Abstract

Kinglet Calyptura Calyptura cristata is one of the great enigmas of the South American avifauna. Endemic to an apparently tiny area of south-eastern Brazil, in the Atlantic Forest biome, the species was not definitely seen between sometime in the second third of the 1800s and 1996, when it was briefly rediscovered in submontane forest north-west of Rio de Janeiro. Since then, C. cristata has been reported several times, but without documentation and always by single observers. It is currently considered Critically Endangered by BirdLife International, and various authors have speculated that the species might already be extinct. Given the extreme paucity of knowledge of this species, we provide a complete inventory of museum material for Kinglet Calyptura - more than 100 specimens are listed, the majority held in European collections, almost doubling previous estimates made in the literature. Several are held in relatively small institutions, thereby suggesting that yet more specimens might still be identified or found. In addition, with the benefit of this large sample of material, we discuss morphological variation in the species and we hypothesise particularly about the appearance of male, female and juvenile plumages.

Keywords
Atlantic Forest; Museum specimens; Endangered species; Collectors’ history; Type specimen; Original description

INTRODUCTION

The genus Calyptura, Swainson, 1832, comprises just a single species, Kinglet Calyptura C. cristata, whose phylogenetic relationships have taken until the 21^st century to resolve. Traditionally, Kinglet Calyptura was treated as a member of the Cotingidae (Sclater, 1888Sclater, P.L. 1888. Catalogue of the birds in the British Museum. London, Trustees of the British Museum. v. 14.; Hellmayr, 1929Hellmayr, C.E. 1929. Catalogue of birds of the Americas and the adjacent islands. Part 6. Oxyruncidae, Pipridae, Cotingidae, Rupicolidae, Phytotomidae. Chicago, Field Museum of Natural History (Zoological Series. Publication 266).; Ames, 1971Ames, P.L. 1971. The morphology of the syrinx in passerine birds. Bulletin of the Peabody Museum of Natural History, 37: 1-194.; Snow, 1973Snow, D.W. 1973. The classification of the Cotingidae (Aves ). Breviora, 409: 1-27., 1979Snow, D.W. 1979. Family Cotingidae, cotingas. In: Traylor, M.A. (Ed.). Check-list of birds of the world. Cambridge, Museum of Comparative Zoology . v. 8, p. 281-308., 1982Snow, D.W. 1982. The Cotinga s: bellbirds, umbrellabirds and other species. London, British Museum (Natural History) & Ithaca, Cornell University Press., 2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.; Kirwan & Green, 2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .), based on its tarsal scutellation (pycnaspidean, not exaspidean like manakins and tyrannids) and foot structure, with the toes free, not more or less united (as in manakins; cf.,Snow, 1982Snow, D.W. 1982. The Cotinga s: bellbirds, umbrellabirds and other species. London, British Museum (Natural History) & Ithaca, Cornell University Press.: 39). In contrast, Olalla (1943Olalla, A.M. 1943. Algumas observações sobre a biologia das aves e mamíferos sul-americanos. Papéis Avulsos do Departamento de Zoologia de São Paulo, 3: 229-236. https://doi.org/10.11606/0031-1049.1940.1p229-236.
https://doi.org/https://doi.org/10.11606... ) suggested that Iodopleura (the purpletufts; now Tityridae) and Calyptura might form a family apart, but this proposal never acquired support, and much earlier Sclater (1888Sclater, P.L. 1888. Catalogue of the birds in the British Museum. London, Trustees of the British Museum. v. 14.: 394) had argued that “I have little doubt that Iodopleura is not its remote ally”, as well as noting, perspicaciously, that Calyptura has “Tyrannine plumage”. In the light of accumulating evidence that several genera long considered to be cotingas actually belong in other families, especially the Tityridae (see, e.g.,Ericson et al., 2006Ericson, P.G.P.; Zuccon, D.; Ohlson, J.I.; Johansson, U.S.; Alvarenga, H. & Prum, R.O. 2006. Higher level phylogeny and morphological evolution of tyrant flycatchers, cotingas, manakins and their allies (Aves: Tyrannida). Molecular Phylogenetics and Evolution, 40(2): 471-483. https://doi.org/10.1016/j.ympev.2006.03.031.
https://doi.org/https://doi.org/10.1016/... ; Ohlson et al., 2008Ohlson, J.I.; Fjeldså, J. & Ericson, P.G.P. 2008. Tyrant flycatchers coming out in the open: phylogeny and ecological radiation in Tyrannidae (Aves: Passeriformes). Zoologica Scripta, 37(3): 315-335. https://doi.org/10.1111/j.1463-6409.2008.00325.x.
https://doi.org/https://doi.org/10.1111/... ; Tello et al., 2009Tello, J.G.; Moyle, R.G.; Marchese, D.J. & Cracraft, J. 2009. Phylogeny and phylogenetic classification of the tyrant flycatchers, cotingas, manakins and their allies (Aves: Tyrannides) . Cladistics, 25(5): 429-467. https://doi.org/10.1111/j.1096-0031.2009.00254.x.
https://doi.org/https://doi.org/10.1111/... ), Ohlson et al. (2012Ohlson, J.I.; Irestedt, M.; Fjeldså, J. & Ericson, P.G.P. 2012. Nuclear DNA from a 180-year-old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides). Ibis, 154(3): 533-541. https://doi.org/10.1111/j.1474-919X.2012.01243.x.
https://doi.org/https://doi.org/10.1111/... ) finally demonstrated that Calyptura cristata is most closely related to the genera Platyrinchus and Neopipo. Furthermore, these three genera constitute a deep branch within the clade sometimes recognised as the family Rhynchocyclidae (tody-tyrants and flatbills). Ohlson et al. (2012Ohlson, J.I.; Irestedt, M.; Fjeldså, J. & Ericson, P.G.P. 2012. Nuclear DNA from a 180-year-old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides). Ibis, 154(3): 533-541. https://doi.org/10.1111/j.1474-919X.2012.01243.x.
https://doi.org/https://doi.org/10.1111/... ) proposed to recognise the clade formed by Platyrinchus + Neopipo and Calyptura at family level, the Platyrinchidae. Nevertheless, most reference works have maintained all of these taxa within the Tyrannidae (Dickinson & Christidis, 2014Dickinson, E.C. & Christidis, L. (Eds.). 2014. The Howard and Moore complete checklist of the birds of the world. 4. ed. Eastbourne, Aves Press. v. 2.; del Hoyo & Collar, 2016del Hoyo, J. & Collar, N.J. 2016. HBW and BirdLife International illustrated checklist of the birds of the world. Barcelona, Lynx Edicions . v. 2.; Gill et al., 2021Gill, F.; Donsker, D. & Rasmussen, P. (Eds.). 2021. IOC world bird list (v11.1). https://doi.org/10.14344/IOC.ML.11.1.
https://doi.org/https://doi.org/10.14344... ; Remsen et al., 2021Remsen, J.V.; Areta, J.I.; Bonaccorso, E.; Claramunt, S.; Jaramillo, A.; Lane, D.F.; Pacheco, J.F.; Robbins, M.B.; Stiles, F.G. & Zimmer, K.J. 2021. A classification of the bird species of South America. American Ornithological Society. Version 15 July 2021. Available: Available: http://www.museum.lsu.edu/~Remsen/SACCBaseline.htm . Access: 07/2021.
http://www.museum.lsu.edu/~Remsen/SACCBa... ). In other words, the size and structure of Calyptura - rather similar to Platyrinchus and Neopipo but abnormally small for a cotinga - while not necessarily taxonomically informative, provided better clues as to its relationships than might have been expected. The work of Ohlson et al. (2012Ohlson, J.I.; Irestedt, M.; Fjeldså, J. & Ericson, P.G.P. 2012. Nuclear DNA from a 180-year-old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides). Ibis, 154(3): 533-541. https://doi.org/10.1111/j.1474-919X.2012.01243.x.
https://doi.org/https://doi.org/10.1111/... ) also reinforced knowledge that tarsal scutellation is not phylogenetically informative in respect of many Tyrannides.

