costa Rica Washington

Nest fledging at least one young 20 22s

Nests fledging no young 73 274

Total nests 93 499

This is highly significant, so nest mortality rates are indeed higher in the tropics for
this and probably most other species. It is well known that clutch sizes are lower in the
tropics, but less well known that they are not only low in lowlands but also in highlands
(Skutch, 1967). Thus, although tropical mountain species diversity is like the temperate,
the clutch sizes are tropical. The highland regions have lower production but probably
not much greater seasonality. They also have a more disturbed history, for warm
interglacial periods probably eliminated some temperate species from the tops of the
mountains.
 IN THE TROPICS
COMMUNITIES 25
The relative abundance pattern is particularly interesting because it seems to be well
correlated with the number of species present. That is, wherever there are few species-
whether in the temperate or on a remote tropical island or, perhaps, on tropical moun-
tains-there are greater extremes of commonness and rarity. Grant (1966) has observed
this for island birds, Patrick (see Fig. 3) for diatoms, E. MacArthur for trees, and Joel

14k A -I
I3 - -
A = Hunting creek (actual)
Hunting creek (fitted) -
-
VI
.-
-
0
a -
L
0 8-
o = Layou stream (actual)
-
z
0
Layou stream (fitted) -
6- -
-
4- 0

3-
0
-
2-

1-
I I I I I I , I A I , r l I
0 1 2 4 8 16 32 64 128 256512 1,024 4,096 16,384
Abundance

FIGURE 3. Truncated lognormal curves for diatom communities calculated by Patrick according
to the method of Preston. Hunting Creek is in Maryland, U.S.A., while Layou Stream is on
the island of Dominica in the Lesser Antilles. Notice that on Dominica, where fewer species
are present, the standard deviation of the fitted curve is much greater. (After MacArthur &
Wilson, 1967.)

Sohn (pers. comm.) for coral reef fish. That these results are not artifacts of sampling
is most easily seen by presenting data as in Fig. 3. and estimating the value of the
variance of the fitted truncated normal curve (Patrick, Hohn & Wallace, 1954, give the
maximum likelihood equations and a graphical solution). The fitted u2 is a function of
the topographic diversity of the area sampled and hence increases with area, but for
fixed areas should be quite independent of adequacy of sample. On fitting truncated
lognormal distributions to bird censuses from tropical and temperate and island
homogeneous forest areas, I have obtained mean u2 values of 1.861 zt0.233 for the
temperate, 0.972 zt0-157 for the low tropics and 3.871 h1.437 for islands. This is
merely a documentation of what plant geographers have long known : there is usually
less ‘dominance’ in the tropics.
Finally, the spotty geographic distribution of tropical species is harder to prove,
since it can be claimed that further search will reveal the species in what are now
26 ROBERT
H. MACARTHUR
regarded as gaps in their distributions. I believe this is not the major explanation and
that tropical species do often have disjunct distributions. These are usually called relict
distributions in temperate regions, but I believe their prevalence in the tropics reflects
a very different cause.

AN HYPOTHESIS TO ACCOUNT FOR THESE PROPERTIES

The simplest-in fact the only-hypothesis I know which can account for all of these
properties of tropical communities is that species interactions are important ; both
competition and predation influence species existence and abundance. The alternative,
with which I cannot account for these data, is that species live their lives independently
of one another. But if interactions are important the explanation proceeds easily and
all of the hypotheses mentioned in the introduction can be amalgamated.
According to this hypothesis the tropical environment is not like a box that will hold
only so many eggs ;rather it is more like a balloon which resists further invasion pro-
portionally to its present contents but which can alwayshold a little bit more if necessary.
Furthermore its contents always try to escape to regions of lower pressure. Again,
according to this hypothesis, if there is a global equilibrium in species diversity, it is
achieved by a balance between speciation and immigration on the one hand and extinc-
tion and emigration on the other. If the tropical i d o w of species were greater, or its
outflow were less, then it would equilibrate with more species (Fig. 4).
Next notice that, in theory at least, there is a duality between predator and prey. In
the classical predator-prey equations the symbols for predator and prey can be inter-
changed with only the values of the constants being altered. This leads to the conclusion
that if ‘Two prey-limited predators cannot coexist in a fine-grained environment
containing one prey’ is a valid statement, then so is ‘Two predator-limited prey cannot
coexist in a fine-grained environment containing one predator.’ Among two predators,
one must be more efficient on a single resource, and among two prey one must be more
efficient at avoiding a single predator. Superficially, this seems to mean there must
be equal numbers of predator and prey species, but this is a naive oversimplification.
Rather there must be equal numbers of predator species and the resources on which
they depend. A resource can be just part of a species (fallen leaves or fruit, for instance)
or it can be several species which are harvested interchangeably (the plankton caught
by a whale, for instance). Hence there should be only a rough correlation between
numbers of species in different trophic levels. But they do grow ‘hand in hand’. In
what follows, I picture resource-limited predators, but everything should apply to
predator limited prey and especially to species simultaneouslylimited by both resources
and predators.
Now consider, as species B, C and D in Fig. 5 , several species living their lives quite
independently. Their abundances will be governed largely by the carrying capacity of
the environment. I find it easiest to picture birds in which the abundances may be
determined by amounts of resources. One species may require insect food of a particular
(a) size, (b) height in the trees and stability of perch, (c) degree of mobility, and (d)lying
in a habitat of particular layer structure. These four (and there are doubtless some more)
requirements are independent; that is, all must be satisfied simultaneously. This
 IN THE TROPICS
COMMUNITIES 27
means the proportion of resource which is suitable is roughly the product of the propor-
tions independently satisfying the four requirements. Thus the abundances of the
species will be determined as the distribution of a product of independent random
variables. Now the logarithm of a product of independent random variables is the sum
of the logarithms of the random variables, which is a sum of different independent
random variables. Such sums become normally distributed, according to the central

