NOTES ON TWO NEOTROPICAL , CAPRIMULGUS ANTHONYI AND C. PARVULUS

PAUL SCHWARTZ Ministerio de Agricultura y Cria Estaci6n Biol6gica Henry Pittier ( Ranch0 Grande ) Apartado 184, Maracay Aragua, Venezuela

Recently while in the American Museum of viewing the species and genera of de- Natural History in New York engaged in vari- scribed as new between 1920 and 1934, ous matters pertaining to the operations of the “reclassified” Chapmans’ as “Caprimulgus Estaci6n Biol6gica Henry Pittier (Ranch0 parvus anth.” without giving any reason what- Grande ), Venezuela, I had occasion to com- ever. The misspelling of the specific name was pare some specimens of the Little , doubtless a typographical error. Apparently Caprimulgus parvulus, from Venezuela with neither Meise nor Peters ever examined any those in the American Museums’ collection. specimens of anthonyi. (No illustration of the As a matter of routine checking I also exam- bird has been published hitherto.) Probably ined the type of the west Ecuadorean form they were influenced by the fact that Chap- listed by Peters (1940:202) as Caprimulgus man (1923:4-5) in his original description par&us anthonyi and was surprised to find compared the new bird with C. parvulus and that this form, although of similar size and later ( 1926 :278 ) stated : “This well-marked proportions, is otherwise strikingly different species, known only from the type, is doubt- from C. parvulus. In this paper I present rea- less a representative of parvulus” sons for considering it a separate species. ( emphasis mine ) . Peters (1940:202) recognized two other Meyer de Schauensee ( 1964: 126, 1966: 151) races of C. par&us, including the nominate evidently followed Peters, for he does not form, par&us Gould, described from Santa mention anthonyi but includes its range in that F6, Rio Paran6, , ranging over most of the species C. parvulus. of south of the Amazon to Ar- Figure 1 shows the most striking differences gentina and and west to eastern among adult males of C. anthonyi and of the PerG, and heterurus (Todd), attributed to the two races of C. parvulus. As the photograph the Santa Marta region of Colombia but known can not show the major differences fully, I de- to occur also in Venezuela. So little has been scribe them as follows. published about the latter form that I include Tail. In anthonyi the two outer pairs of rectrices have here additional data on its morphology and the entire inner web white for its full length, except ecology, and also extend considerably its along the shaft on the subapical half. (Chapman known distribution in Venezuela. stated only that the “outer pair” is so marked; either he overlooked the next outer pair or else he used the word “pair” in a sense different from that usually CAPRZMULGUS ANTHONYZ (CHAPMAN) understood.) The inner three pairs of rectrices have no white areas. There is no barring on the two outer Chapman (1923:4-5) described anthonyi as a pairs, and relatively little, and that only marginally distinct species based on a single adult male on the inner web on the next two pairs of rectrices; from southwestern Ecuador (Portovelo, in the the middle pair is unbarred. upper Tumbez drainage at 750 m). There is By contrast, in C. parvulus there is a roundish white “spot” (smaller in nominate pamdus, larger in heter- another specimen sexed as a female bearing urns) on the distal end of the four outer pairs of identification as this form which probably is rectrices. Also the dark areas are barred, conspicu- correctly allocated. It too is from western ously on the outer rectrices, less conspicuously on the Ecuador but at the northern end (Vaqueria, inner. Furthermore, the inner web of the outer rec- trices in C. anthonyi is more tapered apically and the at the mouth of the Rio Santiago, AMNH no. tip of the tail is thus somewhat square, slightly 477258 [ex Rothschild Coll.], collected 3, Sep- notched, rather than rounded as in C. paruulus. tember 1901, R. Miketta). These seem to be Wing band. In C. anthonyi the five outer prima- ries are totally crossed by a broad white band. In C. the only two known specimens, but it is hoped parvulus the wing band is limited to the four outer that this paper may serve to uncover others. primaries and usually does not cross completely the Why Peters lumped anthonyi with C. par- first, sometimes not the fourth; very rarely there may be a slight indication of it on the fifth primary VU~S is not clear. Perhaps he was following (counting from the outside). Meise ( 1938: 144)) who, in a long paper re- Furthermore, this band in C. anthonyi is located

r2231 The Condor, 70:223-227, 1968 224 PAUL SCHWARTZ

FIGURE 1, left to right. Caprimu& ~U~W~U.Spar~lus, Corrientes, Argentina, Partridge no. 13640 (AMNH, uncatalogued); C. par~lus heterurus, Rio Aricuaisa, Zulia, Venezuela, EBHP no. 4709; C. anthonyi, Portovelo, Prov. de1 Oro, Ecuador, AMNH no. 166785, Type; C. coyenne&.s insu~ris, Curacao, AMNH no. 477208. (The outermost tail feather of the birds’ right side is missing, showing the pattern of the next outer, which has some resemblance to the outermost in C. anthonyi.) All specimens are adult males.

