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British Ornithologists’ Club

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Volume 134 No. 1 March 2014

FORTHCOMING MEETINGS

See also BOC website: http://www.boc-online.org BOC MEETINGS are open to all, not just BOC members, and are free.

Evening meetings are in an upstairs room at The Barley Mow, 104 Horseferry Road, Westminster, London SW1P 2EE. The nearest Tube stations are Victoria and St James's Park; and the 507 bus, which runs from Victoria to Waterloo, stops nearby. For maps, see http://www.markettaverns.co.uk/the_barley_mow.html or ask the Chairman for directions.

The cash bar opens at 6.00 pm and those who wish to eat after the meeting can place an order. The talk will start at 6.30 pm and, with questions, will last c.l hour.

It would be very helpful if those intending to come can notify the Chairman no later than the day before the meeting.

25 February 2014 — 6.30 pm — Richard Porter— Birds of Socotra: populations and distribution

Abstract : The Socotra archipelago lies in the Arabian Sea, c.350 km south of the Yemen mainland. Ecologically heavily influenced by the dry south-west monsoon that batters the islands in May-September, it boasts high endemism in plants, reptiles, insects and birds, which I will summarise. Whilst its avifauna is species-poor, with just 42 regular breeders, 1 1 are endemic, making it the richest area of avian endemism in the Middle East (along with the highlands of south-west Arabia). From 1999 to 2011, I have been engaged in mapping the distribution and determining the populations of the breeding species and I will present some of my findings. I will also talk briefly about the migrants that visit the islands and recent taxonomic studies that have added Socotra Buzzard Buteo socotraensis, Abd A1 Kuri Sparrow Passer hemileucus and Socotra Golden¬ winged Grosbeak Rhynchostruthus socotranus to the species list. My talk will end with my thoughts on future research and conservation.

Biography: Richard Porter has had a continuing involvement in bird research and conservation in the Middle East since 1966, when he spent the autumn studying soaring bird migration over the Bosphorus. Whilst his early exploits were largely in Turkey, since 1979 most of his visits have been to Yemen, particularly Socotra, and Iraq, where he is the bird and conservation adviser to Nature Iraq. Richard, who is author of Birds of the Middle East, advises BirdLife International on their Middle East programme.

20 May 2014 — 5.30 pm — Annual General Meeting, followed at 6.30 pm by Guy Kirwan — Cuban birds at home and abroad, in the field and museum

Abstract: This talk will provide an introduction to the birds of the largest Caribbean island, Cuba, which despite boasting the most speciose avifauna in the West Indies supports fewer endemics than either Hispaniola or tiny Jamaica. Nevertheless, depending on taxonomy, at least seven avian genera occur only on Cuba, as well as the world's smallest bird. Bee Hummingbird Mellisuga helenae, while the country might yet prove to be the last bastion of one of the planet's most iconic birds, Ivory-billed Woodpecker Campephilus principalis. Despite >150 years of ornithological exploration, our knowledge of Cuban birds is still advancing comparatively rapidly, especially with respect to their ecology and conservation, largely via the efforts of a few dedicated researchers, both Cuban and foreign. My talk will focus on the taxonomy, ecology and conservation of some of the most special of Cuba's birds, as well as providing an introduction to travel in what was very briefly part of the British Empire!

Biography: Guy Kirwan has been a regular visitor to the Greater Antilles since the mid-1990s and is a co-author of a forthcoming checklist to Cuban birds. He is a freelance ornithologist and editor, notably of Bull. Brit. Orn. CL, with strong interests in avian taxonomy and the breeding biology of birds in the New World tropics. A Research Associate at the Field Museum of Natural History in Chicago, he recently joined the BOU's Taxonomic Subcommittee and currently works for Lynx Edicions on the HBW Alive project.

23 September 2014 — 6.30 pm — Dr Andrew Gosler—Ethno-ornitIwlogy

22 November 2014 — Joint meeting with the Oriental Bird Club and the Natural History Museum

A one-day meeting in the Flett Theatre, Natural History Museum, South Kensington, London SW7 5BD starting at 10.30 am. Details to be announced

The Chairman: Chris Storey, 22 Richmond Park Road, London SW14 8JT UK. Tel. +44 (0)208 8764728. E-mail: c.storeyl@btinternet.com

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Bull. B.O.C. 2014 134(1)

Bulletin of the

BRITISH ORNITHOLOGISTS’ CLUB

Vol. 134 No. 1 Published 8 March 2014

CLUB ANNOUNCEMENTS

Chairman's message

Since 2003 and the publication of the The birds of Morocco, the BOC and the BOU have jointly published the Checklist series and other books (e.g. The bird atlas of Uganda). However, following a thorough review of its activities, the BOU has decided to withdraw from the publication of such titles with immediate effect. All BOU and joint BOU/BOC titles may of course still be purchased via the BOU website: www.bou.org.uk or from the BOU office, P.O. Box 417, Peterborough PE7 3FX, UK.

The BOC Committee remains of the view that Checklists fit naturally with the Club's aims and objectives, and make an important contribution to ornithological studies. Consequently, it is committed in principle to their continued publication and is now considering in detail the practicalities of undertaking this task as sole publisher, including discussion with the BOU and Dr David R. Wells, Checklist Commissioning Editor. The aim is to seek to resolve the many outstanding issues and to establish our intensions in time for the AGM in May. In the meantime any views and or comments would be gratefully received by David Wells or myself.

Chris Storey

Annual General Meeting

The Annual General Meeting of the British Ornithologists' Club will be held in the upstairs room at The Barley Mow, Horseferry Road, Westminster, London SW1P 2EE, at 5.30 pm on Tuesday 20 May 2014.

Agenda

1. Apologies for absence

2. Minutes of the Annual General Meeting held on 21 May 2013 (see Bull. Brit. Orn. Cl. 133: 165-166, and the BOC website).

3. Receive and consider the Chairman's Review, the Trustees' Report and the Accounts for 2013 (these will be available in the room before the start of the meeting).

4. The Bulletin Editor's report— Mr G. M. Kirwan.

5. Election of Officers. The Committee proposes that:

i. Dr R. P. Prys-Jones be re-elected as Hon. Secretary

ii. Mr D. J. Montier be re-elected as Hon. Treasurer

No other changes to the Committee are proposed as all other members are eligible to serve at least one more year in office.

6. Any other business, of which advance notice has been given.

Gerlof Fokko Mees (1926-2013)

Gerlof Mees, former bird curator at the Rijksmuseum van Natuurlijke Historie (RMNH) at Leiden, Netherlands, was born into a well-off family in Bloemendaal, Netherlands. Both his parents had a keen interest in natural history, and Gerlof was stimulated to study birds by them and by his uncle Jan Verwey, founder of Dutch research into bird migration and ethology. After completing secondary school, he started to study biology in Leiden, but this was soon interrupted by a stay on Java (1946-49) as part of the Dutch forces in the Indonesian independence war. His scarce spare time was spent birdwatching, and problems identifying the local white-eyes Zosterops brought him into contact with George Junge, tbien bird curator at RMNH. Junge permitted Mees to study the white-eyes in his care, and Gerlof also learned the practical work of a museum department. Though he specialised in fish research (thought to be more likely to produce employment than birds), Gerlof visited Trinidad & Tobago when still a student and collected birds during an eight-month stay in 1953/54, reporting on these together with his mentor (Junge & Mees 1958). Between May 1955 and July 1957 he was an assistant at RMNH, working in both the bird and fish departments. He obtained his doctorate in 1956. His study on white-eyes resulted in a thesis on the Indo-Australian Zosteropidae (Mees 1957). However, fishes still held his interest and Mees was appointed a curator at the Western Australian Museum in Perth in 1958 mainly for his fame as an ichthyologist. There he continued

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to work on Zosteropidae, but (with fishes) also on Australian larks and owls, and on the birds of the West Pilbara and Kimberley areas. He described several new taxa for Western Australia and, moreover, met his future wife, Veronica. After Junge's death in 1962 Gerlof was appointed bird curator at RMNH and the family moved to Leiden in 1963.

The rich Indonesian collections at RMNH now formed the main subject of Mees' work, resulting in very thorough publications on the birds of Bangka, the Moluccas, Misool, southern New Guinea and Java. Moreover, he made collecting trips to Nigeria, Guyana, Zimbabwe and (especially) Suriname, often with Veronica, while he also oversaw the acquisition of large bird collections for RMNH from Turkey, Mexico, Taiwan, Kenya, Tanzania and Flores, resulting in further major papers and several revisions and shorter notes. His distributional notes or reviews of Andaman sparrowhawks, Chinese Crested Tern Sterna bernsteini, Large-tailed Nightjar Caprimulgus macrurus, Common Cuckoo Cuculus canorus and Whiskered Tern Chlidonias hybrida are meticulously detailed. His publication list numbers c.100 articles on birds, but Iris list of papers on fishes is also long.

Mees' luxury work room at RMNH, surrounded by shelves with 3,000 bird books and complete series of >200 bird journals with a central desk covered by long rows of bird skins, jars with fishes, and piles of reprints and manuscripts made a lasting impression on most visitors, especially as Gerlof often seemed disturbed when one dared to enter his room. However, visitors with serious interests were always pleasantly provided with books and articles, and given free access to the collections, though they perhaps would still feel themselves rather uneasy because Gerlof often would debate the failure of the political system in the Netherlands at the same time. It was wonderful to see Mees find his way among the 50,000 mounted birds and 200,000 skins and clutches to pick out a special item, especially as large parts of the collections were not yet databased. Within the museum staff he was admired because of his deep knowledge on the history of the collections and of nomenclatorial questions, but some people felt ambiguous towards him. For instance, though fluent in at least four languages, as an employee of a Dutch museum Mees often insisted in writing in his native language when publishing in RMNH journals, using a rather old-fashioned style. As a result, editors of journals, being some of his fellow curators, had to translate his lengthy manuscripts into English, during which small errors frequently crept in. Mees tended to blame his colleagues for this, although his own proof-reading could have prevented the mistakes. Also, he still worked periodically on fishes, even though these had a curator of their own.

