Beginning an End to 63 Years of Uncertainty: The Neotropical Parakeets Known as Pyrrhura picta and...

Academy of Natural Sciences

Beginning an End to 63 Years of Uncertainty: The Neotropical Parakeets Known as Pyrrhurapicta and P. leucotis Comprise More than Two SpeciesAuthor(s): Leo JosephSource: Proceedings of the Academy of Natural Sciences of Philadelphia, Vol. 150 (Apr. 14,2000), pp. 279-292Published by: Academy of Natural SciencesStable URL: http://www.jstor.org/stable/4065072 .

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PROCEEDINGS OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA 150: 279-292. 14 APRIL 2000

Beginning an end to 63 years of uncertainty: The Neotropical parakeets known as Pyrrhura picta and P. leucotis comprise more than two species

LEO JOSEPH Department of Ornithology, Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway, Philadelphia PA 19103-1195, U.S.A.

ABSTRACT - Patterns of geographical variation in plumage and morphometrics of the 13 taxa of Neotropical parakeets currently classified as subspecies of either Pyrrhura picta or P. leucotis were assessed from 257 specimens. The 13 taxa and the species with which they are usually aligned are: "picta" group-picta, amazonum, lucianii, roseifrons, caeruleiceps, subandina, pantchenkoi, eisenmanni; and "leucotis" group-leucotis, pfrimeri, griseipectus, emma and auricularis. Characters varying in clear, if irregular, geographical patterns were colors and patterns of the pileum, lores, feathered ocular ring, ear coverts and cheeks, pattern of feather edging on the underparts, and lengths of maxilla and wing. The main findings are (a) the prevalent taxonomy treating all taxa as subspecies of either picta or leucotis should be discarded because it unduly emphasizes the concordance between patterning on the underparts and cheek color at the expense of variation in other informative characters such as the color and pattern of the pileum, lores, and feathered ocular ring; (b) subandina and pfrimeri are so distinct that phenotypically they stand apart as much from each other as they do from other taxa in the group; (c) treatment of two near-identical taxa with the "leucotis" pattern of underparts and cheeks, eisenmanni from Panama and caeruleiceps from northwestern Colombia, as subspecies of picta has confused understanding of variation, taxonomy and biogeography; (d) auricularis and pantchenkoi are not diagnosable and should be synonymized with, respectively, emma and caeruleiceps; and (e) closer study is needed of the variable western Amazonian populations here tentatively maintained as lucianii and roseifrons to determine how many taxa are involved and the nature of any intergradation between them. Overall, the biological inappropriateness of the prevalent two- species arrangement has been highlighted as has the need for phylogenetic analyses to determine relationships among the taxa studied as well as their relationships to the rest of Pyrrhura. An interim systematic arrangement is suggested based on the phylogenetic species concept. Variation in Amazonian populations will be examined more closely in a separate paper.

INTRODUCTION

Recent texts (Juniper and Parr, 1998; Collar, 1997; Forshaw and Cooper, 1989; Ridgely and Gwynne, 1989) have stressed the unsatisfactory understanding of geographical variation and taxonomy in two supposed- ly closely related Neotropical parakeets, the Painted Parakeet Pyrrhurapicta, widespread in humid lowland South America, and the Maroon-faced or White-eared Parakeet P. leucotis, which occurs in isolated populations around the northern and eastern fringes of South America. The two-species arrangement traces to Peters (1937) who without comment reduced the number of species in the complex to two from the six to eight (picta, leucotis, lucianii, roseifrons, emma, pfrimeri, subandina and griseipectus) that had been variously recognized (see Todd, 1917; Cory, 1918; Miranda- Ribeiro, 1920).

Given that some taxa in the picta-leucotis complex were described as recently as 1977 (pantchenkoi) and 1985 (eisenmanni), it is no surprise that study of geographical variation in this group never has been based on museum material from throughout its

geographical range and including all 13 taxa. The present contribution's main aim, then, is to begin to rectify this situation and provide a foundation for later systematic study of the group. Patterns of variation in individual characters will be described with the aims of (a) locating zones where unique combinations of character states occur, and (b) determining whether gradients of change among characters are concordant geographically. Appendix 1 gives a brief nomenclatural history of the group and notes which taxa have been associated with picta and leucotis. To facilitate reading of the text, Fig. 1 shows the distribution of all 13 taxa currently or recently treated as subspecies of either picta or leucotis. This paper will give an overview of patterns of geographical variation in the whole complex. A second paper will examine Amazonian populations in further detail.

For ease of discussion, most currently recognized names (Appendices, Table 1, Fig. 1; Juniper and Parr, 1998; Collar, 1997; Forshaw and Cooper, 1989) can be used without circularity because of the geographical isolation of the taxa. There is, however, some risk of circularity in using names when trying to establish pat-

279



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terns of geographical variation for the populations of lowland Amazonia and the Guianas because of uncer- tainties concerning range limits (see Fig. 1; Appendi- ces). For simplicity, picta will hereafter apply to populations from the Guianas, Amapa (Brazil) and southern Venezuela, and amazonum will apply to populations of southeastern Amazonia east of and including the Madeira and along and south of the north bank of the Amazon (type locality Obidos). How to group and code western Amazonian populations for analyses posed a challenge from the outset. As this work progressed, lucianii came to be applied provisionally to brown-headed populations along and mostly north of the Amazonas-Solim6es-Maranion rivers west of Manaus and into Peru (i.e., north of roughly 5?00' S), and roseifrons was applied to the generally red-crowned populations from lower southwestern Amazonia into far southwestern Brazil and Bolivia roughly south of 5?00' S. "Western Amaz- onia" will refer to the combined ranges of lucianii and roseifrons as just defined.

