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.3 FIELDIANA ZOOLOGY 

Published by 
CHICAGO NATURAL HISTORY MUSEUM 

Volume 31 September 8, 1950 No. 37 

A STUDY OF THE SPOTTED TINAMOUS 

Genus Nothura 

BOARDMAN CONOVER 
Research Associate, Division of Biros, 1936-50 > 

My interest in the genus Nothura began in 1923, when I first 
became acquainted with some of its members in the field, and 
since then I have acquired as many specimens as possible, with the 
hope of making a special study of the group. In 1942, Dr. C. E. 
Hellmayr and I published a review of the Family Tinamidae (Field 
Mus. Nat. Hist., Zool. Ser., 13, (1), pp. 6-114), including, of course, 
this genus, but we were handicapped by the impossibility, because 
of the war, of comparing certain critical specimens in European and 
American collections. In 1948, I gathered together from the mu- 
seums of this country as many specimens as possible, and I also 
borrowed critical examples from Europe and South America. Alto- 
gether, more than four hundred adult specimens have been examined, 
representing all but one of the known forms. Five species (one 
doubtful), comprising fifteen races, have been recognized. Of 
these, I had for study eight types (in addition to the types of two 
unrecognized forms). Topotypical material of six additional forms 
was also available. I am indebted to the following institutions for 
the loan of specimens: the American Museum of Natural History, 
Carnegie Museum, the Zoologische Sammlung des Bayerisches 
Staates, the Museum of Comparative Zoology, the Philadelphia 
Academy of Sciences, Princeton University, the United States 
National Museum, and the Museum of Vertebrate Zoology, Uni- 
versity of California. 

There has long been a lack of information about the exact ranges 
of the species and subspecies of the genus. There are five species, 
and in several localities their ranges meet (chacoensis and maculosa 
paludivaga, Paraguayan Chaco) or even overlap (darwinii and macu- 
losa, Rio Negro Valley and probably western Argentina; boraquira 
and chacoensis, Paraguayan Chaco; minor and maculosa, Brazil). 

1 Deceased May 5, 1950. 
No. 647 339 iHh LIBRARY OF THE 

I ^ru*^-"*- SEP 2 H950 



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JjfottXf UNIVERSITY OF ILLINOIS 



340 FIELDIANA: ZOOLOGY, VOLUME 31 

1 hope that this paper may correct some of the misconceptions 
as to the number of species and races, and their ranges, and may also 
show the parts of South America from which additional material 
is most urgently needed. It is especially hoped that the paper 
may help others to identify their material more easily than has been 
possible in the past. 

The genus divides into four groups or complexes: boraquira, 
minor, darwinii, and maculosa. The first two are monotypic, the 
third consists of one species divided into five races, and the fourth 
comprises a full species of seven races together with another form 
that must be considered at least an incipient species, until more is 
known of its range and its relationship to the neighboring races of 
maculosa. 

A. Lesser under wing coverts barred with dusky and inner webs of primaries 

immaculate boraquira 

B. Lesser under wing coverts unbarred. 

a. Toes and tarsus relatively short. Middle toe with claw generally under 

28 mm. N. maculosa cearensis has a toe measuring 27 mm., but a short 
wing (110 mm.). 

1. Smaller, wing not over 120; middle toe with claw 21-24 mm minor 

2. Larger, wing over 120; middle toe with claw over 24 but generally not 

over 28 mm darwinii 

b. Toes and tarsus relatively long and stout. Middle toe with claw generally 

over 28 mm. except in N. m. cearensis where the wing is 110 and the 
middle toe 27 mm maculosa and chacoensis 

Nothura boraquira Spix. Marbled Tinamou. 

Tinamus boraquira Spix, Av. Spec. Nov. Bras., 2, p. 63, pi. 79, 1825 "in 
campis petrosis districtus adamantini"[= Minas Geraes], Brazil (type in 
Munich Museum). 1 

Nothura marmorata G. R. Gray, List Spec. Bds. Brit. Mus., 5, p. 104, 1867 
Bolivia (cotypes in British Museum); 2 Salvadori, Cat. Bds. Brit. Mus., 
27, p. 561, pi. 18, 1895 Bolivia. 

Nothura boraquira Hellmayr, Field Mus. Nat. Hist., Zool. Ser., 12, p. 478, 
1929 Quixada and Jua, near Iguatu, Ceara; Peters, Bds. World, 1, 

'Hellmayr and Conover (Field Mus. Nat. Hist., Zool. Ser., 13, (1), p. 107, 
footnote) called attention to the fact that Reinhardt (Vidensk. Medd. Naturhist. 
Foren., 1870, p. 51) denied that this species occurs in Minas Geraes, and they 
therefore concluded that the type locality was probably erroneous. However, 
recently I have examined ten specimens from Janauba, Minas Geraes, so Spix's 
type locality must stand. 

2 Some years ago Dr. Hellmayr compared two specimens in my collection 
(Nos. 4634 and 6828), from Buena Vista, Santa Cruz, Bolivia, with the cotypes 
of Nothura marmorata G. R. Gray in the British Museum. Both, he stated, were 
very like the types, No> 4634 being a trifle lighter and No. 6828 a trifle darker on 
the upper parts. 



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3 1 
Y,3I CONOVER: SPOTTED TINAMOUS 341 

f p. 27, 1931 (range); Laubmann, Verh. Orn. Ges. Bay., 20, pp. 281, 282, 
1934 west of Puerto Casado, Paraguayan Chaco; Pinto, Cat. Aves 
Bras., 1, p. 13, 1938 Parnagua, Piauhy; Hellmayr and Conover, Field 
Mus. Nat. Hist., Zool. Ser., 13, (1), p. 107, 1942 (bibliog.; range); Bond 
and de Schauensee, Proc. Acad. Nat. Sci. Phila., 95, p. 171, 1943 Buena 
Vista, Chiquitos, Bolivia. 

Nothura spixi Miranda-Ribeiro, Rev. Mus. Paul., 23, p. 704, pis. 3 and 4a, 
1938 (new name for Tinamus boraquira Spix on grounds of purism). 

? Nothura schreineri Miranda-Ribeiro, Rev. Mus. Paul., 23, p. 702, pi. 2, 
1938 Minas Geraes (type in Museo Nacional, Rio de Janeiro) ; Hellmayr 
and Conover, Field Mus. Nat. Hist., Zool. Ser., 13, (1), p. 95 (footnote), 
1942 (disc); Zimmer and Mayr, Auk, 60, p. 250, 1943 (disc). 

Nothura maculosa cearensis (not of Naumburg) Lamm, Auk, 65, p. 263, 
1948 Campena Grande, Paraiba, Brazil (specimen examined). 

Range. From northeastern Brazil (Piauhy, Ceara, Parahyba) 
south to Minas Geraes; also eastern Bolivia (Dept. Santa Cruz) 
and the drier parts of the Paraguayan Chaco (Colonia Fernheim 
and west of Puerto Casado). 

Characters. Nothura boraquira can always be identified by the 
fact that its lesser under wing coverts are distinctly barred with 
dusky, a feature that distinguishes it from all the other species in 
the genus. Also, the inner webs of all the primaries are unbarred. 

Specimens from Minas Geraes are very dark (fuscous) on the 
upper parts and in this respect, surprisingly, are nearer to birds 
from Bolivia and Paraguay than they are to those of northeastern 
Brazil (Bahia, Piauhy, Ceard, etc.). Specimens from these latter 
localities tend to be more rufescent dorsally. Out of twenty-two 
examined, seven are indistinguishable from the topotypical series, 
five are slightly lighter-colored, and ten are distinctly more rufescent. 
On the other hand, Bolivian and Paraguayan birds are dorsally 
very like the Minas specimens, but tend to have more buffy upper 
wing coverts and secondaries, this feature showing up most strongly 
in Bolivian examples. However, all these differences are so slight 
and the individual variation is so great that the recognition of any 
races seems hardly justified. 

Downies. A downy specimen from the Paraguayan Chaco differs 
from several of the maculosa complex by having the sides of the 
head back of the eye, the back and sides of the neck, and the light 
streaks on top of the head white or buffy white, not ochraceous 
orange. 

