Geographic and Climatic
Relationships of Avifaunas with
Special Reference to Comparative
Distribution in the Neotropics
PAUL SLUD
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 212
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 2 1 2
Geographic and Climatic
Relationships of Avifaunas with
Special Reference to Comparative
Distribution in the Neotropics
Paul Slud
SMITHSONIAN INSTITUTION PRESS
City of Washington
1976
ABSTRACT
Slud, Paul. Geographic and Climatic Relationships of Avifaunas with Special
Reference to Comparative Distribution in the Neotropics. Smithsonian Contri-
butions to Zoology, number 212, 149 pages, 37 figures, 11 tables, 1976.?A simple
species list of birds contains within itself environmental information which is
analogous to that obtained from a weather station. In both instances the numeri-
cal data do not become meaningful until used comparatively.
The Class Aves is divisible into passerines, comprising two-thirds of the world's
land birds, and nonpasserines, comprising the remaining third. The passerines in
turn are divisible into oscines and suboscines. The suboscines are numerous only
in the Neotropical region, where they outnumber the oscines. Utilization of these
major taxonomic components as though they were ecological assemblages is sub-
stantiated on objective grounds for the mainland but is not justifiable for islands.
In the neotropics, the relative proportions of the major taxonomic components
correlate with the major dimatovegetational parameters of rainfall and tempera-
ture to provide a comparative measure of faunal and environmental similarity or
dissimilarity and of biotic complexity. The suboscine proportion is highest in the
rain-forested Amazonian lowlands, thence decreases radially and altitudinally.
The oscine proportion fluctuates to varying degree in the opposite direction from
that of the suboscine proportion. The nonpasserine proportion decreases with
increasing elevation to tree line and with increasing rainfall altitudinally and
latitudinally.
Crossplotted, the passerine-nonpasserine and suboscine-oscine relationships pro-
duce a two-dimensional "map" on which to visualize the comparative distribution
of neotropical avifaunas. Avifaunas having similar proportions of passerines (or
nonpasserines) and of suboscines (or oscines) form into discrete climatic clusters.
Any faunal level can be accommodated from the zoogeographic region down to
the habitats at a study site, provided the comparisons are restricted to units of
similar size.
The smaller the size of the unit, the more precise is the avifaunal or environ-
mental information it furnishes. Whether for purposes of comparison or predic-
tion, equivalent samples from equivalent points at equivalent times of year are
not only necessary but also feasible to obtain, and a simple methodology by
which to accomplish this is presented. The proposition that site-oriented com-
parisons in which the loci each approach a bioclimatic standard are fundamental
to the choice, study, and preservation of representative natural areas is supported
on theoretical grounds and by examples.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6050. SERIES COVER DESICN:
The coral Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Slud, Paul.
Geographic and climatic relationships of avifaunas with special reference to comparative distri-
bution in the neotropics.
(Smithsonian contributions to zoology, no. 212)
Supt. of Docs, no.: SI 157:212
1. Birds?Latin America?Geographical distribution. 2. Birds?Geographical distribution. S.
Bioclimatology. I. Title. II. Series: Smithsonian Institution.
Smithsonian contributions to zoology, no. 212.
QL1.S54 no. 212 [QL689A1] 591'.08s [5985^8] 75-619130
Contents
Page
Introduction 1
Species Lists 2
Categories 2
Need to Compare Equal Areas 2
Elimination of Water Birds 2
Total Land Birds 5
Migrants 6
The Passerine-Nonpasserine Relationship 10
Recognition of the Relationship 10
Regional Diversity and Climate . . 11
Mainland Environmental Correlations 14
Islands 21
Area, Species and Ratio 22
Turnover 29
Island Types 33
Summary 50
The Suboscine-Oscine Relationship 50
Neotropical Distribution 51
Sample versus Compilation 56
Summary ? 56
The Passerine-Nonpasserine Suboscine-Oscine Relationship 57
Comparative Categories 57
Single-Figure Values 60
Environmental Clusters 64
Summary 66
Requirements for Further Research 70
Sampling 70
Checklists versus Samples 70
Analytical Methods 70
Faunal Resemblance 79
Accuracy of Counts 80
Choice of Locale 81
Amazonia 81
Costa Rica 82
Appendix 1: Bibliographic Key to Geographic Units in the Tables and
Figures 84
Appendix 2: Tables 90
Literature Cited 135

Geographic and Climatic
Relationships of Avifaunas with
Special Reference to Comparative
Distribution in the Neotropics
Paul Slud
Introduction
There are about 8650 living species of birds in
the world (based on Austin, 1971). This total, dif-
fering little from that of other recent authors or
even from one compiled a generation ago by Mayr
(1946), can be accepted as virtually complete. In-
cluding extinctions in the last few centuries (based
on Austin, 1971) the number rises to 8717. This is
the figure used here, inasmuch as recently defunct
species formed part of existing natural assemblages
and often appear in faunal works.
Faunal works, or compilations, are for the most
part freely available in the printed literature (but
not, in my experience, when unpublished or pri-
vate). The boundaries can be regional, geographic,
political, or ecological, the scope supracontinental
to local. Ranging from a set of volumes on a shelf
to a pocket pamphlet or card, a compilation may
or may not inventory an area completely. The ones
which do are the source of species lists that can be
put to comparative use.
On the basis of simple species lists, this paper
attempts to point out elementary sets of avifaunal
relations and to utilize these relations to compare
avifaunas and to link them to their surroundings.
One set of relations is that between passerines and
Paul Slud, Department of Vertebrate Zoology, National Mu-
seum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.
nonpasserines. This is a global relationship mak-
ing it possible to compare the whole with its parts
and the parts with one another. Eliminating the
water birds serves to place the passerines (all of
them land birds) and the nonpasserines (one-
quarter of them aquatic) on the same terrestrial
footing. A second set of relations is that between
oscines and suboscines. This is primarily a neotropi-
cal relationship, which is here further strengthened
and substantiated. The two sets of relations are con-
sidered separately, then together. Cross-plotted,
they produce a two-dimensional distribution of
neotropical avifaunas. As an imitation of nature,
the picture can be appreciated on its own merits or
in relation to vegetation, climate, and topography.
The paper concludes with a rationale in which
avifaunal sampling and choice of locale are con-
sidered interdependent, provided the objective is
the acquisition of comparative data.
ACKNOWLEDGMENTS.?The American Museum of
Natural History and the Carnegie Museum per-
mitted me to examine the extensive Amazonian
ornithological materials in their care. At the Smith-
sonian Institution, H. Daniel Roth, formerly, and
Lee Ann Hayek, at present, in the Information
Systems Division were instrumental in calculating
correlation coefficients and regressions; Robert G.
Tuck, Jr., formerly in the Department of Verte-
brate Zoology, supplied species counts of reptiles
and amphibians in the eastern Caribbean area;
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Robert H. Gibbs, Jr., Chairman of the Department
of Vertebrate Zoology, made many valuable com-
ments aimed at improving the manuscript.
Species Lists
CATEGORIES
A species list may consist of (1) the entire avi-
fauna or (2) the resident, or native, avifauna, (3)
all the land birds or (4) the resident, or native,
land birds. Categories (1) and (3) contain species
whose status is for the most part self-evident. Cate-
gories (2) and (4) include species whose status is
variable, questionable, or unascertainable not only
in poorly known tropical regions but even in the
well-worked temperate north; consistency of treat-
ment is always a problem. Residents, or natives, are
nonintroduced species that occur year-round, the
others are nonresidents. (A compromise between
native land birds and total land birds is presented
in columns 4 and 7 of Table 1, where a resident
has a value of one, a nonresident has a value of
one-half.) Breeding species offer a poor alternative.
Their status is often presumptive and therefore
the most difficult to determine. Some occur, or
have been recorded, irregularly or in very small
numbers, others lack pertinent data. Nonbreeding
species may inhabit an area for as long a period of
year as the breeding species, yet they are often
ignored.
NEED TO COMPARE EQUAL AREAS
The thoroughgoing compiler generally strives to
assemble as complete a faunal list as he can. Par-
ticularly in the tropics he is apt to emphasize the
"richness" of his area by contrasting it with one
that is larger or less complex or lies outside the
tropics: e.g., Mayr, 1941a (New Guinea), Meyer de
Schauensee, 1948, 1964 (Colombia), Phelps and
Phelps, 1950, 1963 (Venezuela), Friedmann and
Williams, 1971 (Bwamba, Uganda), Wetmore,
1972b (Panama), Mayr and Serventy, 1944, and
Keast, 1972 (Australia). The favored area is thus
represented as possessing proportionately or abso-
lutely as many taxa as, if not more than, the moder-
ately to enormously larger area. Obviously, unequal-
size units having similar-size lists are not acceptable
for unit-area comparison (see below).
As Table 1 demonstrates, species densities increase
as the area being considered decreases. Therefore,
in order to compare avifaunal size (species density)
it is necessary to compare units of similar area. It
is equally necessary that the lists be both accurate
and comprehensive. Suitable data for similar-size
units, whether they differ by belonging to differ-
ent biogeographical regions or are similar if only
in being located somewhere in the global tropics,
occur infrequently in the literature. How infre-
quently is made evident by the melange of entries
in Table 1. Indeed, some of the entries are really
a synthesis of two or more geographically contig-
uous units, specially created to serve as areal mates
for otherwise unpaired units. The units are almost
all political; few ecological reports treat an entire
avifauna or mention the size of the area. Even so,
species densities diverge among entries of compar-
able size when these differ in overall environmental
complexity and converge when they are similar.
ELIMINATION OF WATER BIRDS
Water birds are the Spheniscidae, Gaviidae, Podi-
cipedidae, Procellariiformes, Pelecaniformes, Ci-
coniiformes, Anseriformes, Gruidae, Aramidae,
Rallidae, Heliornithidae, Eurypygidae, and the
Charadriiformes minus Burhinidae, Glareolidae,
and Thinocoridae. Arbitrary determination of in-
dividual species as land birds or water birds (viz.
Rallidae and Alcedinidae) is avoided by not going
below the family level (see Moreau, 1966).
Certain generalizations are made apparent by
Table 1 and Figure 1. The relative size of the wa-
ter bird component, whether on a unit-area basis
or as a percentage of the avifauna, is obtainable
from the difference in species density between
Total Avifauna and Total Land Avifauna in a
global survey of checklists (Table 1). A visualized
sampling of these, based on actual numbers of spe-
cies (Figure 1), shows that the ratio of water birds
to land birds broadly correlates with climate. The
ratio decreases drastically with decreasing latitude
from very high in the boreal region (also on iso-
lated oceanic islands) and well marked in the cool-
temperate region to moderate in the warm-
temperate region, low in the subtropical region,
and extremely low in the tropical region; para-
doxically, the water bird ratio is usually smaller in
the humid tropics than in the dry tropics. Among
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NUMBER 212
mainland units of approximately the same size, in-
clusion or exclusion of water birds has little effect
on the total avifauna relative to the total land avi-
fauna: species densities rise or fall equivalent
amounts, provided tropical units are compared
with tropical units and extratropical units with
extratropical units.
On a subregional to regional scale, water bird per-
centages parallel each other in the Old World and
in the New World, not only in size but also in lati-
tudinal sequence. The Congo and Brazil, on opposite
sides of the Atlantic, have respective percentages
of 14 and 12; southern Africa and southern South
America, 26 and 23. Nearctic North America
(United States and Canada) and the Palearctic
Soviet Union each have a percentage of 37.
A survey of water bird percentages in North Amer-
ica shows that they fall into a distributive pattern
(Figure 2). In the temperate part of the continent,
the lowest percentages are found in arid south-
western United States and adjacent Mexico. The
percentages increase northward and eastward into
the Great Basin, the Rocky Mountains, the Great
Plains, and the area between the Mississippi-
Missouri rivers and the Appalachian Mountains.
The eastern seaboard and to some extent the Pa-
cific sector are distinct from the interior. Much the
highest values occur peripherally at boreal lati-
tudes. The neotropical units, southward from Vera-
cruz and Oaxaca, have the lowest values and lie
apart as a cluster of their own.
In view of the fact that aquatic areas are inhabited
by a specialized water bird fauna, it follows that
water birds should, as a matter of normal proce-
dure, be excluded from consideration when com-
paring the land birds: terrestrial environments are
best represented by terra firma faunas.
For example, Hall (in Voous, 1972:585) com-
pared the breeding land bird proportion of the
avifauna in Africa with that in the Palearctic re-
gion without eliminating the water birds. Hall con-
cluded that Africa is richer in "younger" elements,
that is, the passerines and the nonpredatory arbo-
real nonpasserines (pigeons through woodpeckers),
where they comprise 62 percent and 18 percent, re-
spectively, of the avifauna than is the Palearctic,
where the corresponding percentages are 55 and
9.5. If, however, the water birds are removed, Hall's
own figures show that Africa becomes poorer than
the Palearctic in percent passerines (67 versus 69)
and less rich in percent nonpredatory arboreal non-
passerines (19 versus 11). The combined percentage
of "younger" elements is 80 in Africa and 64.5 in
the Palearctic when water birds are included, 86 in
Africa and 80 in the Palearctic when water birds are
excluded.
The need to remove the water birds becomes
increasingly apparent as the units to be compared
become smaller or as their overall environments in-
creasingly differ. Examples are an oceanic rock
frequented by sea birds and, say, a wood lot support-
ing a community of land birds; a place that teems
with water birds in the wet season and the same
place when the water has evaporated in the dry
season; an inland area that lacks wetlands and a
coastal area possessing estuaries, marshes, and
swamps. The problem also arises when comparing
places at very different altitudes. In contrast to the
lowlands, aquatic birds are rare to absent at mon-
tane elevations, except very locally: Sikkim, in the
Himalaya, has a water bird percentage of only 2.6
(based on Ali, 1962); at San Antonio, in the
Colombian Andes, it was 1.2 (based on Miller,
1963).
TOTAL LAND BIRDS
Eliminating the water birds depresses species
densities markedly, sometimes enormously, in the
temperate zone and on oceanic islands, moderately
or insignificantly on the tropical mainland. When
nontropical mainland units are now compared
with tropical ones, the latter appear overwhelm-
ingly rich. Darlington (1957:265) put it this way:
"The land bird faunas of temperate Eurasia and
North America are little more than depauperate
fringes of adjacent tropical faunas." Even those of
luxuriant tropical islands, such as New Guinea,
Borneo, or Java, rate poorly when set against anal-
ogous areas on the tropical mainland. (See Table 1.)
A total land bird figure can be viewed as a
comparative measure of faunal "richness" or en-
vironmental complexity, whether the unit is a
Colombia or an oceanic islet. Nevertheless, multi-
ple factors complicate the interpretation of a single
figure. The figure is uninformative as to the rela-
tive "richness" or "poorness" of subdivisions with-
in the unit. It does not distinguish between com-
mon or regular, rare or irregular, evenly distributed
or disjunct taxa. Equal-size areas having unequal
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
diversities may have similar totals for different
reasons: an area, owing to its position, may be
visited by more species of migrants than it has resi-
dents; an island normally accessible only to aquatic
species may have a list that includes a number of
mainland strays. Many lists are incomplete, and
better knowledge of the migrants and of the na-
tives could increase the species numbers and den-
sities appreciably. Lack of standards permits one
author to list taxa the occurrence of which another
author does not accept as proved.
MIGRANTS
A world count does not distinguish between total
avifauna and native avifauna, residents and mi-
grants, breeders and nonbreeders. A reproducing
resident in one part of the world may be a transient, a
visitant, or an accidental in other parts. A taxon re-
stricted to a remote bit of land in the ocean has the
same value in a count as one which is cosmopoli-
tan and abundant. The zoogeographic regions,
apart from the Holarctic, have such large land avi-
faunas that the presence or absence of migrants is
hardly noticeable percentagewise. Migratory effects
are nevertheless present and can affect the compar-
ability of geographic units on opposite sides of the
equator.
A global survey of migrant percentages (Figure
3) shows that these follow parallel courses in the
eastern and western hemispheres. Old World and
New World counterparts, insofar as these can be
approximately determined latitudinally, climati-
cally, and areally, have migrant percentages which,
if not quite equivalent, occupy similar relative po-
sitions within the progression for their respective
regions. An inverse geometric correlation, in which
the native land avifauna more than doubles in size
each time the migrant percentage is halved, extends
through most of the progression. At either extrem-
ity, i.e., the high latitude-low species density units
and the low latitude-high species density units,
the percentages tend to curve from the alignment.
In the north, this begins to take place when the
number of species has declined to 60 or so; in the
tropics, when the migrant proportion has declined
to less than 5 percent. If, however, the migrants
were each assigned a value of one-half and added
to the number of residents (as in Table 1), the
extremities would follow the alignment. High-
latitude units, having the largest numbers of mi-
grants, would be shifted the most, units in the sub-
tropics and outer tropics very much less, and units
in the inner tropics hardly at all. The avifauna
would increase an average of 1.8 times in size each
time the migrant percentage is halved.
Figure 3 shows further that islands and satellites
of islands lie varying distances "at sea," so to speak,
generally to the left of the mainland units but
never to the right. The distances correspond roughly
to the size of the islands and to their latitudinal
position or degree of isolation. A north-south se-
quence, or set of sequences, is also discernible. Is-
lands, in part because of their small size compared
to the mainland units, tend to have relatively high
migrant percentages.
LATITUDE.?Western Hemisphere: Temperate
North America and tropical to subtropical Middle
America alternate semiannually in gaining and
losing an enormous avian mass. South America
has a much smaller flux of species and individuals,
this principally in the northwest, especially the Car-
ibbean versant. Large areas, virtually amounting
to subregions, show a progressive reduction south-
ward in percentage and number of migrants.
Texas
Mexico
Central America
Northwestern South America
(Colombia, Ecuador, Peru)
Central South America (Brazil)
Southern South America (Uruguay,
Paraguay, Argentina, Chile)
The states and countries that succeed one
another geographically from northwestern Mexico
through Central America to equatorial South Amer-
ica substantiate and refine the progression. In the
tabulation that follows, the figures for Nicaragua
have been compiled largely from Eisenmann
(1955) and for Ecuador and Peru from Meyer de
Schauensee (1970). Those for all the other entries
come from their own published checklists. It need
be emphasized perhaps that the figures are un-
adorned counts of the species listed in standard
works. No assumptions or guesses have been made
or values assigned or formulas devised to modify
the totals or to give added weight to regularity,
Land bird
migrants
170
125
112
78
40
% Migrants/
total land
species
51.3
16.1
12.2
4.2
2.9
14 1.8
O z
90 80 70 60 SO 40 30 20
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In
flu
en
ce
 
o
f 
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s 
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la
tiv
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to
 
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m
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rs
 
o
f 
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o
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nd
 
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un
as
:
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f 
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s 
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o
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t 
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un
as
.
 
(L
ar
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so
lid
 
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de
s 
w
ith
en
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se
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s 
>?
 
th
e 
u
n
itt
 
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bu
la
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d 
in
 
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e 
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n
 
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ge
 
8;
 
sm
al
l 
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nl
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n
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; 
sm
al
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n
 
dr
de
s 
?
 
isl
an
ds
.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
abundance, frequency, environmental impact,
portance," or whatever.
im-
Baja California
Sonora
Yucatan Peninsula
British Honduras
Chiapas
Guatemala
?1 Salvador
Honduras
Nicaragua
Costa Rica
Panama
Colombia
Venezuela
Ecuador
Peru
Brazil
Area
(1000 mi1)
56
70
63
9
29
42
13
59
57
19
29
440
352
101
482
3286
Land bird
migrants
73
90
93
91
103
114
82
109
84
94
94
74
61
39
43
40
Native
land
species
127
210
227
277
431
433
247
432
428
568
622
1351
1090
1161
1291
1323
Migrants I
total
land
species
36.5
30.0
28.0
24.8
19.3
20.8
24.9
205
16.4
145
13.1
55
5.3
355
35
2.9
The migrant percentages are not directly related to
area. They do show an inverse correlation with size
of avifauna: the regression has a correlation coeffi-
cient of 0.99. Even if the sequence were extended
northward into the border states of Texas, New
Mexico, and Arizona, the coefficient would be 0.94.
A decrease of 10 (where feasible) in percent mi-
grants tends to be accompanied by a twofold in-
crease in size of avifauna.
Notwithstanding the extraordinarily high coeffi-
cient, the sequence is not quite right in two places.
Chiapas probably ought to have a somewhat higher
migrant percentage than Guatemala (with which
it is physiographically linked) instead of one that
is barely lower; Nicaragua should perhaps have
more species than neighboring Honduras. The
progression can be almost idealized if Chiapas is re-
placed by a unit that completely crosses southern
Mexico and if British Honduras, on the Caribbean
versant, and ?1 Salvador, on the Pacific versant, are
merged, respectively, with Guatemala and with
Honduras so as to cross-section the Middle Ameri-
can isthmus, as do the countries to the south. The
sequence of percentages now becomes 21.0 for
southern Mexico, 21.2 for Guatemala-British Hon-
duras, 19.8 for Honduras-El Salvador, 16.4 for Nic-
aragua, 14.2 for Costa Rica, and 13.1 for Panama.
Units represented in the literature by two or
more checklists as a rule have the shorter, or ear-
lier, list with a lower migrant land bird percentage
(and passerine percentage) than the later, or
longer, list (Table 2); successively earlier lists fit
less and less well, if at all, into the geographic
sequence; largest numbers of migrants occur in the
block of mountainous units comprising southern
Mexico and northern Central America. With these
things in mind, we can evaluate the relative status
of Nicaragua, the only Middle American country
without its own checklist, from the numbers in the
above tabulation.
Nicaragua has a resident land avifauna nearly the
same size as in Honduras, Guatemala, and Chiapas
to the north but a migrant percentage closer to
that of Costa Rica and Panama to the south. Also,
Nicaragua has 25 fewer migrants than Honduras
and 10 fewer migrants, instead of more, than Costa
Rica. If the Nicaraguan list were increased only by
the 10 required to match the number in Costa Rica,
the migrant percentage would rise to 18.0; if in-
creased by 25 to the number in Honduras, the mi-
grant percentage would nearly equal that of Hon-
duras. Additional native species that might come to
light in a country as sufficiently well known as
Nicaragua would be too few to appreciably alter
the percentage compared to the number of migrants
yet to be expected. The present figures suggest that
Nicaragua is transitional between northern and
southern Central America. An up-to-date checklist
should place it with northern Central America both
in size of avifauna and in migrant percentage.
Before leaving the above tabulation, one might
add that migrant species of passerines, outnumbering
migrant nonpasserines by as much as four to one
(Table 2), correlate almost as well with total pas-
serines as do total migrants with total land avifauna
and much better than do the relatively few migrant
nonpasserines with total nonpasserines.
Eastern Hemisphere: Moreau (1966: chapt. 8)
compared two groups of African and extra-
Ethiopian units to illustrate "the difference be-
tween the land-bird fauna of a tropical area and a
temperate-zone fauna." In the accompanying tabula-
tion, I have combined Moreau's groups, used his
numbers, and listed his units in order of decreasing
latitude.
NUMBER 212
Thames Valley
Arizona
Morocco
Gambia "Valley"
Migrants/
Native total
Area Land bird land land
(1000 mi2) migrants species species
49 67 42.3
114 78 146 34.8
170 44 132 25.0
42 194 17.8
"British" Somaliland 68 24 214 10.1
The progressive reduction in percent migrants cor-
relates roughly with decreasing latitude and, if
Arizona is omitted, directly with number of migrants
and inversely with size of avifauna.
Figures compiled by me from a larger number and
variety of checklists substantiate the above progres-
sion and follow it into southern Africa, where the
migrant percentage rises again. On the west, the per-
centages range from 60 or more in the north (Fin-
land 59, Great Britain 59, Germany 64, Nether-
lands 67) to 45 in Portugal and 42 in North Africa;
in sub-Saharan Africa, from 12 in Nigeria down to
9 in Liberia, 7 in the Congo, and 5 in French
Cameroon and in Angola, then up to 11 in
Bechuanaland, 11.5 in West Cape Province, and 19
in South-West Africa. On the east, the percentages
range from about 50 in the Balkans (Yugoslavia
48.5, Bulgaria 53, Greece 46.5), 46 in Asia Minor
(Turkey), 49 in the Near East, and 45 in Arabia
to 64 in Egypt and 60 in Libya; in sub-Saharan
Africa, from 23 in Eritrea, 19 in British Somali-
land, 15.5 in the Sudan, and 14 in Ethiopia down
to about 7.5 in Central Africa (Kenya-Uganda 8,
Rwanda 7, Burundi 8), then up to 13 or so farther
south (Nyasaland 12, Zambia 14, Rhodesia 13,
Natal-Zululand 14, East Cape Province 13).
A similar progression extends southward to the
equator in eastern Asia. The migrant percentages
drop from 50 or more in the north (Mongolia 52,
Manchuria 60, Hopei 72, Korea 70, Japan 49) to 19
in southern China (Fukien) and in Thailand, 16 in
the Philippines, 12.5 in Burma, 12 in Malaya and in
South Vietnam, about 11 in the Greater Sundas
(Sumatra 9, Java 12, Borneo 12), 8 in the Lesser
Sundas, and 5 in New Guinea.
AREAL SIZE.?Areal size also affects the size of
the migrant percentage on the mainland, hence a
small area and a large area are not really comparable
with respect to migrants. In the wintering range,
small areas have a smaller number but a higher
proportion of migrants than the larger area of
which they are part. The proportional disparity
increases as the difference in area increases: e.g.,
Guatemala versus Pet?n versus Tikal, Costa Rica
versus Finca La Selva, Panama versus the Canal Zone
versus Barro Colorado Island (Table 2). The dispar-
ity becomes more pronounced when the large area
is elongated latitudinally, so that a taxon which
occurs seasonally in a part of the country remains a
resident of the entire country, as in Mexico or
Argentina. Apparent exceptions are the Yucatan
Peninsula and the Guianas (Table 2). Both are
subdivided into thirds, and the difference in size
between the whole and its parts is evidently not
overwhelmingly great.
NATURE OF COMPILATION.?Number or percent-
age of migrants is coupled to the date or size of
the compilation, whether the unit is a country,
such as Honduras or Costa Rica, or a site, such as
Finca La Selva or Barro Colorado Island (Table
2). Accumulating data are accompanied by (a) the
number and percentage of migrants growing faster
than those of residents, (b) passerine migrants
increasingly outnumbering the nonpasserine up
to several times or more, and (c) the percentage
of nonpasserine migrants rising faster than that of
passerine migrants.
ROLE OF MIGRANTS.?Whether migrants benefit
the economy of a natural environment or detract
from it depends upon whether an author is in-
clined to believe they complement the residents or
compete with them (Slud, 1960:144-145, 1964b).
The species counts and percentages (Table 2) sug-
gest it is the former, the increasing complexity of
the resident biota which has evolved under in-
creasingly amenable climatic conditions, rather
than distance, that is responsible for a decreasing
number of ecological opportunities left open to a
diminishing migrant proportion. Both in Africa
and South America it is the richest biotope, the
equatorial rain belt, that acts as a barrier which
many migrants do not cross, very few enter, and the
remaining ones skirt or pass over in order to winter
in the southern third of the continent.
In the Neotropical region, the passerine per-
centages of the total land avifaunas of the sover-
eign states and territories, thanks to the inclusion
of the migrants, all fit into a narrow range (Figure
4). The influence of the migrants declines most
markedly in South America, where the total land
percentages and native land percentages are nearly
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
/ / 1
MEXICO
BRITISH HONDURAS
GUATEMALA
EL SALVADOR
HONDURAS
NICARAGUA
COSTA RICA
PANAMA
COLOMBIA
ECUADOR
PERU
BOLIVIA
VENEZUELA
GUYANA
SURINAM
FRENCH GUIANA
BRAZIL
PARAGUAY
URUGUAY
ARGENTINA
CHILE
TOTAL LAND BIRDS f
NATIVE LAND BIRDS ?
/ / 1
|
o
o
o
o
o
d
' , I /
o ?
?
?
?
?
?6 ?
d
m
cm
>
0 9
?
>
d
?
cm
?
?
t 51 53 55 57 59 61 63 65 67 69
) | i i ,
40 50 60 70 100
FIGURE 4.?Neotropical land birds: percent passerines per
country.
identical. Northward from Panama, the native
land percentages, owing to the increasing size of
the migrant proportion, veer away from the total
land percentages. Figure 4 can be interpreted as
supporting the view that migrants complement
residents and play a constructive ecological role.
The Passerine-Nonpasserine Relationship
RECOGNITION OF THE RELATIONSHIP
Division of the Class Aves into two taxonomic
entities, the order Passeriformes, or passerines, ver-
sus all the other orders, collectively termed the
nonpasserines, permeates the ornithological litera-
ture. Many faunal works are organized into ap-
proximately equal halves, one of nonpasserines,
the other of passerines. The 15-volume checklist of
birds of the world, for example, was planned to
consist of nonpasserines in the first seven volumes
and passerines in the remaining eight (Mayr and
Greenway, 1962). One reason is undoubtedly prac-
tical convenience. A second motivation could lie
in the fact that authors generally agree on the se-
quence and content of the nonpasserine higher
categories but disagree on family limits and ar-
rangements in the passerines.
Because of the fossil record and certain features
of their anatomy, it is assumed that the passerines
were the last order to appear, that they are in the
midst of a radiation complementing that of the
true flowering plants, whereas many nonpasserines
seem to have passed their peak, and that they have
been evolving mostly as small arboreal perching
birds which, having added seeds and nectar to the
diet, have become dominant on land both in spe-
cies and individuals (Amadon, 1957; Darlington,
1957; Storer, in Marshall, 1960). If reliance be
placed on the estimates by Brodkorb (in Farner
and King, 1971) of species numbers in the past,
the passerine percentage of all birds has risen from
54.8 in the Pleistocene to 59.6 today.
"Passerine" and "nonpasserine" carry the connota-
tion of ecological opposites. To some the terms
imply an aggressive rivalry between adversaries, to
others they perhaps bring to mind a dynamic bal-
ance in which the expanding passerines exert the
greater counterpressure, and possibly to others they
suggest a communal sharing of the environment
under long-term or climax conditions. The follow-
ing examples show how authors have seen fit to
employ the passerine-nonpasserine relationship
toward various ends.
Moreau (1966:94) divided the African avifauna
into five "broadly ecological as well as taxonomic"
groups: one of passerines, three of land nonpas-
serines, and one of water birds. This enabled him
to point out that the nonforest fauna south of the
Sahara contains more species of passerines than
any other group and to conclude that passerines
have on the average much smaller geographic
ranges than nonpasserines. Winterbottom (1959:
40) found that in South Africa "a much higher
percentage (61 as against 29) of the Non-passerines
have a distribution all over the Ethiopian Region
or beyond and therefore the Passerines are much
more useful as indicating the faunal affinities of
our birds." Not dissimilarly, Moreau (1966:112)
compared African and South American families,
genera, and species with the result that the degree
of taxonomic affinity between the two regions was
shown to be much greater in the nonpasserines
than in the passerines.
Moreau (1966:10), noting "the fact that with
. . . different techniques the number of families
NUMBER 212 11
recognized in the non-passerines remains almost
unchanged, while the number of passerine families
varies greatly, shows how much more imperfectly
differentiated the passerines are," suggested "one
possible interpretation . . . is that the passerine
species . . . average much younger than those of
other birds." Enlarging upon this premise, Mo-
reau (1966:96) thought that "the passerines, as
apparently the youngest and most rapidly evolving
group of birds, would be the quickest to speciate
in response to the vicissitudes of African ecology.
Indeed, the proportion of passerine species which
belong to superspecies, nearly two-thirds . . . prob-
ably greatly exceeds the proportion in other
groups."
Lack (1968:11), in the style of Moreau, divided
the Class Aves taxonomically into six ecological
groups of subfamilies: one of passerines (93 sub-
families), two of land nonpasserines (94 sub-
families), and three of aquatic birds (49 subfami-
lies). Assuming the order Passeriformes to be the
last to appear and the one which probably "did not
start its adaptive radiation until the mid-Tertiary,"
Lack found, "fitting with this view, the distribu-
tion of passerine subfamilies is more circumscribed
than that of many other nidicolous land birds,
presumably in part because they have not had so
long in which to spread."
Darlington (1957:254), on the other hand, wrote:
That most species and genera (and families) of passerines are
relatively limited in distribution is apparently not (or not
entirely) owing to any peculiarities of classification or rate
of evolution or geological age. Passerines are probably rela-
tively recent birds, but many have been in existence long
enough to spread over the world if they had the power to do
so rapidly, as have the swallows. Apparently most do not
have the power to do so, perhaps partly because they are
mostly small birds.
Klopfer and MacArthur (1960) tried to equate
passerines and nonpasserines with niche size and
faunal diversity. On the basis of selected breeding-
bird censuses, the authors obtained mean-abun-
dance ratios of passerine individuals to nonpas-
serine individuals, indicating that "at all latitudes
. . . non-passerines are less abundant than passer-
ines." Their assumption that "the phylogenetically
younger passerines . . . have a less limited central
nervous capacity than the non-passerines and thus
. . . [are] more capable of modifying their behav-
ior to fit changing environmental stimuli" they
interpreted "as support for the notion that the
phylogenetically older non-passerine species are
insufficiently plastic in their niche requirements to
colonize temperate areas, tropical niches being
smaller and less subject to change."
REGIONAL DIVERSITY AND CLIMATE
Globally the passerines comprise 66 percent of
all the species of recent land birds (based on Austin,
1971), a ratio of two passerines to every nonpasser-
ine. As to the zoogeographic regions, "modern
comparative statistical details of their bird faunas
are not readily available owing to the absence of
good regional surveys" (Serventy, in Marshall,
1960:106). The statement largely still holds true,
especially in the case of the Oriental region. The
passerine percentages that can be derived from
available regional and subregional faunal works,
despite the great differences in area and faunal
magnitude, closely approximate the global per-
centage (Figure 5). Malaysia and possible New
Guinea appear to be somewhat out of alignment.
Only these two of all the subregional entries are
insular, and both lie in the direction of lower pas-
serine percentage.
The largest continuous tropical areas and the
largest avifaunas occur in the Neotropical and
Ethiopian regions. Yet the former has twice as
many species of land birds as the latter. Moreau
(1966:145) was "inclined to find the basis for the
present difference in the Pleistocene history,"
when the vicissitudes of climate affected wide areas
of Africa in particular and caused the extinction
of many species "so recently that there would have
been only limited opportunities for subsequent
evolution of new ones."
Keast (in Dobzhansky et al., 1972:269) con-
trasted the evolution in isolation enjoyed by most
South American groups with the situation in Af-
rica, where nearly all the avian families are shared
with one or more of the other zoogeographic regions.
Africa, in his view, has continually been receiving
colonists that presumably were already highly effi-
cient and competitive, and the avifauna has been
built up as a stratified series of families occurring
together, each of which "tended to become re-
stricted, or confined, to the narrow zone where it
was optimally efficient." Keast concluded (in Dob-
1 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
5000
2000
1000-
200-
WORLD
NEOTROPICAL REGI
NORTHWESTERN SOUTH AMERICA
ETHIOPIAN
/% CENTRAL AND EASTERN SOUTH AMERICA
> EAST AND NORTHEAST AFRICA
.. .NORTHERN SOUTH AMERICA
/>?T-CENTRAL AND WEST AFRICA
. MIDDLE AMERICA
.INDIAN SUBCONTINENT
PALEARCTIC ^ ^
 D
AUSTRALIAN REGION
5 0 0 F " ^ S O U T H E R N SOUTH AMERICA
/ ^ S O V I E T UNION
^FRENCH 1ND0-CHINA
NEARCTIC R E G I O N *
J MALAYSIA
/ ? N O N T R O P I C A L MEXICO
s 'NEW GUINEA
300f- / 'AUSTRALIA
? * UNITED STATES
/
'EUROPE AND NORTH AFRICA
300 500 1000 2000 5000
TOTAL LAND BIRDS
FIGURE 5.?Regional and subregional survey of the passerine percentages of the numbers of total
land birds. (Dashed line = passerine-nonpasserine ratio of 2:1.)
zhansky et al., 1972:274): "It is apparent that all stable in South America, emphasized the drier cli-
available ecological opportunities, and ways of life, mate in Africa: "The evolution of steppe, savan-
are taken up on each continent." nah and desert faunas has always had sufficient
Fittkau (in Fittkau et al., 1969:631), agreeing areas of appropriate environment at its disposal,
that humid-forest conditions remained relatively whereas over wide areas the rain forest fauna that
NUMBER 212 13
was also formerly present has probably yielded be-
cause of increasing aridity." This was seconded by
Keast (in Dobzhansky et al., 1972:272), who
pointed to the "tremendous range of savannah-
adapted animal groups in Africa where that vege-
tation type is particularly prominent."
On the other hand, "abundant evidence from
many fields of research has recently begun to show
that the Quaternary in both South America and
Africa was a time of drastic climatic change" and
that the inclination "to ascribe the discrepancy [in
species numbers] to a supposed stabler climatic
history in South America . . . we now know to be
improbable" (Vanzolini, in Meggers et al., 1973:
255-257). Sufficient time has elapsed for speciation
to have probably taken place both in Africa and
South America from as recently as the late Pleisto-
cene and post-Pleistocene (Haffer, 1969; Keast, in
Dobzhansky et al., 1972; Moreau, 1966:10-11; Van-
zolini, in Meggers et al., 1973).
Accepting neither climate nor gross ecology as af-
fording any prominent clues, Vanzolini (in Meggers
et al., 1973:255-257) attributed the discrepancy in
species numbers to the increased opportunities for
speciation made possible by a ring of humid-forested
refuges around the Amazonian lowlands during dry
climatic cycles compared to only two diametrically
opposed humid refuges during dry cycles in the
Congo. Evidently duplication and reduplication
of species have occurred primarily under humid-
forest conditions in the neotropics but less often
under drier conditions in Africa. Moreover, biotic
complexity may in itself be capable of generating
further biotic complexity: "Interbiotic relation-
ships apparently had increasingly profound effects
on the evolution of floras in the Cretaceous and
later. With more kinds of organisms involved in
a variety of climatological, topographical, and bio-
logical combinations, conditions became ripe for
the proliferation of evolutionary series culminat-
ing in the diversity manifested in the world today"
(Delevoryas, in Meggers et al., 1973:19).
Nevertheless present-day dimatovegetational fac-
tors can indeed be invoked to explain much of
the present difference in faunal size. Wallace (in
Thomson, 1964:516) long ago distinguished the
Neotropical region "from all the other great Zoo-
logical divisions of the globe by the small propor-
tion of its surface occupied by deserts, by the large
proportion of its lowlands, and by the altogether
unequalled extent and luxuriance of its tropical
forests." Moreau (1966:145) thought the decisive
element in the South American predominance, the
abundance of evergreen forest birds (mainly sub-
oscine), reflected the botanical situation: the
African forest is poor and uniform relative to the
tropical American forest, South America is some
two and a half times richer floristically than Africa.
Africa, compared to South America, is a higher,
drier continent. Much of eastern, central, and
southern Africa lies above 1500 meters; frost is reg-
istered at least 100 nights a year, in some places at
least 150, over much of the South African plateau,
and occasionally damages nursery plants at 3000 feet
in the Zambezi Valley only 18? from the equator;
except for a few isolated spots, the equatorial rain
belt receives only 2000-2500 mm of rain a year,
much less than in parts of Amazonia or the East
Indies; the greater part of Africa on both sides of
the equator is dominated by extremely well-marked
dry periods extending unbrokenly from five to nine
months according to the area, deciduous vegetation
(wooded savanna and steppe of various kinds) cov-
ers most of the region, and the seasonal contrast in
appearance is very great (Moreau, 1966, chapt.
2). While "over most of its vast surface the influ-
ence of man on the vegetation has been great,
both directly and indirectly, and overwhelmingly
towards impoverishment," there have probably
been few large-scale changes in geographical range
as a result of the use or even the misuse of land in
Africa, although that stage is beginning (Moreau,
1966:34, 36).
Habitat destruction may not yet have led to the
extinction of species of birds on the African conti-
nent, but it has made comparisons of natural envi-
ronments or analogous localities within Africa, or
between Africa and other biogeographic regions,
enormously difficult if not impossible. The Upper
Guinea Forest in West Africa, formerly some
150,000 square miles in size, has probably less than
half the area in "actual forest" and much of that in
various stages of secondary growth following agri-
cultural occupation; the Nigerian forest block,
mapped as occupying nearly 50,000 square miles,
appears to have hardly any primary forest left,
while fully grown, tall secondary occurs only in
patches; in the Congo, Chapin, 40 years ago, stated
"the virgin forest has been devastated over wide
areas"; only vestiges of Congo-type forest remain
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in Angola and the East African lake region; vestig-
ial forests along the East African coast "have suf-
fered greatly in the last hundred years of 'develop-
ment,' so that several species of birds discovered in
them in the 1880's have not been found there since"
(Moreau, 1966:102, 159-160). "Man, as a fire-raiser
and especially as a cattle-keeper in modern times,
has tended to push upwards the lower edges of the
montane forests and greatly to change the nature
of the other montane vegetation" (Moreau, 1966:
49).
In contrast to the discrepancy in numbers of spe-
cies, the Neotropical and Ethiopian regions, respec-
tively, have passerine percentages of 64.8 and 66.3
or, more usefully for comparative purposes,
passerine-nonpasserine ratios of 1.84 and 1.96.
Their tropical analogs, Brazil and the Congo, have
corresponding ratios of 1.80 and 1.71. The rain-
forest avifaunas in the Amazon basin (compiled
by me from Meyer de Schauensee, 1970) and in
the Congo basin (compiled by Amadon, in Meggers
et al., 1973) have respective ratios of 1.40 and 1.44.
This parallel example of passerine-nonpasserine
environmental convergence on either side of the
Atlantic lends numerical support to a statement of
belief such as the following: "The real value of
comparative ecological studies covering different
continents" is that "observations and data collected
from one continent may be important and relevant
to another, notwithstanding the fact that the faunas
may be distinct taxonomically and their ecological
similarities due to parallelism and convergence"
(Brenan, in Meggers et al., 1973:4).
In the New World, Figures 2 and 6 trace the pas-
serine proportion of the total land avifauna through
North America from the Arctic Circle to Panama.
If we exclude Greenland, the passerine percentages
(Figure 2) are highest in the temperate zone, that
is, continuously through the southern tier of Cana-
dian provinces and virtually all of coterminous
United States, except the Pacific Northwest
(Washington, British Columbia, and Alaska); they
are lowest south of the Isthmus of Tehuantepec at
one extreme and also, apparently, in boreal to arctic
Canada at the other extreme. Constructed from
actual numbers of passerines and nonpasserines,
Figure 6 illustrates how the passerine-nonpasserine
ratio decreases from between 2:1 to 3:1 in temper-
ate and boreal North America to around 1.5:1 in
tropical Middle America; makes apparent the rela-
tively large size of the avifauna in southwestern
United States, particularly the large states border-
ing Mexico; and sets apart the small but avifau-
nally rich Central American states, especially the
two southernmost, Costa Rica and Panama.
MAINLAND ENVIRONMENTAL CORRELATIONS
AREA AND COMPLEXITY.?Figure 7 illustrates
how continued subdivision of an area increasingly
accentuates differences in the passerine-nonpasserine
ratios of the smaller units which have been averaged
out in the ratios of the larger units. Brazil was
chosen because of its very large size, central posi-
tion, few mountains, paucity of migrants, and fair
to good species compilations for a number of states
and territories. Subdivided into states and terri-
tories, Brazil has a ratio range between 1.3 and 2.0
(Figure 7a). When the states and territories are
consolidated into the country's major geographic
sectors, the ratio range becomes extremely narrow,
between 1.5 and 1.7 (Figure 7b).
Colombia, in contrast to Brazil, is physiographically
and environmentally a highly varied and complex
Andean country. In the accompanying tabulation,
the principal faunal sectors are ecological, hence
have neither marked boundaries nor stipulated
areas; passerine-nonpasserine ratios are followed by
numbers of species in parentheses (compiled from
Meyer de Schauensee, 1964).
Sector
Pacific
Caribbean
Central Mountain
Orinocan
Amazonian
Total land birds
1.27 (419)
1.16 (332)
1.84 (896)
1.34 (544)
1.64 (551)
Native land birds
1.15 (384)
1.03 (305)
1.77 (843)
132 (509)
1.52 (523)
Species allocations and counts are almost certainly
inaccurate in a number of instances, owing to in-
sufficient distributional information, whereas the
ratios are less subject to error.
The "Central Mountain" ratios of 1.84 and 1.77,
respectively, are probably representative of the
sector: Cundinamarca (based on Olivares, 1969), a
department in the heart of the massif, has corre-
sponding ratios of 1.79 and 1.72. The 1.16 and 1.03
of the dry Caribbean sector agree with the 1.14 and
1.04 of the department of Atlantico (based on
Dugand, 1947). The Amazonian sector, with
ratios of 1.64 and 1.52, is paralleled by the 1.52 and
8
 "k ~ \
3
S. 3
8 <?ac ?".<
\
\ \ \ \
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\ % l \ X \ \
\ * * * if. f\ \ x^
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A s l l 3 < f< I n \ X-
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 < < ?g ? J N. >v
digit Ii|i|?jj j y | 2 j | {JLuihssi jfiX*u sii?:i9L ; \ ??
ilNiEilil?^isiiil?2iiBiiiii3isiiiiiislsi9iiiE?l?iiiiiiislalsi?ilig \ *.
S 3 N l d 3 S S V d
FIGURE 6.?Numbers of species of land birds in North and Middle America: passerines versus
nonpasserines. (Dashed diagonals = passerine-nonpasserine ratios; solid circles = same political
units (and islands) as in Figure 2; open circles = political units (and islands) additional to
those in Figure 2; enclosed solid circles = political units contained within larger political units;
enclosed open circles ? nonpolitical units contained within larger units.)
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
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250 -
ft 200 -
= 150 -
100 -
50 -
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J?ESPIR ITO SANTO
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? TOTAL LAND BIRDS
O NATIVE LAND BIRDS
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300 -
250 -
u. 200
o
S 150
IE
100 -
50 -
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y
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? M ^ ? ^ ? ? (AMAZONAS-ACRE- J"''t(MATO GROSSO-GOIAS) RORAIMA) /
SOUTHEAST /
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/ (BAHIA-MINAS GERAIS- -
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NORTH / o ? ^
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^ ^ ^ (PARANA-STA. CATARINA-
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-
? TOTAL LAND BIRDS
O NATIVE LAND BIRDS
50 100 150 200 250 300 350 400 450 500
PASSERINES (NO. OF SPECIES)
FIGURE 7.?Numbers of passerine and nonpasserine land birds in Brazil: a, per state or territory;
b, per sector. (Dashed diagonals = passerine-nonpasserine ratios.)
NUMBER 212 17
1.41 of the commissary of Caqueta (based on
Meyer de Schauensee, 1964) and the 1.54 and 1.51
of Amazonian Brazil (based on compiled sources).
The 1.34 and 1.22 of the Orinocan sector are not
so closely matched by the 1.51 and 1.39 of the in-
tendancy of Meta (based on Meyer de Schauensee,
1964). I have no adequate comparative information
for the Pacific sector.
ELEVATION.?Colombia (Meyer de Schauensee,
1964) and Venezuela (Phelps and Phelps, 1958,
1963), as well as isolated Santa Marta (Todd and
Carriker, 1922), have their avifaunas distributed
by their authors according to the altitudinal zona-
tion of Chapman (1917); passerine-nonpasserine
ratios are followed by numbers of species in paren-
theses. (See below.)
Ratio and elevation rise jointly from the tropical
lowlands to the temperate highlands (excluding
paramo). The midpoint between the two follows the
1500-meter mark, an altitudinal line of division
that is paralleled in the mountains of southeast
Asia, New Guinea, and east-central Africa. In the
Congo basin (based on Schouteden, 1961-1966b),
total land and native land ratios together range
between approximately 1.15 and 1.45 in the low-
lands. The easternmost province, Ituri, rises into
humid-forested highlands, and both its ratios rise
to 1.76.
Chapin (1932-1954, 65:98) saw similarities be-
tween the highland zone in the Congo and in
Colombia, where Chapman's subtropical zone "is
dependent upon abundant precipitation, whereas
decreasing rainfall and increasing cold are respon-
sible for the changes marking the temperate and
Paramo zones" and, one might add, the same fac-
tors could be responsible for the fall in ratios. On
Mt. Ruwenzori, between the Congo and Uganda,
the native land ratio works out to 3.46 above 5000
feet (based on Chapin, 1932-1954, 65:252-256) or to
3.42 above 8000 feet (based on Weekes, 1949). In
Uganda, the Bwamba "lowlands," at an elevation
of 2200-3500 feet, have a ratio of 1.35 (based on
Friedmann and Williams, 1971); the Impenetrable
Forest area has a ratio of 1.69 at 3500-4000 feet, 2.75
at 8000 feet, and 3.79 when restricted to typically
montane species (based on Keith et al., 1969). In
Kenya, the coastal Sokoke Forest has a ratio of 1.17,
the Kakamega Forest area at 5000 feet has a ratio
of 1.85 (based on Ripley and Bond, 1971). The
ratio for strictly montane species at Kakamega is
3.25 (based on Zimmerman, 1972).
Another sort of example comes from three "is-
land" mountains in Portuguese East Africa (Vin-
cent, 1934). These have species numbers of 48, 68,
and 65 and ratios of 3.00, 2.24, and 2.61. Taken to-
gether, the number of species rises to 104 and the
ratio falls to 1.89. Isolated Santa Marta (tabulated
above) bears a similar relation to the Colombian or
Venezuelan highlands taken as a whole. It appears
that a list of the species actually recorded from a
locality reflects the environment more faithfully
than does a combined, compiled, much less an as-
sumed, list from a large area.
ARIDITY.?If the entire North American conti-
nent be taken as an example, the distribution of
passerine-nonpasserine ratios of the native land avi-
faunas shows a gross correlation with general envi-
ronment (Figure 8). The plains states and provinces
of the United States and Canada are conspicuous in
having the lowest ratios, ranging from around unity
(1:1) to well below unity. The highest ratios, as
well as highest species numbers (Figure 6), are con-
centrated in mountainous southwestern United
States and adjoining northwestern Mexico. South
of the Isthmus of Tehuantepec, the ratios are low-
est in the dry Pacific lowlands, in the Yucatan
Peninsula, and in the arid interior of Caribbean
Guatemala.
The few comparative data that are available for
dry areas in South America also have low ratios,
that is, they approach unity and are much lower
than in humid areas. These are the department of
Atlantico in Caribbean Colombia (Dugand, 1947),
with a ratio of 1.04; the interior of northeastern
Venezuela (Friedmann and Smith, 1950, 1955),
with a ratio of 1.06; Santa Elena peninsula in
Tropical (0-1500 m.)
Subtropical (1500-2600 m.)
Temperate (2600-3400 m.)
Paramo (3400 m. to snow line)
Total land birds
Columbia
156 (1149)
1.79 (570)
1.81 (253)
152 (40)
Venezuela
1.58
1.82
1.77
2.00
(868)
(583)
(ISO)
(27)
Native land birds
Colombia Santa Marta
1.48
1.72
1.65
1.31
(1082)
(532)
(230)
(37)
1.34
1.67
3.22
2.33
(311)
(193)
(39)
(10)
Venezuela
1.51 (822)
1.75 (552)
1.64 (124)
1.67 (24)
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
130 120 no 100
FIGURE 8.?Passerine-nonpasserine ratios of resident land avifaunas in North and Middle America
and adjacent South America. (Numbers not underlined = nonpolitical areas.)
NUMBER 212 19
southwestern Ecuador (Marchant, 1958), with a
ratio of 1.05; Mollendo District in southwestern
Peru (Hughes, 1970), with a ratio of 1.04.
A "peninsular" effect, identified by Simpson (1964:
74) with reduced species densities of North Ameri-
can mammals on large peninsulas, would appear
to belong in this section. The resident birds likewise
drop markedly, not only in species density, as do the
mammals, but also in passerine-nonpasserine ratio in
Florida and the Yucatan Peninsula (Figure 8). Flor-
ida and the Yucatan Peninsula mostly consist of
physiographically undramatic lowland and the
vegetations display a monotonously simple physiog-
nomy. This is reason enough to make them predict-
ably poor in variety of habitats and numbers of
species. The very low ratio in Florida could corre-
spond to that in the featureless plains states, in the
Yucatan Peninsula to that in other dry neotropical
lowlands.
Two possibly "peninsular" areas are brought to
conjectural light solely on account of their ratios.
One is the mountainous Honduran "hump," jut-
ting into the Caribbean Sea between the Yucatan
Peninsula and the Nicaraguan lowland "break"
(Figure 8), with a relatively low ratio compared
to neighboring units. The other is low-lying north-
eastern Para, opposite the Amazon delta, with a rela-
tively very low ratio of 1.14 (based on compiled
data). Both, however, have species densities at
least as high as if not higher than those in adjoin-
ing areas.
COSTA RICA.?Altitude and Climate: Smaller,
better-known units are preferable to larger, less
well-known units if the object is to make faunal and
environmental comparisons. Costa Rica may be more
useful in this respect than any other neotropical
country. I have species lists for a number of altitti-
dinally representative "areas" or localities (Table
3), few of which have been delimited or described.
They differ from one another in size, topography,
climate, exposure, diversity, and thoroughness of
inventory. I chose them because they appear to have
at least minimally adequate species totals per
Holdridge life zone (formation). (See Slud, 1964a,
map.) Preference was given when possible to places
reported upon more than once as more likely to
have balanced lists than those known from a single
visit.
The localities in the relatively dry northwest
Pacific lowlands (TD) have the nonpasserine pro-
portion (50 percent or more) of the native land avi-
fauna so much larger than in the other life zones
(formations) that they form a distinct group.
All the rest of the country, comprising some 85 per-
cent of the national territory, is humid, and the
species of passerines outnumber the nonpasserines.
The localities in the tropical, or basal, belt (TM,
TW) have the lowest ratios, averaging 1.50 for na-
tive land birds and 1.75 for total land birds (Tables
3 and 4). A rise to 1.60 or more roughly corresponds
to an actual rise in elevation from the coastal plain
onto foothills or ridges and reflects the inclusion of
some subtropical species.
The subtropical tier of localities (SW) is char-
acterized by a rather abruptly higher set of ratios,
particularly along the Caribbean slope. A corrobo-
rative exception is the Turrialba-Angostura area,
located just inside the subtropical belt on the
Holdridge ecological map of 1959 (see Slud, 1964a),
which has since been found to occupy a Tropical
Moist (TM) transitional enclave in a revised ver-
sion (Tosi, 1969). Compared to the Caribbean side,
a lowering in subtropical ratios at analogous alti-
tudes and topographies on the Pacific slope is prob-
ably related to the difference in climatic regime.
The lower-montane belt (LMW, LMR) succeeds
the subtropical belt altitudinally and complements
it faunally. The two belts have species that are
characteristic of, but not exclusive to, one or the
other as well as species that normally range up-
ward or downward from either. Unfortunately, the
lower-montane entries are few in number and have
poor data. It is upon this unsatisfactory basis that
these appear to have somewhat higher average ratios
than do the subtropical entries.
The lower-montane belt is even more closely allied
to the montane belt (MR), to the extent that a fau-
nal resemblance formula favoring similarity would
find them virtually identical. Together these two
belts comprise a homogeneous unit in that they
completely share a species complex undergoing de-
pletion with increasing altitude: no bird in the
montane belt has not also been reported from the
lower-montane belt. This does not mean that typi-
cally montane species are not more common, even
ubiquitous, at montane-belt elevations, where they
form a distinctive assemblage. The montane-belt
entries (Table 3), of which there are only two, fall
so sharply in value that, in a manner of speaking,
they counterbalance the lowest ratios in the basal
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
belt. The decreasing size of the passerine proportion
would appear to correspond to the decreasing com-
lexity of the montane forest with increasing altitude
and to the inclusion, at Cerro de la Muerte, of tree-
less paramo.
FOREST: The countrywide effect of forest on the
passerine-nonpasserine ratios of the native avifauna
at localities in Costa Rica is not readily apparent.
In general, forest-bird ratios do tend to increase
with increasing rainfall horizontally in the lowlands
and with decreasing temperature vertically into
the highlands. My information is limited to a small
number of unreplicated samples in which I indicated
at the time of my visit the species that in my opin-
ion were primarily forest birds within the context
of the local conditions (Table 5). On the Pacific
slope, from the subhumid north to the humid
south, passerine-nonpasserine ratios are consistently
lower for nonforest birds than for forest birds or
for the entire local avifauna. These Pacific localities
have in common as a possible causative factor a
monsoon-type seasonal climate.
On the Caribbean slope, localities in the tropical
and subtropical belts often have a higher ratio for
nonforest birds than for forest birds or for the entire
local avifauna. The large proportion of these in-
stances on the heavily wooded Caribbean side
might appear to be the opposite of what would be
expected, namely, higher passerine-nonpasserine
ratios in forest than in nonforest. Here, however, the
extremely humid climate, lacking an effective dry
season, is responsible for an aboriginal "rain"-forest
cover that even when disturbed has not until recently
begun to be penerated to appreciable extent by spe-
cies from open-country biotopes. Indeed, tree plan-
tations, regenerating clearings, and shrubby second-
ary and and broken primary growth are richer in
opportunities for arboreal, predominantly passer-
ine birds than is closed lowland forest. The apparent
exceptions are the very places it could be rewarding
to investigate comparatively.
In the Old World, by way of comparison, I have
only two forest versus nonforest examples, both of
them areal compilations rather than local samples,
from Africa. There, as elsewhere, the authors have
their own interpretations of the terms forest and
nonforest. The Impenetrable Forest area in Uganda
(Keith et al., 1969), occuping about 150 square miles,
has a native land passerine-nonpasserine ratio of 2.52
for forest species and 2.22 for forest species plus
nonforest species in forest, 1.73 for nonforest spe-
cies and 1.50 for nonforest species plus forest species
in nonforest. The entire African breeding land avi-
fauna south of the Sahara (Moreau, 1966) has a
lowland forest ratio of 1.75 and a lowland nonforest
ratio of 1.70, a montane forest ratio of 4.00 and a
montane nonforest ratio of 2.48.
Recency of Compilation: A later or more com-
plete compilation tends to have a lower passerine-
nonpasserine ratio than does an earlier or less com-
plete compilation, irrespective of the size of the
unit. Put another way, the larger the number of
contributing samples the smaller will be the pas-
serine proportion, especially among the residents.
This is exemplified by the Central American units
that have more than one published checklist: Gua-
temala, Honduras, Costa Rica, Panama Canal Zone,
and Barro Colorado Island (Table 4). The disparity
is usually greater still between a compilation and a
sample, as exemplified by the Costa Rican localities
in the same table.
In the case of Costa Rica and Honduras, the early
checklists were far from complete, yet were already
sufficiently representative to be in passerine-
nonpasserine equilibrium. The two field stations,
Barro Colorado Island and Finca La Selva, are also
well enough known for their ratios to be in equi-
librium, at least for the resident avifauna, and
neither is likely to change without extensive altera-
tion of the environment. At Barro Colorado, the
recent slight increase in total land bird ratio reflects
a mounting record of scarce passerine migrants,
which the addition of a nonpasserine or two would
reverse.
Among native land birds, the one clear instance
of a locality in which the compilation has a higher
ratio than the sample is the Villa Mills-Cerro de la
Muerte entry, but only in one of six monthly sam-
ples (Table 4). Among total land birds, there are
fewer than a handful of such instances: the General
Valley and Monteverde when migrants were ab-
sent, Taboga during the "mid-winter" lull, and
Villa Mills-Cerro de la Muerte during the same
monthly visit mentioned above.
Limitations of Knowledge: Species distributions
in general works are incomplete or inexact, overly
broad and insufficiently detailed. Belts vary in width
and elevation and are apt to be disjunct. The com-
position of the corresponding avifaunas varies ac-
cordingly, and many species are included on assump-
NUMBER 212 21
tion. A species found once or at one place can enter
into a count as though it were a commonly known,
widespread inhabitant. If recorded from separated
points, its political or ecological occurrence either
horizontally or vertically anywhere between may or
may not be taken for granted. So many decisions if
not guesses must be made by the person trying to
compile distributional categories from a faunal work,
intended merely as a digest or guide by its author,
that a second set of totals taken from the same ma-
terial will seldom turn out the same as the first.
The ratios, manifestly so precise, have been derived
from the species totals, which have been synthe-
sized from sources that in turn often depend upon
other sources.
In Costa Rica, a word should be said about the
two Tropical Wet localities, Rinc6n de Osa on the
Pacific side and Finca La Selva on the Caribbean
side (Table 4). Rinc6n de Osa has a larger area,
more complex topography, more seasonal climate,
taller and "richer" forest, and more habitats, yet its
bird list is considerably shorter, not for want of ob-
servers, than at Finca La Selva. On the other hand,
a survey I made at Rincon de Osa, where I was
unacquainted, resulted in a list that was not much
shorter than ones I had obtained in surveys of com-
parable length at Finca La Selva, which I knew
quite well. It is conceivable from this, the only
evidence at hand, that the number of occupied
niches may be approximately the same at any mo-
ment in both places.
The little information that is available for the
damp and chilly Costa Rican highlands is neither
comprehensive nor precise, and it rapidly decreases
in quantity with increasing elevation. Probably the
lower-montane and especially the montane ratios
are too low. I may have unwittingly excluded quite
a few subtropical passerines from both these belts,
as in Table 8, simply because so little is known of
their distributional limits, and included as residents
certain nonpasserines that are actually visitors. A
number of species, particularly among the nonpas-
serines, are scarce, scattered, secretive, or otherwise
hard to find, while wind, rain, and mist often
severely hinder observation. Species recorded dur-
ing different trips usually differ from one another
more in the highlands than at lower, warmer eleva-
tions. Native birds that rise seasonally to higher
altitudes, even if no more abundant in species than
those moving downward, produce a proportion-
ately greater numerical impact on the highland
counts than do the latter on the lowland counts.
Whether compiled, sampled, or made the objects
of special study, the status of a number of kinds as
temporary or permanent inhabitants of the belt, let
alone the site, is not known. Local horizontal move-
ments by some, either alone or mixed with others
and casually or opportunistically noted, can fur-
ther affect already imbalanced results. Only when
proper attention is paid to representative points will
suitable comparative data be forthcoming.
ISLANDS
Insular avifaunas tend to correlate in size with
the area of the island, but only in a general way.
Equal-area islands that appear to be similar to each
other ecologically and are similarly situated with
respect to the mainland are richer in native land
birds in equable warm climates than in seasonally
inhospitable climates. Tropical islands are largely
populated opportunistically by colonizer species
arriving from areas with climates and, inferentially,
ecologies that are like their own. Land migrants in
the tropics originate almost entirely from outside the
tropics, colonizers from inside the tropics. Invasion
of the mainland by insular taxa is at best local or
coastal and seldom successful.
Islands are the preferred source of area-species
formulations, if only because of their clearly
bounded insularity and the apparent lack of success
with mainland situations. Islands come closest to
constituting discrete, independent ecosystems or
natural laboratories: this makes islands desirable
for the study of geographic variation. Insular bio-
tas, however, conform to no universal standard and
their compositions are each the unique result of in-
terplay among many factors that are differentially
peculiar to islands: this makes islands synecologi-
cally nonintercomparable. Moreau (1966:309), for
example, stated he would not follow uniform lines
when discussing the islands around Africa, "since
the circumstances of each African island or archi-
pelago differ a good deal."
If it be assumed that islands do not differ environ-
mentally from comparable mainland under similar
conditions, the impoverishment of insular avifau-
nas becomes a reflection of unfilled niches owing
to difficulties of overwater colonization. But if it be
assumed, as by Lack (1970), "that the primaeval
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
habitats on islands are 'ecologically full' so far as
birds are concerned, and that the reduced numbers
of bird species are due to ecological impoverish-
ment, as exemplified by the reduced numbers of
plant species (with which the numbers of bird spe-
cies are correlated)," an impoverished avifauna is
the sign of an impoverished biota.
Vegetations in different biogeographical regions
have evolved similar physiognomies in response to
correspondingly similar climatic conditions on the
mainland. Modifying the physiognomy is the num-
ber or variety of species in the association or com-
munity. Islands, on the other hand, can be similar to
one another in size, age, physiography, and climate,
but, owing to the degree to which they are inaccessi-
ble to potential colonizers, can have widely divergent
numbers and constellations of species. Some islands
are relatively more impoverished than others. Com-
pared to the mainland, they are all impoverished
botanically and altered physiognomically. Only the
mainland is in full flower, so to speak, and it is there
that environmental standards must be formulated
and established.
Area, Species, and Ratio
AREA-SPECIES.?Figure 9 is the usual double-log
representation of area-species distributions which
has become current in the literature. Here it is
restricted to single islands, whether alone or a mem-
ber of an island group, in tropical and subtropical
waters around the globe, including the southwest-
ern Pacific Ocean. Not included are the myriad
islets for which information is lacking, also islands
at high northern latitudes where species of native
land birds are few and their resident status is often
unknown or hard to decide. The few entries that,
for islands, have high counts are situated on con-
tinental shelves and, except Trinidad, are mostly
large. Islands with very low counts are mostly
small. Both kinds follow a weakly rising gradient.
Viewed in toto, the entries combine a decelerating
increase in number of species with an accelerating
increase in area.
The same array acquires a different appearance
on semilog paper (Figure 10). Most of the islands
under 1000, and all under 100, square miles have
fewer than 50 species of land birds. The propor-
tion of islands with more than 50 species suddenly
rises near the 1000-square-mile mark; even so, few
islands of less than 1000 square miles have as many
as 100 species and none reaches 150. The accelera-
tion and the abruptness of the increase are both
contrary to what happens on the mainland. (Intra-
island comparisons, however, may duplicate if not
exaggerate the mainland situation: Borneo has 367
land species and a ratio of 1.30; Sabah [North
Borneo], one-tenth as large, has 95 percent of the
species and the same ratio.) More than one-fifth of
the islands from approximately 1000 to over 50,000
square miles in area continue to have fewer than 50
species; against this background, Trinidad (number
122), a tropical continental island, is outstandingly
rich. Upward from about the 7000-square-mile
mark (an area smaller than Israel), the number of
entries decreases drastically with increasing area.
The large, luxuriant, tropical continental islands
(Ceylon, the individual Greater Sundas, and New
Guinea) follow a gradient of increasing species
numbers; the others, barely excepting perhaps Min-
danao and Luzon and possibly Celebes, do not.
Figure 11 is a semilog survey of tropical, sub-
tropical, and southern-hemisphere island groups
containing at least two islands. In overall appear-
ance it differs little from that of the single islands.
The majority of the entries fit between the 100 and
10,000 square-mile marks: those under 500 square
miles have less than 50 species; those between 500
and 10,000 square miles in size include fewer than
a handful with 100 or more species, and only one
of these, just barely, exceeds 150 species. The
10,000 square-mile mark represents a dividing line
between island groups with small avifaunas and
those with large avifaunas. Large-size island groups
are few, and they follow a steep species-number
slope.
AREA-RATIO.?Area-ratio distributions of the
single islands (Figure 10) are for the most part
rather evenly dispersed within a horizontal band
spreading across the survey from the smallest to the
largest island; those of the island groups slant very
slightly upward (Figure 11). Insular passerine-
nonpasserine ratio thus appears to be largely inde-
pendent of island size or number of species. Rather,
it is related to or modified by the type of island, its
age, its position with respect to faunal sources and
interchange with other islands, secondary differen-
tiation of its avifauna, and the local environment.
Islands having generally similar topographic and
biotic conditions are often, though not always,
NUMBER 212 23
AREA IN SQUARE MILES
soo
82 "?V*> " ?
or on ?
PB
.
rf ?*
io**ia
73* rt*
?? S, 'm
i ...I
100 O
50
 S
CO
i
1000
AREA IN SQUARE MILES
FIGURE 9.?Area-species distribution of single islands (includ-
ing New Zealand and Tasmania) in the warmer oceans. (1,
Santa Ana (Solomons); 1A, Laysan; 2, Jervis (Galapagos); 3,
Saba; 4, Lord Howe; 5, Barbareta; 6, Annobon; 7, St. Eusta-
tius; 8, Old Providence; 9, Anegada; 10, St. Andrew; 11, Gizo;
12, Virgin Gorda; 12A, Norfolk; 13, Culebra; 14, Maria
Cleofas; 15, Mono; 15A, Henderson; 16, Cocos (E. Pacific);
17, St. John; 18, Bermuda; 19, Tortola; 20, Utila; 20A, Cayo
Largo; 21, Clarion; 22, Mona; 23, San Jos? (Pearl Is.); 24,
Nissan; 25, Guanaja; 26, St. Thomas; 27, Montserrat; 28,
Anguilla; 29, St. Martin; 30, Rodriguez; 31, Vieques; 31A,
Car Nicobar; 32, Aneiteum; 33, Nevis; 34, Marfa Magdalena;
35, Principe; 36, Tutuila; 36A, Camorta; 37, Marie Galante;
38, Aldabra; 39, Barbuda; 40, Christmas (Indian Ocean); 41,
St. Kitts; 42, Aruba; 43, Charles; 44, Roatan; 45, Socorro;
46, Grand Cayman; 47, Guadalupe; 48, St. Croix; 49, Moheli;
50, Bonaire; 51, Rendova; 52, Maria Madre; 52A, Niue; 53,
Antigua; 54, Hierro; 55, Aurora; 56, Tobago; 57, Grenada;
58, Cozumel; 59, St. Vincent; 60, Ponape; 61, Lanai; 62,
Mayotte; 63, Anjouan; 64, Gomera; 65, Coiba; 65A, Enggano;
66, Kandavu; 67, Taveuni; 68, Barbados; 69, Curasao; 70,
Vella Lavella; 71, Efate; 72, Pentecost; 73, Guam; 74, St.
Lucia; 75, Mafia; 76, Kolombangara; 77, Tanna; 78, Molokai;
79, Palma; 80, Gonave; 81, Dominica; 81A, Sao Tom?; 82,
Pemba; 83, Lanzarote; 84, Sao Tiago; 85, Gran Comoro; 86,
Ambrym; 87, Martinique; 88, Tahiti; 89, Rennell; 89A, Great
Nicobar; 90, Upolu; 91, Margarita; 92, Basilan; 93, New
Georgia; 94, Kauai; 95, Guadeloupe; 96, Gran Canada; 97,
Oahu; 97A, Simalur; 98, Zanzibar; 99, Fuerteventura; 100,
Savaii; 101, Mauritius; 102, Maui; 103, Bunguran; 104, Inde-
fatigable; 105, Tenerife; 106, Fernando Po; 107, Erromanga;
108, Japen; 109, Biak; 110, Reunion; 111, Malekula; 112,
Waigeu; 113, Isle of Pines; 114, Socotra; 115, Bohol; 116,
Choiseul; 117, Santa Isabel (Solomons); 117A, Nias; 118, San
Cristobal (Solomons); 119, Albemarle; 120, Cebu; 121, Lom-
bok; 122, Trinidad; 123, Espfritu Santo; 124, Malaita; 125,
Vanua Levu; 126, Bali; 127, Guadalcanal; 128, Leyte; 129,
Bum; 130, Puerto Rico; 131, Bougainville; 132, Mindoro;
133, Hawaii; 134, Vitu Levu; 135, Sumba; 136, Jamaica;
137, Panay; 138, Palawan; 139, Negros; 140, Samar; 141, Flores;
142, Sumbawa; 143, New Caledonia; 144, Ceram; 145, Hal-
mahera; 146, Hainan; 147, Timor; 148, Taiwan; 149, Isla
Grande (Tierra del Fuego); 150, Ceylon; 151, Tasmania; 152,
Hispaniola; 153, Mindanao; 154, Luzon; 155, Cuba; 156,
North Island (New Zealand); 157, Java; 158, South Island
(New Zealand); 159, Celebes; 160, Sumatra; 161, Madagascar;
162, Borneo; 163, New Guinea.)
members of chains or archipelagos and tend to.
follow the same area-ratio tier (Figure 10): Lesser
Antilles, Greater Antilles, Sundas (the larger
islands on a higher level, the smaller islands on a
lower level), Philippines, Solomons, French Poly-
nesia; far above lie islands of the Hawaiian and
Galapagos groups, all with very high ratios; the
few higher northern-latitude islands which have
?
#  
17
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10
.?
Ar
ea
-sp
ec
ies
 
