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Home , Andean swallow, Atticora, Cave swallow, Notiochelidon, Petrochelidon, Progne

October1993] ShortCommunications andCommentaries 947

The Auk 110(4):947-950, 1993

Systematicsand Nomenclature of the Andean , "Petrochelidor•" ar•decola

KENNETH C. PARKES Sectionof ,Carnegie Museum of NaturalHistory, 4400 Forbes Avenue, Pittsburgh,Pennsylvania 15213, USA

The is a relatively little-known verified Chapman's color characters,but found that speciesinhabiting the Puna zone (3,100-4,600 m) of oroyaehad a significantlylonger wing, a difference the Andes of , northern Chile, and Bolivia, and overlookedby both Chapmanand Hellmayr. possiblynorthwestern Argentina (Fjeldsti and Krabbe In hisdescription of oroyae,Chapman (1924:12) made 1990).It was describedby Lafresnayeand d'Orbigny the following comments:"The cliff form in 1837from a Bolivian specimenin the ParisMuseum suchan obviouslynatural group that I hesitateto add and, like mostswallow , was originally placed to their a speciesthat doesnot wear their dis- in the all-encompassingLinnaean swallow genus Hi- tinctive pattern of marking and which is not known rundo,with the specificname andecola.It is a medium- to build their peculiar type of nest. But the factsthat sized swallow (mass14-19 g; Turner and Rose1989), I cannotfind one good generic characterseparating weakly iridescentbluish or greenish above, and gray- this speciesfrom Petrochelidonand that in juvena! ish-white below with a brownish throat. plumageandecola has the upper tail-covertsstrongly Sinceits original description,the Andean Swallow tinged with ochraceous-tawnyindicate that it may be has been moved about in a number of genera and its placedin this genuswithout undue violenceto either relationships debated. Sharpe (1885) synonymized systematicor bio!ogicornithology." andecolawith the Brown-bellied Swallow, which he Todd (1929), definitely a "splitter" at the generic identified with the cinerea of Gmelin, 1789, level, pointed out that Ridgway(1904; erroneously and placedin the genusAtticora, then a sort of waste- cited by Todd as 1902) had "dismembered the old basket genus for a miscellany of Neotropical swal- genusAtticora Bole," placing each speciesin a mono- lows. Hellmayr (1935) consideredGmelin's name to typic genus except for andecola,which, as mentioned be unidentifiable, and used the next oldest species above,he placedin his recentlydescribed genus Oro- name for the Brown-bellied Swallow ( cheildon.Todd completedthe dismembermentof the murinaCassin, 1853), while adoptingfor it the generic old Atticoraby creatinga new monotypicgenus, Hap- name Orochelidonproposed by Ridgway (1903). Al- lochelidon,for andecola.Todd stated he could not fol- though Ridgway mentioned only murinain differ- low Chapmanin referringthis species to Petrochelidon, entiating Orochelidonfrom ,Neochelidon, and "the very pointshe bringsup arguingagainst such a ,Peters (1960) pointed out that, in a later disposition." Nevertheless, all subsequentauthors publication,Ridgway (1904) included andecolain his havefollowed Chapman rather than Todd, and kept new genusas well. Peterscould find no charactersto andecolaeither in Petrochelidon(Peters 1960) or in a separate Orochelidonfrom Notiochelidon,and placed broadened Hirundo that included Petrochelidon(Tur- the Brown-bellied Swallow, but not the Andean Swal- ner and Rose1989, Sibley and Monroe 1990). low, in Notiochelidon,where it remains in the modern Chapman (1924) had pointed out that it was not literature (Turner and Rose 1989). then known whether andecolabuilt a typical Petro- Meanwhile Berlepschand Stolzmann (1896) had chelidonmud-pellet nest. In a landmarkpaper, Mayr demonstratedthat Sharpe was in error in synony- and Bond (1943) presenteda tentative reclassification mizing andecolawith "cinereaGmelin" (=murinaCas- of the swallows, giving major importance to nest sin), and that andecolawas a good species.Possibly structureas a character.They mentioned the fact that influencedby Sharpe,they placedandecola in Atticora. the nesting habits of andecolawere unknown, and That large genus subsequentlyhas been reduced to went on to say: "Its nidification should either prove the extentthat it now includesonly two species,fas- or disproveits relationshipwith