UNIVERSITY OF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
BIOLOGY
MAR 2 6 1985
IELDIANA
Zoology
Published by Field Museum of Natural History
New Series, No. 13
NOTES ON TYRANT FLYCATCHERS
(AVES: TYRANNIDAE)
fHfc UBRARY OF IttE
MELVIN A. TRAYLOR, JR.
' Of ILLINOIS
DEC 9 1982
November 19, 1982
Publication 1338
NOTES ON TYRANT FLYCATCHERS
(AVES: TYRANNIDAE)
FIELDIANA
Zoology
Published by Field Museum of Natural History
New Series, No. 13
NOTES ON TYRANT FLYCATCHERS
(AVES: TYRANNIDAE)
MELVIN A. TRAYLOR, JR.
Curator Emeritus
Division of Birds
Department of Zoology
Field Museum of Natural History
Accepted for publication January 15, 1980
November 19, 1982
Publication 1338
Library of Congress Catalog Card Number: 82-83010
ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA
CONTENTS
Introduction 1
Tyranniscus australis Olrog and Contino 1
Phyllomyias virescens reiseri 1
Zimmerius vilissimus 2
Sublegatus 4
Suiriri suiririlaffinis 10
Elaenia albiceps and £. parvirostris 12
Hemitriccus iohannis and H. striaticollis 15
Ochthodiaeta signatus Taczanowski, 1874 18
Summary 20
Acknowledgments 20
Literature Cited 21
LIST OF ILLUSTRATIONS
1 . Wing length of Zimmerius vilissimus 3
2. Distribution of Sublegatus 6
3. Localities of Suiriri suiriri suiriri and S. s. affinis 11
4. Breeding records of Elaenia albiceps chilensis and E. parvirostris 14
5. Distribution of Hemitriccus striaticollis and H. iohannis 17
LIST OF TABLES
1. Comparative measurements of wing and bill length in Sublegatus obscurior,
S. modestus, and middle Amazon variable population 9
2. Measurements in Elaenia spp 13
Introduction
During the preparation of the manuscript of the Tyrannidae for Volume 8 of
Peters' Check-list of Birds of the World (Traylor, 1979, pp. 1-228), a number of
problems arose which required further study. Ideally, these problems should
have been resolved and the results published before the appearance of Volume
8. However, in order to avoid any further delay of the volume, I included my
tentative conclusions there and am publishing the justifications here. Un-
fortunately, in the case of Sublegatus, subsequent studies have caused me to
revise the classification presented in Peters' Check-list. The nomenclature used
in this paper is that of Peters' Check-list, the rationale for which was explained
in Traylor (1977).
Tyranniscus australis Olrog and Contino
Olrog & Contino (1966, p. 113) described Tyranniscus australis from a single
female from Yuto, Jujuy, Argentina. They considered it most closely related to
Tyranniscus ( = Phyllomyias) cinereiceps, from which it differed in having the
throat and breast clear gray, not yellow washed with olive, and the belly pale
yellowish, not intense yellow as in cinereiceps. The latter species occurs in the
subtropical zone of the Andes south to Cuzco, Peru, so that there was a gap of
some 1,000 km between the ranges of cinereiceps and australis.
Olrog & Contino also included measurements of the type in their description.
The proportion of tail:wing (53:59 or 90%) differed so much from the pro-
portions of our single male of cinereiceps in Field Museum (taihwing = 44:61 or
72%) that I questioned the identification. Through the kindness of Dr. Olrog, I
have been able to examine the type. It proves to be a specimen of Phyllomyias
sclateri sclateri, a species previously known from Jujuy. The type matches sclateri
in both color and size. Comparison of my measurements (in mm) of the type
with those of four females of sclateri in Field Museum are as follows: australis
(type) — wing 62, tail 54.5, culmen 11, tarsus 16.5; sclateri 4$ 9 — wing 61-65 (63),
tail 54-60 (56.8), culmen 11-11.5 (11.3), tarsus 17-17.5 (17.3).
The name Tyranniscus australis Olrog and Contino becomes a synonym of
Phyllomyias sclateri Berlepsch.
Phyllomyias virescens reiseri
Zimmer (1955, pp. 21-23) considered that Phyllomyias reiseri would have to be
treated as a species distinct from P. virescens, since it occurred in Piaui and
Goias and in northern Paraguay, with virescens appearing in eastern Mato
Grosso, between the two populations. He pointed out differences in coloration
2 FIELDIANA: ZOOLOGY
as well as the very short wing length, which is the salient character of reiseri. I
have examined Zimmer's material in the American Museum, as well as that in
Field Museum, and do not see that two taxa segregate out on either color or
measurements.
With 25 specimens from Piaui south through eastern Mato Grosso and
Paraguay to Misiones, the wing measurements are distributed as follows: 11
males— (1) 54, (1) 58, (2) 59, (1) 60, (2) 61, (1) 62, (2) 63, (1) 65; 14 females— (1) 56,
(2) 57, (2) 58, (5) 59, (1) 60, (2) 61, (1) 62.
Only the single male with wing length 54 can be separated on size; it is from
Piaui, a topotype of reiseri. The spread of measurements at some localities
almost encompasses the total spread for the remainder of the species. Four
males from the vicinity of Villa Rica, southern Paraguay, have wings 59-65,
whereas 10 females from Misiones measure 57-62. I consider reiseri to be a
subspecies of P. virescens, with its range restricted to southern Piaui.
Zimmerius vilissimus
There are two generally recognized races of Zimmerius (olim. Tyranniscus)
vilissimus in Central America: nominate vilissimus from Guatemala and adjoin-
ing Chiapas and El Salvador and parvus from Honduras south through
Nicaragua, Costa Rica, and Panama to northwestern Choco, Colombia. As ex-
pressed in its name, parvus is markedly smaller than vilissimus and has the
whitish forehead and superciliaries less developed, the crown paler and more
slaty, less brownish gray, and the yellow edgings on the flight feathers and
coverts darker and brighter. The color characters are constant between the two
races, with slight overlap due to individual variation, but there is considerable
size variation in parvus, which caused Ridgway (1907, p. 409) to say, "If all the
birds of this species from Nicaragua, Costa Rica and Panama are really of one
subspecies the individual variation in size is very remarkable." An examination
of 197 specimens from Central America shows that this remarkable variation in
size is not individual, but is due to an abrupt increase in size with altitude over
a comparatively small geographic range.