a
n
d 
ar
ea
-r
at
io
 
se
m
ilo
g 
di
st
ri
bu
tio
n
 
o
f 
sin
gl
e 
isl
an
ds
: 
a
, 
sp
ec
ies
 
n
u
m
be
rs
sc
al
ed
 
in
 
tw
en
tie
s;
 
b,
 
sp
ec
ies
 
n
u
m
be
rs
 
sc
al
ed
 
in
 
fif
tie
s. 
(E
ntr
ies
 
th
e 
sa
m
e 
a
s 
in
 
Fi
gu
re
 
9;
 
o
pe
n
sy
m
bo
ls 
=
 
n
u
m
be
rs
 
o
f 
n
at
iv
e 
la
nd
 
sp
ec
ies
, 
so
lid
 
sy
m
bo
ls 
=
 
pa
ss
er
in
e-
no
np
as
se
rin
e 
ra
tio
s.
Sy
m
bo
ls 
ar
e:
 
st
ar
 
in
 
cir
cle
 
=
 
Su
nd
a
 
Is
la
nd
s;
 
tr
ia
ng
le
 
po
in
tin
g 
do
wn
wa
rd
 
?
 
M
ol
uc
ca
s-C
ele
be
s*
Ph
ili
pp
in
es
; 
tr
ia
ng
le
 
po
in
tin
g 
u
pw
ar
d 
?
 
N
ew
 
G
ui
ne
a
 
ar
ea
; 
st
ar
 
=
 
so
u
th
w
es
t 
Pa
cif
ic;
 
sq
ua
re
 
in
cir
cle
 
=
 
G
re
at
er
 
A
nt
ill
es
 
an
d 
Vi
rg
in
s; 
sq
ua
re
 
=
 
Le
ss
er
 
A
nt
ill
es
 
an
d 
w
es
te
rn
 
Ca
rib
be
an
; 
di
am
on
d
=
 
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di
an
 
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ce
an
; 
sm
al
l c
irc
les
 
=
 
a
ll 
ot
he
rs
.)
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?
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40
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20
?
3.
00
-
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0 
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0
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0.6
0
?
0.4
0
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
RATIO OF PASSERINES TO NONPASSERINES
S $ S 8 S 8 ? S 8 2 S ? 8 8
n r i m n o i M N N o i ? ? ? ? ?
I I I I I I I I I I I I I I
8 g O Q
o d d
I I I
o om o
(VJ (VJ
? O
?rs
?h
2 ? _ - ?
O Q O o O O O O O O O
S3Nld3SSVdNON 0 1 S3NI?3SSVd JO 0I1V8
FIG
UR
E 
11
.?
Ar
ea
-sp
ec
ies
 
an
d 
ar
ea
-r
at
io
 
se
m
ilo
g 
di
str
ib
ut
io
n
 
of
 
isl
an
d 
gr
ou
ps
 
(in
clu
din
g 
N
ew
Ze
al
an
d) 
in
 
th
e 
w
ar
m
er
 
oc
ea
n
s.
 
(O
pe
n
 
cir
cle
s 
=
 
n
u
m
be
rs
 
of
 
n
at
iv
e 
la
nd
 
sp
ec
ies
; 
so
lid
 
cir
cle
s
=
 
pa
ss
er
in
e-
no
np
as
se
rin
e 
ra
tio
s;
 
do
ub
le
 
cir
cle
s 
=
 
a
 
sm
al
le
r 
gr
ou
p 
an
d 
la
rg
er
 
gr
ou
p 
to
 
w
hi
ch
 
it
be
lo
ng
s. 
1,
 
M
an
ua
; 
2,
 
La
u;
 
3,
 
Al
da
br
a;
 
4,
 
Ba
y 
Is
.
 
of
 
H
on
du
ra
s;
 
5,
 
Se
yc
he
lle
s; 
6,
 
Tr
es
 
M
ar
ia
s; 
7,
Pa
la
u;
 
8,
 
V
irg
in
; 
8A
, 
To
ng
a;
 
9,
 
An
am
ba
; 
10
, 
K
an
ge
an
; 
11
, 
M
ad
eir
a;
 
12
, 
R
ev
ill
a
 
G
ig
ed
os
; 
12
A
,
Tu
am
ot
u;
 
13
, 
Ca
ro
lin
e;
 
14
, 
D
ut
ch
 
Le
ew
ar
d;
 
15
, 
Sa
nt
a
 
Cr
uz
 
(so
uth
we
st 
Pa
cif
ic)
; 
16
, 
M
ar
ia
na
;
16
A,
 
M
ar
qu
es
as
 
(So
uth
 
Pa
cif
ic)
; 
16
B,
 
Pe
ar
l; 
17
, 
K
ei;
 
18
, 
Lo
ui
sia
de
; 
19
, 
Ni
co
ba
r;
 
19
A
, 
So
cie
ty
;
20
, 
Co
m
or
o;
 
21
, 
N
at
un
a;
 
22
, 
A
dm
ira
lty
; 
23
, 
Az
or
es
; 
24
, 
Lo
ya
lty
; 
25
, 
W
in
dw
ar
d 
(L
ess
er 
A
nt
ill
es
);
26
, 
Sa
m
oa
; 
27
, 
D
'E
nt
re
ca
ste
au
x;
 
28
, 
M
icr
on
es
ia
; 
29
, 
Le
ew
ar
d 
(L
ess
er 
A
nt
ill
es
); 
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A
, 
Fr
en
ch
Po
ly
ne
sia
; 
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, 
Ca
pe
 
Ve
rd
e;
 
31
, 
Su
la;
 
32
, 
Le
ss
er
 
A
nt
ill
es
; 
33
, 
A
nd
am
an
; 
34
, 
Ca
na
ry
; 
35
, 
G
al
ap
ag
os
;
36
, 
A
ru
; 
37
, 
Ba
ha
m
as
; 
38
, 
Fa
lk
la
nd
; 
38
A,
 
W
es
t 
Su
m
at
ra
; 
39
, 
Pa
la
w
an
; 
40
, 
N
ew
 
H
eb
rid
es
; 
41
,
H
aw
ai
i; 
42
, 
Fi
ji;
 
43
, 
N
ew
 
Ca
led
on
ia
; 
44
, 
So
lo
m
on
; 
44
A
, 
Le
ss
er
 
Su
nd
a
 
(ex
clu
din
g 
Ba
li 
an
d
K
an
ge
an
 
Is
.);
 
44
B,
 
Le
ss
er
 
Su
nd
a
 
(in
clu
din
g 
Ba
li 
an
d 
K
an
ge
an
 
Is
.); 
45
, 
M
ol
uc
ca
s;
 
46
, 
G
re
at
er
A
nt
ill
es
; 
47
, 
W
es
t 
In
di
es
; 
48
, 
N
ew
 
Ze
al
an
d;
 
49
, 
Ph
ili
pp
in
e 
(ex
clu
din
g 
Pa
la
w
an
); 
50
, 
Ph
ili
pp
in
e;
51
, 
Ne
w
 
G
ui
ne
a;
 
52
, 
G
re
at
er
 
Su
nd
a.)
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
been included also tend toward higher ratios, as
do mainland areas. Passerine-nonpasserine ratios
of island groups likewise tend to remain stable
rather than rise with island size or species numbers
(Figure 11).
RATIO.?An attempt was made to formalize the
relation of the passerines to the nonpasserines on
single islands and on island groups by plotting the
logarithms of the numbers of passerines against the
logarithms of the numbers of nonpasserines and
subjecting them to regression analysis.
For single islands, the complete set of entries
that appears in Figure 10 produced these values:
for the regression of Y (passerines) on X (non-
passerines), 0.33 for the Y-intercept, 0.85 for the
regression slope, and 0.05 for the variance of esti-
mate of Y on X; the correlation coefficient was
0.84. Thus, for a given number of nonpasserines,
say, 100, the predicted mean number of passerines
rounds to 75. Seventy percent of the known varia-
tion in the number of passerines is explainable
by the number of nonpasserines.
Deleting the very small entries, i.e., those with
less than a minimum of ten species of passerines or
nonpasserines, and those lying outside the tropics,
the remaining 116 entries produced these values:
for the regression of Y (passerines) on X (non-
passerines), 0.075 for the Y-intercept, 0.94 for the
regression slope, and 0.016 for the variance of
estimate of Y on X; the correlation coefficient was
0.92. For a given number of nonpasserines, say,
100, the number of passerines rounds to 94. Eighty-
five percent of the known variation in the number
of passerines is explainable by the number of
nonpasserines.
For island groups (Figure 11), excluding those
with fewer than ten species of passerines or non-
passerines, the values were these: for the regression
of Y (passerines) on X (nonpasserines), 0.005 for
the Y-intercept, 0.996 for the regression slope, and
0.015 for the variance of estimate of Y on X; the
correlation coefficient was 0.95. In other words, the
regression line, bisecting the graph into almost
exactly equal halves, represents a near-perfect 1:1
ratio of passerines to nonpasserines. For a given
number of nonpasserines, say, 100, the predicted
mean number of passerines rounds to 100. Ninety
percent of the known variation in the number of
passerines is explainable by the number of non-
passerines.
Several of the above entries, however, form part
of larger entries or incorporate smaller ones?for
example, the Lesser Antilles and its two compo-
nents, the Windward Islands and the Leeward
Islands, were entered as separate entities?while
another, New Zealand, lies well outside the tropics.
But even when the list is purged of these elements
(numbers 11, 18, 25, 27, 29, 36, 47, 48 in Figure
11), the 34 island groups that remain yield virtually
the same values as before: -0.02 for the Y-intercept,
1.009 for the regression slope, 0.017 for the variance
of estimate of Y on X, and 0.94 for the correlation
coefficient. If the given number of nonpasserines is
100, the predicted mean number of passerines
rounds to 100. Eighty-nine percent of the known
variation in the number of passerines is explainable
by the number of nonpasserines.
The above results, ultimately derived from
simple species lists, make it appear that the pre-
dictable ratio of native land passerines to non-
passerines is almost exactly 1:1 on island groups in
the warmer oceans and seas around the globe, irre-
spective of such contributory factors as area, dis-
tance, origin, ecology, number and variety of
islands, or size of avifauna. Island groups are pre-
ferred to single islands for regression analysis on
the assumption that an island group, taken as a
whole, is populated by a more or less representative
spectrum of species. Apart from the fact that
groups with many entries would "swamp" groups
with very few, the member islands may be so
diverse or so small areally or avifaunally as to bear
little relation to the whole. The assumption re-
ceives support from the following example, that
of the Pearl Archipelago in the Bay of Panama
(based on MacArthur et al., 1972, 1973; A. Wet-
more, oral communication).
The Pearl Archipelago is arbitrarily divisible
into (a) a northern sector that consists entirely of
a scattering of islets and (b) a southern sector that
consists principally of the three "large" Pearl Is-
lands but also includes (c) islets ringing the
periphery of the largest island (Rey). For the
northern sector (a), such data as are available for
eight of the islets show a species-number range
from 1 to 24 and passerine-nonpasserine ratios from
0 to 3.50. The three "large" main islands (b), in
order of decreasing area, have species numbers of
44, 42, and 35 and ratios of 1.45, 1.10, and 0.95.
For (c) the islets around Rey, such data as are
NUMBER 212 29
available for six of them show a species-number
range from 4 to 25 and passerine-nonpasserine
ratios from 0 to 3.67.
When the islets of the northern sector (a) are
considered as a group, they have a total of 34
species of resident land birds and a passerine-
nonpasserine ratio of 1.43. In the southern sector
(b), the largest island (Rey) has 44 species and a
ratio of 1.45 (or, if as is probable it also has at least
one of the two vultures found on the other smaller
islands, a ratio of 1.37). The peripheral islets (c),
when taken together, have 34 species, or three-
quarters as many as on the source island (Rey)
and exactly the same number as on the islets in
(a); their ratio of 1.43 is virtually identical to that
of (a) and (b). The other two main islands, San
Jose1 and Pedro Gonzalez (ca. 20 and 8 square
miles, respectively), are a fraction the size of Rey
(ca. 100 square miles). As their progressively re-
duced ratios suggest, San Jose and Pedro Gonzalez
act as satellites of Rey, though they include a few
species not recorded from the latter. Combined,
the three islands have 52 species and a ratio of 1.00
compared to 54 species and a ratio of 1.00 for the
Pearl Archipelago as a whole. Thus the three main
islands as a group are equivalent avifaunally to the
entire archipelago, but each is apparently too
small to be reliably considered so by itself.
Turnover
TURNOVER RATES.?"Modern biogeographic the-
ory states that the number of species in a class or
other large taxonomic unit that inhabits an island
is maintained within narrow bounds by a dynamic
steady state between colonization and extinction
(MacArthur and Wilson 1967)" (Terborgh and
Faaborg, 1973). Terborgh and Faaborg, reporting
on the changes in the birdlife of Mona Island,
Puerto Rico, cited only two previous studies of
avifaunal turnover on islands that had come to
their attention, one involving the California Chan-
nel Islands, the other, the island of Karkar off the
north coast of New Guinea.
The Channel Islands, a temperate-zone archi-
pelago of nine islands or island groups off southern
California, with areas of 1 to 96 square miles and
distances of 8 to 61 miles from the mainland, were
compared for avifaunal turnover between 1917
and 1968, by Diamond (1969). Minimum turnover
rates per island ranged from 17 to 62 percent of
the breeding land and freshwater species over the
intervening 51 years, or 0.33 percent to 1.2 percent
per year. The number of species per island re-
mained about the same, provided the habitat con-
ditions had also remained about the same and the
two surveys were equally thorough.
Diamond did not include species lists in his anal-
ysis. Data from the earlier survey (A. B. Howell,
1917) show that the Channel Islands had at that
time a resident land avifauna of approximately 35
species with a passerine-nonpasserine ratio of 1.92
and a breeding land avifauna of approximately
41 species with a passerine-nonpasserine ratio of
1.73. The only comparative information to have
come to my attention is contained in a statement
by N. K. Johnson (1972:313) that approximately
41 species of land birds breed in the Channel
Islands. Evidently the size of the breeding land
avifauna has not changed since 1917. Whether or
not the passerine and nonpasserine proportions
have changed cannnot be determined in the absence
of comparative species lists.
If one were to sum into an aggregate total the
numbers of species tabulated by Diamond for each
of the nine islands in 1917 and 1968, the minimum
turnover rate for the archipelago would be 29 per-
cent for the 51-year period or 0.57 percent per
year; the aggregate species totals would be 166 in
1917 and 177 in 1968. The increase in species totals
of about six percent seems to be largely due to the
fact that the island (Santa Rosa) with the largest
increase (from 14 species to 25) had been incom-
pletely censused in 1917. Omitting Santa Rosa,
the archipelago shows an identical aggregate total
of 152 species in 1917 and again, a half-century
later, in 1968; the turnover rate remains the same
at 29 percent.
Karker, a volcanic island lying 10 miles off the
nearest point on New Guinea, has an area of 142
square miles, a maximum elevation of 6100 feet,
and 88 percent of its area below 2500 feet; the
fauna has been derived entirely by overwater colo-
nization, mainly from New Guinea but with seven
additional species from New Britain; recent vol-
canic activity has not affected bird distribution
significantly; the original vegetation was princi-
pally rain forest, but habitat conditions were
apparently similar at the two times the island was
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
surveyed, in 1914 and in 1969 (Diamond, 1971:
2742).
Diamond (1971) compared Karkar with Santa
Cruz, the Channel Island most nearly similar to it
in size and position, for extinction rates and immi-
gration rates. Qualifying the actual figures, Dia-
mond concluded that "the figures suggest that an
adequately surveyed tropical island and a temperate
island of identical size and isolation would have
similar avifaunal extinction rates" and that, ex-
pressed as percentage of the source pool, "an
adequately surveyed tropical island would have an
avifaunal immigration rate at least twofold lower
than that of a temperate island of identical size
and isolation."
The actual figures show that the turnover rate
in the breeding land avifauna on Karkar was 15
percent (including sight records) or 13 percent
(excluding sight records) in 55 years, or 0.27 per-
cent per year (including sight records) and 0.24
percent per year (excluding sight records); on
Santa Cruz, the turnover rate was 17 percent in 51
years, or 0.33 percent per year. Thus the actual
figures suggest that the yearly turnover rate is
moderately to appreciably lower on Karkar than
on Santa Cruz.
The extinction rate for the breeding land avi-
fauna on Karkar works out to 0.13 percent per
year with or without sight records; on Santa Cruz,
0.32 percent per year. Thus the extinction rate on
tropical Karkar, far from being similar to that on
temperate Santa Cruz, is more than twofold lower.
Apart from Diamond's immigration rates, which
are calculated in relation to the size of the pre-
sumed source pool whose precise composition is
largely based on surmise, it might be added that
on Karkar, over the 55-year period, and on Santa
Cruz, over the 51-year period, the number of land
species involved in the turnover, 12, was the same
on either island.
In 1914, the number of resident land species
collected on Karkar was 40, with a passerine-
nonpasserine ratio of 0.48. Between 1914 and 1969,
three of the land birds may have become extinct
(all nonpasserines), while ten, presumably breed-
ing, land birds (eight nonpasserines and two pas-
serines) apparently colonized the island. In 1969,
within the same elevational limits as in 1914, the
number of species (including sight records) was 47,
with a virtually unchanged passerine-nonpasserine
ratio of 0.47. On the basis of the two surveys, the
total number of resident land species known from
Karkar is 54, with a passerine-nonpasserine ratio
of 0.46.
Mona Island, with an area of 24 square miles,
"lies nearly midway between Puerto Rico and His-
paniola, being 42 miles from the former and 36
miles from the latter... . About 90% of the island's
surface is contained in a nearly featureless plateau
covered by a thick, mostly evergreen, sclerophyll
scrub. Over most of the plateau the tree stratum
reaches a height of 15-20 feet. . . . A dense layer
of 2-3 foot shrubs forms the understory,. . ." (Ter-
borgh and Faaborg, 1973:760).
In the first comprehensive survey, in 1901, and
again in the last, in 1972, on Mona (see Terborgh
and Faaborg, 1973:761), the nonintroduced resi-
dent land avifauna remained at three passerines
and five nonpasserines, despite two extinctions
and two natural colonizations (all nonpasserines);
the breeding land avifauna consisted of three pas-
serines and seven nonpasserines in 1901, three
passerines and eight nonpasserines in 1972. If a
parrakeet not recorded since 1892 be included, the
number of extinctions is increased by one in either
category; the passerine-nonpasserine ratio for the
native residents was presumably 3:6 in 1892
compared to 3:5 in 1972 and, for the breeding
species, 3:8 in 1892 and 3:8 in 1972. Or, based on
the annotated complete list of the birds of Mona
by Raffaele (1973), the resident land avifauna
presumably consisted of three passerines and eight
nonpasserines in 1892 compared to three passerines
and eight nonpasserines in 1972; the breeding land
avifauna consisted of three passerines and eight
nonpasserines in 1892 compared to three passerines
and nine nonpasserines in 1972.
Between 1901 and 1972, Mona had a yearly turn-
over rate of about 0.34 percent both among the
nonintroduced resident land birds and the breed-
ing land birds, whether based on Terborgh and
Faaborg (1973) or on Raffaele (1973). This yearly
turnover rate, coincidentally or not, closely
matches the 0.33 percent on Santa Cruz but not the
0.27 percent with, or 0.24 percent without, sight
records on Karkar.
The yearly extinction rate on Mona works out
to 0.35 percent (based on Terborgh and Faa-
borg) or 0.27 percent (based on Raffaele) for the
resident land avifauna, 0.27 percent (based on
NUMBER 212 31
Terborgh and Faaborg) or 0.26 percent (based on
Raffaele) for the breeding land avifauna. If the
extinct parrakeet is included, the yearly extinction
rate since 1892 is either 0.44 percent (based on
Terborgh and Faaborg) or 0.34 percent (based on
Raffaele) for the resident land avifauna, and
either 0.34 percent (based on Terborgh and Faa-
borg) or 0.33 percent (based on Raffaele) for the
breeding land avifauna. It appears, therefore, that
the yearly extinction rate of either 0.26 or 0.34
percent (based on Terborgh and Faaborg) for
the breeding land avifauna on tropical Mona is
roughly similar to the 0.32 percent on temperate
Santa Cruz, whereas the 0.13 percent of tropical
Karkar is more than twofold lower than on Santa
Cruz or Mona. The prediction that small, far
islands, such as Mona, and large, near islands, such
as Karkar, should have similar extinction rates "is
gratifyingly upheld by these two tropical islands
on opposite sides of the globe" (Terborgh and
Faaborg, 1973:762) is not supported by the above
data.
COLONIZATION RATES.?On Santa Cruz, Karkar,
and Mona, the islands for which some documenta-
tion of turnover exists, the resident land avifauna
shows a colonization rate of 0.34, 0.34, and 0.35
percent per year, respectively, whereas the corre-
sponding extinction rates are 0.32, 0.13, and either
0.35 or 0.27 percent per year. If it be assumed that
these islands are in dynamic balance between
colonization and extinction, the imbalance in the
case of Karkar is due either to a real difference
between the two rates or, more likely, to the
earlier survey having been less thorough than the
later one. In the latter instance, the larger number
of colonizations than extinctions becomes an arti-
fact, and the colonization rate is probably as low
as the extinction rate.
Diamond (1971) calculated immigration rates,
not on the basis of newly established species rela-
tive to the number of species known to have bred
on Santa Cruz or on Karkar, but as a percentage of
a pool of presumed colonizer species on the main-
land adjacent to either island. Moderately isolated
Mona and well-isolated Cocos, however, provide
examples of the difficulties to be encountered
when attempting to estimate the size, contents, or
even the location of a source pool.
Mona Island was compared with the Guanica
Reserve in southwestern Puerto Rico by Terborgh
and Faaborg (1973:765), who considered the two
very similar ecologically and avifaunally, to the
extent that all the land birds currently resident
on Mona are to be found among those nesting at
Guanica and constitute a subset of the latter.
Guanica thus harbors a species pool of some 43
prospective colonizers (based on Kepler and Kep-
ler, 1970; Cameron B. Kepler, in litt.). Yet four
(all nonpasserines) of the 14 species of land birds
known to have bred on Mona (see Terborgh and
Faaborg, 1973:761) are not included in the Guan-
ica list, two species (both of them Antillean non-
passerines) that have occurred but have not bred
on Mona are not known from Guanica, while
another six species (all Antillean passerines) that
breed at Guanica have reached Mona, where
Terborgh and Faaborg (1973) and Raffaele (1973)
regarded them as accidentals, vagrants, or other-
wise unestablished visitants rather than propagules
emanating from a pool of potential colonizers.
Cocos Island has no record at all of immigrants
that could qualify as potential colonizers. Not in-
cluding the Galapagos Archipelago, the species
pool from which to draw extends through south-
ern Central America and northern South America
and is enormous. Though the distance from the
mainland is not great compared to interisland
distances in the tropical western Pacific and has
not prevented the arrival of numbers of North
American land bird migrants, I would hesitate to
nominate a single species on the tropical American
mainland as a candidate to successfully colonize
Cocos. "One cannot predict on the basis of morpho-
logical, ecological or physiological characteristics
whether or not a given species will be a successful
colonizer" (Mayr, quoted by Moreau, 1966:368).
The chances that islands will be colonized by
species that breed at warmer or cooler latitudes
distant from those in which the islands lie are
close to zero. A way to estimate the probability
that certain islands or island groups are apt to
have higher or lower colonization rates than others
is furnished simply by counting the numbers of
residents and of nonresidents in a species list. With
few exceptions, lists in which the ratio of residents
to nonresidents is less than 1:1 pertain to islands
or island groups outside the tropics, lists in which
the ratio of residents to nonresidents is more than
1:1 pertain to islands or island groups inside the
tropics. Though sufficient examples are hard to
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
find, the ratios of islands that are at least roughly
comparable to one another in position and area or
size of avifauna also tend to follow a latitudinal
progression, as in the accompanying tabulation of
single islands lying off the coast of eastern and
southeastern Asia.
No. of Residents:
Area land No. of non-
(mi*) Latitude birds residents residents
Hokkaido 30,300 42-45? 155 52 031
Honshu 87,000 34-40? 184 84 0.84
Taiwan 13,800 22-25? 219 127 1.38
Hainan 13,000 19? 205 140 2.15
Singapore 217 219 156 2.48
Ratios of island groups increase even more
markedly than do those of the single islands from
the Japanese Empire in the north to the Papuan
subregion at the equator: Japan, 1.4; Philippines,
5.1; Sula Islands, 5.9; Moluccas, 8.3; Lesser Sundas,
13.2 (average of six islands); New Guinea and
satellites, 17.6.
Checklist data that are at hand point to only
three nontropical islands or island groups on
which the residents outnumber the nonresidents.
All three are located peripherally in the Southern
Hemisphere, where they face open, cold oceans
and have land avifaunas consisting largely to
almost entirely of nonmigratory populations: the
Falklands (with a borderline ratio of 1:1), east of
the southern tip of South America; Tasmania, off
southeastern Australia; and oceanic New Zealand.
In the tropics, the few islands on which non-
residents outnumber residents occur mostly in the
Western Hemisphere. Cuba, Cozumel, and the
Swan Islands are easily explainable as stepping-
stones utilized by transients crossing the Gulf of
Mexico and the Caribbean Sea; depauperate Bar-
bados has an "abnormally" small number of resi-
dents compared to the cumulative number of mi-
grants and strays it has been receiving over the
years; the Virgin Islands, as a political entity, have
a ratio of 0.65, but if St. Croix, which lies isolated
off the Puerto Rican shelf, be excluded, the ratio
rises to a "normal" 1.21 (compared to the 1.31 of
Puerto Rico). Elsewhere, the Aldabra archipelago
in the Malagasy area, with a below-unity ratio
based on 12 resident and 15 nonresident land birds,
apparently is in position to intercept transients
and strays to or from East Africa and Madagascar
(Benson and Penny, 1971:512). In the same area,
the Comoros, which lie closer to East Africa and
Madagascar, and the Seychelles and the Mascarene
Islands, which lie farther out in the Indian Ocean,
have more recorded residents than nonresidents.
Mona in the Caribbean area and Cocos in the
eastern Pacific, with resident:nonresident ratios of
0.35 and 0.17, respectively, run counter to expecta-
tion. The situation on Mona may be similar to
that on Barbados, where the number of residents
is small on an island that is accessible to a mount-
ing number of irregular and accidental visitors.
Cocos appears to be too small, too distant, too
isolated, and conceivably too inhospitable for neo-
tropical species of land birds to reach it or to
colonize it. Evidently it is not too distant for strays
and, probably, a number of regular transients and
visitors whose migratory urge causes them to strike
out over open ocean.
TURNOVER AND ENDEMISM.?A relationship be-
tween island size and percent endemism among
the species of land birds was postulated by Mayr
(1965): "The percentage of endemic birds on
islands increases with island area at a double loga-
rithmic rate. This relation is apparently due to
extinction, which is more rapid the smaller the
island." This view was seconded by MacArthur
and Wilson (1967:173-174), who wrote that "the
percentage of non-endemic species is probably a
measure of the turnover rate," that "we know on
both theoretical and empirical grounds that the
turnover rate of species varies inversely with island
area," and that the data in the above-cited note
by Mayr are consistent with the prediction that
percentage endemicity should increase with island
area.
Predicated upon the four islands he selected for
plotting on a double-log graph, Mayr (1965) made
the statement that solitary, well-isolated islands
show only small deviations from expectancy. Read
from his graph, the expectancy is that the endemic
species of land birds range from about seven per-
cent on an island of 10 square miles to about 75
percent on an island of 300,000 square miles.
It happens that the one island on which I pub-
lished a report, Cocos, in the tropical eastern Pa-
cific, I selected precisely because it is a solitary,
well-isolated island (Slud, 1967). Its four species
of resident land birds, made known to science in
the nineteenth century, are the same four species
found there today. No other species of resident land
NUMBER 212 33
bird has ever been reliably reported by any of the
many visitors to the island.
Thus, Cocos Island, surveyed from 1891 to 1963,
may not have come to the attention of Terborgh
and Faaborg (1973) as an example of avifaunal
turnover on islands, because it hasn't had any. The
statement by Mayr (1965) that, the smaller the
island, the turnover "appears to be far more rapid
than hitherto suspected" or the one by Diamond
(1969:57), based upon the Channel Islands, that
"percentage turnover rates vary inversely as insular
species diversities, with no effect of distance ap-
parent" cannot begin to apply to Cocos, with four
species of resident land birds, an area of 10 square
miles, and surveys the extremes of which are spaced
72 years apart. Nor does the statement by Terborgh
and Faaborg (1973:762) that "the equilibrium the-
ory qualitatively anticipates similar extinction
rates on small far islands and large near islands"
apply to Cocos, since it has neither an extinction
rate nor an immigration rate that can be used to
uphold theoretical expectancies.
In regard to the predictably low percentage of
endemic species of land birds on small islands
claimed by Mayr (1965) and by MacArthur and
Wilson (1967), little Cocos Island, instead of an
expected endemicity of seven percent, has an actual
endemicity either of 50 percent (if one excludes
the Cocos Island cuckoo), a higher rate than on
any island or island group, save Madagascar, in any
of the insular categories graphed by Mayr (1965),
or of 75 percent (if one includes the Cocos Island
cuckoo), a rate as high as that of 22,000 times
larger Madagascar.
"A further consequence," wrote Mayr (1965),
"is that the smaller the island the lower the percent-
age of endemic species should be, because most
of the populations become extinct before they
reach species level, or soon thereafter. MacArthur
and Wilson. . . , on the basis of slightly different
considerations, arrived at a similar conclusion."
Again Cocos stands apart. The first land bird to be
discovered there, around 1840, was the Cocos Is-
land cuckoo. A well-marked Pacific isolate immedi-
ately distinguishable from the Middle American
and Caribbean Mangrove cuckoo (Coccyzus minor),
it is either an endemic race of the latter or an
endemic species confined to Cocos. The least com-
mon of the land birds, though by no means rare,
it seems not to have decreased but perhaps even to
have increased in numbers. The Cocos Island fly-
catcher is the sole representative of an endemic
genus (Nesotriccus) restricted to Cocos. Previously
deemed rare to infrequent, it was common and
widespread in 1963 (Slud, 1967:286). The third
endemic is the abundant Cocos Island finch (Pi-
naroloxias), the sole representative of the genus
and found only on Cocos. The fourth species of
land bird on Cocos is a mangrove-type warbler
(Dendroica petechia). Belonging to the race aure-
ola, it is endemic to the Galapagos Archipelago
and Cocos Island, on both of which it is without
doubt a relatively recent arrival. Unlike the other
three species on Cocos, the unknown geographical
origin of which could have been West Indian
rather than continental, aureola is weakly differ-
entiated from the population on the opposite
coast of South America.
" 'There is little doubt that . . . well isolated
islands are evolutionary traps, which in due time
kill one species after another that settles on them' "
(Mayr, 1965). On Cocos, however, sufficient time
has evidently elapsed for two of the land birds to
have evolved, presumably in situ (if not marooned
there as relicts), to the generic level and a third,
almost if not quite to the specific level. Moreover,
the two that are least common are not rare and,
instead of disappearing, have apparently been in-
creasing in numbers. Cocos Island is either an
actual or a potential death trap for all its land bird
arrivals, every one of which, so far as known, is a
northern migrant that has not become established
or even lingered there unseasonably.
Island Types
CONTINENTAL ISLANDS.?"Recent" Continental
Islands: Darlington (1957:480) loosely categorized
obviously recent continental islands as those with
more or less reduced continental faunas, such as
Sumatra, Java, Borneo, New Guinea, Ceylon, and
Trinidad in the tropics, Formosa (Taiwan) on
the edge of the tropics, and the British Isles, Japan,
Newfoundland, Greenland, Tasmania, Tierra del
Fuego, and the Falklands outside the tropics. Mo-
reau (1966:302-303) gave as typical examples the
small islands of Fernando Po, Zanzibar, and Mafia
on the continental shelf of Africa, which, cut off
from the mainland in the Late Pleistocene as a re-
sult of the postglacial rise of ocean level, are too
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
young for evolution to have proceeded far, are
located sufficiently close inshore for gene-flow from
the continent to hamper the divergence of the
insular populations, and at the start of their inde-
pendent existence had a full complement of species.
Recent tropical continental islands have native
land avifaunas with passerine-nonpasserine ratios
that are generally about the same as or somewhat
reduced from the ratio on the opposite mainland
(Figures 13 and 15). The proportional representa-
tions of passerines and nonpasserines on the island
may thus tell us something of the relative age of
the island or the recency of its avifauna.
Tropical American examples are Isla Coiba,
with a ratio of 1.17, compared to the facing slope
of the Azuero Peninsula, 1.29, and the Pearl Archi-
pelago, with a ratio of 1.00 (ranging from 0.95 to
1.44 for the three main islands), compared to the
Canal Zone, 1.46, or all of Panama, 1.43; Margarita
and Trinidad, with respective ratios of 0.93 and
1.09, compared to northeastern Venezuela, with a
ratio of 1.06, while all of Venezuela has a ratio of
1.61. Another probable example of islands on the
continental shelf is the offshore Tres Marias, with
a ratio of 1.26, compared (in lieu of Nayarit, which
lacks a checklist) to Colima and the immediately
adjacent portion of Jalisco, with a ratio of 1.34.
Possibly another example is Isla Cozumel, with a
ratio of 1.00, compared to nearby Quintana Roo,
with a ratio of 1.06, and the Yucatan Peninsula,
with a ratio of 1.03; the adjoining, more humid
Pete"n and British Honduras have respective ratios
of 1.21 and 1.18. In the Old World tropics I have
even fewer examples for which matching data are
available: Ceylon, 0.85, compared to Kerala, at the
tip of peninsular India, 0.89; Sumatra, 1.24, com-
pared to the Malay Peninsula, 1.28, and Singapore,
1.21, compared to Malaya, 1.20.
In the temperate zone, at least at northern lati-
tudes, passerine-nonpasserine ratios on continental
islands exceed those on the neighboring mainland,
but again available matching data are hard to
come by. Examples are Taiwan (Formosa), at the
edge of the tropics, with a ratio of 2.23, and
Fukien, 1.92; Vancouver Island, 1.42, and southern
British Columbia, 1.19; Prince Edward Island, 1.50,
and New Brunswick, 1.28, or Nova Scotia, 1.24;
Great Britain, 2.00 and Ireland, 2.54 compared to
the Netherlands, 1.83, or France (including Bel-
gium and Switzerland), 1.66; Newfoundland Is-
land, 1.48, and Newfoundland Labrador, 0.58, or
the entire Labrador Peninsula, 0.62. At southern
latitudes, my only examples have the ratio smaller
on the continental island: Tasmania, 1.59 and
Victoria, on the north side of Bass Strait, 1.71; Isla
Grande (Tierra del Fuego) and the Falklands (300
miles out in the ocean and with only 19 species
of land birds), respectively, 0.94 and 0.90 compared
to southern Argentina, 1.77. In respect to Tasma-
nia, it could perhaps be a case of one island, Tas-
mania, being satellic to a very much larger island,
Australia (see below). On the other hand, Tasma-
nia, Isla Grande, and the Falklands, lying at the
southern edge of the terrestrial world (they can
expect no land birds from Antarctica) would
appear to correspond in their situation to islands
at far northern latitudes, where the ratios are all
extremely reduced (Greenland, 0.75; Southampton
Island, 0.33; Banks Island, 0.29).
Intraisland comparisons are extremely hard to
find. Only altitudinal data are at hand and solely
for Java (based on Hoogerwerf, 1948): 229 species
and a ratio of 1.01 between sea level and 1500
meters, 111 species and a ratio of 1.52 between
1000 and over 3000 meters. These figures, i.e., rela-
tive numbers of species and passerine-nonpasserine
ratios, correspond to those on the nearest compar-
able mainland, the Malay Peninsula: 342 species
and a ratio of 1.02 in the lowlands (based on
Chasen, 1939), 257 species and a ratio of 1.67 in
the hill country (based on Robinson, 1928).
"Old" Continental Islands: The east African trio
of Zanzibar, Mafia, and Pemba, with respective
areas of 640, 240, and 320 square miles, are low,
generally similar islands, none more than 30 miles
distant from the Tanzania mainland (for which I
have no list). The most striking difference, the fact
that Zanzibar and Mafia lie on the continental
shelf and Pemba does not, is not obvious to the eye
(Moreau, 1966:350-351), but it is certainly sug-
gested by the passerine-nonpasserine ratios: Zanzi-
bar, 1.10; Mafia, less effectively isolated than
Zanzibar, 1.27; old-continental Pemba, 0.65. "The
passerine bird fauna of Pemba . . . looks almost like
that of an oceanic island, and it is extremely diffi-
cult to imagine why this should be so" (Moreau,
1966:352). Evidently great distance is not neces-
sarily required to produce an oceanic effect, partic-
ularly in tropical latitudes.
Socotra, like Pemba, is probably "an 'old' conti-
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ds
 