Attention should be called here to the fact that in the Paraguayan 
Chaco west of Puerto Casado Nothura boraquira is found living to- 
gether with N. (maculosa?) chacoensis. 



342 FIELDIANA: ZOOLOGY, VOLUME 31 

Specimens examined. 66: Brazil (Manga, Maranhao, 1; Quixada, 
Ceara, 1; Jua, Ceard, 1; Santa Lucia, Parahyba, 1; Campena Grande, 
Parahyba, 1; Feoriano, Piauhy, 1; Corrente, Piauhy, 10; Santa Rita, 
Bahia, 5; Lamarao, Bahia, 1; Janauba, Minas Geraes, 10); Bolivia, 
Santa Cruz (Cercado, 2; Buena Vista, 16; unspecified, 1); Paraguay, 
Chaco (265 km. west of Puerto Casado, 7; 120 km. west of Puerto 
Casado, 1; Orloff, 5; Colonia Fernheim, 2). 

Nothura minor Spix. Least Spotted Tinamou. 

Tinamus minor Spix, Av. Spec. Nov. Bras., 2, p. 65, pi. 82, 1825 "in campis 

prope pagum Tejuco" [=Diamantina], Minas Geraes, Brazil (type, in 

Munich Museum, examined). 
Nothura assimilis G. R. Gray, List Spec. Bds. Brit. Mus., 5, p. 105, 1867 

"South America" (type in British Museum). 
Nothura media (not Tinamus medius Spix) Salvadori, Cat. Bds. Brit. Mus., 

27, p. 563, 1895 Itarare and Chapada. 
Nothura minor Peters, Bds. World, 1, p. 27, 1931 (range); Pinto, Cat. Aves 

Bras., 1, p. 13, 1938 Sao Paulo; Hellmayr and Conover, Field Mus. 

Nat. Hist., Zool. Ser., 13, (1), p. 106, 1942 (descr.; range; bibliog.). 

Range. From Minas Geraes (Diamantina, Lagoa Santa, Agua 
Suja near Bagagem) south to Sao Paulo (Itatinga, Itapetininga, 
etc.) and Matto Grosso (Chapada). 

Characters. This species can be identified by its very small size 
(wing 100-120) combined with its short toes (middle toe with claw 
21-24 mm.). While N. maculosa cearensis has as small a wing 
(110 mm.), its middle toe and claw are larger (27 mm.). N. minor 
also has much more finely barred upper wing coverts than any of 
the other species and, in the rufous phase, the intense chestnut- 
rufous upper parts are diagnostic. 

In this species there would appear to be two color phases, one 
chestnut-rufous and the other rufescent buffy. However, in a large 
series it is very probable that the gap between the two would be 
entirely closed by intermediate specimens, as there is great variation 
even in the small series examined. 

In the dark phase the upper parts are chestnut-rufous, vermicu- 
lated and blotched with black. Six of the ten birds examined fall 
into this group, but no two are alike. Two are heavily blotched 
with black and another is finely vermiculated with this color, while 
a fourth is almost solid chestnut-rufous from mantle to rump, except 
for the bufly edges to the feathers and a very occasional spot of black. 
The other two falll between these extremes. Two are whitish buff 
below, three are buff washed with rufous, and one is rufous buff. 



CONOVER: SPOTTED TINAMOUS 343 

The feathers of the chest have dark shaft streaks, very narrow in 
four specimens, much wider in the other two. 

The light phase (to which the type belongs) has the upper parts 
rufescent buff, vermiculated and blotched with black. The under 
parts are light buff with narrow dark shaft streaks in the feathers 
of the lower neck and chest. 

In both phases the upper wing coverts and flanks are much more 
finely barred with black than in any form of maculosa or darwinii. 

Specimens examined. 10: Minas Geraes ("Tejuco"=Diaman- 
tina, 1 [the type] ; Agua Suja, near Bagagem, 2) ; Sao Paulo (Itape- 
tininga, 2; Itatinga, 1); Matto Grosso (Serra da Chapada, 1; 
Chapada, 3). 

Nothura darwinii G. R. Gray. 

Many authors have considered darwinii to be a form of maculosa, 
although long ago W. H. Hudson (Proc. Zool. Soc. Lond., 1872, 
p. 534) pointed out the fact that the two live together in the lower 
Rio Negro Valley. According to his account the "Perdiz comun" 
( Nothura maculosa nigroguttata) frequents the grassy, moister, more 
fertile plains near the river, while the "Perdiz chico" ( N. d. darwinii) 
lives in the higher, drier, brush-covered tablelands bordering the 
valley. I have compared specimens of typical darwinii taken by 
Hudson in the lower Rio Negro Valley ("I advanced altogether not 
much over a hundred miles from the sea") with examples of N. macu- 
losa nigroguttata from Chimpay and Valcheta, Rio Negro, and they 
are very different both in coloration and in the length and stoutness 
of their tarsi and feet. There can be no doubt that the two are 
specifically distinct. Also, I have examined three specimens of 
maculosa from well within the range of N. darwinii salvadorii. Two 
were from Mendoza and one was from Rosario de Lerma, Province 
of Salta. 

That darwinii likes the drier regions seems to be borne out by 
the ranges of its different forms. The species appears to range from 
the Rio Chubut north through the lower Rio Negro Valley to the 
Rio Colorado and then west to Mendoza and north again through 
the western plains of Argentina into the highlands of Bolivia and 
southern Peru, where it is found at altitudes ranging from 6,000 
to 12,000 feet. It has also been supposed to inhabit southern 
Patagonia, but as stated under the nominate form no specimens have 
ever been taken and recorded by competent ornithologists south of 
northern Chubut. 



344 FIELDIANA: ZOOLOGY, VOLUME 31 

The different forms of this species can be distinguished from those 
of maculosa by the shorter, stubbier legs and feet. Measurements 
do not give the entire picture, but in darwinii the length of the 
tarsus usually is from 28 to 34 mm. and that of the middle toe 
and claw from 24 to 29 mm., while in maculosa they are from 34 
to 40 and 28 to 35 mm., respectively. In N. maculosa cearensis, 
however, the type and only known specimen measures tarsus 29, 
middle toe with claw 27; but the race is a very small one, with a 
wing length of only 110 mm. Moreover, no known form of darwinii 
is found within hundreds of miles of Ceara. 

An additional character is the tendency in Darwin's Tinamou 
toward having unbarred inner webs of the outermost primaries. In 
the nominate race the apical half of the first and the apical third 
of the second primary are generally immaculate or the barring of 
the inner web is practically obsolescent. This tendency increases 
in the races toward the north until in the form agassizii, from south- 
ern Peru, the inner webs of the four outermost remiges are un- 
barred or nearly so. Still another feature of this species is the more 
loosely barbed, more hair-like feathers of the rump and sides, an- 
other character that tends to become more noticeable in the northern 
races. 

Nothura darwinii darwinii G. R. Gray. Darwin's Tinamou. 

Nothura darwinii G. R. Gray, List Spec. Bds. Brit. Mus., 5, p. 104, 1867 

Bahia Blanca, southern Buenos Aires Province (type in British Museum) ; 

Sclater, Proc. Zool. Soc. London, 1872, p. 547 Rio Negro. 
Nothura darwinii Salvadori, Cat. Bds. Brit. Mus., 27, p. 562, pi. xix, 1895 

part, Bahia Blanca and Rio Negro (bibliog.). 
Nothura darwinii darwinii Peters, Bds. World, 1, p. 27, 1931 (range). 
Nothura maculosa darwinii Hellmayr and Conover, Field Mus. Nat. Hist., 

Zool. Ser., 13, (1), p. 107, 1942 Bahia Blanca, Rio Colorado, Rio Negro 

and (?)Santa Cruz. 

Range. The high, dry country from eastern La Pampa, Bahia 
Blanca, and the Rio Colorado south through the lower Rio Negro 
Valley to at least the Rio Chubut. 

No properly identified specimens of this form have ever been 
recorded, as far as I can ascertain, from south of the Golfo Nuevo, 
Chubut, or from more than a hundred miles inland. 