Mees retired in 1991 to Western Australia with Veronica, where their children had already settled. Until his death, he maintained a deep interest in ornithology, and despite living in the outback he published several papers and notes, in part with help of friends in Leiden who sent him literature. The change from the research institute RMNH into the public museum with research facilities Naturalis Biodiversity Center and its move within Leiden was followed by him with much interest. In his last years, failing eyesight largely hindered further research, much to his regret. He died peacefully on 31 March 2013. He will be remembered for Monarcha sacerdotum Mees, 1973, from Flores, and in Caprimulgus meesi Sangster & Rozendaal, 2004.

Main publications of Gerlof F. Mees

1946. Courtship feeding of Willow-Warbler. Brit. Birds 39: 280.

1957. A systematic review of the Indo-Australian Zosteropidae (pt. 1). Ph.D. thesis. Univ. of Leiden.

1958. The avifauna of Trinidad and Tobago. Zoo/. Verhand. 37: 1-172 (with G. C. A. Junge).

1961. A systematic review of the Indo-Australian Zosteropidae (pt. II). Zodl. Verhand. 50: 1-168.

1961. An annotated catalogue of a collection of bird skins from West Pilbara, Western Australia. /. Roy. Soc. West. Austral. 44: 97-143.

1964. A revision of the Australian owls (Strigidae and Tytonidae). Zodl. Verhand. 65: 1-62.

1965. The avifauna of Misool. Nova Guinea, Zool. 31: 139-203.

1969. A systematic review of the Indo-Australian Zosteropidae (pt. III). Zoo/. Verhand. 102: 1-390.

1970. On some birds from southern Mexico. Zodl. Meded. 44: 237-245.

1970. Birds of the Inyanga National Park, Rhodesia. Zodl. Verhand. 109: 3-74.

1970. Notes on some birds from the island of Formosa (Taiwan). Zodl. Meded. 44: 287-304.

1971. Systematic and faunistic remarks on birds from Borneo and Java, with new records. Zodl. Meded. 45: 225-244.

1972. Die Vogel der Insel Gebe. Zodl. Meded. 46: 69-89.

1974. Additions to the fauna of Suriname. Zodl. Meded. 48: 55-67.

1975. Identiteit en status van Sterna bernsteini Schlegel. Ardea 63: 78-86.

1977. Geographical variation of Caprimulgus macrurus Horsfield (Aves, Caprimulgidae). Zodl. Verhand. 155: 1-47.

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1977. The subspecies of Chlidonias hybridus (Pallas), their breeding distribution and migration (Aves, Laridae, Sterninae). Zodl. Verhand. 107: 1-64.

1979. Die Nachweise von Cuculus canorus L. im Indo-Australischen Raum. Mitt. Zool. Mus. Berlin 55, Suppl. Ann. Orn. 3: 127-134.

1979. Verspreiding en getalssterkte van de Witwangstern, Chlidonias hybridus (Pallas), in Europa en Noord- Afrika. Zool. Bijdragen 26: 3-63.

1980. The sparrow-hawks ( Accipiter ) of the Andaman Islands. ]. Bombay Nat. Hist. Soc. 77: 371^112.

1982. Bird records from the Moluccas. Zool. Meded. 56: 91-111.

1982. Birds from the lowlands of southern New Guinea (Merauke and Koembe). Zool. Verhand. 191: 1-188. 1986. A list of the birds recorded from Bangka Island, Indonesia. Zool. Verhand. 232: 3-176.

1991. Bemerkungen uber Acrocephalus caffer (Sparrman) in der Tahiti-Gruppe (Aves, Sylviidae). Proc. K. Ned. Akad. Wet., Ser. C 94: 243-256.

1994. Tlie birds of Surinam. Second edn. Vaco, Paramaribo. [Thoroughly updated version of Haverschmidt's original work of 1968.]

1996. Geographical variation in birds of Java. Publ. Nuttall Orn. Cl. 26: 1-119.

2006. The avifauna of Flores. Zool. Meded. 80(3): 1-261.

C. S. (Kees) Roselaar

Naturalis Biodiversity Center, Netherlands, e-mail: cees.roselaar@naturalis.nl

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Some black-and-white facts about the Faeroese white-speckled Common Raven Corvus corax varins

by Hein van Grouw

Received 24 April 2013

Summary. — The white-speckled raven, a colour aberration of the Faroese Raven Corvus corax varius Briinnich, 1764, has occurred in the Faroe Islands since at least the Middle Ages. It has been described in many publications, and was a desired object for collectors of curiosities, especially in the 18th and 19th centuries. Early in the 20th century (1902) the last white-speckled individual was seen in the Faeroes, leaving only about two dozen specimens in museum collections. Although often referred to as albino, the aberration causing the white feathers is not albinism but leucism.

Common Raven Corvus corax Linnaeus, 1758, lias a circumpolar distribution in the Northern Hemisphere. In addition to nominate C. c. corax, ten subspecies have been described, including the slightly larger C. c. varius Briinnich, 1764, which is endemic to the Faeroes and Iceland. The white-speckled variant of C. c. varius was known only from the Faeroes. Salomonsen (1934a) discussed whether white-speckled birds might also have occurred in Iceland, as claimed by Olaus Magnus (1555), but doubted this for several reasons. It is unknown when the mutation first arose, but Ole Worm already had two specimens in his Museum Wormianum in Copenhagen, both collected before 1650 (Worm 1655).

By 1767 Ascanius had written that the white-speckled variant was not a separate species but a variety of Common Raven. However, in later descriptions both Vieillot (1817) and Wagler (1827) referred to it as a distinct species, giving rise to an ongoing debate concerning its taxonomic status.

Salomonsen (1934) described the presumed inheritance of white-speckled plumage based on historical records of the colours of pairs and their offspring. His study suggested a recessive inheritance for the black-and-white pattern, which is a form of leucism (van Grouw 2012, 2013). Recessive genes can be inherited invisibly via many generations before two birds, both carrying the recessive gene, mate, making the white-speckled gene visible in their offspring. Therefore, if this was indeed a recessive gene, it is difficult to be sure whether it has been lost, through time, since the last white-speckled bird was seen, or whether it might still occur in some individuals.

A remarkable bird, occurring in small numbers in a remote place, will inevitably be targeted by collectors. Both live birds and mounted specimens of the white-speckled raven were sent to museums and collectors of curiosities throughout Europe from the 17th century, although many specimens were subsequently lost or destroyed. Only 26 specimens remain in museums (van Grouw & Bloch in press).

Based on the numerous reports in the literature, one can acquire a fairly good idea of the population size of the Faroese raven through time. It is obvious that it was very common until c.1850. Svabo (1783) mentioned 'large flocks', Landt (1800) stated that it was a well-known 'bird of prey', Graba (1830) saw many in 1828, and Atkinson (1989) said the same in 1833. Both Holm (1848) and Muller (1862) regarded the raven as common but Feilden (1872) saw fewer than expected. Annandale (1905) observed hardly any and, although the population

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thereafter increased again, in 1930 Salomonsen (1931) still saw only small numbers, as did Ferdinand (1947) later. Bloch also observed few ravens in 1963, but in 1974 she noted a remarkable increase (Bloch et al. 2010). By 1981 at least 117 breeding pairs were counted on the Faeroes and the population was estimated at 150-350 pairs (Bloch & Sorensen 1984: 59). The recovery was mainly due to food supplies at waste sites of fish factories and incineration plants (Bloch 1981, Bloch & Sorensen 1984). Despite a legal requirement, introduced in 1988, for waste sites to be covered, limiting food for the ravens, the estimated number of breeding pairs in the Faeroes is currently c.500 (J.-K. Jensen pers. comm.).

The sharp decline in numbers on the Faeroes after 1850 was mainly due to the increase in the islands' human population, combined with the local beak tax still in force (Bloch 2012). The Faeroese National Archive contains lists from all six districts, covering most of the period 1742-1934, during which the annual number of ravens destroyed appears to have been c.150-250 (Bloch 2012). Around 1900 the population apparently reached its lowest ebb and the white-speckled variant became extinct.

The white-speckled mutation

Although the terms albino and partial albino are frequently used for the white-speckled ravens (Hartert & Kleinschmidt 1901, Sage 1962), the mutation causing the black-and-white pattern is a form of leucism. Leucism, from the Greek Leukos = white, can be defined as the partial or total lack of melanin in feathers (and skin) (van Grouw 2012, 2013). The lack of melanin is due to the congenital and heritable absence of pigment cells from some or all of those skin areas where they would normally provide the growing feather with colour. Depending on the type of leucism, the amount of white can vary from just a few feathers (= partial leucistic) to all-white individuals, which always possess colourless skin as well. Partially leucistic birds can have a normal-coloured bill and legs depending on where the colourless patches occur. However, leucistic birds always have pigmented eyes. To properly understand the nature of leucism it is necessary to know something of feather pigmentation first.

Melanins are the commonest pigments in birds and play a major role in the coloration of feathers, skin and eyes. They comprise two main types: eumelanin and phaeomelanin, which differ in colour, chemical composition and pigment granule structure (Fox & Vevers 1960). Eumelanin produces black, grey and brown feathers, and eye and skin colour, while phaeomelanin only occurs in feathers, and determines colours from deep reddish brown to pale buff. In all Corvus species only eumelanin is present, but for convenience it will simply be referred to as melanin hereafter.

Melanin is produced by specialised cells in the skin referred to as melanin cells or melanocytes, which develop from melanoblasts formed in the 'neural crest' — the embryonic spinal cord. Normally, melanoblasts migrate at an early embryonic stage to the mesodermal layers of the skin. Finally incorporated in the skin and feather follicles, melanoblasts develop into melanocytes to provide the feather cells with melanin (Crawford 1990).