fr
in
gi
ng
 
th
e 
m
ai
nl
an
d:
 
a.
Sw
an
 
Is
la
nd
s;
 
b,
 
Ba
rb
ar
et
a;
 
c,
 
U
til
a;
 
d,
 
G
ua
na
ja;
 
e,
 
R
oa
ta
n;
 
f, 
Cu
leb
ra
; 
g,
 
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ne
ga
da
; 
h,
 
V
irg
in
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or
da
; 
i, 
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.
 
Jo
hn
; j
, 
To
rt
ol
a;
 
k,
 
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.
 
Th
om
as
; 
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.
 
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m
, 
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on
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n
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V
ieq
ue
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o
, 
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rg
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Ar
ub
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q,
 
Bo
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; 
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s,
 
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ba
go
; 
t, 
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zu
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el;
 
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, 
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on
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; 
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in
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ad
.)
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
nental island dating from Plio-Pleistocene times, so
that opportunity for evolution on its surface has
been considerable" (Moreau, 1966:303). Situated
130 miles of! the Somali coast, it has a resident
land avifauna of 15 passerines and 14 nonpasser-
ines and a dose-to-unity ratio of 1.07. This ratio is
virtually equivalent to the 1.04 of former British
Somaliland, with which Socotra presumably was
once connected. The approximately equal propor-
tion of passerines to nonpasserines also charac-
terizes many isolated islands, particularly in the
Indian Ocean (see below).
FRINGING ARCHIPELAGOS.?Under this heading,
Darlington (1957, Chapter 8) enumerated the
Philippines, the islands of the western Pacific, and
the West Indies as the principal archipelagos of
the world (all tropical),
which, though not simply recent continental, lie close to con-
tinents and receive the fringes of continental faunas and are
extensive enough, with enough comparable islands, to show
informative patterns of distribution. The archipelago of Wal-
lacea [Celebes, Lesser Sundas, Moluccas] . . . might be added
to this list. It lies between the continental shelves of Asia
and Australia and receives the fringes of the faunas of both
continents.
West Indies: Main Islands: Recent attempts at
area-species formulations for islands have come to
be associated most prominently with MacArthur
and Wilson (1963, 1967). The double-log area-
species curve of the West Indian herpetofauna
presented by these authors (1967:8) is a virtually
perfect model, though it only includes the four
major Greater Antillean islands at one end of the
scale of magnitude and three of the smallest and
northernmost Lesser Antillean islets at the other
end.
Using the same coordinates, I plotted the area-
species distribution of the land birds, as well as
their corresponding passerine-nonpasserine ratios,
for the islands of the entire Caribbean area (Figure
12). As have other authors, I excluded migrants.
My reason is that islands range in status from
migrant-free in parts of the tropics to migrant-
dominated outside the tropics, particularly in the
northern hemisphere. Hence the only common
basis upon which to make intercomparisons is the
resident avifauna. Islands 1, 6, 22, 23, 24, and 25
are the same as in the above model, only omitting
the smallest, Redonda (area 1 square mile), for
which I have no list. Remarkably, these islands
align themselves as ideally for the birds as for the
reptiles and amphibians. The neatness of the pic-
ture is spoiled, however, when the other islands
are included.
The two regressions in Figure 12 are constructed
upon the islands numbered from 1 to 25, compris-
ing the Greater and Lesser Antilles and a scattered
few in the western Caribbean Sea (excluding satel-
lites, e.g., Isle of Pines, Gonave, islets on the
Puerto Rican shelf). These satisfy two require-
ments: all are single islands, except the Grenadines,
and all are zoogeographically West Indian (in the
sense of Bond, 1948). Increasing numbers of spe-
cies show a very high, positive correlation with
increasing area. Islands 1 through 14, ranging in
size from 5 to 100 square miles and in number of
species from 11 to 24, lie along approximately the
same level. Their small avifaunas could as well be
explained by their geographic position out at sea
as by their small areas. A step higher lie islands 15
through 21, comprising the principal members of
the Lesser Antilles, that is, Guadeloupe through
Grenada. These range in size from 100 to some 500
square miles and have a compiled species-number
range from 36 to 43 (except for outlying, over-
populated, impoverished island 17, Barbados).
The passerine-nonpasserine ratios correlate neg-
atively with increasing area along a slightly falling
slope. I do not attribute this correlation, though
highly significant statistically, to the geometric
increase in area which the West Indies are, so to
speak, preadapted to show on a graph (Figure 12).
Rather, the four main Greater Antillean islands
(22-25) are sufficiently large and old to have a
faunal evolutionary history of their own. Singly or
collectively they have below-unity ratios, as do
other comparable tropical noncontinental islands.
Cuba (25), the largest of the four, has the lowest
ratio, but it also has the simplest physiography.
The other numbered islands (1-21) are very much
smaller, and all but one have ratios rising above
unity. The exception, Antigua, a low, dry island
converted to sugar production, would reach unity
with the addition of one passerine. Small and iso-
lated Grand Cayman (13) is structurally Greater
Antillean but Lesser Antillean in size and ratio.
Offshore Islands and Satellites: Islands that lie
off the Caribbean shores of Middle America and
northern South America or are satellic to the main
West Indian islands are symbolized by letter in
NUMBER 212 37
Figure 12. Islands that pertain to the same complex
tend to align themselves along a slope which lies
parallel to that of either regression, particularly
the one of area-ratio, and at a different level from
that of another island complex. (An attempt to
cluster all the Caribbean islands two-dimensionally
into natural groupings based on passerine-
nonpasserine and suboscine-oscine ratios is pre-
sented in Figure 30.)
Margarita and Trinidad, off northeastern Vene-
zuela, were included above with continental islands,
as was also Isla Cozumel, off the Yucatan Penin-
sula. Cozumel, strictly speaking, appears to be
somewhat separated from the continental shelf
proper, hence could belong in the offshore cate-
gory, together with the Bay Islands, off Honduras,
and the Dutch Leewards, off northwestern Vene-
zuela.
The Bay Islands of Honduras "are evidently the
tops of a submerged mountain range, which,
through isolation from the mainland, have retained
a number of Central American species, which have
entirely or mostly disappeared from Central Amer-
ica proper and which appear as relicts, not only
on the Bay Islands, but also on islands off the
coast of Yucatan" (Bond, 1936:354). Of the main
islands, "Utila is situated on the Continental Shelf,
separated from the mainland by waters not exceed-
ing 10 meters in depth, while Roatan, Guanaja,
and associated smaller islands are surrounded by
waters at least 275 meters deep" (Monroe, 1968:
11). Utila, however, is the smallest of the major
islands. It reaches an elevation of only 90 meters
and, unlike the others, is otherwise flat. The major
islands are all small, the fauna is very limited, and
the passerine-nonpasserine ratios are extremely
low.
The low-lying, dry Dutch Leewards (Aruba,
Curasao, Bonaire) are situated only 20 to 40
miles off Venezuela. Despite the short distance, the
islands, particularly Curasao and Bonaire, are
separated from the mainland by deep water (Fig-
ure 13). Both geologically and zoogeographically,
all three islands seem never to have been part of
the South American mainland (Voous, 1957:32).
All have below-unity passerine-nonpasserine ratios,
including Aruba, which is closest to shore but is
also the most barren. Possibly the low ratios of the
islands are a reflection of the composition of the
avifauna on the arid opposite coast; endemism on
the islands is extremely low, it is confined to the
subspecific level, and it affects only four of the land
birds. The only figure I have for the arid opposite
coast, 1.14, is based solely on the 24 passerines and
21 nonpasserines reported in the lowlands of the
Paraguana Peninsula, opposite Aruba, by Barnes
(1940). On the other hand, the islands conceivably
show an oceanic effect, manifested by marked re-
duction in ratio compared to the mainland, as do
the following: Guadalupe Island, 180 miles out
at sea and with a ratio of 1.33, compared to Baja
California, with a ratio of 2.02; the Revilla Gigedos
Islands, over 400 miles at sea and with a ratio of
1.00, compared to Colima (and adjacent Jalisco),
with a ratio of 1.34.
Satellites are relatively small to extremely small
islands which have native land avifaunas that
range in passerine-nonpasserine ratio from about
the same as, to very much lower than, that of the
very much larger island to which they pertain.
Examples are the Isle of Pines, with a ratio of 0.60,
compared to the 0.64 of Cuba; Gonave, 0.80, Beata,
0.84, and He a Vache, 0.85, compared to Hispani-
ola, 0.88; Vieques, 0.87, compared to Puerto Rico,
0.86.
The islets, primarily including the Virgin Is-
lands, that are situated on the Puerto Rican shelf
(Figure 12/-/) or, in the case of St. Croix, associ-
ated with it are satellic to Puerto Rico. Though
often considered Lesser Antillean in atlases and
dictionaries, the Virgins have a mixed avifauna
that is predominantly Puerto Rican, while their
low passerine-nonpasserine ratios ally them to the
geologically similar Greater Antilles and separate
them from the Lesser Antilles as surely as does
the Anegada Passage (Figure 13). The islets of
the Puerto Rican shelf follow a pattern typical
of insular satellites in which diminution in area
goes hand in hand with reduction in number of
species of land birds accompanied by falling
passerine-nonpasserine ratios, probably in response
to a broadening in the size and a reduction in the
number of occupied niches.
Supplemental data for the Old World are few,
but these tend to corroborate the situation in the
New World tropics. In the Philippines (Figure
16), Mindoro, with a ratio of 0.78, is satellic to
Luzon, with a ratio of 1.09; Basilan, 0.74, is satellic
to Mindanao, 1.07. In Indonesia (Figure 15), the
West Sumatra Islands have a combined ratio of
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
? i 7 0
27
861
1.00
1.06.
I.IOk - 92^
?4I.4O
1.121
?1.46
1.381
FIGURE 13.?Passerine-nonpasserine native land bird ratios in the eastern Caribbean area.
0.95 (the Mentawei Group, 0.89; Nias, 0.89; Sima-
lur, 0.73; Enggano, 0.80) compared to Sumatra,
1.24; Bali has a ratio of 1.15 compared to that of
Java, 1.26. In the Papuan subregion, New Guinea
has a ratio of 1.71, whereas the coastal islands
range from 1.13 to 0.65. The pattern recurs among
island groups in the western Pacific.
Apparently the trend to smaller ratios may be
reversed when the areal difference is of a lesser
magnitude or when the larger island is under a
certain minimum size: Tobago, with a ratio of
1.28, compared to Trinidad, 1.09; Marie Galante,
1.46, compared to Guadeloupe, 1.12.
Distance and Ecology: MacArthur and Wilson
(1967:20) reproduced a diagram of the number of
bats on islands off Venezuela to demonstrate that
area and ecology are the critical factors affecting
species numbers rather than degree of geographic
isolation when distance can be minimized as an
influence. I have converted the diagram into the
following table in which the herpetofauna and the
avifauna (including passerine-nonpasserine ratios)
have been incorporated.
NUMBER 212 39
Species of
Xerophytic
Aruba
Curasao
Bonaire
Traces of rain
forest
Margarita
Rain forest
Trinidad
Tobago
Grenada
Area
(mi2)
69
173
95
444
1864
116
120
Species
of bats
3
6
3
7
53
16
11
amphib-
ians,
lizards,
snakes
13
11
9
17
85
42
17
Species of
resident
land
birds ;
29
29
27
77
239
82
39
Pas-
serines:
non-
passerines
0.81
0.93
0.93
0.93
1.09
1.28
1.17
With respect to the bats, the above authors pointed
out that "Grenada, which is both small and iso-
lated, still has four more species than the large,
near island of Margarita, and many times more
species than the small, near islands of Aruba and
Bonaire." The statement holds poorly for the her-
petofauna and not at all for the avifauna. Grenada,
instead of more species of birds, has half as many
species as Margarita and only one-third more than
the xerophytic Dutch Leewards.
Distance and faunal origin could be responsible
for the different numbers, whether of bats, of
amphibians, lizards, and snakes, or of birds, on
Tobago and Grenada. Both are rain-forested, high
islands of equal size and with similar passerine-
nonpasserine ratios: Tobago lies close to and is
fed by its parent, Trinidad; Grenada sits apart as
the terminal link in the Antillean chain. In the
case of Margarita, the island may have traces of
rain forest, but these are severely restricted to tiny
areas of high ground; the island is otherwise as
arid as the Araya Peninsula on the opposite main-
land (see Ewel et al., 1968, map). Margarita, apart
from its much larger size and closer distance to
shore, is a continental island, and it has more
than twice as many species of land birds than non-
continental Aruba, Curasao, or Bonaire, individ-
ually or as a group. In the accompanying table, it
appears that the passerine-nonpasserine ratios tend
to correlate with the climato-vegetational condi-
tions of the islands, whereas species numbers,
unless individually qualified, do not.
Southeastern Asia, New Guinea, and the West-
ern Pacific: MacArthur and Wilson (1967:23)
presented an area-species curve of the land and
freshwater birds based on a selection of continental
and noncontinental islands and archipelagos that
included the Sundas, Philippines, and New Guinea
(but not the Moluccas), as well as tiny, isolated
Christmas Island in the Indian Ocean. I was able
to compile lists of resident land birds for 17 of
MacArthur's and Wilson's 23 entries, substituting
the Anamba, Natuna, and Palawan islands for the
omissions. These are represented by the 20 num-
bered entries upon which the regressions are con-
structed in Figure 14. (Also see Figure 15.)
The steepness of the area-species slope is caused
by the very low values of the very small entries and
the high values of the very large entries. If the
extremes were removed, the slope would be de-
pressed toward that of the West Indies; the middle
entries are scattered more or less horizontally, as
in the West Indies (Figure 12). The inclusion of
tiny Christmas Island contributes to the weakly
positive correlation between area and ratio. With-
out Christmas Island, the area-ratio slope would
show a closer approach to the horizontal.
Supplementing the numbered entries are the
lettered entries. These include individual islands
in the Moluccas, in the Philippines, and on the
western New Guinea, or Sahul, shelf. Most of the
lettered entries fall amid the numbered entries
and strengthen the correlations, especially that of
area-ratio. (In the West Indies the lettered entries
weakened the correlations, especially that of
area-ratio.)
The East and West Indies appear to differ mainly
in the steepness of their area-species slopes and in
positive versus negative correlation between area
and ratio. These differences may be directly attribu-
table to their contrasting magnitudes, due to the
paucity of small islands in the East Indies. For
example, Dominica, one of the principal islands
in the Lesser Antilles, is only the size of New York
City. It occurs about midway in Figure 12, but
would be one of the smallest if entered into Figure
14. Thresholds of insular size in relation to step-
wise increase in faunal, and presumably environ-
mental, complexity could be involved, as well as
faunal origin and history and the distances and
directions the islands lie from one another and
from faunal reservoirs.
Philippine Islands: The Philippines are incom-
pletely separated from the Sunda shelf. The
passerine-nonpasserine ratio of 1.30 for the archi-
pelago as a whole, including Palawan, or 1.20
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
AREA IN SQUARE MILES
FIGURE 14.?Malayo-Papuan insular native land avifaunas: correlation between area and numbers
of species (numbered large open circles and dashed regression line) and between area and
passerine-nonpasserine ratios (numbered large solid circles and solid regression line). (1, Christ-
mas; 2, Enggano; 3, Anambas; 4, Simalur; 5, Natunas; 6, Nias; 7, Lombok; 8, Bali; 9, Mentawai;
10, Sumba; 11, Palawan; 12, Flores; 13, Sumbawa; 14, Timor; 15, Java; 16, Celebes; 17, Philip-
pines (minus Palawan); 18, Sumatra; 19, Borneo; 20, New Guinea. Lettered entries (small circles)
include islands in the New Guinea, Moluccan, and Philippine areas: a, Kei Is.; b, Japan; c, Biak;
d, Waigeu; e, Sula Is.; f, Aru Is.; g, Bum; h, Ceram; i, Halmahera; j , Basilan; k, Bohol; I, Cebu;
m, Leyte; n, Mindoro; o, Panay; p, Negros; q, Samar; r, Mindanao; s, Luzon; t, Hainan.)
without Palawan, is slightly less than the average
for the Greater and Lesser Sundas (Figure 15).
The two largest islands, Luzon and Mindanao,
have about the same area and virtually identical
species totals and somewhat above-unity ratios
(Figure 16). Either island features a long, penin-
sular extension which, if lists were available, would
no doubt prove to have a distinctly lower ratio
than the entire island, just as in mainland situa-
tions. The principal smaller islands have the re-
duced ratios that characterize satellites and species
numbers that correspond more or less to individual
island size and position. Only Panay, compared to
the other islands, is impoverished, not, as has been
stated (Delacour and Mayr, 1946:12), the entire
central province. Palawan, lying between Borneo
and the Philippines, is closer to Borneo zoogeo-
graphically but more like similar-size islands of the
Philippines in number of species and below-unity
ratio. In both these respects, however, it also acts
as a satellite of Borneo.
The same variables apply of course to the numer-
ous other small islands lying between southeast
Asia and Australia (Figure 15). These occur as
NUMBER 212 41
loose or tight, linear or clumped configurations
situated beside or between larger masses of land,
compared to which they have markedly lower
ratios. Distance and position, type of island, and
history contribute to and help cause this effect.
Examples are the Andaman, Nicobar, West Su-
matra, and Kangean islands, also to some extent
the Anambas and Natunas, in the Malaysian area,
and the Aru, Admiralty, D'Entrecasteaux, and
Louisiade groups, as well as peripheral, or satellic,
Waigeu, Japen, and Biak, in the New Guinea area.
Wallacea: The Lesser Sundas are situated upon
a projection of the Sunda shelf extending eastward
from Java, a disposition that brings to mind the
Antillean volcanic arc stretching north of Vene-
zuela. They have above-unity ratios, except Lom-
bok and Sumba.
Lombok is narrowly separated from similar-size
Bali yet has a markedly lower ratio and a one-third
smaller avifauna. The smaller size of the avifauna
could be due to the zoogeographic effectiveness of
Lombok Strait, between Bali and Lombok, which
apparently persisted through the Pleistocene while
the strait between Bali and Java dried up at the
height of the glaciation (Mayr, 1944b:7). The low
ratio on Lombok is perhaps explained by the
"heavy Pleistocene eruptions of Mount Rindjani
[which] seem to have destroyed much of the moun-
tain fauna" (Mayr, 1944b:9). The predominantly
montane passerines would thus have been affected
far more adversely than the nonpasserines. The
implication that the number of species on Lombok
ought to be higher is difficult to prove. Lombok, in
fact, has a few more species than thrice-larger Sum-
bawa, an even closer neighbor than Bali, with
which it was presumably connected at the height of
the Pleistocene glaciation (Mayr, 1944b:9).
Sumba, twice as big as Bali and more like Flores
and Sumbawa in size, has fewer species than any
of the Lesser Sundas figured and only two-thirds
the number on Flores 25 miles away. The fact that
Sumba is a low and arid island without a moun-
tain fauna (Mayr, 1944a: 176) could in itself ac-
count for the low species total and low ratio.
According to Mayr (1944a: 181), "lack of pro-
nounced endemism indicates either that Sumba is
a young island, or that it has had a continued
active faunal exchange with neighboring islands,
or both." Applied to Sumba in relation to its
similar-size neighbors, this reasoning would seem
to demand a higher immigration rate and species
number instead of a smaller. If greater endemism
and fewer species are a function of increasing
distance or inaccessibility, expressed as a low
immigration rate (MacArthur and Wilson, 1967),
the more distant island ought to have a larger pro-
portion of endemics and a smaller number of spe-
cies. The converse is true of Sumba with respect
to endemism but not to number of species, hence
other variables need be brought into play.
Celebes and the Moluccas (and to lesser extent
the Philippines) occupy a zone of complicated
large-scale and violent tectonic movement between
the Indo-Malayan and Australo-Papuan continental
shelves (Mayr, 1944b). Celebes and the Moluccas,
even Waigeu off the western tip of New Guinea,
have distinctly smaller avifaunas and sharply lower
ratios than do the shelf islands. This could be a
reflection of a derived fauna on the one hand and
of reduced competitive success of passerines as over-
seas colonists compared to nonpasserines on the
other hand.
A selective brake may at the same time be apply-
ing counterpressure against immigration. To quote
Mayr (1941c: 198):
. . . of the 265 species of land birds which are known from
that part of New Guinea which is opposite New Britain, only
about 80 species have a representative on New Britain. In
other words, the 45-mile-wide stretch of water which separates
the two islands has prevented the crossing over of 70 per
cent of the New Guinea species . . . . Literally hundreds of
similar instances could be listed from the distribution of birds
in the Indo-Australian archipelagos, all of them indicating
the sedentary habits of their avian inhabitants.
In the American tropical Pacific, eight families of
land birds regularly occurring on the Panamanian
mainland have not been found nearby on Isla
Coiba (Wetmore, 1957:12). Tiny, oceanic Cocos
Island has not yielded an additional species of
native land bird since the original four were dis-
covered, yet many migrants arrive alone or in
groups (Slud, 1967). Obviously a distinction must
be made between potential colonizers and visitors,
especially in the tropics.
Tropical Western Pacific: Darlington (1957:
506-507) observed that
considered together, as one fringing archipelago, all the is-
lands of the tropical western Pacific east of Australia and
New Guinea, and as far out into the ocean as terrestrial
vertebrates go . . . . None of these islands is connected with
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Palaus
1.20.'
Marianas '
1.50 \
?0*20
233' ' \ Caroline Islands 2.43
' ' ' ...!'.' '.? s - " .2-00 ?? ?
.89^  ? B l ^ ? 1 . 2 1
W. Sumatravm24^
 fls
- .95V89^
Christmas I. 33
Cocos-Keeling*
7B w ^ ^ - Keils., ^
^ - ^ ? ? '
A r u l s
-
S U N D A S 2
Louisiades
Q kilometers 1000
FIGURE 15.?Southeast Asia to tropical central Pacific: passerine-
nonpasseritye ratios of native land birds.
the continental shelf. Even the Bismarcks and Solomons are
narrowly separated from it and from each other. Zoogeo-
graphically, all these islands should be treated as of unknown
geological history, and their faunas should be allowed to
speak for themselves.
The New Guinea area is the primary center for
the evolution, radiation, and dispersion of birds
over the tropical western Pacific. Two sorts of pas-
serine decrease, relative to the nonpasserines, be-
come apparent (Figure 15). One takes place, as
already mentioned for New Guinea, on peripheral
shelf islands, the other on groups of oceanic islets,
usually low ones, with depauperate avifaunas
derived from much larger and richer, high islands.
Area, avifaunal size, and ratio, relative to in-
creasing distance from Australia and New Guinea
(see Figure 15), are summarized in the following
tabulation.
NUMBER 212 43
Marshall Is*
,1.72
.2.00
'., Gilbert Is*
Phoenix Is*
JSolomons
.94
Sta.Cruz Is.
U .93
Area L12
- . E l I ice Is*
, ..Banks Is.
^}2L Fiji 1.30
New HebridesV\
1.09.
?f Toketou Is*
Samoa L46
1.40^% ^
.
1 3 0 7
* Manuals..57
French Polynesia
,52
Marquesas
.57
j'^Lauls.
 m
r ; j?9 ?"? Tongas
Tuamotu Arch.
.40
Society Is.
.44
Austral Is* Pitcaim Is.
Solomon Is.
(excluding Rennell)
Rennell
New Caledonia Area
New Caledonia
Loyalty Is.
Santa Cruz Is.
New Hebrides Group
New Hebrides
Banks Is.
Fiji
Samoa
French Polynesia
Society Is.
Area
(mi*)
15,700
400
7,325
6,530
1,059
380
6,000
5,700
300
7,069
1,209
1,575
650
Resident
land species
111
33
55
50
34
27
46
46
34
46
27
32
13
Passeri
nonpassi
0.77
0.94
1.12
0.92
127
0.93
1.09
1.09
157
1.30
1.46
052
0.44
Marquesas
Tuamotu Archipelago
Austral Is.
Pitcaim Is.
Easter Island
Area Resident Passerines:
(mi9) land species nonpasserines
480 11 037
330 7 0.40
115 3
18 3 030
50 0 0.00
Northeast from New Guinea, the Admiralties
(and probably New Britain, for which I lack a
compilation) have a very low ratio. The effect
spreads eastward through the Solomon group,
which, as New Britain, has "strictly an impover-
ished Papuan fauna, except for a few endemics of
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
0 . U . l n i . l m . 5Q0
PHILIPPINES
FIGURE 16.?Philippine Islands: passerine-nonpasserine ratios
of native land birds (numbers of species in parentheses).
uncertain relationship" (Mayr, 1944a: 189). The
Solomons also have the low ratios of a faunal satel-
lite (Figures 15, 17). On an equal-area basis,
individual Solomon islands have very much lower
species totals and ratios than do the Philippines
far to the northwest but not much lower than the
Moluccas on the opposite side of New Guinea. The
Solomons have even lower ratios individually than
as a group, indicating that here, as in other archi-
pelagos, forested mountain conditions are more
conducive to passerine than to nonpasserine differ-
entiation. The larger islands generally have larger
avifaunas, but some smaller ones, such as Rendova
and especially tiny Gizo, have many more species
than their sizes might warrant. A similar example
is to be found in the Galapagos, where minute
Jervis has nine species of permanently resident
Darwin's Finches while islands hundreds of times
larger have at most ten (Bowman, 1961:20). Other
such instances, once brought to view (Figure 10),
might then be explained from the vantage of
hindsight.
Rennell Island, though separated by deep water,
appears to belong with the Solomons on a map
(Figures 15, 17). Unlike the Solomons, it has been
populated from several directions, more or less in
agreement with the prevailing winds (Mayr,
1931). It is similar to the Santa Cruz Islands not
only ecologically but also in area, species number,
and ratio.
East of the Solomons, low ratio is still evident in
the Santa Cruz Islands, with avifaunal elements
from the New Hebrides, Fijis, and Solomons (Mayr,
1945:201). Species and ratio keep approximate
pace with decreasing area, at least until the num-
bers become very small.
Resident Passerines:
land species nonpasserines
Santa Cruz Is.
Ndemi
Vanikoro
Utupua
Reef Is.
Duff Is.
Swallow Group
Tikopia
Area
(mi*)
380
215
100
50
27
16
18
15
12
8
6
4
0.93
1.29
0.80
0.67
030
0.33
050
1.00
Fiji, Samoa, New Caledonia, and the New He-
brides contain the principal large islands east of
the Solomons. Fiji and Samoa constitute the avi-
faunal core of Oceania and (in view of the distance
from the nearest continent) have a comparatively
rich, typically Polynesian bird life; both consist of
several old, big, mountainous islands and many
small, mostly coralline, impoverished islands lack-
ing distinctive endemic elements (Mayr, 1941c:
205). The accompanying tabulation contrasts avi-
faunal size and passerine differentiation in the
high islands and in the small or low islands (see
Figure 15).
Fiji Islands
VitiLevu
VanuaLevu
Taveuni
Kandavu
Ovalau
Lau Archipelago
Tonga Archipelago
Samoa
Savaii
Upolu
Tutuila
Manua Is.
Area
(mi*)
7069
4053
2128
166
165
43
45
250
1209
703
430
52
22
Resident Passerines:
land species nonpasserines
46
39
35
37
32
29
22
14
27
24
23
12
11
1.30
159
130
1.31
1.25
0.93
0.69
0.75
1.46
1.40
1.30
0.71
057
NUMBER 212 45
Nissan (23)
Bougainville (77)
V . Mono (15)
? .67*
+, Vella Lavella(52)i Santa Isabel (60)
.71
Malaita(57)
.54
Pavuvu(24)**
.41
Guadalcanal (83)
.66
miles 200
kilometers _200
SOLOMON ISLANDS
* San Cristobal (62)
.69
'Santa Ana (25)
.67
Rennell (33)
3 4
FIGURE 17.?Solomon Islands: passcrine-nonpasserine ratios of native land birds
(numbers of spedes in parentheses).
High, oceanic New Caledonia (Figure 18), the
largest island east of New Guinea and the richest
east of the Solomons, has a relatively high species
count that may be due to the comparative vicinity
of Australia and to an equally strong Papuan
influence; long isolation is indicated by the high
endemism, about one species in three, but "the
number of endemic genera of birds is amazingly
small" (Mayr, 1941c:209; 1945:149). The below-
unity ratio of New Caledonia is "normal" for an
isolated oceanic island.
The Loyalty Islands ought to have at least as
low a ratio as New Caledonia, provided they are
indeed satellites (Figure 18). They are located,
however, on a different, probably younger island
arc, have a weakened Australian element and show
a stronger Papuo-Melanesian influence, share only
four of the New Caledonian endemics, and have
received a number of New Hebridean strays that
might otherwise have reached New Caledonia
(Mayr, 1941c:210). In relation to area, environ-
mental complexity, and species number, the three
Loyalties seem to vary unpredictably, as do the
Banks Islands at the opposite end of the New
Hebrides. In both these groups, sharp reductions
in area are accompanied by equally sharp drops
in ratio.
The New Hebrides as a whole have a close-to-
unity ratio of 1.09 based on 22 nonpasserines and
24 passerines. Individually the islands have an
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ostensibly perplexing range of values. Ratios in
particular but also species numbers present a dis-
tributional pattern of sorts (Figure 18). Excluding
the two largest islands (Espiritu Santo and Male-
kula), avifaunas become impoverished with in-
creasing distance from the center of the arc, ratios
decrease most markedly through the islands trail-
ing southward into completely open ocean.
Approximate
Torres Islands
Banks Islands
Valua (Saddle)
Bligh
Vanua Lava
Gaua (Santa Maria)
area (mi1)
?
10
?
100
125
New Hebrides (excluding
Espiritu Santo and
Malekula)
Aurora (Maewo)
Oba
Pentecost
Ambrym
Epi
Efate
Erromanga
Tanna
Aneiteum
115
100
200
375
150
350
315
250
50
Resident
land species
10
21
22
29
31
24
31
28
32
34
37
33
29
27
Passerines:
nonpasserines
0.67
0.91
1.20
1.23
1.39
1.40
1.21
1.33
1.67
1.13
1.06
0.94
0.71
0.83
The three southernmost islands lack most of the
New Hebridean endemic species and have a distri-
bution indicating accidental dispersal owing to age
(considerably younger than New Caledonia and
Espiritu Santo); also "on Tanna (and perhaps on
Erromanga and Aneityum) volcanic eruptions have
exterminated part of the indigenous fauna . . .
[while] destructive hurricanes are another factor
that should not be ruled out altogether, particu-
larly for smaller islands" (Mayr, 1941c:211). One
might add that the destructive effects would prob-
ably be greater on the passerines than on the
larger, more robust nonpasserines. The two large
islands of Espiritu Santo and Malekula are situ-
ated off to one side from the others. Espiritu San-
to has the richest avifauna, the most endemics,
and a relatively low ratio (21 passerines and 19
nonpasserines). Malekula has, for its size, a low
species total and a much lower ratio than any
nearby island, as if it stood apart from the main-
stream of interisland colonization.
French Polynesia: French Polynesia (based on
Bruner, 1972) has in toto 32 resident land species;
its principal components have the following totals:
? T : . Banks Is. (34) 127
V a a a l ^ ^ - V a w c i ) ^
GauGl) 1.39^ : ""??-?.
l
' "> Espiritu Santo(40) ' '?"??, (
/? -... L ~ \ M I V A .Aurora )^'; \
K 121 I Pentecost (28)
\ ...-...,. ,
 r V *
l>33
*-..???" \ ^ ^ ^Ambrym (32)
Malekula (31 )^ !? l 6 7
,82
1.13 NEW HEBRIDES (46)
Efate(37) ^
1.06
4Erromanga (33)
-e , -
W
/ * \ W Tanna (29)
\ NEW CALEDONIA AREA(55) \ W . 7 |
? '"-. ? Aneiteum (27)
.83
\
New"Ca^!nja(50) 1 3 8 ^ _
f?W M?fi(20)%
miles 200
11000
0 kilometers 200
NEW HEBRIDES-NEW CALEDONIA
FIGURE 18.?New Hebrides and New Caledonia: passerine-
nonpasserine ratios of native land birds (numbers of species
in parentheses).
Society Islands, 13; Marquesas, 11; Tuamotu Archi-
pelago, 7; the Austral Islands, 3. Compared to
island groups less distant from Australo-Papua,
French Polynesia, judged by the foregoing figures,
has a somewhat larger avifauna than expected,
whereas its four component groups each have a
smaller avifauna than expected, at least on the
high islands. Visual inspection of the figures gives
rise to the suspicion that representative forms may
have been raised to specific rank. In other insular
groups, the aggregate number of species obtained
by summing the components' totals is very much
larger than the number in a checklist of the insular
group as a whole. Here, in contrast, the aggregate
total of the four components is hardly larger than
that of French Polynesia as a whole. The suspicion
is bolstered by Mayr (1941c: 209), who regarded
the avifauna of eastern Polynesia as exceedingly
poor, with a high degree of slight, and a very low
NUMBER 212 47
degree of strong, endemism, and with little dif-
ference between the component groups.
Micronesia: Micronesia has few resident land
birds and generally high passerine-nonpasserine
ratios. The Palaus, on the near side of the Caro-
lines least distant from a faunal source, have the
most species (10 nonpasserines and 12 passerines)
and a birdlife and ratio similar to the Philippines'
(Figure 15). Ponape, on the far side of the Caro-
lines, with three fewer nonpasserines and the same
number of passerines, has a much higher ratio
(1.72). The three remaining Caroline entries
(Yap, Truk, Kusaie) and four in the southern
Marianas (Saipan, Tinian, Rota, Guam) are very
small and support only 9 to 12 species each. All
the above-named are high islands.
The Carolines are strongly dominated by passer-
ines, the Marianas less strongly. Individually the
islands have avifaunas that are perhaps too small
for their ratios to be meaningful. Nevertheless the
high ratios of Micronesia as a whole and of each
of its component groups suggest that a suitable set
of explanatory circumstances probably does exist
and waits to be found.
OTHER ISLANDS AND ARCHIPELAGOS.?Following
Darlington (1957:480), this category consists of
"single islands or isolated archipelagos not falling
into either of the preceding categories, but diverse
otherwise, from large islands like Madagascar to
the smallest oceanic ones": Celebes (previously
included under fringing archipelagos); Madagas-
car, Mascarenes, Seychelles; New Caledonia (pre-
viously included under fringing archipelagos),
Lord Howe, Norfolk; New Zealand; Hawaiian
Islands; Galapagos; Atlantic islands; Antarctica
(lacking land birds). Their existing faunas have all
been derived overseas.
Indian Ocean: The islands in the Malagasy
region?Madagascar, the Comoros, and the out-
lying Aldabras, Seychelles, and Mascarenes?and
Christmas Island, near Indonesia, for the most part
have native land avifaunas with approximately
equal numbers of passerines and nonpasserines. All
have the total land bird ratio lower than the native
land bird ratio, suggesting that in this part of the
world the islands are reached more easily or more
often by more kinds of nonpasserines than of pas-
serines, despite the larger species pools and greater
numbers of individuals of the latter.
Madagascar, one of the world's great islands,
dating from the Secondary and possessing a variety
of climates, altitudes, and environments, has a
smaller avifauna than on mainland areas or even
on continental islands a tiny fraction of its size,
while passerines comprise a much smaller propor-
tion than on the mainland and include hardly
any nonendemics (Moreau, 1966, chapt. 17). That
Madagascar is sufficiently old and distant to have
an "impoverished" avifauna and a reduced number
of passerines is in keeping with its below-unity
ratio of 0.94, which is considerably smaller than
on comparable mainland but typical of oceanic
islands in general. "The most likely explanation,"
according to Moreau, "is that there have been
widespread extinctions, not only of the sub-oscines
. . . but also of the 'modern' birds, especially the
passerines," the cause of which must have been
climatic.
Moreau pointed to three features of the Mada-
gascan bird fauna. "The first is its poverty and
unbalance compared to African areas." But these
are the very characteristics one associates with
chance dispersal from the mainland. Further, Mo-
reau noted that over one-third of the passerines
belong to two families and all but one to endemic
genera. Elsewhere, however, this would be a cer-
tain sign of local radiation in an insular environ-
ment. "The second is the low average number of
species per genus, which, compared with the
African figures, suggests that in Madagascar more
than twice as many species as now exist there may
have recently been lost." An alternative explana-
tion, relating reduction in number of species per
genus to reduction in number of niches (Slud,
1960:141), could also apply here. "Thirdly, the
bird species in Madagascar are so imperfectly
segregated into each of the main habitats . . . that
a considerable proportion of them occupy both
forest and also one or both of the drier zones . . . a
degree of adaptability extremely rare in Africa
. . . and it is difficult to imagine how it can have
come about in Madagascar except by such casual-
ties in the bird fauna of one or other of the habi-
tats that numerous niches became vacant." Yet
Moreau's reference to the "obvious broadening of
niche . . . occupied by the same species in Mada-
gascar or in Africa" can be interpreted to mean
that there are fewer niches on the island than on
the mainland.
The remaining islands are the Comoros, with 46
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
species and a ratio of 1.19 for the group (or 0.97
and 1.04 for forested Gran Comoro and Moheli,
0.62 and 0.59 for deforested Anjouan and May-
otte); the Aldabras, with only eight passerines and
six nonpasserines; the Seychelles, with eight pas-
serines and seven nonpasserines; the Mascarenes,
with seven species each of passerines and non-
passerines on Reunion and on Mauritius, two
passerines and four nonpasserines on Rodriguez;
and Christmas Island, with two passerines and six
nonpasserines.
Lord Howe Island and Norfolk Island: "Lord
Howe is a small volcanic island about 300 miles
east of Australia and about twice as far south (and
a little west) of New Caledonia, 5 square miles in
area and nearly 3000 feet in greatest altitude"
(Darlington, 1957:525). It has a native land bird
total of eight passerines and four nonpasserines or,
excluding recent extinctions, three passerines and
two nonpasserines. Of the land and fresh-water
species, "two or three seem to have come from
Australia, three from New Caledonia, and one
from New Zealand, and the geographical origin
of the rest is doubtful" (Darlington, 1957:525).
"Norfolk Island is a small volcanic island about
800 miles east of Australia and midway between
New Caledonia and New Zealand, about 13 square
miles in area, and 1000 feet high" (Darlington,
1957:526). It has a list of nine passerines and five
nonpasserines, though two of the latter are now
extinct.
It would appear that Lord Howe and Norfolk,
both of them high nontropical islands and with
lists of almost equal length, are in position to re-
ceive colonists from at least three directions. That
they are dominated by passerines is due in part to
the sympatric presence on Lord Howe and Norfolk,
respectively, of two and three species of Zosterops,
almost certainly the result of separate colonizations
(Lack, 1947:150). With respect to total land birds,
at least on Lord Howe, where some data are
available (Hindwood, 1940), the ratio is close to
unity (11 passerines and 12 nonpasserines), as in
the Indian Ocean.
New Zealand: New Zealand is located about
1000 miles east-southeast of Australia; the two
main islands have a combined area of more than
100,000 square miles; the islands are old, with high
mountains, and are ecologically diverse within
the limits of their temperate climate; the existing
vertebrate fauna appears to have accumulated
gradually across wide water gaps (Darlington,
1957:526-527).
The two main islands have a combined list of
only 20 passerines and 16 nonpasserines (based on
Fleming et al., 1953). The number of nonpasser-
ines, however, considerably exceeds that of the
passerines if recent species of moas are included:
"The living birds were well known to the Maoris;
and extinction became complete only within the
last few centuries" (Thomson, 1964:477). As on
little Lord Howe and Norfolk islands to the west,
the total land bird ratio on New Zealand, exclud-
ing moas, is much lower than the resident land
bird ratio: nonresident nonpasserines outnumber
nonresident passerines by about two to one.
Cool New Zealand and warm Madagascar, with
the oldest vertebrate island faunas (Darlington,
1957:536), both have avifaunas and passerine-
nonpasserine ratios that are greatly reduced from
those on comparable mainland; they receive ex-
tremely few potential colonizers; and both have
given rise, especially on more isolated New Zealand,
to a variety of flightless birds, including spectacu-
larly gigantic ones.
Hawaiian Islands and Galapagos: Adaptive radia-
tion from a single ancestor into a number of re-
lated species on oceanic archipelagos has taken
place most dramatically in these two groups, prob-
ably because newer, more efficient colonists have
been arriving too infrequently to eliminate the
earlier inhabitants; similar considerations probably
account for the absence of local adaptive radiations
among the land birds of archipelagos off Europe,
Africa, America, or even in Polynesia (Lack, 1947:
150-151).
On the forested Hawaiian Islands, at least 2000
miles away from North America, 28 of 35 native
birds (80 percent) have evolved from three propa-
gules (a drepanid, a meliphagid, and a turdid),
all passerines and presumably the first successful
land birds to have arrived (see Darlington, 1957:
528); only two species are nonpasserines. On the
dry Galapagos, two-thirds the area of the Hawai-
ians and one-third their closest distance from the
American mainland, some 13 of 24 native land
species (54 percent) have evolved in this manner,
again from an ancestral passerine (a fringillid)
that presumably was the first land bird to success-
fully colonize the islands; only five of the land
NUMBER 212 49
Azores (Portugal)
Madeira Is. (Morocco)
Canary Is. (Spanish Sahara)
Cape Verde Is. (Gambia "Valley") 1557
Closest Resident
Area distance land Passerines: Residents:
(mi2) Latitude to coast birds nonpasserines migrants
888 38? 800 mi. 14 1.80 (1.61) 0.42
302 33? 400 23 1.30 (1.46) 0.32
2807 28? 60 49 0.96 (156) 0.77
16? 350 25 0.92 (1.06) 1.79
birds are nonpasserines.
If one could hypothesize a geographical situation
which prohibited specific radiation on these islands,
the Hawaiians would have five passerines and two
nonpasserines, the Galapagos seven passerines
and five nonpasserines. As to total land birds, the
Hawaiians as a whole would have either ten pas-
serines and eight nonpasserines or, excluding the
westernmost atolls, five passerines and eight non-
passerines (based on Berger, 1972); the Galapagos
would have 11 passerines and eight nonpasserines.
It is paradoxical, also difficult to explain, why six
nonresident nonpasserines and no nonresident
passerines have been recorded from the main Ha-
waiian chain (the State of Hawaii) when the
residents are so heavily passerine.
Atlantic Islands: The North Atlantic groups?
Azores, Madeiras, Canaries, and Cape Verdes?lie
at different distances off southern Europe and
western Africa. (See above.)
Their passerine-nonpasserine ratios, as on the
mainland, decrease with latitude from well above
unity on the Azores and Madeira Islands in cooler
waters to below-unity on the Canary and Cape
Verde islands in warmer waters. The ratio of resi-
dents to nonresidents, increasing equatorward (as
on the island groups off eastern Asia), remains low
on the Azores and Madeiras, mounts on the sub-
tropical Canaries, and rises high above unity on
the tropical Cape Verdes. The Canaries have two
to over three times the number of species on the
other groups, but they are also the closest to shore.
The two innermost Canary Islands are dry and have
passerine-nonpasserine ratios under 0.90, the five
other main islands are much higher, more humid,
and have ratios ranging upwards of 1.20 (based
on Moreau, 1966:360).
The four tropical Gulf of Guinea islands "lie on
a straight line, diverging somewhat from the main
north-and-south direction of the African coast"
(Moreau, 1966:318). In the following tabulation,
the numbers of resident land species and passerine-
nonpasserine ratios are based on Moreau, 1966;
the other entries are based on Amadon, 1953.
Fernando Prin- Sao Anno-
Po cipe Tome" ban
Area (mi1) 800 50 400 7
Distance from mainland (mi) 20 140 175 210
Distance from preceding
island (mi) - 140 90 115
Number of species 140 23 32 6
Percent endemic species 1 20 38 33
Passerine-nonpasserine ratio 2.11 1.30 129 030
Percent passerines on
Fernando Po 100 33 23 25
Percent nonpasserines on
Fernando Po 100 76 48 50
Fernando Po is the largest, highest, and by far
the wettest of the four. Situated on the coastal
shelf and of recent continental origin, it has a
well-balanced avifauna, three-quarters of which
are typically forest birds; it bears a very close fau-
nistic resemblance to Cameroon Mountain and its
immediately surrounding lowlands only 20 miles
away; though it has only one endemic species,
fully one-quarter of the species show some diver-
gence from the mainland populations, a propor-
tion close to that shown by montane species on
Cameroon Mountain compared to other montane
populations in the Cameroons (Moreau, 1966:320).
The high passerine-nonpasserine ratio unquestion-
ably is a reflection of and is probably reduced from
that on Cameroon Mountain as a whole (for
which I have no list).
The other three islands stretch some 300 miles
southward of Fernando Po and cross the equator.
Of oceanic origin, they have been populated over-
seas, probably in larger part from the mainland
than from one another. Taking into account their
different sizes and positions, they follow a trailing
sequence of greatly reduced species numbers and
sharply decreasing passerine-nonpasserine ratios.
The extent to which the species that reached these
islands are the same as those on Fernando Po,
particularly among the nonpasserines (see accora-
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
panying tabulation), is suggestive of the degree to