Kinglet Calyptura is endemic to an ostensibly very restricted range just north of the city of Rio de Janeiro, in south-east Brazil; the type is stated to be from ‘Rio de Janeiro’ (Hellmayr, 1929Hellmayr, C.E. 1929. Catalogue of birds of the Americas and the adjacent islands. Part 6. Oxyruncidae, Pipridae, Cotingidae, Rupicolidae, Phytotomidae. Chicago, Field Museum of Natural History (Zoological Series. Publication 266).), which has generally been interpreted as meaning the immediate environs of the city itself (e.g.,Pinto, 1944Pinto, O.M.O. 1944. Catálogo das aves do Brasil. 2ª parte. São Paulo, Departamento de Zoologia.; Sick & Pabst, 1968Sick, H. & Pabst, L.F. 1968. As aves do Rio de Janeiro (Guanabara). Lista sistemática anotada. Arquivos do Museu Nacional do Rio de Janeiro, 53: 99-160.; Snow, 1982Snow, D.W. 1982. The Cotinga s: bellbirds, umbrellabirds and other species. London, British Museum (Natural History) & Ithaca, Cornell University Press.). Ruschi (1953Ruschi, A. 1953. Lista das aves do Estado do Espírito Santo. Boletim do Museu de Biologia Professor Mello Leitão, série Zoologia, 11: 1-21.) listed the species for the state of Espírito Santo, but this was doubted by King (1978-1979King, W.B. 1978-1979. Red Data book, 2. Aves. 2. ed. Morges, IUCN.) and rejected firmly by Collar et al. (1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.) and Pacheco & Bauer (2001Pacheco, J.F. & Bauer, C. 2001. A lista de aves do Espírito Santo de Augusto Ruschi (1953): uma análise crítica. In: Albuquerque, J.L.B., Cândido, J.F.; Straube, F.C. & Roos, A.L. (Eds.). Ornitologia e conservação: da ciência às estratégias. Tubarão, Editora Unisul. p. 261-278.). A specimen, purported to have been collected somewhere in the state of São Paulo between May 1819 and April 1820, was discovered in the Museum für Naturkunde, Berlin, in 2007 (Stopiglia et al., 2009Stopiglia, R.; Straker, L.C. & Raposo, M.A. 2009. Kinglet Calyptura Calyptura cristata (Vieillot, 1818): documented record for the state of São Paulo and taxonomic status of the name Pipra tyrannulus Wagler, 1830. Bulletin of the British Ornithologists’ Club , 129(3): 185-188.); however, its true provenance has not been satisfactorily established (Rego et al., 2013Rego, M.A.; Moreira-Lima, L.; Silveira, L.F. & Frahnert, S. 2013. On the ornithological collection of Friedrich Sellow in Brazil (1814-1831), with some considerations about the provenance of his specimens. Zootaxa, 3616(5): 478-484. https://doi.org/10.11646/zootaxa.3616.5.4.
https://doi.org/https://doi.org/10.11646... ). Given the number of specimens mentioned in the previous literature, the species was not uncommon, even in secondary forested habitats, in the early to mid-19^th century (Collar et al., 1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.; Kirwan & Green, 2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .), but virtually the only available information concerning its habits/ecology is the testimony of Descourtilz (1852Descourtilz, J.T. 1852. Ornithologie Brésilienne, ou histoire des oiseaux du Brésil remarquables par leur plumage, leur chant ou leurs habitudes. Rio de Janeiro, Thomas Reeves.). The latter author mentioned C. cristata as feeding on fruit and insects, usually in pairs in the midstorey, maintaining contact with surprisingly loud vocalisations. A trickle of specimens continued to reach overseas museums with the last said to have been collected around 1890 (Snow, 2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.: 88, repeated by Kirwan & Green, 2011), but as we show in this paper several were accessioned even later than this, although we generally lack robust knowledge of when they were actually collected. Thereafter the species went unrecorded until two individuals were seen in the Serra dos Órgãos, at Guapimirim, in the environs of Teresópolis, by multiple observers on several days in late October 1996 (Sick, 1997Sick, H. 1997. Ornitologia brasileira. Rio de Janeiro, Editora Nova Fronteira.; Pacheco & Fonseca, 2000Pacheco, J.F. & Fonseca, P.S.M. 2000. A admirável redescoberta de Calyptura cristata por Ricardo Parrini no contexto das preciosidades aladas da Mata Atlântica. Atualidades Ornitológicas , 93: 6-7., 2001Pacheco, J.F. & Fonseca, P.S.M. 2001. The remarkable rediscovery of the Kinglet Calyptura Calyptura cristata. Cotinga, 16: 48-51.). Most of the few specimens with reasonably precise locality data come from this general region. There have been no reliable records since 1996 despite searches, mainly in Rio de Janeiro state, for example in the Reserva Ecológica Guapiaçu, the Teresópolis area, the foothills of the Serra do Mar, and between Nova Friburgo and Sumidouro, during September-November 2006 (Lambert & Kirwan, 2010Lambert, F. & Kirwan, G.M. 2010. The twice-vanishing ‘pardalote’: what future for the Kinglet Calyptura? Neotropical Birding, 6: 4-17.) and in October 2016 (https://ebird.org/news/kingletcalyptura2016?tagId=128). Nevertheless, there have been several claimed records of the species, all by single observers and from the broad environs of Ubatuba in the state of São Paulo, in July 1990, March 1997, March 2006 and September 2008 (Sigrist, 2006Sigrist, T. 2006. Aves do Brasil: uma visão artística. Vinhedo, Editora Avis Brasilis.; Lambert & Kirwan, 2010Lambert, F. & Kirwan, G.M. 2010. The twice-vanishing ‘pardalote’: what future for the Kinglet Calyptura? Neotropical Birding, 6: 4-17.; Kirwan & Green, 2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .). Earlier, D.F. Stotz (inRidgely & Tudor, 1994Ridgely, R.S. & Tudor, G. 1994. The birds of South America. Austin, University of Texas Press. v. 2.) had speculated that the species might eventually be found in this region. Whilst BirdLife International (2023BirdLife International. 2023. Species factsheet: Calyptura cristata. Available: Available: http://datazone.birdlife.org/species/factsheet/kinglet-calyptura-calyptura-cristata . Access: 27/01/2023.
http://datazone.birdlife.org/species/fac... ) currently treats the species as Critically Endangered, with a population expected to number fewer than 50 individuals, some authors have speculated that the lack of definite records in the last c. 25 years suggests that C. cristata is likely to be extinct (Lees & Pimm, 2015Lees, A.C. & Pimm, S. 2015. Species, extinct before we know them? Current Biology, 25(7): R177-R180. https://doi.org/10.1016/j.cub.2014.12.017.
https://doi.org/https://doi.org/10.1016/... ).

Given that virtually our entire knowledge of this species is based on the specimen record, it is pertinent to assemble a complete inventory of such material and its provenance. Various authors have commented on the number of specimens in natural history museums. For example, Snow (1982Snow, D.W. 1982. The Cotinga s: bellbirds, umbrellabirds and other species. London, British Museum (Natural History) & Ithaca, Cornell University Press.: 39) stated that “there exist only a handful of specimens in a few museums”, whereas Collar et al. (1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.: 726) mentioned that there were more than 45 specimens, preserved in AMNH, ANSP, NHMUK, RBINS, NML-VZ, MCZ, MNHN, NMW, UMZC, USNM, ZMB and ZMUC (for museum acronyms, see Methods). The figure of 45 was repeated by Pacheco & Fonseca (2000Pacheco, J.F. & Fonseca, P.S.M. 2000. A admirável redescoberta de Calyptura cristata por Ricardo Parrini no contexto das preciosidades aladas da Mata Atlântica. Atualidades Ornitológicas , 93: 6-7.), but a year later the same authors stated that the total was approximately 50 (Pacheco & Fonseca, 2001Pacheco, J.F. & Fonseca, P.S.M. 2001. The remarkable rediscovery of the Kinglet Calyptura Calyptura cristata. Cotinga, 16: 48-51.). Presumably drawing on the latter, Snow (2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.: 88) repeated the c. 50 assertion. Tobias et al. (2006Tobias, J.A.; Butchart, S.H.M. & Collar, N.J. 2006. Lost and found: a gap analysis for the Neotropical avifauna. Neotropical Birding , 1: 4-22.) did not speculate on the actual number of specimens, but suggested that their number did provide a hint as to the species’ former abundance; Krabbe (2007Krabbe, N. 2007. Birds collected by P.W. Lund and J.T. Reinhardt in south-eastern Brazil between 1825 and 1855, with notes on P.W. Lund’s travels in Rio de Janeiro. Revista Brasileira de Ornitologia, 15(3): 331-357.) also reported that there are “nearly 50 specimens”. Subsequently, Lambert & Kirwan (2010Lambert, F. & Kirwan, G.M. 2010. The twice-vanishing ‘pardalote’: what future for the Kinglet Calyptura? Neotropical Birding, 6: 4-17.), Kirwan & Green (2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .: 578) and Ohlson et al. (2012Ohlson, J.I.; Irestedt, M.; Fjeldså, J. & Ericson, P.G.P. 2012. Nuclear DNA from a 180-year-old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides). Ibis, 154(3): 533-541. https://doi.org/10.1111/j.1474-919X.2012.01243.x.
https://doi.org/https://doi.org/10.1111/... ) all mentioned the existence of c. 55 specimens; this revised total was based on work conducted by GMK’s own studies in museums additional to those mentioned by Collar et al. (1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.), as well as specific publications in the interim (e.g.,McGhie, 2005McGhie, H.A. 2005. Specimens of extinct and endangered birds in the collections of the Manchester Museum. Bulletin of the British Ornithologists’ Club , 125(4): 247-252.). Finally, Hume & Walters (2012Hume, J.P. & Walters, M. 2012. Extinct birds. London, T. & A.D. Poyser.: 354) noted the presence of specimens in various museums of which those in Florence (MZUF), in Italy and Kiel (ZMK), in Germany, were extra to the inventory of Collar et al. (1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.) and those additional institutions visited by GMK.