The migration process is genetically determined and any (inheritable) change can affect the final distribution of the melanoblasts. If, due to a mutation, the melanoblasts are unable to migrate from the neural crest to the skin, there will be no melanocytes present to produce melanin, resulting in a completely un-pigmented (white) bird with pink skin. However, the eyes are not red (the crucial difference from albinos). The embryonic origin of eye pigments partially differs from that of the rest of the body; eye pigments are formed mainly from the outer layer of the optic cup (Lamoreux et al. 2010) and as leucism affects only the migration of melanoblasts originating from the neural tube it has no influence on eye pigmentation with an optic cup origin.

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In addition to being all white, leucistic birds can be partially white, with colourless feathers adjacent to normal ones. The pied appearance can be caused by a delay in the migration of the melanoblasts from the neural tube to the skin (Wagener 1959, Wendt- Wagener 1961). Because of the delay, some melanoblasts reach certain parts of the body where the skin is too far developed to incorporate them, resulting in these parts lacking colour. Another possibility is that, from the outset, insufficient melanoblasts develop in the neural crest and therefore not all parts of the body are provided with pigment cells (Daneel & Schumann 1961, 1963).

The white in leucistic birds is often patchy and bilaterally symmetrical due to the way melanoblasts migrate to the rest of the body, leaving certain areas without pigmentation. The white pattern is already present in juvenile plumage and the amount of white does not change with age. The commonest form of leucism affects pigment in body parts furthest from the neural crest: the face, the 'hand' of the wings, the feet and belly. This form caused the white-speckled plumage of Faroese ravens.

Although this form of leucism is commonest and occurs in many bird species (Fig. 4), leucism in general is uncommon. A far more widespread cause for the lack of pigment in feathers is progressive greying (van Grouw 2012, 2013) but this is not discussed further here.

Presumed inheritance and leucism in related species

The black-and-white pattern in ravens was uniform and symmetrical, following the pattern of the commonest form of leucism in birds; absence of pigment in the body parts furthest from the pigment cells' origin (Figs. 1-2). The total area of affected skin parts may differ, with smaller or larger white patches as a result, but generally the head and throat are white, as are the belly, primaries and primary-coverts, and claws. In extreme cases almost the entire wings are white, and the white throat extends to the belly breaking the black pectoral band.

Depending on where in the face pigment is lacking, the melanins in the eyes may be absent too, resulting in only those pigments that formed in the optic cup being visible. These appear to be much paler as, according to Graba (1830), who examined two freshly shot white-speckled ravens, the eyes were grey greenish-white (‘Iris graugriinlichweiss'). In other species, e.g. Rook Corvus frugilegus and Greylag Goose Anser anser, bluish- white eyes occur in leucistic individuals.

Different genes in birds result in this leucistic pattern. In Mallard Anas platyrhynchos it is due to a dominant gene, while in Japanese Quail Coturnix japonica it is recessive. It is partially dominant in Helmeted Guineafowl Numida meleagris : if the gene is present just once (heterozygous) the bird will possess the white pattern, while the presence of two genes (homozygous) produces an all-white bird. The same is true in Zebra Finch Poephila guttata (Fig. 3), although in this case another recessive gene is the cause.

Nothing is known with certainty concerning inheritance in genus Corvus. It has not been recorded in ravens except in the Faeroes (van Grouw 2012). Nor is it known to have occurred in Carrion Crow Corvus cor one, but is found very rarely in Hooded Crow C. cornix and Jackdaw C. monedula. In Rook, however, it is reasonably frequent, though in some cases only a small patch of white feathers is present on the chin, occasionally with 1-2 white claws. Given that Rooks lose most of their facial feathering on reaching adulthood, leucism is probably under-recorded. A juvenile leucistic Rook was described as a separate species by Sparrman (1786), who named it Corvus clericus (Latin for priest) on account of its white 'bib' (Fig. 8). There is, however, considerable variation in the white chin's size and, the larger it is, the more likely it is to be accompanied by white primaries and claws. More extreme

Figure 1. Drawing of a life-like white-speckled Common Raven Corvus corax in flight (Katrina van Grouw)

Figure 2. White-speckled Common Raven Corvus corax specimen in the Uberseemuseum Bremen, Germany (UMB 3800), collector and collection date unknown, but probably pre-1870 (Gabriele Wamke)

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cases follow the same pattern as in Common Raven (Fig. 5).

According to Salomonsen (1934) the inheritance of leucism in Common Raven was recessive, based on different crossings mentioned in the literature: two black parents may produce white-speckled offspring (Ascanius 1767, Graba 1830, Holm 1848, Muller 1862) and a white-speckled crossed with black may also yield white-speckled young (Graba 1830, Holm 1848, Muller 1862). In all cases white-speckled ravens hatched only when black parents were heterozygous (= carrying the allele for white- speckled once). Unfortunately, no records are available of crossings between two white- speckled birds. Based on these pairings, provided they are correct, straightforward recessive inheritance for white- speckled appears to be true. However, that this genetic variety was lost during the period that the variants were hunted to extinction, c.110 years ago, is in contrast to the recessive trait. Many black ravens in the Faeroes must have been heterozygous for leucism and the gene would be carried for many generations after the last white-speckled bird was seen. Sooner or later two heterozygous individuals would presumably have paired, producing white- speckled offspring. However, during the last century no pied ravens have been reported, but this might be explained by the severe decline in the population as a whole in the second half of the 19th century.

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Figure 3. Comparable leucism in different bird species: above, Bengalese finch Lonchura domestica ; bottom. Zebra Finch Poephila guttata (Pieter van den Hooven)

Figure 4. Comparable leucism in different bird species: Common Moorhen Gallinula chloropus, Durham, UK, January 2010 (Glen Roberts)

Figure 5. Leucistic Rooks Corvus frugilegus: left, juvenile. State Darwin Museum Moscow (SDM OF61) (Igor Fadeev); right, adult, formerly at Zoological Museum Amsterdam (ZMA 54604, now at NBC Naturalis) (Hein van Grouw)

As there are no reliable estimates of the proportions of white-speckled ravens and black ones, it is impossible to calculate the presumed gene frequency in a specified timeframe. However, with some assumptions we can acquire an impression of how common the allele for white-speckled could have been. Given that ravens were still common in the mid 1800s (Holm 1848, Muller 1862), we might assume that the pre-1850 population comprised c.120 breeding pairs (240 individuals). White-speckled individuals were not uncommon then (Graba 1830), so let us assume they comprised 10% of the population (24 birds). Of the 216 black birds a percentage were heterozygous. Knowing that the mutation had been present for centuries and that white-speckled individuals were breeding, it is plausible that 50% carried the allele for white-speckled (108 birds). Symbolising black as A and white-speckled as a, the number of genotypes in the population were: 108 AA, 108Aa and 24 aa, i.e. 45% did not carry the white-speckled gene (AA), 45% were carriers (Aa) and 10% were white- speckled (aa), thus the white-speckled gene was present in 55% of the raven population.

If we consider all white-speckled individuals eliminated, the a-allele in single form is still present in 50% of the population; that is, 108 pairs of black ravens (216 individuals; 108 AA and 108 Aa). In this scenario, four different crosses are possible and all equally probable (AA x AA = 100% AA; AA x Aa = 50% AA and 50% Aa; Aa x AA = 50% AA and 50% Aa; Aa x Aa = 25% AA, 50% Aa and 25% aa). Expressed as a percentage, the genotype of the offspring of these 108 pairs is: 56.25% black and lacking the gene for white-speckled (AA); 37.5% black and carrying the gene for white-speckled (Aa) and 6.25% white-speckled (aa). By removing the white-speckled individuals, a-gene frequency will be reduced, but if the population is sufficiently large the gene will still be present. However, the natural process of genetic drift can cause loss of certain alleles, especially in small populations.

Genetic drift is the random change in allele frequency in a population and is, along with selection, mutation and migration, a basic mechanism of evolution. These mechanisms cause changes in genotypes and phenotypes over time, and determine the degree of genetic variation within a population. In a textbook scenario, the alleles in the offspring form a representative sample of those in parents. However, in practice, certain alleles may be unrepresented in the next generation, especially if the number of offspring is low.

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C’rnu On mu

Figure 6. Diluted Common Raven Corvus cornx, Velbastadur, 2 March 2008 (Hans Eli Sivertsen)

Figure 7. Diluted Common Raven Corvus cornx specimen shot at Fugloy in 2008, prepared by J.-K. Jensen and donated to the Museum of Natural History, Torshavn (Jens-Kjeld Jensen)

Figure 8. Corvus clericus, described by Sparrman in 1786 in Museum Carlsonianum, was in fact a leucistic Rook C. frugilegus (Harry Taylor / © Natural History Museum)

TTiese alleles are therefore lost to subsequent generations.

The degree of loss of genetic variation (dF) is inversely proportional to the number of breeding individuals (N). The following formula illustrates the correlation between dF and N: dF = 1 / (Ouborg 1988). This shows that the loss of alleles is higher in proportion to the number of breeding birds. Moreover, in small populations genetic variation decreases more rapidly. Besides that white-speckled individuals were hunted because of their value to collectors, the beak tax caused an overall decline in the raven population. In all, it is plausible that the allele for

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white-speckled was recessive and was lost when the raven population reached its lowest abundance.

Last observations of white-speckled ravens

Records in the literature always referred to white-speckled or pied ravens, which were known to the Faroese people by the name Hvitravnur. Occasionally 'white' specimens may have been reported, as Debes (1673) suggested, but he also stated that 'the white ravens are not entirely white, but speckled with black feathers'. Because of their curiosity, white speckled-ravens were always highly prized. In the latter half of the 19th century especially, when the raven population was suffering intense persecution, white-speckled birds were even more severely hunted, leading to their extinction in the early 20th century. Several observations of presumed white-speckled birds were made post-1900: (1) 2 November 1902, Mykines (Andersen 1905, Salomonsen 1934), (2) autumn 1916, Velbastadur and Koltur (Ryggi 1951), (3) winter 1947 and again in late 1948, Nolsoy (Ryggi 1951), and (4) before Easter 1965, Sandvik (Nolsoe & Jespersen 2004).