MATERIALS AND METHODS

All specimens of the 13 taxa (total - 257; Table 1; Appendix 2) held in the collections of the Academy of Natural Sciences, Philadelphia (ANSP), the American

Museum of Natural History, New York (AMNH), and the United States National Museum, Washington (USNM) were examined. A further three specimens from the Carnegie Museum, Pittsburgh (CM) of caeruleiceps (CM 54727) and subandina (CM 52828, 52829), and 22 western Amazonian specimens from the Field Museum of Natural History, Chicago (FMNH) were examined on loan. As the study progressed, it became necessary to borrow still more specimens from other institutions so that problems that arose in the present study could be examined more closely. The results of that work will be reported elsewhere.

Characters were selected after an initial examination of 85 specimens of 11 of the 13 taxa (pantchenkoi and caeruleiceps not included). Although geographical patterns of variation in some characters had been claimed in the literature, the initial sample showed that some of the variation was individual, not geographical. Accordingly, those characters were omitted (see Table 2). Soft part colors also were not included because of too few appropriately labeled specimens. Remaining characters showed variation that was either less obvi- ously individual or clearly geographical and were chosen for description and analysis. These were (1) length of maxilla measured with calipers in millimeters (mm) to the nearest 0.1 mm as a straight line from the anterior edge of the cere to the tip, (2) length of



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flattened right wing measured with a wing rule to the nearest mm, (3) head pattern and color described in terms of pileum i.e., fore- and hindcrown and frontal region, which in turn was divided into anterior frons and posterior forehead, (4) face color and pattern in terms of cheeks, feathered ocular ring, lores and ear coverts, (5) ventral patterning, i.e., pattern on the feathers of the breast and throat (barring or chevrons), (6) base color of throat and breast feathers, (7) presence or absence of red shoulder patch, and (8) color to the cast over the breast. Measurements were repeated on randomly chosen specimens at intervals of six months and variation was ? 0.1 mm.

Because of the small sample sizes for some taxa, the nonparametric Mann-Whitney U-test was used in pairwise comparisons of morphometrics among taxa using Statistica (StatSoft 1994). Sufficient numbers of specimens of both sexes were available from picta, amazonum, lucianji, roseifrons, leucotis and auricularis to test for differences between sexes in wing and bill measurements. All differences were non-significant (p > 0.05) so sexes were pooled in later analyses. Although variation in plumage characters might ideally be described quantitatively with a character index, I chose to use qualitative descriptions. The example of color of ear coverts can be used to explain this. Variation within taxa arises from the quality of specimen preparation and the degree of wear on the plumage when the bird was collected. If one is using a character index, problems would arise in deciding how to allow for those sources of variation when assigning a quantitative value to an individual specimen. Further, it was clear from the initial examination of 85 specimens that this variation was less than that between taxa. Therefore, qualitative description was considered reasonable given the scope of this study.

All latitudes and longitudes were derived from the ornithological gazetteers of Paynter (1992), Paynter and Traylor (1991) and Stephens and Traylor (1983).

RESULTS

Plumage

Table 2 summarizes trends in plumage variation among the 13 taxa. Notable variations are now described.

Western Amazonian populations currently treated as lucianii or roseifrons are variable in color and pattern of the pileum, especially with regard to the presence and extent of red. They will be examined in more detail elsewhere, but the following summarizes observations to date. Of 57 specimens examined from western Amazonia, 29 have been tentatively identified as roseifrons. All 29 are from south of the Marafnon- Amazonas rivers and 26 of them have extensive bright

red in the crown (Table 3). The remaining 28 western Amazonian birds were tentatively identified as lucianii and were mostly from north of the Marafnon-Ama- zonas rivers. They are brown-crowned and have maroon, not bright red, ocular rings. Notably in 20 of these 28 birds, the frons was wholly or partly bright red in seven of eight examined from Shanusi (6?07' S, 76015' W; AMNH 474697-474698 and 474700-474702; USNM 145680-81), in all nine from the Orosa River (3?26' S, 72008' W; AMNH 238066-238074), in all three from Santa Cecilia (3036' S, 72?57' W; ANSP 176010-176012), and in one from Sarayacu (6044' S, 75006' W; AMNH 237723). Many of these same birds had some bright or rosy red, not maroon, feathers in the ocular ring and they will be a focus of further study. A series from several localities along the Rio Santiago in far northwestern Peru (FMNH 296580, FMNH 278312, FMNH 299022-299025) had the pileum brown with no bright red in the frons or feathered ocular ring and are typical lucianii. Still concerning western Amazonia, the bright yellow ear coverts reported and illustrated in the literature for roseifrons (e.g., Forshaw and Cooper 1989; Juniper and Parr 1998) were not seen in any specimens but may have been affected by fading of specimens; interesting- ly, Gyldenstolpe (1945) described the fresher roseifrons he examined as having warm buff, not yellow, ear coverts. The yellowest ear coverts I observed were in the following specimens: AMNH 474703, an unsexed bird from Palcazu, Peru, collected in November 1902, FMNH 222875 and 222879, both males collected on 16 October 1954 at Itahuania, Peru, and specimen 81446 in the collection of the Peabody Museum, Yale Uni- versity, from Altamira, Peru collected on 2 June 1962 [examined as this paper went to press]. In all, the ear- coverts were buffy yellow, not bright yellow.