Nominate vilissimus is found over a wide altitudinal range, from 500 to 2,600
m among specimens examined, without any size variation correlated with
altitude. Wing lengths of nine males are 57-64 (av. 60.0) and of 15 females 52-57
(av. 54.4). The northern populations of parvus in Honduras and Nicaragua, all
in the lowlands below about 500 m, are strikingly smaller, with wings of six
males 49-51 (av. 49.5) and wings of five females 43-46 (av. 44.8). These figures
are fairly typical of the populations of the lowlands of Costa Rica and Panama be-
low 600 m, where the average wing lengths, respectively, are males 51.5 and 49.7
and females 45.6 and 44.8 (fig. 1). These lowland records are from the Caribbean
and Pacific slopes of both countries, although parvus avoids the dry Pacific slope
of western Costa Rica. However, in the highlands of Costa Rica at the west end
of the Cordillera de Talamanca around Santa Maria de Dota, and in the high-
lands of western Panama around Volcan de Chiriqui and Cerro Horqueta, both
above 1,500 m, occur long-winged populations that overlap vilissimus in size.
Wing lengths of the Costa Rican populations are 14 males 55-58 (av. 56.1) and 11
females 49-54 (av. 51.3), and those of the Panama populations are 24 males
53-59 (av. 56.1) and 12 females 48-52 (av. 50.5). Although these two populations
Males
vilissimus
Guatemala and Salvador
parvus
Honduras and Nicaragua
Costa Rica < 600 m
Costa Rica 600-1500 m.
Costa Rica > 1500 m.
Panama- 600m.
Panama 600-1500 m.
Panama -1500m.
No. 40
6
27
13
14
18
1
24
vilissimus
Guatemala and Salvador
15
parvus
Honduras and Nicaragua
5
Costa Pica < 600 m.
15
Costa Rica 600-1500 m.
7
Costa Rica > 1500 m.
11
Panama < 600 m.
16
Panama 600-1500 m.
4
Panama > 1500 m.
12
WING LENGTH
45 50 55
60
65
Fig. 1. Wing length of various populations of Zimmerius vilissimus. Wing lengths of
nominate vilissimus are strikingly longer than those of parvus from adjoining Nicaragua
and Honduras and from lowland Costa Rica and Panama. However, highland popula-
tions of parvus from Panama and Costa Rica approach vilissimus in wing length.
4 FIELDIANA: ZOOLOGY
appear isolated, they are connected by continuous highlands above 1,500 m,
and the range of the large populations may also be continuous.
Although the large-sized populations of parvus from Costa Rica and Panama
overlap vilissimus in wing length, there is no evidence that they are biologically
related. The former are identical in color with lowland parvus, and birds from
intermediate localities, between 600 and 1,500 m, particularly in Costa Rica
from which we have more extensive collections, are exactly intermediate in size
(fig. 1). The large size of the highland birds appears to be the result of higher
altitudes acting on the smaller birds of the adjacent lowlands. The latter, "typi-
cal" parvus, stretch from eastern Panama to the Caribbean coast of Honduras
virtually unchanged. The only real biological break is between the northern
populations of parvus in Nicaragua and Honduras and nominate vilissimus in
Guatemala. Although only 320 km separate the two taxa in this area, there is no
evidence of any intergradation between the two, and even the smallest females
of vilissimus are larger than the largest males of northern parvus.
The present evidence strongly suggests that vilissimus and parvus have had
separate evolutionary histories for some time and that their present proximity is
of recent origin. Presumably, in an earlier warm, wet period, tropical forest,
their primary habitat, was continuous through Central America, and proto-
vilissimus had an unbroken range from southern Mexico to South America. In a
subsequent cooler, drier period, this range was fragmented, with one popula-
tion in Guatemala and a second in Costa Rica, Panama, or even northwestern
Colombia. During this period of isolation, they diverged morphologically into
the present vilissimus and parvus. With amelioration of conditions, the forest
again 'became continuous, and parvus extended its range west and north to
Honduras. However, even today, Honduras and Nicaragua are not a favored
habitat for parvus, for the taxon is much less common there than in Costa Rica or
Panama, where it is one of the most common forest-edge species.
The abrupt change between vilissimus andparvus in Guatemala and Honduras
suggests that they might well behave as distinct species if they were to come in
contact. However, the distinction between parvus and improbus of northeastern
Colombia is said to be equally as great (Hellmayr, 1927, p. 471), and until all the
related taxa can be compared, I keep them conspecific. Biological data on voice
and behavior will be as important as morphological data in determining the
status of the various taxa.
Sublegatus
Sublegatus is a widespread genus of small flycatchers found in open wood-
land, scrub, and mangroves from Costa Rica and northern Colombia east to the
Guianas and south, east of the Andes, to northern Argentina. In this century it
was considered monotypic with several well-marked races until Zimmer (1941b,
pp. 1-7) suggested that there were two species. Since that time, treatment of the
taxa has varied considerably. Meyer de Schauensee (1950, p. 861) and Phelps &
Phelps (1963, p. 235) recognized Zimmer's two species and suggested that a
third might have to be recognized, since two races of one of Zimmer's species
occurred together in Colombia and Venezuela, respectively. Haverschmidt
(1970, p. 358) also believed that the coastal and inland populations of Surinam
belonged to different species, both probably resident. However, Meyer de
Schauensee (1966, p. 384) and Meyer de Schauensee & Phelps (1978, p. 277)
TRAYLOR: TYRANT FLYCATCHERS 5
recognized only a single species and considered the apparent cases of sympatry
to be caused by wandering or migrant individuals.