In winter 1988/89, a 'white' raven was seen around Torshavn (J.-K. Jensen pers. comm.) and again at Velbastadur throughout winter 2007/08. It was photographed on 2 March 2008 (Fig. 6), clearly revealing that it was not a white-speckled individual. In this case, and almost certainly in three of the four cases listed above, a different mutation was involved, namely dilution. Dilution is a quantitative reduction of melanins (van Grouw 2013) meaning that the number of pigment granules is reduced, but the pigment itself is unchanged. Therefore, due to the lower concentration of granules than normal, a 'weaker' or 'diluted' colour occurs. This can be compared to a photograph in a newspaper; a high concentration of black ink dots close together is perceived as black, while fewer black dots in the same-sized area appear grey. Dilution is not uncommon in Carrion Crow, Hooded Crow and Rook.

The bird at Velbastadur was last seen in April 2008, and this, or another diluted individual, later in 2008 at Suduroy. Earlier the same year another was shot at Fugloy and donated to the Museum of Natural History in Torshavn (Fig. 7). Additionally, in August 2008, one was at Vidareidi and two more at Fugloy. In December 2009 another was seen twice at the south end of Suduroy. All those seen in 2007-09 appeared to be young (J.-K. Jensen pers. comm.). Their fate is unknown.

Obviously, the gene for dilution is present in Faroese ravens and probably has been for a considerable period. Returning to the observations in 1916, 1947, 1948 and 1965, all of these records were listed as 'white' ravens and not 'white-speckled'. Given that the gene for dilution is present and this aberration causes a solid grey-white plumage coloration, it is probable that these records were all diluted individuals. Only the record of 1902 mentioned white-speckling and therefore this date is considered the official date of extinction of the white-speckled variant of the Faeroes raven.

Conclusion

Despite being an icon for several centuries, the last white-speckled Faeroes raven was shot in 1902. One can assume that, along with the bird, the recessive allele for white-speckled plumage was also lost as a result of genetic drift combined with the severely depleted population. In addition to the species being hunted for pest control purposes, especially in the 19th century, white-speckled birds were consistently targeted by bird collectors.

The white-speckled plumage was caused by a recessive gene mutation known as leucism: a lack of melanin pigment due to the congenital and heritable lack of pigment cells from some or all of the skin parts where they would normally provide the growing feather

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with pigment. Inheritable leucism in birds is rare, but the black-and-white pattern in ravens (lack of pigment in the face, the 'hand' of the wings, the feet and belly) is the commonest form and occurs in many bird species.

Although the allele for leucism is apparently lost, another inheritable, recessive colour mutation now occurs in Faeroes ravens; dilution— a quantitative reduction of melanin pigment that produces a solid silvery grey plumage. Dilution is less rare and is occasionally found in other raven populations in North America and Europe.

To date, there are no confirmed records of diluted ravens breeding in the Faeroes, or elsewhere, but there is no reason to suppose that is impossible. Therefore, given that the Faeroe ravens represent an island population, the allele for dilution may become established in the future, just as the allele for leucism was in the past. Other island species offer examples of inheritable colour aberrations in their populations, of which the best is probably the flightless Weka Gallirallus australis in New Zealand. Three independent inheritable colour aberrations, melanism, leucism and progressive greying, occur in the South Island population (pers. obs.), and the extinct White Gallinule Porphyria albus of Lord Howe Island was the result of inherited progressive greying in the local Purple Gallinule P. porphyrio population (Hume & van Grouw submitted).

For at least a century, carriers of the dilution allele have been present on the Faeroes and a new icon in the island's avifauna is likely to arise, albeit only if diluted individuals are not targeted as a collector's item. The continued occurrence of 'white' ravens on the Faeroes is conditional upon naturalists of the future learning a lesson from the past.

Acknowledgements

First, 1 thank Dorete Bloch of the Faroese Museum of Natural History for her valuable input. This paper originally formed part of a joint, larger paper concerning the history of white-speckled ravens and museum specimens. Other results will be published in Archives of Natural History. Second, I thank Jens-Kjeld Jensen for photographs of the mounted diluted raven and for sharing his personal observations. Hans Eli Sivertsen provided photographs of the live diluted raven and Glen Roberts, Pieter van den Hooven and Igor Fadeev photographs of other leucistic birds. Last, but by no means least, I thank my wife, Katrina van Grouw, for the excellent drawing of the white-speckled raven (probably the first-ever depiction in a life-like pose) and for transforming my text into accessible English. Where unreferenced statements concerning colour aberrations in birds (inheritance, pigmentation, frequency) are made in this paper, they are based on personal findings during unpublished research into this subject over the last 19 years, involving both practical breeding experiments with domesticated birds and examination of >3,000 aberrantly coloured bird specimens in museum collections.

References:

Andersen, K. 1905. Meddelelser om Faeroernes fugle. 6te Raekke. Efter sktiftlige oplysninger fra P. F.

Petersen, Nolso, og S. Niclasen. Vidensk. Medd.fra den naturh. Foren. i Kbhvn 1905: 53-110.

Annandale, T. N. 1905. The Faroes and Iceland. Studies in island life. Clarendon Press, Oxford.

Ascanius, P. 1767. leones rerum naturalium, on figures enluminees d'histoire naturelle du nord, pt. 1. Copenhagen. Atkinson, G. C. 1989. journal of an expedition to the Feroe and Westman Islands and Iceland, 1833. Bewick-Beaufort Press, Newcastle-upon-Tyne.

Bloch, D. 1981. Fugletaelling pa Faeraerne sommeren 1981 - forelobig rapport. Dansk Orn. Foren. Tidsskr. 75: 1-6.

Bloch, D. 2012. Beak tax to control predatory birds in the Faroe Islands. Arch. Nat. Hist. 39: 126-135.

Bloch, D. & Sorensen, S. 1984. Yvirlit yvir Foroya fuglar. Frodskaparrit 31: 36-67.

Bloch, D., Skarup, P. & Grouw, H. van. 2010. Den feeroske hvidbrogede ravn. Foroya Natturugripasavn, Torshavn.

Crawford, R. D. 1990. Poultry breeding and genetics. Elsevier, Amsterdam.

Daneel, R. & Schumann, H. 1961. Uber die Entstehung und Vererbung der Blesse bei Mausen. Zeitschrift f. Vererbungslehre 92: 69-72.

Daneel, R. & Schumann, H. 1963. Die Entstehung des Farbmusters beim Lakenfelder Huhn. Roux' Archiv fur Entwicklungs Mechanik 154: 405-406.

Debes, L. 1673. Feeroee et Fseroa Reserata. Copenhagen.

Feilden, H. W. 1872. The birds of the Faeroe Islands. Zoologist (2)84: 3210-3225; 85: 3245-3257; 86: 3277-3295. Ferdinand, L. 1947. Studier af fuglelivet paa Faeroerne. Dansk Orn. Foren. Tidsskr. 41: 1-37.

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Fox, H. M. & Vevers, G. 1960. The nature of animal colours. Sidgwick & Jackson, London.

Graba, C. J. 1830. Tagebuch gefiihrt aufeiner Reise nach Faro im Jahre 1828. Hamburg.

Grouw, H. van. 2012. Geen kraai zo bont of er zit wel een vlekje aan; het fenomeen 'witte veren' in kraaiachtigen, Het Vogeljaar 60: 3-20.

Grouw, H. van. 2013. What colour is that bird? The causes and recognition of common colour aberrations in birds. Brit. Birds 106: 17-29.

Grouw, H. van & Bloch, D. submitted. The history of the remaining Faroese White-Speckled Raven specimens. Arch. Nat. Hist.

Hartert, E. & Kleinschmidt, O. 1901. Uebersicht liber den Formkreis Corvus corax. Novit. Zool. 8: 40^8.

Holm, P. A. 1848. Ornithologiske Bidrag til Faeroernes Fauna. Naturh. Tidsskr. 2: 465-525.

Hume, J. P. & van Grouw, H. submitted. Colour aberrations in some extinct and endangered birds. Bull. Brit. Orn. Cl.

Lamoreux, M. L., Delmas, V., Larue, L. & Bennett, D. C. 2010. The colors of mice: a model genetic network. Wiley- Blackwell, Chichester.

Landt, J. 1800. Forsog til en beskrivelse over Fseroerne. Einars Prent, Torshavn.

Muller, H. C. 1862. Faeroernes Fuglefauna med Bemaerkninger om Fuglefangsten. Vidensk. Medd. fra Dansk Naturh. Foren. 24: 1-78.

Nolsoe, J. P. A. & Jespersen, K. 2004. Havnar sega I. Torshavnar Kommuna, Torshavn.

Olaus Magnus. 1555. Historin de gentibus septentrionalibus. Rome.

Ouborg, N. J. 1988. Genetsche verarming: de problematiek van het Beheer van kleine plantenpopulaties. De Levende Natuur 89: 7-13.

Ryggi, M. A. 1951. Fuglabokin. Mentunargrunnur Logtingsins, Torshavn.

Sage, B. L. 1962. Albinism and melanism in birds. Brit. Birds 55: 201-220.

Salomonsen, F. 1931. Beretning om en Rejse til Faeroerne. Dansk Orn. Foren. Tidsskr. 25: 3-37.

Salomonsen, F. 1934. Aves. Pp. 1-277 in Jensen, A. S., Lundbeck, W., Mortensen, T. & Sparck, R. (eds.) Tlte zoology of the Faroes, vol. 3(2). Copenhagen.

Sparrman, A. 1786. Museum Carlsonianum, in quo novas et selectas aves, coloribus ad vivum brevique description illustrates, suasu et sumptibus generossimi possessoris. Fascuculus I. Stockholm.