Concerning shoulder color, some otherwise green- shouldered specimens do have a few red shoulder feathers (e.g., amazonum-AMNH 285834; roseifrons- AMNH 406879, FMNH 222875-80 and subandina- ANSP 160659). Noticeably in roseifrons, the red feathers are sometimes more flame-orange than the scarlet seen in picta.

Concerning ventral patterning, the extent of the pale chevron and the base color of the feathers vary for the most part individually. However, variation of these characters in some populations can lend a distinctive pointed appearance to the breast and throat feathers. The effect is most marked in westernmost samples of amazonum, e.g., from the Rio Madeira and Rio Roosevelt (Appendix 2). In these birds the chevron's outer edge is far more extensive as well as often paler than in other taxa; as a result, the dark base to the feather, which is also contrastingly dark choco- late-brown in amazonum, is very narrow. Further, Phelps and Phelps (1949) cited this same pointed appearance of the ventral patterning as a diagnostic



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character of P. picta cuchivera from southern Venezuela but in the material I have examined of picta it varies individually. For example, the ventral patterning of AMNH 272374 from Valle de los Monos in southern Venezuela has the pointed appearance seen in the holotype of P. p. cuchivera, whereas AMNH 272371 from the same locality does not. The chevron's outer edge is also extensive in subandina, whereas it is generally reduced in western Amazonia, especially in the most red-crowned individuals. Conversely, variation in the extent of barring is markedly geographical. Barring on the feathers comprises a narrow black terminal band and a broader subterminal greyish white band (Fig. 2), which is irregularly tinged with yellow or rosy, as in chevroned specimens, yellow being most consistently developed in emma and auricularis. The subterminal band itself, however, is broadest and pale creamy in eisenmanni, caeruleiceps and griseipectus, reduced in leucotis and virtually absent in pfrimeri. Fig. 2 illustrates some major patterns of breast feathers.

Comment is warranted on the single specimen examined of pantchenkoi, its holotype, AMNH 73168, from the Sierra de Perij'a, Venezuela (10?51' N, 72?57' W). The specimen is a male (gonads 5.5 x 2.5 mm) in extremely worn plumage, though some pin feathers are visible in the frons. Allowing for the condition of the specimen's plumage, I note that ventrally it is barred with broad grey subterminal bars characteristic of caeruleiceps and eisenmanni, that its lores and ocular ring are of the same dull red as in those two taxa, and that its pileal pattern of blue frons and forecrown, brown hindcrown, and blue nuchal collar are also shared with caeruleiceps (see also Appendix 1).

Finally, one of the two emma examined, USNM 389625, and two of the eleven caeruleiceps, USNM 372618 and 372619, had the entire pileum blue.

Morphometrics

In a comparison between picta and leucotis sensu Peters (1937), wing length differed significantly (p < < 0.001) but maxilla length did not. In pairwise comparisons of wing and maxilla lengths (Table 4), subandina, emma, auricularis and amazonum were shorter winged than other populations. Amazonum and leucotis are short-billed and griseipectus is large-billed, the latter as found by Olmos et al. (1997).

DISCUSSION

A picture emerges of some characters varying in a clear geographical pattern but with only few concordances. A clear example of concordant geographical patterns is evident in cheek color and ventral patterning. All populations with barred underparts have maroon cheeks, and all those with chevroned underparts have bicolored cheeks. In contrast, neither

shoulder color nor fore- and hindcrown colors are concordant with cheek color, ventral patterning and loral coloring. For example, "picta"-type birds with bicolored cheeks and chevrons have the shoulders red (picta) or green (subandina, amazonum, western Ama- zonia), and "leucotis"-type birds with maroon cheeks and barred underparts likewise have the shoulders either red (caeruleiceps, emma, auricularis, griseipectus, pfrimeri, leucotis) or green (eisenmanni). Similarly, among small, short-winged populations, subandina and amazonum have ventral chevrons whereas emma and auricularis have bars. Individual variation is in the extent of yellow and rosy washes on the breast, the base color of breast and throat feathering and the presence of both chevrons and bars on the ventrum.