which these are good colonizers in general or have
successfully occupied similar habitats on these is-
lands in particular. Principe and Sao Tom? (based
on the island lists of Amadon, 1953) today share
only 12 out of a total of 50 species of resident land
birds, yet have similar passerine-nonpasserine ratios.
In the tropical South Atlantic, Ascension and
St. Helena have no known members of strictly land
bird families.
The three islands of the south-temperate Tristan
da Cunha group, with an area of 52 square miles
and nearly 2000 miles distant from the tip of Africa
and farther from South America, have four resi-
dent passerines and no nonpasserines. With respect
to adaptive radiation and passerine predominance,
"Tristan da Cunha is Galapagos in miniature. At
the other extreme . . . the Hawaiian Islands are
as remote in the Pacific as is Tristan da Cunha in
the Atlantic, and only five passerine forms have
succeeded in reaching them" (Lack, 1947:152),
compared to the original two or perhaps three that
reached Tristan da Cunha.
SUMMARY
Division of the Class Aves into two elements, the
passerines and the nonpasserines, has enabled vari-
ous authors to utilize the reciprocal relation of one
to the other toward distributional, environmental,
or evolutionary ends. The interrelationship is here
justified on objective grounds.
Over the terrestrial globe, the species of passer-
ines outnumber those of nonpasserines two to one.
The ratio remains about the same for the zoogeo-
graphic regions: slightly lower in the tropics,
slightly higher outside the tropics. Continued sub-
division into smaller and increasingly homogene-
ous units expands the ratio range from very high
above to far below unity (1:1). The passerines and
the nonpasserines both have species densities that
are considerably higher in the total land avifaunas
and very much higher in the native land avifaunas
of tropical units than of comparable extratropical
units. Climatovegetational factors can explain
much of the difference in faunal size that occurs
latitudinally and biogeographically.
Migrants of course account for the difference in
size between the total land bird and native land
bird categories. Their percentage of the avifauna
shows a parallel decrease equatorward along
either side of the Atlantic and Pacific oceans in
inverse proportion to the size of the avifauna.
Wintering migrants hardly add to the residents in
tropical "rain" forest but occur in greatest abun-
dance under brokenly forested and disturbed
conditions.
In the tropics, the passerine-nonpasserine ratio
rises jointly with altitude from the lowlands to the
temperate highlands. An avifauna under forested
conditions has a higher proportion of passerines
than one under nonforested or mixed conditions
in the lowlands or in the highlands. A highland
avifauna under forested conditions has a higher
passerine proportion than a neighboring lowland
avifauna under forested conditions. For purposes
of faunal or environmental comparison, well-
known small units are preferable to necessarily
less well-known large units. A list of species actually
recorded at a locality reflects the environment more
faithfully than does an amalgamated list from a
large area.
Islands are each unique and conform to no
universal standard. Two islands the same in size,
age, physiography, and climate will have different
constellations of taxa, numbers of species, and
vegetational physiognomies to the degree that they
are inaccessible to potential colonizers; an impov-
erished avifauna is surely the sign of an impover-
ished biota. Comparative judgments must relate
ultimately to standards set on the mainland.
The Suboscine-Oscine Relationship
The Passeriformes, also known as the passerines
or perching birds, contains 5168 species or two-
thirds of the world's recent land birds (based on
Austin, 1971). Almost four-fifths belong to a single
suborder, the oscines or true songbirds. The several
remaining suborders comprise the antithetically
designated suboscines and contain approximately
1100 species. About 50 species of suboscines are
known from the Old World, the rest are American
and are mostly confined to the Neotropical region.
The complementary nature of the neotropical
suboscine-oscine relationship had been suggested
by Slud (1960), largely on the comparative basis
of compilations from the distributional literature.
Since then new faunal works have appeared which
treat several major units for the first time and bring
several others up to date.
NUMBER 212 51
MEXICO
BRITISH HONDURAS
GUATEMALA
HONDURAS
66 ))100
FIGURE 19.?Neotropical suboscines: percent of the spedes of
passerines per country.
NEOTROPICAL DISTRIBUTION
GENERAL.?The nearly 1050 New World sub-
oscines are equally divided between the strictly
neotropical Furnarii, including woodhewers, oven-
birds, antbirds, and tapaculos, and the more wide-
spread Tyranni, principally cotingas, manakins,
and tyrant-flycatchers. Suboscines amount to more
than half the species of native passerines or total
passerines on the neotropical mainland (i.e., south
of the Isthmus of Tehuantepec), three-fifths of the
species in South America, two-fifths of the species
in Central America, and between one-quarter and
one-fifth of the species in Mexico.
The suboscine proportion of the passerines
among the individual Latin American countries,
both with and without extralimital migrants, is
shown in Figure 19. Suboscines are dominant over
oscines throughout mainland South America, ex-
cept perhaps in deforested Uruguay, and keep
approximately the same proportion in Colombia
in the north as in Chile in the south, despite the
sevenfold difference in number of species. Percent-
agewise the suboscines decline linearly from north-
ern South America northward through Middle
America, except for displacement toward lower
percentage in "dry" El Salvador on the Pacific side
and toward higher percentage in the British Hon-
duran lowland on the Caribbean side. If El Salva-
dor were united with adjoining Honduras, and
British Honduras with adjoining Guatemala, so as
to cross-section the Central American isthmus and
thus embrace a wider range of environments, as do
Panama, Costa Rica, and Nicaragua, there would
be no displacement. Mexico's suboscine percent-
ages would not fall nearly so low if the country
were represented only by its neotropical portion,
that lying south of the Isthmus of Tehuantepec.
Figure 20 compares absolute numbers of sub-
oscines and oscines, shows their relative magni-
tudes, and relates them to suboscine-oscine ratios.
Further, it illustrates the relative effect of the
presence or absence of migrants on numbers of
species and on suboscine-oscine ratios. South-
western United States and adjoining Mexico have
been included to emphasize the suboscine decline
northward into North America. The Middle
American units and the South American units are
separated by, and face one another across, the
suboscine-oscine line of unity (1:1).
BRAZIL.?Brazil, the country with the largest
extent of lowland "rain" forest, has the largest
number of suboscines and the highest ratio of sub-
oscines to oscines. As with passerines and non-
passerines, the individual states and territories,
excluding those in Amazonia (Figure 21a), have
a wider range of suboscine-oscine ratios than do
the country's principal geographic sectors, exclud-
ing Amazonia (Figure 21&). Whether plotted by
individual state and territory or divided into
halves, Amazonia lies in a suboscine direction well
away from the other units. Throughout the "rain"-
forested Amazon drainage, suboscines outnumber
oscines up to two to one, locally up to three to
one (Table 6), in species and comprise two-fifths
or more of the entire land avifauna.
The suboscine-oscine relationship within the
Amazonian sector of Brazil is presented in Table 6.
Only those "areas" and places are included that
have a minimum of 150 recorded species of pas-
serines. All have suboscine-oscine ratios which
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
600
550
500
450
400
350
300
250
200
150
? TOTAL LAND PASSERINES
O NATIVE LAND PASSERINES
ILOMBIA
50 100 150 200 250
OSCINES (NO. OF SPECIES)
300 350 400
FIGURE 20.?Neotropical passerines (including southwestern United States): number of species
of suboscines and oscines. (Radiating lines = subostine-osdne ratios.)
NUMBER 212
?5 -xn ?
1 1
? ? TOTAL SPECIES
O NATIVE SPECIES
_
1 1 1 1
A M A Z O N A S ^
PARA
> M A 0 PAULO
MNAS 6ERAIS / ^
BAHIA^
PARANA So*
MARANNAO \ S^ RIO DE JANEIRO
AMAPAl^ \ f ESPIRITO SANTO
ACREo. V ^ * " * " *
STA. CATARINA ??"pufui *KRNAMIUCO
y S ? ? A L A G O A S
yr ? MRAIIA
/ CEARA
X 1 1 1 1
'RIO GRANDE DO SUL
1 1
1
/
1
MATO GROSM-RONDONIA
1
?
a
i
100 125
OSCINES (NO. OF SPECIES)
? TOTAL SPECIES
O NATIVE SPECIES
I I l? * I
WESTERN AMAZONIA
(AMAZONAS-ACRE-
EASTERN AMAZONIA o ? ? RORAMA)
(PARA-AMAM-MARANHAO)
SOUTHWEST (MAT0 6R0SS0-R0NDONIA)
EAST-CCNTRAL (BAHIA-SOIAS)
SOUTHEAn (SAOPAUIO).
"EAST MWASGERAIS-
ESHRTTO STO.-RIO DE JANEIRO)
SOUTH 9ARANA-STA.CATARINA-
RIO GRANDE 0 0 SOD
NORTHEAST PIAUI-CEARA-PARAIIA-
KRNAMIUC0-ALA60AS)
b
100 125
OSCINES (NO. OF SPECIES)
FICURE 21.?Numbers of species of subosdnes and osdnes in Brazil: a, per state and territory;
b, per sector. (Diagonal = suboscine-oscine ratio of 1.4.)
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
exceed that of the sector as a whole (1.85), except
northeastern Par A (1.49), at the mouth of the
Amazon, and the Manaus-Itacoatiara area (1.46,
1.70), along the juncture of the Rio Negro and the
Rio Madeira with the Amazon. In northeastern
Para, where there is an admixture of species from
the dry northeast of Brazil and the climate and
vegetation appear to be broadly transitional (see
Weber, in Fittkau et al., 1969:476-479, maps),
the reduced suboscine proportion forms part of an
eastward-spreading decrease that seems to originate
in the neighborhood of the right bank of the
lower Tapajos (2.12) or possibly nearer the Xingu
(2.15). At Manaus-Itacoatiara the drop in ratio is
abrupt, as though the area were isolated from the
surrounding high levels. The great width of the
Rio Negro at its confluence with the Amazon can-
not be invoked here as a distributional barrier to
birds. Igarape1 Cacau Pereira (not included in
Table 6), situated on the opposite side of the Rio
Negro from Manaus, has total and native ratios of
1.42 and 1.43 (based on 133 passerines, including
two migrants) that are virtually identical to the 1.46
and 1.48 of Manaus. The area around Manaus, it
appears, is one of bleached sandy soil associated
with a distinctive type of rain forest (Richards, in
Meggers et al., 1973:21).
COLOMBIA.?More diverse environments should
show greater avifaunal differences than those dis-
cernible in the relatively uniform Amazon basin.
In the accompanying tabulation, the principal
faunal sectors are the same ones previously em-
ployed for passerines and nonpasserines in Co-
lombia; suboscine-oscine ratios are followed by
numbers of species of passerines in parentheses
(compiled from Meyer de Schauensee, 1964).
Pacific
Caribbean
Central Mountain
Orinocan
Amazonian
Total passerines
1.11
0.79
0.91
1.38
1.80
(234)
(178)
(578)
(311)
(342)
Native passerines
1.31
0.94
0.96
159
1.97
(204)
(155)
(537)
(280)
(315)
Suboscines predominate in the humid lowlands of
Colombia, whether Amazonian, Orinocan, or Pa-
cific, but fall below unity on the dry Caribbean
versant and in the mountains. As was the case with
passerine-nonpasserine ratios, the total and native
suboscine-oscine ratios of 0.91 and 0.96, respec-
tively, for the "Central Mountain" sector appear
to be a good representative average: the corre-
sponding figures for Cundinamarca are 0.88 and
0.94. The 0.79 and 0.94 of the Caribbean sector
agree with the 0.82 and 0.98 of Atlantico. The
ratios of 1.80 and 1.97 of the Amazonian sector are
virtually duplicated by the 1.80 and 1.92 of Ca-
queta and the 1.85 and 1.97 of Amazonian Brazil.
The 1.38 and 1.59 of the Orinocan sector are ap-
proached by the 1.33 and 1.49 of Meta. The 1.11
and 1.31 of the declivitous Pacific sector compare,
respectively, less well with the 1.29 for total pas-
serines but extremely well with the 1.38 for native
passerines of a collection from Guapi, on the coast,
reported by Olivares (1957-1958).
OTHER AREAS.?The few adequate data available
indicate that as a general rule the suboscine pro-
portion decreases with increasing elevation (Table
7). Exceptions, i.e., lowland areas with low sub-
oscine proportions, are the interior valleys of east-
ern Guatemala, Caribbean Colombia, northeastern
Venezuela, southwestern Ecuador, the department
of Lima in coastal Peru, and southwestern Peru.
These units are doubly distinguishable, on the one
hand by the fact that they are climatically dry,
on the other hand by their small avifaunas, which,
together with low ratio, produce a combination
found elsewhere only at high altitudes.
The relatively high suboscine proportion for
Peru, a typically Andean country, is explainable
by the fact that it possesses extensive Amazonian
lowland teeming with suboscines. The relatively
low ratio for the Apurimac valley in eastern Peru
is in itself sufficient reason to suspect a subtropical
influence, the actual presence of which is borne
out by the valley's description as "a broad finger
of Amazonian lowland that penetrates 150 miles
into the Andes between two high ranges" (Ter-
borgh and Weske, 1969:765). Also in Peru, the
Urubamba valley passes through tropical Amazonia
and would be expected to have higher ratios than
shown (Table 7), but the low ones are in accord
with its having been collected most heavily at its
middle and upper reaches (Chapman, 1921:11).
COSTA RICA.?Altitude: Whether or not a cli-
matic pattern emerges from the distribution of the
suboscine-oscine ratios of the same "areas" and
localities in Costa Rica previously compared for
their passerine-nonpasserine ratios can be deter-
mined from Table 3. Strictures to be kept in mind
are that some places have been visited more often
or at more different times of year than others, that
NUMBER 212 55
for many places the avifaunal representation has
passed little beyond the stage of a sample, and that
the major components of the native avifauna are
far less affected proportionally by local movements
of native species than are those of the total avi-
fauna by the occurrence of extralimital migrants.
Native ratios indicate in the main that the sub-
oscine proportion does decrease from one set of
localities with rise to another set in the next higher
belt. The humid tropical entries all have ratios
above unity, except Los Chiles and La Lola, both
of which have been extensively deforested. Two
other areas, Boruca-Paso Real on the Pacific slope
and Sixaola on the Caribbean slope, each have a
close-to-unity ratio of 1.03. (The two also have iden-
tical passerine-nonpasserine ratios.) The Boruca-
Paso Real area is located in edaphically poor, cut-
over savanna-type country; the Sixaola area has
been heavily deforested by commercial interests and
includes old records that probably apply to the sub-
tropical belt. Their low ratios would appear to ally
the above-mentioned localities to those in the
Tropical Dry Life Zone, but they are immediately
separable from the latter by their considerably
longer species lists and high passerine-nonpasserine
ratios.
The subtropical entries (Table 3) fall below
unity, with two exceptions: Juan Vinas-Tucurriqui
on the Caribbean slope and Las Mellizas-Cot6n
on the Pacific slope. I combined lists for Juan
Vinas and Tucurriqui, which I had compiled from
the literature (primarily Carriker, 1910), in hope
of achieving a better representation than separate
treatment would furnish. Either the combined list
continues to be unrepresentative or the Reventa-
zon valley, in which the two localities lie, affords
suboscine-dominated birds from the tropical belt
unimpeded access to the interior. In the case of
Las Mellizas-Coton, the data appear to reflect the
climatic regime governing the humid Pacific
southwest, "where the suboscine proportion re-
mains at approximately the same level from the
Osa Site [Rinc6n de Osa] upward to the Rio
Coton Site. On the Caribbean side the suboscines
suffer approximately a 40 percent drop in relative
proportions between the lowlands and the equiva-
lent altitude of the Rio Cot6n Site" (Slud, 1965:
117). Still, I place little confidence in this list,
which consists of a wet-season sample from Las
Mellizas combined with a dry-season sample from
Cot6n. Cot6n, judged by its physiognomy, assort-
ment of birds, and climate, has no Caribbean
counterpart. At higher elevations, the suboscine-
oscine ratios in the lower-montane and montane
belts plunge to half their value or less in the sub-
tropical belt.
Forest versus Nonforest: How the suboscine-
oscine relationship fares at a site under forested
and under unforested conditions cannot be deter-
mined reliably, comprehensively, or comparatively
from the literature. Some information is available
for Costa Rica, based almost entirely on what I
considered to be a forest bird at the time and place
of my visits (Table 5). The impression that the
suboscine proportion increases inside woods is
amply substantiated, with three apparent excep-
tions. One, the Bagaces site, is perhaps an "abnor-
mal" example as it was very small, very disturbed,
and depauperate (Slud, 1965:6). Second, the Rio
Suab locality, with only four nonforest passerines
and 56 forest-inhabiting passerines, was poorly
censused, as was the third, the San Jos? de la
Montana site; neither would remain an exception
if it had one more suboscine.
Forest-inhabiting avifaunas have suboscine-
oscine ratios that tend to increase with increasing
elevation (Table 5). Whether for climatic or
edaphic reasons, as the case may be, humid-forested
tropical-belt localities have above-unity forest ratios
(except La Lola and Los Chiles, both extensively
deforested and poorly represented by forest
species). Subtropical localities with a Caribbean
climate, viz. trade-wind influenced and a very
short effective dry season, have forest ratios rang-
ing downward from unity. Apparent exceptions
are Hacienda Santa Maria and Silencio de
Tilaran, both of which, however, occupy low gaps
in the continental divide and thus are subject to
faunal overlap from the Caribbean and the Pacific
lowlands on either side. In the subtropical belt
in the Pacific southwest, unlike the situation at
analogous elevations along the Caribbean slope,
humid-forest ratios tend to remain at least as high
as in the lowlands, apparently in correlation with
the more marked seasonality of the Pacific slope in
general. At still higher altitudes, localities for
which data are available are few. These are virtu-
ally restricted to the Caribbean slope and the
Caribbean-influenced central mountains, and their
ratios are distinctly low. An exception, La Hon-
56
dura, situated on a long, steep slope, would surely
have its ratio fall below unity if its list did not
include species more properly known from lower,
warmer elevations.
SAMPLE VERSUS COMPILATION
Table 4 compares suboscine-oscine ratios of com-
pilations with those of samples for countries and
localities. Countries, such as Honduras and Costa
Rica, but also the Canal Zone territory, apparently
have a large enough area and a sufficiently well-
known birdlife for the relative proportions of the
avifaunal components to have become fixed some
time ago. The localities have suboscine-oscine
ratios that, unlike their passerine-nonpasserine
ratios, vary little between compilation and sample
or, more often, differ increasingly as the difference
in size between the compilation and the sample
increases. The only group of related localities in
which the ratio decreases from sample to compila-
tion is confined to, and apparently characterizes,
the Tropical Dry sector.
Barro Colorado Island and Finca La Selva, the
best known of the localities, still have not reached
suboscine-oscine stability (Table 4). Barro Colo-
rado, in particular, shows a progressive decrease in
ratio that correlates roughly with increase in num-
ber of passerines. The decrease is due in part to
foreign and native visitants, the majority of which
are oscine. As a result, the growth of the oscine
proportion has increasingly outstripped that of the
suboscine proportion:
Total suboscines
Native suboscines
1970
1952
1938
1929
84 = 1.15
73
78 = 1.11
70
67 = 1.05
64
61 = 1.05
58
Total oscines
Native oscines
101 = 1.68
60
85 = 1.57
54
71 = 1.54
46
58 = 1.45
40
Total passerines
Native passerines
185 = 1.39
133
163 = 1.31
124
138 = 1.25
110
119 = 1.21
98
Finca La Selva, with a much shorter history, has
suboscine-oscine ratios that have changed little
(Table 4). However, the sequence of periodic in-
crease in numbers of suboscines, oscines, and pas-
serines is similar at Barro Colorado Island (above)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
and Finca La Selva (below) and the sequence of
ratios of total to native suboscines, oscines, and
passerines, respectively, is practically identical:
Total suboscines
Native suboscines
1970 89 =
80
1960 88 =
79
1957 69 =
66
1.11
1.11
1.05
Total oscines
Native oscines
110 = 1.62
68
105 = 1.64
64
84 = 1.47
57
SUMMARY
Total passerines
Native passerines
199 = 1.35
148
193 = 1.35
143
153 = 1.24
123
The suboscines decline from three-fifths of the
native passerines in South America to one-fifth in
Mexico. Brazil, more than any other country, and
Amazonia, more than any other area, have the most
species of suboscines and the highest suboscine-
oscine ratios. Suboscines not only outnumber
oscines or nonpasserines in the "rain"-forested
Amazon basin, but they are also very successful
under analogous climatovegetational conditions in
other tropical American lowlands, from Central
America to western Ecuador and southeastern
Brazil.
Broadly viewed, suboscine-oscine ratios in the
neotropics are lower in the dry than in the humid
life zones of the tropical belt on the one hand, and
higher in the lowlands than in the highlands on
the other hand. Two further types of ecologically
based ratio are obtainable. One type is derived
from the sum of reports for an entire belt or life
zone. Altitudinally to tree line, the compiled data
become progressively poorer in quantity and qual-
ity, yielding ratios that decrease irregularly or may
even level out in the highlands. Thus qualified,
suboscine-oscine ratios correlate inversely with ris-
ing elevation upon mountain slopes facing the
lowlands, not upon isolated hills or ridges. The
ratio tends to be higher than expected in inter-
montane situations, as in the central highlands of
Colombia or Costa Rica, where the climatic and
topographic conditions are complex and the avail-
able data imprecise. The other type of ratio is
derived from a sample or from a sum of samples
for a particular locality or "area." Individual locali-
ties in Costa Rica have suboscine-oscine ratios that
vary inversely with rise from one altitudinal, or
NUMBER 212 57
thermal, belt to the next, at least on the Caribbean
versant. On the Pacific versant, the pattern is only
weakly distinguishable, perhaps as a consequence
of the markedly seasonal climate, not only here
but possibly in monsoonal climatic regimes in
general. Other ratios which are lower than ex-
pected pertain to places that at present are exten-
sively unforested when climatically they ought to
be forested.
Countries or other sizeable units with large avi-
faunas that are represented by lists compiled a
generation apart show little change in suboscine-
oscine ratio from one to the other. Few localities
or sites possess information, adequate or not, that
permits their being compared against themselves
chronologically. The well-worked ones are well
along the road to suboscine-oscine stability, and the
way from sample to checklist is marked by decreas-
ing ratio, at least in humid localities. At all magni-
tudes of faunal or areal size, suboscines increase
at the expense of the oscines in the forested por-
tion of an environment compared to the un-
forested portions. The increase becomes relatively
greater as the climate becomes warmer and wetter.
Possibly the suboscines will yet be found to show a
decrease under conditions that are warm but
increasingly dry.
The Passerine-Nonpasserine
Suboscine-Oscine Relationship
COMPARATIVE CATEGORIES
Just as passerines and nonpasserines comprise an
avifauna, suboscines and oscines complement each
other to comprise the neotropical passerines. Con-
verted into percentages or ratios and cross-plotted,
the two relationships produce a graph or scatter-
gram permitting measurement and visual apprecia-
tion of the degrees of similarity or dissimilarity
among avifaunas. An intraregional comparison
might involve such major areas as Middle America,
northwestern South America, eastern South Amer-
ica (Brazil), and southern South America (Figure
22). At a lesser scale of magnitude, a strictly politi-
cal comparison would involve all the neotropical
countries, regardless of size (Figure 23). Similarly,
political comparisons within a major area, such as
Brazil, would involve all the states and territories,
again regardless of size.
The treatment becomes increasingly effective
ecologically, rather than geographically, when
applied locally to points on the map, whether these
are scattered over different political units (Figure
27) or confined to a single political unit (Figures
28 and 29). Potentially the most useful subdivision
is that created by parceling a locality into homoge-
neous environments. Comparisons could then be
made of similar habitats at different places or dis-
similar habitats at the same place. The factors
limiting further refinement are those which so
reduce a list of names that upon replication a
small change in the number of species causes dis-
proportionately large changes in the percentages
or ratios.
SUBREGIONAL SECTORS.?Four sorts of lists for
each of the major areas?Middle America, north-
western South America, Brazil, southern South
America?are plotted in Figure 21. The lists sepa-
rate into two sets, one with total avifauna and
resident avifauna closely paired, the other set with
total land avifauna and native land avifauna closely
paired. Taking into account the prominent water-
bird element in southern South America, the lists
bear an overall similarity to one another despite
their considerable neotropical spread. The straight-
ness of the vertical alignment is due to the very
small difference in passerine percentages (1.2 per-
cent in the case of total passerines).
COUNTRIES.?In Figure 23, the unit of compari-
son is the individual Latin American countries
(exclusive of the Antilles, Figure 30). The units
are represented by their total land avifaunas in
part a and by their native land avifaunas in part
6. Changes in position of the units in the two parts
of the figure are an indication of the relative size
of their migrant proportions. In the Central Ameri-
can units, the distribution of the native land avi-
faunas, compared to that of the total land avifau-
nas, shows an equatorward shift toward the high
suboscine proportions of the South American units,
which remain stationary (except temperate-zone
Chile and Uruguay), and a shift toward higher
nonpasserine proportions in a direction away from
those of the South American units, which again
remain nearly stationary (except Chile).
Brazil lies closer to the Andean countries than
to the Guianas (Figure 23). If, however, Brazil
were sectioned geographically, its principal sectors
would help fill the gap between Ecuador, Peru,
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
30
35
40
w 45
50
? B 6
60
65
55
% NONPASSERINES/ AVIFAUNA
45 40
? Brazil
? Middle America
^ Southern South America
A Northwestern South America
Total Avifauna
Resident Avifauna
? Total Land Avifauna
? - ? Native Land Avifauna
30 70
65
60
55
50
45 v>
40
35
45 50 55 60
% PASSERINES/ AVIFAUNA
65 70
FIGURE 22.?Relative positions of four neotropical subregions: passerine (or nonpasserine)
percentage of avifauna versus suboscine (or oscine) percentage of passerines.
Bolivia, and Paraguay on one side and the Guianas
on the other side. The individual states and terri-
tories, especially the drier ones or those with little
forest, would be dispersed more widely than the
sectors, because a smaller unit is less heterogeneous
than the larger unit to which it belongs. In prin-
ciple, further subdivision into a countrywide
array of small units would cause them to more fully
occupy a still wider expanse. Graphed, a small
unit tends to lie to one side of the larger unit of
which it is part; the less it is known and the more
unrepresentative is its list, the farther the small
unit should lie from the large unit.
Figure 23 bears a distorted likeness to Middle
America and to a condensed South America, the
outline of which would be more complete if Brazil
were divided into its country-size sectors. The simi-
larity may not be entirely accidental, provided
one is impressed by the correct alignment of Mid-
dle American units, including the pairing of
southern Mexico with Guatemala, Honduras with
Nicaragua, and Costa Rica with Panama; the loca-
tion of the Yucatan Peninsula "offshore" from
the neighboring mainland; the sequence of Andean
NUMBER 212 59
% N O N P A S S E R I N E S / A V I F A U N A
35 40 45 50
15
20
25
3 0 -
3 5 -
45
50
o 55-
6 0 -
5 5 -
5 0 -
45
?Nontropical Mexico
Tropica. Mexico x
 6UATEMALA_BR> H0NDURAS
Guatemala* \ \ucatan Peninsula
HONDURAS-EL SALVADOR
Honduras
?Trinidad
COLOMBIA
VENEZUELA'
.PERU
^ B O L I V I A
^ N * ARGENTINA
/ "
J .
65 60 55
% P A S S E R I N E S / A V I F A U N A
50
85
80
65
60
55
50
45 f
40
40
1 5 -
2 0 -
3 0 -
3 5 -
4 0 -
4 5 -
Z 50
o 55
6 5 -
60
5 5 -
5 0 -
- 55
% N O N P A S S E R I N E S / A V I F A U N A
35 40 45 SO
Nontropical Mexico
?jTropiC3l Mexico
*EI Salvador
HONDURAS-EL SALVADOR
NICARAGUA
?Trinidad
COLOMBIA,
VENEZUEL
PERU
N^BOLIVIA
TMtQENTINA
65 60 55 50
% P A S S E R I N E S / A V I F A U N A
- 4 5
? 35
FIGURE 23.?Scattergram of neotropical countries based on avifaunal components (suboscine or
oscine percentage of passerines versus passerine or nonpasserine percentage of avifauna; horizon-
tal dashed line = equatorial transition between northern and southern South America): a, total
land birds; b, native land birds.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
countries on the west; the location of the Guianas
on the opposite side of the continent, with Guyana
followed at a distance by Surinam and French
Guiana, closely paired and in proper order; the
placement of Brazil closer to its Andean neighbors,
with which it shares the Amazonian watershed,
than to the Guianas, which occupy a different
versant.
The suboscine-oscine ratio of 65:35 (approxi-
mately 2:1) was chosen arbitrarily to represent
the line of division between the northern and
southern hemispheres in Figure 23. A strict latitu-
dinal accounting would, of course, require that
southeastern Colombia, most of Ecuador, all of
Peru, and the greater part of Brazil, including most
of the Amazon and Rio Negro drainages, be trans-
posed equivalent distances south of the equator.
A way to circumvent the difficulty is to devise some
sort of representative single figure and place it
directly on a map (Figures 24-26).
SINGLE-FIGURE VALUES
The components contributing to the land bird
passerine-nonpasserine ratio, designated (a), and
to the suboscine-oscine ratio, designated (b), can
be interrelated directly and reciprocally in several
ways. In my opinion, the simple (a): (b) relation-
ship furnishes a single-figure value that is not only
the best representative average of a unit, whatever
its magnitude, but is also the most usefully compre-
hensible for comparative purposes (Table 8 and
Figure 24; also Tables 3-5). At low elevations, the
lowest (a): (b) values to be found anywhere in the
neotropics occur in the Amazon valley (except
the Manaus-Itacoatiara area) and the next lowest
in the Guianas and along the "rain"-forested
coastal strip of eastern Brazil. The highest (a): (b)
values occur in arid areas?northeastern Venezuela,
southwestern Ecuador, coastal Peru?and also in
Uruguay (which is largely unforested and where
the resident status of a number of species is not
clear). The (a): (b) value increases with increas-
ing elevation, reaching an extreme in the high
cordilleras, and with increasing latitude, particu-
larly in Middle America.
A list which one has reason to suspect is deficient
usually yields an (a): (b) value that looks exagger-
ated or "wrong." A value that looks "wrong"
should make one wonder about the representative-
ness of a list which had not previously come under
suspicion. (Incidentally, "wrong" (a): (b) values
can be intensified to stand out all the more when
(a) and (b) are multiplied.) If, in place of ratios,
the passerine (or nonpasserine) percentage of the
land avifauna is termed (a) and the suboscine (or
oscine) percentage of the passerines is termed
(b), the (a): (b) values now tend to emphasize
the similarities instead of the dissimilarities among
units. Either way, one is provided with a measure
of humid tropicality that relates to units of known
area and to quantifiable climatic parameters.
The (a): (b) interaction apparently has the
potential of bringing a degree of order into the
distribution of similar avifaunas. For example,
Vuilleumier (1970) analyzed 15 paramo "islands"
in northwestern South America. An unpublished
appendix supplied by the author enabled me to
total 56 land species for one "island," 25 to 32
species (average 28.8) for each of five "islands,"
and 11 to 17 species (average 13.6) for each re-
maining "island" (disregarding one "island" with
only four species): a virtually perfect geometric
progression. Table 9 lists the "islands" individually,
followed by five discrete physiographic groupings
into which the "islands" separate according to the
contours on the author's map. The groupings re-
veal an exponential (a): (b) increase from central
Ecuador to central Colombia, thence to eastern
Colombia, then to western Venezuela and to Santa
Marta in northern Colombia. This pattern would
not have come to light if reliance had been placed
solely on correlation coefficients. Ratios for high
Andean Ecuador, Colombia, and Venezuela are
presented in the following tabulation.
Ecuador
Colombia
Venezuela
Passerines:
nonpasserines
(a)
1.00
1.28
2.00
Suboscmes:
oscines
(b)
1.55
1.29
1.00
(a): (b)
0.65
1.00
2.00
The discrepancy, noted earlier, between the par-
amo zones of Colombia and Venezuela might now
be regarded instead as support for an "island"
sequence in the high mountains of northwestern
South America.
Table 9 goes on to contrast the correlation co-
efficients of the 15 "islands" with those of the five
groups. The coefficients of the 15 "islands" are
fair to good for species and distance and for species
NUMBER 212 61
FIGURE 24.?South American native land birds: ratio of passerines-nonpasserines to suboscines-
oscines. (Numbers in large type = countries; numbers in smaller type = political subdivisions.)
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 25.?Native suboscine-oscine ratios in South America. (Numbers in boxes = countries;
underlined numbers = political subdivisions; unmarked numbers = smaller areas or general
localities.)
NUMBER 212 63
033 ? _
FIGURE 26.?Ratios of species of Formicariidae to those of native oscines in South America.
(Numbers in boxes = countries; underlined numbers = political subdivisions; unmarked
numbers = smaller areas or general localities.)
64
and area. The coefficients of the five groups are
extraordinarily high in these respects. In addition,
they range from significantly high to very high in
categories where those of the 15 "islands" correlate
poorly or not at all. Provided it is statistically
permissible to do so, a reduction from five to four
groups, owing to the removal of isolated, non-
Andean Santa Marta, would result in even
higher correlations, most strikingly with passerine-
nonpasserine ratios and suboscine-oscine ratios.
The (a): (b) single-figure value would appear
to have advantages that are best suited to site-
oriented situations. Unfortunately, it is often the
local list that engenders doubts as to its reliability
and comprehensiveness. A telltale sign is the pres-
ence of few birds of prey, a deficiency which
indicates in turn that the entire nonpasserine com-
ponent may be too small. Yet the same lists can
have good representations of suboscines and oscines.
Elimination of the nonpasserines, therefore, actu-
ally increases their accuracy and usefulness. Indeed,
suboscine-oscine ratios (Figure 25) provide single-
figure comparative values that may be only little
less representative of the distribution and affinities
of South American land avifaunas than are the
(a): (b) values. Perhaps this should have been
expected from a "bird continent" that is also a
suboscine stronghold.
A single-figure example of more restricted utility
is furnished by the Formicariidae, or antbirds. The
distribution of this large suboscine family, from
southern Mexico to northern Argentina (Figure
26; Table 10), coincides with that of frost-free
broadleaf woodland. Centered in Upper Amazonia,
where the species of antbirds per locality number
three-fifths to nine-tenths those of all the oscines,
the family acts as a comparative climatovegeta-
tional indicator of biotic richness in the New
World tropics, exclusive of the Antilles (all of
which lack this sedentary family). Since the main-
land distribution of the Formicariidae, or of any
other selected group of "rain"-forest families, is
ecological rather than political, comparisons be-
tween localities should be based upon environ-
mental data, but few are available. In any event,
one ought to be wary of relying on a single group,
whether a large family or several "characteristic"
small families, as representative of an avifauna.
For instance, Costa Rica and Nicaragua approach
each other in species of antbirds but are separated
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
by a gap between their total numbers of suboscines.
ENVIRONMENTAL CLUSTERS
Altitudinal and latitudinal relations that result
from cross-plotting passerine-nonpasserine and
suboscine-oscine ratios are visualized in Figure 27.
Avifaunas having similar relative proportions form
a cluster, irrespective of shared or unshared taxa
and of geographical proximity or distance. The
assortment of entries, even though based on what
was available rather than on true areal, faunal, or
climatic comparability, is poor. Notwithstanding,
the entries separate into two sets of environmental
clusters, one set for the highlands, the other for the
lowlands, in both of which the member groups
succeed one another latitudinally. The dry low-
lands in South America for which data are
available?Caribbean Colombia, northeastern Vene-
zuela, southwestern Ecuador, and coastal Peru?
would, if entered in Figure 27, form an "out-of-
place" cluster of their own in the area already
occupied by the humid Central American lowlands,
hence were omitted for the sake of clarity. But
even if they had been included, the drastic reduc-
tion in numbers of species in the dry climates
could be invoked as a third dimension that would
keep them spatially distinct. An analogous concept
is the "complexity index" of Holdridge (Holdridge
et al., 1971:545-546) in which the climatic plant
associations can have the same numerical value in
different life zones lying along different tempera-
ture belts or in different humidity provinces.
"Areas" and localities within a single political
unit are compared in Figures 28 and 29. Costa Rica
was selected, because it is the unit for which I have
the most information. The entries differ in size,
complexity, range of elevation and climate, degree
of disturbance, and extent to which they are
known. A few consist of neighboring localities with
inadequate lists, which I combined in hope of
obtaining a more representative list of the local
avifauna. Sometimes this procedure produces the
opposite effect, but the alternative was to utilize
lists that are too skimpy or not use them at all. In
any event, the entries that are close to one another
geographically but not climatically lie separated to
the extent that differences in climate are reflected
by changes in proportions of the avifaunal com-
ponents.
NUMBER 212 65
10
15
20
25
30
?/>
- 3 5
- 4 0
UJ
> 45
- 5 0
55
60
65
70
75
25
?/o N O N P A S S E R I N E S / A V I F A U N A
30 35 40 45 50
90
85
80
75
70
Northern
Soloma.
MEXICAN HIGHLANDS
\
ntral
Southern
GUATEMALAN NIGNLANDS\_^Cerro de la Muerte-Villa Mills NaGulf ?
lowlands
x Wetmore. I94L
COSTA RICAN HIGHLANDS \ / N ^ P o a s Volcano (upper)
Sierra las Minas
Poas Volcano (n. w. slope)
.Upper Urubamba, PERU San Antonio
Upper Auyan-tepui Upper Chiriquf PANAMANIAN!
^ ? Highlands ? ? HIGNLANDS~~*Cerro Campana
COLOMBIAN HIGHLANDS
Cent. Gulf
? lowlands
Yucatan Peninsula
60 m
? Mt. Roraima
VENEZUELAN HIGHLANDS
Cerro Neblina
Mt.Duida?
CUNOINAMARCA
N.e.NiCARAGUA ^fc.fertral Awrka
^GUATEMALA
S.e.NICARAGUA ^ \
HONDURAS^
So. COSTA RICA
(Highland BOLIVIA
AMAPA
?W.PANAMA\ LOWLANK
?.COSTARICA I . _ _ . , . .(CanalZone \St.CMtnl AMnca
E. PANAMA
region
No. SURINAM
Bartica. GUYANA
FR. GUIANA
Lr.Tocantins* . A C R E > ^
/ * N a BRAZIL-So.VENEZUELA
/MiddleMadeira. #s.e.COLOMBIA
Lr Xincu * ^ ^Rio Purus f
RioJurua* 'UTapajos
AMAZONIAH LOWLANDS
Lr. Madeira
I
50
45
40
30
25
75 70 65 60
? / ? P A S S E R I N E S / A V I F A U N A
55 50
FIGURE 27.?Latitudinal and altitudinal distribution of Central American and northern South
American areas and localities based on percentages of major avifaunal components.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Figure 28 is restricted to Costa Rican localities
with a typically Caribbean, almost constantly
humid, climatic regime. These form into groups
the placement of which corresponds to the altitu-
dinal zonation of the temperature belts. Not to
be passed over is the apparently misplaced trio of
entries consisting of Angostura (at an elevation of
600 m) and Los Chiles and La Lola (in the low-
lands). These agree in having little primary
woodland coupled with a reduced proportion of
suboscines from that normally present at tropical-
belt elevations. Another entry, Hacienda Altamira,
situated in the northern lowlands, also ought to
occur lower in the tropical belt. It is only known,
however, from a one-week sample. Further work
would almost surely bring its suboscine proportion
up to expectation, provided the remaining forest
has not been destroyed in the meantime.
Figure 29 plots localities on the more seasonal,
partially monsoonal, Pacific slope of Costa Rica.
These form into clusters that are disposed more or
less horizontally, unlike the vertical arrangement
on the Caribbean side. This pattern on the Pacific
side suggests that the avifauna may be affected
more by rainfall than by temperature. In the
northern, seasonally arid sector of the slope, two
entries, Miravalles and Rinc6n de la Vieja, lie
above the others- Actually, both these "areas"
range from the climatically dry lowlands upward
into the Caribbean-influenced humid subtropical
belt along the facing continental divide. In the
southern, humid sector of the slope, the transition
from tropical to subtropical is rapid and short in
terms of suboscine or oscine percentage. Inter-
mediately placed entries occupy an area the nature
of which may not be so much transitional as con-
fused, owing to seasonal effects having been
blended into an annual average in some instances,
while the majority are represented each by a single
sample.
The fact that environmentally similar units form
discrete clusters can be utilized to evaluate the
degree to which a list may or may not be represent-
ative of an avifauna. For example, reports on gen-
eral collections, such as those of Dearborn in Gua-
temala and of Sassi or Ferry in Costa Rica (Table
4), yield ratios that favor the climatovegetational
zones and altered countrysides in which the authors
spent most of their time. If plotted, these reports
would lie outside the cluster of native land bird
ratios formed by the checklists that are available
for either country, whether in the direction of
higher passerine percentages or of lower suboscine
percentages. The (a): (b) values further emphasize
the extent of the gap between the checklists and
these reports, especially the one by Ferry, who did
not work in the lowlands. Another example has
recently come to hand (Table 4), thanks to Burt L.
Monroe, Jr., who sent me the results of a midwinter
tour he led in Costa Rica. The itinerary allowed
little time in the humid tropical belt compared to
that spent in the higher or drier parts of the
country. This is reflected in the increased passerine
proportion, the decreased suboscine proportion,
and the very high (a): (b) value.
In like manner, it is conceivable that a measure
of the environmental changes caused by man can
be obtained from the numerical changes in the
avifaunal values, especially in small units such as
localities or sites, through a chronological sequence
of checklists or of representative samples.
SUMMARY
Cross-plotted, the passerine-nonpasserine and
suboscine-oscine relationships produce a geographi-
cal or ecological "map" showing the comparative
distribution of neotropical avifaunas. Any faunal
level can be accommodated, from the subregional
down to the parceling of habitats in a study site.
At the scale of magnitude represented by the indi-
vidual Latin American countries, the avifaunal
"map" takes on the appearance of a political map
of the region, with the line of division between the
northern and southern hemispheres corresponding
to a suboscine-oscine ratio that peaks at about 2:1
in the Amazon basin. At a lesser scale of magnitude,
local avifaunas having similar proportions of pas-
serines (or nonpasserines) and of suboscines (or
oscines), irrespective of faunal resemblance or
geographical distance, form into discrete ecological
or climatic assemblages or clusters.
When the passerine-nonpasserine ratio (a) is
combined with the suboscine-oscine ratio (b) into
a simple (a): (b) relationship, the result is a single-
figure value that can be put to comparative use.
Placed directly on a map, it permits one to see
whether the unit it represents follows certain
trends or fits into a general pattern. From a low
in the humid equatorial lowlands, the (a): (b)
NUMBER 212 67
45
25
30
- 3 5
\
- 4 0
45
50
55
N O N P A S S E R I N E S / A V I F A U N A
4 0 3 5 3 0 25
> ? < ? I i i i i I i 1?1 r
? VILU MILLS-CERRO DE LA MUERTE
POAS VOLCANO (UPPER) MONTANE
LOWER MONTANE
SUBTROPICAL -
LOWER MONTANE
? AZAHAR-ESTRELLA DE CARTAGO
. SAN JOSE DE LA MONTANA
LA CHONTA*
? Bajos del Toro
SUBTROPICAL
? LA HONDURA
? MONTEVERDE DE TILARAN
.Silencio de Tilaran
? Pavones ridge Virgen*
del Socorro ?
?BONILLA ? CAR I BLANCO
LOS C H I L E S . .
LA LOLA* Hda. Altamira
.CARRILLO
?GUAYABO-AQUIARES
.SIXAOLA
FINCA LA SELVAU972)
? F I N C A U SELVAU960)
i i . I I I
? FINCA LA SELVAU957)
?OLD LINE
?between Siquirres and
Turrialba
TROPICAL
75
70
65
6 0 "
55
50
-45
55 60 65 70
" a P A S S E R I N E S / A V I F A U N A
75
FIGURE 28.?Altitudinal disposition of areas and localities on the Caribbean slope of CosU
Rica based on percentages of major avifaunal components.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
55
% N O N P A S S E R I N E S / A V I F A U N A
50 45 40 35 30.
2 5 -
3 0 -
35
N
.40
? VILLA MILLS-CERRO DE LA MUERTE
? 0 N T A N E
75
L O W E I M O N T A N E
LA CHONTA?"
70
65
\
60 -o
?MONTEVEROE DE TILARAN
.SAN VITO Silencio de Tilaran
55
5 0 -
Rincon de la Vie ja . .
MIRAVALLES. /
TROPICAL /
TO SUBTROPICAL '
HDA. SANTA /
MARIA AREA .
PELON DE LA ALTURA . .
 B A R R A N C A SECTOR ? , HELECHALES " B O R U C A * T i n a m a 5tas de San Isidro 50
TROPICAL TO
SUBTROPICAL
TROPICAL BELT
? PALO VERDE
GENERAL VALLEY
SUBTROPICAL
Mellizas-Coton
Sta. Teresa
de San Isidro
LR. COTO BRUS, . vijagual de Buenos Aires
BUENOS AIRES*
45
? RINCON DEOSA TROPICAL BELT
? VOLCAN DE OSA
SUBHUMID NORTHWEST
I ? ?
H U M I D S O U T H W E S T
? POZO AZUL-POZO PITAL
-I 1 1 1 1 1 1 1 1 1 I ? ' ' I ?45 50 55 60
% P A S S E R I N E S / A V I F A U N A
65 70
FICURE 29.?Latitudinal and altitudinal disposition of areas and localities on the Pacific slope
of Costa Rica based on percentages of major avifaunal components.
AY
 