Here, we report the existence of 104 specimens and provide as complete as possible an inventory of museum material, with details on its provenance and dating, based on research conducted by ourselves, separately and collaboratively. We also present a review of the species’ morphological (sexual and age-related) variation based on photographs and personal examination of the specimens located to date. We also present photographs of all of these specimens, as well as historical information concerning their provenance.

MATERIAL AND METHODS

Building on the brief history concerning our collective knowledge of specimens in the world’s museums outlined above, we used a combination of personal contacts and consultation, published information (e.g.,Fisher, 1981Fisher, C.T. 1981. Specimens of extinct, endangered or rare birds in the Merseyside County Museums, Liverpool. Bulletin of the British Ornithologists’ Club, 101(2): 276-285.; Lima, 2005Lima, L.M. 2005. O acervo ornitológico brasileiro do Finnish Museum of Natural History, Helsinki, Finlândia. In: Congresso Brasileiro de Ornitologia, Belém, 13º. Resumos. Belém, Sociedade Brasileira de Ornitologia. p. 160.; McGhie, 2005McGhie, H.A. 2005. Specimens of extinct and endangered birds in the collections of the Manchester Museum. Bulletin of the British Ornithologists’ Club , 125(4): 247-252.; Ghiraldi & Aimassi, 2019Ghiraldi, L. & Aimassi, G. 2019. Extinct and endangered (‘E&E’) birds in the ornithological collection of the Museum of Zoology of Torino University, Italy. Bulletin of the British Ornithologists’ Club , 139(1): 28-45. https://doi.org/10.25226/bboc.v139i1.2019.a2.
https://doi.org/https://doi.org/10.25226... ), as well as web searches of online museum catalogues and broader compendia (e.g., VertNet, GBIF, etc.), plus appeals on internet fora, e.g., the eBEAC mailing list. In particular, we sought information from smaller museums in France, Germany, Italy and Switzerland, given the evidence already furnished by Hume & Walters (2012Hume, J.P. & Walters, M. 2012. Extinct birds. London, T. & A.D. Poyser.) and Waldeck (2018Waldeck, M. 2018. The mysterious Kinglet Calyptura. Available: Available: http://thelittlestint.blogspot.com/2018/02/the-mysterious-kinglet-calyptura.html . Access: 06/05/2021.
http://thelittlestint.blogspot.com/2018/... ) that such institutions harboured previously unnoticed specimens of C. cristata. Our requests for information were coupled with solicitations for photographs of any specimens in order to assess plumage variation in the species. These specimen images originated from many museums, each one of which produced their photographs in different ways, ranging from professional standard, with excellent quality artificial lighting and in high resolution, to photographs made with mobile phones under natural light conditions.

The acronyms of those museums wherein specimens of Calyptura cristata were located are as follows: AMNH = American Museum of Natural History, New York, USA; ANSP = Academy of Natural Sciences at Drexel University, Philadelphia, USA; CCECL = Centre de Conservation et d’étude des Collections, Musée des Confluences, Lyon, France; CUMV = Cornell University Museum of Vertebrates, Ithaca, USA; IZH = Institute für Zoologie, Martin-Luther-Universität, Halle-Wittenberg, Germany; LMNM = Landesmuseum Natur und Mensch, Oldenburg, Germany; LUOMUS = Finnish Museum of Natural History, Helsinki, Finland; MAB = Musée de l’Areuse Boudry, Neuchâtel, Switzerland; MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, USA; MHH = Museum Heineanum, Halberstadt, Germany; MHNC-UP = Museu de História Natural e Ciências da Universidade do Porto, Portugal; MHNGr = Muséum d’Histoire Naturelle de Grenoble, France; MHNMON = Muséum d’Histoire Naturelle Victor Brun, Montauban, France; MHNN = Muséum d’Histoire Naturelle de Neuchâtel, Switzerland; MHNVT = Musée d’histoire Naturelle et Vivarium de Tournai, Belgium; MM = Manchester Museum, University of Manchester, UK; MNHN = Muséum National d’Histoire Naturelle, Paris, France; MNRJ = Museu Nacional, Rio de Janeiro, Brazil; MPUW = Muzeum Przyrodnicze, Uniwersytet Wroctawski, Poland; MSNT = Museo Civico di Storia Naturale, Trieste, Italy; MSNTP = Museo di Storia Naturale e del Territorio, Università di Pisa, Italy; MWNH = Landesmuseum Wiesbaden, Naturwissenschaftlichen Sammlung, Wiesbaden, Germany; MZLU = Zoologisk Museum, Universitet fran Lund, Sweden; MZS = Musée Zoologique de la ville de Strasbourg, France; MZUF = Museo di Storia Naturale dell’Università di Firenze, Italy; MZUT = Museo di Zoologia dell’Università di Torino, Italy; NHMO = Naturhistorisk Museum, Universitetet i Oslo, Norway; NHMUK = Natural History Museum, Tring, UK; NLMH = Niedersächsisches Landesmuseum, Naturkunde-Abteilung, Hannover, Germany; NMB = Naturhistorisches Museum Basel, Switzerland; NMBE = Naturhistorisches Museum Bern, Switzerland; NML-VZ = World Museum, National Museums Liverpool, UK; NMSG = Naturmuseum Sankt Gallen, Sankt Gallen, Switzerland; NMW = Naturhistorisches Museum Wien, Austria; NRM = Naturhistoriska Riksmuseet, Stockholm, Sweden; RBINS = Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium; RMNH = Naturalis Biodiversity Center, Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands; SMF = Senckenberg Naturmuseum Frankfurt, Germany; SMNG = Staatliches Museum für Naturkunde Görlitz, Germany; SMNS = Staatliches Museum für Naturkunde, Stuttgart, Germany; UMMZ = University of Michigan, Museum of Zoology, Ann Arbor, USA; UMZC = University Museum of Zoology Cambridge, UK; USNM = United States National Museum, Washington, USA; ZIMG = Zoologisches Museum, Institut für Zoologie und Anthropologie der Universität Göttingen, Göttingen, Germany; ZISP = Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia; ZMB = Museum für Naturkunde, Berlin, Germany; ZMK = Zoologisches Museum der Christian-Albrechts-Universität Kiel, Germany; ZMMU = Moscow Lomonosov State University, Russia; ZMUC = Zoologisk Museum i Københavns Universitet, Denmark; ZMUL = Aquarium-Muséum de l’Université de Liège, Belgium.

RESULTS

Inventory

To date 104 specimens of C. cristata have been identified in 47 museums, the majority in European collections (Tables 1 and 2). Of this material, 86 specimens are held in a total of 39 different European institutions, 17 at seven collections in the USA, and one specimen is held in Brazil (a second, currently ZMB 2000.12102, is due to be sent to MZUSP in the forseeable future; P. Eckhoff in litt., 2023). Among the specimens listed herein, ten are in eight different smaller museums, considered part of the European ‘B’ list of institutions each harbouring fewer than c. 4,000 skins, or c. 5,000 bird items in total (Roselaar, 2003Roselaar, C.S. 2003. An inventory of major European bird collections. In: Collar, N.J.; Fisher, C.T. & Feare, C.J. (Eds.). Why museums matter: avian archives in an age of extinction. Bulletin of the British Ornithologists’ Club Supplement, 123A. p. 253-337.), or not mentioned in the latter inventory (e.g., MAB). An additional 12-15 specimens (Table 3) are known or are currently believed to be lost (see footnotes 7 and 8 to Table 3, and note 45 to Table 2, for reasons to consider that the total of “lost” specimens is 12, rather than 15).