I
s I

FIGURE 4. R is the rate of extinction plus emigration (curve E ) and immigration plus speciation
(curve I). On a global level, lower extinction and higher speciation rates will cause a higher
equilibrium as shown for the solid lines supposedly representing the tropics. The dotted curves
represent the temperate and the arrows show how the numbers of species (S)grow in each area.
If history is important the numbers will not be at equilibrium but only part way along the
arrows.
In the local graph the rate, I, of immigration falls as the number of species increases because
competition can act at this level. U and P are the extinction rates in unproductive and pro-
ductive environments, respectively.

limit theorem, and hence the logarithm of the product is similarly distributed. (For
a more exact distribution, see Feller, 1966, chapter I, section 8.) This means the abund-
ances are lognormally distributed, which is the pattern Preston (1948) found to fit
the data and which we have used here (Fig. 3). As speciation adds more species, as in
Fig. 5, many have difficulty in establishing a colony, and they become crowded in.
The species marked with arrows in the figure, and which are too tightly crowded in,
actually become eliminated by competition (MacArthur & Levins, 1967) except in
habitats which happen to have an excess of those species’ favoured resources. These
28 H . MACARTHUR
ROBERT
species will tend to have very spotty distributions. Eventually, as enough species are
tried in the area, it will become fairly uniformly packed with species and two features
then become apparent. First, the relative abundances become much more uniform
as the species overlap to this extent and share the common and rare resources. (The
overlapping species act as a sort of smoothing device, smoothing the irregularities
in abundance ;when packed even tighter, relative abundances again become uneven,
this time due to competition.) Second, with further speciation, additional species find
it difficult to enter our area, although there may be some substitutions. The new species

FIGURE 5. The utilization efficiency of various species (small bell-shaped curves) is plotted
against a spectrum of resources, R, in a hypothetical community. The high curves indicated
densities of the resources in productive and unproductive environments and T is the threshold
density necessary for species to maintain their populations. See text for further details.

more easily persist by further subdivision of the geography of the country into small
ranges. These two properties are of course exactly what were documented in the last
section, and competition would tend to confine species to smaller horizontal layers in
the forest as has been documented by MacArthur, Recher & Cody (1966). Within each
area there are marginal niches which cannot at present support a species, but which
could with an increase in resource production. These are species A, D and E in the
figure. In the more productive tropics these species would be present, usually repre-
senting new genera, and would constitute part of the increase in species diversity
which was found even within a habitat. And there will be more cases of large, medium
and small relatives subdividing the food by size. Offshore islands, with fewer species
of competitors and predators will, according to our hypothesis, be released to expand
their niches. This surely happens (Wetmore, 1957 ;Grant, 1966; MacArthur & Wilson,
1967). Recher (pers. comm.) has actually shown that Australian habitats of the same
layer structure as North American ones contain the same diversity of breeding birds,
giving further evidence that there is a global equilibrium of species, in birds at least.
The reduced seasonality in the tropics would have two effects. Like a chain which is
 I N THE TROPICS
COMMUNITIES 29
as strong as its weakest link, the predators can plausibly be commoner where no season
is difficult. Also selection will tend to favour small clutch sizes since only these parents
can feed their young successfully, whereas in varying climates there is a great spring
bloom of food, which allows the parents with large clutches to outproduce those with
small and gives these species a greater opportunity to recolonize each spring. And the
increased predation reduces the optimal clutch size by causing disproportionately heavy
mortality in large clutches. The increased predation influences the species diversity
by reducing the food competition. Since the predation rate is empirically (and theoreti-
cally) greater in the tropics with their low seasonality, and since predators tend to
focus their attention on common prey, the competition is eased and more species can
coexist. Thus, plausibly, a greater historical accumulation of species in an area where
production is greater and seasonality is less should, coupled with the effects of com-
petition and predation, produce all of the characteristics of tropical communities.
The arguments can be made mathematical, which serves to prove that all of the neces-
sary ingredients are included. (See MacArthur, 1968 ; MacArthur & Wilson, 1967;
MacArthur & Levins, 1967; Levins, 1968 for these mathematics.)

ACKNOWLEDGEMENTS
I have discussed most of these ideas profitably with Richard Levins, Henry Horn
and Steven Fretwell, and with Gordon Orians who has kindly supplied me with some
data from his forthcoming paper on bird species diversity in Costa Rica. I have also
been privileged to see Terborgh and Weske's important account of habitat selection and
species diversity in Peruvian birds.

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