typical 'Petrocheli- ciata and melanoleuca,both of which differ from an- don.' If the nesting habits of this swallow are like decolain severalways, including possessionof long, those of the speciesincluded above under 'A', the forked tails. recognitionof Todd'sHaplochelidon would seemjus- Chapman (1924) divided andecolainto two subspe- tified." Their category"A" inc!udedspecies with the cies,naming oroyaebased on three adults and an im- "nestplaced in a creviceamong rocks or in a building, maturebird from Oroyaand Chipa, Peru,using color ho!e in a tree, or in a burrow; no mud used in con- charactersand the greaterbill width of oroyaein his struction." diagnosis.With only a pair before him, Hellmayr The first descriptionof the nestinghabits of the (1935)was skepticalabout the validity of oroyae.Zim- Andean Swallow appearedin the first Spanish-lan- mer (1955) reexaminedthe type seriesand not only guage edition of LasAves de Chileby Goodall et al. 948 ShortCommunications andCommentaries [Auk, Vol. 110

(1946). In my own translation,their statementwas as cies otherwiseclosely resembles the rough-winged follows: "As is known, all of the swallows of this swallowsmorphologically; it hasalready been placed genus[i.e. Petrochelidon]have the peculiarity of nest- in Stelgidopteryxby Short (1975), followed by Ridgely ing in colonies,in crevicesor caves,on the vertical and Tudor (1989) and Sibley and Monroe (1990). It is facesof rocky cragsor canyons,and it is evident that true that the Andean Swallow is the only one of this this speciesis faithful to this tradition, seeingthat we group to exhibit dorsaliridescence, although it is only found it nesting thus in Chismisa,although unfor- weakly developed. However, within the rough- tunately on an inaccesiblecliff." As Zimmer (1955) winged swallowsuperspecies, the exceptionallydark pointed out, this seemed to settle the question of Central American subspeciesS. ridgwayistuarti shows whether andecolareally was a (! use the a faint iridescencewhen in freshly molted plumage. uncapitalized name "cliff swallow" to refer to mem- There are iridescent and noniridescent swallows al- bers of the genus or subgenusPetrochelidon in gen- ready acceptedas closely related if not congeneric, eral), although he alsocautioned that one could only suchas the Progneand Phaeoprognegroups of martins infer from the authors' statement that andecola built and the noniridescentPtyonoprogne, now generally mud nests,as this was not explicitly stated. placed in the genus Hirundo with many iridescent Niethammer (1956) describedan decola as nesting in species.Resemblances among the Andean, Tawny- holes or crevices ("L6chern") in a house wall on a headed, and rough-winged swallows are further finca in Bolivia, with no further details. This could strengthened by comparing juvenile specimens. conceivablybut improbablyrefer to mud nestsof the Chapman(1924) had invoked the reddish-brownrump Petrochelidontype. of juvenile andecolaas his only evidencefor putting The matter was finally settled when the English- the speciesin Petrochelidon.This color, however, is languageedition of TheBirds of Chileappeared (John- completely compatible with its being a member of son 1967). The accountof the nesting of andecolawas the Stelgidopteryxgroup. Within this group there is expanded to read, "Like the related Cave and Cliff alsoa strongtendency for the underpartsof juveniles Swallow that breed in the northern hemisphere, the to exhibit a rusty color, or at least a rusty wash. The Andean Cliff Swallows nest in colonies in crevices or flanksof juvenilesof andecolahave a light rustywash holes in the vertical surfacesof bluffs or escarpments; not present in adults. I have examined juveniles of when the chosenlocation is soft enough the birds all South American swallows,either at CarnegieMu- excavatethe holes themselves,breaking the surface seum of Natural History or the American Museum of with their bills, and scrapingor pushing the earth Natural History (AMNH). None are rustier or redder away with their feet like the Sand Martins [=] in juvenal plumagethan asadults except Alopochelidon of Europe. Frequently the holes are so high up that fucata, ruficollis, and "Petrochelidon"an- they can only be reachedwith the aid of ropes or decola. ladders."It