Particularly notable is the striking and hitherto unremarked resemblance between eisenmanni, caeruleiceps (including pantchenkoi, which is not diagnosably different from caeruleiceps) and, though it is less pronounced, between these two taxa and griseipectus (Fig. 3). This is due largely to the combination of broad, pale creamy subterminal bars on their ventral surfaces and their maroon cheeks. Eisenmanni and caeruleiceps further have the lores, ocular rings and frons dull red, being separable from each other by eisenmanni having no blue in the pileum and green, not red, shoulders. Similarly, eisenmanni and griseipectus differ in eisenmanni having (a) darker brown crowns, the difference perhaps accentuated due to wear in the specimens of griseipectus examined, (b) red rather than maroon frons and lores, (c) darker green dorsally, and (d) green, not red, shoulders. Phylogenetic analysis is necessary to determine whether the similarities among these three are due to relationship or convergence. Next, it is worth noting that in any assemblage of specimens, subandina and pfrimeri stand apart from each other and from all other taxa (Fig. 3; Todd, 1917; Olmos et al., 1997). Subandina is small (Tables 1 and 4), its ventral patterning is consistently and heavily washed brown, its ear coverts are russet- brown (diul ochraceous tawny of Todd, 1947), it has a unique bicolored frontal band and pace Delgado (1985) subandina and eisenmanni bear little resemblance to each other (Fig. 3). Unfortunately for such a distinctive bird (Todd, 1917, 1947), subandina has never been illustrated well in the literature. Pfrimeri has the pileum and chest entirely blue, the latter almost unbarred, and continuously maroon ear coverts and cheeks; Olmos et al. (1997, fig. 3) showed this striking pattern well.

Among the lowland Amazonian-Guianan populations, all of which have chevroned underparts, at least three groupings are diagnosable: (1) red-shouldered birds of the Guianas, Amapa (Brazil) and southern Venezuela (picta), (2) green-shouldered, dark-faced, short-winged and short-billed birds along and south of the Rio Amazonas and east of and including the



286 L. JOSEPH

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Fig. 2. Patterns of individual breast feathers of (A) picta, ANSP 188409; (B) lucotis, ANSP 175865; (C) eisenmanni, ANSP 189116; and (D) caeruleiceps, CM 54727. Note the two main patterns: squamate chevrons in pinca, and bars in the other taxa, as well as the strong similarity due to the broad subterminal band in eisenmanni and caeruliceps. Illustrations prepared by Dana Cohen.




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Madeira (amazonum), and (3) western Amazonian populations, which are variable in color and pattern of the pileum. Further study is needed of each these groupings. Present evidence strongly suggests that picta and amazonum are isolated and that amazonum may comprise disjunct populations (see Appendix 2); these inferences may of course simply reflect inade- quate sampling north and south of the Amazonas in Para, Brazil. Also, patterns of variation and taxonomy of populations in western Amazonia are not fully resolved. Pending further study, I note that brown- and red-crowned specimens so far examined have been collected to within ca. 40 km of each other at Shanusi and Tarapoto (6030' S, 760251 W), respectively. Ques- tions concerning western Amazonian populations that this study has highlighted are: what is the significance of the presence or absence of bright red in the frons in brown-crowned birds (this variation apparently not having been previously described) is the extent of red in the crown related to age as Hellmayr (1919) and Gyldenstolpe (1945) argued, how many taxa occur there, are they anywhere sympatric, and are the populations of western Amazonia more closely related to each other than to any others in the complex? Until these matters are studied afresh and settled, it is premature to merge lucianii and roseifrons. Although roseifrons might for now appropriately be applied to the often red-crowned populations south of ca. 50 S in western Amazonia, difficulties arise in determining to which populations lucianii should be applied. All such issues surrounding Amazonian populations will be examined more closely elsewhere.

The main findings from this study, which has been intended as an overview of the whole complex, are as follows. Peters' (1937) reduction of the number of species in the complex to two emphasized the few and more obvious concordances in character variation at the expense of oversimplifying the variation present in the group and the often more subtle discordances. His recognition of ventrally chevroned and bicolored- cheeked "picta" and ventrally barred and maroon- cheeked "leucotis" had the further attraction in 1937 of apparent biogeographical unity. Thus "leucotis" was distributed around the fringe of South America, while "picta" was centered in humid lowlands of Amazonia and the Guianas. A negative consequence of this taxonomy has been a tendency to overlook (a) the distinctiveness of subandina and pfrimeri (but see Todd 1917, 1947, and Olmos et al., 1997, respectively) and (b) the similarities among eisenmanni, caeruleiceps and griseipectus (Fig. 3), the first two of which, it should be noted in fairness to Peters, were described after 1937. In the context of these findings, Ridgely and Gwynne's (1989) desire to query Delgado's (1985) description of ventrally barred and maroon-cheeked eisenmanni as a subspecies of picta rather than leucotis is easy to appreciate. Nonetheless, this study shows that pending

further study of Amazonian populations, the taxa currently recognized (Table 1) are all diagnosable except (a) auricularis, which should be synonymized with the older name of emma from which it was at best only weakly diagnosed (see Zimmer and Phelps 1949) and (b) pantchenkoi, which is not diagnosable relative to the geographically close caeruleiceps with which it was not compared when described by Phelps (1977; Appendix 1).