Characters. Typical darwinii is characterized by its small size 
and gray coloration. The dorsal surface is gray, finely vermiculated 
with black, with few transverse markings of ferruginous and black. 
The lower neck and chest are pale grayish brown with narrow, trans- 



CONOVER: SPOTTED TINAMOUS 345 

verse, dark markings. The flanks are narrowly but thickly barred 
with dusky. 

N. d. darwinii is nearest to N. d. salvadorii but is paler and grayer 
above, with the vermiculations finer and the black and ferruginous 
transverse markings fewer. The lower neck and chest are grayer, 
with the dark markings less conspicuous. The abdomen is slightly 
lighter, less buffy. This form differs from N. maculosa nigroguttata 
(the race of the maculosa complex that shares its range, at least in 
part) by having the general tone of its coloration gray, not yellow; 
the light edges of the feathers of the upper parts white, not yellow 
or yellowish white; the light vermiculations of the dorsal surface 
grayish or ferruginous, not yellowish ochraceous; the dark markings 
of the lower neck and chest in the form of crossbars on a light 
reddish brown background, not rounded spots or longitudinal streaks 
on a yellowish background; and the abdomen much whiter, less 
yellowish. The size is also smaller and the tarsus and toes are 
much shorter: wing 130-137, against 138-153; tarsus 31-33, against 
37-40; middle toe with claw 25-27, against 30-34 mm. 

In the maculosa complex, this form most nearly resembles in 
coloration the race N. m. submontana, whose range it or N. d. 
salvadorii may overlap. However, in darwinii the upper parts are 
purer, less olivaceous gray, much more finely vermiculated and less 
blotched with black. The feathers of the flanks and thighs are more 
hair like. It is also smaller, wing 130-137, against 154 mm., with 
the tarsus and toes much shorter and lighter. 

Specimens examined. 10: La Pampa (Laguna Colorado Grande, 
Dept. Caleu-Caleu, 2); Buenos Aires (San Bias, 1); Rio Negro 
("not much over 100 miles from the sea," 4); Chubut (Golfo Nuevo, 
1; between Gaiman and Trelew, 1); "Patagonia," 1. 

Nothura darwinii salvadorii Hartert. Salvadori's Tinamou. 

Nothura salvadorii Hartert, in Hartert and Venturi, Nov. Zool., 16, p. 266, 
1909 Arenal, Salta, Argentina (type, in Tring Collection, now in the 
American Museum of Natural History, New York, examined). 

Nothura boraquira (not of Spix) Salvadori, Cat. Bds. Brit. Mus., 27, p. 561, 
1895 part, western Argentina near Mendoza. 

Nothura darwini mendozensis Chubb, Bull. Brit. Orn. Club, 38, p. 31, Dec. 29, 
1917 Mendoza, Argentina (type in British Museum); Wetmore, Bull. 
U. S. Nat. Mus., 133, p. 39, 1926 part, Tunuyan, Mendoza and Cordoba 
(specimen examined). 

Nothura darwinii salvadorii Peters, Bds. World, 1, p. 27, 1931 (range). 



346 FIELDIANA: ZOOLOGY, VOLUME 31 

Nothura maculosa salvadorii Laubmann, Verh. Orn. Ges. Bay., 20, p. 280, 
1934 Salta; Hellmayr and Conover, Field Mus. Nat. Hist., Zool. Ser., 
13,- (1), p. 97, 1942 part, except northern Neuquen (Chos-Malal). 

Nothura maculosa mendozensis Laubmann, Verh. Orn. Ges. Bay., 20, p. 280, 
1934 Mendoza. 

Range. Western Argentina from Salta and Tucuman south to 
Mendoza (Tunuyan) and east to western Santiago del Estero 
(Lavalle), western Cordoba (Sierra de Cordoba; Cosquin) and central 
La Pampa. 

Characters. Very close to the typical race but general tone of 
the dorsal surface darker, more ferruginous, less grayish. The 
upper parts are more coarsely vermiculated and have more numerous 
and heavier transverse markings of ferruginous and black; the dark 
markings on the lower neck and chest are more conspicuous and in 
general take the form of longitudinal streaks rather than transverse 
bars; the dark bars on the flanks are coarser and the abdomen is 
slightly more buffy. Averages larger, wing 130-152 mm. 

Differs from N. d. boliviana by having the upper parts and the 
lower neck and chest darker and grayer, much less rufescent, the 
abdomen lighter, less buffy, and the flanks perhaps slightly more 
heavily barred. Size larger, wing 130-152, against 126-138 mm. 

Remarks. The specimens from San Juan, surprisingly, are notice- 
ably lighter dorsally than those from Salta, Tucuman, and even 
Tunuyan, Mendoza. They rather resemble in coloration the two 
lightest of four specimens of N. d. boliviana examined from Tarija, 
Bolivia. It may be that west of the range of salvadorii, in the foot- 
hills of the Andes, there is another race of darwinii, lighter in colora- 
tion and somewhat resembling the form boliviana. 

Of seven specimens with the locality marked only as Mendoza, 
one, a bird collected by Chapman and Miller in 1916, resembles the 
San Juan series, while the other six are darker and very like a series 
that I collected in Tunuyan, Mendoza. These six darker specimens, 
however, may not have come from the immediate vicinity of the 
city of Mendoza, as they were collected by Carlos Reed, who, as 
I know from personal experience, sometimes bought specimens in 
the market. Chapman and Miller's example, therefore, may more 
truly show the characters of the birds from near that city. 

Specimens examined. 44: Tucuman (Tapia, 1; unspecified, 1); 
Salta (Arenal, 1; Rosario de Lerma, 3); Santiago del Estero (Lavalle, 
4); San Juan (Media Agua, 2; Angaco Sud, 12); Mendoza (Mendoza, 
7; Tunuyan, 8); Cordoba (Cordoba, 1; El Carrizal, Sierra de Cordoba, 



CONOVER: SPOTTED TINAMOUS 347 

1; Cosquin, 1); San Luis (unspecified, 1); La Pampa ("Region 
central," 1). 

Nothura darwinii boliviana Salvadori. Bolivian Tinamou. 

Nothura boliviana Salvadori, Cat. Bds. Brit. Mus., 27, p. 561, 1895 Bolivia 
and Cinti [=Camargo] (cotypes from "Bolivia" in British Museum). 

Nothura maculosa agassizii (not of Bangs) Peters, Bds. World, 1, p. 26, 1931 
part, highlands of southwestern Bolivia. 

Nothura maculosa boliviana Hellmayr and Conover, Field Mus. Nat. Hist., 
Zool. Ser., 13, (1), p. 97, 1942 (range; descr.); Bond and de Schauensee, 
Proc. Acad. Nat. Sci. Phila., 95, p. 171, 1943 Tiraque, San Lorenzo. 

Range. Highlands of southwestern Bolivia from Cochabamba 
south to Tarija at elevations from 6,000 to 11,500 feet. 

Characters. This race of darwinii is characterized by its small 
size, rufescent upper parts, ochraceous abdomen and scantily barred 
flanks. Its feathers are very hairlike and the first four primaries 
usually have little or no barring on their inner webs. 

Closest to N. d. agassizii, from which it differs by being more 
rufescent above and more ochraceous below. It has less barring 
on the flanks, and the dark markings on the neck, chest and upper 
parts are much more reduced. Size much smaller, wing 126-138 
(against 130-147), tarsus 28-34, middle toe (with claw) 24-28. 

Differs from N. d. salvadorii by being lighter in coloration, more 
rufescent on the upper parts, lower neck and chest, more ochraceous 
on the lower breast and abdomen, and less barred on the flanks. Size 
smaller, wing 126-138 (against 130-152). 

Remarks. Before the war two specimens in my collection, Nos. 
6830 and 6831, from Tiraque, Cochabamba, Bolivia (alt. 10,000 
feet), were compared by Dr. Hellmayr with the cotypes in the 
British Museum. According to him these specimens "agree perfectly 
in coloration except that your birds, due probably to their being in 
fresher plumage, have the white lateral edges to the interscapulars 
more pronounced." As several forms of Nothura are known from 
Bolivia, I suggest Tiraque, Cochabamba, as a restricted type 
locality for N. boliviana Salvadori. T. Bridges, who collected the 
cotypes, is known to have traveled in that region and his specimens 
could well have come from that vicinity. 