is obviousthat the acceptanceof andecola Zimmer (1955), while admitting that "the generic as a Petrochelidonin TheBirds of Chilewas based solely separationof Alopochelidonfrom Stelgidopteryxis not on the fact that its nesting is colonial, the only thing very marked" except for the structural modification that andecola has in common with cliff swallows. of the primaries of the latter, added a secondcharacter If the Andean Swallow is not a cliff swallow, what to "help maintain the genus[Alopochelidon] as at pres- is it? The descriptionof its nestinghabits--sometimes ent recognized."lie statedthat the bill of Alopochel- in preexistingholes, sometimes in holes it excavates idon is notably weaker than that of Stelgidopteryx. for itself--immediatelysuggests another Neotropical However, this difference is bridged by the two sub- genus, the rough-winged swallows (Stelgidopteryx). speciesof andecola.The bill of oroyaeis even stronger Upon making comparisonsof external morphology, than that of Stelgidopteryx,whereas that of nominate I found nothing that would be incompatiblewith a andecolais much weaker and approachesthat of Al- Stelgidopteryxrelationship for andecola,and several opochelidon. points in common,such as the exceptionallylarge In view of the evidenceafforded by nesting habits undertail coverts, to which Todd (1929) called atten- and morphologicalcharacters, I regard the Andean tion in his descriptionof the genusHaplochelidon. Of Swallow as a member of a group that includes the course, andecolalacks the recurved barbs along the Tawny-headedand rough-winged swallows.In view outer edge of the outer primary that give the rough- of the gapsbetween other recognizedgenera of swal- winged swallows their name, but this characteris lows, I proposeto include all of thesein an expanded present only in adults, and is better developed in genusStelgidopteryx--those who wish to callattention malesthan in females--its significanceas a taxonomic to the distinctivecharacters of the three components character is uncertain (it also is present in Psalido- may considerthe genusto consistof three subgenera, procne,an otherwisevery differentgenus of African namely Stelgidopteryx,Alopochelidon, and Haplocheli- swallows). The recurved barbs are absent in the spe- don. ciesgenerally accepted(Peters 1960, Turner and Rose One might ask whether there is anything partic- 1989)as the closestrelative of Stelgidopteryx,the Taw- ularly important about shifting the generic affilia- ny-headedSwallow (Alopochelidonfucata). This spe- tions of a rather obscure South American swallow. October1993] ShortCommunications andCommentaries 949

This question leads to an aspect of the question of the 1976 meeting of the American Ornithologists' the relationshipsof the Andean Swallow not yet con- Union. Its conclusions were made known to several sidered, namely zoogeographicimplications. In re- ornithologists,as suggestedby the "pets. comm." in cent years there has been interest in a synthetic ap- Sibley and Monroe (1990), although those authors proach that includesdeductions about zoogeographic appear to credit me with a statementthat the vocal- relationshipsamong entire avifaunas,and about or- izations of andecolaresemble those of other Stelgidopr igins and movementsof families and genera of birds. teryx. I have made no such observations.Indeed, Every speciesthat is wrongly classifiedcreates noise Ridgely and Tudor (1989),although expressingdoubt in the system, and can lead to unwarranted conclu- that andecolatruly belongsto the Hirundo/Petrocheli- sions. For example, whether Petrochelidonis consid- doncomplex, left it there "in light of reported voice ered as a valid genus or as a subgenusof Hirundo,it similarities to other Petrochelidon(fide T. Parker)." is a group that is virtually worldwide in distribution, The enlarged Stelgidopteryxmay be defined as a but certainly of Old World origin. By the removal of group of primarily Neotropical swallows that exca- andecolafrom Petrochelidon,the only representativein vate nest holes or nest in previously existing cavities. (other than North American migrants) They differ from other swallows in the Neotropical of the entire Hirundo complex is the isolated popu- radiation (Atticora,Neochelidon, Pygochelidon, Notio- lation in Peru