The present study, based on some 200 adult skins, seems to support the
recognition of two species in northern South America, although all those taxa
were included in a single species by Zimmer. On the other hand, there appears
to be extensive intergradation between Zimmer's two species along the lower
Amazon, so that I am unable to recognize his separation of them. Un-
fortunately, the two species here recognized are not those accepted in Peters'
Check-list (Traylor, 1979, p. 18). For this I can only apologize and say that I hope
I have learned something in the meantime.
Within Sublegatus, there are three fairly well-marked entities or groups of
subspecies. The first is the arenarum group which occurs from Costa Rica
through Panama, northern Colombia, and Venezuela to coastal Guianas (fig. 2)
and on most of the islands off the Venezuelan coast (not shown). It is charac-
terized by a comparatively long bill and brighter underparts, with the pale gray
breast sharply distinct from the bright yellow belly, and the throat whitish. At
the extremes of its range, in Costa Rica and the Guianas, it is confined to
mangroves, but from Panama to Venezuela it is also found in dry woodland and
scrub. There are four mainland races recognized within this group, arenarum of
Costa Rica and western Panama, atrirostris of eastern Panama and northern
Colombia, glaber from northern Venezuela through the Guianas, and orinocensis
in the lower Orinoco valley and in Meta, Colombia.
The second entity is the modestus group from southern South America, rang-
ing from southern Peru across northern Bolivia to Mato Grosso, north in the
campos region to Piaui, Maranhao, and Mexiana Island, and south to northern
Argentina and western Uruguay and Rio Grande do Sul, Brazil. This group is
characterized by short bills, fairly bright underparts, and in one race, clear
white wing bars; its habitat is dry woodland and scrub. There are two races,
modestus found from southern Peru through northern Bolivia to Mato Grosso
and eastern Brazil, and brevirostris from central Bolivia and Paraguay south to
western Uruguay and northern Argentina. The latter is migratory, and winter
specimens have been taken in eastern Peru and central Amazonas.
The third group, obscurior, occupies a central position between arenarum and
modestus, mostly peripheral to the Amazonian and Guianan forests. It has been
taken in the north from the lowlands of eastern Colombia east through Ven-
ezuela to interior Guyana and Surinam and the coast of French Guiana, and
south through eastern Ecuador and Peru to northern Bolivia; it also occurs in
eastern Para along the Tocantins and at Belem. The group is characterized by
duller underparts, with the throat grayish like the breast, and the darker gray of
the breast blending into the paler yellow abdomen. It is presumably a bird of
forest clearings and river edges, which accounts for its rather patchy distribu-
tion around the forest. I consider obscurior to be monotypic, although there is
considerable variation in depth of coloration; synonyms of obscurior are sordidus
and peruvianus.
The evidence supporting the concept of two species is found in the distribu-
tion of glaberlorinocensis and obscurior in the region from eastern Colombia to
French Guiana. Meyer de Schauensee (1950, p. 861) reported both glaber and
sordidus ( = obscurior) from Villavicencio, Meta, Colombia. I have examined the
specimen of obscurior, a young unsexed bird in the American Museum, just
completing its molt into adult plumage, and it is unquestionably this form. I
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TRAYLOR: TYRANT FLYCATCHERS 7
have not been able to examine the specimen of glaber, but there are three
specimens of orinocensis in Field Museum from Carimagua, eastern Meta, and
probably the Villavicencio bird was this subspecies. The specimen o( obscurior,
in postjuvenal molt, is almost certainly from a resident population. The
Carimagua birds were in nonbreeding condition, but there is no evidence that
orinocensis is ever migratory. There is no possibility that the dull plumage of the
obscurior could be an immature stage of orinocensis. Its wing length is 74 mm,
exactly the average of 19 males of obscurior, whereas the observed range of wing
lengths of 18 males of orinocensis is only 66-71 mm.
From Venezuela I have two specimens of obscurior (Phelps Coll.), one taken
adjacent to the range of orinocensis and the other well within the range of glaber.
The first is a male from Salto Guaiquinima, Rio Paragua, south of the Orinoco,
less than 160 km from La Paragua where orinocensis has been taken. It differs
from orinocensis not only in color but in much larger size (wing 76). The second
obscurior is a male from Los Altos, Sucre, well within the range of glaber. It is
primarily identifiable on color, since glaber is much larger than orinocensis;
wings of 14 males of glaber were 68-75 mm. The Los Altos male has a wing of 74
mm. Its bill length is 12.5 mm, typical of obscurior but outside the mea-
surements of 14 glaber (13-14 mm), although the difference is not statistically
significant. Phelps & Phelps (1963, p. 235) record a second obscurior from
Guaraunos, Sucre, which I have not seen. They also list a specimen from Isla
Tobeina, Amacuro, as obscurior. I have examined this bird, a female, and be-
lieve it to be an aberrant glaber. It has a paler yellow belly and somewhat darker
gray breast than typical glaber, but it is nearer to the latter than to the two
obscurior which have the dark gray of the breast bleeding over the belly so that
the yellow is hardly apparent.
From Guyana I have examined a single specimen from Rockstone, in the
interior, which is pure obscurior. However, Snyder (1966, p. 222) records two
types of habitat for the species in Guyana: mangroves and swamp, and dry
scrub and open forest. The dry scrub and forest population is obscurior, as shown
by the Rockstone specimen; those from mangroves are presumably glaber, by
extrapolation from adjoining Surinam. Haverschmidt (1970, p. 358) has de-
scribed the distribution of the two forms in Surinam. He found glaber (called
obscurior by Haverschmidt) common in coastal mangroves throughout the year
and obscurior (called sordidus) in inland forest at Phedra in June, August, and
October. To the latter may be added a December specimen from nearby
Brownsweg, taken by Mees. The spread of dates strongly suggests that these
inland birds represent a resident population. I have examined all this material,
and the two series are typical of their respective taxa.