Specimens examined. 33: Bolivia (Tiraque, Cochabamba, 9; 
Vacas, Cochabamba, 5; Cerro San Benito, Cochabamba, 2; Zotora, 
Cochabamba, 2; Chocaya, Cochabamba, 1; Chuchacancha, Cocha- 



348 FIELDIANA: ZOOLOGY, VOLUME 31 

bamba, 4; Cochabamba, 2; Mizque, Cochabamba, 1; Coloni, Co- 
chabamba, 3; San Lorenzo, Tarija, 3; Tarija, 1). 

Nothura darwinii agassizii Bangs. Agassiz's Tinamou. 

Nothura agassizii Bangs, Proc. Biol. Soc. Wash., 23, p. 107, June 24, 1910 
Moho, northern border of Lake Titicaca, Peru (type in Museum of Com- 
parative Zoology, Cambridge, Mass.). 

Nothura maculosa agassizii Peters, Bds. World, I, p. 26, 1931 part, highlands 
of southeastern Peru and northwestern Bolivia; Hellmayr and Conover, 
Field Mus. Nat. Hist., Zool. Ser., 13, (1), p. 96, 1942 (range; descr.); Bond 
and de Schauensee, Proc. Acad. Nat. Sci. Phila., 95, p. 170, 1943 Calli- 
pampa, Oruro, Bolivia. 

Nothura maculosa oruro Bond and de Schauensee, Not. Nat. Acad. Sci. Phila., 
No. 93, p. 2, Oct. 14, 1941 Callipampa, Lake Poopo, Oruro, Bolivia, 
alt. 12,200 feet (type, in the Academy of Natural Sciences of Philadelphia, 
examined). 

Range. Highlands of extreme southeastern Peru from the south- 
ern Cuzco region (where it is said to intergrade with peruviana) 
and the Titicaca basin, south through western Bolivia at least to 
Lake Poopo (Callipampa, Oruro), where specimens show certain 
characters that indicate intergradation with boliviana. 

Characters. The chief characters of this race are the very hair- 
like texture of the feathers, the large size, and the yellowish rufescent 
coloration of the upper parts, which are also conspicuously blotched 
with dark brown. The first four primaries have their inner webs 
immaculate or very scantily barred with buff. Wing 130-147, 
tarsus 29-35, middle toe with claw, 24-29. 

Nothura darwinii agassizii is closest in coloration and size to 
N. d. salvadorii, from which it differs by having the upper parts 
yellower and more rufescent, less grayish. It also has the dark 
markings of the upper parts, lower neck and chest larger and more 
conspicuous and the barring on the flanks heavier. The feathers 
on the flanks and thighs are also much more hair-like. 

It is much less rufescent than boliviana and has the dark markings 
both above and below larger and more conspicuous. The abdomen 
is paler, less ochraceous. Size larger (wing 130-147 against 126- 
138). 

No specimens of peruviana have been available for comparison 
and there may be no difference between the two. However, Ber- 
lepsch and Stolzmann (Ornis, 13, p. 105, 1906) seem to think that 
a specimen from Ptmo was not the same as others from Santa Ana, 
Urubamba Valley, Cuzco, Peru. If a difference does exist it would 



CONOVER: SPOTTED TINAMOUS 349 

seem that agassizii is lighter, less ochraceous on the abdomen, and 
larger in size (wing 130-147 against 126-133). 

Specimens examined. 24: Peru (Puno, Puno, 7; Lake Titicaca, 
3; Zotorani, Puno, 3; Sorapa, Puno, 1; Chucuito, Puno, 1); Bolivia 
(Desaguadero, La Paz, 2; Mercado, La Paz, 3; Guaqui, La Paz, 1; 
Callipampa, Lake Poopo, Oruro, 3). 

Nothura darwinii peruviana Berlepsch and Stolzmann. Peruvian 
Tinamou. 

Nothura maculosa peruviana Berlepsch and Stolzmann, Ornis, 13, p. 101, 
1906 Santa Ana, Urubamba Valley, Cuzco, Peru (type in collection of 
Count Branicki, formerly in Warsaw Museum, now probably destroyed); 
Peters, Bds. World, 1, p. 26, 1931 (range); Hellmayr and Conover, Field 
Mus. Nat. Hist., Zool. Ser., 13, (1), p. 95, 1942 (range; descr.). 

Range. Southeastern Peru in the Department of Cuzco. 

Characters. No specimens of this race were available for examina- 
tion. From the remarks of others it would seem to be very similar 
to, if not identical with, N. d. agassizii. It is said, however, to be 
smaller (wing 126.5-133) and to have a darker abdomen. The inner 
webs of the primaries are immaculate, rarely barred with reddish 
buff. 

Nothura maculosa Temminck. 

Nothura maculosa, as stated before, differs from darwinii by its 
stouter and longer tarsi and toes. 

This species covers a greater extent of territory and lives in a 
wider variety of climates than the preceding one. For these reasons 
it breaks up into more forms. However, since the range is practically 
continuous from the Rio Negro, Argentina, to at least Minas Geraes, 
Brazil, the boundaries of the races are often hard to define, as most 
forms grade gradually from one to the other. Individual variation, 
which is very great, adds to the confusion. When more specimens 
are at hand from such places, among others, as central Argentina 
(Santiago del Estero to La Pampa) the ranges of some of the sub- 
species will undoubtedly have to be revised. 

In this paper Nothura chacoensis is kept in the maculosa complex 
but given specific status. The truth or falsity of this concept prob- 
ably will have to be worked out by ornithologists in the field. How- 
ever, it seems to me that a very good case can be made out for the 
belief that the parent stock of this species was long ago completely 
cut off from all other forms of maculosa by the flood basins of the 



350 FIELDIANA: ZOOLOGY, VOLUME 31 

Pilcomayo and Paraguay rivers. Later, as these basins became silted 
in, spotted tinamous from the east bank of the Paraguay probably 
drifted across and developed into the form here named N. maculosa 
paludivaga. That this separation lasted long enough for speciation 
to occur in chacoensis is proved, I believe, by the fact that in this 
same arid Chaco is found another tinamou, Eudromia mira, which 
was probably cut off from its parent stock at the same time. This 
E. mira is very distinct from its only known relative, E. elegans, 
which inhabits the more arid parts of Argentina, from eastern 
Tucuman and Santiago del Estero south. 

Nothura maculosa nigroguttata Salvadori. Patagonian Spotted 
Tinamou. 

Nothura nigroguttata Salvadori, Cat. Bds. Brit. Mus., 27, p. 560, 1895 
"Central Pampas, Argentine Republic" (type from Choele Choel, Rio 
Negro Territory, Argentina, formerly in Warsaw Museum [cf. Stolzman 
and Domaniewski, Ann. Zool. Mus. Pol. Hist. Nat., 6, p. 98, 1927], now 
probably destroyed). 

Nothura maculosa (not of Temminck) Wetmore, Univ. Calif. Pub. Zool., 24, 
p. 410, 1926 Valcheta, Rio Negro, specimen examined. 1 

Nothura maculosa nigroguttata Peters, Bds. World, 1, p. 27, 1931 part, Rio 
Negro only; Deautier, Obr. Cine. Mus. La Plata, 1, p. 147, 1935 part, 
Bahia de San Bias; Hellmayr and Conover, Field Mus. Nat. Hist., Zool. 
Ser., 13, (1), p. 100, 1942 part, Chimpay, Rio Negro and Collon Cura, 
Neuquen. 

Range. From the valley of the Rio Negro and perhaps the Rio 
Colorado south an unknown distance, but probably as far as the 
Rio Chubut; and from near the Atlantic coast west to the western 
part of the Territory of the Rio Negro and southeastern Neuquen, 
where it appears to intergrade with N. m. submontana. 