and Ecuador of the essentially Carib- chelidon)in having more or less rusty coloration in bean , H. (Petrochelidon)fulva, consid- the juvenal plumage, and essentiallysquare-ended, ered an allospecies,H. rufocollaris,by some authors not forked tails (difference in length between out- (Ridgely and Tudor 1989). More plausible zoogeo- ermost and central rectrices of unworn adult males graphically is the placementof andecolain a wholly lessthan 6 mm vs. 11-42 mm in other genera).The Neotropical radiation of swallows, one member of rectrices are also relatively broader, although ap- which (Stelgidopteryxserripennis) has reached the North proachedby Notiochelidonmurina in this respect.The TemperateZone relatively recently. Along the same undertail covertsare relatively long and broad. Zim- line of the effect of classificationon zoogeographic mer (1955) has already shown that the toe characters analyses,Earl• (1987)mentioned in passingin a paper used by Ridgway (1904) are not consistent. on the affinities of the African Hirundo (Petrochelidon) A referee has suggestedthat the relationshipsof fuliginosathat andecolamight not be a cliff swallow, andecola lie with Notiochelidon murina rather than with and went on to say that it "may be a Ripariaor close Stelgidopteryx.The resemblancesbetween murinaand to it, since, though it is a colonial nester,apparently andecola,however, are superficial. In addition to the it does not use mud for nest building (Zimmer 1955, tail shapeand color of juvenal plumage characters Johnson 1967)." This lumping was actually carried cited above, the wing shapesof the two genera are out by Phillips et al. (1964), who placed the rough- quite different. All Stelgidopteryxspecies in the sense winged swallow (without discussion)in Riparia,the of this paper have broadtertials, and their primaries genus of the Bank Swallow or . As the extendbeyond the tertialsby only 40 to 45% of wing classificationnow stands,the only representativeof length. In N. murinathe tertials are narrow and the Ripariain the New World, R. riparia,is such a recent primaries long, extending beyond the tertials 51 to immigrant from the Old World that the New World 56% of wing length. The undertail covertsof Stelgi- population is consideredby most authors to be sub- dopteryxare white (serripennis,fucata), white with a specificallyidentical with the European population. subterminalblack band (ruficollis,ridgwayi), or pale to The addition of the Neotropical Stelgidopteryxto Ri- dark gray with a white tip (andecola).Those of N. pariawould, of course,necessitate a completely dif- murinaare jet black,with iridescenttips, quite unlike ferent interpretation of the distributional history of thoseof Stelgidopteryx.The tertials and undertail co- Riparia.There are considerationsother than zoogeo- verts of two other speciesplaced by Sibley and Mon- graphicalagainst this particularlumping. Shortly af- roe (1990)in Notiochelidon,cyanoleuca and pileata,match ter the publication of Phillips et al. (1964), Gaunt thoseof murina(I have not examined N. fiavipes). (1965), who assessedfossorial adaptations in the anat- As a peripheral aspectof this study, I investigated omy of Ripariariparia, showed that the resemblances the validity of Chapman's(1924) subspecificdivision between the Bank and rough-winged swallows were of Stelgidopteryxandecola. I examined the four speci- superficialand the differencesmajor. More recently, mens (including holotype) at the AMNH, assigned Monson and Phillips (1981) reverted to using Stelgi- by Chapman to his new subspeciesoroyae. The three dopteryxfor the rough-winged swallows. After the specimensfrom La Oroya, Junln, appearedto be in- publicationof Earl•'s paper I sent him a copy of the tergradesbetween andecolafrom farther south and a script of my 1976 oral presentationon this subject, form representedby AMNH 174,266from Chipa, Pas- whereupon Dr. Earl• wrote me (letter of 18 December co, farther north. The Chipa specimen is distinctly 1987),"I enjoyedreading the copyof your oral paper, more sooty,less pale gray, on the abdomen,and has more so as it was an exactreflection of my thoughts blackish undertail coverts, tipped with buff. In an- on the position of andecola." decolathe abdomen is whitish to very pale gray, and A shorter version of this