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70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
value increases altitudinally, latitudinally, and, at
least in South America, horizontally in the dry
tropics. The (a): (b) value can be used to best
advantage in site-oriented situations.
Requirements for Further Research
SAMPLING
Checklists versus Samples
The lists employed in this report have proven
useful in providing a basis upon which to made
bioclimatic generalizations, but they also point to
the need of something better. Long-term compila-
tions, or checklists, no matter how comprehensive,
cannot tell us the species composition of a unit at
any given time past or present. A checklist for a
large unit incorporates seasonal and ndnregular
taxa scattered over a wide range of conditions;
one for a small unit has fewer nonresident taxa
and a limited environmental range that can be
defined by the local microclimate or by the domi-
nant vegetation.
A checklist for a small unit, that is, a point on
the map, such as a locality or a study site, occurs in
the literature rarely compared to one for a large
unit, such as a country. The few localities that do
occur have been so unequally investigated that
their lists range in status from that of a single col-
lection to that of a long-term compilation. Parti-
cularly in the neotropics, few places are known
to have a reliably comprehensive list, even fewer
have been delimited areally and altitudinally or
are truly representative of a natural life area or
habitat type on the mainland. To help determine
the status of a local list on a comparative basis,
another list from the same place is needed. Rarely
if ever is a replication available for the place in
question or even for its general vicinity. Indeed,
no tropical American field station to my knowledge
ever inventories its biotic stock.
On the other hand, a sample from a small area,
such as a locality or site, is considerably more re-
strictive as it tends to include few nonregular taxa
and to group them seasonally. Moreover, a short-
term survey not only yields a sample of what is
actually present but it can only census a small area
adequately. Further, the small area, i.e., the locality
or site, is manageable in the sense that it can be
worked at will to provide samples at different
times in the same year or at the same time in dif-
ferent years. Sample can be compared against sam-
ple from the same place or from different places.
Thus the elaboration of a bioclimatic scheme by
which to plot the comparative distribution of neo-
tropical avifaunas comes to depend upon a supply
of representative points in much the same manner
as a climatic map is constructed upon the data
supplied by a network of weather stations. The
fundamental requirement, therefore, is a site-
oriented method that produces equivalent samples
and can easily be put into effect, whether apart
from or in conjunction with general or special
field studies.
Analytical Methods
Prime considerations in making a survey, besides
deciding where to do it, are how to go about it
and knowing when to stop. Ideally, a group of
competent persons employing a variety of data-
gathering means would work cooperatively re-
gardless of cost. In practice an investigator usually
works alone and has limited resources. As to the
size of his area, it ought to have a radius about as
long as a half day's bird-seeking amble, while the
pace at which he proceeds continually adjusts to
the nature of his surroundings, whether closed or
open, broken or level. The unit should not be so
small or unbuffered as to be exposed to internal
contamination, so to speak, by the dissimilar biota
of a neighboring unit.
LENGTH OF SURVEY.?The rate at which addi-
tional species continue to be met, rather than an
arbitrary allocation of time, determines the length
of the survey. One way to do this is to plot the
daily cumulative species totals. The result is a
steeply rising arc which soon declines toward the
horizontal (Figures 31, 32). The straightening of
the curve signifies the onset of unprofitable yield
relative to time invested. Or one can decide to wait
until it takes a few days to record another species.
Instead of continuing this survey it is better to
start a new one and obtain a second sample.
The point of diminishing returns is reached first
with the native land avifauna, last with the total
avifauna (Figure 32). The native land avifauna
has the fewest species; nevertheless, it is the least
variable of the four categories of list. A rapidly
RIO COTON SITE
SAN CARLOS SITE
y VOLCAN SITE (DRY)
/ 'SARAPIQUI SITE SIQUIRRESSITE'' ?I
^VIRGEN DEL SOCORRO SITE
OSASITE
VOLCAN SITE (WET
TAPANTI SITE
BARRANCA SITE (WET)
? BAGACES SITE
>BARBA SITE (DRY)
> LA CHONTA SITE
RIO LA HOJA SITE
' BARBA SITE (WET)
3 4 5 6 7
NO. OF DAYS
FIGURE 31.?Cumulative native land birds recorded at Costa Rican localities by P. Slud
in 1964-1965.
acquired sample is preferable to a slowly acquired
one, because of the reduced chances of incorporat-
ing seasonal effects. The better the investigator's
acquaintance with the birds of an area the sooner
the curves approach the asymptote.
Figures 32 and 33 illustrate what should have
been done but was not during a trip I made to
Barro Colorado Island in the Panama Canal
Zone. It was my first visit, yet I had preset a de-
parture date on the assumption that the time
allotted would be ample. During the penultimate
two days I found only one more species, and the
curve appeared well on its way toward apogee.
The upturn produced by four additional species
the final day makes it clear I should have stayed
longer. The same conclusion can be drawn from
some of the Costa Rican localities in Figure 31.
Another way to determine the length of a survey
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
200
190
180
170
160
150
140
130
120
I I I II I I I I I I I I I I I I
10
TOTAL AVIFAUNA
NATIVE LAND NONPASSERIN
TOTAL LAND NONPASSERINES
I I I I I I I I I I I I I I I
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
NO. OF DAYS
FIGURE 32.?Cumulative native land birds recorded at Barro Colorado Island, Panama Canal
Zone, by P. Slud, 16 March to 8 April 1966.
NUMBER 212 73
i 4 4
u
? 42
40
38
36
34
32
30
n
j i i i i i i i i i i i I i i -
TOTAL PASSERINES/
TOTAL LAND AVIFAUNA
NATIVE PASSERINES/
NATIVE LAND AVIFAUNA
NATIVE SUIOSCINES/
NATIVE PASSERINES
NATIVE PASSERINES/
TOTAL LAND AVIFAUNA
TOTAL SUMSCINES/
TOTAL PASSERINES
NATIVE SUIOSCINES/
TOTAL PASSERINES
NATIVE SUSOSCIRES/
NATIVE LAID AVIFAUNA
TOTAL OSCINES /TOTAL
LAND AVIFAUNA
NATIVE OSCINES/TOTAL
PASSERINES
NATIVE SUBOSCMES/
TOTAL LAND AVIFAUNA
TOTAL SUIOSCINES / TOTAL
LAND AVIFAUNA
NATIVE OSCINES /NATIVE
LANS AVIFAUNA
NATIVE OSCINES/TOTAl
LAND AVIFAUNA
9 10 11 12 13 14 IS 16 17 18 19 20 21 22 23
FIGURE 33.?Cumulative daily percentages of avifaunal components recorded at Barro Colorado
Island, Panama Canal Zone, by P. Slud, 16 March to 8 April 1966.
is to convert the daily cumulative totals into their
component percentages of oscines, suboscines, pas-
serines, and nonpasserines. The percentages mostly
level out while the numbers of species are still
climbing (Figures 32, 33). It is a rapid method
which usually becomes effective as soon as a major-
ity of the sample has been gathered. For the
experienced person this happens as a rule early in
the survey. Whether obtained this way or the
other, the sample should be as representative of the
composition and relative size of the avifauna at
the time and place of the survey as is any other
similarly acquired sample at any other time or
place.
In Figures 34 to 37, the abscissa stands for num-
ber of species instead of number of days. Figure
34 cumulatively plots the proportions of the major
components. It also shows the exponential in-
crease in the number of days that are needed to
record in the field each additional equal-size
increment in number of species. The appearance
of the components plotted by percentage (lower
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NATIVE LAND AVIFAUNA (NO. OF DAYS)
2 3 4 . .5 10 IS 20 23
55 -
50
45
7Z 40
S 35
30
2 25
O
20
15
NATIVE LAND BIRDS
? NONPASSERINES
O SUBOSCINES
e OSCINES
36
34
32
30
28
2 6 -
24
2 2 -
20
-J55
50
45
40
35
30
25
20
15
50 60 70 80 90 100 110 120
NATIVE LAND AVIFAUNA (NO. OF SPECIES)
130 140
FIGURE 34.?Cumulative number of species (upper) and cumulative percentages (lower) of avi-
faunal components per day and per cumulative number of species in total avifauna during visit
to Barro Colorado Island, Panama Canal Zone, by P. Slud, 16 March to 8 April 1966.
NUMBER 212 75
65
60
CO
j? 55
?
LU
^ 50
Q.
E
Z
 45
40
35
65
60
55
CO
LU
1 50
om
CO
45
40
35
50
S3NI0S0
35
30
7
j-
'-
7
- ?
7
7
7
7
7
L
 ?
7
-
- ?
1 ..
m
. . i 1 . .
<
?
?
ft
?
?
. . . . . . . 1 . . . .
?
?
?
. . . . 1 . . . . ! . .
?
?
. . i . . . . 1 . . . . i . .
?
-
-
-
-
? -
-
-
..1
65
60
5 5 |
?o
50 jjjj
x
45^1
40
35
65
100 110 130 140 150
TOTAL LAND AVIFAUNA (NO. OF SPECIES)
160 170 180
FIGURE 35.?Average of nine 2-week surveys at Finca La Selva, Costa Rica, made by P. Slud in
1957-1958: cumulative number of species of avifaunal components per cumulative number of
species in total avifauna.
part of Figure 34) complements their appearance
when plotted by days (Figure 32). Taken together
they enable the investigator to view a survey in
progress and to estimate its stage of completion.
That the components were perhaps being en-
countered at an approximately constant rate dur-
ing the above-mentioned reconnaissance I was
making at Barro Colorado Island, Panama, as
suggested by their appearance in Figure 34, is
substantiated by the rigid alignment of the totals
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
when averaged for a number of surveys I had made
at Finca La Selva, Costa Rica (Figure 35).
As a rule the nonpasserine proportion of the
land birds is considerably smaller than the passer-
ine proportion in compilations or checklists gen-
erally, even more so in samples from the humid
tropics. In full-term samples (those with a flattened
cumulative-species curve) the nonpasserines, after
a retarded start, tend to increase faster than do
the other two components, and the rate accelerates
once the majority of the sample has been obtained
(Figures 34, 35). This is the logical pattern to
expect since the nonpasserine component, owing
to the larger size and smaller numbers of its mem-
bers, consists principally of low-frequency species
and includes hard-to-find terrestrial and nocturnal
groups as well as wide-ranging elements and those
most heavily persecuted by man. The nonpasserine
upturn at the end of a graph, when the total
avifauna has been leveling out, indicates that for
practical purposes the survey has come to a close.
It can happen that the cumulative-total curve
which has been gradually approaching the hori-
zontal suddenly shoots upward. An example comes
from the Carnegie Museum catalog of collections
made by Samuel L. Klages and party on 81 days
between 22 December 1920 and 4 May 1921 at
Obidos on the north side of the Amazon River in
western Pard (Table 11). The insurgence began
abruptly the 68th day of collecting. During the
ensuing 13-day period, that is, until the next to last
day in the field, the increment amounted to 66
species (including two migrants). During the pre-
ceding 13-day period it had been only eight species
(none migrant). Obviously the increase was not
caused by an influx of migrants. It is possible that
the skipping of only one collecting day between
the two 13-day periods was enough to permit dis-
mantling and reestablishing camp at a new site
not noted on the labels or to exploit a habitat not
previously visited in 68 days of collecting. (The
Obidos data do not include the Obidos Islands,
treated separately in Figure 36.)
Usually a renewed rise in the cumulative species
total is a sign of incomplete sampling. The per-
centages of the components comprising the Obidos
native land sample are, however, affected slightly
if at all (Table 11): fluctuations exceeding 1.5
percent above or below the average of the remain-
der of the survey do not occur after the 22nd day,
when 55 percent of the sample had been obtained,
for the nonpasserines; after the 32nd day, when
62.5 percent of the sample had been obtained, for
the suboscines; and after the 33rd day, when 63
percent of the sample had been obtained, for the
oscines. The suboscine proportion of the passerines
does not level out until the final 13 days of collect-
ing, after most of the sample had been obtained.
This leveling out is apparent on a daily basis, not
from the 10-day averages in Table 11. The Obidos
sample of 281 native land species yields higher
ratios than does the compiled Obidos total of 354
(Table 8), as is usual when a sample is compared
to a compilation: 2.16 versus 1.70 in the passerine-
nonpasserine ratio (a), 2.49 versus 2.23 in the
suboscine-oscine ratio (b), and 0.87 versus 0.76 in
the (a): (b) value. The degree of difference be-
tween the sample and the compilation can be
magnified by multiplying (a) and (b), the product
of which is 5.38 in the sample and 3.80 in the
compilation.
The Obidos sample calls for further remark. For
one thing, competent observers probably could
have obtained a list in a fraction of the time
needed by the collectors, and the cumulative spe-
cies curve would have started out steeper and
higher and become flattened much sooner. Whether
the observers might have ultimately amassed a
higher total than the collectors is problematical.
Most likely the proportional representation would
have been similar since either method records the
components at a regular rate (Figures 34-36). For
another thing, the Obidos data follow a pattern
that is common to comprehensive samplings. The
dominant component, here the suboscines, is met
most frequently at the beginning of the survey.
Afterwards its proportion of the total either levels
out or undergoes a barely perceptible decline
within a percent or two of the horizontal. En-
counters with the other two components tend to
be relatively infrequent at the start of the survey,
then increase more rapidly at the expense of the
dominant component. The minority components,
here the nonpasserines and the oscines, follow this
trend at Obidos, except when interrupted tempo-
rarily by the late surge already discussed. Graphed,
the nonpasserine and oscine percentages more or
less mirror each other as long as the suboscine
proportion remains fairly constant.
The suboscine-oscine ratio can be particularly
NUMBER 212 77
PASSERINES (NO. OF SPECIES)
7Q 80 go loo n o
 120 130 140 1S0 160 170 180
1
 ' i i i 1 1 1
OBI DOS ISLANDS
. APACY
MANACAPURU
SANTAREM
TONANTINS
HYUTANAHA
OBIDOS
S. PAULO OLIVENCA
147
I3S~ SANTAREM
? 53 -5J
3S * ?
' I I I L. I I 1 1 I I I I I I L
40 50 60 70 80 90 100 110 120 130 140 ISO 160 170 180 190 200 210
PASSERINES (NO. OF SPECIES)
FIGURE 36.?Collecting localities of S. M. Klages in Amazonian Brazil: cumulative number of
subosdnes per cumulative number of passerines. (Slopes determined visually to emphasize con-
stant rate of increase in number of subosdnes. Diameter of dots equivalent to two spedes on
absdssa.)
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
0
0
0
I I I I I I ? i ? I ? ? i I ' ' ' ' I ' ' ' ' I ' '
PASSERINES NO. or SPECIES
fc
1
 
.
 
.
 