Characters. The distinctive feature of this race is its rather 
bright yellow and black appearance. When compared with a series 
of annectens from the provinces of Buenos Aires and Cordoba, the 
seven specimens from the Rio Negro Territory have the upper parts 
lighter and much yellower; the upper wing coverts with the light 
bars wider and yellower and the dark bars narrower; the flanks 
slightly less heavily barred; the lower neck and chest less heavily 
spotted; and the under parts brighter, more ochraceous buff. 

1 Wetmore speaks of this specimen as being very immature, although it was 
taken on May 7, which is late fall for those latitudes. While it has the appearance 
of a bird of the year, it seems fully grown, having a wing of 139 mm. The example 
agrees in coloration with the series from Chimpay. 



CONOVER: SPOTTED TINAMOUS 351 

Nothura m. nigroguttata differs from N. m. submontana by its 
lighter, yellower, much less olivaceous appearance. It is also much 
darker and more yellowish, less grayish, than N. d. darwinii, whose 
range it shares in part, and the upper parts are more blotched and 
much less vermiculated. 

Remarks. The name nigroguttata heretofore has generally been 
assigned to the birds ranging from the Rio Negro north through 
the provinces of Buenos Aires and Cordoba. However, six practically 
topotypical specimens of nigroguttata from Chimpay, Rio Negro 
Territory, about thirty miles west of Choele Choel, are much 
yellower than birds from the more northern localities mentioned. 
As a single specimen from Valcheta, a locality somewhat to the south 
of Choele Choel, is even yellower, it would appear that the true range 
of the form nigroguttata is from the Rio Negro (or perhaps the Rio 
Colorado) south. In this connection attention is called to the fact 
that Salvadori himself in the British Museum Catalogue lists ex- 
amples from Uruguay and Buenos Aires as typical of maculosa, al- 
though he must have had the opportunity of comparing these speci- 
mens with the types of nigroguttata. Two specimens in my collec- 
tion from Collon Cura, southern Neuquen, are somewhat darker 
than those from Chimpay and are probably intergrades with N. m. 
submontana. 

Specimens examined. 9: Argentina (Chimpay, Rio Negro, 6; 
Valcheta, Rio Negro, 1; Collon Cura, Neuquen, 2). 

Nothura maculosa submontana subsp. nov. Neuquen Spotted 
Tinamou. 

Nothura maculosa salvadorii (not of Hartert) Hellmayr and Conover, Field 
Mus. Nat. Hist., Zool. Ser., 13, (1), p. 97, 1942 part, northern Neuquen 
(Chos-Malal). 

Type. From Chos-Malal (37 20' S.-70 15' W.), Rio Neuquen, 
Neuquen, Argentina, elevation 2,500 feet; No. 4796, adult male in 
the Conover Collection, Chicago Natural History Museum; collected 
in May, 1926, by E. Budin. Original number 3138. 

Characters. The distinctive characters of this race are its light 
olive grayish appearance above, coupled with its pale whitish buff 
coloration below. It differs from nigroguttata, its nearest neighbor 
geographically, by having the general tone of the upper parts olive 
gray rather than yellow, with the light longitudinal streaks to the 
feathers much whiter, less yellowish; the under parts very much 
paler, whitish buff rather than ochraceous yellow; the dark markings 



352 FIELDIANA: ZOOLOGY, VOLUME 31 

of the chest much duller, less clearly defined, and the flanks less 
heavily barred. It is much lighter (olive gray rather than ochraceous 
brown) dorsally when compared to N. m. annectens from Buenos 
Aires Province and also much paler ventrally. From all the members 
of the darwinii complex, submontana can be easily told by its much 
longer tarsus and toes. 

Description of type. Top of head dark brown, each feather tipped 
with brownish buff; throat buffy white; sides of head, superciliary 
stripe and neck all around light dull buff with dark brown shaft 
streaks to each feather; feathers of mantle, back, scapulars and upper 
tail coverts dark brown, vermiculated with rufous brown and olive 
gray, with a broad edge of olive gray inside of which is generally a 
longitudinal streak of white that is often yellowish toward the base; 
tail feathers light buff, faintly barred with dusky; upper wing 
coverts barred with dark brown and ochraceous buff; primaries 
dark brown, the outer web broadly notched with buffy white and 
the inner web broadly barred with ochraceous buff; under wing 
coverts and axillaries rich ochraceous buff, the outermost coverts 
dotted with dusky; secondaries broadly barred with dark brown 
and ochraceous buff; chest whitish buff, each feather with a dark 
brown shaft streak that is spotted with rufous brown; flanks and 
thighs dirty light buff, the former broadly barred with dusky; 
abdomen, vent, and under tail coverts dirty light buff. Wing (flat) 
153, tarsus 41, middle toe (with claw) 35. 

Range. Northern Neuquen. Known only from the type. 

Remarks. It is with some hesitation that I describe this new 
form from a single specimen. However, several facts point to the 
probability that the individual at hand represents a new race and 
is not simply an aberrant example. In the first place the type is 
very different in coloration from any other specimen of the genus 
that I have examined, and the two specimens from Collon Cura, 
Neuquen, listed under the previous race, are somewhat intermediate 
between it and nigroguttata, as their under parts are lighter and the 
light longitudinal streaks on the feathers of the mantle, back, etc., 
are whiter than those of the Chimpay birds. Finally, the locality 
at which the type was taken is much farther to the west than any 
from which specimens of the maculosa complex have previously 
been known. In fact, except for Collon Cura, mentioned above, the 
nearest places at which examples of Nothura maculosa have been 
taken are some thr^e or four hundred miles to the east at Chimpay, 
Rio Negro Territory, or in the western part of Buenos Aires Province. 



CONOVER: SPOTTED TINAMOUS 353 

Indeed from such a western and Andean locality as Chos-Malal one 
would rather expect to find a form of darwinii. 

Specimens examined. 1: Argentina (Chos-Malal, Neuquen, 1). 

Nothura maculosa annectens subsp. nov. Pampa Spotted 
Tinamou. 

Nothura maculosa (not Tinamus maculosus Temminck) Salvadori, Cat. Bds. 
Brit. Mus., 27, p. 559, 1895 part, Buenos Aires. 

Nothura maculosa nigroguttata (not of Salvadori) Wetmore, Bull. U. S. Nat. 
Mus., 133, p. 33, 1926 Buenos Aires (Dolores to Lavalle; Carhu6; 
Guamini) (crit.); Peters, Bds. World, 1, p. 27, 1931 part, southern Santa 
Fe, Cordoba and Buenos Aires; Laubmann, Verh. Orn. Ges. Bay., 20, 
p. 280, 1934 part, southern Santa F6 (Las Rosas) to southern Buenos 
Aires Province; Steullet and Deautier, Obr. Cine. Mus. La Plata, 1, 
p. 147, 1935 part, Buenos Aires (Pigue, Christiano Muerto, La Plata, 
Bahia de San Bias) ; Hellmayr and Conover, Field Mus. Nat. Hist., Zool. 
Ser., 13, (1), p. 100, 1942 part, southern Santa Fe to southern Buenos 
Aires Province and eastern Cordoba. 

Nothura darwini salvadorii (not of Hartert) Laubmann, Wiss. Erg. Deuts. 
Gran Chaco Exp., Vogel, p. 74, 1930 part, Est. La Germania and Est. 
Wildermuth, near Las Rosas, Santa Fe. 

Type. From Cambaceres, Province of Buenos Aires, Argentina; 
No. 2556, adult female in the Conover Collection, Chicago Natural 
History Museum; collected on June 22, 1923, by Boardman Conover. 
Original number 697. 

Characters. Nearest to typical maculosa, from which it differs 
by having the upper parts less rufescent, more ochraceous brown; 
the dark markings on the chest and flanks darker (browner, less 
tinged with rusty), heavier, and more numerous; and the belly and 
abdomen lighter, more ochraceous, less rufescent. Size slightly 
larger (wing 132-146 mm.). 

Differs from nigroguttata by having the dorsal surface darker, 
browner and grayer, much less yellowish, and the under parts dull 
buff rather than bright ochraceous, with the dark markings much 
more numerous, especially on the chest. 

