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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SMITHSONIA N CONTRIBUTION S TO ZOOLOG Y • NUMBE R 21 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 Contributions 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 numerical 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 substantiated 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 temperature 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 produce 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 environmental information it furnishes. Whether for purposes of comparison or prediction, 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 comparisons 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 distribution 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, differing little from that of other recent authors or

even from one compiled a generation ago by Mayr

(1946), can be accepted as virtually complete. Including 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 private). 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 Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

nonpasserines. This is a global relationship making 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 (onequarter of them aquatic) on the same terrestrial

footing. A second set of relations is that between

oscines and suboscines. This is primarily a neotropical relationship, which is here further strengthened

and substantiated. The two sets of relations are considered 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 considered interdependent, provided the objective is

the acquisition of comparative data.

ACKNOWLEDGMENTS.—The American Museum of

Natural History and the Carnegie Museum permitted me to examine the extensive Amazonian

ornithological materials in their care. At the Smithsonian 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 Vertebrate 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 comments aimed at improving the manuscript.

Species Lists

CATEGORIES

A species list may consist of (1) the entire avifauna 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. Categories (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 treatment 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. Particularly 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 absolutely as many taxa as, if not more than, the moderately to enormously larger area. Obviously, unequalsize 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 different biogeographical regions or are similar if only

in being located somewhere in the global tropics,

occur infrequently in the literature. How infrequently 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 contiguous 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 comparable size when these differ in overall environmental

complexity and converge when they are similar.

ELIMINATION OF WATER BIRDS

Water birds are the Spheniscidae, Gaviidae, Podicipedidae, Procellariiformes, Pelecaniformes, Ciconiiformes, Anseriformes, Gruidae, Aramidae,

Rallidae, Heliornithidae, Eurypygidae, and the

Charadriiformes minus Burhinidae, Glareolidae,

and Thinocoridae. Arbitrary determination of individual 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 water 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 species (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 isolated oceanic islands) and well marked in the cooltemperate region to moderate in the warmtemperate region, low in the subtropical region,

and extremely low in the tropical region; paradoxically, the water bird ratio is usually smaller in

the humid tropics than in the dry tropics. Among
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FIGURE 2.—Scattergram of North and Middle American political divisions: percent water bird

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NUMBER 212

mainland units of approximately the same size, inclusion or exclusion of water birds has little effect

on the total avifauna relative to the total land avifauna: 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 percentages parallel each other in the Old World and

in the New World, not only in size but also in latitudinal 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 America shows that they fall into a distributive pattern

(Figure 2). In the temperate part of the continent,

the lowest percentages are found in arid southwestern 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 MississippiMissouri rivers and the Appalachian Mountains.

The eastern seaboard and to some extent the Pacific sector are distinct from the interior. Much the

highest values occur peripherally at boreal latitudes. The neotropical units, southward from Veracruz 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 procedure, be excluded from consideration when comparing the land birds: terrestrial environments are

best represented by terra firma faunas.

For example, Hall (in Voous, 1972:585) compared the breeding land bird proportion of the

avifauna in Africa with that in the Palearctic region without eliminating the water birds. Hall concluded that Africa is richer in "younger" elements,

that is, the passerines and the nonpredatory arboreal nonpasserines (pigeons through woodpeckers),

where they comprise 62 percent and 18 percent, respectively, 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 nonpasserines (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 increasingly differ. Examples are an oceanic rock

frequented by sea birds and, say, a wood lot supporting 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 montane 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 overwhelmingly 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 analogous areas on the tropical mainland. (See Table 1.)

A total land bird figure can be viewed as a

comparative measure of faunal "richness" or environmental complexity, whether the unit is a

Colombia or an oceanic islet. Nevertheless, multiple factors complicate the interpretation of a single

figure. The figure is uninformative as to the relative "richness" or "poorness" of subdivisions within the unit. It does not distinguish between common 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 residents; 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 natives could increase the species numbers and densities 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 migrants, 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 restricted to a remote bit of land in the ocean has the

same value in a count as one which is cosmopolitan and abundant. The zoogeographic regions,

apart from the Holarctic, have such large land avifaunas that the presence or absence of migrants is

hardly noticeable percentagewise. Migratory effects

are nevertheless present and can affect the comparability 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, climatically, and areally, have migrant percentages which,

if not quite equivalent, occupy similar relative positions 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 extremity, 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. Highlatitude units, having the largest numbers of migrants, would be shifted the most, units in the subtropics 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 sequence, or set of sequences, is also discernible. Islands, 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 Caribbean versant. Large areas, virtually amounting

to subregions, show a progressive reduction southward 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 America 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 unadorned 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

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125

112

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73 KERALA

74 MYSORE

75 BRITISH SOMALILAND

76 PHILIPPINES

77 VERACRUZ

78 GUATEMALA

79 HONDURAS

80 CHIAPAS

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82 NICARAGUA

83 THAILAND

84 CHINA

83 MEXICO

86 SUDAN

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88 CERAM

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4 MEW BRUNSWICK

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7 NEWFOUNDLAND I.

8 KOREA

9 NEBRASKA

10 KANSAS

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12 HOPE I o*

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13 NETHERLANDS

14 PENNSYLVANIA

15 FLORIDA

16 EGYPT

17 GERMANY

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19 ALASKA

20NEW HAMPSHIRE

21 FINLAND

22 BRITISH ISLES

23 ONTARIO

24 MANCHURIA

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27 ANDAMAN IS.

28 BULGARIA

29BRITISH COLUMBIA

30 OREGON

31 MONGOLIA

32 JAPAN

33 WASHINGTON

34 PORTUGAL

35 GREECE

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37 YUGOSLAVIA

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47 CAPE VERDE IS.

48 SOKOTRA

49BAJA CALIFORNIA

50 BALUCHISTAN

51 GREATER ANTILLES

32 HAINAN

53 YUCATAN STATE

54 AFGHANISTAN

55 IRAN

56TAMAULIPAS

57 TIBET

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20 2

NATIV E LAN D BIRD S

FJGUU S.—Influence of migrants relative to numbers of species and location of land avifaunas:

global survey of percent migranu versus size of resident avifaunas. (Large solid drdes with

endosed stars >• the unitt tabulated in the text on page 8; small solid drdes »• mainland

unitt; small open drdes — islands.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

abundance, frequency, environmental impact,

portance," or whatever.

imBaja 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 coefficient 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 migrants tends to be accompanied by a twofold increase in size of avifauna.