Figure 1
Holotype of Calyptura cristata (MNHN ZO-MO-2004-300), (A) dorsal view; (B) ventral view; (C) lateral view. Photos: Muséum National d’Histoire Naturelle, Paris.

Figure 2
Extant mounted specimens of Calyptura cristata, acronyms are listed in Methods. (1) IZH-V 3260, photo: Frank Steinheimer/IZH; (2) LUOMUS 1573, photo: Hanna Laakkonen/LUOMUS; (3) MHNN 926029, photo: Celia Bueno/MHNN; (4) MNHN 3137, photo: Vitor Piacentini; (5) NRM 535288, photo: Ulf Johansson/NRM; (6) MHNGr OR.8050, photo: MHNGr; (7) MHNVT R1-E6-C151-001, (8) MHNVT R1-E6-C151-002, photos: Christophe Remy/MHNVT; (9) NMW 17350, photo: Hans-Martin Berg/NMW; (10) MSNT, photo: Nicola Bressi; (11) MZLU Aves L848/6080, photo: Maria Mostadius/MZLU; (12) MZS Ave 08286, photo: Marie Meister/MZS; (13) NMSG 5741, photo: Lorenzo Vinciguerra/NMSG; (14) MPUW 203024, photo: Jan Lotkowski/MPUW; (15) MHNMON O.2427, (16) MHNMON O.2426, photos: Aude Medina/MHNMON; (17) ZISP 1819, photo: Vladimir Loskot/ZISP; (18) LMNM 5741, photo: Mathieu Waldeck; (19) NMBE 1033790, photo: Manuel Schweizer/NMBE; (20) MAB, photo: Pierre-Henri Béguin; (21) RMNH.AVES.172607, (22) RMNH.AVES.172611, (23) RMNH.AVES.172610, (24) RMNH.AVES.172609, (25) RMNH.AVES.172608, photos: Robson Silva e Silva; (26) MSNTP AV3082, (27) MSNTP AV3083, (28) AV3081, photos: Simone Farina & Lorenzo Vanni/MSNTP; (29) NMB 2042, (30) NMB 2041, photos: Dieter Thomas Tietze/NMB; (31) SMNG A07725a, photo: Diana Jeschke/SMNG; (32) ZMUC 105507, photo: Jon Fjeldså/ZMUC; (33) MHNC-UP AVE-1125, (34) MHNC-UP AVE-1126, photos: Mariana Costa/MHNC-UP.

Figure 3
Extant skin specimens of Calyptura cristata, acronyms are listed in Methods. (1) AMNH 5156, (2) AMNH 494721, (3) AMNH 494720, (4) AMNH 494719, (5) AMNH 43795, photos: Jason Sheldon/AMNH; (6) ANSP 8366, (7) ANSP 8367, (8) ANSP 8368, photos: Nate Rice/ANSP; (9) CCECL 41007379, photo: Cédric Audibert/CCECL; (10) MCZ 75787, (11) MCZ 85035, photos: MCZ © President and Fellows of Harvard College; (12) LUOMUS 4950, (13) LUOMUS 4949, photos: Hanna Laakkonen/LUOMUS; (14) MHH 5667, (15) MHH 5666, (16) MHH 5665, photos: Rüdiger Becker/MH; (17) MNHN ZO-MO-2002-661, (18) MNHN ZO-MO-2000-2155, (19) MNHN ZO-MO-2000-2154, (20) MNHN ZO-MO-2000-2153, (21) MNHN ZO-MO-1931-1285, (22) MNHN ZO-MO-1845-440, photos: Vitor Piacentini; (23) NHMUK 1895.4.1.731, (24) NHMUK 1895.4.1.730, (25) NHMUK 1888.1.20.972, (26) NHMUK 1888.1.13.1675, (27) NHMUK 1881.5.1.3739, photos: Jonathan Jackson/NHMUK; (28) MZUT AV16101, photo: Luca Ghiraldi/MZUT; (29) NMW 17351, (30) NMW 17349, photos: Hans-Martin Berg/NHMW; (31) NML-VZ 5055a, (32) NML-VZ 3029, (33) NML-VZ 1980-70a, photos: John-James Wilson/NML-VZ; (34) SMF 41364, photo: Gerald Mayr/SMF; (35) RBINS 10126B, photo: Olivier Pauwels/RBINS.

Figure 4
More extant skin specimens of Calyptura cristata, acronyms are listed in Methods. (1) UMZC 27/Cot/4/a/2, (2) SMNS 38253, (3) SMNS 33477, (4) SMNS 33478 photos: Guy M. Kirwan; (5) SMNS 114246, photo: Friederike Woog/SMNS; (6) UMMZ 134367, photo: Brett Benz. Image used with permission of University of Michigan Museum of Zoology Bird Division; (7) ZISP 117201, photo: Robson Silva e Silva; (8) USNM 33161, (9) USNM 33162, (10) USNM A15195, (11) USNM A15224, (12) USNM 145362, photos: Brian Schmidt/USNM; (13) ZISP 117204, photo: Robson Silva e Silva; (14) MWNH AV 5323, photo: Fritz Geller-Grimm/MWNH; (15) MNRJ 33299, photo: Claydson Assis/MNRJ; (16) NHMO BI-66671/1-P, (17) NHMO BI-66672/1-P, photos: Lars Erik Johannessen/NHMO; (18) ZMB 2000.12101, (19) ZMB 2000.12102, (20) ZMB 2000.12103, (21) ZMB 2000.12104, (22) ZMB 2305, (23) ZMB 2306, (24) ZMB 7567, photos: Vitor Piacentini; (25) ZMMU R525, photo: Pavel Smirnov/ZMMU; (26) CUVM 48403, photo: Vanya Gregor Rohwer/CUMV; (27) MHNN 92.6030, photo: Celia Bueno/MHNN; (28) MM B6327, (29) MM B6328, photos: MM; (30) NRM 90127687, photo: Ulf Johansson/NRM; (31) UMZC 27/Cot/4/a/1, photo: Nigel Collar; (32) ZISP 117202, (33) ZISP 117203, photos: Robson Silva e Silva; (34) ZMUC 105508, photo: Guy M. Kirwan; (35) MNHN ZO-MO-2004-300, photo: Muséum National d’Histoire Naturelle, Paris.

One of the specimens added to the list in a small Swiss museum (MAB) was previously misidentified as a species from North America, a Regulidae, the Ruby-crowned Kinglet, Corthylio calendula (Linnaeus, 1766), labelled: “Regulus calendula ♂, le Roitelet, Amerique du Nord”. It was discovered by Julien Mazenauer who recognised the specimen in the museum’s public exhibition as a Kinglet Calyptura.

One of the two RBINS specimens (RBINS 10126A) was removed from the list, because it is not a C. cristata, but rather a Tyrannidae, possibly a Yellow-crowned Tyrannulet Tyrannulus elatus (Latham, 1790), despite its label declaring it to be C. cristata.