medially slightly more than 50 per cent from the tip having the back barred and a broad band of the folded wing, whereas in C. porwlus it is closer crossing the five outer primaries, midway up to the wing tip (less than 40 per cent from the tip). Throat patch. One of the conspicuous features of the folded wing, although this band is buffy the adult male C. purouZu.s is the proportionately large rufous rather than white. The outer prima- white throat patch composed of dense, silky textured ries are otherwise immaculate brownish-black feathers. Meyer de Schauensees’ ( 1964: 126) failure as in the male. The females of C. pamulus to mention the white throat patch is obviously an have the primaries broadly mottled or brokenly oversight. In the only adult male C. anthonyi available for barred (usually mostly basally) with buffy comparison, the white throat patch is less extensive, rufous and lack a wing band. not so profusely feathered, and the feathersmuch less The tail of this bird differs from that of fe- dense. male C. parvulus by having a large, dull buffy Back. In C. onthonyi the back, especially the inter- scauular area and the rump, is finely barred with black patch extending about 20 mm up the inner and ochraceous, with little or no suggestion of streak- web of the outer pair of rectrices. This inner ing. In part&us these areas are speckled and con- web also has a dull buffy edge about 1.5 mm spicuously streaked; only occasionally is there a sug- wide running from the tip almost halfway up gestion of barring. the feather. Otherwise the rectrices are mot- The other specimen thought to be C. anthonyi, tled and barred as in C. parvulus. The differ- sexed as female, is similar to the male type in ences between this specimen and the type of NOTES ON NEOTROPICAL NIGHTJARS 225

TABLE 1. Measurements(range and mean) of specimensof Caprimulgusspp.

Wing R&O TP.XOP (chord ) , mm Tail, mm wing/tail

C. anthonyi 135 98 1.38 “0 (PT) 132 95 1.39 C. p. parvulus LT (20) 134-143 ( 138.5) 94-105 (97.5) 1.33-1.49 (1.42) 0 (13) 128-141 (135.8) 93-105 (98.1) 1.32-1.44 (1.38) c. p. heterurus $ (lo) 135-146 (140.8) 89-98 ( 93.2 ) 1.46-1.56 (1.51) 0 (3) 134-137 (136) 87-93 (90) 1.48-1.54 t 1.51) subadult (3) 134-137 (136) 87-93 (89) 1.48-1.58 i 1.53j c. c. cayeWW?Wk$ (12) 133-148 (140) 111-126 (117) 1.16-1.24 (1.2) 0 (11) 132-144 (139) 94-108 (102) 1.28-1.47 (1.36) * Number of specimens in parentheses.