Notwithstanding the extraordinarily high coefficient, 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 replaced 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 American isthmus, as do the countries to the south. The

sequence of percentages now becomes 21.0 for

southern Mexico, 21.2 for Guatemala-British Honduras, 19.8 for Honduras-El Salvador, 16.4 for Nicaragua, 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 earlier, 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 increased by 25 to the number in Honduras, the migrant percentage would nearly equal that of Honduras. 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 passerines 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 extraEthiopian units to illustrate "the difference between the land-bird fauna of a tropical area and a

temperate-zone fauna." In the accompanying tabulation, 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 correlates 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 progression and follow it into southern Africa, where the

migrant percentage rises again. On the west, the percentages range from 60 or more in the north (Finland 59, Great Britain 59, Germany 64, Netherlands 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 Somaliland, 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 disparity 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 percentage 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 inclined to believe they complement the residents or

compete with them (Slud, 1960:144-145, 1964b).

The species counts and percentages (Table 2) suggest it is the former, the increasing complexity of

the resident biota which has evolved under increasingly 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 percentages of the total land avifaunas of the sovereign 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, versus all the other orders, collectively termed the

nonpasserines, permeates the ornithological literature. Many faunal works are organized into approximately 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 practical convenience. A second motivation could lie

in the fact that authors generally agree on the sequence and content of the nonpasserine higher

categories but disagree on family limits and arrangements 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 species 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 connotation of ecological opposites. To some the terms

imply an aggressive rivalry between adversaries, to

others they perhaps bring to mind a dynamic balance 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 following 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 nonpasserines, 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, Moreau (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 . . . probably 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 subfamilies), two of land nonpasserines (94 subfamilies), and three of aquatic birds (49 subfamilies). 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 distribution 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 relatively 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 breedingbird censuses, the authors obtained mean-abundance ratios of passerine individuals to nonpasserine individuals, indicating that "at all latitudes

. . . non-passerines are less abundant than passerines." 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 behavior 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 nonpasserine. 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 percentage (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 passerine 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) contrasted the evolution in isolation enjoyed by most

South American groups with the situation in Africa, 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 efficient and competitive, and the avifauna has been

built up as a stratified series of families occurring

together, each of which "tended to become restricted, 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

500F " ^SOUTHERN SOUTH AMERICA

/^SOVIE T UNION

^FRENCH 1ND0-CHINA

NEARCTIC REGION *

J MALAYSIA

/•NONTROPICAL 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 cliavailable ecological opportunities, and ways of life, mate in Africa: "The evolution of steppe, savanare 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 because of increasing aridity." This was seconded by

Keast (in Dobzhansky et al., 1972:272), who

pointed to the "tremendous range of savannahadapted animal groups in Africa where that vegetation 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 Pleistocene and post-Pleistocene (Haffer, 1969; Keast, in

Dobzhansky et al., 1972; Moreau, 1966:10-11; Vanzolini, in Meggers et al., 1973).

Accepting neither climate nor gross ecology as affording 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 humidforest 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 relationships 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 biological combinations, conditions became ripe for

the proliferation of evolutionary series culminating in the diversity manifested in the world today"

(Delevoryas, in Meggers et al., 1973:19).

Nevertheless present-day dimatovegetational factors 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 Zoological divisions of the globe by the small proportion 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 suboscine), 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 registered 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) covers 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 influence 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 continent, but it has made comparisons of natural environments 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 agricultural 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; vestigial forests along the East African coast "have suffered greatly in the last hundred years of 'development,' 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 species, the Neotropical and Ethiopian regions, respectively, 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 rainforest 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 passerine 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 Canadian 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 temperate and boreal North America to around 1.5:1 in

tropical Middle America; makes apparent the relatively large size of the avifauna in southwestern

United States, particularly the large states bordering Mexico; and sets apart the small but avifaunally 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 position, few mountains, paucity of migrants, and fair

to good species compilations for a number of states

and territories. Subdivided into states and territories, 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 insufficient 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 corresponding 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 §".<

\

\ \ \ \

\ *, \ \ \ f \ * . \ \ \

\

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\ * * * if. f\ \ x^

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A sll 3 < f< I n \ X-

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digit Ii|i|§jj jy | 2 j | {JLuihssi jfiX*u sii»:i9L ; \ •»

ilNiEilil£^isiiil§2iiBiiiii3isiiiiiislsi9iiiE§l§iiiiiiislalsi»ilig \ *.

S3Nld3SSV 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

300 -

250 -

ft 200 -

= 150 -

100 -

50 -

111 '

a

GOIAS^

MARANHAO

RIO GRANDE DO SUL Vtf

SANTA^TARi^^J^^1100

CEARA'a^" / ^ACRE

/ /^/^ALAGOAS

S" . . . •

1 I >

/

MATOGROSSO ,

/ >

/ /

BAHIA. ^ / -

>^ O ^SAO PAULO

MINASGERAIS<*//' / ^

^°*RJO DE JANEI^O^^

^SRORAIMA-^

J»ESPIRITO SANTO

PARANA

i l l

1 / l \ '

1.3:1 L5:l

J> ° AMAZONAS

/ ^ ^

• TOTAL LAND BIRDS

O NATIVE LAND BIRDS

I f !

>

7:i

^2:1'

-

-

300 -

250 -

u. 200

o

S 150

IE

100 -

50 -

b

-

y

• i i i i 1

 ' 1.5:1'

„.„ _ .. UPPER AMAZONIA'S--*,..

?M^?^ « » (AMAZONAS-ACRE- J"''t

(MATO GROSSO-GOIAS) RORAIMA) /

SOUTHEAST /

(SAO PAULO^T

^^-•LOWE R AMAZONIA

' m (PARA-AMAPA)

y ° EAST y

/ (BAHIA-MINAS GERAIS- -

^ ESP IRITO SANTO-

^ RIO DE JANEIRO)

NORTH / o» ^

(MARANHAO-PIAUI-CEARA)^ / SOUTH

^ ^ ^ (PARANA-STA. CATARINA-

/ y RIO GRANDE DO SUL)

/ y

• ^ ^'NORTHEAST

^ / ^ (PERNAMBUCO-PARAIBA-ALAGOAS)

S y^ y>

i i 1 i |

-

• 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 intendancy 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 zonation of Chapman (1917); passerine-nonpasserine

ratios are followed by numbers of species in parentheses. (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 lowlands. The easternmost province, Ituri, rises into

humid-forested highlands, and both its ratios rise

to 1.76.