.
-
?
-
-
?
" 11
b
1 1 1
?
?
, 1 , , , ,
?
1
?
..1 1 1. t 1
?
, . , , 1
?
' ? 1 ' ' ' ? ?
?
1 1 1 1 1 1 1 1 1
SUBOSCINES NO. OF SPECIES
FIGURE 37.?Average of nine 2-week surveys at Finca La Selva, Costa Rica, made by P. Slud in
1957-1958 (cumulative numbers of species per major component): a, passerines versus nonpasser-
ines (numbers in circles = avifaunal totals); b, suboscines versus oscines (numbers in circles =
passerine totals). (Diameter of circles equivalent to one species on either coordinate.)
NUMBER 212 79
useful distributionally when the representative-
ness of the nonpasserine component is in doubt, if
only because of its unusually small size. This is the
common situation at Amazonian localities, many
of which are known mostly from a single collection.
Whether actually or only apparently "poor" in
nonpasserines, these localities are evidently "rich"
in passerines. The passerines, more abundant and
more easily found and presumably better sampled
than the nonpasserines, are in turn "rich" in sub-
oscines (or "poor" in oscines). There is little rea-
son to doubt that this is the true state of affairs.
Indeed, many species of suboscines can be harder
to find than many of the oscines. It follows that
suboscine-oscine ratios in Amazonia vary much
less than do the passerine-nonpasserine ratios.
Figure 36 cumulatively plots the number of sub-
oscine species versus that of total passerines at
localities in Amazonian Brazil. The localities are
each represented by samples obtained during a
single continuous stay, though not necessarily on
consecutive days. When total passerines reach the
150-species mark, the proportion of suboscines
and oscines is as follows: 97 and 53 at Santar^m,
116 and 34 at Sao Paulo de Oliven^a, 110 and 40
at Hyutanaha, 108 and 42 at Obidos, 109 and 41
at Vila Braga, 104 and 46 at Arima, and approxi-
mately 110 and 40 at Nova Olinda and at Mirituba.
These "rich" samples almost all attain a constant
rate of suboscine increase long before the number
of passerines reaches 100; the smaller, or "poor,"
samples mostly do so before the number of passer-
ines reaches 50. One ought not, however, suppose
a survey has been completed simply because the
suboscines have settled into a regular rate of
increase.
Figure 37a and b plots the passerine-nonpasserine
and suboscine-oscine species averages of a number
of surveys at a locality, Finca La Selva, in Costa
Rica. Ratios are preferred, because they have a
greater range of variation, hence show a finer re-
gard for departures from the norm than do per-
centages (Figure 35). The passerine-nonpasserine
ratio stabilizes by the time approximately two-
thirds of the sample has been obtained, usually
well within a week; the nonpasserine upturn at
the end of the graph signifies the effective end of a
survey. The suboscine-oscine ratio stabilizes even
earlier, when one-half of the sample has been
obtained. Thereafter the rate of increase remains
constant for the suboscines and for the oscines.
Faunal Resemblance
A more conventional analysis of comparative
distribution with ecological overtones is that of
taxonomic faunal resemblance as determined by
formula (see Hagmeier and Stults, 1964; Peters,
1968). Individually the formulas tend to emphasize
similarities, to emphasize dissimilarities, or to
strike a balance between the two tendencies,
though they all produce a similar pattern when
graphed and none eliminates the effects caused by
the different sizes of the faunas being compared.
Whereas a proportional relationship among its
major components is an innate property of a list
which the passerine-nonpasserine suboscine-oscine
interrelationship brings to light, faunal resem-
blance tells us nothing about an individual list
but can only compare it with other lists. From the
faunal-resemblance point of view, comparison is
based on the number of shared taxa relative to
total taxa; two lists can be dissimilar if only be-
cause one has more entries than the other; the fact
that the internal proportions may be similar is
immaterial.
Faunal resemblance formulas and the passerine-
nonpasserine suboscine-oscine interrelationship
both treat the same raw material, i.e., lists of taxa.
Either method can be applied from the level of the
continent or faunal realm down to the habitats at
a site. When the avifaunas to be compared are
taxonomically similar, the two reinforce each other
as they perform much the same function. When the
two avifaunas are taxonomically dissimilar, the
methods interact. Faunal resemblance now becomes
a measure of degree of difference, the passerine-
nonpasserine suboscine-ocine interrelationship con-
tinues to be concerned with relative proportions of
the major components. Now if two lists that are
similar in proportions of components but dissimilar
in taxonomic content should be matched against
each other, a residue of unshared taxa would be
left over in one list that conceivably is equivalent
ecologically to the residue in the other list.
Further, interplay between the two methods
names the taxa and the units, the study of which
could develop insight into such matters as replace-
ment and displacement or convergence and div?r-
gence. The result would be constant improvement
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in the making of quantitative comparisons between
geographically changing faunas. The ability to dis-
cern taxonomic derivatives and ecological counter-
parts could be used qualitatively in helping to plan
a numerical framework for the housing of niches
in the neotropics. Superimposed upon the Old
World tropics, the framework would serve to com-
partmentalize environmental equivalence for un-
related taxa around the globe.
Accuracy of Counts
A twofold difficulty involving taxonomic status
arises, due to the occurrence of migrants and to
the size and location of the faunal unit. With re-
gard to the former, two or more closely related
forms considered conspecies by one author and
separate species by another author can be present
in the same list. How to treat them is an avoidable
issue in short-term samples, few of which ever in-
clude more than one. In long-term compilations the
chances are that all will have been recorded, if only
once. The rule that the smaller area will have
fewer kinds of North American migrants than the
larger area becomes virtually absolute at points
in Amazonia, where few migrants occur in compila-
tions and none in most of the samples. In general,
a summation of samples from one or more sites or
a checklist for a large unit overemphasizes the
proportion of nonregular taxa. The number of
nonregular taxa is minimized in a sample.
For another thing, an increase in area or in
number of distributional barriers fosters geographic
variation and results in subspecies tending increas-
ingly to outnumber species. For instance, Costa
Rica, oversimplified, consists of Caribbean and
Pacific halves. Widespread taxon A, deemed a
monotypic species, would be counted once in a
species list for the Caribbean side, once in a species
list for the Pacific side, and once in a species list
for the entire country. Taxon B, deemed a poly-
typic species with a Caribbean race and a Pacific
race, would be counted once in each of the three
species lists and twice in a subspecies list for the
entire country. Taxon, C, deemed a superspecies
with one Caribbean species and one Pacific species,
would be counted once in a species list for either
slope and twice in a species list for the country.
Species confined to one slope would be counted
once for the slope and once for the country, thus
increasing the disparity between the country and
either slope. Actually, the disparity between the
country and either slope or between the two
slopes is even greater, because some species are
represented by two races on one slope and by one
or none on the opposite slope.
The disparities dissipate, especially on the sub-
specific level, as the units diminish in size down to
a point on the map. If Costa Rica happened to
occupy only one slope, the lists of species and of
subspecies for the country would be the same as
for the slope. A sector of the slope would be ex-
pected to have fewer species and relatively still
fewer subspecies than the slope. A subdivision of
the sector would have both lists further reduced,
and so on, down to the locality or site, at which
point the two lists become identical. Incidentally,
the degree of difference between the corresponding
proportions of any two of the above lists would be
less than that suggested by faunal resemblance
formulas.
Counts are further affected by differences in the
taxonomic outlook of catalogers, particularly when
treating large or complex units inhabited by poly-
typic species. Counts would become more truly
comparable if ecological equivalence were better
understood and if superspecies were no longer
determined intuitively and could be assigned a
binomial for comparative purposes.
Since by definition there can be no allopatry at
a site, at least during the reproductive period, one
need not rely upon intuition to decide the speci-
ficity of closely related birds. Put another way, the
site is the testing ground for the belief that sym-
patric populations do not interbreed and are spe-
cifically distinct in the area of geographic overlap
irrespective of their ecological requirements. The
site is also the place at which to evaluate notions
concerning character displacement and to look for
traits that distinguish sibling species in life. Implicit
is the assumption that in birds the species is the
basic, the only biologically definable, taxonomic
unit. It follows that the number of forms, i.e.,
species and subspecies, at a site is tantamount to
the number of species. The statement, for example,
that "Chapman records 1508 species and subspecies
of birds from Ecuador, while I record 736 from
Guatemala" (Griscom, 1932a:3) is the sort that has
appeared commonly in faunal works, yet it tells
nothing about the number of species in either
NUMBER 212 81
country and would be meaningless for a site. Again
the conclusion is reached that only site-oriented
studies can be truly intercomparable and provide
suitable material for environmental constructs.
CHOICE OF LOCALE
Amazonia
Equatorial Amazonia approaches the theoretic
ideal of a locale in which to begin to compare avi-
faunas. The very vastness virtually guarantees it
will be a heartland of evolutionary dynamism and
ecological radiation, the proliferation of species
that it can supply good-size samples. These two
properties, promising variety and quantity, are in-
dispensable to the investigation of a "rain"-forest
lowland in which the only physical barriers appear
to be the great waterways. A dual objective would
be to determine the extent to which the faunas,
represented by samples, differ taxonomically and
in proportions of major components on opposite
sides of rivers. Another objective would be to
evaluate the extent to which the differences may
have evolved disjunctly in similar environments or
in climatically or vegetationally dissimilar envi-
ronments. Evidence that can be interrelated should
be forthcoming from the vegetation, the birds, and
the weather records.
Biologically complex Amazonia seems to be
macroclimatically simple over much of its basin.
The basic parameters are average annual precipi-
tation, which varies geographically in quantity and
seasonal distribution, and average temperature,
which seems to be fairly uniform annually and
seasonally. Variation in one of the parameters
provides the simplest condition for making envi-
ronmental and faunal comparisons. Responses to
significantly different ranges of rainfall should be
recognizable in the physiognomies of the mature
natural vegetation (perhaps the intermediate
stages, too).
The evolutionary goal of undisturbed vegetation seems to be
the attainment of the most complete utilization of the quali-
ties of the site as possible. Each association is, in effect, a
natural mechanism endeavoring to sustain the greatest possi-
ble amount of living matter per unit area. To do this, the
vegetation adjusts to the given total set of environmental
factors, i.e., both to the general climatic pattern and also to
special conditions existing in the local area. The natural
vegetation anywhere in the world is, in a metaphorical sense,
an organic computer that evaluates all the controlling en-
vironmental conditions in an association, assigns proper
weights to each, and produces an answer in terms of a vege-
tative cover. (Holdridge et al., 1971:16.)
Responses of the vegetation-dependent birds are
secondary or indirect.
The investigator will be confronted by biocli-
matic alternatives in Amazonia. Should average
rainfall prove insufficiently variable geographically
to produce measureable differential effects on the
biota, the entire region could be regarded as a
single unit. Should it prove to be sufficiently vari-
able the responses of the biota might then be
agglomerated into life-zone type units. Should it
be clinal the problem arises whether a climatic
continuum is paralleled by a continuum in the
vegetation and in the composition of the avifauna.
Should it take the form of a step-gradient the op-
portunity arises to test the efficacy of the passerine-
nonpasserine suboscine-oscine method within nar-
row climatic limits.
The primary unit to investigate, the standard
against which to make comparisons, is the "climatic
association" of Holdridge (1947) or the at-times
analogous "climatic climax" of other authors.
Theoretically only one climatic association can
exist anywhere in a natural division of climate, i.e.,
the life zone (plant formation) of Holdridge, from
which the mature vegetations of all the other
associations can deviate almost infinitely in kind
and degree. The climatic association should in
principle be the same wherever found, that is, in
predictable conjunction with quantitative climatic
parameters and be recognizable physiognomically
and by the presence of shared taxa or ecological
counterparts.
A phytoecological reconnaissance of Thailand
by personnel with neotropical experience elicited
the following observations (Holdridge et al.,
1971:684):
The life form, as a morphological expression of the plant's
physiological adaptation to a given habitat, tends to be
similar (but not identical) wherever the same habitat con-
ditions prevail anywhere in the world. This ecological-
evolutionary principle proved true to a remarkable degree
for these widely separated and floristically unique regions.
Uncounted cases arose in which trees, taxonomically unknown
to the observers, possessed such a striking morphological
resemblance to tropical American species with which they
were familiar that they independently recorded their life
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
form in terms of the same American species name. Moreover,
these taxonomic prototypes were observed to occupy equiva-
lent ecological niches, by Life Zone and association, in both
hemispheres.
The Holdridge life-zone system provides equally
weighted natural areas among and within which
to select representative points, not only for tax-
onomic, faunal, and comparative ecological study
but also, ever more urgently, for preservation.
Selections are thus made from a theoretical
global perspective rather than randomly and
opportunistically.
Amazonia need be treated no differently for com-
parative purposes than an area with strong climatic
contrast. By concentrating on representative points
one should be able to reassemble thermopluvial
divisions of the earth's surface on the basis of the
avian samples as well as from the weather data.
The opposite course, which is the one we are usu-
ally compelled to follow, is to obtain the average
of a single master list for a large block of terrain
and try to gauge microclimates from generalizations
about the macroclimate. For the Amazonian
hylea, a comparison of the proportional representa-
tions of the major components over a network of
points should indicate the location of centers of
avian speciation and ecological radiation. This
approach also serves to associate recurrent migrants
with clearly defined environments, whereas cover-
age of a large unit records even a lone migrant as
inhabiting the entire area.
There are several reasons that make it prefer-
able to institute a comparative survey in monoton-
ously immense Amazonia than, say, in complex
little Costa Rica, the other area figuring promi-
nently in this report.
Costa Rica
Costa Rica in an area less than one percent that
of Amazonia possesses a number of altitudinal and
latitudinal life zones (plant formations) so closely
juxtaposed that an observer can quickly accumulate
a long list of birds. The country would appear,
therefore, to be eminently desirable for compara-
tive life-zone treatment on theoretical or practical
grounds. Yet the intermixture of biotic richness,
telescoped physiography, and rapid changes in
climate can produce a perplexing jumble. The
transitional zones, some of relatively great breadth
for such a rugged little country, are so numerous
(see Tosi, 1969) that attempting to preselect a site
as a climatic standard often becomes guesswork.
But even if a representative set of climatic points
were reliably determined, distances remain so
short that a brief flight can carry a bird into a
different environmental context. Vertically a differ-
ence of a few hundred meters in the occurrence of
a species could be interpreted as a dislocation of
the altitudinal belts or an example of avian dis-
tribution not in accord with vegetation or climate.
The difficulty in knowing precisely which species
are resident and which are irregular or seasonal
may not be inconsiderable at such climatic extremes
as the lowlands and the mountain tops. It is very
considerable throughout the intermediate eleva-
tions, where environmental influences are multi-
directional and knowledge of faunal movements
is rudimentary. By contrast, the Amazonian low-
land is essentially free of altitudinal complexities,
and it apparently contains huge, climatically simi-
lar blocks of vegetation: "In regard to the large-
area vegetation types it is possible to define a
series of zoogeographic provinces. Within the vast,
climatically very homogeneous, tropical rain-forest
area several faunal areas can be observed to re-
spond to the degree of the fertility of the soil and
to the geological structure of the region" (Fittkau,
in Fittkau et al., 1972:652). Large size alone should
protect them from other than marginal infiltration
by external biotic elements, unlike the usual situa-
tion in Costa Rica.
Costa Rica, at ten degrees above the equator,
has an avifauna whose reproductive period gener-
ally coincides with that in the temperate north.
"It seems . . . that the majority of the birds in
Central America breed during the second quarter
of the year" and "the peak . . . is March to June or
July, inclusive, with a reduced but still important
amount of nesting activity in August and Septem-
ber, whereas only a small proportion of the birds
have nests from October to February" (Skutch,
1950). Amazonia, on the other hand, straddles the
equator and is analogous to the Congo. "In no part
of the Congo is there any approach to a universal
breeding season, or even a time of year when the
great majority of birds are nesting. The region of
least seasonal change in this respect is unquestion-
ably the equatorial belt of heavy forest, though
NUMBER 212 83
conditions may be equally stable in the mountain
forests" (Chapin, 1932-1954, 65:320). In those
areas, therefore, where breeding activity is spread
over the year, no survey should on this account be
affected more than any other.
In Amazonia, long-distance migrants are few
in kinds and numbers: a list of the total avifauna
hardly differs in length from a list of the native avi-
fauna. In Costa Rica, the two lists match each
other closely during the approximately three
migrant-free months of the year and fluctuate from
little different to considerably different the other
nine months. Percent total migrants, including
water birds, in the two areas are as follows:
Total Avifauna
Nonpasserines
Suboscines
Oscines
Costa Rica
20
23
8
25
Amazonia
5
8
1
7
Restricted to land birds, migrants make up 14 per-
cent of the avifauna in Costa Rica and 2.5 percent
in Amazonia, 8 percent of the nonpasserines in
Costa Rica and 1 percent in Amazonia; suboscines
and oscines remain the same.
At a Costa Rican locality, Finca La Selva, in the
wet Caribbean lowlands, the migrant percentages
over a 12-month period (Slud, unpublished) are
as follows:
Total Avifauna
Nonpasserines
Suboscines
Oscines
12
months
20
9
11
41
Sept.-
Oct.
17
8
10
32
Jan.-
Feb.
10
4
4
21
Mar.-
Apr.
14
6
8
29
June
July
0
0
0
0
Incidentally, I know of no comparable seasonal
information that is available for any other locality
in the neotropics. It can be obtained only through
site-oriented studies.
Finally, transitions between life zones (forma-
tions) should be amenable to analysis, provided
these have not been unduly disturbed. Findings
having to do with the bird-life, even if obtained
as here proposed, would be secondary to, depend-
ent upon, and variously intermediate between the
adjacent climatic standard of the life zone, or
ecosystem, on either side. Transects made horizon-
tally, say, in Amazonia or vertically in Costa Rica
should, if properly planned, pass through a natural
sequence of life zones (see Holdridge, 1947; Hold-
ridge et al., 1971). The transitions ought then be
made recognizable by the occurrence of a small
contingent of indicator species associated with the
next succeeding humidity province or thermal belt,
by the sympatric occurrence of congeneric or closely
related counterparts normally restricted to opposite
sides of the transition, and perhaps, as the case may
be, by a larger number or by a smaller number of
species forming a hump or a trough in the smooth
curve expected from a climatic continuum.
Appendix 1
Bibliographic Key to Geographic Units in the Tables and Figures
Acre, Brazil (Pinto, 1938, 1944, 1954; Novaes, 1957; Meyer de
Schauensee, 1970)
Admiralty Islands, New Guinea region (Rothschild and
Hartert, 1914; Ripley, 1947)
Alabama (Imhof, 1962)
Alagoas, Brazil (Pinto, 1954; Meyer de Schauensee, 1970)
Alajuela Province, Costa Rica (Carriker, 1910; Slud, 1964a;
compiled sources)
Alaska (Gabrielson and Lincoln, 1959)
Albania (Ticehurst and Whistler, 1932)
Albemarle, Galapagos Islands (Swarth, 1931)
Alberta, Canada (Salt and Wilk, 1961)
Aldabra Islands, Indian Ocean (Benson and Penny, 1971)
Amapa, Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970; compiled sources)
Amazonas, Brazil (compiled sources)
Amazonian Brazil (compiled sources)
Ambrym, New Hebrides (Mayr, 1945)
Anamba Islands, South China Sea (Oberholser, 1917)
Andaman Islands, Bay of Bengal (Abdulali, 1965, 1968)
Andros, Bahamas (Bond, 1971)
Anegada, Virgin Islands (Bond, 1971)
Aneiteum, New Hebrides (Mayr, 1945)
Anglo-Egyptian Sudan (Cave and Macdonald, 1955)
Angola (Traylor, 1963; Ripley and Heinrich, 1966)
Anguilla, West Indies (Peters, J. L., 1927; Bond, 1971)
Anjouan, Comoro Islands (Benson, C. W., 1960)
Annobon, Gulf of Guinea (Moreau, 1966)
Antigua, West Indies (Danforth, 1934; Bond, 1971)
Arabia (Meinertzhagen, 1954)
Arctic Canada (Snyder, L. L., 1957)
Arctic slope of Alaska (Bailey, 1948)
Argentina (Olrog, 1963; Meyer de Schauensee, 1970)
Arizona (Phillips et al., 1964)
Arkansas (Baerg, 1951)
Am Islands, New Guinea (Rand and Gilliard, 1967)
Aruba, Netherlands Antilles (Voous, 1957)
Atlantico, Colombia (Dugand, 1947)
Aurora (Maewo), New Hebrides (Mayr, 1945)
Austral Islands, French Polynesia (Bruner, 1972)
Australia (Slater, 1971)
Australian Region (Mayr, 1941a; Rand and Gilliard, 1967;
Slater, 1971)
Azores (Bannerman and Bannerman, 1965-1968)
Azuero Peninsula (Aldrich and Bole, 1937)
Bahamas (Bond, 1971)
Bahfa, Brazil (Pinto, 1938, 1944; Meyer de Schauensee, 1970)
Baja California, Mexico (Grinnell, 1928; Edwards, 1972)
Bali, East Indies (Kuroda, 1936)
Baluchistan (Vaurie, 1959-1965)
Banks Island, Canada (Manning et al., 1956)
Banks Islands, New Hebrides (Mayr, 1945)
Barbados, West Indies (Bond, 1971)
Barbareta, Bay Islands of Honduras (Bond, 1936; Monroe,
1968)
Barbuda, West Indies (Bond, 1971)
Barro Colorado Island (Eisenmann, 1952; compiled sources)
Bartica, Guyana (Beebe, 1925; Beebe et al., 1917)
Basilan, Philippines (duPont, 1971)
Bay Islands, Honduras (Bond, 1936; Monroe, 1968)
Bechuanaland (Smithers, 1964)
Belem, Brazil (Novaes [MS.]; compiled sources)
Belgian Congo (Chapin, 1932-1954; White, 1960-1965)
Bermuda (Wingate, 1959)
Biak Island, New Guinea (Mayr and Meyer de Schauensee,
1939; Rand and Gilliard, 1967)
Big Bend National Park, Texas (Wauer, 1973)
Black Hills, So. Dakota-Wyoming (Pettingill and Whitney,
1965)
Bohol, Philippines (duPont, 1971)
Bolivia (Meyer de Schauensee, 1966, 1970)
Bonaire, Netherlands Antilles (Voous, 1957)
Borneo (Smythies, 1968)
Bougainville, Solomon Islands (Mayr, 1945)
Brazil (Pinto, 1938, 1944; Meyer de Schauensee, 1970)
British Columbia, Canada (Munroe and Cowan, 1947)
British Honduras (Russell, 1964)
British Isles (British Ornithologists' Union, 1952)
British Somaliland (Archer and Godman, 1937-1961)
Bulgaria (Pateff, 1950)
Bunguran, Natuna Islands (Oberholser, 1932)
Burma (Smythies, 1953)
Buru, Moluccas (van Bemmel, 1948)
Burundi (Schouteden, 1966b)
Bwamba County, W. Uganda (van Someren and van Someren,
1949; Friedmann and Williams, 1971)
California (McCaskie et al., 1970)
Camorta, Nicobar Islands (Abdulali, 1968)
Campeche, Mexico (Paynter, 1955; Storer, 1961)
Canada (Godfrey, 1966)
Canary Islands (Bannerman, 1963)
Cape Verde Islands (Bannerman and Bannerman, 1965-1968)
Car Nicobar, Nicobar Islands (Abdulali, 1968)
Caroline Islands, Micronesia (Baker, 1951)
Cartago Province, Costa Rica (Carriker, 1910; Slud, 1964a;
compiled sources)
Cayo Largo, Cuba (Bond, 1950)
Ceara, Brazil (Hellmayr, 1929; Pinto, 1938, 1944; Meyer de
Schauensee, 1970)
Cebu, Philippines (duPont, 1971)
84
NUMBER 212 85
Celebes (Stresemann, 1939-1941)
Central America (Eisenmann, 1955)
Central and eastern South America (Pinto, 1938, 1944; Meyer
de Schauensee, 1970)
Central Brazil (Pinto, 1938, 1940, 1944; Meyer de Schauensee,
1970)
Ceram, Moluccas (van Bemmel, 1948)
Ceylon (Henry, 1955)
Charles, Galapagos Archipelago (Swarth, 1931)
Chiapas, Mexico (Alvarez del Toro, 1964; Edwards, 1972)
Chile (Johnson, 1965-1967)
China (Cheng, 1955-1958)
Choiseul, Solomon Islands (Mayr, 1945)
Christmas Island, Indian Ocean (Pearson, 1966)
Churchill region, Manitoba (Jehl and Smith, 1970)
Clarion, Revilla Gigedos Islands, Mexico (Brattstrom and
Howell, 1956)
Cocos Island, E. Pacific (Slud, 1967)
Cocos-Keeling Islands, Indian Ocean (Watson et al., 1963)
Coiba Island, Panama (Wetmore, 1957)
Colima and adjacent Jalisco, Mexico (Schaldach, 1963)
Colombia (Meyer de Schauensee, 1964)
Colorado (Bailey and Niedrach, 1965)
Comoro Islands, Indian Ocean (Benson, C. W., 1960)
Costa Rica (Slud, 1964a)
Cozumel Island, Mexico (Paynter, 1955)
Crete (Stresemann, 1956)
Cuba (Bond, 1971)
Culebra Island, Puerto Rico (Leopold, 1963; Bond, 1971)
Cundinamarca, Colombia (Olivares, 1969)
Curasao, Netherlands Antilles (Voous, 1957)
Cyprus (Cyprus Ornithological Society, 1972)
Darjeeling area, India (Matthews and Edwards, 1944)
Death Valley, California (Wauer, 1962)
Delaware (Barnhill, 1972)
D'Entrecasteaux Archipelago, New Guinea (Rand and Gil-
liard, 1967)
Dominica, West Indies (Bond, 1971)
East and northeast Africa (Mackworth-Praed and Grant,
1952-1955)
East Cape Province, South Africa (Skead, 1967)
East-central Brazil (Meyer de Schauensee, 1970; compiled
sources)
Eastern China (La Touche, 1925-1934)
Eastern Nigeria (Serle, 1957)
Ecuador (Meyer de Schauensee, 1970)
Efate, New Hebrides (Mayr, 1945)
Egypt (Etchecopar and Hue, 1967)
Ellice Islands, W. Pacific (Child, 1960)
El Salvador (Rand and Traylor, 1954)
Enggano, W. Sumatra Islands (Ripley, 1944)
Epi, New Hebrides (Mayr, 1945)
Equateur-Tshuapa, Congo (Schouteden, 1961)
Eritrea (Smith, K. D., 1957)
Erromanga, New Hebrides (Mayr, 1945)
Espfrito Santo, Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970)
Espfritu Santo, New Hebrides (Mayr, 1945)
Ethiopia (Urban and Brown, 1971)
Ethiopian Region (White, 1960-1965)
Eurasian tundra (Pleske, 1928)
Europe (Bruun, 1970)
Europe and North Africa (Heinzel et al., 1972)
Falkland Islands (CawkeU and Hamilton, 1961)
Fernando Po, Gulf of Guinea (Moreau, 1966)
Fiji Islands (Mayr, 1945)
Finca La Selva, Costa Rica (Slud, 1960)
Finland (Bruun, 1970)
Flores, East Indies (Kuroda, 1936)
Florida (Sprunt, 1954)
France-Belgium-Switzerland (Barruel, 1949)
French Cameroon (Good, 1952)
French Guiana (Berlepsch, 1908; Meyer de Schauensee, 1970;
compiled sources)
French Indo-China (Delacour and Jabouille, 1931)
Fuerteventura, Canary Islands (Moreau, 1966)
Fukien, China (Cheng, 1955-1958)
Galapagos Islands (Swarth, 1931; Leveque et al., 1966)
Gambia Valley (Moreau, 1966)
Gaua (Sta. Maria), Banks Islands (Mayr, 1945)
Georgia (Burleigh, 1958)
Germany (Niethammer et al., 1964)
Ghana (Karr, [MS.])
Gilbert Islands, W. Pacific (Amerson, 1969)
Gizo, Solomon Islands (Mayr, 1945)
Goias, Brazil (Pinto, 1938, 1944; Meyer de Schauensee, 1970)
Gomera, Canary Islands (Moreau, 1966)
Gonave, Hispaniola (Wetmore and Swales, 1931; Bond, 1971)
Gran Canaria, Canary Islands (Moreau, 1966)
Gran Chaco (Laubmann, 1930)
Gran Comoro, Comoro Islands (Benson, C. W., 1960)
Grand Cayman, West Indies (Cory, 1886; Bangs, 1916; Bond,
1971)
Great Britain (British Ornithologists' Union, 1952)
Great Nicobar, Nicobar Islands (Abdulali, 1968)
Greater Antilles (Bond, 1971)
Greater Sundas (Delacour, 1947)
Greece (Lambert, 1957)
Greenland (Salomonsen, 1950)
Grenada, West Indies (Wells, 1887; Bond, 1971)
Grenadines, West Indies (Bond, 1971)
Guadalcanal, Solomon Islands (Mayr, 1945; Cain and Gal-
braith, 1956)
Guadalupe Island, Mexico (Howell and Cade, 1954)
Guadeloupe, West Indies (Danforth, 1939a; Bond, 1971)
Guam, Marianas (Baker, 1951)
Guanacaste Province, Costa Rica (Carriker, 1910; Slud, 1964a;
compiled sources)
Guanaja, Bay Islands of Honduras (Bond, 1936; Monroe,
1968)
Guatemala (Land, 1970)
Gujarat, India (Ali, 1954-1955)
Guyana (Snyder, D. E., 1966; Meyer de Schauensee, 1970)
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Hainan, China (Hachisuka, 1939)
Halmahera, Moluccas (van fiemmel, 1948)
Hawaii (Peterson, 1961)
Hawaiian Islands (Peterson, 1961)
Henderson Island, Central Pacific (Williams, 1960)
Heredia Province, Costa Rica (Carriker, 1910; Slud, 1960,
1964a; compiled sources)
Hierro, Canary Islands (Moreau, 1966)
Hispaniola (Wetmore and Swales, 1931; Bond, 1971)
Hokkaido, Japan (Ornithological Society of Japan, 1958)
Honduras (Monroe, 1968)
Hong Kong Colony (Macfarlane and Macdonald, 1966)
Honshu, Japan (Ornithological Society of Japan, 1958)
Hopei, China (Shaw, 1936)
Hungary (Keve, 1960)
Ibadan, Nigeria (Elgood and Sibley, 1964)
Iberian Peninsula (Bruun, 1970)
Idaho (Burleigh, 1972)
Illinois (Smith and Parmalee, 1955)
Impenetrable Forest, Uganda (Keith et al., 1969; Friedmann
and Williams, 1970)
Indefatigable, Galapagos Islands (Swarth, 1931)
Indian subcontinent (Ripley, 1961)
Indiana (Mumford, [MS.])
Iowa (Grant, 1963)
Iran (Hue and Etchdcopar, 1970)
Iraq (Hue and Etch&opar, 1970)
Ireland (Kennedy, 1961)
Isia Grande, Tierra del Fuego (Humphrey et al., 1970)
Isla Margarita, Venezuela (Fernandez and Benedetti, 1940;
Yepez, 1957)
Isle of Pines, Cuba (Bangs and Zappey, 1905; Bond, 1971)
Ituri, Congo (Schouteden, 1963b)
Jamaica (Bond, 1971)
Japan (Austin and Kuroda, 1953; Ornithological Society of
Japan, 1958)
Japen Island, New Guinea (Rand and Gilliard, 1967)
Java (Kuroda, 1936; Delacour, 1947)
Jervis, Galapagos Archipelago (Swarth, 1931)
Jodhpur, India (Whistler, 1938)
Joshua Tree Monument, California (Miller and Stebbins,
1964)
Kamchatka, Siberia (Bergman, 1935)
Kanab area, Utah (Behle et al., 1958)
Kandavu, Fiji Islands (Mayr, 1945)
Kansas (Johnston, 1960)
Kasai, Congo (Schouteden, 1964)
Katmai, Alaska (Cahalane, 1959)
Kauai, Hawaiian Islands (Peterson, 1961)
Keewatin District, Canada (Godfrey, 1966)
Kei Islands, Moluccas (van Bemmel, 1948)
Kentucky (Mengel, 1965)
Kenya-Uganda (Jackson, 1938)
Kerala (Ali, 1969)
Kolombangara, Solomon Islands (Mayr, 1945)
Korea (Austin, 1948; Gore and Won, 1971)
Kurile Islands (Bergman, 1935)
Kusaie, Caroline Islands (Baker, 1951)
Kutch, India (Ali, 1945)
Kwango-Kwilu, Congo (Schouteden, 1965)
Labrador Peninsula, Canada (Todd, 1963)
Lanai, Hawaiian Islands (Peterson, 1961)
Lanzarote, Canary Islands (Moreau, 1966)
Lau Islands, Southwest Pacific (Mayr, 1945)
Lebanon (Benson, S. V., 1970)
Laysan, Hawaiian Islands (Peterson, 1961)
Leeward Islands, Lesser Antilles (Bond, 1971)
Lesser Antilles, West Indies (Clark, 1905; Bond, 1971)
Lesser Sundas, East Indies (Kuroda, 1936)
Leyte, Philippines (duPont, 1971; Parkes, 1973)
Liberia (Forbes-Watson, [1970])
Libya (Toschi, 1969)
Lifu, Loyalty Islands (Mayr, 1945; Delacour, 1966)
Lima Department, Peru (Koepcke, 1964)
Lombok, East Indies (Kuroda, 1936)
Lord Howe Island, So. Pacific (Hindwood, 1940; Darlington,
1957)
Louisiade Archipelago, New Guinea (Rand and Gilliard,
1967)
Louisiana (Lowery, 1955)
Lower Amazonia (Snethlage, 1914; Griscom and Greenway,
1941; Pinto, 1938, 1944; Meyer de Schauensee, 1970; com-
piled sources)
Loyalty Islands, New Caledonia (Mayr, 1945; Delacour, 1966)
Luzon, Philippines (duPont, 1971)
Mackenzie District, Canada (Godfrey, 1966)
Madagascar (Rand, 1936)
Mafia Island, Tanzania (Moreau, 1966)
Maine (Palmer, 1949)
Malaita, Solomon Islands (Mayr, 1945)
Malay Peninsula (Glenister, 1951)
Malaya (Delacour, 1947)
Malaysia (Delacour, 1947)
Malekula, New Hebrides (Mayr, 1945)
Manchuria (Vaurie, 1959-1965)
Manitoba, Canada (Godfrey, 1966)
Manua Islands, Samoa (Mayr, 1945)
Maranhao, Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970)
Mard, Loyalty Islands (Mayr, 1945; Delacour, 1966)
Margarita I., Venezuela (Fernandez and Benedetti, 1940;
Ye-pez, 1957)
Marfa Cleofas, Tres Marias Islands, Mexico (Stager, 1957)
Maria Madre, Tres Marias Islands, Mexico (Stager, 1957)
Maria Magdalena, Tres Marias Islands, Mexico (Stager, 1957)
Mariana Islands, Micronesia (Baker, 1951)
Marie Galante, West Indies (Danforth, 1939a; Bond, 1971)
Marquesas, French Polynesia (Bruner, 1972)
Marshall Islands, Micronesia (Baker, 1951)
Martinique, West Indies (Bond, 1971)
Maryland (Stewart and Robbins, 1958)
NUMBER 212 87
Massachusetts (Griscom and Snyder, 1955)
Mato Grosso, Brazil (Pinto, 1938, 1940, 1944; Sick, 1955;
Meyer de Schauensee, 1970; Fry, 1970)
Maui, Hawaiian Islands (Peterson, 1961)
Mauritius, Indian Ocean (Watson et al., 1963)
Mayotte, Comoro Islands (Benson, C. W., 1960)
Mentawai group, W. Sumatra Islands (Ripley, 1944)
Meta, Colombia (Meyer de Schauensee, 1964)
Mexico (Edwards, 1972)
Michigan (Zimmerman and Van Tyne, 1959)
Micronesia (Baker, 1951)
Middle America (Eisenmann, 1955; Edwards, 1972)
Middle East (Hue and Etchecopar, 1970)
Minas Gerais, Brazil (Pinto, 1952; Meyer de Schauensee,
1970)
Mindanao, Philippines (duPont, 1971)
Mindoro, Philippines (duDont, 1971)
Minnesota (Roberts, T. S., 1949)
Misiones, Argentina (Olrog, 1963; Meyer de Schauensee,
1970)
Mississippi (Mississippi Ornithological Society, 1962)
Mississippi Coast (Burleigh, 1944)
Missouri (Widmann, 1907)
Moheli, Comoro Islands (Benson, C. W., 1960)
Mollendo, Peru (Hughes, 1970)
Molokai, Hawaiian Islands (Peterson, 1961)
Moluccas (van Bemmel, 1948)
Mona Island, Puerto Rico (Raffaele, 1973)
Mongala-Ubangi, Congo (Schouteden, 1962b)
Mongolia (Vaurie, 1959-1965, 1964)
Mono, Solomon Islands (Mayr, 1945)
Montana (Skaar et al., 1971)
Montserrat, West Indies (Danforth, 1939b; Bond, 1971)
Morocco (Etchecopar and Hue, 1967)
Mushie, Congo (Schouteden, 1962a)
Mysore, India (AH, 1943-1944)
Natal-Zululand, South Africa (Clancey, 1964)
Natuna Islands, South China Sea (Oberholser, 1932)
Navajo country, Utah-Arizona (Woodbury and Russell, 1945)
Near East (Hue and Etchecopar, 1970)
Nearctic Region (American Ornithologists' Union, 1957; Ed-
wards, 1972)
Nebraska (Rapp et al., 1958)
Negros, Philippines (duPont, 1971)
Neotropical Region (Edwards, 1972; Eisenmann, 1955; Meyer
de Schauensee, 1970)
Netherlands (Commissie Nederlandse Avifauna, 1962)
Nevada (Linsdale, 1936)
Nevis, West Indies (Danforth, 1936; Bond, 1971)
New Brunswick, Canada (Squires, 1952)
New Caledonia, Indian Ocean (Delacour, 1966)
New Georgia, Solomon Islands (Mayr, 1945)
New Guinea (Rand and Gilliard, 1967)
New Hampshire (Richards, 1964)
New Hebrides, Southwest Pacific (Mayr, 1945)
New Jersey (Fables, 1955)
New Mexico (Hubbard, 1970)
New York (Reilly and Parkes, 1959)
New Zealand (Fleming et al., 1953)
Newfoundland Island, Canada (Peters and Burleigh, 1951)
Newfoundland Labrador, Canada (Austin, 1932)
Nias, W. Sumatra Islands (Ripley, 1944)
Nicaragua (Eisenmann, 1955)
Nicobar Islands, Bay of Bengal (Abdulali, 1968)
Nissan, Solomon Islands (Mayr, 1945; Ripley, 1947)
Niue, South Pacific (Wodzicki, 1971)
Nontropical Mexico (Edwards, 1972)
Norfolk Island, South Pacific (Darlington, 1957)
North Africa (Etchecopar and Hue, 1967)
North America (Robbins et al., 1966; Eisenmann, 1955)
North America (A.O.U.) (American Ornithologists' Union,
1957)
North Carolina (Pearson et al., 1959)
North Dakota (Stewart, [MS.])
North Island, New Zealand (Fleming et al., 1953)
Northeastern Brazil (Pinto, 1938, 1944, 1954; Pinto and
Camargo, 1961; Meyer de Schauensee, 1970)
Northeastern Para (compiled sources)
Northeastern Venezuela (Friedmann and Smith, 1950, 1955)
Northern Bolivia (Gyldenstolpe, 1945b)
Northern Brazil-Southern Venezuela (Friedmann, 1948)
Northern Libya (Stanford, 1954)
Northern South America (Phelps and Phelps. 1958, 1963;
Meyer de Schauensee, 1970)
Northern Territory, Australia (Condon, 1944-1945)
Northern Thailand (Deignan, 1945)
Northwest Kansu and Kuku Nor (Stresemann et al., 1937-
1938)
Northwest South America (Meyer de Schauensee, 1970)
Nova Scotia, Canada (Tufts, 1961)
Nyasaland (Benson, C. W., 1953)
Oahu, Hawaiian Islands (Peterson, 1961)
Oaxaca, Mexico (Friedmann et al., 1950; Miller et al., 1957;
Edwards, 1972; Peterson and Chalif, 1973)
Oba, New Hebrides (Mayr, 1945)
Obidos, Brazil (compiled sources)
Ohio (Trautman and Trautman, 1968)
Oklahoma (Sutton, 1967)
Old Providence, West Indies (Bond, 1950, 1971)
Ontario, Canada (Snyder, L. L., 1951)
Oregon (Bertrand and Scott, 1971)
Orinoco region, Venezuela (Cherrie, 1916)
Ovalau, Fiji Islands (Mayr, 1945)
Palau Islands, Micronesia (Baker, 1951)
Palawan, Philippines (Delacour and Mayr, 1946; duPont,
1971)
Palearctic Region (Vaurie, 1959-1965)
Palma, Canary Islands (Moreau, 1966)
Panama (Griscom, 1935; Eisenmann, 1955; Wetmore, 1965-
1972a)
Panama Canal Zone (Eisenmann and Loftin, 1968)
Panay, Philippines (duPont, 1971)
Para, Brazil (Pinto, 1938, 1944; compiled sources)
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Paraguana Peninsula (Barnes, 1940)
Paraguay (Laubmann, 1939-1940; Meyer de Schauensee, 1970)
Paraiba, Brazil (Pinto and Camargo, 1961; Meyer de Schauen-
see, 1970)
Parana, Brazil (Pinto, 1938, 1944; Pinto and Camargo, 1955;
Meyer de Schauensee, 1970)
Patagonia, Argentina (Olrog, 1959)
Pavuvu, Solomon Islands (Mayr, 1945)
Pearl Archipelago (MacArthur et al., 1972; Wetmore, 1965-
1972a)
Pemba Island, Tanzania (Moreau, 1966)
Pennsylvania (Poole, 1964)
Pentecost, New Hebrides (Mayr, 1945)
Perija, Venezuela (Phelps, 1943; Gines et al., 1953)
Pernambuco, Brazil (Pinto, 1938, 1944; Berla, 1946; Meyer
de Schauensee, 1970)
Peru (Meyer de Schauensee, 1970)
Pet?n, Guatemala (Smithe, 1966; Land, 1970)
Philippines (duPont, 1971)
Phoenix Islands, Central Pacific (Roger Clapp, pers. comm.)
Piauf, Brazil (Pinto, 1938, 1944; Meyer de Schauensee, 1970)
Pitcairn Islands, Central Pacific (Williams, 1960)
Ponape, Caroline Islands (Baker, 1951)
Portugal (Themido, 1952)
Prince Edward Island, Canada (Godfrey, 1954)
Principe, Gulf of Guinea (Moreau, 1966)
Puerto Rico (Wetmore, 1927; Leopold, 1963; Bond, 1971)
Puntarenas Province, Costa Rica (Carriker, 1910; Slud, 1964a;
compiled sources)
Quintana Roo, Mexico (Paynter, 1955)
Rancho Grande, Venezuela (Schafer and Phelps, 1954)
Rendova, Solomon Islands (Mayr, 1945)
Rennell, Solomon Islands (Mayr, 1931, 1945)
Reunion (Watson et al., 1963)
Revilla Gigedos Islands, Mexico (Brattstrom and Howell,
1956)
Rio de Janeiro, Brazil (Pinto, 1938, 1944; Meyer de Schauen-
see, 1970)
Rio Grande do Sul, Brazil (Pinto, 1938, 1944; Camargo, 1962;
Meyer de Schauensee, 1970)
Roatan, Bay Islands of Honduras (Bond, 1936; Monroe, 1968)
Rodriguez, Indian Ocean (Watson et al., 1963)
Roraima, Brazil (Pinto, 1966)
Rota, Marianas (Baker, 1951)
Rwanda (Schouteden, 1966a)
Saba, West Indies (Voous and Koelers, 1967)
Saipan, Marianas (Baker, 1951)
Samar, Philippines (duPont, 1971)
Samoa (Ashmole, 1963)
San Cristobal, Solomon Islands (Mayr, 1945; Cain and Gal-