Svabo, J. C. 1783. Indberetninger fra en Reise i Fsero 1781 og 1782. Copenhagen.

Vieillot, L. J. P. 1817. Nouveau dictionnaire d'histoire naturelle, vol. 8. Paris.

Wagener, G. 1959. Die Entstehung der Scheckung bei der Haubenratte. Biol. Zentralblad 78: 451-460.

Wagler, J. 1827. Systema avium, vol. 1. Stuttgart & Tubingen.

Wendt-Wagener, G. 1961. Untersuchungen fiber die Ausbreitung der Melanoblasten bei Einfarbig Schwarzen Ratten und bei Haubenratten. Zeitschrift f. Vererbungslehre 92: 63-68.

Worm, O. 1655. Museum Wormianum. Amsterdam.

Address: Bird Group, Dept, of Life Sciences, Natural History Museum, Akeman Street, Tring, Herts. HP23 6AP, UK, e-mail: h.van-grouw@nhm.ac.uk

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Breeding biology of the Grey-breasted Flycatcher Lathrotriccus griseipectus in south-west Ecuador

by Harold F. Greeney

Received 3 May 2013

Summary. — I studied two nests of Grey-breasted Flycatcher Lathrotriccus griseipectus in seasonally deciduous dry forest in south-west Ecuador. Nests were open cups constructed in natural depressions, one in the buttress of a large tree and one in a clump of bromeliads. Construction of one nest was completed in five days. Clutch size was two at one nest, and the eggs were pale beige with sparse, red-brown blotching. Eggs at both nests were laid 48 hours apart, and at one nest both eggs hatched 16 days after clutch completion. One nest was depredated immediately after the second egg was laid, but both nestlings fledged after 14 days at the other. Only one adult incubated, but both provisioned nestlings. The species' breeding biology is similar in all respects to that of the congeneric Euler's Flycatcher L. euleri, as well as to members of the closely related genus Empidonax of temperate and subtropical America.

Grey-breasted Flycatcher Lathrotriccus griseipectus is a monotypic species restricted to the Tumbesian region of western Ecuador and Peru (Fitzpatrick 2004). Within its small range, the species is generally uncommon and has apparently declined in recent years, consequently Bird life International (2013) treat it as Vulnerable. The species' only congeneric, Euler's Flycatcher L. euleri , is comparatively widespread and its breeding biology well known (Allen 1893, Euler 1900, Belcher & Smooker 1937, Aguilar et al. 1999, Aguilar & Marini 2007, Marini et al. 2007). The breeding biology of L. griseipectus, however, is wholly unknown. Here I describe the species' nest, eggs and nestlings from south-west Ecuador.

Methods and Results

I studied two nests of L. griseipectus at the Jorupe Reserve (Jocotoco Foundation), near Macara, Toja province, south-west Ecuador (04°23'S, 79°57'W; 600 m). Jorupe encompasses tropical deciduous forest typical of the Tumbesian bioregion (Best & Kessler 1995) and protects several other range-restricted species that have only recently had their breeding biology described (Miller et al. 2007, Rheindt 2008, Gelis et al. 2009). I took linear measurements of eggs to the nearest 0.1 mm and weighed them periodically during incubation using an electronic balance sensitive to 0.001 g. Eggs were individually marked using a permanent marker. I equate loss of mass during incubation with loss of water from the embryo (Ar & Rahn 1980). I made nest measurements to the nearest 0.5 cm. I collected nests after fledging or abandonment and let them dry ex-situ for two weeks before taking them apart and weighing their components to the nearest 0.001 g.

On 10 February 2010 I discovered the first nest (Fig. 1) at 18.00 h. It was empty and no adults were nearby. When I returned next day at 14.00 h, it contained a single egg, which was dry and cool to the touch. I returned six times during the afternoon but I did not observe an adult and the egg was cold and unattended until after dark. Next morning, pre-dawn, the egg was still cold, indicating that an adult had not spent the night at the nest. During seven visits on 12 February I encountered an adult only twice, suggesting that little time

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was devoted to incubation that day. The nest still held a single egg at 18.00 h and next morning at 05.00 h the egg was cold, again indicating that no adult had spent the night at the nest. When I returned that afternoon the second egg had been laid. I checked the nest contents daily until, on 1 March at 10.45 h, I discovered that the first egg to have been laid was hatching. The second hatched a few hours later. These observations indicate an incubation period of 16 days from the laying of the second egg until hatching of both eggs. Both nestlings left the nest at c.11.00 h on 15 March, giving a nestling period of 14 days.

I found the second nest on 16 February at 08.00 h, when I flushed an adult from a natural indentation at the base of a bromeliad (see below). On closer inspection I was unable to detect a nest, but 3M dead leaves inside the depression had possibly been brought by the adult. By the morning of 18 February, however, the nest cup was well formed and a few rootlets had been added to the lining. The first egg was laid on the morning of 20 February, giving a building period of no more than five days.

The nest still contained one egg at 18.00 h on 21 February and the second egg was laid

between 06.15 and 10.00 h on 22 February. On 23 February, however, the nest was empty at 16.15 h. Both adults were still present and swooped close to me while bill snapping. There were no signs of eggshells and the nest was undamaged.

Both nests were open cups composed externally of dead leaves and bark, with a relatively sparse inner lining of fungal rhizomorphs, dark rootlets and pale grass stems and fibres. Mean (± SD) total dry weight of both nests was 7.2 ± 1.9 g. The relative contribution of materials is presented in Table 1. The first nest (Fig. 1) was placed 1 m above ground in a shallow, upward-opening cavity in the buttress root of a canopy-emergent Ceiba tree. The cavity was longer than it was wide, and the nest completely filled it. The second nest (Fig.

Figure 1. Position of nest 1 of Grey-breasted Flycatcher Lathrotriccus griseipectus in a shallow cavity on a tree root, with inset showing a detail of the nest, Jorupe Reserve, Loja province, Ecuador (Harold F. Greeney)

TABLE 1

Dry weight (g) of materials comprising the nests of two Grey-breasted Flycatchers Lathrotriccus griseipectus

in south-west Ecuador.

	flexible bark	stiff bark	total	dead	fungal	dark	thin	Total
	strips	chips	bark	leaves	rhizomorphs	rootlets	stems
Nest 1	0.295	1.752	2.047	0.549	1.289	0.527	0.111	4.523
Nest 2	3.693	3.064	6.757	1.380	0.521	0.703	0.518	9.879
Mean	1.994	2.408	4.402	0.965	0.905	0.615	0.315	7.201
SD	1.201	0.464	3.330	0.294	0.272	0.062	0.144	1.894
% of total	27.7	33.4	61.1	13.4	12.6	8.5	4.4

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Figure 2. Position of nest 2 of Grey-breasted Flycatcher Lathrotriccus griseipectus in a depression amongst epiphytic bromeliads, with inset showing three eggs from two separate clutches, Jorupe Reserve, Loja province, Ecuador (Harold F. Greeney)

Figure 3. Nestlings of Grey-breasted Flycatcher Lathrotriccus griseipectus: upper row two days after hatching, and lower row nine days after hatching, Jorupe Reserve, Loja province, Ecuador (Harold F. Greeney)

2) was 4 m above ground, filling a slightly oblong hollow between the leaf bases of two bromeliads in a suspended clump of epiphytes. The external shape of the nests was oblong, fitting their respective cavities. The egg cups, however, were almost circular. Mean (cm ± SD) measurements were: greatest external diameter, 8.3 ± 0.4; smallest external diameter, 5.5 ± 0.7; external height, 5.3 ± 0.4; internal diameter, 4.8 ± 0.4; internal depth, 3.3 ± 0.4. Post-

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fledging, the internal cup of the first nest was slightly stretched and the internal diameter (measured at perpendicular angles) was 6 * 5 cm.

All four eggs had a pale beige ground colour with sparse cinnamon and orange-brown spotting, heaviest near the larger end (Fig. 2). Mean (± SD) size of three eggs was 18.0 ±1.4 x 13.5 ± 0.5 mm. Mean fresh weight of these eggs was 1.76 ± 0.27 g, all measured on the day they were laid. During incubation the two eggs at the first nest lost mass at a mean rate of 0.9 ± 0.1% / day of their original mass (range = 0.8-1 .0% / day).

I was unable to obtain linear measurements or weights of the nestlings but here provide a qualitative description. On hatching, nestlings bore a dorsal covering of pale brown natal down, which was paler at the base of the plumes and even paler posteriorly, affording them a camouflaged, sun-dappled appearance. Their skin was dusky, pinker ventrally and their legs yellowish. Their hills were similar in coloration to their legs and bore contrasting, bright yellow-white rictal flanges. The mouth lining, throughout the nestling period, was bright yellow-orange. Two days after hatching their appearance had changed little. By nine days of age, however, the nestlings were well feathered, with only sparse tufts of natal down remaining (Fig. 3).

Adults were extremely wary around the nests, generally flushing when approached to within 10 m, disappearing silently into the nearest dense vegetation. Using a tripod- mounted video camera I was able to ascertain that only one adult (presumably the female as in other tyrannids: Fitzpatrick 2004) incubated. Fiowever, both sexes provisioned the young.

Discussion

Unsurprisingly, all aspects of the biology of L. griseipectus described here are similar to those of L. euleri (Aguilar et al. 1999, Aguilar & Marini 2007, Marini et al. 2007). In fact, the nest and eggs of L. euleri photographed by Buzzetti & Silva (2008) are extremely similar in colour, placement and construction to one of the two nests described here. While a sample size of two nests precludes any firm comparisons, it appears that L. griseipectus may lay fewer eggs than L. euleri, which is usually reported as having a modal clutch size of three (Aguilar et al. 1999, Auer et al. 2007). The incubation and nestling periods documented here at a single nest of L. griseipectus are both c.l day shorter than mean durations reported for L. euleri (Aguilar et al. 1999, Auer et al. 2007). Both species of Lathrotriccus are also similar in their breeding habits to members of the closely related genus Empidonax of temperate regions (Lanyon 1986, Cicero & Johnson 2002), within which they have been placed in the past (Cory & Hellmayr 1927, Traylor 1979). Among other aspects, both genera share similar, cream-coloured or pale white, lightly spotted eggs, and well-constructed open-cup nests using a variety of substrates, but seemingly preferring well-supported sites such as ledges, thick branches, branch forks and even man-made structures (Russell & Woodbury 1941, King 1955, Walkinshaw & Henry 1957, Bowers & Dunning 1994, Briskie 1994, Wilson & Cooper 1998, Lowther 2000, Dobbs 2005).