Chapin (1932-1954, 65:98) saw similarities between 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 responsible for the changes marking the temperate and

Paramo zones" and, one might add, the same factors 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 "island" mountains in Portuguese East Africa (Vincent, 1934). These have species numbers of 48, 68,

and 65 and ratios of 3.00, 2.24, and 2.61. Taken together, 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 assumed, list from a large area.

ARIDITY.—If the entire North American continent be taken as an example, the distribution of

passerine-nonpasserine ratios of the native land avifaunas shows a gross correlation with general environment (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 concentrated in mountainous southwestern United

States and adjoining northwestern Mexico. South

of the Isthmus of Tehuantepec, the ratios are lowest 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 American 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). Florida and the Yucatan Peninsula mostly consist of

physiographically undramatic lowland and the

vegetations display a monotonously simple physiognomy. This is reason enough to make them predictably poor in variety of habitats and numbers of

species. The very low ratio in Florida could correspond 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," jutting 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 northeastern Para, opposite the Amazon delta, with a relatively 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 adjoining 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 altittidinally 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 proportion (50 percent or more) of the native land avifauna 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 percent 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 native 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 characterized by a rather abruptly higher set of ratios,

particularly along the Caribbean slope. A corroborative 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 version (Tosi, 1969). Compared to the Caribbean side,

a lowering in subtropical ratios at analogous altitudes and topographies on the Pacific slope is probably 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 upward 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 faunal 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 depletion with increasing altitude: no bird in the

montane belt has not also been reported from the

lower-montane belt. This does not mean that typically 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 comlexity of the montane forest with increasing altitude

and to the inclusion, at Cerro de la Muerte, of treeless 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 opinion 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 instances 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 species from open-country biotopes. Indeed, tree plantations, regenerating clearings, and shrubby secondary and and broken primary growth are richer in

opportunities for arboreal, predominantly passerine 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 species and 1.50 for nonforest species plus forest species

in nonforest. The entire African breeding land avifauna 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 complete compilation tends to have a lower passerinenonpasserine ratio than does an earlier or less complete compilation, irrespective of the size of the

unit. Put another way, the larger the number of

contributing samples the smaller will be the passerine proportion, especially among the residents.

This is exemplified by the Central American units

that have more than one published checklist: Guatemala, 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 passerinenonpasserine equilibrium. The two field stations,

Barro Colorado Island and Finca La Selva, are also

well enough known for their ratios to be in equilibrium, at least for the resident avifauna, and

neither is likely to change without extensive alteration 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 samples (Table 4). Among total land birds, there are

fewer than a handful of such instances: the General

Valley and Monteverde when migrants were absent, 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 composition of the corresponding avifaunas varies accordingly, 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 material will seldom turn out the same as the first.

The ratios, manifestly so precise, have been derived

from the species totals, which have been synthesized 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 observers, 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 comparable 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 moment 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 nonpasserines, are scarce, scattered, secretive, or otherwise

hard to find, while wind, rain, and mist often

severely hinder observation. Species recorded during different trips usually differ from one another

more in the highlands than at lower, warmer elevations. Native birds that rise seasonally to higher

altitudes, even if no more abundant in species than

those moving downward, produce a proportionately 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 movements by some, either alone or mixed with others

and casually or opportunistically noted, can further 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 biotas, however, conform to no universal standard and

their compositions are each the unique result of interplay among many factors that are differentially

peculiar to islands: this makes islands synecologically 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 archipelago differ a good deal."

If it be assumed that islands do not differ environmentally from comparable mainland under similar

conditions, the impoverishment of insular avifaunas 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 impoverishment, as exemplified by the reduced numbers of

plant species (with which the numbers of bird species 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 number or variety of species in the association or community. 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 inaccessible to potential colonizers, can have widely divergent

numbers and constellations of species. Some islands

are relatively more impoverished than others. Compared 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 member of an island group, in tropical and subtropical

waters around the globe, including the southwestern 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 continental 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 proportion 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 acceleration and the abruptness of the increase are both

contrary to what happens on the mainland. (Intraisland 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 Mindanao and Luzon and possibly Celebes, do not.

Figure 11 is a semilog survey of tropical, subtropical, and southern-hemisphere island groups

containing at least two islands. In overall appearance 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 passerinenonpasserine ratio thus appears to be largely independent 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 differentiation 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 (including 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. Eustatius; 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, Indefatigable; 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, Lombok; 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, Halmahera; 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.

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Antilles, Greater Antilles, Sundas (the larger

islands on a higher level, the smaller islands on a

lower level), Philippines, Solomons, French Polynesia; far above lie islands of the Hawaiian and

Galapagos groups, all with very high ratios; the

few higher northern-latitude islands which have
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FIGURE 10.—Area-species and area-ratio semilog distribution of single islands: a, species numbers

scaled in twenties; b, species numbers scaled in fifties. (Entries the same as in Figure 9; open

symbols = numbers of native land species, solid symbols = passerine-nonpasserine ratios.

Symbols are: star in circle = Sunda Islands; triangle pointing downward — Moluccas-Celebes*

Philippines; triangle pointing upward « New Guinea area; star = southwest Pacific; square in

circle = Greater Antilles and Virgins; square = Lesser Antilles and western Caribbean; diamond

= Indian Ocean; small circles = all others.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

RATIO OF PASSERINES TO NONPASSERINES

S$S8S8?S82S?8 8

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FIGURE 11.—Area-species and area-ratio semilog distribution of island groups (including New

Zealand) in the warmer oceans. (Open circles = numbers of native land species; solid circles

= passerine-nonpasserine ratios; double circles = a smaller group and larger group to which it

belongs. 1, Manua; 2, Lau; 3, Aldabra; 4, Bay Is. of Honduras; 5, Seychelles; 6, Tres Marias; 7,

Palau; 8, Virgin; 8A, Tonga; 9, Anamba; 10, Kangean; 11, Madeira; 12, Revilla Gigedos; 12A,