The situation in French Guiana is more confusing. The type of obscurior is
from Cayenne, on the coast, and two other obscurior are from Mana, further
west on the coast. However, there are four specimens from Isle Le Fere, not far
east of Cayenne, which more nearly resemble glaber. Zimmer (1941b, p. 3) noted
this anomaly when he stated that "four adults, all sexed as males though two are
small enough to belong to the other sex [ = Isle Le Pere series], are much
brighter above and below, though the upper parts are not as light nor as
brownish as those of most glaber. They are, however, not far removed from the
Venezuelan form." At the time Zimmer wrote, there were no specimens of
glaber from between Sucre, Venezuela, and French Guiana, and he was natu-
8 FIELDIANA: ZOOLOGY
rally hesitant to assign the Isle Le Pere specimens to that form. Now, however,
glaber is known to be common in the coastal mangroves of Surinam, and I
believe the Isle Le Pere birds belong to that form.
When we consider the examples of parapatric and sympatric distributions of
glaberlorinocensis and obscurior listed above, I believe they must be considered
distinct species. The range of obscurior elsewhere is primarily equatorial, and
there is no reason to expect that it is migratory. Certainly the wide spread of
dates on which obscurior has been taken in inland Surinam argues for a resident
population. I would make S. arenarum and its mainland races, atrirostris, glaber,
and orinocensis, a species separate from S. obscurior.
As the map shows, obscurior is widely found in central South America,
although there is no evidence to show that it is anywhere common. It is re-
markably stable in size throughout its range, showing much less variation than
that between glaber and orinocensis, or between the highland and lowland
populations of brevirostris in Bolivia. Wing lengths of 19 obscurior males from
southeastern Peru to French Guiana show an observed range of 70-78 mm, with
a coefficient of variation (C/V) of 2.71%, comparable with the variation shown
in 14 males of glaber from northern Venezuela (range 68-75, C/V 2.43%). Plum-
age variation, however, is considerable, appearing in the saturation of grays
rather than in pattern. The palest, least saturated populations are found in
French Guiana and adjoining Surinam, topotypical obscurior. The most heavily
saturated specimens are from eastern Amazonia around Belem, and birds from
the upper Rio Negro and Rio Vaupes are almost as dark; these are the popula-
tions called sordidus by Zimmer. Specimens from the eastern edge of the Andes
and adjoining lowlands, from Colombia to northern Bolivia, are intermediate;
this group was called peruvianus by Zimmer. Although these plumage dif-
ferences are readily apparent, I prefer to treat all these populations as a single
taxon, obscurior, until good series are available. The most specimens I have seen
from any one locality are three, and most of the dots on the map represent single
specimens.
Zimmer treated obscurior as conspecific with arenarum and allies, but consid-
ered them distinct from modestus because obscurior and modestus appeared to
overlap in southeastern Peru. In southern Peru, obscurior has been taken on the
middle Rio Ucayali, and in Junin, Madre de Dios, and Puno, and there is a
population of modestus in the Urubamba Valley at Sta. Ana and Maranura. I
have examined all these specimens except the type of peruvianus from Rio Ta-
vara, Puno, and the two Maranura birds in the British Museum. They are all as
Zimmer described them, and there does appear to be an overlap of the two taxa
in southeastern Peru. However, along the lower Amazon the resident popula-
tions are quite variable and appear to be intergrades between obscurior and
modestus. At the eastern extreme of the range of obscurior there is another exam-
ple of apparent overlap. A single specimen from Mexiana Island at the mouth of
the Amazon taken 13 November is modestus. Its range is separated from the
nearest populations of modestus in Maranhao by the presence of obscurior in
Para. It might possibly be a vagrant or migrant from further south, but the date
is quite late for a wintering bird.
Nominate modestus is found from the Urubamba valley in Peru east through
Beni, Bolivia, to Mato Grosso, Brazil; and north in the dry interior of eastern
Brazil to Maranhao, Piaui, and Pernambuco; and south to Sao Paulo and
TRAYLOR: TYRANT FLYCATCHERS 9
Parana. It has the shortest bill length of any form, 15 males having a range of
10-12 mm and an average of 10.8 mm. The closely related brevirostris, rec-
ognized by Zimmer but not by Pinto (1944, p. 283), ranges from central Bolivia
through Paraguay to western Rio Grande do Sul, Brazil, and western Uruguay,
and south to Buenos Aires, La Pampa, and Mendoza, Argentina. The main
distinction between brevirostris and modestus is the clear white wing-barring of
the former. This is important, because the wing bars of the obscurior and the
arenarum groups are generally dull gray, and it is possible to distinguish win-
tering brevirostris in the breeding range of obscurior with a good degree of
certainty. I have examined wintering brevirostris from Huanuco, Loreto, Junin,
and Puno, Peru, and Novo Olindo, Manaus, and Villa Bella Imperatriz, Amazo-
nian Brazil. It appears to be the only migratory taxon in the genus. Vertical
distribution and size variation in brevirostris are unique in the genus. Speci-
mens have been taken up to 2,300 m in Cochabamba and Tarija, Bolivia, and
populations from above 1,200 m are markedly longer-winged than lowland
ones. Wing lengths of six highland males are 72-82 mm (av. 76.5 mm), while 13
lowland males measure 69-73 mm (av. 70.8 mm). The highland birds are the
largest population within the genus, while the lowland are one of the smallest.
The highland population is not migratory.
Along the Amazon, from Manacapuru just west of Manaus to Santarem, lives
a variable population that seems to be intergrades between obscurior and mo-
destus. There is a fine series of 11 specimens from Obidos in the Carnegie Mu-
seum which shows the full extent of variation. As a whole the series is darker
gray on the breast than modestus, but not as dark or with as extensive a gray on
the belly as obscurior. The color of the belly is particularly variable, going from
clear pale yellow, as in modestus, to almost white. The extremes of variation are
not far from typical obscurior and modestus, respectively. CM 84649 looks like a
pure obscurior, with a dark gray breast extending over a pale whitish belly and
the throat grayish, whereas CM 84477 has a pale breast, whitish throat, and
yellow belly, only slightly darker on the breast than modestus. Single birds from
Manacapuru and Santarem fit into this series, as does a female from Villa Bella
Imperatriz in the American Museum. Bill length and wing length are also
intermediate between obscurior and modestus, particularly in males; females are
closer to obscurior. Comparative measurements are given in Table 1.