braith, 1956)
San Jose\ Pearl Islands, Panama (Wetmore, 1946)
San Jos? Province, Costa Rica (Carriker, 1910; Slud, 1964a;
compiled sources)
Santa Ana, Solomon Islands (Mayr, 1945)
Santa Catarina, Brazil (Pinto, 1938, 1944; Meyer de Schauen-
see, 1970)
Santa Cruz Islands, Southwest Pacific (Mayr, 1945)
Santa Elena Peninsula, Ecuador (Marchant, 1958)
Santa Isabel, Solomon Islands (Mayr, 1945)
Santa Marta area, Colombia (Todd and Carriker, 1922; Dar-
lington, 1931)
Santarem, Brazil (Riker, 1890-1891; Griscom and Greenway,
1941; Pinto, 1938, 1944; Meyer de Schauensee, 1970; com-
piled sources)
Sao Paulo, Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970)
Sao Tiago, Cape Verde Islands (Moreau, 1966)
Sao Tome\ Gulf of Guinea (Moreau, 1966)
Saskatchewan, Canada (Houston et al., 1958)
Savaii, Samoa (Mayr, 1945)
Scandinavian Peninsula (Bruun, 1970)
Seychelles (Gaymer et al., 1969; Watson et al., 1963)
Sierra Macarena, Colombia (Blake, 1962)
Simalur, W. Sumatra Islands (Ripley, 1944)
Sinaloa, Mexico (Friedmann et al., 1950; Miller ct al., 1957;
Edwards, 1972; Peterson and Chalif, 1973)
Singapore (Gibson-Hill, 1950a, 1950b)
Society Islands, French Polynesia (Bruner, 1972)
Socorro, Revilla Gigedos Islands, Mexico (Brattstrom and
Howell, 1956)
Socotra, Indian Ocean (Ripley and Bond, 1966)
Solomon Islands, S. W. Pacific (Mayr, 1945; Cain and Gal-
braith, 1956)
Sonora, Mexico (van Rossem, 1945; Edwards, 1972)
South America (Meyer de Schauensee, 1966, 1970)
South Australia (Condon, 1962)
South Carolina (Sprunt and Chamberlain, 1949)
South China (Caldwell and Caldwell, 1931)
South Dakota (Over and Thorns, 1946)
South Island, New Zealand (Fleming et al., 1953)
South Vietnam (Wildash, 1968)
Southampton Island, Canada (Sutton, 1932)
Southeastern Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970)
Southeastern Transbaikal (Gagina, 1960)
Southeastern Utah (Behle, 1960)
Southern Africa (Roberts, A., 1970)
Southern Brazil (Pinto, 1938, 1944; Meyer de Schauensee,
1970)
Southern Burma (Smythies, 1953)
Southern California (Pyle, 1953)
Southern Cameroon (Germain et al., 1973)
Southern Illinois (George, 1968)
Southern Patagonia, Argentina (Olrog, 1959)
Southern Rhodesia (Smithers et al., 1957)
Southern South America (Meyer de Schauensee, 1970)
Southern third of Africa (Mackworth-Praed and Grant, 1962-
1963)
South-West Africa (Hoesch and Niethammer, 1940)
Southwestern Ohio (Kemsies and Randle, 1953)
Soviet Union (Dementiev and Gladkov, 1951-1954)
Spanish Sahara (Valverde, 1957)
NUMBER 212 89
St. Andrew, West Indies (Bond, 1950, 1971)
St. Croix, Virgin Islands (Wetmore, 1918; Beatty, 19S0; Leo-
pold, 1963)
St. Eustatius, West Indies (Voous and Koelers, 1967)
St. John, Virgin Islands (Nichols, 1943; Robertson, 1962;
Bond, 1971)
St. Kitts, West Indies (Danforth, 1936; Bond, 1971)
St. Lucia, West Indies (Danforth, 1935; Bond, 1971)
St. Martin, West Indies (Voous and Koelers, 1967)
St. Thomas, Virgin Islands (Wetmore, 1918; Nichols, 1943;
Leopold, 1963)
St. Vincent, West Indies (Bond, 1971)
Sudan (Cave and Macdonald, 1955)
Sula Islands, Moluccas (van Bemmel, 1948)
Sumatra (Delacour, 1947)
Sumba, East Indies (Mayr, 1944a)
Sumbawa, East Indies (Kuroda, 1936)
Sunda Islands, East Indies (Kuroda, 1936; Mayr, 1944a; Dela-
cour, 1947)
Surinam (Haverschmidt, 1968)
Swan Islands, West Indies (Paynter, 1956)
Tabasco, Mexico (Friedmann et al., 1950; Miller et al., 1957;
Peterson and Chalif, 1973)
Tahiti, French Polynesia (Bruner, 1972)
Taiwan (Hachisuka and Udagwa, 1950-1951; Severinghaus et
al., 1970)
Tamaulipas, Mexico (Friedmann et al., 1950; Miller et al.,
1957; Edwards, 1972; Peterson and Chalif, 1973)
Tanna, New Hebrides (Mayr, 1945)
Tasmania (Sharland, 1945)
Taveuni, Fiji Islands (Mayr, 1945)
Tenasserim, Burma (Smythies, 1953)
Tenerife, Canary Islands (Moreau, 1966)
Texas (Peterson, 1963)
Thailand (Lekagul, 1968)
Thames Valley, England (Moreau, 1966)
The Guianas (Meyer de Schauensee, 1966, 1970)
Tibet (Vaurie, 1972)
Tikal, Guatemala (Smithe, 1966)
Timor, East Indies (Mayr, 1944a)
Tinian, Marianas (Baker, 1951)
Tobago, West Indies (Herklots, 1961; ffrench and ffrench,
1966)
Tonga Islands, Southwest Pacific (Mayr, 1945)
Torres Islands, New Hebrides (Mayr, 1945)
Tortola, Virgin Islands (Bond, 1971)
Tranninh, Laos (David-Beaulieu, 1944)
Tres Marias Islands, Mexico (Stager, 1957)
Trinidad, West Indies (Herklots, 1961; ffrench and ffrench,
1966)
Truk, Caroline Islands (Baker, 1951)
Tuamotu Archipelago, French Polynesia (Bruner, 1972)
Tunisia (Etch?copar and Hue, 1967)
Turkey (Hue and Etchecopar, 1970)
Tutuila, Samoa (Mayr, 1945)
Uele, Congo (Schouteden, 1963a)
United States (American Ornithologists' Union, 1957)
Upolu, Samoa (Mayr, 1945)
Upper Amazonia (Pinto, 1938, 1944; Gyldenstolpe, 1945a,
1951; Meyer de Schauensee, 1970; compiled sources)
Uruguay (Cuello and Gerzenstein, 1962; Meyer de Schauensee,
1970)
Utah (Woodbury et al., 1969)
Utila, Bay Islands of Honduras (Bond, 1936; Monroe, 1968)
Uvea, Loyalty Islands (Mayr, 1945; Delacour, 1966)
Valua (Saddle I.), Banks Islands (Mayr, 1945)
Vanua Lava, Banks Islands (Mayr, 1945)
Vanua Levu, Fiji Islands (Mayr, 1945)
Vaup&, Colombia (Olivares, 1955, 1964a, 1964b; Olivares and
Hernandez, 1962)
Vella Lavella, Solomon Islands (Mayr, 1945)
Venezuela (Phelps and Phelps, 1958, 1963)
Veracruz, Mexico (Friedmann et al., 1950; Miller et al., 1957;
Edwards, 1972; Peterson and Chalif, 1973)
Victoria, Australia (Wheeler, 1967)
Vieques, Puerto Rico (Wetmore, 1927; Danforth, 1937; Leo-
pold, 1963)
Virgin Gorda, Virgin Islands (Bond, 1971)
Virgin Islands (Wetmore, 1927; Leopold, 1963; Bond, 1971)
Virginia (Murray, 1952)
Vitu Levu, Fiji Islands (Mayr, 1945)
Waigeu Island, New Guinea (Rand and Gilliard, 1967)
Washington (Alcorn, 1971)
West Chiriquf Highlands (Eisenmann and Loftin, 1967a)
West Indies (Bond, 1971)
West Sumatra Islands (Ripley, 1944)
West Virginia (Hall, 1971)
West-central and west Africa (Mackworth-Praed and Grant,
1970-1973)
Western Australia (Serventy and Whittell, 1951)
Western China-eastern Tibet (SchSfer, 1938)
Windward Islands, Lesser Antilles (Bond, 1971)
Wisconsin (Gromme, 1963)
World (Austin, 1971)
Wyoming (McCreary, 1939)
Yap, Caroline Islands (Baker, 1951)
Yucatan, Mexico (Paynter, 1955)
Yucatan Peninsula, Mexico (Paynter, 1955)
Yugoslavia (Matvejev and Vasic, 1973)
, Yukon Territory, Canada (Godfrey, 1966)
Zambia (Benson et al., 1971)
Zanzibar Island, Tanzania (Moreau, 1966)
Zaria, Northern Nigeria (Fry, 1966)
Appendix 2
Tables
TABLE 1.?World survey of species densities and passerine-nonpasserine ratios
(TL = total land species, NL = native land species)
World (l iving and recently extinct)
Soviet Union
North America (mainland)
Ethiopian Region (mainland)
Neotropical Region (mainland south
of Isthmus of Tehuantepec)
Temperate North America (excluding
Middle America and Greenland)
China (mainland)
Europe
Canada
Brazil
Australia
United States (coterminous)
Indian Subcontinent (mainland)
Russia in Europe
Southern South America (Paraguay,
Uruguay, Argentina, Chile)
Northwestern South America (Colombia,
Ecuador, Peru, Bolivia)
Southern South America (Argentina,
Chile)
Southern Africa
Northwest Africa (Morocco,
Algeria, Tunisia)
Amazonian Brazil (Amazonas, Para,
Amapa)
Bolivia-Mato Grosso (including
Rondonia)
Argentina
Northwest South America (Colombia,
Ecuador, Peru)
Mongolia
Arabia
Area
(mi2)
52,410,000
8,354,200
8,150,000
8,000,000
7,280,000
7,156,000
3,878,150
3,800,000
3,610,000
3,286,000
2,974,500
2,959,500
1,730,650
1,595,500
1,588,300
1,536,650
1,359,150
1,312,000
1,141,600
1,121,700
1,095,400
1,072,750
1,023,500
1,016,100
1,000,000
Total
Avifauna
0.0165
0.0084
0.018
0.021
0.045
0.0093
0.027
0.015
0.014
0.047
0.023
0.022
0.067
0.063
0.15
0.073
0.061
0.034
0.088
0.12
0.086
0.20
0.038
Species/100 mi2
Land Avifauna
Total
(TL)
0.015
0.0053
0.015
0.018
0.040
0.0059
0.022
0.0087
0.0079
0.041
0.016
0.014
0.054
0.016
0.049
0.13
0.055
0.046
0.020
0.079
0.11
0.067
0.18
0.022
0.025
NL+(TL-NL)
0.015
0.0040
0.014
0.018
0.039
0.0052
0.017
0.0067
0.0057
0.041
0.016
0.012
0.051
0.013
0.048
0.13
0.054
0.044
0.016
0.078
0.11
0.066
0.18
0.016
0.020
Native
(NL)
0.015
0.0026
0.014
0.017
0.038
0.0044
0.013
0.0046
0.0035
0.040
0.016
0.010
0.047
0.0090
0.048
0.13
0.053
0.042
0.012
0.077
0.10
0.065
0.17
0.010
0.014
Passerines:Nonpasserines
Total
(TL)
1.94
2.05
1.71
1.96
1.84
2.08
2.51
1.97
2.52
1.80
1.87
2.08
2.06
1.78
1.71
1.76
1.73
1.55
1.72
1.54
1.75
1.71
1.75
1.76
1.41
Land Avifauna
NL+(TL-NL)
1.94
1.88
1.70
1.96
1.80
2.00
2.62
1.70
2.06
1.79
1.90
1.96
2.00
1.63
1.72
1.73
1.73
1.59
1.60
1.53
1.72
1.71
1.72
1.57
1.33
Native
(NL)
1.94
1.60
1.68
1.95
1.77
1.79
2.75
1.39
1.35
1.78
1.93
1.82
1.93
1.39
1.73
1.70
1.74
1.64
1.43
1.51
1.70
1.71
1.69
1.25
1.19
90
NUMBER 212 91
TABLE 1.?Continued
Middle America (mainland)
Anglo-Egyptian Sudan
Belgian Congo
Greenland
Tibet
Western Austral 1a (minus
Kimberley Division)
Mexico
Libya
Iran
Malaysia
Amazonas, Brazil
Alaska (mainland)
Ecuador-Peru
Mato Grosso (including Rondonia)
Venezuela-The Guianas
Labrador Peninsula
Para-Amapa, Brazil
Mackenzie District, Canada
Northern Territory, Australia
Bolivia
Peru
Angola
Para, Brazil
East-central Brazil (Bahfa-Goias)
Colombia
Manchuria
Egypt
South Australia
Area
(mi*)
992.952
950,950
895.348
839.800
835,660
831,920
763,944
679,358
628,000
617,924
595,474
586,400
583.738
582,391
531,504
530,000
526,241
525,000
523,620
513,086
482,257
481,226
470,752
459,659
439,825
408,162
386,200
380,070
Total
Avifauna
0.14
0.092
0.11
0.023
0.060
0.042
0.13
0.041
0.071
0.12
0.14
0.047
0.30
0.13
0.25
0.059
0.16
0.041
0.067
0.23
0.32
0.18
0.17
0.15
0.36
0.095
0.10
Species/100 mi2
Land Avifauna
Total
(TL)
0.11
0.074
0.098
0.0088
0.048
0.024
0.10
0.027
0.049
0.098
0.13
0.023
0.27
0.11
0.23
0.033
0.14
0.023
0.049
0.20
0.28
0.15
0.15
0.13
0.32
0.059
0.058
0.065
NL+(TL-NL)
0.11
0.069
0.095
0.0050
0.040
0.024
0.093
0.018
0.041
0.094
0.12
0.016
0.26
0.11
0.22
0.020
0.14
0.014
0.049
0.20
0.27
0.15
0.15
0.13
0.32
0.041
0.040
0.063
Native
(NL)
0.10
0.063
0.091
0.00083
0.032
0.024
0.085
0.0093
0.032
0.090
0.12
0.0095
0.26
0.11
0.21
0.0064
0.14
0.0053
0.049
0.20
0.27
0.14
0.15
0.13
0.31
0.023
0.021
0.060
Passerines:Nonpasserines
Total
(TL)
1.71
1.61
1.71
3.63
3.14
1.54
1.72
1.76
1.81
1.39
1.62
1.96
1.80
1.45
1.68
2.67
1.49
2.30
1.45
1.78
1.87
1.53
1.51
1.41
1.61
1.79
1.63
1.55
Land Avifaurv
NL+(TL-NL)
1.63
1.58
1.71
3.05
3.24
1.58
1.59
1.66
1.74
1.38
1.59
1.62
1.79
1.44
1.65
2.04
1.48
1.77
1.45
1.76
1.86
1.54
1.50
1.41
1.58
1.69
1.43
1.57
l
Native
(NL)
1.54
1.55
1.72
0.75
3.41
1.62
1.47
1.47
1.64
1.37
1.57
1.06
1.78
1.43
1.61
0.62
1.48
0.65
1.45
1.73
1.85
1.56
1.49
1.41
1.54
1.47
1.01
1.60
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Continued
Ontario
Bri t ish Columbia
Nigeria
Venezuela
New Guinea (Including coastal Islands)
Southeastern Brazil (SIo Paulo
through Rio Grande do Sul)
New Guinea (mainland)
Kenya-Uganda
Ethiopia
South-West Africa (excluding Caprivi
Str ip and Okavango)
Zambia
Borneo
Chile
Turkey (Asia Minor)
Scandinavian Peninsula
Texas
Burma
Afghanistan
Alberta
Goias, Brazil
France-Bel gi um-Swi tzerland
Saskatchewan
Central America
Keewatin D is t r ic t . Canada
Madagascar
Iberian Peninsula
Minas Gerais, Brazil
Southern Brazil (Parana, Sta.
Catarina, Rio Grande do Sul)
Bechuanaland
Area
(ml2)
363,282
359,279
356.667
352,141
322,044
319,468
306,600
306,000
305,731
296,887
290,320
289,993
286,396
285,162
277,480
262,398
261,610
250,965
248,800
244,330
240,369
237,975
228,578
228,000
227,678
227,384
226,179
224,009
222,000
Total
Avifauna
0.078
0.094
0.22
0.37
0.21
0.24
0.20
0.35
0.27
0.15
0.24
0.19
0.13
0.13
0.18
0.37
0.16
0.13
0.18
0.14
0.13
0.47
0.045
0.10
0.25
0.28
0.21
Spedes/100 mi2
Land Avifauna
Total
(TL)
0.051
0.057
0.18
0.33
0.17
0.19
0.17
0.30
0.21
0.11
0.20
0.14
0.070
0.086
0.064
0.13
0.31
0.12
0.082
0.16
0.082
0.086
0.40
0.018
0.059
0.083
0.22
0.23
0.17
NL+(TL-NL_)
0.036
0.042
0.17
0.32
0.17
0.19
0.16
0.29
0.20
0.10
0.19
0.14
0.066
0.066
0.047
0.11
0.29
0.099
0.051
0.16
0.066
0.052
0.37
0.012
0.058
0.067
0.22
0.23
0.16
Native
(NL)
0.021
0.027
0.16
0.31
0.16
0.19
0.16
0.28
0.18
0.091
0.17
0.13
0.061
0.046
0.030
0.06
0.27
0.076
0.020
0.16
0.050
0.018
0.35
0.0061
0.056
0.051
0.22
0.23
0.15
Passerines:Nonpasserines
Total
(TL)
2.62
2.14
1.54
1.68
1.54
1.61
1.65
1.78
1.49
1.25
1.55
1.30
1.74
1.78
1.45
2.48
1.86
2.30
2.62
1.33
1.65
2.53
1.60
1.67
0.93
1.84
1.43
1.70
1.30
Land Avifauna
NL+(TL-NL)
2.00
1.77
1.55
1.64
1.59
1.60
1.68
1.77
1.44
1.32
1.63
1.30
1.64
1.76
1.37
2.16
1.78
2.32
2.08
1.33
1.65
2.13
1.51
1.16
0.93
1.88
1.43
1.69
1.33
Native
(NL)
1.12
1.23
1.55
1.61
1.65
1.59
1.71
1.77
1.38
1.40
1.72
1.30
1.55
1.71
1.22
1.64
1.68
2.34
0.90
1.32
1.66
1.00
1.42
0.40
0.94
1.93
1.43
1.67
1.37
NUMBER 212 93
TABLE 1.?Continued
Manitoba
Bahfa, Brazil
Yukon Terr i tory, Canada
Thailand
Germany
Patagonia (Santa Cruz, Chubut)
The Guianas
Morocco
East Cape Province, South Africa
Sumatra
French Cameroon
California
Paraguay
Northeastern Brazil (Ceara
through Sergipe)
P1auf-Ceara, Brazil
Southern Rhodesia
Montana
Guyana-Surinam
Japan
Kasai, Congo
Baluchistan
Maranhao, Brazil
Finland
Near East (Syria, Lebanon, Israel ,
Transjordan)
New Mexico
British Isles
Iraq
Philippines
Northern Central America
(Guatemala, Honduras, El Salvador)
Area
(?12)
219,723
215,329
205,346
198,247
182,426
180,398
179,363
173,700
167,000
166,789
166,489
158,693
157,006
156,865
152,190
150,333
147,138
144,623
141,086
136,222
134,638
133,674
130,165
122,905
121,266
120,459
116,600
114,830
114,380
Total
Avifauna
0.14
0.28
0.42
0.24
0.44
0.20
0.33
0.42
0.31
0.41
0.26
0.25
0.38
0.23
0.53
0.30
0.39
0.32
0.30
0.32
0.33
0.30
0.44
0.63
Spedes/100 ml2
Land Avifauna
Total
(TL)
0.085
0.24
0.055
0.35
0.14
0.063
0.37
0.12
0.23
0.25
0.36
0.19
0.34
0.22
0.21
0.30
0.15
0.45
0.15
0.33
0.15
0.28
0.12
0.20
0.24
0.19
0.20
0.32
0.52
NL+(TL-NL)
0.053
0.24
0.034
0.32
0.098
0.061
0.36
0.10
0.21
0.25
0.36
0.14
0.33
0.22
0.21
0.28
0.095
0.44
0.11
0.32
0.13
0.28
0.085
0.15
0.18
0.13
0.14
0.30
0.47
Native
(NL)
0.020
0.24
0.014
0.28
0.051
0.060
0.35
0.076
0.20
0.24
0.35
0.097
0.32
0.22
0.21
0.26
0.039
0.43
0.075
0.31
0.097
0.28
0.049
0.10
0.12
0.070
0.076
0.27
0.43
Passerines .-Nonpasserines
Total
(TL)
2.48
1.35
2.14
1.50
1.94
1.72
1.48
1.64
1.31
1.28
1.60
2.60
1.51
1.49
1.33
1.45
2.36
1.48
1.90
1.36
1.54
1.52
1.68
1.74
2.46
2.20
1.66
1.42
1.53
Land Avifauna
NL+(TL-NL)
2.07
1.35
1.66
1.43
1.77
1.74
1.47
1.57
1.36
1.26
1.61
2.18
1.51
1.49
1.32
1.50
1.98
1.47
1.72
1.36
1.49
1.53
1.54
1.71
2.18
2.15
1.51
1.36
1.39
Native
(NL)
1.05
1.34
0.65
1.36
1.37
1.77
1.45
1.46
1.41
1.24
1.61
1.59
1.50
1.48
1.31
1.57
1.11
1.45
1.43
1.35
1.41
1.53
1.24
1.65
1.72
2.00
1.19
1.30
1.23
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Continued
Algeria (minus Saharan Departments)
Arizona
Newfoundland Labrador, Canada
Nevada
Rio Grande do Sul , Brazil
West Cape Province, South Africa
Spanish Sahara
Colorado
New Zealand
Ecuador
Wyomi ng
Roraima, Brazi l
Oregon
Yugoslavia
Sao Paulo, Brazi l
Piauf , Brazil
Uele, Congo
Surinam-French Guiana
Guyana
Great Br i ta in
V ic tor ia , Australia
Honshu, Japan
Korea
Utah
Greater Ant i l les
Minnesota
Idaho
Parana, Brazi l
Kansas
Southern Burma
Nebraska
Area
(n.12)
113,912
113,909
112,630
110,540
110,150
110,000
105,409
104,247
103,410
101,481
97,914
97,438
96,981
96,201
95,459
94,819
91,897
89,883
89,480
88,619
87,884
87,000
85,225
84,916
84,750
84,068
83,557
82,741
82,276
80,874
77,227
Total
Avifauna
0.30
0.38
0.13
0.28
0.47
0.38
0.20
0.41
0.21
1.34
0.33
0.44
0.35
0.40
0.71
0.35
0.71
0.76
0.83
0.48
0.44
0.44
0.38
0.48
0.39
0.36
0.48
0.47
0.71
0.49
Species/100 mi2
Land Avifauna
Total
(TL)
0.19
0.27
0.055
0.20
0.39
0.28
0.11
0.28
0.052
1.18
0.22
0.39
0.22
0.25
0.58
0.29
0.60
0.64
0.70
0.26
0.30
0.21
0.26
0.25
0.33
0.26
0.24
0.43
0.31
0.56
0.33
NL+(TL-NL)
0.15
0.21
0.036
0.14
0.37
0.26
0.076
0.19
0.046
1.16
0.15
0.39
0.16
0.19
0.58
0.29
0.57
0.62
0.68
0.18
0.28
0.15
0.17
0.18
0.26
0.16
0.17
0.43
0.19
0.52
0.20
Native
(NL)
0.11
0.14
0.017
0.080
0.36
0.25
0.041
0.099
0.039
1.14
0.072
0.38
0.11
0.13
0.58
0.29
0.54
0.61
0.66
0.091
0.25
0.096
0.078
0.12
0.20
0.062
0.095
0.43
0.078
0.47
0.078
Pa s ser1nes:Non pa s ser1nes
Total
(TL)
1.78
2.36
1.58
2.22
1.70
1.24
1.37
2.41
0.93
1.62
2.56
1.61
2.28
1.60
1.51
1.32
1.37
1.39
1.42
2.20
1.54
1.69
1.85
2.20
1.60
2.70
2.26
1.79
2.90
1.57
2.66
Land Avifauna
NL+(TL-NL)
1.69
2.23
1.25
2.08
1.69
1.29
1.34
2.03
1.00
1.61
2.07
1.60
1.97
1.60
1.51
1.32
1.37
1.38
1.40
2.15
1.60
1.94
1.59
1.94
1.29
2.08
1.99
1.80
2.12
1.46
2.02
Native
(NL)
1.54
2.00
0.58
1.78
1.68
1.35
1.26
1.31
1.11
1.59
1.15
1.58
1.49
1.60
1.50
1.31
1.38
1.37
1.37
2.00
1.68
2.07
0.99
1.49
0.93
0.91
1.47
1.80
0.74
1.35
0.73
NUMBER 212 95
TABLE 1.?Continued
South Dakota
Southern Central America
(Nicaragua, Costa Rica)
Mysore, India
Uruguay
Northeastern Brazil (Pernambuco,
Paraiba, Alagoas)
North Dakota
Sonora, Mexico
Malay Peninsula
Oklahoma
Missouri
Celebes
Washington State
British Somaliland
Honduras-El Salvador
Kwango-Kwilu, Congo
South Vietnam
Gujarat, India
Yucatan Peninsula, Mexico
Fukien, China
Hopei, China
Honduras
Georgia
Florida
Michigan
Vaupes, Colombia
Ceara, Brazil
Nicaragua
Illinois
Iowa
Wisconsin
Baja California, Mexico
Area(mi2)
77,047
76,381
74,326
72,172
70,937
70,665
70,477
70,000
69,919
69,674
69,255
68,192
67,936
67,420
65,400
65,000
64,000
63,034
61,259
59,341
59,160
58,876
58,560
58,216
57,857
57,371
57,143
56,400
56,290
56,154
55,629
Total
Avifauna
0.42
1.09
0.47
0.50
0.54
0.45
0.58
0.82
0.56
0.46
0.40
0.49
0.61
0.99
0.55
0.90
0.65
0.68
1.09
0.57
0.64
0.58
0.60
0.44
1.04
0.66
0.57
0.62
0.60
Species/100 mi2
Land Avifauna-
Total
(TL)
0.28
0.93
0.38
0.31
0.47
0.29
0.43
0.64
0.39
0.30
0.29
0.28
0.46
0.83
0.46
0.72
0.45
0.51
0.48
0.44
0.91
0.34
0.37
0.37
0.55
0.35
0.90
0.41
0.36
0.38
0.36
NL+(TL-NL)
0.17
0.87
0.34
0.29
0.47
0.17
0.36
0.60
0.25
0.19
0.27
0.21
0.42
0.74
0.45
0.68
0.39
0.43
0.36
0.28
0.82
0.24
0.26
0.25
0.54
0.35
0.82
0.26
0.22
0.26
0.29
Native
(NL)
0.057
0.80
0.31
0.27
0.47
0.048
0.30
0.56
0.12
0.077
0.25
0.14
0.37
0.66
0.45
0.63
0.34
0.36
0.24
0.12
0.73
0.13
0.15
0.13
0.54
0.35
0.75
0.10
0.080
0.13
0.23
Passerines:Nonpasserines
Total
(TL)
2.58
1.55
1.33
1.60
1.60
2.51
1.97
1.23
2.62
2.68
0.90
1.76
1.26
1.44
1.12
1.49
1.27
1.42
2.10
1.92
1.44
2.96
2.47
2.80
1.79
1.44
1.47
2.75
2.69
2.72
2.33
.and Avi faun:
NL+(TL-NL)
2.09
1.47
1.23
1.58
1.59
2.04
1.80
1.22
2.08
2.16
0.87
1.66
1.16
1.30
1.14
1.47
1.17
1.24
2.03
1.79
1.32
2.38
1.84
2.32
1.76
1.44
1.36
2.20
2.07
2.16
2.21
l
Native
(NL)
0.83
1.39
1.12
1.55
1.58
0.68
1.59
1.20
1.10
1.04
0.84
1.48
1.04
1.16
1.16
1.44
1.04
1.03
1.92
1.40
1.19
1.43
0.89
1.42
1.73
1.44
1.23
1.02
0.75
1.19
2.02
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Continued
Surinam
Arkansas
North Carolina
Alabama
Guatemala-British Honduras
New York State
Java
Louisiana
Southern Central America
(Costa Rica, Panama)
Tunisia
Mississippi
Greece
Eritrea
Pennsylvania
Cuba
Liberia
Bulgaria
Newfoundland Island
Guatemala
Ohio
Virginia
Kentucky
Pemambuco, Brazil
Tenasserim, Burma
Nyasaland
Oaxaca, Mexico
Indiana
Hungary
Natal-Zululand, South Africa
Zaria, Nigeria
French Guiana
Area
(mi5)
55,143
53,104
52,712
51,609
50,732
49,576
48,830
48,523
48,462
48,300
47,716
46,948
45,754
45,333
44,164
43,000
42,796
42,734
42,044
41,222
40,815
40,395
38,315
37,614
37,374
36,371
36,291
35,918
35,284
35,000
34,740
Total
Avifauna
1.19
0.61
0.67
0.66
1.34
0.77
0.92
0.75
1.93
0.68
0.66
0.72
1.15
0.79
1.00
0.72
0.50
1.57
0.82
0.82
0.74
0.75
1.57
1.63
1.60
0.88
0.92
1.68
0.95
1.79
Species/100 mi2
Land Avifauna
Total
(TL)
0.99
0.41
0.38
0.40
1.08
0.42
0.68
0.46
1.63
0.42
0.41
0.45
0.87
0.47
0.39
0.84
0.47
0.23
1.28
0.51
0.48
0.48
0.67
1.31
1.31
1.37
0.55
0.55
1.24
0.79
1.52
NL+(TL-NL)
0.97
0.26
0.31
0.27
0.98
0.29
0.64
0.30
1.53
0.33
0.27
0.35
0.77
0.32
0.28
0.80
0.35
0.15
1.15
0.32
0.33
0.31
0.66
1.23
1.24
1.26
0.36
0.39
1.15
0.68
1.50
Native
(NL)
0.95
0.12
0.14
0.14
0.87
0.15
0.60
0.13
1.42
0.23
0.14
0.24
0.67
0.16
0.17
0.76
0.22
0.073
1.02
0.14
0.17
0.15
0.66
1.15
1.16
1.15
0.15
0.22
1.06
0.56
1.48
Passerines:Nonpasserines
Total
(TL)
1.36
2.78
2.90
3.00
1.54
2.84
1.25
2.44
1.56
1.64
3.02
1.65
1.26
2.86
1.81
1.36
1.58
2.33
1.56
2.67
2.92
2.82
1.74
1.38
1.46
1.67
2.88
1.52
1.30
1.25
1.35
.and Avifauna
NL+(TL-NL)
1.35
2.19
2.35
2.37
1.40
2.31
1.25
2.03
1.48
1.63
2.31
1.63
1.22
2.25
1.30
1.35
1.50
2.08
1.42
2.20
2.26
2.12
1.74
1.32
1.47
1.56
2.11
1.51
1.33
1.36
1.35
Native
(NL)
1.34
1.08
1.45
1.34
1.25
1.39
1.26
1.12
1.40
1.59
1.15
1.61
1.16
1.20
0.64
1.33
1.33
1.48
1.27
1.19
1.22
0.93
1.73
1.24
1.50
1.44
0.84
1.45
1.37
1.52
1.34
NUMBER 212
TABLE 1.?Continued
Portugal
Maine
R1o de Janeiro-Espfrito Santo, Brazi l
Meta, Colombia
Moluccas
Ireland
Santa Catarina, Brazi l
South Carolina
Tamaulipas, Mexico
Hokkaido, Japan
H1span1o1a
Sabah (North Borneo)
Panama
Chiapas, Mexico
Veracruz, Mexico
New Brunswick, Canada
Banks Is land, Canada
Tasmania
Ceylon
Northern Thailand
I t u r i , Congo
West V i rg in ia
Yucatan State, Mexico
Sinaloa, Mexico
Paraiba, Brazi l
Nova Scotia, Canada
Campeche, Mexico
Quintana Roo, Mexico
Costa Rica
Southampton Island, Canada
Is la Grande (Tierra del Fuego)
Espi'rito Santo, Brazi l
Area
fmi2)
34,240
33,215
32,915
32,903
32,300
31,840
31,118
31,055
30,731
30,300
29,979
29,388
29,224
28,729
27,736
27,473
26,400
26,304
25,332
25,000
24,840
24,181
23,926
22,580
21,251
20,743
19,670
19,438
19,238
19,100
18,530
16,543
Total
Avifauna
0.93
1.04
1.86
1.02
1.07
0.86
1.09
1.44
1.04
1.69
2.92
2.18
2.26
1.19
0.22
0.80
1.50
2.12
2.68
1.19
1.30
1.68
0.97
1.55
1.53
1.73
4.05
0.31
0.86
2.56
Spec1es/l00 n>12
Land Avifauna
Total
(TL)
0.54
0.56
1.56
1.44
0.69
0.55
0.73
0.64
1.08
0.51
0.37
1.34
2.45
1.87
1.85
0.66
0.068
0.40
0.98
1.81
2.36
0.77
0.98
1.22
0.91
0.83
1.33
1.39
3.44
0.094
0.41
2.32
NH-(TL-NL)
0.42
0.36
1.55
1.40
0.66
0.37
0.73
0.45
0.89
0.34
0.32
1.26
2.29
1.69
1.67
0.41
0.043
0.33
0.86
1.61
2.28
0.52
0.83
1.06
0.91
0.52
1.17
1.23
3.20
0.058
0.29
2.32
Native
(NL)
0.30
0.17
1.54
1.36
0.62
0.19
0.73
0.26
0.71
0.17
0.26
1.19
2.13
1.51
1.49
0.15
0.017
0,26
0.74
1.40
2.20
0.27
0.67
0.90
0.90
0.21
1.01
1.07
2.95
0.021
0.17
2.32
Passerines:Nonpasser1nes
Total
(TL)
1.71
2.76
1.60
1.50
0.75
2.12
1.65
2.64
1.64
1.82
1.15
1.36
1.59
1.51
1.70
2.50
1.57
1.12
0.98
1.72
1.76
3.06
1.53
1.74
1.97
2.44
1.46
1.44
1.53
2.00
1.34
2.01
Land Avifauna
NL+(TL-NL)
1.67
2.23
1.60
1.44
0.75
2.23
1.65
2.12
1.46
1.67
1.03
1.33
1.52
1.38
1.59
2.18
1.14
1.29
0.92
1.61
1.76
2.50
1.32
1.56
1.97
2.11
1.28
1.26
1.45
1.44
1.21
2.00
Native
(NL)
1.61
1.20
1.61
1.37
0.74
2.54
1.65
1.30
1.22
1.31
0.88
1.29
1.43
1.24
1.46
1.28
0.29
1.59
0.85
1.48
1.76
1.48
1.06
1.35
1.96
1.24
1.08
1.06
1.36
0.33
0.94
1.99
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Continued
Area
(mi2)
16,515
16,372
15,052
15,035
13,843
13,808
13,433
13,094
13,000
12,500
11,506
11,031
10,747
Total
Avifauna
1.79
3.28
2.08
2.53
2.59
2.72
2.61
2.92
4.39
2.20
3.68
Spedes/100 mi2
1
Total
(TL)
1.16
2.74
1.17
1.88
2.26
1.60
1.46
0.96
1.58
2.42
3.57
1.89
2.83
.and Avifauna
NL-KTL-NL)
0.72
2.72
1.15
1.71
2.00
1.25
0.97
0.92
1.33
2.36
3.22
1.89
2.71
Native
{NLi
0.29
2.70
1.14
1.54
1.74
0.91
0.48
0.88
1.08
2.30
3.13
1.89
2.59
Passerines:Nonpasserines
Total
(TL)
2.98
1.56
1.71
1.04
1.68
2.17
1.88
1.29
1.45
1.34
1.36
1.90
1.34
Land Avifauna
NL+(TL-NL)
2.32
1.56
1.76
0.97
1.45
2.21
1.87
1.34
1.35
1.33
1.30
1.90
1.38
Native
(NL)
1.00
1.55
1.80
0.89
1.21
2.23
1.83
1.40
1.20
1.31
1.23
1.90
1.43
Southern Illinois
Rio de Janeiro, Brazil
Lima Department, Peru
Kerala, India
Peteh, Guatemala
Taiwan
Netherlands
Timor, East Indies
Hainan Island, China
Mushie Territory, Congo
Misiones, Argentina
Alagoas, Brazil
Burundi
Maryland (including District of
Columbia)
Rwanda
Tabasco, Mexico
New Hampshire
Cundinamarca, Colombia
British Honduras
Kutch, India
El Salvador.
Massachusetts
New Jersey
Tranninh, Laos
Halmahera, Moluccas
Ceram, Moluccas
New Caledonia
Hawaiian Islands
Black Hi l ls , South Dakota-Wyoming
Sumbawa, East Indies
Palawan Group, Philippines
10,646 3.13 1.86 1.33 0.80 3.08 2.40 1.48
10,166
9,782
9,304
9,108
8,688
8,461
8,268
8,257
7,836
7,500
6,928
6,621
6,531
6,435
6,000
5,693
5,690
5.15
3.10
6.88
5.41
3.25
4.80
4.76
4.61
6.35
1.69
2.10
3.62
4.24
2.56
1.82
6.06
4.24
2.09
3.98
2.66
2.62
5.60
1.67
1.60
0.87
0.59
2.86
1.87
2.76
4.10
2.28
1.27
5.81
3.71
1.71
3.48
1.70
1.72
5.03
1.57
1.56
0.82
0.56
1.81
1.81
2.49
3.96
2.00
0.70
5.57
3.19
1.34
2.99
0.74
0.82
4.45
1.47
1.51
0.77
0.53
0.75
1.76
2.22
1.27
1.59
2.83
1.79
1.54
1.24
1.47
2.86
3.02
1.90
0.68
0.91
0.87
5.33
2.66
1.10
0.92
1.28
1.42
2.48
1.75
1.37
1.11
1.30
2.36
2.40
1.83
0.64
0.89
0.89
8.00
2.24
1.15
0.87
1.28
1.23
1.80
1.72
1.18
0.94
1.11
1.30
1.27
1.75
0.59
0.86
0.92
16.0
1.25
1.17
0.83
NUMBER 212 99
TABLE 1.?Continued
Mores, East Indies
Connecticut
Falkland Islands
West Sumatra Islands
Jamaica
Bahama Islands
Sumba, East Indies
San Diego County, California
Puntarenas Province, Costa R1ca
Guanacaste Province, Costa Rica
Alajuela Province, Costa Rica
Cyprus
Puerto R1co
Buru, Moluccas
Aru Islands, New Guinea
Galapagos Archipelago
Death Valley, California
Canary Islands
Leyte, Philippines
Sikkim
Gambia Valley
Andaman Islands, Bay of Bengal
Lesser Antilles
Prince Edward Island, Canada
Bali, East Indies
Delaware
San Jose Province, Costa R1ca
Sula Islands, Molucca Sea
Trinidad, West Indies
Lombok, East Indies
Cape Verde Islands
Area
(mi2)
5,509
5,009
4,618
4,600
4,450
4,400
4,305
4,258
4,210
4,020
3,670
3,572
3,423
3,400
3,305
3,029
2,892
2,807
2,785
2,745
2,500
2,500
2,420
2,184
2,146
2,057
2,010
1,872
1,864
1,825
1,557
Total
Avifauna
6.24
2.71
4.41
6.44
9.61
9.41
5.61
7.81
7.19
6.45
15.2
12.0
6.88
9.56
8.80
16.2
20.7
7.56
6.36
Spec1es/l00 ml2
Land Avifauna
Total
(TL)
2.48
3.72
0.82
3.74
2.65
3.66
2.16
5.70
11.9
10.1
13.3
5.85
2.69
3.18
4.35
1.02
5.32
4.02
5.31
14.7
9.44
4.56
4.84
5.62
7.36
9.39
26.4
3.68
14.7
6.08
2.50
NL+(TL-NL)
2.35
2.48
0.62
3.51
2.10
2.34
2.08
4.19
11.1
9.29
12.4
3.61
2.20
2.99
4.25
0.89
2.82
2.88
5.08
14.1
8.60
3.71
4.05
3.38
7.17
6.01
24.4
3.42
13.8
5.86
2.06
Native
(NL)
2.22
1.24
0.41
3.28
1.55
1.02
2.00
2.68
10.2
8.46
11.5
1.37
1.72
2.80
4.15
0.77
0.31
1.75
4.84
13.5
7.75
2.86
3.26
1.14
6.98
2.62
22.5
3.15
12.9
5.65
1.61
Passerines:Nonpasserines
Total
(TL)
1.11
2.87
1.24
1.05
1.68
3.35
0.98
2.80
1.59
1.45
1.64
1.82
1.57
0.74
1.08
2.88
3.28
1.51
1.00
2.82
1.09
0.97
1.60
2.51
1.16
3.20
1.52
0.77
1.20
0.98
0.86
Land Avifauna
NL+(TL-NL)
1.14
2.30
1.11
1.02
1.36
2.55
0.95
2.40
1.53
1.37
1.55
1.80
1.24
0.72
1.11
3.16
2.97
1.32
0.93
2.84
1.08
0.84
1.40
2.29
1.16
2.45
1.46
0.74
1.14
0.98
0.88
Native
(NL)
1.18
1.30
0.90
0.99
0.94
1.13
0.91
1.78
1.46
1.27
1.46
1.72
0.86
0.70
1.13
3.60
0.80
0.96
0.85
2.86
1.06
0.66
1.14
1.50
1.15
1.10
1.39
0.70
1.09
0.98
0.92
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Continued
Socotra, Indian Ocean
Southwestern Ohio
Atlantico, Colombia
Micronesia
Heredia Province, Costa Rica
Big Bend National Park, Texas
Joshua Tree Monument, California
Cartago Province, Costa Rica
Reunion, Indian Ocean
Biak Island, New Guinea
Japen Island, New Guinea
Azores
Natuna Islands, South China Sea
Comoro Islands, Indian Ocean
Mauritius, Indian Ocean
Nicobar Islands, Bay of Bengal
Santa Elena Peninsula, Ecuador
Isia Margarita, Venezuela
Bwamba County, W. Uganda
Hong Kong Colony
Panama Canal Zone (land area)
Rancho Grande, Venezuela
Dominica, Lesser Antilles
Madeira Islands
Ibadan, Nigeria
Darjeeling area, India
Curasao, Netherlands Antilles
Seychelles, Indian Ocean
Impenetrable Forest area, Uganda
Cozumel Island, Mexico
Area
(mi2)
1,400
1,400
1,340
1,335
1,120
1,107
1,025
1,020
970
948
936
888
815
740
720
635
475
444
400
391
372
325
305
302
300
275
173
156
150
125
Total
Avifauna
6.36
20.9
21.4
14.9
31.7
16.2
4.85
19.5
15.1
10.4
8.05
19.5
29.3
106.
89.0
114.
156.
34.1
67.5
94.3
163.
68.8
35.3
189.
Spedes/100 mi2
Land Avifauna
Total
(TL)
2.93
13.5
15.8
4.27
42.2
24.8
14.7
48.3
1.96
6.96
12.8
5.29
12.8
6.75
2.64
12.6
16.2
18.7
95.0
56.2
94.6
134.
20.0
31.8
81.6
152.
34.1
13.5
183.
80.6
NL+(TL-NL)
2.50
8.33
15.2
3.63
39.1
15.3
~9.5
44.8
1.76
6.85
12.7
3.43
12.6
6.55
2.29
9.85
14.5
18.0
92.1
35.7
86.7
129.
16.7
19.7
74.4
130.
25.4
11.5
181.
61.9
Native
(NL)
2.07
3.18
14.5
3.07
36.0
6.15
4.20
41.3
1.55
6.75
12.6
1.58
12.4
6.35
1.94
7.08
12.8
17.4
89.3
15.1
78.8
123.
13.4
7.61
67.3
108.
16.8
9.61
179.
43.2
Passerines:Nonpasserines
I
Total
(TL)
1.16
2.94
1.14
1.60
1.64
2.48
2.98
1.75
0.61
0.61
0.97
0.96
1.12
1.00
0.58
0.82
1.19
0.89
1.66
2.01
1.71
1.62
1.54
1.91
1.40
2.16
1.36
0.91
2.20
2.11
.and Avifauna
NL+(TL-NL)
1.11
2.31
1.09
1.38
1.56
2.30
2.50
1.67
0.74
0.63
0.95
1.10
1.09
1.06
0.74
0.76
1.12
0.91
1.65
1.89
1.59
1.56
1.37
1.77
1.45
2.20
1.20
1.00
2.20
1.72
Native
(NL)
1.07
0.98
1.04
1.49
1.48
1.72
1.46
1.59
0.93
0.65
0.93
1.80
1.06
1.14
1.00
0.67
1.05
0.93
1.64
1.51
1.46
1.49
1.16
1.30
1.51
2.28
0.93
1.14
2.20
1.00
NUMBER 212 101
TABLE 2.?Numbers and percentages of land bird migrants in Middle America
(and adjacent southwestern United States) and in South America
California
Texas
New Mexico
Arizona
Mexico
Baj a California
Tamaulipas
Sonora
Sinaloa
Oaxaca
Veracruz
Chiapas
Tabasco
Yucatan Peninsula
Yucatan State
Campeche
Quintana Roo
Cozumel Island
Central America
British Honduras
Gust eroala
Peten
Tikal
Honduras (1968)
(1955)
(1932)
El Salvador
Nicaragua
Costa Rica (1970)
(19&)
" (1910)
Finca La Selva (1970
(I960
(1957
Total
Non-
asserines
83
95
86
93
286
60
125
101
101
187
190
213
97
132
93
106
111
34
353
145
2l4
117
89
222
192
138
133
207
262
259
235
125
116
83
Land Spp.
Passerines
216
236
212
219
4 90
140
205
199
175
312
324
321
154
188
142
155
160
76
566
223
333
196
159
319
287
209
196
305
Uoo
393
372
199
195
153
Total
299
331
298
312
776
200
330
300
276
1+99
514
53^
251
320
235
261
271
no
919
368
547
313
248
541
479
347
329
512
662
652
607
324
3H
236
Migrant
Non-
passerines
23
34
32
41
22
18
27
20
15
IB
22
21
9
20
15
11
10
8
19
18
23
8
10
25
15
8
16
15
22
21
16
14
9
3
Land Spp.
Passerines
120
136
119
115
103
55
85
70
58
68
79
82
46
73
59
52
53
49
93
73
91
64
61
84
73
44
66
69
72
69
56
51
52
32
Total
143
170
151
156
125
73
112
90
73
86
101
103
55
93
74
63
63
57
112
91
114
72
71
109
88
52
82
84
94
90
72
65
61
35
<f> Migrants/Total Land
Non-
passerines
27.7
35.8
37.2
44.1
7.7
30.0
21.6
19.8
14.9
9.6
11.6
9.9
9.3
15.1
16.1
10.4
9-1
23.6
5.4
12.4
10.7
6.8
11.2
11.3
7.8
5.8
12.0
7.3
8.4
8.1
6.8
11.2
7.8
3.6
Passerines
55.6
57.6
56.1
52.5
21.0
39.3
41.5
35.2
33.2
21.8
24.4
25.6
29.9
38.8
41.5
33.6
33.1
64.5
16.4
32.8
27.3
32.6
38.4
26.4
25.4
21.0
33.7
22.6
18.0
17.6
15.1
25.6
26.7
20.9
Spp.
Total
47.8
51.3
50.6
50.0
16.1
36.5
34.0
30.0
26.4
17.2
19.7
19.3
21.9
29.1
31.5
24.2
23.3
51.8
12.2
24.8
20.8
23.0
28.6
20.2
18.U
15.0
24.9
16.4
14.2
13.8
11.9
20.1
19.6
14.8
102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.?Continued
Panama (1970)
" (1935)
Canal Zone (1970)
(1928)
Barro Colorado I.
(1970)
(1952)
" (1938)
(1929)
Colombia
Santa Marta area
Atlantico
Cundinamarca
Venezuela
Rancho Grande
Perija
Trinidad
The Guianas
Guyana
Surinam
French Guiana
Ecuador
Peru
Bolivia
Brazil
Southern South America
Paraguay
Uruguay
Argentina
Misiones
Chile
Isla Grande (Tierra
del Fuego)
Falkland Islands
Total
Non-
passerines
276
261
130
126
122
110
90
73
546
170
99
198
1*30
167
160
125
269
257
233
224
458
465
375
487
283
213
88
265
175
74
32
17
Land Spp.
Passerines
44o
1+23
222
209
185
163
138
119
879
286
113
354
721
271
21*5
ll*9
399
365
316
302
71*2
869
667
876
1*87
322
139
452
238
127
43
21
Total
716
684
352
335
307
273
228
192
1425
456
212
552
1151
438
405
274
668
622
549
526
1200
1334
1042
1363
770
535
227
717
413
201
75
38
Migrant
Non-
passerines
20
17
11
8
10
5
5
4
14
5
4
11
12
5
3
10
10
7
9
4
10
11
7
11
6
11
11
8
13
5
16
7
Land Spp.
Passerines
74
60
48
42
52
39
28
21
60
42
14
33
49
29
28
24
23
22
16
7
29
32
31
29
8
19
20
12
38
20
28
12
Total
94
76
59
50
62
44
33
25
74
47
18
44
61
34
31
34
33
29
25
11
39
43
38
40
14
30
31
20
51
25
44
19
% Migrants/Total Land
Non-
passerines
7.3
6.5
8.5
6.4
8.2
4.5
5.6
5.5
2.6
2.9
4.0
5.6
2.8
3.0
1.9
8.0
3.7
2.7
3.9
1.8
2.2
2.4
1-9
2.3
2.1
5.2
12.5
3.0
7.4
6.8
50.0
41.2
Passerines
16.8
14.2
21.6
20.1
28.1
24.0
20.3
17-7
6.8
14.7
12.4
9.3
6.8
10.7
11.4
16.1
5.8
6.0
5.1
2.3
3.9
3.7
4.7
3.3
1.6
5.9
14.4
2.7
16.0
15.7
65.1
57.2
Spp.
Total
13.1
11.1
16.8
14.9
20.2
16.1
14.5
13.0
5.2
10.3
8.5
8.0
5.3
7.8
7.7
12.4
4.9
4.7
4.6
2.1
3.25
3.2
3.6
2.9
1.8
5.6
13.7
2.8
12.4
12.4
58.6
50.0
NUMBER 212 103
TABLE 3.?Passerine-nonpasserine and suboscine-oscine ratios in Costa Rica (P =
passerines; N = nonpasserines; S = suboscines; O = oscines; (a):(b) =
passerine-nonpasserine ratio/suboscine-oscine ratio; life zones (formations): TD =
Tropical Dry, TM = Tropical Moist, TW = Tropical Wet, SW = Subtropical
Wet, LMW = Lower Montane Wet, LMR = Lower Montane Rain, MR =
Montane Rain)
Lower Tempisque valley (TD Pacific)
Barranca area (TD Pacific)
El Pel6n de l a Altura (TD Pacific)
Miravalles (TD-SW Pacific)
Los Chiles-Rio Frio (TM Carib.)
La Lola (TM Carib.)
Rinc6n de Osa (TW Pacific)
Finca La Selva (TW Carib.)
Pozo Azul-Pozo Pi ta l (TM Pacific)
Buenos Aires area (TM Pacific)
Boruca-Paso Real area (TM Pacific)
Sixaola area (TM-TW Carib.)
Old Line area (TW Carib.)
El General valley (TM-SW Pacific)
Turrialba-Angostura area (TM-SW Carib.)
Juan Viflas-Tucurriqui (SW Carib.)
Las Mellizas-Cot6n (SW Pacific;
San Vito (SW Pacific)
Bonilla (SW Carib.)
Guayabo (SW Carib.)
Matriculas de Pavones-Silencio de Sit io
Mata (SW Carib.)
Cariblanco (SW Carib.)
Carrillo (SW Carib.)
Monteverde (SW Pacific)
Silencio de Tilaran (SW Pacific-Carib.)
La Hondura (SW-IMR Central Mountains)
Bajos del Toro (SW-IMR Central
Mountains)
Land Avifauna
Total
P:NP
1.19
1.30
1.12
1.55
1.70
1.93
1.48
1.59
1.67
I.69
1.81
1.88
1.83
1.81
1.78
2.18
2.26
2.19
2.36
2.34
2.32
2.28
2.41
2.16
2.42
2.11
2.20
(No. of
Species)
(204)
(186)
(108)
(263)
(200)
(214)
(238)
(324)
(184)
(258)
(233)
(276)
(280)
(286)
(308)
(191)
(188)
(236)
(265)
(214)
(206)
(213)
(225)
(161)
(130)
(146)
(99)
Native
P:NP
0.88
O.96
1.00
1.26
1.46
1.48
1.19
1.33
1.55
1.45
1.56
1.56
1.60
1.45
1.48
1.90
1.93
1.94
1.90
2.07
1.98
2.14
2.17
1.90
2.33
1.96
2.10
(No. of
Species)
(156)
(143)
(100)
(212)
(162)
(171)
(203)
(259)
(176)
(213)
(197)
(220)
(244)
(230)
(245)
(168)
(161)
(212)
(220)
(187)
(176)
(195)
(206)
(1^5)
(120)
(130)
(90)
Passerines
Total
(No. of
S:0 Species)
O.76 (111)
0.72 (105)
1.11 (57)
0.70 (160)
0.68 (126)
0.72 (141)
1.00 (142)
0.81 (199)
1.30 (115)
0.86 (162)
0.81 (150)
0.75 (180)
0.91 (181)
0.79 (184)
O.67 (197)
0.93 (131)
0.88 (130)
0.71 (162)
0.81 (186)
0.79 (150)
0.64 (144)
0.81 (148)
0.85 (159)
0.62 (110)
0.77 (92)
0.62 (99)
0.45 (68)
Native
(No. of
S:0 Species)
1.09 (73)
1.00 (70)
1.00 (50)
0.90 (118)
0.92 (96)
O.96 (102)
1.29 (110)
1.18 (148)
1.38 (107)
1.17 (126)
1.03 (120)
1.03 (134)
1.14 (150)
1.19 (136)
0.92 (146)
1.12 (110)
1.08 (106)
0.77 (140)
0.85 (144)
0.94 (126)
0.80 (117)
0.85 (133)
0.93 (141)
0.73 (95)
0.83 (84)
0.72 (86)
0.49 (61)
(a):(b)
0.81
O.96
1.00
1.40
1.59
1.54
0.92
1.13
1.12
1.24
1.51
1.51
1.40
1.22
1.61
1.71
1.79
2.52
2.24
2.21
2.47
2.53
2.34
2.60
2.82
2.72
4.30
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 3.?Continued
San Jos? de la Montana (IMR Central
Mountains)
La Chonta (LMR Central Mountains)
Rio La Hoja (LMW Central Mountains)
Poas Volcano (LMW-MR Central Mountains)
Villa Mllls-Cerro de la Muerte
(MR Central Mountains)
Land Avifauna
Total
P:NP
2.22
3.14
2.81
1.60
1.74
(No. of
Species)
(71)
(58)
(61)
(65)
(85)
Native
P:NP
2.04
2.72
2.07
1.40
1.59
(No. of
Species)
(64)
(52)
(43)
(60)
(75)
Passerines
Total
(No. of
S:0 Species)
0.36 (49)
0.42 (44)
0.22 (45)
0.33 (40)
0.29 (54)
Native
S:0
0.39
0.4l
0.26
0.35
0.28
(No. of
Species)
(43)
(38)
(29)
(35)
(46)
(a):(b)
5.27
6.68
7.95
4.05
5.70
TABLE 4.?Passerine-nonpasserine and suboscine-oscine ratios in Central America
and in Costa Rica: checklists versus earlier lists or samples (P = passerines; N =
nonpasserines; S = suboscines; O = oscines; (a):(b) = passerine-nonpasserine
ratio/suboscine-oscine ratio; life zones (formation): TD = Tropical Dry, TM =
Tropical Moist, TW = Tropical Wet, SW = Subtropical Wet, LMR = Lower
Montane Rain, MR = Montane Rain)
Guatemala
1970 (Land)
1932a (Griscom)
I879-I904 (Salvin and Godman)
Dearborn (1907)
Honduras (mainland)
1968 (Monroe)
1955 (Ei8enmann)
1932 (Stone)
Panama
1972 (compiled)
1935 (Griscom)
1879-1904 (Salvin and Godman)
Land Avifauna
Total
P:NP
1.56
1.62
1.50
1.76
1.1+4
1.50
1.52
1.59
1.62
1.49
(No. of
Species)
(547)
(528)
(502)
(281)
(541)
(479)
(347)
(716)
(684)
(564)
Native
P:NP
1.27
1.31
1.23
1.40
1.19
1.21
1.23
1.42
1.47
1.39
(No. of
Species)
(433)
(409)
(403)
(226)
(432)
(391)
(299)
(622)
(608)
(513)
Passerines
Total
S:0
0.43
0.44
0.4l
0.42
0.55
0.55
0.56
O.78
0.83
0.80
(No. of
Species)
(333)
(326)
(301)
(179)
(319)
(287)
(209)
(440)
(423)
(337)
Native
S:0
0.51
0.53
0.47
0.47
O.69
O.69
0.68
O.96
0.99
0.86
(No. of
Species)
(242)
(232)
(221)
(132)
(235)
(211+)
(165)
(366)
(363)
(298)
(a):(b)
2.45
2.34
2.60
2.98
1.72
1.77
1.80
1.48
1.48
1.61
NUMBER 212
TABLE 4.?Continued 105
Panama Canal Zone
I968 (Eisenmann and Loftin):
including doubtful and
unusual records.
1968 (Eisenmann and Loftin):
definitely known and
modern presence.
1928 (Sturgis)
Barro Colorado Island
Total
1952 (Eisenmann)
1938 (Chapman)
1929 (Chapman)
P. Slud, March l6-April 7 , 1966
Costa Rica (mainland)
Total
1964 (Slud)
1910 (Carriker)
1879-1904 (Salvin and Godman)
I887 (Zeledon)
I869 (Frantzius)
Sassi (1938-1939)
Ferry (1910)
Monroe, Burt L., J r . (pers. cctnm.)
Taboga, Brosimum woodland area
(TD Pacific)
Total
P. Slud, Jan. 24-28, 30, 1965
El Pelon de l a Altura (TD Pacific)
Total
P. Slud, May 26-30, 1961
Scheelia woodland s i t e , northwest
from Barranca (TD Pacific)
Total
Land Avifauna
Total
P:NP
1.59
1.71
1.66
1.52
1.48
1.53
1.63
1.81
1.53
1.52
1.58
1.54
1.51
1.56
1.48
2.80
1.93
1.51
1.39
1.12
1.21
1.39
(No. of
Species)
(427)
(352)
(335)
(307)
(273)
(228)
(192)
(180)
(662)
(652)
(607)
(539)
(568)
(440)
(226)
(171)
(296)
(153)
(136)
(108)
(93)
(141)
Native
P:NP
1.34
1.46
1.42
1.19
1.18
1.29
1.42
1.51
1.37
1.36
1.44
1.40
1.35
1.39
1.40
2.52
I.69
1.07
1.06
1.00
1.05
0.93
(No. of
Species)
(348)
(293)
(285)
(245)
(229)
(195)
(167)
(143)
(568)
(562)
(535)
(474)
(494)
(384)
(214)
(148)
(256)
(122)
(109)
(100)
(86)
(108)
Passerines
Total
S:0
0.86
O.87
0.94
0.83
0.92
0.94
1.05
0.90
0.71
O.76
0.74
0.74
0.72
0.68
0.65
0.60
0.52
O.92
0.88
1.11
1.32
0.86
(No. of
Species)
(262)
(222)
(209)
(185)
(163)
(138)
(H9)
(116)
(400)
(393)
(372)
(327)
(343)
(268)
(135)
(126)
(195)
(92) *
(79)
157)
(51)
(82)
Native
S:C
1.19
1.20
1.23
1.22
1.30
1.39
1.45
1.46
0.88
0.88
0.88
0.86
0.86
0.81
0.64
0.74
0.58
1.25
1.44
1.00
1.20
1.36
(No. of
Species)
(199)
(174)
(167)
(133)
(124)
(HO)
(98)
(86)
(328)
(324)
(316)
(277)
(285)
(224)
(125)
(106)
(161)
(63)
(56)
(50)
(44)
(52)
(a):(b)
1.12
1.22
1.15
0.97
0.91
0.93
O.98
1.03
1.55
1.54
1.63
1.61
1.57
1.74
2.20
3.42
2.92
0.86
0.74
1.00
O.87
0.68
106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 4.?Continued
P. Slud, Aug. 30-Sept. 2, 1964;
Jan. 18-20, Apr. 20-22, I965
Miravalles (TD-SW Pacific)
Total
P. Slud, Apr. 20-May 5, 1955
Underwood (1896)
Hacienda Santa Maria (TD-SW, Pacific)
Total
Wetmore (1944) plus P. Slud,
Nov. 20-24, 1954
Rincon de Osa (TW Pacific)
Total
P. Slud, Mar. 12-27, 1962;
Oct. 26-31, 1964
Finca La Selva (TW Carib.)
Total to 1970
Total to i960 (Slud)
Total to 1957, P. Slud
4 40-day t o t a l s , 1957-58
Sept.-Oct. 1957
Jan.-Feb. 1958
Mar.-May 1958
July-Sept. 1958
A er e
9 2-week t o t a l s , 1957-58
Sept. 20-Oct. 4, 1957
Jan. 13-27, 1958
Feb. 11-25, 1958
Mar. 17-29, Apr. 3-4, 1958
Apr. 6-20, 1958
Apr. 24-May 9, I958
Land Avifauna
Total
P:NP
1.49
1.55
1.70
1.72
1.58
1.76
1.48
1.45
1.59
1.63
1.80
1.90
1.87
1.86
1.68
2.17
1.92
2.05
1.90
1.76
1.71
(No. of
Species)
(337)
(263)
(193)
(152)
(152)
(124)
(238)
(202)
(324)
(311)
(238)
(249)
(235)
(249)
(217)
(206)
(219)
(198)
(209)
(221)
(217)
Native
P:NP
O.96
1.26
1.34
1.42
1.36
1.60
1.19
1.23
1.33
1.34
1.52
1.55
1.61
1.56
1.57
1 57
1.85
1.65
1.86
1.64
1.57
1.63
(No. of
Species)
(102)
(212)
(159)
(126)
(132)
(109)
(203)
(174)
(259)
(250)
(204)
(199)
(209)
(210)
(203)
(205}
(171)
(196)
(180)
(182)
(193)
(192)
Passerines
Total
S : 0
0.86
0.70
0.70
0.66
0.86
0.88
1.00
1.05
0.81
0.83
0.82
0.81
1.01
0.95
1.23
0.91
1.03
0.99
0.93
1.02
1.11
(No. of
Species)
(82)
(160)
(122)
(96)
(93)
(79)
(142)
(120)
(199)
(193)
(153)
(163)
(153)
(162)
(136)
(141)
(144)
(133)
(137)
(141)
(137)
Native
S:0
1.50
0.90
0.78
0.90
1.11
1.09
1.29
1.23
1.18
1.24
1.12
1.16
1.30
1.27
1.30
1.26
1.18
1.30
1.21
1.22
1.27
1.25
(No. of
Species)
(50)
(118)
(91)
(74)
(76)
(67)
(110)
(96)
(148)
(143)
(123)
(121)
(129)
(128)
(124)
(126)
(HI)
(122)
(117)
(113)
(118)
(119)
(a):(b)
0.64
1.40
1.72
1.58
1.22
1.47
O.92
1.00
1.13
1.07
1.36
1.32
1.24
1.23
1.21
1 25
X . ^~y
1.58
1.27
1.54
1.35
1.24
1.31
NUMBER 212 107
TABLE 4.?Continued
May 31-June H+, 1958
July 13-26, 1958
Aug. 3-9, 13-17, 23-25, 1958
Average
Buenos Aires area (TM Pacific)
Total
P. Slud, Jan. 7-25, 1955
Boruca (TM Pacific)
Total
P. Slud, Feb. 2-6, 1955