Tuamotu; 13, Caroline; 14, Dutch Leeward; 15, Santa Cruz (southwest Pacific); 16, Mariana;

16A, Marquesas (South Pacific); 16B, Pearl; 17, Kei; 18, Louisiade; 19, Nicobar; 19A, Society;

20, Comoro; 21, Natuna; 22, Admiralty; 23, Azores; 24, Loyalty; 25, Windward (Lesser Antilles);

26, Samoa; 27, D'Entrecasteaux; 28, Micronesia; 29, Leeward (Lesser Antilles); 29A, French

Polynesia; 30, Cape Verde; 31, Sula; 32, Lesser Antilles; 33, Andaman; 34, Canary; 35, Galapagos;

36, Aru; 37, Bahamas; 38, Falkland; 38A, West Sumatra; 39, Palawan; 40, New Hebrides; 41,

Hawaii; 42, Fiji; 43, New Caledonia; 44, Solomon; 44A, Lesser Sunda (excluding Bali and

Kangean Is.); 44B, Lesser Sunda (including Bali and Kangean Is.); 45, Moluccas; 46, Greater

Antilles; 47, West Indies; 48, New Zealand; 49, Philippine (excluding Palawan); 50, Philippine;

51, New Guinea; 52, Greater Sunda.)
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 (nonpasserines), 0.33 for the Y-intercept, 0.85 for the

regression slope, and 0.05 for the variance of estimate 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 variation 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 (nonpasserines), 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. Eightyfive 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 nonpasserines, 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 nonpasserines.

Several of the above entries, however, form part

of larger entries or incorporate smaller ones—for

example, the Lesser Antilles and its two components, 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 predictable ratio of native land passerines to nonpasserines is almost exactly 1:1 on island groups in

the warmer oceans and seas around the globe, irrespective of such contributory factors as area, distance, origin, ecology, number and variety of

islands, or size of avifauna. Island groups are preferred 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 receives support from the following example, that

of the Pearl Archipelago in the Bay of Panama

(based on MacArthur et al., 1972, 1973; A. Wetmore, 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 Islands 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 passerinenonpasserine 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 threequarters 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 reduced 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 theory 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 Channel Islands, the other, the island of Karkar off the

north coast of New Guinea.

The Channel Islands, a temperate-zone archipelago 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 remained about the same, provided the habitat conditions had also remained about the same and the

two surveys were equally thorough.

Diamond did not include species lists in his analysis. 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 percent 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 incompletely 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 colonization, mainly from New Guinea but with seven

additional species from New Britain; recent volcanic activity has not affected bird distribution

significantly; the original vegetation was principally 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 immigration rates. Qualifying the actual figures, Diamond 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, expressed 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 percent 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 avifauna 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 presumed 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 passerinenonpasserine ratio of 0.48. Between 1914 and 1969,

three of the land birds may have become extinct

(all nonpasserines), while ten, presumably breeding, land birds (eight nonpasserines and two passerines) 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 Hispaniola, 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,. . ." (Terborgh 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 resident 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 passerines 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 turnover rate of about 0.34 percent both among the

nonintroduced resident land birds and the breeding 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 Faaborg) 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 Faaborg) 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 documentation 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 corresponding 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 artifact, 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 relative 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 mainland 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 Kepler, 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 Guanica list, two species (both of them Antillean nonpasserines) 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 otherwise 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 including the Galapagos Archipelago, the species

pool from which to draw extends through southern 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 morphological, 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 nonresidents outnumber residents occur mostly in the

Western Hemisphere. Cuba, Cozumel, and the

Swan Islands are easily explainable as steppingstones utilized by transients crossing the Gulf of

Mexico and the Caribbean Sea; depauperate Barbados has an "abnormally" small number of residents compared to the cumulative number of migrants 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 expectation. 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 mounting number of irregular and accidental visitors.

Cocos appears to be too small, too distant, too

isolated, and conceivably too inhospitable for neotropical 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 between 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 logarithmic 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 percent 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 published a report, Cocos, in the tropical eastern Pacific, 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 apparent" 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 theory 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 percentage 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 Island cuckoo. A well-marked Pacific isolate immediately 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 common 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 flycatcher 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 (Pinaroloxias), 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 aureola, 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 differentiated 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 increasing 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. Moreau (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 result 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 independent 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 representations 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 Archipelago, 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, compared to the Malay Peninsula, 1.28, and Singapore,

1.21, compared to Malaya, 1.20.

In the temperate zone, at least at northern latitudes, 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 Belgium and Switzerland), 1.66; Newfoundland Island, 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 Tasmania, it could perhaps be a case of one island, Tasmania, being satellic to a very much larger island,

Australia (see below). On the other hand, Tasmania, 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., relative numbers of species and passerine-nonpasserine

ratios, correspond to those on the nearest comparable 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 suggested by the passerine-nonpasserine ratios: Zanzibar, 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 difficult to imagine why this should be so" (Moreau,

1966:352). Evidently great distance is not necessarily required to produce an oceanic effect, particularly in tropical latitudes.

Socotra, like Pemba, is probably "an 'old' conti-
AREA IN SQUARE MILES

FIGURE 12.—West Indian native land avifaunas: correlation between area and numbers of

species (numbered large open drdes and dashed regression line) and between area and passerinenonpasserine ratios (numbered large solid cirdes and solid regression line). (Numbered entries

include islands of the Greater and Lesser Antilles and in the open Caribbean Sea: 1, Saba; 2,

St. Eustatius; 3, Old Providence; 4, St. Andrew; 5, Grenadines; 6, Montserrat; 7, Anguilla; 8, St.

Martin; 9, Nevis; 10, Marie Galante; 11, Barbuda; 12, St. Kitts; 13, Grand Cayman; 14, Antigua;

15, Grenada; 16, St. Vincent; 17, Barbados; 18, St. Luda; 19, Dominica; 20, Martinique; 21,

Guadeloupe; 22, Puerto Rico; 23, Jamaica; 24, Hispaniola; 25, Cuba. Lettered entries (small

circles) stand for satellite islands (induding the Virgins) and islands fringing the mainland: a.