It seems, therefore, that despite the apparent geographical overlap between
obscurior and modestus in southeastern Peru and near the mouth of the Amazon,
the two taxa intergrade extensively along the middle Amazon, and they must be
Table 1. Comparative measurements (in mm) of wing and bill length in Sublegatus
obscurior and Sublegatus modestus and in the middle Amazon variable population.
6 6 N Wing Bill
obscurior 19 70-78 (74.1) 11.5-14 (12.7)
middle Amazon 6 69-74 (71.3) 11-13 (12.0)
modestus 15 65-72 (68.2) 10-12 (10.8)
99
obscurior 6 67-71 (68.8) 11.5-13 (12.2)
middle Amazon 7 67-73 (68.9) 11.5-13 (12.1)
modestus 9 65-68 (66.7) 10-11 (10.6)
10 FIELDIANA: ZOOLOGY
considered conspecific. The classification that I presently recognize for con-
tinental South America is:
Sublegatus arenarutn atrirostris
glaber
orinocensis
modestus obscurior (syn. sordidus, peruvianus)
modestus
brevirostris
Any advances in our understanding of this genus will have to come from
behavioral studies rather than the amassing of more skins. The most convenient
place to start would probably be Surinam, where marked ecological differences
exist between glaber and obscurior and their ranges are near. If investigations
using taped vocalizations and playbacks show that there are indeed two species
there, then the study can be extended to include the relations between modestus
and obscurior in Brazil and southeastern Peru.
Additional note: In Field Museum there are two specimens that do not fit the
above pattern, and for which I have neither a name nor an explanation. They are
from the upper Rio Branco in northwestern Brazil — a male taken 11 December at
Boa Vista and a female taken 12 March at Serra de Lua. In coloration they are
nearest modestus, although somewhat darker, but in size they are smaller than
even the smallest modestus (male wing 64 mm, bill 10 mm; female wing 61 mm,
bill 10 mm). They are far too small to be obscurior, which is where Hellmayr
(1927, p. 447) placed them. They almost certainly represent a breeding popula-
tion, because nominate modestus is on its breeding grounds in southern Brazil
in December. Considering the uncertainties within the genus, it would be
premature to try to name them now.
Suiriri suiriri/affinis
Suiriri suiriri suiriri and S. s. affinis had been treated as separate species until
Zimmer (1955, p. 18) suggested that they were conspecific. Zimmer pointed out
that all available specimens (19 in AMNH) from northern Paraguay were inter-
mediate in one degree or another between typical suiriri and typical affinis and
that this area was one of intergradation between the two taxa. Evidence of
intergradation was found north of this area in a specimen from Companario,
southern Mato Grosso. Meyer de Schauensee (1966, p. 384) followed Zimmer in
treating the two as conspecific. However, Short (1975, p. 283) stated that he had
failed to find any specimens showing intergradation and reverted to treating
suiriri and affinis as separate species.
I have examined the Paraguayan material in the American Museum and find
it exactly as described by Zimmer. The specimens are highly variable both in
size and color, and none is typical of either suiriri or affinis. Average wing length
is almost exactly intermediate, but bill length is more nearly like that of affinis
rather than that of suiriri as stated by Zimmer. The three localities in Paraguay
(fig. 3) at which these specimens were collected, La Fonciere ( = San Luis de la
Sierra), Zanja Moroti, and Belen (not listed by Zimmer) form a triangle about
100 km on each side, and Companario is another 150 km to the northeast.
Considering the extensive area over which intergradation occurs and the fact
that the whole population is affected, I consider that suiriri and affinis are con-
specific.
TRAYLOR: TYRANT FLYCATCHERS
11
Zimmer also discussed five specimens of affinis in the American Museum
which were separable from typical specimens by their short broad bills and
broad pale tips to the rectrices. I have examined these birds, which were all
taken from localities where typical affinis occurs. There is a sixth specimen, from
Tranqueira Maranhao, in Field Museum, which was described by Hellmayr
(1929, p. 329) and is identical with Zimmer's birds. There are other characters
that correlate with the short bill and pale tips to the rectrices. In typical affinis
the pale gray crown and nape are sharply separated from the pale olive back,
but in the short-billed birds, the gray extends over the upper back and changes
gradually into olive. In affinis, and even more in suiriri, the central pairs of
rectrices have narrow pale edgings on both webs, but in the short-billed birds,
there are no pale edgings at all, and the central rectrices are distinctly broader.
On the other hand, except for the short bill, measurements of the two groups are
identical.
Ordinarily the close correlation of two such discrete and unrelated characters
as the short bill and the coloration and shape of the rectrices would be strong
evidence that we have two sibling species. I think that eventually this will prove
to be true, but the possibility that the short-billed birds may somehow be
x suiriri — spec, examined
x intergrades
• affinis — spec, examined
0 affinis — from literature
Fig. 3. Localities of Suiriri suiriri suiriri (crosses) and S. S. affinis (closed circles)
from which specimens have been examined. Open circles are localities of affinis from
the literature, to show its range in southern Brazil. The population of northeastern Para-
guay and adjoining Mato Grosso is composed of intergrades between suiriri and affinis.