TAXONOMIC CONCLUSIONS

Species concepts continue to be debated actively in the literature (e.g., Cantino et al., 1999). The biological inappropriateness under any species concept of recognizing two species, picta and leucotis, for the 11 taxa considered valid here is clear. So too is the need for a better understanding of phylogenetic relationships within the complex. Viewed in this way, the picta-leucotis complex might be an ideal test case for a recently proposed protocol aimed at reconciling the phylogenetic and biological species concepts (Avise and Ball, 1990; Avise, 2000). By that protocol, the species category would continue to refer, in principle, to reproductively isolated units; reproductive barriers generate genealogical depth and concordance across the different transmission pathways tracked by different pieces of DNA, i.e., different loci. They are therefore seen to be important even in strict phylogenetic frameworks. This concept is neatly illustrated in Avise (2000, fig. 6.7). Taxonomic subspecies should mirror any pronounced and concordant phylogeographic partitions observed across multiple genetic traits within a species (Avise, 2000). Species-level recognition presumably would be warranted if a reproductive barrier could also be indicated.

At present, however, it is difficult to apply this protocol to the picta-leucotis complex because of several unresolved problems. Do reproductive barriers exist among the mostly geographically isolated and diagnosable taxa of the complex? Are those taxa genealogically separated, i.e., reciprocally monophyletic for any character set under study? What are the character gradients and phylogenetic relationships among the Amazonian populations currently recognized as lucianii, roseifrons and amazonum? Nonethe- less, and given the inappropriateness of the Peters (1937) two-species arrangement, I suggest that the geographical and genetic isolation of most of the taxa coupled with their diagnosability dictates that the most useful interim taxonomy for the group ought to be devised under the phylogenetic species concept. Thus, eisenmanni, caeruleiceps-pantchenkoi, emma-auricularis, griseipectus, pfrimeri, leucotis, picta and subandina warrant recognition as species. (I question whether subandina and pfrimeri are as closely related to all other members of the complex as the latter are to each



290 L. JOSEPH

other.) Not so easy to deal with are lucianii, roseifrons and amazonum. Present evidence strongly suggests that the morphologically distinct amazonum is genetically and geographically isolated from lucianii and roseifrons as well as from picta (Appendices; Fig. 1). This brings us back to a key finding of the present study: distribu- tional and character gradients as well as phylogenetic relationships among all Amazonian populations need to be clarified. A separate paper examining the variation among Amazonian populations in more detail is in preparation. I hope that that paper as well as the present one will be steps towards the field, museum and phylogenetic studies that are necessary to resolve these issues. When all such issues associated with this complex are better understood, it should be possible to end the 63 years of uncertainty that have so far clouded the systematics of this complex. Then its systematics will adequately describe relationships among all taxa within the complex as well as between it and the rest of Pyrrhura.

ACKNOWLEDGMENTS

I wish to express my gratitude to the collectors of all specimens examined in this study as well as to curatorial staff past and present who have cared for them. For making specimens available for study and for answering queries, I thank the curators and collection managers at the following museums (abbreviations as in Materials and Methods): AMNH, USNM, CM, as well as the Natural History Museum of Los Angeles County, Los Angeles, California, the Museum of Natural Science, Louisiana State University, Baton Rouge, the Swedish Museum of Natural History, Stockholm, Sweden, the Natural History Museum, Tring, England, and the Musee National d'Histoire Naturelle, Paris, France. Comments on drafts of the manuscript and other assistance for which I am grate- ful came from L. Belasco, C. T. Fisher, A. T. Peterson, C. Reimer, N. Rice, R. Ridgely, R. Schodde, and B. Whitney. T. S. Schulenberg's critical review was especially welcome and created weeks of further, enjoyable work. T. Wilke, D. Cohen and B. Tepper (Academy of Natural Sciences, Philadelphia) kindly prepared Figs. 1-3, respectively.

LITERATURE CITED

Avise, J.C. 2000. Phylogeography; the history and forma- tion of species. Harvard University Press, Cambridge, Mass.

Avise, J.C., and R.M. Ball. 1990. Principles of genealogical concordance in species concepts and biological taxonomy. Oxford Surveys in Evolutionary Biology 7: 45-67.

Cantino, P. D., Bryant, H. N., de Queiroz, K., et al. 1999. Species names in phylogenetic nomenclature. Systematic Biology 48: 790-807.

Collar, N.J. 1997. Family Psittacidae (Parrots). Pp. 280-477 in: del Hoyo, J., Elliott, A. and Sargatal, J. eds. (1997).

Handbook of the Birds of the World. Vol. 4. Sandgrouse to Cuckoos. Lynx Ediciones: Barcelona.

Cory, C. B. 1918. Catalogue of Birds of the Americas. Volume XIII. Part II. Number 1. Field Museum of Natural History: Chicago. Publication 197.

Delgado B., F.S., 1985. A new subspecies of the Painted Parakeet (P3yrrhura picta) from Panama. Ornithological Monographs 36: 16-20.

Deville, M.E. 1851. Note sur quatre especes d'oiseaux provenant de l'expedition de M. Castelnau. Revue et Magasin de Zoologie Pure et Appliquee. Second series, 3: 209-215.