Description of type. Top of head dark brown, each feather tipped 
with rusty buff and edged with light buff; throat white; sides of head, 
superciliary stripe and neck all around light buff with dark brown 
shaft streaks to each feather; feathers of mantle, back, scapulars 
and upper tail coverts dark brown, vermiculated with rufous brown, 
and with broad edges of grayish buff inside of which are longitudinal 
streaks of white often tinged with dull buff; tail feathers light buff 



354 FIELDIANA: ZOOLOGY, VOLUME 31 

barred with dusky; upper wing coverts barred with dark brown and 
ochraceous buff; primaries dark brown, the outer web broadly 
notched with buffy white and the inner web broadly barred with 
ochraceous buff; under wing coverts and axillaries ochraceous buff, 
the very outermost coverts spotted with dusky; secondaries broadly 
barred with dark brown and ochraceous buff; chest dull buff, the 
feathers with longitudinal streaks and spots of dusky; flanks dull 
buff, heavily marked with broad bars of dark brown; abdomen, vent 
and under tail coverts dull ochraceous buff. Wing (flat) 141, tarsus 
41, middle toe with claw 29 mm. 

Range. Argentina, from central Cordoba (Noetinger) and south- 
ern Santa F^ (Las Rosas) south through Buenos Aires Province to 
Bahia Blanca. 1 

Remarks. The group of birds here named annectens has formerly 
been included in the race nigroguttata by all authors except Salvadori 
(Cat. Bds. Brit. Mus., 27, p. 560, 1895). However, as explained 
under that race, a series from near the type locality of nigroguttata 
proves to be quite different from birds taken in the Province of 
Buenos Aires. 

This is an intermediate form, showing great individual variation 
and possessing rather slight characters of its own. Yet it cannot 
be united with the races found to the north or south and inhabits 
too great an extent of country to be ignored or considered simply 
as a series of intergrades. 

Specimens examined. 49: Cordoba (Noetinger, 9); Santa F^ 
(Las Rosas, 5); Buenos Aires (Cambaceres, 3; Henderson, 1; Papin, 
near Bonifacio, 3; Las Ingleses, 1; Torrecito, 1; Alvarez Yonte, 2; 
General Lavalle, 3; San Vicente, 1; Buenos Aires, 2; Barracas al 
Sud, 2; Quilmes, 1; Conchitas, 1; Mar del Plata, 7; Dorrego, 5); 
(?)Mendoza, 2. 

Nothura maculosa maculosa Temminck. Spotted Tinamou. 

Tinamus maculosus Temminck, Hist. Nat. Pig. Gall., 3, pp. 557, 748, 1815 
based on "Ynambui" Azara, No. 327, Paraguay and two specimens (of 
unrecorded locality) in the Paris Museum (type locality restricted to 
Bernalcue, near Asuncion, Paraguay; cf. Laubmann, Verh. Orn. Ges. 
Bay., 20, p. 273, 1934). 

Cryptura fasciata Vieillot, Nouv. Diet. Hist. Nat., nouv. ed., 34, p. 109, 
1819 based on "Ynambui" Azara, No. 327; Paraguay. 

1 This form may range west through San Luis and La Pampa to Mendoza, 
though two specimens from the latter province, one in my collection and one 
(No. 14113) in the Munich Museum, while not typical, are closest to annectens. 



CONOVER: SPOTTED TINAMOUS 355 

Nothura maculosa savannarum Wetmore, Journ. Wash. Acad. Sci., 11, p. 435, 
Nov. 4, 1921 San Vicente, Rocha, Uruguay (type, in U. S. National 
Museum, examined); Peters, Bds. World, 1, p. 27, 1931 (range); Laub- 
mann, Verh. Orn. Ges. Bay., 20, p. 279, 1934. 

Nothura maculosa maculosa Peters, Bds. World, 1, p. 27, 1931 range, in part; 
Laubmann, Verh. Orn. Ges. Bay., 20, p. 279, 1934 part, except Minas 
Geraes; idem, Vogel Paraguay, p. 120, 1939 part, except Minas Geraes; 
Hellmayr and Conover, Field Mus. Nat. Hist., Zool. Ser., 13, (1), p. 102, 
1942 part, except the Paraguayan and Argentine Chaco and Formosa. 

Nothura maculosa nigroguttata (not of Salvadori) Laubmann, Verh. Orn. Ges. 
Bay., 20, p. 275, 1934 part, Santa Fe (Estancias Ines and La Geraldina, 
near Hersilia; Galvez). 

Nothura maculosa boliviano (not of Salvadori) Laubmann, Verh. Orn. Ges. 
Bay., 20, p. 280, 1934 part, northern and northeastern Paraguay; idem, 
Vogel Paraguay, p. 120, 1939 part, except Paraguayan Chaco. 

Range. Southern Brazil from southern Matto Grosso (Vaccaria), 
Sao Paulo and Rio de Janeiro (Campos) south through Paraguay 
and Uruguay to northeastern Argentina in the provinces of Misiones, 
Corrientes, Entre Rios and northern Santa Fe" (south to Galvez). 1 

Characters. Typical maculosa is characterized by its rufescent 
(mikado brown) and black appearance above and its ochraceous 
(sometimes rufescent) buff coloration below. The dark spotting 
on the chest and the barring of the flanks is moderately heavy. 
There is, however, great variation even in specimens from the same 
locality, but, in general, examples from the range given above have 
the rusty yellow coloration of this form. 

Specimens from Colonia Nueva Italia (near Villeta) and Villa 
Rica, Paraguay, are the most rufescent of any examined, and for 
purposes of comparison are taken as being typical of the race. 
Examples from Rosario are darker (blacker) on the dorsal surface, 
approaching in coloration the race from the west bank of the Rio 
Paraguay (that is, iV. m. paludivaga) in this respect, but they are 
much brighter ochraceous below than that form. Birds from 
Horqueta are very variable, some being typically dark rufescent 
and others considerably lighter, especially on the breast. Specimens 

1 Some form of maculosa probably inhabits the country from western Salta 
through Santiago del Estero. One specimen from Rosario de Lerma, Salta, belong- 
ing to the American Museum, has been examined. The example is in rather poor 
condition, but the upper parts are quite rusty, as in typical maculosa, while the 
under parts are light ochraceous with heavy dark spotting on the chest and lower 
neck. It may represent an unnamed form, as it is not at all like paludivaga, its 
nearest neighbor geographically. 

It may be well to call attention to the fact that Miller and Boyle, the collectors 
of the above specimen, also obtained three examples of Nothura darwinii salvadorii 
at this same locality. 



356 FIELDIANA: ZOOLOGY, VOLUME 31 

in the series from Capitan Bado, Cerro Amambay, are as rufescent 
as Villa Rica birds above, but on the average are lighter below and 
have the flanks and chest less heavily marked. 

Examples from Vaccaria, Matto Grosso, are typically rufescent 
above, but slightly lighter below. The birds from Sao Paulo on 
the average are not quite so typically rufous above, average lighter 
below and show a tendency toward N. m. major. Dorsally, speci- 
mens from Rio Grande do Sul are like typical Paraguayan examples, 
although birds from the southern part of the state average lighter 
than a series from the north. The under parts are slightly lighter 
buff but the spotting is as heavy as that of Villa Rica birds, and much 
heavier than that of those from Sao Paulo. 

Birds from northern and western Uruguay (9 specimens) are 
like typical maculosa in their rufescent coloration, although one or 
two show a tendency toward specimens from Buenos Aires Province. 

Four specimens (including the type of N. m. savannarum) from 
San Vicente, Rocha, Uruguay, have been examined. The type and 
one other are yellower, less rufescent, dorsally than typical Para- 
guayan examples and are lighter buff, less heavily marked with dusky 
below. A third has the typically rufescent coloration of maculosa 
while the fourth is intermediate between the two extremes. Despite 
the fact that two out of four birds from Rocha are very light-colored 
and a third has tendencies that way, savannarum would not seem 
to be a good race. It is probably only a local color phase. Birds 
from western and northern Uruguay are typical of maculosa, as are 
a series of six specimens from the Rio Grande do Sul seacoast (Santa 
Vitoria) near the Uruguayan border. 