C. anthonyi tend to follow the pattern of sex- birds is still too incomplete to permit sweeping ual differences usual in the genus. However, generalizations, but from many field observa- the specimen shows signs of immaturity, and tions of a number of species I have come to the consideration should be given to the possibility conclusion that in the activity of the male birds that it is a young male, improperly sexed. of this group there are two outstandingly In these respects the male C. anthonyi re- important factors serving signal functions for sembles adult males of the widely distributed territory and mating. One is the primary song C. cayennensis: white running the full length (as distinguished from other vocal or mechan- of the rectrices (although in C. cayennensis ical sounds); the other is the pattern and loca- this is on the four outer pairs and the outer- tion of the white areas. The white areas in- most pair is also white on the outer web with clude “steady flags” and “flash patches.” a diagonal dark bar toward the base), square- White throat areas are primarily “steady flags”; tipped tail (in cayenneds it is even more broad, white wing bands may work either square), type and position of wing band (but way; white tail areas are primarily “flash in cayenne&s it crosses only the four outer patches” and as such must show a distinctive primaries), and fine barring of the back. HOW- pattern if they are to serve a function in spe- ever, there is a very great difference in general cies recognition. That they do is suggested by color pattern and tail length, as can be seen the variety of such patterns within the capri- from figure 1 and table 1. mulgids. Even some tail patterns that may An occasional subadult male C. cuyennensis appear quite similar in inert museum speci- does have white restricted to the inner web of mens will be found to appear quite different the outer rectrices, but such individuals can as flash patterns in life. easily be distinguished from C. unthonyi by There is no doubt that the more important having the outer web barred, by the longer of the two factors is the voice. Unfortunately, tail, and by the absence or reduction of the we know nothing of the voice of C. anthonyi, wing band (outer primaries barred). so that factor must be omitted from present In ecological preferences too it appears that considerations. (For those who may be inter- C. unthonyi may differ from C. purvulus. Of ested, the songs of both C. purvulus heterurus the type of C. unthonyi Chapman (1923:5) and C. c. cuyennetis have been published wrote: “the specimen . . . was shot . . . on a in a recording: “Bird Songs from the Tropics,” trail running through open, grassy, arid coun- Naturaleza Venezolana No. 1, Instituto Neo- try. * . *” According to my experience with tropical. ) C. parvulus heterurus throughout Venezuela, However, as pointed out previously, the it is not a bird of open, grassy country but white areas of C. anthonyi are quite different rather inhabits the more sparse parts of scrubby from those of C. purvulus, especially in the or deciduous woodlands, together with the tail pattern. Actually in this character there adjacent or included clearings, either natural is a radical morphological difference of a type or man-made. It is interesting to note that in basic to this group. Also, in this and other its apparent ecological preference, C. anthonyi features C. unthonyi differs from any other again seems to resemble C. cuyennensis, a bird known form of nightjar. Therefore, in spite of of completely open areas. However, I d6 not the inability to make comparisons of voice at suggest by this or previous statements that I present, it seems best that we consider Cupri- consider unthonyi more closely related to C. mulgus unthonyi a valid, distinct species and cayennensis than to C. purvulus. not a race of C. parvulus. After examination Our knowledge of the nightjars as living of the specimens in question, Eugene Eisen- 226 PAUL SCHWARTZ mann has indicated his agreement with this surements, although the combination of slightly opinion, longer wing and somewhat shorter tail does result in wing/tail ratios that are significantly C. PARVULUS HETERURUS (TODD) larger in heterumcs (see table 1). This form was described as differing from The published distribution of heterurus in- nominate pur~ulus chiefly in the greater ex- dicates its presence in the north-central region tent of white on the tail, based on one adult of Venezuela (Distrito Federal, Aragua, and male specimen from the Santa Marta region Miranda) as well as in the Santa Marta and of Colombia. Except for distributional data, Cucuta regions of Colombia (Phelps and the only other information about this race that Phelps 1958:171; Meyer de Schauensee 1964: I have found in the literature refers to this 126, 1966: 151). In the last-mentioned citation, same specimen ( Todd and Carriker 1922: 220)) where only species are treated, heterurus is so it has hardly been possible to judge fairly not definitely specified as the form inhabiting the differences between heterurus and the the CGcuta region of Colombia, but I have nominate form. examined specimens from Villa Felisa, Norte I have examined a number of specimens of de Santander, Colombia, in the United States heterurus, among them 10 adult males includ- National Museum, and find that they corre- ing the type, as well as no less than 40 nomi- spond to this race. nate putd~s. There is much individual vari- I now add the following new localities ation in size in this species, as