Forshaw, J.M. and Cooper, W.T. 1989. Parrots of the World. Third Edition. Landsdowne: Melbourne.

Gmelin, J.F. 1788. [Not examined. Cited in Teixeira (1991).] Gray, G.R. 1859. List of the specimens of birds in the British

Museum. Part Im, Section II, Psittacidae. British Muse- um: London.

Gyldenstolpe, N. 1945. The bird fauna of Rio Jurua in western Brazil. Kungl. Svenska Vetenskapsakademiens Handlingar 22 (3): 1-338.

Hellmayr, C.E. 1906. [Untitled. Description of Pyrrhura picta amazonum.] Bulletin of the British Ornithologists' Club 19: 8.

Hellmayr, C.E. 1907. Another contribution to the ornithology of the Lower Amazons. Novitates Zoologicae 14: 1- 39.

Hellmayr, C.E. 1919. Ein Beitrag zur Ornithologie von Sudost-Peru. Archiv fur Naturgeschichte 85 (10): 1-131.

Hilty, S.L. and Brown, W.L. 1986. A Guide to the Birds of Colombia. Princeton University Press: Princeton.

Ihering, H. 1904. 0 Rio Jurua. Revista do Museu Paulista 6: 385-460.

Juniper, T. and Parr, M. 1998. Parrots. A Guide to Parrots of the World. Yale University Press: New Haven.

Kuhl, 1820. [Not examined. Cited in Collar (1977) and Peters (1937) as Nova Acta Acad. Caes. Leop. Carol. 10: 21.]

Meyer de Schauensee, R. 1949. The Birds of the Republic of Colombia. Their distribution and keys to identification. Caldasia 5: 381-644.

Miranda-Ribeiro, A. 1920. Revisao dos psittacideos brasil- eiros. Revista do Museu Paulista 12 (Part 2): 1-82.

Muller, P.L.S., 1776. Systema Naturae Supplement. Gabriel Nildaus Kalpe: Nurnberg.

Olmos, F., Martuscelli, P. and Silva e Silva, R. 1997. Distribution and dry-season ecology of Pfrimer's Conure Jyrrhura pfrimeri, with a reappraisal of Brazilian Pyrrhura "leucotis". Ornitologia Neotropical 8: 121-132.

Paynter, R. A. 1992. Ornithological gazetteer of Bolivia. 2nd edition. Museum of Comparative Zoology, Cam- bridge, Mass.

Paynter, R. A. and Traylor, M.A. 1991. Ornithological Gazetteer of Brazil. Museum of Comparative Zoology: Cambridge, Mass.

Peters, J.L. 1937. Check-List of Birds of the World. Vol- ume Ill. Harvard University Press: Cambridge.

Phelps, W.H. and Phelps, Jr, W.H. 1949. Eleven new subspecies of birds from Venezuela. Proceedings of the Biological Society of Washington. 62: 109-124.

Phelps, W.H. and Phelps, Jr, W.H. 1958. Lista de las Aves de Venezuela con su Distribucion. Tomo II, Parte 1. No Passeriformes. Editorial Sucre: Caracas.




Phelps, Jr, W.H. 1977. Una nueva especie y dos nuevas subespecies de aves (Psittacidae, Furnariidae) de la Sierra de Perija cerca de la divisoria Colombo-Venezolana. Boletino de la Sociedad Venezolana de Ciencias Naturales 134: 43- 53.

Pinto, O.M. de 0. 1938. Catalogo das Aves do Brasil. Sao Paulo.

Pinto, O.M. de 0. 1978. Novo Catalogo das Aves do Brasil. Sao Paulo.

Ridgely, R.S. and Gwynne, J.A. 1989. A Guide to the Birds of Panama with Costa Rica, Nicaragua, and Honduras. Princeton University Press: Princeton.

Ridgway, R.A. 1912. Color Standards and Color Nomencla- ture. Smithsonian Institution: Washington, DC.

Salvadori, T. 1891. Catalogue of Birds in the British Muse- um, Volume 20. Psittaci. British Museum (Natural History): London.

Salvadori, T. 1900. On some additional species of parrots in the genus Pyrrhura. Ibis, 7th series, 6: 667-674.

StatSoft 1994. Statistica for the Macintosh. StatSoft: Tulsa. Stephens, L. and Traylor, M.A. 1983. Ornithological Gazet-

teer of Peru. Museum of Comparative Zoology: Cam- bridge, Mass.

Teixeira, D. M. 1991. Revalidagao de Pyrrhura anaca (Gmelin, 1788), do nordeste do Brasil (Psittaciformes: Psittacidae). Ararajuba 2: 103-104.

Todd, W.E. 1917. Preliminary diagnoses of apparently new birds from Colombia and Bolivia. Proceedings of the Biological Society of Washington. 30: 3-6.

Todd, W.E. 1947. New South American parrots. Annals of the Carnegie Museum 30: 331-338.

Zimmer, J.T. and Phelps, W.H. 1949. Four new subspecies of birds from Venezuela. American Museum Novitates 1395.