In Argentina three specimens from Santa Ana, Misiones, are 
very rufescent and typical in every respect, and a specimen from 
Corrientes, although old and rather faded, appears to belong to this 
form. Five examples from Entre Rios (Conception del Uruguay, 4; 
Santa Elena, 1), while most like this race, show strong tendencies 
toward N. m. annectens from Buenos Aires Province. Six others, 
however, from La Soledad, Entre Rios, are very aberrant, being 
bright buff below with very sparse dark markings on chest and flanks 
and yellowish brown above except for one specimen, which is rather 
blackish. However, they have an appearance of being young and 
as they were collected during midsummer (January and February) 
this may account for their curious coloration. In Santa F, three 
specimens from Hersilia are very dark above and approach N. m. 
paludivaga in coloration, while one from Galvez is very rufescent, 



CONOVER: SPOTTED TINAMOUS 357 

like typical maculosa. Five others, however, from Las Rosas, only 
fifty kilometers farther south, can be referred only to N. m. an- 
nectens. 

Downies. Five specimens from Villa Rica, Paraguay, are much 
more rufescent about the head and neck than four specimens of 
chacoensis and one of boraquira. 

Specimens examined. 133: Paraguay (Colonia Nueva Italia, 
near Villeta, 5; Asuncion, 1; Bernalcue, near Asuncion, 1; Villa 
Rica, 12; Colonia Independencia, east of Villa Rica, 1; Rosario, 5; 
Horqueta, 12; Colonia Nuevo Germania, 1; San Luis de la Sierra, 
Apa Hills, 3; Sapucay, 1; Cambyreta, 1; Capitan Bado, Cerro 
Amambay, 12; La Fonciere, San Luis de la Sierra, 1; Caaguasu, 
eastern Paraguay, 1); Brazil (Vaccaria, Matto Grosso, 8; Aracassu, 
Sao Paulo, 5; Rio Tiete, Sao Paulo, 6; Ypiranga, Sao Paulo, 1; 
Itapetininga, Sao Paulo, 2; Victoria, Sao Paulo, 1; Campos, Rio de 
Janeiro, 1; Santa Vitoria, Rio Grande do Sul, 6; Tahym, Rio Grande 
do Sul, 2; Palmares, Rio Grande do Sul, 6; Vaccaria, Rio Grande 
do Sul, 2; Candiota, Rio Grande do Sul, 1; Conceicao do Arroyo, 
Rio Grande do Sul, 1; Passo Fundo, Rio Grande do Sul, 1; Rio 
Jaguarao, Rio Grande do Sul, 1); Uruguay (Mercedes, Soriano, 4; 
Caraguata, Tucarembo, 3; Fraile Muerto, Cerro Largo, 2; San 
Vicente, Rocha, 4); Argentina (Santa Ana, Misiones, 3; Corrientes, 
1; Conception del Uruguay, Entre Rios, 4; Santa Elena, Entre Rios, 
1; La Soledad, Entre Rios, 6; Hersilia, Santa F, 3; Galvez, Santa 
F<5, 1). 

Nothura maculosa paludivaga subsp. nov. Swamp Spotted 
Tinamou. 

Nothura maculosa boliviana (not of Salvadori) Wetmore, Bull. U. S. Nat. Mus., 
113, p. 36, 1926 Argentina (Las Palmas, Chaco; Kilometer 182 [Riacho 
Pilaga], Formosa) and Paraguay (Kilometer 80, west of Puerto Pinasco) ; 
Peters, Bds. World, 1, p. 26, 1931 part, Argentine Chaco and eastern 
Paraguayan Chaco; Laubmann, Verh. Orn. Ges. Bay., 20, p. 280, 1934 
part, Formosa, Argentina and eastern Paraguayan Chaco; Steullet and 
Deautier, Obr. Cine. Mus. La Plata, 1, p. 146, 1935 Chaco and Formosa, 
Argentina; Laubmann, Vogel Paraguay, pp. 120, 121, 1939 part, Formosa, 
Argentina and eastern Paraguayan Chaco (145 km. west of Puerto Casado). 

Nothura darwini salvadorii (not of Hartert) Laubmann, Wiss. Erg. Deuts. Gran 
Chaco Exp., Vogel, p. 74, 1930 part, Formosa (San Jos6; Mision Tacaagte; 
Lapango). 

Nothura maculosa chacoensis Conover, Proc. Biol. Soc. Wash., 50, p. 227, 
1937 part, Paraguay (Puerto Pinasco), Argentina (Formosa and the 
Chaco). 



358 FIELDIANA: ZOOLOGY, VOLUME 31 

Nothura maculosa maculosa (not of Temminck) Hellmayr and Conover, Field 
Mus. Nat. Hist., Zool. Ser., 13, (1), p. 102, 1942 part, Paraguayan 
and Argentine Chaco and Formosa. 

Type. From seven kilometers southwest of Laguna General 
Diaz, Paraguayan Chaco (22 15' S.-59 5' W.); No. 17653, adult 
male in the Conover Collection, Chicago Natural History Museum; 
collected July 24, 1945, by Jacob Unger. Original number 119. 

Characters. The distinctive feature of this new race is its dark, 
rather grayish black and dull yellow appearance dorsally. It is 
nearest to typical maculosa and like that form has the upper parts 
and chest rather heavily blotched with dark brown, but it is much 
less rufescent brown above and duller (grayer) much less yellowish 
ochraceous below. It is darker than N. chacoensis (whose range it 
may slightly overlap), the dorsal surface being much more heavily 
blotched, the chest more heavily streaked and the flanks more 
heavily barred with dark brown. The under parts are duller, paler, 
and less ochraceous; in many specimens the center of the abdomen 
is whitish. Its general appearance is grayish black, in contrast to 
the ochraceous buff of chacoensis. 

Description of type. Top of head dark brown, each feather tipped 
with brownish buff; throat white; sides of head, superciliary stripe 
and neck all around light yellowish buff with broad dark brown shaft 
streaks to each feather; feathers of mantle, back, scapulars and upper 
tail coverts blotched with dark brown and unevenly vermiculated 
with grayish buff, with a broad edge of smoky grayish buff, inside 
of which is generally a longitudinal streak of buff y white ; tail feathers 
hair-like, and light smoky buff broadly barred with dusky; upper 
wing coverts barred with dark brown and ochraceous buff; primaries 
dark brown, the outer web broadly notched with buffy white and 
the inner web broadly barred with ochraceous buff; secondaries 
broadly barred with dark brown and ochraceous buff; under wing 
coverts and axillaries whitish buff; chest smoky ochraceous buff, 
each feather with a broad dark brown shaft streak on basal two thirds 
and a dark brown spot near tip; flanks and thighs smoky ochraceous 
buff, the former broadly barred with dark brown; abdomen smoky 
whitish buff (many individuals are smoky buff without any whitish 
tinge). Wing (flat) 130, culmen (exposed) 20, tarsus 33, middle toe 
(with claw) 28. 

Range. The marshy country along the west bank of the Para- 
guay River from Puerto Casado (and probably the Bolivian border) 
south at least to the Argentine Chaco (Las Palmas; General Pinedo). 



CONOVER: SPOTTED TINAMOUS 359 

Also west up the valley of the Pilcomayo for 235 kilometers and 
probably farther. 

In the northern Paraguayan Chaco, west of Puerto Casado, the 
range of this tinamou does not seem to extend west from the River 
Paraguay much over 150 kilometers, where it meets and may overlap 
the range of N. chacoensis. At this point, according to the collectors, 
the climate, soil and vegetation change rather abruptly, although 
there is no appreciable change in elevation. "The part near the 
river," they write, "is swampy and covered with a sweet grass, 
whereas the more western portion is dry, and the soil alkaline and 
covered with what they call a salt grass. Other types of vegetation 
change also." 

Specimens examined. 29: Paraguayan Chaco (145 km. west of 
Puerto Casado, 1; 30 km. northeast of V. Militar [Islapoi], 1; 80 km. 
west of Puerto Pinasco, 1; Laguna General Diaz, 6; 17 km. west of 
Laguna General Diaz, 5; 7 km. southwest of Laguna General Diaz, 2; 
Fort Wheeler, 1; Makthlawarija, Mission Vieja, 3; 235 km. west 
of Riacho Negro, 2; Monte Alto, 1); Argentina (10 miles northwest 
of Riacho Pilago, Kilometer 182, Formosa, 2; San Jos, Formosa, 1; 
Las Palmas, Chaco, 1; General Pinedo, Chaco, 2). 