Acknowledgements

Field work by HFG and the purchase of equipment used in this study were supported by Matt Kaplan, John V. Moore, the Population Biology Foundation, the Maryland Ornithological Society, and Field Guides, Inc. I thank the Jocotoco Foundation for access to their reserve and Eli & Issac Marck-Lichter for assistance in the field. J. Freile, A. Solano-Ugalde, and M. A. Crozariol suggested improvements to the submitted manuscript.

References:

Aguilar, T. M. & Marini, M. A. 2007. Nest and nest-site reuse within and between breeding seasons by three

Neotropical flycatchers (Tyrannidae). Brazil. ]. Biol. 67: 537-540.

Aguilar, T. M., Leite, L. O. & Marini, M. A. 1999. Biologia da nidificagao de Lathrotriccus euleri (Cabanis, 1968)

(Tyrannidae) em fragmentos de mata de Minas Gerais. Ararajuba 7: 125-133.

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Allen, J. A. 1893. On a collection of birds from Chapada, Matto Grosso, Brazil, by Mr. H. H. Smith. Part III.

Pipridae to Rheidae. Bull. Amer. Mus. Nat. Hist. 5: 107-158.

Ar, A. & Rahn, H. 1980. Water in the avian egg: overall budget of incubation. Amer. Zool. 20: 373-384.

Auer, S. K., Bassar, R. D., Fontaine, J. J. & Martin, T. E. 2007. Breeding biology of passerines in a subtropical montane forest in northwestern Argentina. Condor 109: 321-333.

Belcher, C. & Smooker, G. D. 1937. Birds of the colony of Trinidad and Tobago. Part VI. Ibis (14)1: 504-550. Best, B. J. & Kessler, M. 1995. Biodiversity and conservation in Tumbesian Ecuador and Peru. BirdLife International, Cambridge, UK.

BirdLife International. 2013. Species factsheet: Lathrotriccus griseipectus. www.birdlife.org/datazone/species/ index.html (accessed 25 February 2013).

Bowers, R. K. & Dunning, J. B. 1994. Buff-breasted Flycatcher (Empidonax fulvifrons). In Poole, A. (ed.) The birds of North America online. Cornell Lab of Orn., Ithaca, NY.

Briskie, J. V. 1994. Least Flycatcher (Empidonax minimus). In Poole, A. (ed.) Tire birds of North America online. Cornell Lab of Orn., Ithaca, NY.

Buzzetti, D. & Silva, S. 2008. Berios da vida, ninhos de aves brasileiras. Ed. Terceiro Nome, Sao Paulo.

Cicero, C. & Johnson, N. K. 2002. Phylogeny and character evolution in the Empidonax group of tyrant flycatchers (Aves: Tyrannidae): a test of W. E. Lanyon's hypothesis using mtDNA sequences. Mol. Phyl. & Evol. 22: 289-302.

Cory, C. B. & Hellnrayr, C. E. 1927. Catalogue of birds of the Americas and the adjacent islands, pt. V. Pubi. Field Mus. Nat. Hist. Zool. Ser. 13(5).

Dobbs, R. C. 2005. Breeding biology of Dusky Flycatchers in a southern Utah mixed conifer-aspen woodland. /. Field Orn. 76: 183-192.

Euler, C. 1900. Descrip^ao de ninhos e ovos das aves do Brazil. Rev. Mus. Paulista 4: 9-148.

Fitzpatrick, J. W. 2004. Family Tyrannidae (tyrant-flycatchers). Pp. 170M62 in del Hoyo, J., Elliott, A. & Christie, D. A. (eds.) Handbook of the birds of the world, vol. 9. Lynx Edicions, Barcelona.

Gelis, R. A., Greeney, H. F. & Miller, E. T. 2009. Further observations on nesting of Slaty Becard Pachyramphus spodiurus. Cotinga 31: 70-71.

King, J. R. 1955. Notes on the life history of Traill's Flycatcher ( Empidonax traillii) in southeastern Washington. Auk 72: 148-174.

Lanyon, W. E. 1986. A phylogeny of thirty-three genera in the Empidonax assemblage of tyrant flycatchers. Amer. Mus. Noi’it. 2846: 1-64.

Lowther, P. E. 2000. Pacific-slope Flycatcher Empidonax dijficilis / Cordilleran Flycatcher Empidonax occidentals.

In Poole, A. (ed.) The birds of North America online. Cornell Lab of Orn., Ithaca, NY.

Marini, M. A., Aguilar, T. M., Andrade, R. D., Leite, L. O., Anciaes, M., Carvalho, C. E. A., Duca, C., Maldonado-Coelho, M., Sebaio, F. & Gongalves, J. 2007. Biologia da nidifica^ao de aves do sudeste de Minas Gerais, Brasil. Rev. Bras. Orn. 15: 367-376.

Miller, E. T., Greeney, H. F., Zyskowski, K. & Gelis, R. A. 2007. First description of the nest and eggs of the Gray-and-gold Warbler (Basileuterus fraseri). Orn. Neotrop. 18: 617-621.

Rheindt, F. E. 2008. Description preliminar del nido de Pachyramphus spodiurus, especie amenazada. Cotinga 29: 162-163.

Russell, H. N. & Woodbury, A. M. 1941. Nesting of the Gray Flycatcher. Auk 58: 28-37.

Traylor, M. A. (ed.) 1979. Check-list of birds of the world, vol. 8. Mus. Comp. Zool., Harvard, Cambridge, MA. Walkinshaw, L. H. & Henry, C. J. 1957. Yellow-bellied Flycatcher nesting in Michigan. Auk 74: 293-304. Wilson, R. R. & Cooper, R. J. 1998. Breeding biology of Acadian Flycatchers in a bottomland hardwood forest. Wilson Bull. 110: 226-232.

Address: Yanayacu Biological Station & Center for Creative Studies, c/o Foch 721 y Amazonas, Quito, Ecuador, e-mail: revmmoss@yahoo.com

John P. Dumbacher

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A taxonomic revision of the genus Pitohui Lesson, 1831 (Oriolidae), with historical notes on names

by John P. Dumbacher

Received 13 May 2013

Summary. — Since 1925, the genus Pitohui has included six species: Hooded Pitohui dichrous, Variable P. kirhocephalus, Crested P. cristatus, Black P. nigrescens, Rusty P. ferrugineus and White-bellied Pitohuis P. incertus. Recent molecular work has shown that the genus, as thus circumscribed, is polyphyletic, and includes four different lineages of disparate taxonomic affinities. Here, I review the taxonomic placement of the pitohuis and discuss appropriate names for each taxon. Each monophyletic group has an available generic name, and the true pitohuis (genus Pitohui) include only P. kirhocephalus and P. dichrous. A recommended genus- level taxonomy is proposed, including a new genus-species combination for the Morningbird Pachycephala tenebrosa of Palau.

The genus Pitohui currently includes six species of medium-sized (60-100 g) forest¬ dwelling birds restricted to New Guinea and nearby continental islands. They are: Hooded Pitohui Pitohui dichrous, Variable Pitohui P. kirhocephalus, Crested Pitohui P. cristatus, Black Pitohui P. nigrescens, Rusty Pitohui P. ferrugineus and White-bellied Pitohui P. incertus (Stresemann 1925, Mayr 1941). All have jay-like omnivorous habits, are gregarious (Diamond 1987) and recent work has demonstrated that they carry potent neurotoxins in their skin and feathers that may function in chemical defence in some species (Dumbacher et al. 1992, 2000, Dumbacher 1999). Because of recent interest in the evolution of toxicity, they have been the subject of phylogenetic studies (Dumbacher & Fleischer 2001, Jonsson et al. 2007, Dumbacher et al. 2008); these have suggested that the genus Pitohui is currently misclassified. Below, I summarise the taxonomic history of the genus and propose a classification based upon the most recent available genetic data.

Variable Pitohui Pitohui kirhocephalus (Lesson & Garnot, 1827) was first described in 1827 in the zoological reports of the French exploration ship, the Coquille (Lesson & Garnot 1827), which sailed in 1822-25. The ship's zoological findings were published in several volumes and an atlas (Lesson & Garnot 1826-30), in which the written description of this species was given under the name Vanga kirhocephalus in livr. 14, pp. 632-633, attributed to Lesson alone. Zimmer (1926) reported that this part was published on 9 January 1930. However, a figure of the species was published earlier under the authorship of Lesson & Garnot with the name 'Pie-grieche cap-gris Lanius kirhocephalus' on pi. 11 of the Atlas, released on 17 January 1827 (Zimmer 1926). Thus, this name has precedence, with Lesson & Garnot as authors (ICZN 1999: Art. 12.2.7). In his written description, Lesson recorded that 'Le vanga cap-gris habite les forets de la Nouvelle-Guinee, aux alentours de Dorery, ou les Papous le nomment Pitohoui.' Thereafter, Lesson (1831) recognised the distinctness of the grey-capped bird and placed it in a new genus, Pitohui. No explanation was given for the origin of the name, but it presumably came from the local 'pitohoui' used by New Guinean villagers near Dorey (now Manokwari), New Guinea, Indonesia (Lesson & Garnot 1827). Thus, the name is properly cited as Pitohui kirhocephalus (Lesson & Garnot, 1827).