Of the 104 extant specimens, just eight (IZH-V 3260, NHMUK 1888.1.20.972, NMB 2041, NMB 2042, NMSG 5741, SMNG A07725a, ZMUC 105508 and ZMUC 105507) possess locality information that is any way specific (“Nova Friburgo”, “Cantagalo” or “Rosário”). These three localities lie within 45 km of each other. Nova Friburgo (also Neu Freiburg or Novo Fribourgo) is at c. 850 m elevation in the Serra dos Órgãos, in north-central Rio de Janeiro state (22°16′S, 42°32′W). This general area was visited by several relevant collectors, including Peter Wilhelm Lund, Jean-Théodore Descourtilz, Karl Hermann Konrad Burmeister and, but see below, John Youds (see Paynter & Traylor, 1991Paynter, R.A. & Traylor, M.A. 1991. Ornithological gazetteer of Brazil. Cambridge, Museum of Comparative Zoology., and references and notes therein). Cantagalo (formerly spelt Cantagallo) lies c. 40 km north-east of Nova Friburgo, at c. 21°58′S, 42°22′W, c. 405 m, is where Carl Hieronymus Euler owned a large farm, Fazenda Bom Valle (c. 21°56′S, 42°16′W, c. 355 m), in what is now the small town of Euclidelândia, 44 km from the centre of Nova Friburgo. Both Lund and Burmeister also visited Cantagalo (see Paynter & Traylor, 1991). Rosário (sited at c. 22°16′S, 42°32′W, c. 855 m, according to Paynter & Traylor, 1991, but 22°06′S, 42°25′W, c. 655 m, by Krabbe, 2007Krabbe, N. 2007. Birds collected by P.W. Lund and J.T. Reinhardt in south-eastern Brazil between 1825 and 1855, with notes on P.W. Lund’s travels in Rio de Janeiro. Revista Brasileira de Ornitologia, 15(3): 331-357.) was another farm in the environs of Nova Friburgo; alternatively, JFP and others have visited a locality that is still called Rosário, at the border of the municipalities of Bom Jardim and Duas Barras, at c. 22°08′S, 42°29′W, c. 1,110 m, and just 15 km north-east of Nova Friburgo. Krabbe (2007) indicated that Lund was at Rosário almost permanently between 8 February 1827 and late June 1828, and that the fazenda was somewhere between Nova Friburgo and Cantagalo, but not as close to the former as the coordinates given by Paynter & Traylor (1991) suggested. Krabbe (2007) also noted that Lund may well have taken many of the birds labelled Rosário on a forested mountain named “Morro Queimado” (“Burnt Hill”), perhaps as far as several hours walk from the farm. A “Morro Quemado” (presumably the same locality) was also mentioned by another contemporary observer, Descourtilz. However, D. Miller (inLambert & Kirwan, 2010Lambert, F. & Kirwan, G.M. 2010. The twice-vanishing ‘pardalote’: what future for the Kinglet Calyptura? Neotropical Birding, 6: 4-17.) ascertained that Morro Queimado was the headquarters of a fazenda of the same name depicted in the background of an aquatint produced in the late 1820s, and that “Morro Queimado” was sometimes used synonymously with “Nova Friburgo” when George Gardner visited in 1840 (Miller et al., 2006Miller, D.; Warren, R.; Moura Miller, I. & Seehawer, H. 2006. Serra dos Órgãos. Sua história e suas orquídeas. Rio de Janeiro, Editora Soart.). According to Miller, “Morro Queimado” refers to the area around the farm headquarters (probably the present base of Anchieta College, Nova Friburgo; https://pt.wikipedia.org/wiki/Fazenda_do_Morro_Queimado), and the probable elevation of the original forest in this area would have been 600-900 m (Lambert & Kirwan, 2010).

Some specimens lack any indication of locality, while others are erroneous (e.g., AMNH 5156, labelled “Guiana, Suriname”; NML-VZ 3029, “Bogotá”; RMNH AVES.172607, “México”), but most are labelled simply Brazil (Brésil, Brasilia, Brasilien, Brasile) (n = 57), “southeast Brazil” (n = 6), or occasionally “Rio de Janeiro” (n = 11). Others are, inaccurately, attributed more generally to the Americas (CCECL 41007379, MHNVT R1-E6-C151-002). Two (MSNTP AV3081, AV3083) are labelled “Nuova Olanda” (= New Holland), in an apparent (but clearly erroneous) allusion to the area of north-east Brazil, stretching from Sergipe to Maranhão, and centred on Recife, which was formerly administered by the Dutch, until it was ceded to the Portuguese crown in 1661 (Lockhart & Schwartz, 1983Lockhart, J. & Schwartz, S.B. 1983. Early Latin America. New York, Cambridge University Press.). One specimen (ZMB 2306) stemming from Friedrich Sellow mentions São Paulo (“San Paulo”) on the label, and Stopiglia et al. (2009Stopiglia, R.; Straker, L.C. & Raposo, M.A. 2009. Kinglet Calyptura Calyptura cristata (Vieillot, 1818): documented record for the state of São Paulo and taxonomic status of the name Pipra tyrannulus Wagler, 1830. Bulletin of the British Ornithologists’ Club , 129(3): 185-188.) contested that this provided the first incontrovertible proof that C. cristata is not endemic to the state of Rio de Janeiro. Rego et al. (2013Rego, M.A.; Moreira-Lima, L.; Silveira, L.F. & Frahnert, S. 2013. On the ornithological collection of Friedrich Sellow in Brazil (1814-1831), with some considerations about the provenance of his specimens. Zootaxa, 3616(5): 478-484. https://doi.org/10.11646/zootaxa.3616.5.4.
https://doi.org/https://doi.org/10.11646... ), however, advocated caution in placing too much faith in the label data because Sellow sent multiple shipments of specimens to ZMB during the relevant period, and because of gaps in the original documentation in the museum’s archive concerning these arrivals.

Only a small percentage of the material can be more or less accurately dated as to when it was collected, all of which was probably acquired during the first two-thirds of the 19^th century, for example IZH-V 3260, MCZ 75787, MNHN 3137, MSNT uncatalogued, ZMB 2305 and 2306, and ZMUC 105507 and 105508 (for the latter two specimens, see Krabbe, 2007Krabbe, N. 2007. Birds collected by P.W. Lund and J.T. Reinhardt in south-eastern Brazil between 1825 and 1855, with notes on P.W. Lund’s travels in Rio de Janeiro. Revista Brasileira de Ornitologia, 15(3): 331-357.). For the majority of traced specimens, the date of registration in a European or North American institution, or the date of acquisition, donation or exchange makes identifying when they were collected extremely difficult, probably impossible, especially given a lack of knowledge of the collectors involved. Nevertheless, where archival material might be available, we encourage curators and researchers at the museums concerned to pursue such avenues of investigation as may be open to them. Although some material was certainly not accessioned until the last third of the 19^th century, e.g., NMBE 1033790 (sometime in the 1870s) and all of that at NHMUK (in the 1880s and 1890s), and a few specimens (MM B6327, MNHN ZO-MO-2000-2153 and MNHN ZO-MO-1931-1285) were not registered in other European collections until the first third of the 20^th century, in none of these cases do we possess definite evidence that they were collected so late. For example, the NHMUK specimens came to the British Museum via P.L. Sclater (n = 1), John Gould (n = 1), the Salvin-Godman collection (n = 1) and Alexander Fry (n = 2). That originally in Sclater’s private collection must have been collected prior to 1862, given that it was listed in his own catalogue published in that year (Sclater, 1862: 247), and the specimen reached him via the natural history specimen dealer, James Argent, about whom very little seems to be known, but the British Museum purchased a total of 500 specimens directly from him between 1843 and 1854, including a batch of 30 from Brazil in 1846 (Sharpe, 1906Sharpe, R.B. 1906. The history of the collections contained in the natural history departments of the British Museum, vol 2. Birds. London, Trustees of the British Museum . https://doi.org/10.5962/bhl.title.14168.
https://doi.org/https://doi.org/10.5962/... : 300-301). We lack any knowledge concerning the provenance of Gould’s specimen, which was purchased by the British Museum in 1881, on his death; neither a dealer nor a collector is mentioned on the museum’s label (no original label is attached to the specimen). A trawl of Gould’s correspondence (reproduced by Sauer, 1998a">Sauer, G.C. 1998a. John Gould the bird man: correspondence. Mansfield Centre, CT, Maurizio Martino . v. 1., 1998bSauer, G.C. 1998b. John Gould the bird man: correspondence . Mansfield Centre, CT, Maurizio Martino . v. 2., 1999Sauer, G.C. 1999. John Gould the bird man: correspondence . Mansfield Centre, CT, Maurizio Martino . v. 3.) has failed to located any reference to this bird, but Sauer’s volumes only cover his life up to 1845. Future researchers might wish to study the Gould correspondence held in the UK National Archives (https://discovery.nationalarchives.gov.uk/details/r/55f98d6c-8dac-4163-bafe-70414ef1c494) for information pertaining to this bird. The specimens from the Salvin-Godman collection came via John Youds, another person about whom very little is known, especially as the family (originally from Cheshire, in north-west England, but emigrated to Bahia, Brazil, around the 1820s) numbered multiple members named John, at least one of whom was involved in the slave trade. Ihering (1900Ihering, H. von. 1900. Aves observadas em Cantagalo e Nova Friburgo. Revista do Museu Paulista, 4: 149-164.) reported many species/specimens from the vicinity of Nova Friburgo attributable to Youds, but gave (or knew) no further details about him. Given the complexities, it may prove impossible to elucidate (even to the decade) when this specimen was collected, especially as Youds presumably was responsible only for trading it. Fry lived in Brazil between 1838 and 1854, and it seems most likely that his specimens were collected during this 16-year period, but he did continue to visit the country in the years thereafter (see Table 2, note 28).