Swan Islands; b, Barbareta; c, Utila; d, Guanaja; e, Roatan; f, Culebra; g, Anegada; h, Virgin

Gorda; i, St. John; j, Tortola; k, St. Thomas; 1, St. Croix; m, Mona; n, Vieques; o, Cayo Largo;

p, Aruba; q, Bonaire; r, Curacao; s, Tobago; t, Cozumel; u, Gonave; v, Margarita; w, Isle of

Pines; x, Andros; y, Trinidad.)
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 nonpasserines 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 proportion of passerines to nonpasserines also characterizes 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 continents and receive the fringes of continental faunas and are

extensive enough, with enough comparable islands, to show

informative patterns of distribution. The archipelago of Wallacea [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 areaspecies 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 areaspecies 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 migrantdominated 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 picture 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, comprising the Greater and Lesser Antilles and a scattered

few in the western Caribbean Sea (excluding satellites, e.g., Isle of Pines, Gonave, islets on the

Puerto Rican shelf). These satisfy two requirements: all are single islands, except the Grenadines,

and all are zoogeographically West Indian (in the

sense of Bond, 1948). Increasing numbers of species 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, overpopulated, impoverished island 17, Barbados).

The passerine-nonpasserine ratios correlate negatively 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 isolated 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 passerinenonpasserine and suboscine-oscine ratios is presented in Figure 30.)

Margarita and Trinidad, off northeastern Venezuela, were included above with continental islands,

as was also Isla Cozumel, off the Yucatan Peninsula. Cozumel, strictly speaking, appears to be

somewhat separated from the continental shelf

proper, hence could belong in the offshore category, together with the Bay Islands, off Honduras,

and the Dutch Leewards, off northwestern Venezuela.

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 America 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 exceeding 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 (Figure 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 reduction 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 Hispaniola, 0.88; Vieques, 0.87, compared to Puerto Rico,

0.86.

The islets, primarily including the Virgin Islands, that are situated on the Puerto Rican shelf

(Figure 12/-/) or, in the case of St. Croix, associated 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; Simalur, 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

amphibians,

lizards,

snakes

13

11

9

17

85

42

17

Species of

resident

land

birds ;

29

29

27

77

239

82

39

Passerines:

nonpasserines

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 isolated, 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 herpetofauna 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 passerinenonpasserine 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 mainland (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 noncontinental Aruba, Curasao, or Bonaire, individually or as a group. In the accompanying table, it

appears that the passerine-nonpasserine ratios tend

to correlate with the climato-vegetational conditions of the islands, whereas species numbers,

unless individually qualified, do not.

Southeastern Asia, New Guinea, and the Western 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 numbered entries upon which the regressions are constructed 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 depressed 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. Without 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 attributable 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 stepwise increase in faunal, and presumably environmental, 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 incompletely separated from the Sunda shelf. The

passerine-nonpasserine ratio of 1.30 for the archipelago 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, Christmas; 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, Philippines (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, peninsular extension which, if lists were available, would

no doubt prove to have a distinctly lower ratio

than the entire island, just as in mainland situations. The principal smaller islands have the reduced 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 zoogeographically 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 numerous 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 Sumatra, 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 Venezuela. They have above-unity ratios, except Lombok 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 mountain 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 Sumbawa, 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 mountain fauna (Mayr, 1944a: 176) could in itself account for the low species total and low ratio.

According to Mayr (1944a: 181), "lack of pronounced 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 proportion of endemics and a smaller number of species. 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 overseas colonists compared to nonpasserines on the

other hand.

A selective brake may at the same time be applying 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 discovered, 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 islands 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^ •Bl^«1.2 1

W. Sumatravm24^ f ls

- .95V89^

Christmas I. 33

Cocos-Keeling*

7B

 w ^ ^ - Keils., ^

^ - ^ ••' Aruls-

S U N DA S

2

Louisiades

Q kilometers 1000

FIGURE 15.—Southeast Asia to tropical central Pacific: passerinenonpasseritye ratios of native land birds.

the continental shelf. Even the Bismarcks and Solomons are

narrowly separated from it and from each other. Zoogeographically, 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 passerine decrease, relative to the nonpasserines, become 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 increasing 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

-.El I ice Is*

, ..Banks Is.

^}2L Fiji 1.30

New HebridesV\

1.09.

•f Toketou Is*

Samoa L46

1.40^% ^

.

1307 * 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 impoverished Papuan fauna, except for a few endemics of
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0 . U.lni.lm . 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 satellite (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 archipelagos, forested mountain conditions are more

conducive to passerine than to nonpasserine differentiation. 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 numbers 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 Hebrides contain the principal large islands east of

the Solomons. Fiji and Samoa constitute the avifaunal 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 lacking distinctive endemic elements (Mayr, 1941c:

205). The accompanying tabulation contrasts avifaunal 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 belowunity 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, environmental 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-tounity 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 distributional pattern of sorts (Figure 18). Excluding

the two largest islands (Espiritu Santo and Malekula), avifaunas become impoverished with increasing distance from the center of the arc, ratios

decrease most markedly through the islands trailing 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 distribution 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, particularly for smaller islands" (Mayr, 1941c:211). One

might add that the destructive effects would probably be greater on the passerines than on the

larger, more robust nonpasserines. The two large

islands of Espiritu Santo and Malekula are situated off to one side from the others. Espiritu Santo 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 mainstream 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:

TMT: . Banks Is. (34) 127

Vaaal^^-Vawci ) ^

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)^ !•

 l6 7

,82

1.13

NEW HEBRIDES (46)

Efate(37) ^

1.06

4

Erromanga (33)

-e , -

W

/ * \ W Tanna (29)

\ NEW CALEDONIA AREA(55) \ W. 7 |

— '"-. • Aneiteum (27)

.83

\

New"Ca^!nja(50)

 138 ^ _

f¥W M»fi(20)%

miles 200

11000

0 kilometers 200

NEW HEBRIDES-NEW CALEDONIA

FIGURE 18.—New Hebrides and New Caledonia: passerinenonpasserine ratios of native land birds (numbers of species

in parentheses).