is true with which extend the range of heterurw both west many caprimulgids. There is less variation in and southeast in Venezuela, based on speci- color than in some other species, but heterurus mens in the collection of the Estacion Biolo- does display a considerable range between the gica Henry Pittier (EBHP) : Rio Aricuaisa, darkest and lightest individuals, even within Zulia (la ); Calle Larga, Zulia (28 3, 19 ); a given population. Cumboto, Aragua ( 10 ); Santa Maria (Upata), The one character that serves unquestion- Bolivar (28 3 ). ably to separate the two races is the much In addition there is an adult male from San greater extent of the white tipping of the rec- Pedro, Caura, Bolivar, Venezuela, in the trices in heterurus. This form has the outer- AMNH. In the USNM an immature female most pair completely (or almost so) white from Valle, Merida, Venezuela, bears an iden- tipped; the second and third pairs from the tification as heterurus. outside usually have at least some white on From personal experience, I know C. pur- the outer web, in addition to that normally vulus to be more widely distributed in Vene- expected on the inner web; the fourth outer zuela than the few specimen localities indi- pair has white only on the inner web. In the cate. I have not so far encountered it in the nominate form the white is usually restricted vast lluno country, and the indications are that to the inner web throughout, although it is not it prefers at least slightly hilly terrain, shying too unusual to find birds that also have some away from extensive flatlands. I do not doubt white on the outer web of the outermost rec- that in the proper habitat it may be found trices. The middle pair has no white in either throughout the entire northern part of Vene- form, Furthermore, in heterurus the white ex- zuela. Also it is probably more widely distrib- tends 14-20 mm along the shaft of the outer uted in Colombia than presently thought, and two pairs of rectrices, whereas in nominate very likely is to be found throughout northern parvulus it is I2 mm or less, so that the total South America east of the Andes in ecologi- area of white is conspicuously greater in heter- cally suitable areas, connecting with the known urus (see fig. 1). range south of the Amazon River. However, There are other slight differences between there is a tremendous gap with no available the two races, but they are so masked by over- specimens between about 7.5” N latitude and lapping variation that they have little useful- the Solimoes-Amazon River; neither are there ness for identification. specimens recorded from the Guianas. Some individuals of heterurus do have larger An adult male from Amorin, Igarape, lower white wing bands, and there is a tendency in Rio Tapajoz, (AMNH no. 288296), this race for some individuals to have the band seems to have a bit more white on the tail extend to the outer web of the outer primary than normal nominate parvulus and may rep- (sometimes only on one wing, as in fig. 1). resent a tendency toward heterurus. If so, it is Rarely it may be partially present on the fifth the only indication we have that these two outer primary. forms may actually intergrade, although with The slight differences in wing and tail the information available and considering their lengths are hardly significant as absolute mea- extreme similarity morphologically, there is NOTES ON NEOTROPICAL NIGHTJARS 227 certainly no reason at this time to question tional specimen localities are cited, extending their conspecific relationship. considerably its range in Venezuela. SUMMARY ACKNOWLEDGMENTS On the strength of basic morphological differ- I express sincere appreciation to Dean Ama- ences, the Ecuadorian Nightjar, Caprimulgus don for permission to use the collection and anthonyi (Chapman), seems to be a valid spe- facilities of the American Museum of Natural cies and not a race of C. parvulus as currently History and for supplying the photograph for treated by Peters and others. It is here illus- figure 1. I also thank the authorities of the trated for the first time, in comparison with its Carnegie Museum, Coleccion Ornitologica allies. Phelps, and the United States National Mu- With more material than has been previ- seum for permission to review the material of ously available, C. purvulus heterurus (Todd) those collections. Eugene Eisenmann has been is compared with the nominate form and very helpful, as always, and I am further in- found to differ from it principally in the debted to him for the critical reading of the amount of white tipping of the rectrices. Addi- manuscript.

LITERATURE CITED

CHAPMAN, F. M. 1923. New South American birds. Amer. Mus. Nov. no. 67. CHAPMAN, F. M. 1926. The distribution of bird-life in Ecuador. Bull. Amer. Mus. Nat. Hist. 50. MEISE, W. 1938. Fortschritte der ornithologische Systematik seit 1920. Proc. VIII Intern. Omithol. Congr., Oxford Univ. Press. MEYER DE SCHAUENSEE, R. 1964. The birds of Colombia. Livingston Publishing Co., Narberth, Penn. MEYER DE SCHAUENSEE, R. 1966. Birds of South America. Livingston Publishing Co., Narberth, Penn. PETERS, J. L. 1940. Check-list of the birds of the world, vol. 4. Harvard Univ. Press, Cambridge, Mass. PHELPS, W. H., and W. H. PHELPS, JR. 1958. Lista de las aves de Venezuela con su distribution. Tomo 2, Parte 1, Passerifonnes. Bol. Sot. Ven. de Ciencias Nat., 19, no. 96. TODD, W. E. C., and M. A. CARRIKER, JR. 1922. The birds of the Santa Marta region of Colombia. Ann. Carn. Mus. 14. Accepted for publication 30 June 1967.