12 FIELDIANA: ZOOLOGY
related to the intergradation between affinis and suiriri cannot be ignored at this
time. The absolute bill length in the short-billed birds is the same as that in
suiriri, but the width is the same as in affinis, giving the bill a short, broad form
more like that of Sublegatus modestus. The broad rectrices without edgings,
however, are unlike either affinis or suiriri. A further complication is the status
of bahiae, a subspecies of affinis recorded from eastern Bahia, Pernambuco, and
eastern Piaui. Suiriri bahiae is like affinis in having a yellow belly and long
slender bill, but has the rump and upper tail coverts brown, in little contrast to
the back, lacks the yellow bases to the rectrices, and has a shorter wing. Zimmer
tentatively places two specimens from between Pindahyba and Gilbues, eastern
Piaui, in bahiae, despite their peculiar coloration. They have the belly white, as
in suiriri, but have dull outer webs to the outer rectrices and long slender bills as
in affinis. Zimmer thought they might be lipochrome-deficient specimens of
bahiae, but also noted that there was a typical specimen of affinis from Gilbues.
Without adequate series of specimens, particularly of bahiae, and, most impor-
tant, without field studies of the various taxa, it is useless to speculate.
Elaenia albiceps and E. parvirostris
Elaenia albiceps and parvirostris are two widespread species whose breeding
ranges for the most part replace each other in southern South America (fig. 4).
Elaenia albiceps is a temperate form found in the Andes from southwestern
Colombia to Tierra del Fuego and in the temperate lowlands of Argentina and
Chile. Elaenia parvirostris is a lowland species, breeding from central Bolivia
east to southeastern Brazil, and south to Buenos Aires and Cordoba, Argentina.
Their breeding ranges abut along the foothills of the Andes from central Boliva
to Tucuman and overlap in Santa Fe, San Luis, and Cordoba. Elaenia parvirostris
is probably wholly migratory and is found throughout tropical South America
east of the Andes in winter. The southern race of £. albiceps, chilensis, is also
migratory for the most part. It breeds in southern Bolivia, in Argentina from
Salta, Santa Fe, and Cordoba south to Tierra del Fuego, and in Chile, and
winters commonly north to the Amazon and east of the Andes. The populations
from the coastal areas of southern Chile must cross the Andes at some point on
migration, since there are no known coastal wintering birds. Worn and unsexed
specimens, as in any group of closely related Elaenia spp., may be difficult to
identify, but except for Olrog (1963, p. 269), no one has suggested that E.
albiceps and E. parvirostris are conspecific. However, in recent collections from
southern Bolivia, there are intergrades from intermediate altitudes along the
Andes, suggesting that there may be a local zone of hybridization in this re-
gion. The representatives of E. albiceps involved in this hybridization are a
sedentary population of chilensis which was previously unrecognized.
Fresh specimens of parvirostris and chilensis are readily separable on a number
of characters. Elaenia parvirostris is greener, less brownish above, and has the
white crown patch smaller and more completely concealed. Elaenia parvirostris
usually has three well-marked whitish wing bars, and E. a. chilensis, never more
than two. The form of the wing tip is an excellent character for distinguishing
migratory chilensis, not only from parvirostris but from other races of E. albiceps
in whose ranges it winters. As pointed out by Zimmer (1941a, p. 8), in these
chilensis the 10th primary is almost always longer than the fifth, whereas in all
other taxa of this complex, it is shorter. In size, chilensis is larger than parviros-
tris, although there is overlap in all dimensions. The populations of chilensis in
Males
chilensis, migrant
chilensis, Bolivia
albiceps
parvirostris
Females
chilensis, migrant
chilensis, Bolivia
albiceps
parvirostris
TRAYLOR: TYRANT FLYCATCHERS
Table 2. Measurements (in mm) in Elaenia spp.
N
26
8
7
14
Wing
76-82 (78.7)
77-81 (78.9)
80-87 (83.0)
69-76 (73.3)
10 71-74 (73.2)
7 71-75 (73.1)
6 78-85 (80.2)
9 68-73 (70.2)
Tail
61-71 (64.3)
60-75 (68.6)
72-78 (74.1)
58-66 (62.6)
58-63 (60.4)
58-64 (61.1)
71-77 (73.2)
57-62 (60.6)
Culmen
13-15 (14.2)
13-14 (13.7)
14-15 (14.6)
12-13 (12.6)
13-15 (14.1)
13-14.5 (13.7)
13-14.5 (13.5)
12-14 (12.9)
13
Tarsus
17.5-20.5 (19.3)
18.5-21 (19.1)
18.5-21 (19.4)
17-19 (18.0)
17.5-20 (18.4)
16.5-19 (17.5)
18.5-20 (19.0)
16.5-18 (17.6)
the mountains along the eastern slope of the Andes in southern Bolivia are
identical with the migratory chilensis in every way except that they lack the
distinctive wing tip. On the assumption that the more pointed wing of Argen-
tinian and Chilean birds is associated with their migratory habits, then the
Bolivian populations are probably sedentary. The latter intergrade with nomi-
nate albiceps in northern Chuquisaca and adjoining Cochabamba; albiceps is the
brownest race, with a large white crown whose lateral edges are washed with
brown, and is the largest race, with a proportionately long tail. Measurements
for the various taxa are given in Table 2.
These measurements point up the fact that parvirostris is smallest ( = shortest
winged), with a disproportionately short bill, and nominate albiceps by far the
largest, with an extra long tail. Specimens from Chuquisaca and Tarija have
been lumped in the Bolivian chilensis, but the more northern birds are slightly
larger and definitely longer tailed as they approach albiceps.