Society Islands, 13; Marquesas, 11; Tuamotu Archipelago, 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 difference 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 Carolines 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 Carolines, 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 passerines, 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); Madagascar, Mascarenes, Seychelles; New Caledonia (previously 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 outlying 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 passerines, 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 proportion 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 Madagascan 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, Moreau 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 certain sign of local radiation in an insular environment. "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 explanation, 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 casualties in the bird fauna of one or other of the habitats that numerous niches became vacant." Yet

Moreau's reference to the "obvious broadening of

niche . . . occupied by the same species in Madagascar 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 Mayotte); the Aldabras, with only eight passerines and

six nonpasserines; the Seychelles, with eight passerines and seven nonpasserines; the Mascarenes,

with seven species each of passerines and nonpasserines 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 receive 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 nonpasserines, 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, excluding 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 passerinenonpasserine ratios that are greatly reduced from

those on comparable mainland; they receive extremely few potential colonizers; and both have

given rise, especially on more isolated New Zealand,

to a variety of flightless birds, including spectacularly gigantic ones.

Hawaiian Islands and Galapagos: Adaptive radiation from a single ancestor into a number of related species on oceanic archipelagos has taken

place most dramatically in these two groups, probably 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 propagules (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 Hawaiians 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 successfully 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 passerines and eight nonpasserines or, excluding the

westernmost atolls, five passerines and eight nonpasserines (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 Hawaiian 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 residents to nonresidents, increasing equatorward (as

on the island groups off eastern Asia), remains low

on the Azores and Madeiras, mounts on the subtropical 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 passerinenonpasserine ratios are based on Moreau, 1966;

the other entries are based on Amadon, 1953.

Fernando Prin- Sao AnnoPo 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 faunistic 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 divergence from the mainland populations, a proportion 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 unquestionably 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 overseas, 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 islands 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 resident 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 various 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 passerines outnumber those of nonpasserines two to one.

The ratio remains about the same for the zoogeographic regions: slightly lower in the tropics,

slightly higher outside the tropics. Continued subdivision into smaller and increasingly homogeneous 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 abundance 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, wellknown 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 impoverished avifauna is surely the sign of an impoverished 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 twothirds 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 suboscines are equally divided between the strictly

neotropical Furnarii, including woodhewers, ovenbirds, antbirds, and tapaculos, and the more widespread 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, except 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. Percentagewise the suboscines decline linearly from northern 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 Honduran lowland on the Caribbean side. If El Salvador 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 percentages 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 suboscines and oscines, shows their relative magnitudes, 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. Southwestern 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 suboscines to oscines. As with passerines and nonpasserines, 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, excluding 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 passerines. 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

AMAZONAS ^

PARA

>MA 0 PAULO

MNAS 6ERAIS / ^

BAHIA^

PARANA So*

MARANNAO \ S^ RIO DE JANEIRO

AMAPAl^ \ f ESPIRITO SANTO

ACREo. V^*"*" *

STA. CATARINA ••"pufui *KRNAMIUCO

y S ••ALAGOA 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-ACREEASTERN AMAZONIA o— • RORAMA)

(PARA-AMAM-MARANHAO)

SOUTHWEST (MAT0 6R0SS0-R0NDONIA)

EAST-CCNTRAL (BAHIA-SOIAS)

SOUTHEAn (SAOPAUIO).

"EAST MWASGERAISESHRTTO STO.-RIO DE JANEIRO)

SOUTH 9ARANA-STA.CATARINARIO GRANDE 0 0 SOD

NORTHEAST PIAUI-CEARA-PARAIIAKRNAMIUC0-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 cannot 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 discernible in the relatively uniform Amazon basin.

In the accompanying tabulation, the principal

faunal sectors are the same ones previously employed for passerines and nonpasserines in Colombia; 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 Pacific, 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, respectively, for the "Central Mountain" sector appear

to be a good representative average: the corresponding 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 Caqueta and the 1.85 and 1.97 of Amazonian Brazil.

The 1.38 and 1.59 of the Orinocan sector are approached 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 passerines 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 proportion decreases with increasing elevation (Table

7). Exceptions, i.e., lowland areas with low suboscine proportions, are the interior valleys of eastern 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" (Terborgh 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 climatic 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 determined 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 avifauna by the occurrence of extralimital migrants.

Native ratios indicate in the main that the suboscine 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 identical passerine-nonpasserine ratios.) The BorucaPaso Real area is located in edaphically poor, cutover savanna-type country; the Sixaola area has

been heavily deforested by commercial interests and

includes old records that probably apply to the subtropical 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 Reventazon 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 remains 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 equivalent 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, assortment of birds, and climate, has no Caribbean

counterpart. At higher elevations, the suboscineoscine ratios in the lower-montane and montane

belts plunge to half their value or less in the subtropical belt.

Forest versus Nonforest: How the suboscineoscine relationship fares at a site under forested

and under unforested conditions cannot be determined 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 exceptions. One, the Bagaces site, is perhaps an "abnormal" 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 suboscineoscine 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 ranging 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 virtually 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 compilations 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 wellknown 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 compilation 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 Colorado, in particular, shows a progressive decrease in

ratio that correlates roughly with increase in number 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 increase in numbers of suboscines, oscines, and passerines 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 suboscineoscine 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 quality, yielding ratios that decrease irregularly or may

even level out in the highlands. Thus qualified,

suboscine-oscine ratios correlate inversely with rising elevation upon mountain slopes facing the

lowlands, not upon isolated hills or ridges. The

ratio tends to be higher than expected in intermontane situations, as in the central highlands of

Colombia or Costa Rica, where the climatic and

topographic conditions are complex and the available 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 localities 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 expected pertain to places that at present are extensively unforested when climatically they ought to

be forested.

Countries or other sizeable units with large avifaunas that are represented by lists compiled a

generation apart show little change in suboscineoscine 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 decreasing ratio, at least in humid localities. At all magnitudes of faunal or areal size, suboscines increase

at the expense of the oscines in the forested portion of an environment compared to the unforested 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. Converted into percentages or ratios and cross-plotted,

the two relationships produce a graph or scattergram permitting measurement and visual appreciation 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 America (Brazil), and southern South America (Figure

22). At a lesser scale of magnitude, a strictly political 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 homogeneous environments. Comparisons could then be

made of similar habitats at different places or dissimilar 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 disproportionately large changes in the percentages

or ratios.

SUBREGIONAL SECTORS.—Four sorts of lists for

each of the major areas—Middle America, northwestern South America, Brazil, southern South

America—are plotted in Figure 21. The lists separate 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 waterbird element in southern South America, the lists

bear an overall similarity to one another despite

their considerable neotropical spread. The straightness of the vertical alignment is due to the very

small difference in passerine percentages (1.2 percent in the case of total passerines).