In 1850, Reichenbach proposed the name Rectes for this genus in his Avium systema naturale, but proffered no explanation for the name change, just a single black-and-white

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figure (no. 65) derived from Lesson & Garnot (1827), with the caption 'Redes'. At the time, no other species in the genus was described, so Redes was proposed effectively as a replacement name for Pitohui, with P. kirhocephalus the type species by monotypy. Shortly thereafter, Bonaparte (1850) supported this, saying of P. kirhocephalus, 'C'est le type d'un genre distinct pour lequel nous preferons adopter le nom de Redes, propose par Reichenbach.' Bonaparte (1850) also added two other species to the genus Redes, R. dichrous and R. ferrugineus, but P. kirhocephalus remained the type species as confirmed by multiple authors (Gray 1855, Salvadori 1881, Sharpe 1877). Although no justification was provided for their preference for Redes, it was customary among taxonomists at the time to prefer scientific names with Latin roots over those without and to provide new names for those that had been 'improperly' formed. Sharpe later confirmed that ‘Pitohui is doubtless an older name than Redes, but can surely be laid aside as a barbarous word' (Sharpe 1903).

The genus Rectes was in general usage after 1850, and two additional species were discovered and added to it. Redes nigrescens, Black Pitohui (Schlegel, 1871), and R. cristatus, Crested Pitohui (Salvadori, 1875). When Rothschild & Hartert (1903) resurrected the name Pitohui for the genus, they treated all other generic names that had been proposed for its various species as synonyms. Stresemann (1925) then reduced the number of species recognised by Sharpe (1877), Salvadori (1881) and others from 16 to six, relegating many taxa to subspecies but accepting those mentioned above and P. incertus van Oort (1909). For the remainder of the 20th century, the genus Pitohui was restricted to the above six species (Mayr 1941, Rand & Gilliard 1967, Beehler & Finch 1985, Beehler et al. 1986).

But generic dismemberment had been foreshadowed. Pseudocodes was introduced by Sharpe (1877) for Rusty Pitohui Pitohui ferrugineus (C. L. Bonaparte, 1850). With Rectes ferrugineus as its type species, Pseudorectes is a valid available name for the genus now known to include the species ferrugineus and incertus (Dumbacher et al. 2008). In recent phylogenies, Pseudorectes is strongly supported as a monophyletic genus sister to Colluricincla (shrike- thrushes) and distant from Pitohui (P. kirhocephalus) at family level (Jonsson et al. 2007, Dumbacher et al. 2008).

Black Pitohui Rectes nigrescens Schlegel, 1871, was also placed in its own genus, Melanoredes, by Sharpe (1877). Recent molecular work Iras shown that it too is not closely related to Pitohui but is sister to the whistler genus Pachycephala (Jonsson et al. 2007, Dumbacher et al. 2008) and should be recognised generically as well.

Crested Pitohui has also been recognised as distinct, and Iredale (1956) proposed the generic name Ornorectes for it. Molecular analysis has revealed that Crested Pitohui is most closely related to Crested Bellbird Oreoica gutturalis of Australia and Rufous-naped Whistler Aleadryas rufinucha of New Guinea (Jonsson et al. 2007, Dumbacher et al. 2008). Nevertheless, it is quite distinct morphologically and behaviourally from these relatives, so recognition of Ornorectes is recommended and the genus has recently been resurrected (Norman et al. 2009). These three taxa — Oreoica, Aleadryas and Ornorectes — have historically been placed in the polyphyletic Pachycephalidae or, possibly incorrectly, in the Colluricinclidae. Recent analyses suggest that they are distantly related to other members of the basal core Corvoidea, and the family name Oreoicidae has been applied to this group (Norman et al. 2009).

Molecular work also suggests that P. kirhocephalus and P. dichrous are indeed sister taxa (Dumbacher & Fleischer 2001, Jonsson et al. 2007, Dumbacher et al. 2008) and that this clade is probably a basal member of the Oriolidae, and perhaps sister to the figbirds, Sphecotheres Vieillot (Jonsson et al. 2010). Thus, the genus Pitohui contains only P. dichrous and P. kirhocephalus, and the best-available data place Pitohui within the Oriolidae, although more work is needed to confirm this.

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I recommend that common names remain unchanged, because, although the 'pitohuis' are not monophyletic, they do form an ecological group, much like warblers around the world that are not monophyletic either. Pitohuis also resemble one another in their use of potent toxins (Dumbacher et al. 1992, 2000, 2008), in their participation and movements in mixed-species flocks (Diamond 1987), their geographic restriction to New Guinea, and in morphology and other behaviours (Stresemann 1925). Tire only proposal for an alternative common name has been 'wood-shrike' (Iredale 1956), an epithet already in use for Tephrodornis (Prionopidae).

The Morningbird {Rede s tenebrosns Hartlaub & Finsch, 1868) of Palau has been placed with the genus Pitohui by some authors. Although currently placed in Colluricincla, two independent DNA studies suggest that it is nested well within the whistler clade, Pachycephala (Dumbacher et al. 2008, Jonsson et al. 2008), and should be transferred to that genus.

A recommended taxonomy of pitohuis follows:

Family Oriolidae

Genus Pitohui Lesson, 1831

Pitohui dichrous (Bonaparte, 1850), Hooded Pitohui

Pitohui kirhocevhalus (Lesson & Garnot, 1827), Variable Pitohui

Family Colluricinclidae

Genus Pseudoredes Sharpe, 1877

Pseudoredes ferrugineus (C. L. Bonaparte, 1850), Rusty Pitohui Pseudoredes incertus (van Oort, 1909), White-bellied Pitohui

Family Pachycephalidae

Genus Melanoredes Sharpe, 1877

Melanorectes nigrescens (Schlegel, 1871), Black Pitohui Genus Pachycephala Vigors, 1825

Pachycephala tenebrosa (Hartlaub & Finsch 1868), Morningbird (nov. comb.)

Family Oreoicidae

Genus Ornoredes Iredale, 1956

Ornoredes cristata (Salvadori, 1875), Crested Pitohui

Acknowledgements

I thank the editor, Guy Kirwan, for his help with this manuscript, and several people who read earlier drafts or provided critical reviews, including Bruce Beehler, Guy Dutson, Thane Pratt, Wojciech J. Pulawski and Richard Schodde.

References:

Beehler, B. M. & Finch, B. W. 1985. Species checklist of the birds of New Guinea. Royal Australasian Ornithologists' Union, Victoria.

Beehler, B. M., Pratt, T. K. & Zimmerman, D. A. 1986. Birds of New Guinea. Princeton Univ. Press.

Bonaparte, C. L. J. L. 1850. Note sur plusieurs families naturelles d'oiseaux, et descriptions d'especes nouvelles. Compt. Rend. Acad. Sci. Paris 31: 562-564.

Diamond, J. 1987. Flocks of brown and black New Guinean birds: a bicoloured mixed-species foraging association. Emu 87: 201-211.

Dumbacher, J. P. 1999. The evolution of toxicity in pitohuis: I. Effects of homobatrachotoxin on chewing lice (Order Phthiraptera). Auk 116: 957-963.

Dumbacher, J. P. & Fleischer, R. C. 2001. Phylogenetic evidence for colour pattern convergence in toxic pitohuis: Mullerian mimicry in birds? Proc. Roy. Soc. Lond. Ser. B 268: 1971-1976.

Dumbacher, J. P., Beehler, B. M., Spande, T. F., Garraffo, H. M. & Daly, J. W. 1992. Homobatrachotoxin in the genus Pitohui: chemical defense in birds? Science 258: 799-801.

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Dumbacher, J. P.( Spande, T. F. & Daly, J. W. 2000. Batrachotoxin alkaloids from passerine birds: a second toxic bird genus ( Ifrita kowaldi) from New Guinea. Proc. Natl. Acad. Sci. USA 97: 12970-12975.

Dumbacher, |. P., Deiner, K., Thomsen, L. & Fleischer, R. C. 2008. Phylogeny of the avian genus Pitohui and the evolution of toxicity in birds. Mol. Phyl. & Evol. 49: 774-781.

Gray, G. R. 1855. Catalogue of the genera and subgenera of birds contained in the British Museum. Brit. Mus. (Nat. Hist.), London.

Hartlaub, G. & Finsch, 0. 1868. On a collection of birds from the Pelew Islands. Proc. Zool. Soc. Loud. 1868: 4-9.

International Commission on Zoological Nomenclature (ICZN). 1999. International code of zoological nomenclature. Fourth edn. International Trust for Zoological Nomenclature, London.

Iredale, T. 1956. The birds of New Guinea. Georgian House, Melbourne.

Jonsson, K. A., Bowie, R. C. K., Norman, J. A., Christidis, L. & Fjeldsa, J. 2007. Polyphyletic origin of toxic Pitohui birds suggests widespread occurrence of toxicity in corvoid birds. Biol. Lett. 4: 71-74.

Jonsson, K. A., Irestedt, M., Fuchs, J., Ericson, P. G. P., Christidis, L., Bowie, R. C. K., Norman, J. A., Pasquet, E. & Fjeldsa, J. 2008. Explosive avian radiations and multi-directional dispersal across Wallacea: evidence from the Campephagidae and other crown Corvida (Aves). Mol. Phyl. & Evol. 47: 221-236.

Jonsson, K. A., Bowie, R. C. K., Moyle, R. G., Irestedt, M., Christidis, L., Norman, J. A. & Fjeldsa, J. 2010. Phylogeny and biogeography of Oriolidae (Aves: Passeriformes). Ecography 33: 232-241.

Lesson, R. P. 1831. Traite d'ornithologie. F. G. Levrault, Paris.

Lesson, R. P. & Garnot, P. 1826-30. Voyage autour du monde execute par ordre du Roi, sur la Corvette de Sa Majeste, La Coquille, pendant les annees 1822, 1823, 1824, et 1825. Arthus Bertrand, Paris.

Mayr, E. 1941. List of New Guinea birds: a systematic and faunal list of the birds of New Guinea and adjacent islands. Amer. Mus. Nat. Hist., New York.