In Argentina there is positive evidence that E. parvirostris and £. albiceps
chilensis behave as distinct species. The points on the map (fig. 4) are actual
breeding localities, based on personally examined specimens or on accounts in
the literature. The breeding ranges overlap from Salta south to San Luis and east
to Santa Fe and eastern Cordoba, and I have found no suggestion of intergrada-
tion in any specimens from that area. Not only are the species distinguishable
on the morphological characters listed above, but they build different types of
nests. Hoy (1971, p. 160) found parvirostris nesting at San Lorenzo in the Lerma
Valley, Salta, where chilensis was common. He found parvirostris to be a bird of
tall woodland, which reached its upper limit at 1,500 m at San Lorenzo. Elaenia
chilensis occurred more in small trees and around agriculture, and was found
from 1,500 m up to 3,500 m. The nest of parvirostris was a small neat cup of straw
and plant fibers covered on the outside with moss and lichens, much like that of
a hummingbird. It was built on a large branch or on a fork and sat up on its
foundation. The nest of chilensis was built of dried grasses or straw and lined
with feathers; it was constructed on the outer branches of low bushes, with the
nest woven into their twigs. These descriptions agree with others from
elsewhere. Pereyra (1942, p. 229) and Eisentraut (1935, p. 428), writing of par-
virostris, mention the small nests covered with lichens on the outer branches of
the trees, and Goodall et al. (1957, p. 193) describe the nest of chilensis as built of
various plant fibers and lined with feathers and placed in small dense trees or
most especially in young pines. Despite Olrog's suggestions, there is no evi-
dence that parvirostris and chilensis are other than good sympatric species in
Argentina.
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18 FIELDIANA: ZOOLOGY
striaticollis occurs south and east of Victoria and at Pampas de Heath in south-
eastern Peru where the Madre de Dios crosses the Bolivian border. Further up
the same river at Manu, typical iohannis is found. I have examined the specimen
of striaticollis from Victoria, where it was taken alongside iohannis. It is a worn
skin, but it has the white lores and eye ring and unmarked wing coverts that are
diagnostic of striaticollis. Since the two taxa have been taken at the same locality,
Victoria, and replace each other along the Rio Madre de Dios, I consider them
distinct species. Being essentially allopatric or parapatric in distribution, they
constitute a superspecies.
Size has not been mentioned as a character because of the remarkable uni-
formity within the superspecies. There is no difference in absolute size, as
measured by wing length, between the species, nor is there any difference in
proportions, except for a slightly shorter tail in iohannis. Any identification
based on size would be suspect.
My classification is:
Hemitriccus striaticollis striaticollis (syn. obscuriceps)
griseiceps
iohannis (syn. amazonicus)
Ochthodiaeta signatus Taczanowski, 1874
In 1976, while reviewing Zimmer's manuscript notes on the Tyrannidae, I
found a short report, presumably prepared with the thought of publication, on
the status of Ochthodiaeta signatus. In it he pointed out the proper type locality
of signatus and suggested the possible transfer of the species to Knipolegus,
probably near cabanisi, a suggestion that subsequent collections have amply
confirmed. I was fortunate to find Zimmer's report and am grateful for his good
judgment, for it saved me the possibility of renaming the species.
Taczanowski (1874, p. 532) described Ochthodiaeta signatus on the basis of a
female and a juvenal in the Warsaw Museum, taken by Jelski at Auquimarca
and Ninabamba, respectively, Junin, Peru. Only the Ninabamba specimen re-
mains in Warsaw, and it was considered the type by Sztolcman & Domaniewski
(1927, p. 137), but, as Zimmer noted, Taczanowski (1889, p. 15) had already
designated the Auquimarca female as the type, so that Auquimarca must be
considered the type locality, even though the type is no longer extant. Zimmer
also drew attention to a third Jelski specimen recorded by Ridoutt (1941, p. 248)
in Museo Historia Natural "Javier Prado" in Lima, a female from Ninabamba.
When Zimmer analyzed the descriptions of Taczanowski and the color notes
sent him by Maria Koepke of the Javier Prado bird, he realized that signatus
differed from the species of Ochthodiaeta, not only in much smaller size but also
in most details of color, and he suggested that it was a Knipolegus, probably near
cabanisi whose females it most nearly resembled.
The three specimens taken by Jelski were the only ones known from 1873
until 1973, when a party from Louisiana State University, Museum of Zoology,
led by John O'Neill, collected a female and immature male in the Carpish area of
Huanuco. The party returned the following year and collected the first adult
male. This last specimen removed any doubt that they had a representative of
Knipolegus cabanisi; it was solid black with white wing linings, comparable with
the dark, slate gray male of cabanisi with similar wing markings. Subsequent
collecting by O'Neill, Peter Hocking for Field Museum, and John Fitzpatrick for
TRAYLOR: TYRANT FLYCATCHERS 19
the Museum of Comparative Zoology has taken signatus at San Jose de Lourdes
in Cajamarca, Abra Patricia on the road to Rioja in San Martin, and in the Panao
region of Huanuco. Altogether there are now four adult males, three adult
females, and three subadult males available for study.
Any doubt that the recently collected specimens are identical with Tacza-
nowski's signatus has been removed through the kindness of Charles Munn,
who made meticulous color notes, measurements, and color photographs of the
Ninabamba juvenal in the Warsaw Museum. Except that it appears slightly
more reddish brown on the upper parts, a character also found in juvenal K.
cabanisi, it is virtually identical in size and color with the northern Peruvian
specimens. The taxa will have to be known as:
Knipolegus signatus signatus — mountains of northern Peru from Cajamarca
to San Martin and south to Junin
Knipolegus signatus cabanisi — mountains from southern Peru to northern
Argentina
Wolters (1977, p. 182) has also recognized that signatus is a species of
Knipolegus, and in his list of species, he placed it next to cabanisi. Although I
include cabanisi as a subspecies of signatus, the two are quite distinct, and a case
can be made for calling them two species. Either way, they certainly form a
well-marked zoogeographical species, much closer to each other than to any
other Knipolegus. In the discussion below, I compare signatus with the better-
known cabanisi.
In both sexes, signatus is markedly darker than cabanisi. Males of signatus are
uniformly black rather than dark, slate gray, but in both taxa the inner webs of
the secondaries and of the proximal half of the primaries are broadly edged with
white. In females the upper parts of signatus are a darker, warmer olive-brown
than those of cabanisi, and the same is true of the heavy flammulations of the
underparts. Subadult males are similar to females. In female cabanisi, the outer
webs of the rectrices are narrowly edged rufous, and the inner webs are more
than half rufous, this color almost reaching the shaft at the base of the feathers.