paper was presented at the shorter (anterior) undertail coverts are whitish, 950 ShortCommunications and Commentaries [Auk, Vol. 110 the longer onesvarying from equally whitish to light GOODALL,J. D., A. W. JOHNSON,AND R. A. PHILIPPI B. brownish gray. In the birds from La Oroya, the un- 1946. Las aves de Chile, vol. 1. Platt, Buenos dertail covertsand belly are darker than in andecola, Aires, Argentina. but the undertail coverts are still no darker than HELLMAYR,C. H. 1935. Catalogue of birds of the brownish gray. In the Chipa , the undertail co- Americasand the adjacentislands, part 8. Field verts are distinctlydarker than any other area of the Mus. Nat. Hist. Zool. Ser. 13, part $. underparts,even the throat. The Chipa specimen JOHNSON,A. W. 1967. The birds of Chile and adja- matchesthe descriptionof oroyaeand the La Oroya centregions of Argentina, Boliviaand Peru. Platt, specimensin the slightly more purplish (lessgreen- Buenos Aires, Argentina. ish)blue of the dorsaliridescence, and in having pale MAYR,E., ANDJ. BOND. 1943. Notes on the generic shafts to the primaries. As the specieshas been re- classification of the swallows, Hirundinidae. Ibis corded as far north as Artcash (Fjeldsfiand Krabbe 85:334-341. 1990), an effort should be made to obtain more north- MONSON, G., AND A. g. PHILLIPS. 1981. Annotated ern specimensto see whether they match that from checklist of the birds of Arizona, 2nd ed. Univ. Chipa. Arizona Press, Tucson. The reexaminationof avian classificationat higher NIETHAMMER,G. 1956. Zur Vogelwelt Boliviens(Tell levels has received much attention in recent years, 2: Passeres). Bonn. Zool. Beitr. 7:84-150. and hashad an impacton attemptsto constructplau- PETERS,J. L. 1960. Family Hirundinidae. Pages80- sible distributional historiesof major taxa. The swal- 129 in Check-list of birds of the world, vol. 9 (E. low cases considered here indicate that the erroneous Mayr and J. C. Greenway,Jr., Eds.).Museum of classificationof even a single speciescan have a sig- Comparative Zoology, Cambridge, Massachu- nificanteffect on zoogeographicconclusions. "Clean- setts. ing up" such errors will improve the basic distribu- PHILLIPS, A., J. MARSHALL, AND G. MONSON. 1964. tional datafundamental to future syntheticstudies of The birds of Arizona. Univ. Arizona Press, Tuc- several kinds. son. Acknowledgments.--Iam indebted to the staffof the RIDGELY, R. S., AND G. TUDOR. 1989. The birds of Bird Departmentof the AmericanMuseum of Natural South America, vol. 1. Univ. Press, Austin. History for accessto their collection.1 have enjoyed RIDGWAY,R. 1903. Descriptionsof new genera spe- stimulating correspondenceabout swallows with R. ciesand subspeciesof Americanbirds. Proc.Biol. A. Earl6 of the National Museum of South Africa, and Soc. Wash. 16:105-112. conversations with Frederick H. Sheldon of the Acad- RIDGWA¾,R. 1904. The birds of North and Middle emy of Natural Sciences,Philadelphia. America, part 3. Bull. U.S. Nat. Mus. 50, part 3. SHORT,L.L. 1975. A zoogeographicanalysis of the LITERATURE CITED South American Chaco avifauna. Bull. Am. Mus. Nat. Hist. 154:163-352. BERLEPSCH,H. VON, AND J. STOLZMANN. 1896. On the SHARPE,g. B. 1885. Catalogueof the birds in the ornithological researchesof M. JeanKalinowski British Museum, vol 10. British Museum, Lon- in central Peru. Proc. Zool. Soc. Lond. 1896:322- don. 388. SIBLEY,C. G., AND B. L. MONROE, JR. 1990. Distri- CHAPMAN,F. M. 1924. Descriptionsof new birds bution and taxonomyof the birds of the world. from Ecuador, Colombia, Peru, and Bolivia. Am. Yale Univ. Press, New Haven, Connecticut. Mus. Novit. 138:1-16. TODD,W. E.C. 1929. Haplochelidon,a new genusof EARL•,R.A. 1987. Noteson Hirundofuliginosa and its swallows. Auk 46:245. status as a "cliff swallow." Bull. Br. Ornithol. Club TURNER, A., AND C. ROSE. 1989. Swallows and mar- 107:59-63. tins. An identification guide and handbook. FJELDS•,J., AND N. KRABBE.1990. Birds of the high Houghton Mifflin, Boston. Andes. ZoologicalMuseum, Univ. Copenhagen, ZIMMER,J. T. 1955. Studies of Peruvian birds. No. Denmark. 66. The swallows (Hirundinidae). Am. Mus. Nov- GAUNt,A.S. 1965. Fossorialadaptations in the Bank it. 1723:1-35. Swallow, Ripariariparia (Linnaeus). Univ. Kans. Sci. Bull. 46:99-146. Received21 August1992, accepted 28 January1993.