Even Snow’s (2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.) suggestion that the species’ last known specimen was collected around 1890 appears to be based more on assumption than definite knowledge (and his basis is unfortunately unknown). Indeed, it is unclear whether any of the 100+ specimens was taken later than the 1860s. One of the last that can be pinpointed with any certainty is NRM 90127687, which was acquired by Gunnar Olof Hyltén-Cavallius sometime between 1860 and 1864 during his tenure in the Swedish consulate in Rio de Janeiro (see Table 2, footnote 37).

The specimens from Florence (MZUF) and Kiel (ZMK), cited by Hume & Walters (2012Hume, J.P. & Walters, M. 2012. Extinct birds. London, T. & A.D. Poyser.), could not be re-located. After contact by e-mail with Fausto Barbagli (University of Firenze), it proved impossible to confirm the presence of a specimen of the species in MZUF, only that the collection is currently stored in a warehouse due to renovation (Fausto Barbagli in litt., 21 October 2019). Neither could we confirm the existence of a specimen in Kiel; according to the curator the species is not represented in the collection (Malte Seehausen in litt., 20 March 2023).

Plumage variation

Although a much larger number of specimens preserved in museums is now known, the state of conservation of these specimens varies greatly, reflecting the conditions they have been exposed to over the last 150-200 years.

In the 19^th century it was common to publicly display birds in cases, with the result that many specimens have faded plumage, whereas others were quickly incorporated into scientific collections, remaining better protected inside cabinets unexposed to the light, but still others remain on display (e.g., LUOMUS 1573, MAB, ZISP 1819), despite the species being considered Critically Endangered and thus of special significance and very rare in natural history museums. Changes to original colours were caused not only by exposure to light, as Jon Fjeldså (in litt., 21 August 2019) at ZMUC, said: “The mounted specimen therefore is faded and stained grey by carbon particles, as the old exhibition rooms were heated by burning coal.”

To know a little more about the plumage of a recently prepared specimen (or a live bird), we consulted the original description of the species (Vieillot, 1818Vieillot, L.P. 1818. Oiseaux. In: Nouveau dictionnaire d’histoire naturelle. Nouvelle édition. Paris, Deterville. v. 24.; pp. 528-529): “Le Pardalote huppé, Pardalotus cristatus, se trouve au Brésil, d’où il a été apporté par M. Delalande fils. La huppe qui orne sa tête est rouge et près l’occiput, comme dans le roitelei rubis; la gorge et toutes les parties inférieures sont d’un beau jaune, plus foncé sur le devant du cou et sur la poitrine; les pieds noirs; le bec est de cette couleur à sa base et à sa pointe, et couleur de corne sur le milieu et en dessous; la tête, le dessus du cou et du corps d’ un vert olive tirant au jaune; les plumes du milieu de la tête, du front et de l’ occiput, terminées de brun-noir; les petites couveriures des ailes moitié blanches à l’extérieur; les pennes brunes et bordées de vert-olive en dehors; celles de la queue du même vert et strê-courtes; taille à peu prés pareille à celle du pardalote pointillé”.

Our translation: “The Kinglet Calyptura, Pardalotus cristatus, is found in Brazil, from where it was brought by the younger Mm. Delalande. The crest which adorns its head is red and close to the rear head, as in the roitelei rubis; the throat and all the lower parts are of a beautiful yellow, darker on the foreneck and on the breast; the feet are black; the bill is of this colour at its base and tip, and horn-coloured in the middle and below; the head, nape and upperparts are olive-green to yellow; the feathers on the middle of the head, forehead and rear head are brownish-black; the small wing coverts are half white on the outside; the remiges are brown and edged with olive-green on the outside; those on the tail are of the same green and very short; the size of the bird is about the same as that of pardalote pointillé”. Roitelei rubis is a reference to the bird we now know as Ruby-crowned Kinglet Corthylio calendula (Linnaeus, 1766), whilst pardalote pointillé is the Spotted Pardalote Pardalotus punctatus (Shaw, 1792).

The holotype of Calyptura cristata (MNHN ZO-MO-2004-300) used in Vieillot’s description was collected sometime in 1816 in Rio de Janeiro by Pierre Antoine Delalande, and is preserved in the Muséum National d’Histoire Naturelle, Paris, France (Fig. 1). It is possible to note clearly the effects of time, more than 200 years, on this specimen, which has faded coloration, mainly its yellow crest, as opposed to the red described by Vieillot.

A feature that draws attention in the Kinglet Calyptura specimens (checked personally or via photographs) is the great variation in the colour of the crest, as well as the area or number of feathers with this coloration. The crest colour ranges from bright red (e.g., MHH 5665; MNHN ZO-MO-2000-2153; ZMB 2000/12103) through orange (e.g., LUOMUS 4949; MNRJ 3137; RMNH.AVES.172608; ZMB 2306) to yellow (e.g., AMNH 5156; MSNTP AV3082; NMBE 1033790; NRM 535288). Given that red was the original colour in the crest of the holotype, according to Vieillot’s description, we might assume that other colours (orange and yellow) are only the result of discoloration over the intervening years, in line with research into the effects of time on the colour of museum specimens, perhaps especially carotenoid-based colours (such as reds and yellows) (McNett & Marchetti, 2005McNett, G.D. & Marchetti, K. 2005. Ultraviolet degradation in carotenoid patches: live versus museum specimens of wood warblers (Parulidae). Auk , 122(4): 793-802. https://doi.org/10.1093/auk/122.3.793.
https://doi.org/https://doi.org/10.1093/... ; Armenta et al., 2008Armenta, J.K.; Dunn, P.O. & Whittingham, L.A. 2008. Effects of specimen age on plumage color. Auk, 125(4): 803-808. https://doi.org/10.1525/auk.2008.07006.
https://doi.org/https://doi.org/10.1525/... ; Riedler et al., 2014Riedler, R.; Pesme, C.; Druzik, J.; Gleeson, M. & Pearlstein, E. 2014. A review of color-producing mechanisms in feathers and their influence on preventive conservation strategies. Journal of the American Institute for Conservation, 53(1): 44-65. https://doi.org/10.1179/1945233013Y.0000000020.
https://doi.org/https://doi.org/10.1179/... ). However, this cannot be used to explain why a large number of specimens, all in excess of 100 years old, exhibit such variation in the colour of the coronal patch, especially as some specimens that are known to have been exposed to direct natural light for considerable periods of time, e.g., that in IZH, still have a bright red crown. However, we lack details of how every specimen has been stored throughout the last two centuries, and it is plausible that specimen storage and individual/sex-related differences could be among the factors to explain the variation that is currently observed.

The distinction between the sexes in C. cristata is based mainly on the yellow coloration in the forehead and forecrown in males and dark olive-green in the females, and the volume of red feathers in the crest, with males (e.g., MNHN ZO-MO-1854-440) having more red feathers, and bolder black lateral borders (Snow, 2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.; Kirwan & Green, 2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .). Examination of a larger number of specimens allowed us to pinpoint other plumage characteristics that might help to identify males and females, and distinguish between adults and immatures. We noticed that in males (e.g., NHMO 66671; NMB 2041; NHMUK 1895.4.1.730) the black band starts just in front of the eye and continues to border the red crest on the nape, whereas in females (e.g., NHMUK 1895.4.1.731; NMB 2042; ZISP 117202; ZMB 2306) there is only a small yellow spot behind and below the eye, and the black borders to the red/orange/yellow coronal patch are much less well defined.

Juveniles/immatures might be those individuals with few, or only a very small number of red feathers in the crest (e.g., RMNH AVES.172608), and which also lack yellow in the forehead and forecrown, perhaps with immature males possessing more yellow and red feathers compared to immature females. Analysing tail feather shape, which can be useful for separating adults and juveniles of passerines (being narrower with more pointed tips in the latter), is not always easy in very old museum specimens, rather than live birds, especially when attempting to base the identification solely on photos. Feathers can be lost, become frayed or otherwise changed in shape, however, RMNH.AVES.172608 does appear to have narrower rectrices. Among other specimens that share the full suite of these immature characters are ZMB 2306, ZMB 2000, 12104, ZMUC 105507, and NMBE 1033790.