Appendix 1

Nomenclatural history of the parrots currently classified as subspecies of Pyrrhura picta

and P. leucotis

Pyrrhura picta (P. L. S. Muller, 1776) was the first taxon in the entire complex to be described (for all original citations of names see references above, Peters 1937 and Collar 1997). Three names have been assigned to populations of northern Amazonia and the Guianas: nominotypical picta, orinocensis Todd, 1947 and cuchivera Phelps and Phelps, 1949, the latter two having been described as subspecies of P. picta. At least since Phelps and Phelps (1958), only P. p. picta has been recognized for these populations.

Much confusion has attended study of the nomenclature and distribution of all Amazonian populations, which will be considered in more detail elsewhere. Here it is appropriate to note the following. From southeastern Amazonia, amazonum Hellmayr, 1906 was named as a subspecies of P. picta with type locality Obidos on the Amazon's north bank. From it microtera Todd, 1947 was later separated, as a subspecies of P. picta, with type locality Santarem on the south bank. The latter is now merged in P. p. amazonum. From western Amazonia, lucianii (Deville, 1851) and roseifrons (G. R. Gray, 1859) have been described. Lucianii has

been recognized consistently since its description but mostly as a subspecies of P. picta. In contrast, most authors to 1945 synonymized roseifrons with lucianii. Thus Miranda-Ribeiro (1920) recognized roseifrons but neither Hellmayr (1907, 1919), Cory (1918), Peters (1937) nor Pinto (1938) did. The latter authors therefore included the Rio Jurua in far southwestern Brazil and all Peruvian records in the range of lucianii. Gyldenstolpe (1945) resurrected roseifrons because specimens from the Rio Jurua have what he implied to be age-related but consistent red in the crown. He also noted that they differed from brown-crowned specimens on the Rio Madeira at Porto Velho. In broad terms, then, lucianii had by 1945 come to be associated with brown-crowned populations from the northern parts of western Amazonia and roseifrons with rose-crowned ones from more southern parts of that region. All later authors appear to have followed Gyldenstolpe (1945) in recognizing roseifrons Ouniper and Parr, 1998; Collar, 1997; Forshaw and Cooper, 1989; Pinto 1978).

From northern South America, subandina Todd, 1917 was described as a species isolated in northwestern Colombia. Cory (1918) followed this arrangement, but Peters (1937) treated subandina without comment as a subspecies of P. picta. Todd (1947) strongly rejoined that subandina should be accorded specific status and also introduced caeruleiceps Todd, 1947, another northwestern Colombian isolate, as a subspecies of P. subandina. Two years later and without comment, Meyer de Schauensee (1949) treated subandina and caeruleiceps as subspecies of P. picta, an arrangement followed by later authors (Hilty and Brown, 1986; Juniper and Parr, 1998; Forshaw and Cooper, 1989) with at most tentative question- ing (e.g., Collar, 1997). From the Sierra de Perija on the Venezuela-Colombia border and from the Azuero Peninsula, Panama, pantchenkoi Phelps, 1977 and eisenmanni Delgado, 1985,- respectively, were both introduced as subspecies of P. picta. Though Phelps (1977) did not compare pantchenkoi with the geographically close caeruleiceps, I note in fairness to Phelps that comparative material of caeruleiceps is scarce. Ridgely and Gwynne (1989) noted that eisenmanni may be better aligned with the leucotis group.

Nominotypical leucotis (Kuhl, 1820) occurs in southeastern Brazil north to Bahia, and griseipectus Salvadori, 1900, which Teixeira (1991) argued should be known as anaca (Gmelin, 1788), is found in northeastern Brazil. (Note that by Teix- eira's recommendation the subspecies of P. leucotis sensu Peters (1937) would become subspecies of P. -anaca.) From coastal and near-coastal central northern Venezuela, two taxa have been described: emma Salvadori, 1891, which was described as a species, and auricularis Zimmer and Phelps, 1949, which was described as a subspecies of P. leucotis. Though emma and auricularis have been recorded to within 200 kilometres of each other (Phelps and Phelps 1958), the intervening arid coastal region is probably an effective barrier. Lastly, one of the most distinctive taxa in the entire picta-leucotis complex, pfrimeri Miranda-Ribeiro, 1920, is isolated in southern central Brazil. Cory (1918) had recognized all of these as separate species, except auricularis which was then undescribed, when again Peters (1937) without comment reduced them all to subspecies of P. leucotis. Olmos et al. (1997) argued that pfrimeri merits species-level recognition.