In signatus, the edging on the outer webs is virtually obsolete, and that on the
inner webs is duller and darker and occupies much less than half the web. In
both taxa, females and subadult males have two well-marked wing bars that
vary from white to buff. This variation is not correlated with age or sex, but
seems to be dependent on wear. In two specimens that have a mixture of white
and buff feathers in the wing bars, the buff feathers are fresh and the white ones
worn.
There is little difference in measurements between the two taxa, except that
signatus has a distinctly longer bill in both sexes. The 10th (outermost) primary
in signatus is markedly shorter than that in cabanisi, being equal to or shorter
than the first rather than about equal to the fourth. The outer primaries of
signatus are broader than those of cabanisi, and the 10th lacks the slight notch
that is found in cabanisi. There is no sexual variation in the form of the pri-
maries. The sequence of plumages appears to be identical in cabanisi and sig-
natus. The juvenal plumage is similar to that of the adult female, except that
there is a slight rusty wash on the upper parts. The postjuvenal molt is in-
complete, involving only the body feathers. Subadult females molt into a plum-
age that is indistinguishable from that of the adult female. Males molt into a
subadult plumage that is also like that of the adult female, but they can always
20 FIELDIANA: ZOOLOGY
be distinguished by their greater size. The subadult plumage is worn for at least
a year, and the adult plumage is presumably assumed at the time of the first
regular postnuptial molt. It is not known if males breed in subadult plumage,
but those taken in the breeding season in Bolivia had testes as much enlarged as
fully plumaged males.
Moist montane forest is the preferred habitat for both signatus and cabanisi.
The former has been taken from 1,900 to 3,050 m in northern and central Peru,
and the latter from 1,100 to 2,500 m in Bolivia and northern Argentina.
Berlioz (1959, p. 217) described Knipolegus subflammulatus on the basis of four
males from Alto Palmar, Cochabamba, Bolivia. He characterized it as similar in
plumage to the female of cabanisi, but with males retaining the female plumage
at maturity, as in K. poecilurus, rather than assuming a gray, male-type plum-
age, as in cabanisi. I have examined three cabanisi from Alto Palmar, including
two full-plumaged males, and I think that Berlioz described subflammulatus from
subadult males of cabanisi. This was also the conclusion of Mayr (1971, p. 313),
who noted that Meyer de Schauensee had examined the types. Parkes, who first
suggested this identification of subflammulatus to Mayr, has shown me his cor-
respondence with Berlioz, who was quite willing to accept Parkes' suggestions
when he realized that young males of Knipolegus species have a first-year plum-
age similar to that of the adult female. The measurements given by Berlioz,
70-72 mm, are much too small for any age or sex of cabanisi. However, Dr.
Christian Erard has been kind enough to remeasure the type series for me, and
his wing measurements are 76, 77.5, 79, and 79.5 mm. The two smaller figures
are typical of female cabanisi, and the two larger are smaller than my immature
male cabanisi, but not significantly different.
Summary
Tyranniscus australis is shown to be a synonym of Phyllomyias sclateri. Phyl-
lomyias reiseri is shown to be a race of P. virescens. Extensive altitudinal varia-
tion in size is demonstrated in Zimmerius vilissimus. The genus Sublegatus is
shown to be composed of two species, arenarum and modestus. Suiriri suiriri and
S. affinis intergrade extensively in northern Paraguay and should be considered
conspecific. Elaenia albiceps and E. parvirostris are widely sympatric in Argen-
tina, but hybridize in Bolivia. Hemitriccus iohannis is shown to be a good species
and not a race of striaticollis. Ochthodiaeta signatus belongs in the genus
Knipolegus, where it is conspecific with cabanisi.
Acknowledgments
No study of this type would be possible without the generous cooperation of
one's colleagues. For their kind permission to examine the material in their
institutions or to borrow critical specimens, I would like to thank James Bond
and Frank B. Gill, Academy of Natural Sciences, Philadelphia; Wesley E. Lan-
yon, American Museum of Natural History, New York: William Belton, per-
sonal collection in Gramado, Rio Grande do Sul; Kenneth C. Parkes, Carnegie
Museum of Natural History, Pittsburgh; William H. Phelps, Jr., Coleccion
Phelps, Caracas; Claes C. Olrog, Instituto Miguel Lillo, Tucuman; Raymond A.
Paynter, Jr., Museum of Comparative Zoology, Cambridge; the late George H.
Lowery and John P. O'Neill, Museum of Zoology, Louisiana State University,
Baton Rouge; George E. Watson and Richard L. Zusi, National Museum of
TRAYLOR: TYRANT FLYCATCHERS 21
Natural History, Washington, D.C.; G. F. Mees, Rijksmuseum van Natuurlijke
Historie, Leiden; Carl Edelstam, Royal Natural History Museum, Stockholm;
and John S. Weske, personal collection on deposit at the American Museum of
Natural History, New York. George Lowery and John O'Neill were particularly
generous in allowing me to use recent, unstudied material from Peru which
they were working on themselves. I am also most grateful to Charles A. Munn,
who examined the type of Ochthodiaeta signatus in Warsaw, to Christian Erard
for measuring the type series of Knipolegus subflammulatus, and to Eugene
Eisenmann and John Weske who shared their field notes on several species.
Throughout the preparation of this paper, I have benefitted from discussion
with my colleagues at Field Museum, Emmet R. Blake and John W. Fitzpatrick,
although they cannot be blamed for any shortcomings. And, finally, I owe
special thanks to my wife Marjorie who sacrificed many sunny afternoons re-
cording measurements in dark bird ranges when she might have been out
enjoying new sights.
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22 FIELDIANA: ZOOLOGY
Pinto, O. M. de O., and E. A. de Camargo. 1961. Resultados Ornithologicos de quatro
recentes Expeditoes do Departmento de Zoologia ao nordeste do Brasil, com a Descricao
de seis novas Subspecies. Arq. Zool. S. Paulo, 11: 193-284.
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Am. Mus. Nat. Hist., 154: 163-352.
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