Nothura maculosa major Spix. Spix's Spotted Tinamou. 

Tinamus major Spix, Av. Spec. Nov. Bras., 2, p. 64, pi. 80, 1825 "in campis 
Minas Geraes, prope pagum Tejuco [=Diamantina] et Contendas" 
Brazil (type, in Munich Museum, examined). 

Tinamus medius Spix, Av. Spec. Nov. Bras., 2, p. 65, pi. 81, 1825 "in campis 
prope pagum Tejuco" [=Diamantina], Minas Geraes, Brazil (type, in 
Munich Museum, examined). 

Nothura maculosa maculosa (not of Temminck) Laubmann, Verh. Orn. Ges. 
Bay., 20, pp. 279, 281, 1934 part, Minas Geraes (Tejuco, Agua Suja, 
Pocos de Caldas); Pinto, Rev. Mus. Paul., 20, p. 30, 1936 Fazenda 
B6a Vista, Jaragua, Goyaz. 

Nothura maculosa major Hellmayr and Conover, Field Mus. Nat. Hist., Zool. 
Ser., 13, (1), p. 104, 1942 (range; bibliog.). 

Range. Interior of Brazil in the states of Minas Geraes (Dia- 
mantina, Agua Suja, Pocos de Caldas), Goyaz (Jaragua, Veadeiros) 
and probably southern Bahia. 

Characters. This is a very poor subspecies. When compared 
with the typical race it has the dorsal surface slightly lighter and 
yellower; under parts paler; dusky markings on foreneck more in- 
distinct and flanks less heavily barred. Even these differences are 



360 FIELDIANA: ZOOLOGY, VOLUME 31 

apparent only when series are compared. Wing 124-140, tarsus 
36-39, middle toe (with claw) 30-33. 

While individual specimens of this form may not appear to be 
very different when compared to specimens of maculosa from Sao 
Paulo, it is a fact that birds of this complex average lighter in colora- 
tion as one goes north from Paraguay. Therefore the name major 
may be useful in designating these paler northern examples from 
Minas Geraes, Goyaz, and southern Bahia. 

Remarks. The types of Tinamus major and T. medius Spix 
look somewhat aberrant when compared to the other seven speci- 
mens in the series examined and they are entirely unlike each other, 
although they come from the same locality. The type of major 
has the dorsal surface blotched, with very few ochraceous vermicula- 
tions, while the type of medius has these parts strongly vermiculated. 
For some reason this latter specimen has its label marked "juv." 
and the description of medius has been quoted in synonymy as being 
that of an immature specimen of major. However, to my mind there 
is nothing about this example that speaks of immaturity. The 
feathers of its dorsal surface are very badly worn, and its wing, 
although the primaries are much frayed at the tip, measures 140 mm., 
the longest in the series examined. 

Specimens examined. 10: Brazil (Agua Suja, near Bagagem, 
Minas Geraes, 4; Pocos de Caldas, Minas Geraes, 1; Diamantina, 
Minas Geraes, 2; Pirapora, Minas Geraes, 1; Veadeiros, near Caval- 
canti, Goyaz, 2). 

Nothura maculosa cearensis Naumburg. Ceara Spotted Tinamou. 

Nothura maculosa cearensis Naumburg, Amer. Mus. Nov., 554, p. 1, Aug. 22, 
1932 Lavras, Ceara, Brazil (type, in the American Museum of Natural 
History, examined); Hellmayr and Conover, Field Mus. Nat. Hist., Zool. 
Ser., 13, (1), p. 105, 1942 (disc). 

Range. Known only from the type specimen taken at Lavras, 
Ceard, Brazil. 

Characters. Size very small. Upper parts ochraceous, with 
dark brown and rufous brown blotches. Neck and upper chest 
with narrow longitudinal dark streaks. Rest of under parts light 
buff. Sides of chest lightly barred, flanks practically immaculate. 
Wing 110, tarsus 29, middle toe (with claw) 27. 

In size closest to N. minor Spix but inner webs of outer primaries 
broadly notched with buff, and upper wing coverts much more 
coarsely barred. Upper parts more ochraceous and rufous brown 



CONOVER: SPOTTED TINAMOUS 361 

blotches much less rufescent. Lower breast and abdomen brighter, 
cream-colored rather than dull buff. 

In some respects such as size and general appearance the type is 
very like some specimens of N. darwinii boliviano,, but it can be 
distinguished immediately by the barred inner webs of the outer 
primaries and the much less hair-like texture of the flank feathers. 
In general the dorsal surface is much less rufescent. 

From N. maculosa major, its nearest relative geographically, 
it can be told by its diminutive size, lighter, much more buffy ochra- 
ceous upper parts, and lighter, creamy buff breast and belly. 

Specimens examined. 1: Brazil (Lavras, Ceara, 1). 

Nothura chacoensis Conover. Chaco Spotted Tinamou. 

Nothura maculosa chacoensis Conover, Proc. Biol. Soc. Wash., 50, p. 227, 
Dec. 28, 1937 265 km. west of Puerto Casado, Paraguayan Chaco (type 
in the Conover Collection, Chicago Natural History Museum) (exclusive 
of Puerto Pinasco and Argentine specimens); Hellmayr and Conover, 
Field Mus. Nat. Hist., Zool. Ser., 13, (1), p. 98, 1942 (range). 

Nothura maculosa boliviano (not N. boliviano Salvadori) Peters, Bds. World, 
1, p. 26, 1931 part, western Paraguayan Chaco. 

Range. Arid part of the Paraguayan Chaco, which begins about 
150 kilometers west of the Rio Paraguay. Known only from west 
of Puerto Casado and Puerto Pinasco but probably ranges west 
and north to at least the Bolivian border and south to the more 
humid strip of country along the north bank of the Rio Pilcomayo. 

Characters. Differs from both N. m. maculosa and N. m. 
paludivaga by its much lighter appearance. Where the general 
type of coloration of the upper parts is dusky rufescent in typical 
maculosa and grayish black in paludivaga, in chacoensis it is ochra- 
ceous buff. On the under side the dark markings of the chest are 
not nearly as bold and prominent, being duller and generally taking 
the form of narrow shaft streaks, and the lower breast and belly 
are much lighter and brighter (purer buff). 

Downies. Four specimens from 195 kilometers west of Puerto 
Casado, Paraguayan Chaco, are much less rusty about the head and 
neck and paler on the dorsal surface than a series of the young of 
typical maculosa. 

Remarks. The exact relationship of chacoensis is very problem- 
atical. While in general appearance, size and proportion of tarsus 
and toes it seems to belong to the maculosa complex, the fact must 
be taken into consideration that its range abruptly meets and may 



362 FIELDIANA: ZOOLOGY, VOLUME 31 

slightly overlap that of N. m. paludivaga and that there appears to 
be no area of intergradation between the two. However, in my 
collection there are two specimens of paludivaga that could be hy- 
brids. They are much grayer, less blackish, dorsally than usual, 
because their upper parts are very finely vermiculated (not blotched), 
which is a characteristic of many specimens of chacoensis. The dark 
markings of their chests are also less prominent than is normal for 
the race. 

Specimens examined. 29: Paraguayan Chaco (265 km. west of 
Puerto Casado, 14; 195 km. west of Puerto Casado, 5; 170 km. west 
of Puerto Casado, 1; 120 km. west of Puerto Pinasco, 4; 1 Orloff, 2; 
16 km. east of Philadelphia, 3). 

1 These four specimens are typical of Nothura chacoensis and may have been 
taken farther west of Puerto Pinasco than indicated. If not, they were collected 
fifty kilometers east of the western boundary of the range of Nothura maculosa 
paludivaga, as shown by actual specimens examined (Fort Wheeler; 30 km. north- 
east of V. Militar [=Islapoi]).