292 L. JOSEPH

Appendix 2

Specimens Examined

picta-Guyana: A. Imprecise localities-ANSP 22380, ANSP 22382, AMNH 474670; B. Clearer localities-Baramita: USNM 586307; Demerara: AMNH 474669; Ouruma: ANSP 50594; Quonga: ANSP 50595, USNM 145687-92; Iwokrama: ANSP 187554-56, ANSP 188407-11; Annai: AMNH 474668; Surinam: A. Imprecise localities-AMNH 474665-66; ANSP 22384; B. Clearer localities-Kwata, AMNH 474661-64; vicinity of Paramaribo: AMNH 313405-06, AMNH 474667; French Guiana: A. Imprecise localities-ANSP 73992, B. Clearer localities-Pied Saut: AMNH 233719; Maura River: AMNH 233717; Cayenne: ANSP 22381, ANSP 22383; Venezuela: Valle de los Monos, Mte Duida: AMNH 272371, AMNH 272373-75; La Pricion, Caura R., AMNH 474656-59; Cano Seco, Mte Duida: AMNH 270445; Suapure: AMNH 474660; La Union, Caura: AMNH 76096-105; Mt Duida, Foothills Camp: AMNH 272377; Camp Jaime Benitez, Mt Marahua- cia: USNM 444105-06; Cerro El Negro: AMNH 42336, (holotype of P. p. cuchivera-see Appendix 1); Brazil: Cayari Island, Uassa Swamp, Amapa: AMNH 233718; Amapa, Rio Tracajatuba, 25 km E Ferreira Gomes: USNM 51469093.

amazonum-Brazil: Obidos AMNH 474671 (holotype), AMNH 474672-74; Igarape Brabo: AMNH 285834-39; Santarem: AMNH 50216, AMNH 50222; Caxiracatuba: ANSP 129260, AMNH 285833; Tauari: AMNH 285823- 32; 52 km SSW Altamira: USNM 572507-08; Humaita: AMNH 474675; Roosevelt River: AMNH 127347-49; Calama, Rio Madeira: AMNH 474677-82; Allianca Rio Madeira: AMNH 474676; Porto Velho: AMNH 148193; Diamantina: USNM 121052-55.

lucianii (tentative)-Brazil: Santo Isidoro: AMNH 308975; Peru: Orosa: AMNH 230866-74; Santa Cecilia: ANSP 176010-12; Shanusi, near Yurimaguas: AMNH 474697-702, USNM 145680-81; Betel, Amazonas, Rio Santiago: FMNH 296580; Puerto Galilea, Rio Santiago: FMNH 278362; Villa Gonzalo, Amazonas, Rio Santiago FMNH 299022-25; Sarayacu, Rio Ucayali: AMNH 237723.

roseifrons (tentative)-Brazil: Joao Pessoa, Rio Jurua: AMNH 474705, FMNH 183717; Peru: Cerro de Pantiacolla, above R. Palatoa: FMNH 300430; Taropoto: USNM 108269; Requena Ucayali: AMNH 406878-79; Itahuania, Madre de Dios: FMNH 222875-222880; Paleazu [= Palcazu of Stephens and Traylor (1983)]: AMNH 74703-04; Yurinaqui Alto: ANSP 176390, FMNH 278311; Aguas Calientes, Contamana: FMNH 320233-4; Nusiniscato Rio, Balcead- ero, Marcapata, Cuzco: FMNH 208169-70; Tsioventeni, Pasco: FMNH 297882; Nevati, Pasco: FMNH 297883; Conchapen Mt, Junin: FMNH 285078; Rio Ene at mouth of R. Quipachiari: AMNH 820834; 6 km E Luisiana: AMNH 819871; 2 km E Luisiana: AMNH 819816; La Pampa: AMNIH 145927, ANSP 103849; Bolivia: Teoponte, Rio Kaka: ANSP 121087.

subandina-Colombia: Quimari: ANSP 160658-65; Murucucui: ANSP 160666; Jaraquiel: CM 52828-29; Rio Sinu, 12 mi NW Tierra Alta: USNM 410662-67

caeruleiceps-Colombia: El Cauca: CM 54727; below Airoca: USNM 372610; Guamalito: USNM 372611-19.

pantchenkoi-Venezuela-Colombia: Frontera, Sierra de Perija: AMNH 73168 (holotype).

eisenmanni-Panama: Cascajillaso: ANSP 189116-17; Las Piraguales, El Cortezo de Tondsi: AMNH 824933, AMNH 824181 (holotype).

Ieucotis-Brazil: Rio [de Janeiro]: AMNH 474693; Bahia AMNH 474694, USNM 115223-25; Rio Doce: AMNH 139941; Baixo Girardi [?Guandu-see Paynter and Traylor 1991: 48] AMNH 317275; Casajeiras, Rio Gongojuy: AMNH 241752; Lagoa Juparana: AMNH 317276-81; Linhares: ANSP 175865, USNM 368164-65; Imprecise locality: ANSP 22378.

griseipectus-Brazil: Quixada': AMNH 241753, AMNH 241755-59; Sa do Baturite, Guaramiranga: USNM 370348- 49; Ceara: ANSP 70199.

pfrimeri-Brazil: Nova Roma, Rio Parana: AMNH 474695-96. auricularis-Venezuela: Falda Sur Cerro Humo, AMNH

41008 (holotype); Cariaquito: ANSP 58307, 58309, 58310, 58311; Cristobal Colon: AMNH 120353-61; Rio Neveri: AMNH 188162-65; Quebrada Seca, S of Cumana: AMNH 474683-90; Trinidad: USNM 145684-85.

emma-Venezuela: Los Caracas, DF: AMNH 387973; Sta Lucia, Miranda: USNM 389625.



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