DISCUSSION

Our inventory almost doubles previous estimates of the number of Calyptura cristata specimens held in the world’s natural history museums. We have identified at least 104 specimens, the majority of which are held in European collections. Given that several specimens were found in relatively small institutions, there is still the potential for additional material to come to light in collections subject to less intensive and detailed curation work.

To try and contextualise just how remarkably large this total is, one useful comparison to make is with another species endemic to a tiny area (estimated at 410 km²) of southeast Brazil, in the lowlands of Rio de Janeiro state, and which was also “lost” for more than a century, Black-hooded Antwren Formicivora erythronotos.Pacheco (1988Pacheco, F. 1988. Black-hooded Antwren Formicivora [Myrmotherula] erythronotos re-discovered in Brazil. Bulletin of the British Ornithologists’ Club , 108(4): 179-182.) related the species’ dramatic rediscovery in the environs of Angra dos Reis in September 1987, slightly less than a decade prior to the multi-observer record of Calyptura cristata near Teresópolis in October 1996. Unlike the latter, however, the Formicivora continues to be seen, some additional (albeit nearby) localities have been identified for it, and there are now literally hundreds of photographs of the species on citizen science databases, attesting to its local abundance. Specimens in museums are, however, exceptionally few in number: Pacheco (1988), repeated by Collar et al. (1992Collar, N.J.; Gonzaga, L.P.; Krabbe, N.; Madroño Nieto, A.; Naranjo, L.G.; Parker, T.A. & Wege, D.C. 1992. Threatened birds of the Americas: the ICBP/IUCN Red Data book. Cambridge, International Council for Bird Preservation.) postulated there are “about 20 nineteenth century skins in European and American museums”, but in fact we have found evidence of just ten or 11: one or two of them lost, five extant in Europe, one in the USA, and three in Brazil (MNRJ 33300); see below, and a pair held at the Museu de Zoologia da Universidade de São Paulo, MZUSP 76678 and 76679; (L.F. Silveira in litt., 2023). Hartlaub’s holotype, a male, is in the Zoologisches Institut und Zoologisches Museum, Hamburg (Bolau, 1898Bolau, H. 1898. Die Typen der Vogelsammlung des Naturhistorischen Museums zu Hamburg. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 15: 47-71.); another male from Burmeister is held in IZH (Pacheco, 1988; GMK pers. obs.), an adult male reported to have been collected in February 1879 (?) is in the Naturmuseum Senckenberg, Frankfurt am Main (SMF 24044), and there are two specimens (one male, one female) in NHMUK (Knox & Walters, 1994Knox, A.G. & Walters, M.P. 1994. Extinct and endangered birds in the collections of The Natural History Museum. British Ornithologists’ Club Occasional Publications , 1: 1-292.; H. van Grouw in litt., 2023). Another specimen, reported to have been either a female or immature male, was presumably formerly in IZH (e.g., Burmeister, 1856), but seems to be no longer present there. The specimen in SMF has apparently never been specifically referred to in the literature before; it was purchased from Gustav Schneider (1834-1900), the Basel-based taxidermist. At one time or another, the museum in Tring had five specimens (Sclater, 1890Sclater, P.L. 1890. Catalogue of the birds in the British Museum . London, Trustees of the British Museum . v. 15.; Knox & Walters, 1994Knox, A.G. & Walters, M.P. 1994. Extinct and endangered birds in the collections of The Natural History Museum. British Ornithologists’ Club Occasional Publications , 1: 1-292.), the others being the male now in MNRJ (exchanged in 1984), another male, from Gould, that was one of the four listed by Sclater (1890Sclater, P.L. 1890. Catalogue of the birds in the British Museum . London, Trustees of the British Museum . v. 15.) but is now lost, and a third male, originally in the Sclater collection, that was exchanged with AMNH in 1921. The female still at NHMUK was also originally in Sclater’s private collection and came to him via a dealer named Warwick (it thus seems unlikely to be the same specimen once at IZH). Searches of VertNet and GBIF, as well as databases of several of the large North American museums (AMNH, FMNH, MCZ, NMNH), have revealed no additional material.

The most detailed prior analysis of plumage variation in Calyptura cristata is that by Kirwan & Green (2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .). This was based on the earlier literature and analysis of specimens by GMK in eight museums (IZH, MNHN, MNRJ, NHMUK, RMNH, UMZC, USNM and ZISP). These authors suggested that females differ from males by having the forehead and forecrown largely dark olive-green (rather than yellow), with paler orange-red restricted to the rear crown and any black not forming well-defined lateral crown-stripes, narrower white tips to the wing-coverts and tertials, and a paler yellow rump patch. Furthermore, in common with other authors (e.g.,Snow, 2004Snow, D.W. 2004. Family Cotingidae (cotingas). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 32-108.), they noted that any non-adult plumages are undescribed, but that ZMUC 105507 (photos of which were analysed as part of the present study) said to be an immature male was illustrated (by J. Fjeldså inKrabbe, 2007Krabbe, N. 2007. Birds collected by P.W. Lund and J.T. Reinhardt in south-eastern Brazil between 1825 and 1855, with notes on P.W. Lund’s travels in Rio de Janeiro. Revista Brasileira de Ornitologia, 15(3): 331-357.) as having a pale throat and a greenish cast to the flanks (Kirwan & Green, 2011Kirwan, G.M. & Green, G. 2011. Cotingas and manakins. London, Christopher Helm .). This specimen does indeed appear to have the narrower and slightly more pointed rectrices that might be expected to characterise a younger bird.

Sex and age-related plumage variation in Kinglet Calyptura’s closest relatives are not always well known. In Cinnamon Neopipo Neopipo cinnamomea, the sexes are similar, but the female’s yellow coronal patch is smaller, and there is no published information concerning juvenile plumage (Farnsworth & Lebbin, 2004Farnsworth, A. & Lebbin, D.J. 2004. Cinnamon Tyrant Neopipo cinnamomea. In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world , Barcelona, Lynx Edicions . v. 9, p. 352-353.). Among the seven species of spadebills (genus Platyrinchus), the sexes are separable based on the colour and pattern of the coronal patch in three, Cinnamon-crested Spadebill P. saturatus, Golden-crowned Spadebill P. coronatus and White-crested Spadebill P. platyrhynchos (Hilty, 2003Hilty, S.L. 2003. Birds of Venezuela. Princeton, NJ, Princeton University Press.; Johnson & Wolfe, 2018Johnson, E.I. & Wolfe, J.D. 2018. Molt in Neotropical birds. Life history and aging criteria. Boca Raton, Taylor & Francis. (Studies in Avian Biology Vol. 51). https://doi.org/10.4324/9781315119755.
https://doi.org/https://doi.org/10.4324/... ), whilst less detailed but similar differences have been reported in coronal patch size and colour (smaller and often paler in females) of two other species, Stub-tailed Spadebill P. cancrominus and White-throated Spadebill P. mystaceus (Tello, 2004Tello, J.G. 2004. Genus Platyrinchus (spadebills). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world . Barcelona, Lynx Edicions . V. 9. p. 340-343.). In contrast, the sexes are reported to be alike in Yellow-throated Spadebill P. flavigularis and Russet-winged Spadebill P. leucoryphus (Tello, 2004Tello, J.G. 2004. Genus Platyrinchus (spadebills). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world . Barcelona, Lynx Edicions . V. 9. p. 340-343.), although this perhaps requires re-evaluation in light of the details presented by Johnson & Wolfe (2018Johnson, E.I. & Wolfe, J.D. 2018. Molt in Neotropical birds. Life history and aging criteria. Boca Raton, Taylor & Francis. (Studies in Avian Biology Vol. 51). https://doi.org/10.4324/9781315119755.
https://doi.org/https://doi.org/10.4324/... ). Among Platyrinchus, the juveniles of Yellow-throated and Russet-winged Spadebills are both undescribed, whilst some of the other species are reported to differ principally in that they lack any evidence of a coronal patch, with reduced facial markings (usually well marked in adults) and generally paler and duller underparts, but at least in the hand ageing is probably more reliably based on relatively pointed versus dull and rounded rectrices, molt limits in the wing-coverts, and subtle differences in the colour of some feathers of the wing (Tello, 2004Tello, J.G. 2004. Genus Platyrinchus (spadebills). In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world . Barcelona, Lynx Edicions . V. 9. p. 340-343.; Johnson & Wolfe, 2018Johnson, E.I. & Wolfe, J.D. 2018. Molt in Neotropical birds. Life history and aging criteria. Boca Raton, Taylor & Francis. (Studies in Avian Biology Vol. 51). https://doi.org/10.4324/9781315119755.
https://doi.org/https://doi.org/10.4324/... ). It is also interesting to compare another, slightly more distantly related tyrannid (based on the Ohlson et al., 2012Ohlson, J.I.; Irestedt, M.; Fjeldså, J. & Ericson, P.G.P. 2012. Nuclear DNA from a 180-year-old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides). Ibis, 154(3): 533-541. https://doi.org/10.1111/j.1474-919X.2012.01243.x.
https://doi.org/https://doi.org/10.1111/... phylogeny), Many-coloured Rush Tyrant Tachuris rubrigastra, in which females are characterised by generally duller colours and a smaller coronal patch, and juveniles are distinguished by their lack of wingbars, duller underparts, some yellow scaling on the green upperparts and a lack of blue in the facial mask (Clock, 2004Clock, B.M. 2004. Many-coloured Rush-tyrant Tachuris rubrigastra. In: del Hoyo, J.; Elliott, A. & Christie, D.A. (Eds.). Handbook of the birds of the world. Barcelona, Lynx Edicions. v. 9. p. 315.).