Los Chiles-Rio Frio area (TM Carib.)
Total
P. Slud, April 2-8, 1962
La Lola area (TM Carib.)
La Lola, Total
P. Slud, Sept. 11+-21+, 1953
Sixaola area (TM Carib.)
Total
P. Slud, October I95I+
El General valley (TM-SW Pacific)
Total
P. Slud, June 23-July 9, 195^
Monteverde (SW Pacific)
Total
P. Slud, June 17-July 1, 1961
Cariblanco (SW Carib.)
Total
P. Slud, March 23-30, 1955
La Hondura (SW-LMR
Central Mountains)
Land Avifauna
Total
P:NP
1.55
1.61
1.78
I.69
1.90
1.96
2.01+
1.70
1.81+
1.89
1.93
2.26
1.88
2.22
1.81
1.73
2.16
2.02
2.28
2.72
(No. of
Species)
(186)
(188)
(186)
(258)
(217)
(219)
(161+)
(200)
(156)
(23U)
(211+)
(176)
(276)
(21+8)
(286)
(161+)
(161)
(139)
(213)
(160)
Native
P:NP
1.53
1.61
1.70
1.67
1.1+5
1.72
1.66
1.76
1.65
1.1+8
1.1+8
1.79
1.56
1.91
1.1+5
1.70
1.90
1.96
2.11+
2.1+9
(No. of
Species)
(185)
(188)
(181)
(185)
(213)
(183)
(186)
(138)
(162)
(130)
(191)
(171)
(11+8)
(220)
(195)
(230)
(162)
(136)
(195)
(1^3)
Passer ines
Total
(No", of
S:0 Species)
1.31 (113)
1.32 (n6)
1.25 (119)
0.86 (162)
0.88 (11+3)
O.79 (ll+5)
0.75 (no)
0.68 (126)
0.63 (101)
0.7I+ (153)
O.72 (ll+l)
O.72 (122)
0.75 (180)
0.71 (171)
0.79 (18*0
1.21 (10l+)
0.62 (no)
O.69 (93)
0.81 (ll+8)
0.67 (n7)
Native
(No. of
S:0 Species)
1.29 (112)
1.32 (n6)
1.28 (ni+)
1.26 (n6)
1.17 (126)
1.17 (n7)
1.00 (n6)
1.00 (88)
0.92 (96)
0.8U (81)
0.97 (nU)
O.96 (102)
0.86 (95)
1.03 (23k)
0.9^ (128)
1.19 (136)
1.22 (102)
0.73 (95)
O.67 (90)
0.85 (133)
0.70 (102)
(a):(b)
1.20
1.22
1.33
1.21+
1.1+7
1.68
1.76
1.59
1.96
1.53
1.51+
2.08
1.51
2.03
1.22
1.39
2.60
2.93
2.53
3.56
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 4.?Continued
Total
P. Slud, July 9-17, 1955
La Chonta (LMR Central Mountains)
Total
P. Slud, Sept. 9-12, 1964
Villa Mills-Cerro de la Muerte
(MR Central Mountains)
Total
P. Slud, Apr. 26-30, 1951;
Oct. 13-14, 1964
L. L. Wolf, part of Jan. 1967
" " Feb. "
" " Mar. "
" " Apr. "
" May "
" " June "
" Average Jan.-June 1967
" Total " "
Land Avifauna
Total
P:NP
2.11
2.48
3.14
4.00
1.74
2.36
1.87
1.83
1.05
2.12
2.12
2.00
1.78
1.61
(No. of
Species)
(146)
(<*)
(58)
(*5)
(85)
(47)
(43)
(34)
(45)
(50)
(53)
(45)
(45)
(73)
Native
P:NP
1.96
2.44
2.72
3.33
1.59
1.93
1.79
1.75
1.17
1.93
1.88
2.00
1.73
1.54
(No. of
Species)
(130)
(93)
(52)
(39)
(75)
(41)
(39)
(33)
(39)
(44)
(^f-5 ^
(41.5)
(61)
Passerines
Total
S:0
0.62
0.81
0.42
0.39
0.29
O.38
O.27
0.10
0.21
0.31
0.29
0.30
0.25
0.29
(No. of
Species)
(99)
(67)
(44)
(36)
(54)
(33)
(28)
(22)
(23)
(34)
(36)
(30)
(28.8)
(*+5)
Native
S:0
0.72
O.78
0.4l
0.37
0.28
0.29
0.32
0.11
0.23
0.32
0.23
0.30
0.25
0.28
(No. of
Species)
(86)
(66)
(38)
(30)
(46)
(27)
(25)
(21)
(21)
(29)
(32)
(30)
(26.3)
(37)
(a):(b)
2.72
3.11
6.68
9.15
5.70
6.75
5.67
16.8
4.98
6.06
8.14
6.58
6.81
5.58
NUMBER 212 109
TABLE 5.?Passerine-nonpasserine and suboscine-oscine ratios of Costa Rican
localities: forest versus non-forest ((a):(b) = passerine-nonpasserine ratio/
suboscine-oscine ratio; life zones (formations): TD = Tropical Dry, TM =
Tropical Moist, TW = Tropical Wet, SW = Subtropical Wet, SR = Sub-
tropical Rain, LMR = Lower Montane Rain, MR = Montane Rain)
Bagaces si te (TD Pacific): Slud (1965)
Forest
Non-forest
Taboga, Brosimum site (TD Pacific): Slud (1965)
Forest
Non-forest
G. H. Orians and D. R. Paulson (pers. cctnm.)
Forest (Orians, 1969)
Barranca, Scheelia site (TD [* TM (Tosi, I969)]
Pac i f ic ) : Slud (1965)
Forest
Non-forest
Aug. 30-Sept. 2, I961*
Forest
Non-forest
Jan. 18-20, 1965
Forest
Non-forest
Apr. 20-22, I965
Forest
Non-forest
Orians (1969): Forest
Rincon de la Vieja-Guachipilln (TD-SW Pacific):
P. Slud, May 13-20, 1955
Forest
Non-forest
Miravalles (TD-SW Pacific): P. Slud,
Apr. 20-May 5, 1955
Forest
Non-forest
(a)
Native Land Avifauna
Passerines:
Nanpasserines
0.93
1.57
0.7I*
1.06
1.21
0.97
1.09
1.17
O.96
1.17
0.85
1.06
1.29
0.91
1.07
1.1*3
O.89
1.26
1.1*3
1.17
1.26
1.13
1.32
0.96
1.31*
1.58
1.19
(No. of
Species)
(58)
(18)
(1*0)
(105)
(1*2)
(63)
(9?0
(66)
(102)
(39)
(63)
(72)
(32)
(1*0)
(85)
(3*)
(51)
(86)
(31*)
(52)
(61)
(113)
(58)
(55)
(159)
(67)
(92)
(b)
Native Passerines
Suboscines:
Oscines
1.16
0.83
1.1*3
1.1*6
1.88
1.21
1.58
2.27
1.50
2.50
1.07
1.29
2.60
0.73
1.1*1*
2.33
0.85
1.29
1.50
1.15
1.1*3
0.88
1.06
O.69
O.78
1.16
0.56
(No. of
Species)
(28)
( H )
(17)
(5U)
(23)
(3D
(1*9)
(36)
(50)
(21)
(29)
(37)
(18)
(19)
(1*1*)
(20)
(21*)
(1*8)
(20)
(28)
(31*)
(60)
(33)
(27)
(91)
(1*1)
(50)
(a):(b)
0.81
1.88
0.52
0.73
0.61*
0.80
0.68
0.52
0.61*
0.1*7
0.80
0.82
0.50
1.2b
0.71*
0.61
1.05
O.98
0.95
1.02
0.88
1.29
1.25
1.1*0
1.72
1.36
2.12
110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 5.?Continued
Hacienda Sta. Maria (TD-SW Pacific): P. Slud,
Nov. 20-24, 1954
Forest
Non-forest
La Lola (TM Carib.): P. Slud, Sept. 14-24, 1953
Forest
Non-forest
Los Chiles-Rio Frio area (TM Carib.): P. Slud,
Apr. 2-8, 1962
Forest
Non-forest
Sixaola area (TM Carib.): P. Slud, October 1954
Forest
Non-forest
Chase: Oct. 7-9
Forest
Non-forest
Rio Suab (TM-SW): Oct. 10-16
Forest
Non-forest
Volio: Oct. 18-24 1
Forest
Non-forest
Cahuita: Oct. 25-31
Forest
Non-forest
Hacienda Altamira, San Carlos (TM Carib.): Slud (1965)
Forest
Non-forest
Siquirres site (TM [= TW (Tosi, I969)] Carib.):
Slud (I965)
Forest
Non-forest
(a)
Native Land Avifauna
Passerines:
Nonpasserines
1.79
2.75
0.88
1.81
1.32
2.68
I.69
1.47
1.80
1.90
1.82
2.00
2.06
1.28
2.94
1.47
1.87
3.63
1.72
1.36
2.14
1.98
1.68
2.44
1.86
1.66
2.18
2.16
1.83
3.36
(No. of
Species)
(92)
(60)
(32)
(149)
(79)
(70)
(129)
(42)
(87)
(194)
(no)
(84)
(104)
(41)
(63)
(101)
(86)
(15)
(125)
(59)
(66)
(137)
(75)
(62)
(163)
(93)
(70)
(164)
(116)
(48)
(b)
Native Passerines
Suboscines:
Oscines
1.03
1.20
O.67
0.86
O.96
O.76
0.84
O.92
0.81
0.94
1.22
0.75
1.19
2.28
0.88
1.31
1.24
3.00
1.19
1.62
O.96
0.90
1.14
O.69
1.08
1.64
0.66
1.13
1.27
0.85
(No. of
Species)
(59)
(44)
(15)
(96)
(45)
(51)
(81)
(25)
(56)
(127)
(71)
(56)
(70)
(23)
(47)
(60)
(56)
(4)
(79)
(34)
(45)
(91)
(47)
(44)
(106)
(58)
(48)
(112)
(75)
(37)
(a):(b)
1.74
2.29
1.32
2.08
1.38
3.54
2.01
1.60
2.23
2.03
1.49
2.67
1.73
O.56
3.34
1.12
1.51
1.21
1.44
0.84
2.24
2.20
1.48
3.52
1.72
1.01
3.33
1.86
1.44
3.95
NUMBER 212
TABLE 5.?Continued
111
Finca La Selva (TW [? SW (Tosi, I969)] Carib.)
Average of 9 2-week surveys 1957-58, P. Slud
Forest
Non-forest
Slud (I965)
Forest
Non-forest
Orians (1969): Forest
Rinc6n de Osa (TW Pacific)
P. Slud, Mar. 12-27, 1962
Forest
Non-forest
Slud (I965)
Forest
Non-forest
Gromaco, Rio Coto Brus (TM Pacific): P. Slud,
Feb. 15-23, 1962
Forest
Non-forest
La Granja, Tilaran (SW [= TM (Tosi, 1969)]
Pacific-Carib.): P. Slud, Jan. 21-29, 1951*
Forest
Non-forest
Volcan s i te (TM [? SW (Tosi, I969)] Pacific):
Slud (1965)
Forest
Non-forest
Oct. 7-12, 1961*
Forest
Non-forest
Mar. 9-13, 1965
Forest
Non-forest
(a)
Native Land Avifauna
Passerines:
Nonpas serines
1.67
1.63
1.79
1.75
1.72
1.75
1.79
1.25
1.3*
1.15
1.32
1.1*3
1.16
1.1*0
1.54
1.25
2.35
1.91
3.17
x.8l
1 .81+
1.76
2.1*7
2.65
2.25
1.79
1.79
1.79
(No. of
Species)
(185)
(118)
(67)
(150)
(95)
(55)
(61*)
(169)
(96)
(73)
(139)
(85)
(5?0
(156)
(81*)
(72)
(111*)
(6?*)
(50)
(157)
(88)
(69)
(125)
(73)
(52)
(1W)
(81)
(67)
(b)
Native Passerines
Suboscines:
Oscines
1.26
1.50
0.91
1.29
1.61
0.95
2.16
1.19
1.62
0.77
1.39
1.9*
0.8l
1.12
1.55
0.7I*
1.00
1.33
0.73
1.27
1.72
0.83
1.17
1.65
0.72
1.32
1.7I*
0.96
(No. of
Species)
(116)
(73)
(U3)
(95)
(60)
(35)
(1*1)
(<*)
(55)
(39)
(79)
(50)
(29)
(91)
(51)
(1*0)
(80)
(1*2)
(38)
(101)
(57)
(1*1*)
(89)
(53)
(36)
(95)
(52)
0*3)
(a):(b)
1.3"*
1.09
1.97
1.36
1.07
1.85
0.83
1.05
0.83
1.1*9
0.95
0.71*
1.1*3
1.26
0.99
I.69
2.35
1.1*1*
1*.35
1.1*3
1.07
2.11
2.11
1.61
3.11*
1.35
1.03
1.57
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 5.?Continued
Matriculas de Pavones (SW Carib.): P. Slud,
Oct. 2-15, 1953
Forest
Non-forest
Silencio de Sitio Mata (SW Carib.): P. Slud,
July 29-Aug. 1, 1955
Forest
Non-forest
Cariblanco (SW Carib.): P. Slud, Mar. 23-30, 1955
Forest
Non-forest
Virgen del Socorro (SW [? SR (Tosi, 1969)] Carib.):
Slud (1965)
Forest
Non-forest
Silencio de Tilaran (SW Pacific-Carib.): P. Slud,
Feb. 2-8, 1954
Forest
Non-forest
Monteverde (SW Pacific): P. Slud, June 17-July 1, 1961
Forest
Non-forest
Helechales de Potrero Grande (TM-SW Pacific):
P. Slud, July 14-29, 1961
Forest
Non-forest
Rio Cot6n (SW Pacific): Slud (1965)
Forest
Non-forest
La Hondura (SW-LMR Central Mountains): P. Slud,
July 9-17, 1955
Forest
Non-forest
(a )
Native Land Avifauna
Passerines:
Nonpasserines
2.02
2.06
1.93
2.80
2.67
3.10
2.49
2.03
4.10
2.61
2.38
3.40
2.12
1.80
3.00
1.96
1.86
2.11
1.66
2.06
1.21
2.00
2.08
1.78
2.44
2.56
2.22
(No. of
Species)
(3*5)
(104)
(41)
(118)
(77)
(41)
(3*3)
(92)
(51)
(148)
(104)
(44)
(106)
(70)
(36)
(136)
(80)
(56)
(157)
(95)
(62)
(102)
(77)
(25)
(93)
(64)
(29)
(b)
Native Passerines
Suboscines:
Oscines
0.73
1.00
0.29
0.74
0.81
O.63
0.70
0.85
0.52
0.81
O.87
0.70
0.80
1.14
0.42
0.67
0.93
o.4i
1.08
1.67
0.48
1.34
1.74
0.60
O.78
1.09
0.33
(No. of
Species)
(97)
(70)
(27)
(87)
(56)
(31)
(102)
(61)
(41)
(107)
(73)
(34)
(72)
0*5)
(27)
(90)
(52)
(38)
(98)
(64)
(34)
(68)
(52)
(16)
(66)
(46)
(20)
(a):(b)
2.76
2.06
6.75
3.78
3.30
4.90
3.56
2.31
7.88
3.21
2.74
4.86
2.65
1.58
7.12
2.93
2.00
5.18
1.53
1.23
2.53
1.49
1.20
2.97
3 . H
2.35
6.67
NUMBER 212 113
TABLE 5.?Continued
Bajos del Toro (LMR Central Mountains): P. Slud,
Mar. 10-21, 1955
Forest
Non-forest
San Jos? de l a Montafia (LMR Central Mountains):
Slud (1965)
Forest
Non-forest
La Chonta (LMR Central Mountains): Slud (1965)
Forest
Non-forest
Villa Mills - Cerro de l a Muerte (MR Central
Mountains): P. Slud, Apr. 26-30, 1951; Oct. 13-ll*, 19ft
Forest
Non-forest
(a)
Native Land Avifauna
Passerines:
Nonpasserines
2.10
2.00
2.30
2.00
2.78
1.30
3.33
10.0
1.1*3
1.93
2.00
1.86
(No. of
Species)
(90)
(57)
(33)
(57)
(3U)
(23)
(39)
(22)
(17)
(1*1)
(21)
(20)
(b)
Native Passerines
Suboscines:
Oscinea
0.1*9
0.65
0.35
0.36
0.32
0.1*1*
0.37
0.1*3
0.25
O.29
0.17
0.1*1*
(No. of
Species)
(61)
(38)
(23)
(38)
(25)
(13)
(30)
(20)
(10)
(27)
(11*)
(13)
(a):(b)
l*.3O
3.08
6.52
5.60
8.70
2.93
9.15
23.3
5.73
6.75
11.9
l*.19
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 6.?Suboscine-oscine ratios in Amazonian Brazil
Amazonian Brazil (Amazonas-Para)
Lower Amazonia (Para)
Northeastern Para (excluding Bel&n area)
Belem area
Ins t i tu to Agrcndmico do Norte
Lower Rio Tocantins
Baiao area
Lower Rio Xingu
Lower Rio Tapajos
Right bank
Santarem area
Mirituba area
Left bank
Itaituba area
Vila Braga
Obidos area
Faro area
Upper Amazonia (Amazonas)
Lower Rio Madeira
Parintins area
Rosarinho-Borba areas
Rosarinho
Borba
Middle Rio Madeira
Calama area
Itacoatiara area
Manaus
Manacapuru area
Rio Negro
Tatu-S. Gabriel area
Tefe
Tonantins
S. Paulo de Olivenca
Total Pas
Suboscines:
Oscines
1.85
1.87
1.49
1.59
1.90
1.82
1.90
2.15
2.12
2.10
2.00
2.82
2.58
2.51
2.91
2.15
2.08
1.94
2.4l
2.45
2.67
2.37
1.99
2.10
1.70
1.46
2.17
2.06
2.18
2.35
2.49
2.79
serines
(No. of
Species)
(535)
(430)
(182)
(236)
(168)
(217)
(180)
(151)
(350)
(313)
(276)
(153)
(268)
(200)
(168)
(227)
(231)
(462)
(198)
(262)
(165)
(192)
(209)
(158)
(194)
(160)
(184)
(260)
(165)
(164)
(150)
(197)
Native Passerines
Suboscines
Oscines
1.97
1.94
1.51
I.62
1.90
1.81
I.89
2.10
2.16
2.11
2.01
2.80
2.60
2.65
3.05
2.23
2.18
2.09
2.46
2.44
2.62
2.39
2.02
2.06
1.69
1.48
2.18
2.20
2.18
2.32
2.48
2.94
: (No. of
Species)
(516)
(420)
(181)
(231)
(168)
(216)
(179)
(149)
(341)
(305)
(271)
(152)
(263)
(197)
(166)
(223)
(226)
(445)
(197)
(258)
(163)
(190)
(205)
(156)
(188)
(156)
(181)
(250)
(165)
(159)
(146)
(193)
NUMBER 212 115
TABLE 6.?Continued
Western Amazonas (Rio Jurua plus Rio Purus)
Rio Purus
Lower Purus (Jaburu-Nova Olinda-Arima)
Arima
Middle Purus
Canutama-Oco do Mundo
Hyutanaha
Upper Rio Jurua
Joao Pessoa area
Total Passerines
Suboscines
Oscines
2.30
2.09
2.33
2.15
2.33
2.38
2.07
1.99
: (No. of
Species)
(304)
(2&7)
(213)
(167)
(223)
(210)
(267)
(248)
Native Passerines
Suboscines
Oscines
2.37
2.11
2.37
2.22
2.28
2.35
2.08
1.98
: (No. of
Species)
(300)
(283)
(209)
(164)
(220)
(208)
(259)
(241)
TABLE 7.?Suboscine-oscine ratios in Central America and
northern South America
Guatemala (Land, 1970)
Lowlands:
Peten (Smithe, 1966)
Northern Peten (Van Tyne, 1935)
Eastern lowlands (Land, 1963)
Arid in te r ior to 1000 m.
(Land, 1962a)
Highlands:
15OO-3OOO m. (Land, 1970)
Above 1000 m. (Wetmore, 194l)
Sierra de las Minas (Land, 1962b)
Restricted to highlands ( ib id . )
Soloma region, Huehuetenango
(Baepler, 1962)
Honduras (mainland) (Monroe, I968)
Caribbean lowlands (Peters, 1929)
Nicaragua (based on Kisenmann, 1955)
Total
Ratio
0.43
0.59
0.55
0.64
0.4l
0.32
0.26
0.30
?
0.20
0.55
0.70
0.59
Passerines
(No. of spp.)
(333)
(187)
(130)
(177)
(62)
(174)
(92)
(105)
~
(73)
(319)
(102)
(305)
Native
Ratio
0.51
O.78
O.69
0.91
O.36
0.37-
0.31
0.38
0.26
0.15
O.69
1.12
0.73
Passerines
(No. of spp.)
(242)
(123)
(103)
(122)
(34)
(122)
(63)
(73)
(M5)
(235)
(72)
(236)
116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 7.?Continued
Northeastern lowlands (Huber, 1932)
Southeastern lowlands (Nutting, 1884;
Richmond, 1893; Howell, 1957)
Panama (1970, compiled)
Lowlands:
Almirante-Changuinola area
(compiled sources)
Canal Zone, total (Eisenmann and
Loftin, 1968)
Canal Zone, definitely known and
modern presence (ibid.)
Extreme eastern Caribbean
(Griscom, 1932b)
Cana, Darien (Griscom, 1929)
Highlands:
W. Chiriqul, below 5000 ft.
(Eisenmann and Loftin, 1967a)
W. Chiriqui, above 4000 ft. (ibid.)
Chiriqul Volcano, above 5000 ft.
(Blake, 1958)
Cerro Campana (Eisenmann and
Loftin, 1967b)
Colombia (Meyer de Schauensee, 1964)
Lowlands:
Pacific coast, Guapi
(Olivares, 1957-1958)
Southeast (Meyer de Schauensee,
1964)
Meta (ibid.)
Sierra Macarena area
(Blake, 1962)
Caqueta (Meyer de Schauensee,
1964)
Putumayo (ibid.)
Vaupes (compiled sources)
Caribbean slope (Meyer de
Schauensee, 1964)
Depto. Atlantico (Dugand, 1947)
Rio Frio, Magdalena
(Darlington, 1931)
Total
Ratio
0.75
0.72
O.78
0.71
0.86
O.87
1.04
1.55
0.66
0.53
0.57
0.55
1.11
I.29
1.86
1.33
1.43
1.80
1.62
1.69
0.79
O.82
0.73
Passerines
(No. of spp.)
(103)
(165)
(44o)
(169)
(262)
(222)
(151)
(130)
(179)
(144)
(152)
(121)
(868)
(87)
(334)
(284)
(182)
(224)
(144)
(204)
(178)
(113)
(107)
Native
Ratio
0.82
0.94
O.96
1.03
1.19
1.20
1.34
I.67
0.90
O.69
0.72
0.72
1.19
1.38
1.99
1.49
1.70
1.92
1.66
1.76
O.94
O.98
0.91
Passerines
(No. of spp.)
(89)
(126)
(366)
(122)
(199)
(174)
(124)
(123)
(135)
(108)
(124)
(93)
(814)
(81)
(284)
(259)
(167)
(207)
(1*1)
(196)
(155)
(99)
(86)
NUMBER 212 117
TABLE 7.?Continued
Arid upper Magdalena vaj_Ley
(Miller, 1947, 1952)
Highlands (Meyer de Schauensee, 1964)
Cundinamarca (Olivares, I969)
Santa Marta area (Todd and
Carriker, 1922)
San Antonio, altitude 2000 m.
(Miller, 1963)
Venezuela (Phelps and Phelps,
1958, 1963)
Lowlands:
Orinoco region (Cherrie, 1916)
Northeast (Beebe, 1909)
" (Friedmann and Smith,
1950, 1955)
Highlands:
Perijd (Phelps, 1943; Gines et al.,
1953)
Rancho Grande (Schafer and Phelps,
1951*)
Cerro Neblina (Phelps and Phelps,
1965)
Mt. Duida (Chapman, 1931)
Mt. Roraima (ibid.)
Mt. Auyan-tepui:
460 m. level (Gilliard, 194l)
1100 m. level (ibid.)
1500-2200 m. (ibid.)
Upper Mt. Auyan-tepui
(Chapman, 1939)
Ecuador (Meyer de Schauensee, 1966,
1970)
Southwest lowlands (Marchant, 1958)
Peru (Meyer de Schauensee, 1966, 1970)
Lowlands:
Apurlmac valley (Terborgh and
Weske, 1969)
Depto. Lima (Koepcke, 1964)
Total
Ratio
O.76
0.91
0.88
0.85
O.65
1.20
1.34
0.60
0.65
0.95
0.82
O.78
0.85
0.83
1.17
1.02
0.86
O.67
1.32
0.62
1.55
1.27
O.98
Passerines
(No. of spp.)
(102)
(578)
(354)
(286)
(1C7)
(718)
(288)
(56)
(117)
(245)
(271)
(91)
(48)
(97)
(104)
(117)
(26)
(30)
(742)
(42)
(869)
(134)
(Hi)
Native
Ratio
O.89
O.96
0.94
1.02
0.62
1.32
1.45
0.64
0.70
1.11
0.97
0.95
O.92
0.86
1.37
1.09
0.92
0.71
1.36
0.70
1.57
1.27
1.00
Passerines
(No. of spp.)
(85)
(537)
(321)
(244)
(95)
(672)
(277)
(54)
(112)
(217)
(242)
(37)
(46)
(95)
(97)
(113)
(25)
(29)
(713)
(39)
(837)
(134)
(110)
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 7.?Continued
Southwest (Hughes, 1970)
Highlands:
Urubamba valley (Chapman, 1921)
Bolivia (Bond and Meyer de Schauensee,
1942-191+3)
Northern lowlands (Gyldenstolpe,
19W
Primarily highlands (Niethammer,
1953-1956)
Total
Ratio
0.7^
O.92
1.31
1.66
0.97
Passerines
(No. of spp.)
(33)
(227)
(601)
(303)
(225)
Native
Ratio
0.86
0.9lt
1.3*
I.69
0.99
Passerines
(No. of spp.)
(26)
(223)
(593)
(296)
(223)
TABLE 8.?Numbers of land species and (a):(b) values in the neotropics ((a):(b)
= passerine-nonpasserine ratio /suboscine-oscine ratio)
Middle American mainland (Blake, 1953; Eisenmann,
1955)
Mexico (Blake, 1953; Edwards, 1968)
" (Beristain and Laurencio, 1894)
( i b i d . , keyed to Blake, 1953)
Nontropical Mexico (Edwards, 1972)
Highlands ( ib id. )
Northern
Central
Southern
Lowlands (Edwards, 1972)
Northern Pacific slope
Gulf
Central Pacific "
Gulf
Southern Pacific "
Gulf
Baja California (based on Grinnell, 1928)
No.
Total Land
Birds
1145
776
739
619
542
389
282
313
215
297
290
248
355
256
352
200
of Species
Native Land
Birds
1035
651
642
534
427
295
191
220
142
186
169
158
244
191
254
127
(a)
Total Land
Birds
3.76
6.28
5.36
5.54
10.2
11.4
13.6
9.62
8.04
8.14
6.45
4.50
3.27
2.34
2.74
19.4
:(b)
Native Land
Birds
3.00
4.90
4.23
4.40
9.41
10.6
13.0
8.04
6.82
4.69
3.80
2.43
1.90
1.46
1.51
26.6
NUMBER 212 119
TABLE 8.?Continued
Sonora (van Rossem, 1945; Edwards, 1972)
Tamaulipas (based on Edwards, 1972)
Sinaloa (based on Edwards, 1972)
Colima and adjacent Jalisco (Schaldach, 1963)
Oaxaca (Edwards, 1972; Peterson and Chalif, 1973)
Veracruz (Edwards, 1972; Peterson and Chalif, 1973
Tropical Mexico-Central America (based on Edwards,
1972; Eisenmann, 1955)
Tropical Mexico, (Edwards, 1972)
Chiapas (based on Alvarez del Toro, 1964)
Southern Veracruz (Wetmore, 1943)
Tabasco (compiled sources)
Yucatan Peninsula (based on Paynter, 1955)
Yucatan (Paynter, 1955)
Campeche (ibid; Storer, 1961)
Quintana Roo (Paynter, 1955)
Isla Cozumel (based on Paynter, 1955)
Central America (based on Eisenmann,1955)
British Honduras-Guatemala (Russell, 1964; Land,
1970)
British Honduras (Russell, 1964)
Gallon Jug
Guatemala (Land, 1970)
" (Griscom, 1932a, keyed to Eisenmann,
1955)
Peten (Smithe, 1966; Land, 1970)
Northern PetA
Tikal
Caribbean lowlands (Land, 1970)
(Land, 1963)
Arid interior (Land, 1970)
Humid subtropics (ibid.)
Highlands (ibid.)
(Wetmore, 1941)
Sierra de las Minas (Land, 1962)
No. of
Total Land
Birds
300
330
276
279
499
514
944
563
534
239
251
320
235
261
271
111
919
558
368
236
547
528
313
274
248
309
275
126
380
259
141
158
Species
Native Land
Birds
210
218
203
215
413
413
832
445
431
180
196
227
161
198
208
54
807
440
277
180
433
409
241
208
177
239
211
86
285
199
107
120
(a)
Total Land
Birds
10.7
6.98
7.50
6.54
4.54
4.70
2.61
3.98
3.60
3.88
2.63
3.26
4.33
3.16
3.08
8.15
2.60
3.60
3.23
2.46
3.62
3.65
3.04
2.78
2.64
2.51
2.82
3.30
4.55
6.43
7.70
7.10
(b)
Native Land
Birds
8.18
5.75
4.95
5.31
3.-83
3.81
1.93
2.81
2.71
2.11
1.43
1.66
1.98
1.64
1.64
2.42
1.96
2.40
1.85
1.09
2.45
2.34
1.51
1.33
1.08
1.29
1.51
1.95
2.96
4.27
4.45
4.31
120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Continued
No. of Species
Total Land
Birds
Upper elevations
Soloma, Huehuetenango (Baepler, 1962)
El Salvador-Honduras (Rand and Traylor, 1954;
Monroe, 1968)
El Salvador (Rand and Traylor, 1954)
Arid lowlands ( ib id . ; Dickey and van Rossem,
Honduras (Monroe, 1968)
11
 (Stone, 1932)
Humid Caribbean lowlands (Monroe, 1968)
(Peters, J . L., 1929)
Subtropical Caribbean slope (Monroe, 1968)
Pacific " ( ibid.)
Highlands ( ib id . )
Nicaragua-Costa Rica (Eisenmann, 1955; Slud, 1964a)
Nicaragua (based on Eisenmann, 1955)
Northeast (Huber, 1932)
Southeast (Nutting, 1884; Richmond, 1893;
Howell, 1957)
Costa Rica (based on Slud, 1964a)
(Carriker, 1910)
( i b i d . , keyed to Slud. 1964a)
" (Salvin and Godman, 1879-1904)
11
 ( i b i d . , keyed to Slud, 1964a)
(Zeledrfn, 1887)
( i b i d . , keyed to Slud, 1964a)
(Frantzius, 1869)
( i b i d . , keyed to Slud, 1964a)
" (Lawrence, 1868-70; Salvin, 1869, 1870)
(ibid.. , keyed to Slud, 1964a)
Provinces and zones (Carriker, 1910; Slud, 1964a;
compiled sources)
Provinces
Alajuela
Cartago
-
101
556
329
202
541
347
338
167
404
297
214
710
512
172
264
662
636
607
574
539
611
568
476
440
473
440
488
493
Native Land
Birds
73
70
446
247
147
432
299
257
133
321
223
179
611
428
154
218
568
560
535
516
474
533
494
418
384
416
384
421
421
Total Land
Birds
-
13.7
2.70
3.91
2.27
2.62
2.71
2.00
2.24
2.76
3.91
4.97
2.40
2.49
2.00
2.32
2.14
2.20
2.14
2.03
2.05
2.08
2.11
2.14
2.30
2.19
2.31
2.28
2.38
Native Land
Birds
8.25
12.2
1.70
2.35
1.29
1.72
1.80
1.06
1.05
1.79
2.41
3.57
1.73
1.68
1.68
1.46
1.55
1.68
1.63
1.76
1.61
1.55
1.57
1.63
1.74
1.63
1.72
1.72
1.75
NUMBER 212 121
TABLE 8.?Continued
No. of Species
Total Land
Birds
Guanacaste
Heredia
Limon
Puntarenas
San Jose"
Zones
Tropical belt
Subhumid, following map of Holdridge, 1959
Tos1, 1969
Humid
Pacific southwest
Caribbean slope
F1nca La Selva, 1970 (based on Slud, 1960)
, 1958 (Slud, 1960)
, 1957 (Slud, unpublished)
Subtropical belt
Pacific southwest
Caribbean slope
Lower Montane belt
Montane belt
Costa Rica-Panama (based on Slud, 1964a;
Eisenmann, 1955; Wetmore, 1965-1972a)
Panama (based on Eisenmann, 1955;
Wetmore, 1965-1972a)
" (Griscom, 1935)
Almirante (Peters, J . L., 1931; compiled sources)
W. Chiriquf highlands (Eisenmann and Loft in, 1967a)
Below 5000 f t .
Above 4000 f t .
Chiriquf* Volcano (based on Blake, 1958)
(Blake, 1958)
Cerro Campana (based on Eisenmann and Loft in.
1967b)
Canal Zone (Eisenmann and Loft in, 1968:
defini tely known and modern presence)
407
472
360
502
531
-
-
-
-
-
-
324
311
236
-
-
-
-
-
790
716
684
248
382
281
202
296
227
181
352
Native Land
Birds
340
403
293
431
453
443
228
191
410
273
343
259
250
201
437
248
399
201
84
688
622
608
197
324
228
162
260
193
148
293
Total Land
Birds
1.91
2.28
2.16
1.85
2.08
-
-
-
-
-
-
1.97
2.02
2.24
?
-
-
-
-
2.08
2.05
1.95
2.46
2.58
2.67
4.67
2.80
3.58
3.66
1.99
Native Land
Birds
1.32
1.66
1.27
1.39
1.58
1.47
1.12
1.03
1.36
1.13
1.32
1.13
1.07
1.30
1.52
1.43
1.59
2.30
3.38
1.52
1.48
1.48
1.35
1.76
1.61
2.91
2.07
2.49
2.34
1.22
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Continued
No. of Species
Total Land
Birds
Native Land
Birds
Total Land
Birds
1.86
1.78
1.82
1.61
1.63
1.55
1.38
2.40
1.30
1.40
1.43
1.87
Native Land
Birds
1.12
1.15
0.97
0.91
0.93
0.98
0.96
1.23
1.19
1.28
1.23
1.73
" (ibid.: including doubtful and unusual]
(Sturgis, 1928)
Barro Colorado I. (based on Eisenmann, 1952)
" (Eisenmann, 1952)
Barro Colorado I. (Chapman, 1938)
(Chapman, 1929)
Eastern Caribbean lowlands (Griscom, 1932b)
Isi a Coiba (Wetmore, 1957)
South American mainland (Meyer de Schauensee, 1970)
Northwestern South America (ibid.)
Colombia (based on Meyer de Schauensee, 1964)
(Chapman, 1917)
Zones and sectors (Meyer de Schauensee, 1964)
Zones
Tropical
Subtropical
Temperate
Paramo
Sectors
Pacific sector
Guapi (011vares?1957-58)
Central mountain sector
Cundinamarca (Olivares, 1969)
San Antonio (Miller, 1963)
Caribbean sector
Atla'ntico (Dugand, 1947)
Rio Fri'o, Magdalena (Darlington, 1931)
Arid tropical upper Magdalena valley
(Miller, 1947, 1952)
Santa Marta area (based on Todd and Carriker,
1922, and keyed to Meyer de Schauensee, 1970)
Eastern sector
Orinoco area
427
335
307
273
228
192
256
95
2567
2031
1425
1024
168
456
718
544
348
285
245
229
195
167
220
76
2494
1949
1351
990
1149
570
253
40
419
144
896
552
165
332
212
195
1082
532
230
37
384
137
843
508
151
305
194
174
151
409
676
509
1.30
2.33
2.41
1.02
1.14
1.19
2.00
2.04
2.84
1.46
1.39
1.69
1.10
2.05
1.92
0.99
0.99
1.06
1.82
1.83
2.76
1.09
1.06
0.98
2.07 1.45
1.
0.
0.
98
96
95
1
0
0
.46
.79
.75
NUMBER 212 123
TABLE 8.?Continued
No. of Species
Meta
Sierra Macarena (Blake, 1962)
Southeast
Vaupe's (Olivares, 1955, 1964a, 1964b;
Olivares and Hernandez, 1962)
Caqueta' (based on Meyer de Schauensee, 1964)
Ecuador-Peru (based on Meyer de Schauensee, 1970)
Ecuador (ibid.)
(Chapman, 1926)
Southwest (Marchant, 1958)
Peru (based on Meyer de Schauensee, 1970)
Depto. Lima (Koepcke, 1964)
Southwest (Hughes, 1970)
Northeast (Traylor, 1958)
Apurfmac Valley (Terborgh and Weske, 1969)
Urubamba Valley (Chapman, 1921)
Bolivia (based on Meyer de Schauensee, 1970)
11
 (Bond and Meyer de Schauensee, 1942-43)
Northern lowlands (Gyldenstolpe, 1945b)
Primarily highlands (Niethammer, 1953-56)
Venezuela-The Guianas (based on Meyer de Schauensee
1970)
Venezuela (based on Phelps and Phelps, 1958, 1963)
Tropical Zone
Subtropical Zone
Temperate Zone
Paramo Zone
Orinoco Region (Cherrie, 1916)
Northeast (Beebe, 1909)
(Friedmann and Smith, 1950, 1955)
Central Anzoategui (Smith, F. D., Jr., 1952)
Perija' (Phelps, 1943; Gines et a!., 1953)
Rancho Grande (Scha'fer and Phelps, 1954)
Highlands
Total Land
Birds
1
473
304
551
318
371
1560
1200
1223
76
1334
176
59
282
219
343
1042
916
469
326
1198
1151
868
583
130
27
475
116
224
163
405
438
Native Land
Birds
445
289
523
309
354
1513
1161
1191
70
1291
171
47
276
219
339
1004
907
461
323
1132
1090
822
552
124
24
463
114
218
154
374
404
Total Land
Birds
1.12
1.04
0.89
1.07
0.84
1.23
1.23
1.47
2.01
1.21
1.77
1.73
0.93
1.24
2.15
1.28
1.46
1.10
2.28
1.38
1.41
1.38
1.90
3.12
4.00
1.15
1.56
1.69
1.77
1.62
1.99
Native Land
Birds
0.92
0.81
0.74
0.99
0.72
1.18
1.17
1.27
1.81
1.18
1.80
1.45
0.94
1.24
2.08
1.27
1.41
1.06
2.25
1.19
1.23
1.18
1.70
2.57
2.50
1.03
1.42
1.52
1.60
1.25
1.54
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Continued
No. of Species
Total Land
Birds
Mt. Duida (Chapman, 1931)
Mt. Roraima ( ib id . )
Mt. Auyan-tepui (G i l l i a rd , 1941)
469 m. level
1100 m. level
1500-2200 m. level
Cerro Neblina (Phelps and Phelps, 1965)
The Guianas (based on Meyer de Schauensee, 1970)
Guyana-Surinam
Guyana (based on Snyder, 1966)
(Chubb, 1916-21)
Bartica d is t r ic t (Beebe, 1925; Beebe et a l . , 1917)
Acary Mts. (Blake, 1950)
Surinam-French Guiana (based on Meyer de Schauensee,
1970)
Surinam (based on Haverschmidt, 1968)
" (Bangs and Penard, 1918)
So. Surinam (Blake, 1963)
French Guiana (based on Meyer de Schauensee, 1970)
(Berlepsch, 1908)
Trinidad (based on Herklots, 1961)
11
 (Belcher and Smooker, 1934-37)
No. Brazil-So. Venezuela (Friedmann, 1948)
Brazil (based on Meyer de Schauensee, 1970)
(Pinto, 1938, 1944)
" (Ihering and Ihering, 1907)
Amazonia (based on Meyer de Schauensee, 1970)
Amazonas-Rora1ma-Acre
Amazonas
S. Paulo de Olivenca (compiled sources)
Tonanti'ns ( ib id . )
Rio Jurua-Rio Puru's (Gyldenstoipe, 1945a, 1951)
Upper Jurua-upper Purus ( ib id . )
Rio Jurua (Gyldenstoipe, 1945a)
73
137
284
104
117
34
57
668
654
622
615
411
151
573
549
266
273
526
497
274
252
416
1363
1318
1391
940
836
748
269
216
488
450
394
Native Land
Birds
71
135
276
97
113
33
53
635
621
593
597
394
151
548
524
264
271
515
491
240
223
408
1323
1300
1391
916
816
729
264
212
484
445
387
Total Land
Birds
2.27
2.01
1.42
1.11
1.64
3.78
3.28
1.06
1.07
1.09
1.17
1.01
0.80
0.93
0.89
0.84
0.93
0.83
0.83
1.66
1.49
1.08
1.02
0.90
0.99
0.92
0.95
0.84
0.98
0.93
0.72
0.72
0.80
Native Land
Birds
2.01
2.05
1.22
0.88
1.49
3.39
2.44
0.96
0.94
0.95
1.03
0.86
0.80
0.84
0.81
0.80
0.89
0.80
0.80
1.23
1.25
0.97
0.98
0.85
0.99
0.86
0.86
0.75
0.92
0.94
0.69
0.69
0.77
NUMBER 212 125
TABLE 8.?Continued
No. of Species
Upper Junta (based on Gyldenstolpe, 1945a)
JoSo Pessoa area
Tefe' (compiled sources)
R1o Purus (Gyldenstolpe, 1951)
Lr. Middle Purus
Nova 0l1nda-Ar1ma
Nova OUnda
Arima
Middle Purus
Hyutanaha
Manacapuru (compiled sources)
R1o Negro (1b1d.)
Lower R1o Negro
Upper R1o Negro
Tahuapunto-Vaupes (compiled sources)
Manaus ( ib id. )
Itacoatiara ( ib id . )
Parintins ( ib id. )
R1o Madeira (Hellmayr, 1910)
Lower Madeira (compiled sources)
Borba area
Borba
Igarape" Auara
Rosarinho area
Rosarinho
Middle to upper Madeira (compiled sources)
Calama
Acre (compiled sources)
Roraima (based on Pinto, 1966)
Amazonas-Para (compiled sources)
Lower Amazonia (Griscom and Greenway, 1941)
(Snethlage, 1914)
Para-Amapa (based on Meyer de Schauensee, 1970)
Total Land
Birds
414
386
255
453
314
303
200
240
345
300
285
417
203
243
450
253
298
321
424
433
357
300
207
324
264
338
242
251
379
880
678
947
726
Native Land
Birds
406
379
250
449
310
299
198
237
342
298
281
405
203
243
432
248
292
320
420
428
353
298
206
321
262
333
240
250
371
856
668
932
712
Total Land
Birds
0.88
0.90
0.78
0.83
0.91
1.01
1.06
1.08
0.78
0.98
0.84
0.84
0.90
0.98
0.94
1.19
1.10
0.67
0.71
0.63
0.76
0.75
0.84
0.58
0.63
0.81
0.91
1.00
1.38
0.84
0.79
0.73
0.81
Native Land
Birds
0.85
0.88
0.76
0.81
0.88
0.98
1.06
1.03
0.79
0.98
0.83
0.74
0.90
0.98
0.80
1.15
1.07
0.65
0.68
0.62
0.75
0.74
0.84
0.58
0.63
0.79
0.91
0.98
1.24
0.77
0.76
0.69
0.77
126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Continued
No. of Species
Total Land
Birds
Para ( ib id. )
(Pinto, 1938, 1944)
Faro (compiled sources)
Obidos ( ib id . )
Lower Rio Tapajos ( ibid.)
Right bank
Santaran
(Riker, 1890-91)
Mirituba
Left bank
Itaituba (excluding islands)
Vila Braga
Lower Rio Xingu (compiled sources)
Lower Rio Tocantins ( ibid.)
Baiao area
Extreme n.e. Para ( ibid.)
Bel era (based on Novaes, [MS.])
11
 (Stone, 1928)
Institute) Agronomico do Norte (compiled
sources)
Station "A"
Southeastern Para' (Novaes, 1958)
Amapa (based on Meyer de Schauensee, 1970)
Northeastern Brazil (based on Meyer de Schauensee,
1970)
Maranha'o-Piauf-Ceara' ( ibid.)
(Hellmayr, 1929)
Piauf through Alagoas (based on Meyer de Schauensee,
1970)
Maranhao
Piaui'-Ceara'
Piauf
Ceara*
Ceara'-Paraiba-Alagoas-part of Bahi'a (Pinto and
Camargo, 1961)
716
657
350
358
569
522
453
234
198
418
202
241
229
337
288
437
421
220
291
214
212
351
643
457
440
431
375
319
276
198
274
Native Land
Birds
702
649
345
354
559
513
447
233
197
413
199
239
227
336
287
432
416
219
291
214
212
349
633
453
440
428
372
317
275
198
274
Total Land
Birds
0.80
0.79
0.93
0.81
0.75
0.71
0.78
0.74
1.24
0.70
0.81
0.79
0.90
1.00
0.87
0.73
0.81
1.02
0.72
0.61
1.41
1.08
0.79
0.85
0.90
0.96
0.97
0.99
1.01
1.19
1.28
Native Land
Birds
0.77
0.75
0.87
0.76
0.72
0.69
0.77
0.74
1.20
0.68
0.76
0.75
0.91
1.00
0.88
0.70
0.77
1.01
0.72
0.61
1.41
1.07
0.78
0.86
0.90
0.96
0.96
0.99
1.02
1.19
1.28
NUMBER 212 127
TABLE 8.?Continued
No. of Species
Total Land
Birds
Pernambuco-Alagoas-Paraiba (based on Meyer de
Schauensee, 1970)
Pernambuco
Alagoas
Paraiba (ibid.; Pinto and Camargo, 1961)
Mato Grosso-Goia's (based on Meyer de Schauensee, 1970)
Mato Grosso (ibid.)
11
 (Pinto, 1938, 1944)
" (Naumburg, 1930)
Northeast (Fry, 1970)
(Sick, 1955)
Central sector (Pinto, 1940)
Goia's (based on Meyer de Schauensee, 1970)
" (Pinto, 1938, 1944)
Bahia (based on Meyer de Schauensee, 1970)
Southeastern Brazil (ibid.)
Minas Gerais (ibid.)
11
 (Pinto, 1952)
(Pinto, 1938, 1944)
Espirito Santo-Rio de Janeiro (based on Meyer de
Schauensee, 1970)
Espfrito Santo (ibid.)
(Pinto, 1938, 1944)
Rio de Janeiro (based on Meyer de Schauensee, 1970)
Itatiaya (Holt, 1928; Pinto, 1951; Mitchell, 1957)
SHo Paulo (based on Meyer de Schauensee, 1970)
11
 (Ihering, 1898)
Parana-Sta. Catarina-Rio Grande do Sul (based on
Meyer de Schauensee, 1970)
Parana (ibid.)
(Pinto, 1938, 1944)
Santa Catarina (based on Meyer de Schauensee, 1970)
(Pinto, 1938, 1944)
Rio Grande do Sul (based on Meyer de Schauensee,
1970)
335
255
209
196
723
657
554
584
246
217
215
398
357
521
681
508
479
448
517
388
304
452
249
561
499
515
357
300
228
194
415
Native Land
Birds
333
254
209
195
705
640
543
572
237
217
215
395
357
513
670
505
479
448
511
386
301
446
249
554
497
505
356
300
228
194
401
Total Land
Birds
1.21
1.54
1.63
1.68
0.98
1.06
1.15
1.24
1.23
1.09
1.70
1.10
1.28
0.91
1.05
0.95
0.93
1.08
1.08
1.49
1.51
1.18
1.23
1.05
1.07
1.26
1.27
1.38
1.11
1.35
1.31
Native Land
Birds
1.21
1.51
1.63
1.67
0.97
1.04
1.09
1.18
1.24
1.09
1.70
1.12
1.28
0.89
1.04
0.94
0.93
1.08
1.05
1.45
1.48
1.14
1.23
1.05
1.06
1.25
1.28
1.38
1.11
1.35
1.29
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Continued
No. of Species
Total Land
Birds
Native Land
Birds
Total Land
Birds
Native Land
Birds
(Pinto, 1938, 1944)
So. South America (based on Meyer de Schauensee, 1970)
Paraguay ( ib id. )
" (Laubmann, 1939-40)
Gran Chaco (Laubmann, 1930)
Uruguay (based on Meyer de Schauensee, 1970)
M
 (Cuello and Gerzenstein, 1962)
Argentina-Chile (based on Meyer de Schauensee, 1970)
Argentina ( ib id. )
(Olrog, 1963)
(Dabbene, 1910)
Minus subtropical northeast (based on Olrog, 1963)
Misiones ( ib id . )
So. Patagonia (Olrog, 1959)
Chile (based on Johnson, 1965-67)
Isla Grande, Tierra del Fuego (Humphrey et a l . ,
1970)
326
771
535
484
280
227
213
747
717
689
692
506
413
114
201
75
322
756
505
480
278
196
188
724
697
671
684
492
362
108
176
31
1.41
1.29
1.20
1.23
1.63
1.58
1.79
1.32
1.28
1.30
1.27
1.45
1.18
0.91
1.49
1.17
1.35
1.27
1.21
1.21
1.65
1.80
1.90
1.31
1.28
1.25
1.24
1.38
1.09
0.83
1.26
1.07
TABLE 9.?Paramo "islands" of
ratio; (b) = suboscine-oscine
suboscine-oscine ratio)
Vuilleumier {1970) ((a) = passerine-nonpasserine
ratio; (a):(b) = passerine-nonpasserine ratio/
Paramo "islands" of
Vuilleumier (1970)
1. Ecuador
2. Chiles
3. Las Papas-Coconuco
1+. Sumapaz
5. Tolima-Quindio
6. Paramillo
7. Cocuy
8. Pamplona
9. Cachira
10. Tama
No. of
Species
56
28
25
32
32
11
17
11
13
Ik
Area
(ta2)
3487
326
501
2 0 3 1
989
25
2168
217
11+3
1+6
Distance
from
"island"
no. 1 (km)
1
36
231+
3h3
551
773
801
950
958
995
(a)
1.00
1.55
O.92
I.67
I.67
4.50
1.1+3
1.20
1.17
2.50
(b)
1.55
1.83
1.1+0
1.00
1.50
0.80
2.33
1.00
0.75
0.1+3
(a)-(b)
O.65
0.81+
0.66
1.67
1.11
5.63
0.6l
1.20
1.56
5.83
NUMBER 212 129
TABLE 9.?Continued
11. Batallon
12. Merida
13. Period
lU. Santa Marta
15. Cende
"Island" Groups (grouped by
P. Slud)
1-3, Central Ecuador to southern
Colombia
1+-6, Central Colombia
7-10, Northeastern Colombia
11-12, 15, Venezuela
ll+, Santa Marta
No. of
Species
12
27
1+
17
Ik
57
1+2
29
27
17
Area
(km2)
66
1798
16 7
606
70
1+3H+
301+5
257^
193^
606
Distance
from
"island"
no. 1 (km)
1065
II67
U 8 2
1238
1380
1
5U3
925
U 9 5
U 9 5
(a)
2.00
2.00
1.00
3-25
3.67
l .C*
1.21
1.61+
2.00
3.25
0>)
1.00
0.80
0.00
1.67
0.82
1.1+2
1.09
1.25
0.80
1.17
(a):(b)
2.00
2.50
0.00
2.79
1+.1+0
0.73
l . U
1.31
2.50
2.78
Correlation Coefficients
Paramo "islands"
Distance
Square Root of Distance
log of Distance
Area
Square Root of Area
log of Area
No. of Species
log of Species
"Island" Groups
Distance
Square Root of Distance
log of Distance
Area
Square Root of Area
log Area
No. of Species
log Species
No. of
Species
-0.73
-0.79
-0.83
0.81
0.79
0.72
?
?
-0.97
-0.95
-O.87
0.97
0.93
O.87
?
?
Log of
Species
-0.68
-O.69
-0.65
0.69
0.71
0.68
?
?
-O.92
-O.87
-0.77
0.99
0.97
0.9I+
?
?
(a)
0.1+1+
0.1+1
0.35
-0.35
-0.1+1
-0.53
-0.32
-0.18
0.78
0.69
0.57
-0.95
-O.98
-O.99
-O.87
-0.95
(b)
-0.59
-O.56
-0.1+1+
0.1+9
0.5I+
O.56
0.50
0.63
-0.70
-0.70
-0.68
0.50
0.1+0
0.29
0.52
0.56
(a): 00
0.1+2
0.U1
0.35
-O.38
-0.1+8
-O.63
-0.31
-0.17
0.89
0.79
0.66
-0.93
-O.92
-0.89
-0.88
-O.91
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 10.?Comparative distribution of Formicariidae
Mexico
Chiapas
Yucatan Peninsula
Central America
Br. Honduras-Guatemala
British Honduras
Guatemala
Honduras-El Salvador
Honduras
El Salvador
Nicaragua
Costa Rica
Finca La Selva
Panama
Canal Zone
Barro Colorado I .
Colombia
Vaupes
Atlantico
Cundinamarca
Ecuador-Peru
Ecuador
Peru
Lima
Bolivia
Venezuela
Trinidad
The Guianas
Guyana
Surinam
French Guiana
Brazil
Acre
Alagoas
Area (mi2)
763,9^
28,729
63,031+
228,578
50,732
8,688
1+2,01+1+
67 ,1+20
59,160
8,260
57,H+3
19,238
2
29,221+
372
6
1+39,825
57,857
1,31+0
9,108
583,738 .
101,1+81
1+82,257
15,052
513,086
352,11+1
1,861+
179,363
89,1+80
55,11+3
31+,7^0
3,286,000
57,153
11,031
Species of
Formicariidae
9
9
5
38
11
9
11
19
19
2
21
29
20
37
25
17
128
1+0
6
22
11+1+
109
118
0
69
88
9
53
1+8
!+7
1+7
159
31+
16
i of
Suboscines
10.1
11.8
11.1+
19.1
13.3
15.0
23.1+
19.8
19.8
1+.8
21.2
19.0
25.0
21.2
23.1
23.3
28.5
32.0
12.3
11+.2
21+.8
26.6
23.2
0.0
18.9
23.2
15.3
23.1
23.6
25.2
25.1+
29.0
33.1+
21.6
% of
Passerines
2.3
3.7
1+.3
8.0
4.5
6.0
4.6
8.0
8.1
1.5
8.9
8.9
13.5
10.1+
12.6
12.8
15.7
20.1+
6 .1
6.9
11+.9
15.3
l 4 . i
0.0
10.9
13.1
7.2
ll+.l
ll+.O
15.7
15.9
18.8
21.1+
11.7
% of
Avifauna
1.1+
2 . 1
2.3
4.8
2.5
3 .1
2.6
4.3
4.5
0.8
5.0
5.2
7.9
6.2
7.3
7.5
9.5
13.2
3 . 1
1+.1+
9.6
9.h
9.2
0 .0
6.9
8 . 1
3.8
8.1+
8.2
9 .1
9.2
1 2 . 1
13.8
7.8
Ratio to
Oscines
0.03
0.05
0.07
0.14
0.07
0.10
0.07
0.13
0.11+
0.02.
0.15
0.17
0.29
0.24
0.27
0.28
0.35
0.56
0.12
0.07
0.37
0.36
O.36
0.00
0.26
0.30
0.11+
0.36
0.34
0.1+2
0.1+3
0.53
0.60
0.25
NUMBER 212 131
TABLE 10.?Continued
Amapa
Amazon as
Bahia
Ceard
Esp i r i to Santo
Goias
Maranhao
Mato Grosso
Minas Gerais
Para
Paraiba
Parana
Pernambuco
Piaui
Rio de Janeiro
Rio Grande do Sul
Roraima
Santa Catarina
Sao Paulo
Paraguay
Uruguay
Argentina
Misiones
Chile
Area (mi2)
55,1*89
595,1*71*
215,329
57,371
16,5^3
2Mt,33O
133,67!*
485,1*05
226,179
1+70,752
21,251
82,71*1
38,315
9^,819
16,372
110,150
97,1*38
31,118
95,1*59
157,006
72,172
1,072,750
11,506
286,396
Species of
Formicariidae
kh
92
23
11
37
16
26
53
32
78
11*
21
18
12
35
15
28
18
3k
21
2
23
17
0
I of
Suboscines
31.1*
30.6
13.0
17.2
2I+.8
13.1
18.8
21+.2
17.9
28.2
20.3
15.7
20.9
13.6
22.4
13.0
22.0
21.2
17.1*
12.5
3.6
9-1
16.0
0.0
$ of
Passerines
19.7
20.6
7-8
9.1*
11+A
7 . 1
11.5
14.1
10.8
18.6
10.9
9.2
11.2
7.7
12.9
6.9
12.3
12.7
10.2
6.9
1.8
5.3
8.5
0.0
% of
Avifauna
12.8
12.7
1*.5
5.6
9.7
l*.l
7 . 1
8.3
6.1*
11.2
7.3
6 .0
7 .2
1+.4
7.9
U.3
7.6
8.0
6.2
1+.2
1.0
3.3
M
0.0
Ratio t o
Oscines
0.52
0.6U
0.20
0.21
0.31*
0.16
0.30
0.3!+
0.27
0.55
0.23
0.22
0.21+
0.18
0.30
0.15
0.28
0.32
0.25
0.16
0.03
0.12
0.18
0.00
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 11.?Cumulative daily total of species of land birds collected at Obidos,
Para, Brazil, by 5. M. Klages in 1920-1921
Collecting
Day
1 .
2 .
3 .
4.
5.
6.
7.
8.
9.
10 .
1 1 .
12.
1 3 .
14 .
1 5 .
16 .
17 .
1 8 .
19 .
20.
2 1 .
22 .
2 3 .
24.
2 5 .
26.
27 .
28 .
29.
30.
3 1 .
32 .
Number of
Cumulative
Totals
18
30
kk
55
63
66
77
87
94
103
115
117
123
124
129
133
138
142
146
150
151
154
158
160
162
165
168
169
169
172
173
176
Species
Daily
Increment
18
12
Ik
11
8
3
11
10
7
9
12
2
6
1
5
k
5
k
k
k
1
3
k
2
2
3
3
1
0
3
1
3
Non-
Percentages of Components
passerines Suboscines Oscines
Total
11.1
13.3
13.6
20.0
17.5
18.2
23.it
21.8
20.2
20.lt
22.6
22.2
2it.lt
2lt.2
25.6
26.3
28.2
29.6
28.8
28.7
29.1
29.9
29.8
29.lt
29.6
29.7
30.lt
30.6
30.6
30.6
30.lt
30.5
Total
77.8
73.3
68.2
63.7
66.6
68.1
62.3
63.2
64.9
63.I
59.1
59-8
58.5
58.1
57.lt
56.lt
54.U
53.5
54.1
54.0
53.7
53.3
53.8
54.4
54.3
53.3
52.7
52.3
52.3
51.5
51.8
51.5
Total
11.1
13.4
18.2
16.3
15.9
13.7
14.3
16.0
14.9
16.5
18.3
18.0
17.1
17.7
17.0
17.3
17.4
16.9
17.1
17.3
17.2
16.8
16.4
16.2
16.1
17.0
17.1
17.1
17.1
17.9
17.8
18.0
Suboscines
Passerines
87.5
84.6
79-0
79-5
8O.7
81.9
81.3
8O.9
81.3
79-2
76.5
76.9
77-4
76.6
77.0
76.5
75.8
76.O
76.0
75.7
75.7
75.9
76.6
77.0
77.2
75.9
76.0
76.O
76.4
74.2
74.4
74.0
NUMBER 212 133
TABLE 11.?Continued
Collecting
Day
33.
34.
35.
36.
37.
38.
39-
40.
41.
42.
43.
hk.
45.
46.
?*7.
48.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59-
60.
61.
62.
63.
64.
Number of
Cumulative
Totals
177
180
181
182
182
186
188
190
190
190
192
193
195
195
195
196
200
201
203
204
206
209
210
211
212
212
203
213
213
213
215
216
Species
Daily
Increment
1
3
1
1
0
4
2
2
0
0
2
1
2
0
0
1
4
1
2
1
2
3
1
1
1
0
1
0
0
0
2
1
Percentages of Components
Non-
passerines Suboscines Oscines Suboscines
Total
30.3
30.4
30.2
30.6
30.6
30.0
30.2
30.4
30.4
30.4
30.0
30.4
30.1
30.1
30.1
29.9
29.8
30.0
30.2
30.6
30.8
31.3
31.6
31.4
31.8
31.8
32.1
32.1
32.1
32.1
32.2
32.1
Total
51.2
50.8
51.1
50.8
50.8
51.4
51.4
50.8
50.8
50.8
50.8
50.5
51.0
51.0
51.0
50.8
50.2
50.2
50.2
50.0
49.6
48.3
49.1
49.3
49.1
49.1
48.8
48.8
48.8
48.8
48.9
49.1
Total Passerines
18.5
18.8
16.7
1B.6
18.6
18.6
18.4
18.8
18.8
18.8
19.2
19.1
1B.9
18.9
18.9
19.3
20.0
19.8
19.6
19.4
19.6
20.4
19.3
19.3
19.1
19.1
19.1
19.1
19.1
19.1
18.9
18.8
73.4
73.0
73.2
73.2
73.2
73.3
73.5
72.9
72.9
72.9
72.6
72.6
73.0
73.0
73.0
72.5
71.7
71.9
72.1
72.1
71.5
71.8
71.8
72.0
72.0
72.0
72.0
72.0
72.0
72.0
72.2
72.4
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 11.?Continued
Collecting
Day
65.
66.
67.
68.
69.
70.
71.
72.
73.
7>+.
75.
76.
77.
78.
79-
80.
8 1 .
Number of
Cumulative
Totals
217
217
217
225
235
2*+2
2kk
252
256
261
266
272
273
275
279
281
281
Species
Daily
Increment
1
0
0
8
10
7
2
8
1+
5
5
6
1
2
if
2
0
Percentages
Non-
passerines
Total
32.0
32.0
32.0
31.6
30.6
30.5
30.6
29.7
30.0
29.8
30.4
30.1
30.0
30.1
30.7
31.2
31.2
of Components
Suboscines Oscines
Total
1+8.8
1+8.8
1+8.8
1+8.7
I+8.9
1+9.2
1+8.8
1+9.2
I+8.9
1+9.5
1+8.5
1+8.9
1+9.1
1+9.1
1+8.8
1+8.5
1+8.5
Total
19.2
19.2
19.2
19.7
20.5
20.3
20.6
21.0
21.1
20.7
21.1
21.0
20.9
20.9
20.5
20.3
20.3
Suboscines
Passerines
71.9
71.9
71.9
71.2
70.5
70.8
70.1+
70.0
69.8
70.5
69.7
70.0
70.1
70.3
70.1+
70.1+
70.1+
1-10.
11-20.
21-30.
31-1+0.
1+1-50.
51-60.
61-70.
71-80.
61+
132
163
182
195
209
221
266
10-Day
10.3
U.7
2.2
1.8
1.1
1.2
2.9
3.9
Averages
18.0
26.1
29.9
30.1+
3 0 . 1
31.1+
31.8
30.3
67.1
56.5
53.1
51.2
50.7
1+9.2
1+8.9
1+8.9
15.0
17.1+
17.0
18.5
19.2
19.1+
19.1+
20.8
81.6
76.1+
75.7
73A
72.6
71.9
71.7
70.2
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