COUNTRIES.—In Figure 23, the unit of comparison 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 American units, the distribution of the native land avifaunas, compared to that of the total land avifaunas, 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 territories, 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 principle, 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 similarity may not be entirely accidental, provided

one is impressed by the correct alignment of Middle American units, including the pairing of

southern Mexico with Guatemala, Honduras with

Nicaragua, and Costa Rica with Panama; the location of the Yucatan Peninsula "offshore" from

the neighboring mainland; the sequence of Andean
NUMBER 212 59

% NONPASSERINES/AVIFAUN 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

^BOLIVI A

^ N * ARGENTINA

/ "

J .

65 60 55

% PASSERINES/AVIFAUN 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

% NONPASSERINES/AVIFAUN 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

% PASSERINES / AVIFAUN 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; horizontal 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 (approximately 2:1) was chosen arbitrarily to represent

the line of division between the northern and

southern hemispheres in Figure 23. A strict latitudinal 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 transposed 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) relationship 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 comprehensible 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 increasing elevation, reaching an extreme in the high

cordilleras, and with increasing latitude, particularly in Middle America.

A list which one has reason to suspect is deficient

usually yields an (a): (b) value that looks exaggerated or "wrong." A value that looks "wrong"

should make one wonder about the representativeness 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 remaining "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 reveal 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 paramo 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 coefficients 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 suboscinesoscines. (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, nonAndean Santa Marta, would result in even

higher correlations, most strikingly with passerinenonpasserine ratios and suboscine-oscine ratios.

The (a): (b) single-figure value would appear

to have advantages that are best suited to siteoriented situations. Unfortunately, it is often the

local list that engenders doubts as to its reliability

and comprehensiveness. A telltale sign is the presence of few birds of prey, a deficiency which

indicates in turn that the entire nonpasserine component may be too small. Yet the same lists can

have good representations of suboscines and oscines.

Elimination of the nonpasserines, therefore, actually increases their accuracy and usefulness. Indeed,

suboscine-oscine ratios (Figure 25) provide singlefigure 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 climatovegetational indicator of biotic richness in the New

World tropics, exclusive of the Antilles (all of

which lack this sedentary family). Since the mainland distribution of the Formicariidae, or of any

other selected group of "rain"-forest families, is

ecological rather than political, comparisons between localities should be based upon environmental 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 lowlands in South America for which data are

available—Caribbean Colombia, northeastern Venezuela, southwestern Ecuador, and coastal Peru—

would, if entered in Figure 27, form an "out-ofplace" 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 reduction 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 temperature 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 components.
NUMBER 212 65

10

15

20

25

30

«/>

- 3 5

- 4 0

UJ

> 45

- 5 0

55

60

65

70

75

25

°/o NONPASSERINES/AVIFAUN 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^Poa 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* .

ACRE > ^

/ *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

•/• PASSERINES/AVIFAUN 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 altitudinal 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 lowlands). These agree in having little primary

woodland coupled with a reduced proportion of

suboscines from that normally present at tropicalbelt 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. Intermediately placed entries occupy an area the nature

of which may not be so much transitional as confused, 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 representative of an avifauna. For example, reports on general collections, such as those of Dearborn in Guatemala 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 geographical 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 individual 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 passerines (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 singlefigure 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

NONPASSERINE S / AVIFAUN 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 CHILES. .

LA LOLA* Hda. Altamira

.CARRILLO

•GUAYABO-AQUIARES

.SIXAOLA

FINCA LA SELVAU972)

•FINC 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 PASSERINE S / AVIFAUN 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

%NONPASSERINES / AVIFAUN 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

LOWE I MONTAN E

LA CHONTA«"

70

65

\

60 -o

•MONTEVEROE DE TILARAN

.SAN VITO

Silencio de Tilaran

55

5 0 -

Rincon de la Vieja. .

MIRAVALLES. /

TROPICAL /

TO SUBTROPICAL '

HDA. SANTA /

MARIA AREA .

PELON DE LA ALTURA . . BARRANC A SECTOR • , HELECHALES "BORUC A * Tinama5 t as 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 • •

HUMI D SOUTHWES T

• POZO AZUL-POZO PITAL

-I 1 1 1 1 1 1 1 1 1 I • ' ' I • 45 50 55 60

% PASSERINES/AVIFAUN 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 ISLANDS Of '

HONDURAS

VIROM ISLANDS

HUMNIM.A

GREATER ANTILLES

I I

ft» 0.M 0.10

PASSERINES: NONPASSERINES

FIGURE 30.—Scattergram of islands in the Caribbean area: passerine-nonpassenne ratios versus

suboscine-oscine ratios.
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 compilations, 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 dominant 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 collection to that of a long-term compilation. Particularly 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 restrictive as it tends to include few nonregular taxa

and to group them seasonally. Moreover, a shortterm 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 different years. Sample can be compared against sample from the same place or from different places.

Thus the elaboration of a bioclimatic scheme by

which to plot the comparative distribution of neotropical 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 siteoriented 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 datagathering means would work cooperatively regardless 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 additional 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

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>BARBA SITE (DRY)

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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 incorporating 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 departure 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

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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, passerines, 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 majority 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 number of species instead of number of days. Figure

34 cumulatively plots the proportions of the major

components. It also shows the exponential increase 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

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NATIVE LAND BIRDS

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-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 avifaunal 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

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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 encountered at an approximately constant rate during 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 passerine proportion in compilations or checklists generally, 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 members, 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 horizontal 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 preceding 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 dismantling 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 percentages 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 remainder 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 collecting, 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 passerinenonpasserine 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 between 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 species 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. Encounters 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 temporarily 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

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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 constant rate of increase in number of subosdnes. Diameter of dots equivalent to two spedes on

absdssa.)
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0

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I I I I I I • i • I • • i I ' ' ' ' I ' ' ' ' I ' '

PASSERINES NO. or SPECIES

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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 nonpasserines (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 representativeness 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 suboscines (or "poor" in oscines). There is little reason 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 suboscine 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 approximately 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 passerines 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 regard for departures from the norm than do percentages (Figure 35). The passerine-nonpasserine

ratio stabilizes by the time approximately twothirds 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 resemblance 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 because one has more entries than the other; the fact

that the internal proportions may be similar is

immaterial.