We yet again confirm the vital importance of scientific collections in museums around the world, not only the large and famous, but also smaller and little-known institutions that house much smaller amounts of material. With a considerably larger number of specimens preserved in museums, C. cristata was evidently an even commoner species than was previously imagined, despite its seemingly restricted distribution, thereby making the reasons for its apparently imperilled conservation situation even more mysterious.

Acknowledgments

Pepijn Kamminga and Steven van der Mije (RMNH, now Naturalis Biodiversity Center), and the late Vladimir Loskot (ZISP) are thanked for their hospitality and courtesy to RSS and GMK during our separate visits to these institutions. Julian Hume sent information concerning the specimens reported (perhaps erroneously) to be in Florence and Kiel. The following curators provided data concerning specimens of Calyptura cristata in the collections under their care (GMK has personally examined relevant material at those institutions marked with an asterisk): Paul Sweet and Jason Sheldon (AMNH), Nate Rice (ANSP), Cédric Audibert (CCECL), Vanya Gregor Rohwer (CUMV), Frank Steinheimer (IZH*), Christina Barilaro (LMNM), Alexandre Aleixo (LUOMUS), Pierre-Henri Béguin (MAB), Jeremiah Trimble (MCZ), Bernd Nicolai and Rüdiger Becker (MHH), Ricardo Jorge Lopes and Mariana Costa (MHNC-UP); Philippe Candegabe (MHNGr), Aude Medina (MHNMON), Celia Bueno and Martin Zimmerli (MHNN), Christophe Remy (MHNVT), Henry McGhie, Rachel Petts and David Gelsthorpe (MM*), Patrick Boussès (MNHN*), Marco Aurélio Crozariol and Claydson Assis (MNRJ*), Jan Lontkowski (MPUW), Paul Tout and Nicola Bressi (MSNT), Simone Farina and Lorenzo Vanni (MSNTP), Fritz Geller-Grimm (MWNH), Maria Mostadius (MZLU), Marie Meister (MZS), Luca Ghiraldi (MZUT), Lars Erik Johannessen (NHMO), Hein van Grouw (NHMUK*), Christiane Schilling (NLMH); David Marques, Dieter Thomas Tietze and Urs Wüest (NMB), John-James Wilson (NML-VZ), Lorenzo Vinciguerra (NMSG), Hans-Martin Berg (NMW), Ulf Johansson (NRM), Olivier Pauwels and Annelise Folie (RBINS), Gerald Mayr (SMF), Hjalmar S. Kühl and Diana Jeschke (SMNG), Friederike Woog (SMNS*), Janet Hinshaw and Brett W. Benz (UMMZ), Michael del Brooke, Daniel Field and Matthew Lowe (UMZC*), Brian K. Schmidt (USNM*), Pascal Eckhoff and Sylke Frahnert (ZMB*), Malte Seehausen (ZMK), Pavel Smirnov (ZMMU), Pete Hosner and Jon Fjeldså (ZMUC) and Marie Bournonville (ZMUL). Julien Mazenauer identified the specimen in the Musée de l’Areuse Boudry, Neuchâtel, Switzerland. Nigel Collar photographed one of the specimens at the UMZC on our behalf. The following confirmed the lack of specimens of Kinglet Calyptura at their institutions: Jakob Pöhacker (Haus der Natur, Museum für Natur und Technik, Salzburg, Austria), Peter Sackl (Universalmuseum Joanneum, Graz, Austria), Stephan Weigl (Biologiezentrum des Oberösterreichisches Landesmuseums, Linz, Austria), Alain Reygel (Royal Museum of Central Africa, Tervuren, Belgium), Luís Fábio Silveira (Museu de Zoologia da Universidade de São Paulo, Brazil); Maria de Fátima Cunha Lima (Museu Paraense Emílio Goeldi, Belém, Brazil), Zlatozar Boev (National Museum of Natural History, Bulgarian Academy of Sciences, Sofia, Bulgaria), Jan Hušek and Jiří Mlíkovský (Museum of Natural History, Prague, Czech Republic), Miroslav Sebela and Martin Cerny (Moravské Zemské Muzeum, Brno, Czech Republic), Margus Ots (University of Tartu Natural History Museum, Estonia), Mathilde Schneider (Muséum d’Histoire Naturelle, Rouen, Normandy, France), Muriel Lecouvez (Musée d’Histoire Naturelle, Lille, France), Nathalie Mémoire (Musée d’Histoire Naturelle de Bordeaux, France), Andreas Bick (Zoologischen Sammlung der Universität Rostock, Germany), Cordula Bracker (Zoologisches Institut und Zoologisches Museum, Hamburg, Germany), Dietrich von Knorre (Jena Phyletisches Museum, Germany), Albrecht Manegold (Staatliches Museum für Naturkunde, Karlsruhe, Germany), Matthias Mäuser (Naturkunde-Museum Bamberg, Germany), Carsten Ritzau (Naturkunde-Museum Coburg, Germany), Christiane Schilling (Niedersächsisches Landesmuseum, Hannover, Germany), Melita Vamberger and Martin Päckert (Staatliches Museum für Tierkunde, Dresden, Germany), Till Töpfer (Leibniz Institute for the Analysis of Biodiversity Change, Museum Koenig, Bonn, Germany), Markus Unsöld (Zoologische Staatssammlung München, Germany), Erich Weber (Institute für Evolution und Ökologie, Vergleichende Zoologie, Eberhard-Karls-Universität Tübingen, Germany), Tibor Fuisz (Magyar Nemzeti Múzeum, Budapest, Hungary), Andrea Benocci (Museo Zoologico dell’Accademia dei Fisiocritici, Siena, Italy), Carla Marangoni (Museo Civico de Zoologia, Rome, Italy), Stefano Maretti (Museo di Storia Naturale Università di Pavia, Italy), Alain Frantz and Francesco Vitali (Musée National d’Histoire Naturelle, Luxembourg), Terje Lislevand (Zoologisk Museum, University of Bergen, Norway), Tomasz Huflejt and Dominika Mierzwa-Szymkowiak (Museum and Institute of Zoology, Warsaw, Poland), Josefina Barreiro Rodríguez (Museo Nacional de Ciencias Naturales, Madrid, Spain), Alice Cibois (Museum d’Histoire Naturelle de Genève, Switzerland), Sarah Kenyon (Saffron Walden Museum, UK), Zena Timmons (National Museums of Scotland, Edinburgh, UK), and Stephen P. Rogers (Carnegie Museum of Natural History, Pittsburgh, USA). Dione Seripierri and Martha Zamana in the Museu de Zoologia da Universidade de São Paulo library enabled access to various references. Vitor Piacentini provided additional photographs of Calyptura specimens held at AMNH, MNHN and ZMB. We are grateful to two referees for their comments on the submitted version of the manuscript.

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