Faunal resemblance formulas and the passerinenonpasserine 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 passerinenonpasserine suboscine-ocine interrelationship continues 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 replacement and displacement or convergence and div«rgence. The result would be constant improvement
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in the making of quantitative comparisons between

geographically changing faunas. The ability to discern taxonomic derivatives and ecological counterparts 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 compartmentalize environmental equivalence for unrelated 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 regard 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 include 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 compilations 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 increasingly 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 polytypic 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 subspecific 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 expected 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 polytypic 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 specificity of closely related birds. Put another way, the

site is the testing ground for the belief that sympatric populations do not interbreed and are specifically 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 avifaunas. 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 indispensable 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 environments. 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 precipitation, 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 environmental 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 qualities of the site as possible. Each association is, in effect, a

natural mechanism endeavoring to sustain the greatest possible 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 environmental conditions in an association, assigns proper

weights to each, and produces an answer in terms of a vegetative cover. (Holdridge et al., 1971:16.)

Responses of the vegetation-dependent birds are

secondary or indirect.

The investigator will be confronted by bioclimatic 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 variable 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 opportunity arises to test the efficacy of the passerinenonpasserine suboscine-oscine method within narrow 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 conditions prevail anywhere in the world. This ecologicalevolutionary 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 equivalent 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 taxonomic, 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 comparative 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 usually 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 representations 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 coverage of a large unit records even a lone migrant as

inhabiting the entire area.

There are several reasons that make it preferable to institute a comparative survey in monotonously immense Amazonia than, say, in complex

little Costa Rica, the other area figuring prominently 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 comparative 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 difference 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 distribution 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 elevations, where environmental influences are multidirectional and knowledge of faunal movements

is rudimentary. By contrast, the Amazonian lowland is essentially free of altitudinal complexities,

and it apparently contains huge, climatically similar blocks of vegetation: "In regard to the largearea 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 respond 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 situation in Costa Rica.

Costa Rica, at ten degrees above the equator,

has an avifauna whose reproductive period generally 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 September, 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 unquestionably 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 avifauna. 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 percent 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 (formations) 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, dependent upon, and variously intermediate between the

adjacent climatic standard of the life zone, or

ecosystem, on either side. Transects made horizontally, say, in Amazonia or vertically in Costa Rica

should, if properly planned, pass through a natural

sequence of life zones (see Holdridge, 1947; Holdridge 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 Gilliard, 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 Galbraith, 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; compiled 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; Edwards, 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 Schauensee, 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 Schauensee, 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 Galbraith, 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 Schauensee, 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; Darlington, 1931)

Santarem, Brazil (Riker, 1890-1891; Griscom and Greenway,

1941; Pinto, 1938, 1944; Meyer de Schauensee, 1970; compiled 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 Galbraith, 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; Leopold, 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; Delacour, 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; Leopold, 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 (living 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

British 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

Strip 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 District. 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 Territory, 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, Brazil

Oregon

Yugoslavia

Sao Paulo, Brazil

Piauf, Brazil

Uele, Congo

Surinam-French Guiana

Guyana

Great Britain

Victoria, Australia

Honshu, Japan

Korea

Utah

Greater Antilles

Minnesota

Idaho

Parana, Brazil

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 AvifaunaTotal

(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, Brazil

Meta, Colombia

Moluccas

Ireland

Santa Catarina, Brazil

South Carolina

Tamaulipas, Mexico

Hokkaido, Japan

H1span1o1a

Sabah (North Borneo)

Panama

Chiapas, Mexico

Veracruz, Mexico

New Brunswick, Canada

Banks Island, Canada

Tasmania

Ceylon

Northern Thailand

Ituri , Congo

West Virginia

Yucatan State, Mexico

Sinaloa, Mexico

Paraiba, Brazil

Nova Scotia, Canada

Campeche, Mexico

Quintana Roo, Mexico

Costa Rica

Southampton Island, Canada

Isla Grande (Tierra del Fuego)

Espi'rito Santo, Brazil

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 Hills, 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

Nonasserines

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

Nonpasserines

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

Nonpasserines

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

Nonpasserines

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

Nonpasserines

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

Nonpasserines

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 Pital (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 Sitio

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., Jr. (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 site , 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 totals, 1957-58

Sept.-Oct. 1957

Jan.-Feb. 1958

Mar.-May 1958

July-Sept. 1958

A er e

9 2-week totals, 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)

Passerines

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 = Subtropical Rain, LMR = Lower Montane Rain, MR = Montane Rain)

Bagaces site (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 ific): 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 l a 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 site (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

Instituto 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 interior 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 (ibid.)

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)

(ibid. , keyed to Blake, 1953)

Nontropical Mexico (Edwards, 1972)

Highlands (ibid.)

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 (ibid.; 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 (ibid.)

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)

(ibid. , keyed to Slud. 1964a)

" (Salvin and Godman, 1879-1904)

11

 (ibid. , keyed to Slud, 1964a)

(Zeledrfn, 1887)

(ibid. , keyed to Slud, 1964a)

(Frantzius, 1869)

(ibid. , 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 Loftin, 1967a)

Below 5000 ft .

Above 4000 ft .

Chiriquf* Volcano (based on Blake, 1958)

(Blake, 1958)

Cerro Campana (based on Eisenmann and Loftin.

1967b)

Canal Zone (Eisenmann and Loftin, 1968:

definitely 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 (ibid.)

Mt. Auyan-tepui (Gilliard, 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 district (Beebe, 1925; Beebe et al. , 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 (ibid.)

Rio Jurua-Rio Puru's (Gyldenstoipe, 1945a, 1951)

Upper Jurua-upper Purus (ibid.)

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 (ibid.)

Itacoatiara (ibid.)

Parintins (ibid.)

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 (ibid.)

(Pinto, 1938, 1944)

Faro (compiled sources)

Obidos (ibid.)

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 (ibid.)

" (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 (ibid.)

(Olrog, 1963)

(Dabbene, 1910)

Minus subtropical northeast (based on Olrog, 1963)

Misiones (ibid.)

So. Patagonia (Olrog, 1959)

Chile (based on Johnson, 1965-67)

Isla Grande, Tierra del Fuego (Humphrey et al. ,

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

203 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

1 67

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

l4.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

12. 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

Espirito 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.

1 0.

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

NonPercentages 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

Nonpasserines 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

Nonpasserines

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+

30. 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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