Norman, J. A., Ericson, P., Jonsson, K., Fjeldsa, J. & Christidis, L. 2009. A multi-gene phylogeny reveals novel relationships for aberrant genera of Australo-Papuan core Corvoidea and polyphyly of the Pachycephalidae and Psophodidae (Aves: Passeriformes). Mol. Phyl. & Evol. 52: 488M97.

van Oort, E. D. 1909. Birds from southwestern and southern New Guinea. Nova Guinea Zool. 9: 51-107.

Rand, A. L. & Gilliard, E. T. 1967. Handbook of New Guinea birds. Weidenfeld & Nicolson, London.

Reichenbach, H. G. L. 1850. Avium systema naturale. Das Natiirliche System der Vogel. Friedrich Hofmeister, Dresden & Leipzig.

Rothschild, W. & Hartert, E. 1903. Notes on Papuan birds. Novit. Zool. 10: 65-116, 196-231, 435-480.

Salvadori, T. 1875. Descrizione di cinquantotto nuove specie di uccelli, ed osservazioni intorno ad altre poco note, della Nuova Guinea e di altre Isole Papuane, raccolte dal D.'Odoardo Beccari e dai cacciatori del Sig. A. A. Bruijn. Ann. Mus. Civ. Stor. Nat. Genova 7: 896-977.

Salvadori, T. 1881. Ornitologia della Papuasia e delle Molucche. Stamperia Reale Della Ditta G. B. Paravia & Comp., Turin.

Sharpe, R. B. 1877. Catalogue of the birds in the British Museum, vol. 3. Trustees of the Brit. Mus., London.

Sharpe, R. B. 1903. A hand-list of the genera and species of birds. Trustees of the Brit. Mus. London.

Stresemann, E. 1925. Die gattung Pitohui. Mitt. Zool. Mus. Berlin 11: 413-416.

Zimmer, J. T. 1926. Catalogue of the Edward E. Ayer Ornithological Library, pt. 1. Publ. Field Mus. Nat. Hist. 16: 1-364.

Address: Curator of Ornithology & Mammalogy, Dept, of Vertebrate Zoology & Anthropology, Institute of Biodiversity Science & Sustainability, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA, e-mail: jdumbaclier@calacademy.org

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Notes on the birds of Parque Nacional Saslaya, Reserva de

Biosfera Bosawas, Nicaragua

by Andrew C. Vallely & Liliana Chavarria-Duriaux

Received 13 June 2013

Summary. — We detail records of 23 rare or poorly known species from Parque Nacional Saslaya in the Reserva de Biosfera Bosawas of the Region Autonoma del Atlantico Norte in Nicaragua. These include the northernmost records of Violaceous Quail-Dove Geotrygon violacea, Black-headed Tody-Flycatcher Todirostrum nigriceps and Scarlet-thighed Dacnis Dacnis venusta. These findings clarify the northern range limits of several Caribbean slope species and highlight the importance of Central America's largest remaining wilderness area for biodiversity conservation.

Recent field work in eastern Honduras has documented northward range extensions for several Central American Caribbean slope species previously thought to reach only to Costa Rica or southern Nicaragua (Marcus 1983, Anderson et nl. 2004, Vallely et al. 2010). Northern Nicaragua also hosts extensive Caribbean slope humid forest but remains little surveyed since the collections of W. W. Huber (1932) at El Eden on 20 March-10 April 1922. In particular, the extensive Reserva de Biosfera Bosawas (RBB), including parts of dpto. Jinotega and the Region Autonoma del Atlantico Norte (RAAN), and spanning elevations of 200-1,650 m, is poorly explored. RBB encompasses 19,922 km2 including buffer zones and several core areas of 8,060 km2. This protected area, with those in adjacent eastern Honduras, forms the largest remaining wilderness in Central America. Although the northern RBB is sparsely populated, Parque Nacional Saslaya (PNS) encompassing 631 km2 in the south-eastern RBB is threatened by deforestation from small-landholder claims, some well inside the park boundaries (Kaimowitz et al. 2003, Stocks et al. 2007; ACV, LC-D pers. obs.). Anthropogenic landscapes now almost completely surround the eastern and southern borders of PNS (Stocks et al. 2007). While management challenges facing PNS are formidable, the area hosts many rare and poorly known species, and offers valuable opportunities for research, conservation and tourism. We are unaware of previously published reports on the avifauna of PNS and the larger core areas of the RBB to the north are virtually unknown to ornithologists. To date, 262 bird species have been reported in PNS (pers. obs.). Here, we detail observations of 23 rare or poorly known species in PNS, mostly in the rio Labu drainage, but also at Cerro El Torito in the western PNS.

Methods

We visited the PNS on 14-20 April 2009 (LC-D), 30 April-8 May 2012 (LC-D) and 3-13 March 2013 (ACV, LC-D). We searched for birds primarily in humid lowland rainforest and along adjacent watercourses. In May 2012 and March 2013 we used a dozen 12 x 2.5 m mist-nets with 30-36 mm mesh size deployed in forest understorey. On 2-8 May 2012 we amassed 546.33 mist net hours, and on 4-12 March 2013 another 582 mist-net hours. Observations were gathered opportunistically along the limited network of existing trails and passable watercourses. Coordinates for sites within PNS are: Cerro El Torito 13°42'50"N, 85°02'50"W, rio Labu drainage 13°41'20"N, 85°01'20"W, Camp La Vaquita 13°43'50"N, 85°01'49"W. The locations of these and most other localities mentioned in the text are shown in Fig. 1.

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Species accounts

Figure 1. Map showing the location of Parque Nacional Saslaya (PNS, dark shaded area) within the Reserva de Biosfera Bosawas (pale shaded area) in Nicaragua. Inset shows topography and individual sites within PNS. Numbers refer to sites mentioned in the text: (1) Cerro El Torito, PNS; (2) rio Labu drainage, PNS; (3) Camp La Vaquita, PNS; (4) El Eden, Region Autonoma del Atlantico Norte (RAAN); (5) rio San Juan drainage, dpto. Rio San Juan; (6) rio Grande, dpto. Matagalpa; (7) Penas Blancas, dpto. Jinotega; (8) Reserva El Jaguar, dpto. Jinotega; (9) Cerros Las Chachagiias, dpto. Jinotega; (10) Cerro Musun, dpto. Matagalpa; (11) Santo Domingo, dpto. Chontales; (12) Alamikangban / Layasiksa area, RAAN; (13) San Rafael del Norte, dpto. Jinotega.

SOLITARY EAGLE Buteogalliis solitarius

Considered Near Threatened (IUCN 2013) and very rare and local throughout its range. T. R. Howell (in Martinez-Sanchez & Will 2010) considered B. solitarius hypothetical in Nicaragua and knew of no records. Martinez-Sanchez (2007) mentioned a sight record from the north-central highlands of Nicaragua (dpto. Nueva Segovia) on 18 January 2004. On 3 May and 6 May 2012, LC-D & G. Duriaux observed singles over the rio Labu drainage at 425 m. On 11 March 2013 we observed a pair in courtship flight near Camp La Vaquita at c.650 m. A few minutes later a single, persistently vocalising, adult was seen at close range flying low over the canopy. On 13 March 2013 we observed an adult circling over the rio Labu drainage at c.400 m. D. Hille (pers. comm.) also observed one at PNS on 15 January 2013. Given the few reports in recent years from northern Central America (outside Belize) we suspect that PNS (and perhaps the greater RBB) may represent an important regional stronghold.

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BLACK-AND-WHITE HAWK-EAGLE Spizaetus melanoleucus

T. R. Howell (in Martinez-Sanchez & Will 2010) listed just two historical records from Nicaragua and the species is generally considered scarce and local over its extensive range. Three records from the rio Labu drainage appear to be the first in northern Nicaragua: ACV & LC-D observed a single adult over the rio Labu drainage, at 425 m, on 5, 7 and 8 March 2013.

SLATY-BACKED FOREST FALCON Micrastur mirandollei

Poorly known in Central America and long thought to range no further north than Costa Rica (AOU 1998). One seen by LC-D on 17 April 2009 at Cerro El Torito, PNS. It clearly showed plain white underparts and a yellow cere, eliminating Semiplumbeous Hawk Leucopternis semiplumbeus. ACV & LC-D also heard the distinctive song of M. mirandollei in the pre-dawn darkness at c.600 m on the mornings of 9-10 March 2013 at Camp La Vaquita. These are among the northernmost records, as the species has also recently been reported 140 km to the north in dpto. Gracias a Dios in eastern Honduras (Jones & Komar 2013b).

VIOLACEOUS QUAIL-DOVE Geotrygon violacea

Particularly rare in Central America, where unrecorded north of Nicaragua. Collected by Richardson early in the 20th century at Penas Blancas in eastern dpto. Jinotega and in the rio San Juan drainage at Los Sabalos (Martinez-Sanchez & Will 2010). More recently found in central Nicaragua at Cerro Musun by LC-D (eBird 2013). On 5 March 2013 ACV observed a single adult female for several minutes at close range in the rio Labu drainage at 425 m, which is the northernmost record ever.

SCARLET MACAW Ara macao

Described by Huber (1932) as common around El Eden. Decades later T. R. Howell suggested that it was declining on the Caribbean slope of Nicaragua (Martinez-Sanchez & Will 2010). We encountered singles and pairs at Cerro El Torito and in the rio Labii drainage, PNS. A. Farnsworth (eBird 2013) also reported it at rio Labu on 27 March 1996, and D. Hille (pers. comm.) encountered two at PNS on 18 January 2013. Though the species persists at PNS, large macaws are usually easily detected and our few encounters do not suggest the presence of a large population.

RUFOUS-VENTED GROUND CUCKOO Neomorphus geoffroyi

Very poorly known in Nicaragua, with historical specimens from the central foothills at Chontales (Salvin & Godman 1896), rio Tuma and Penas Blancas, and from southern Nicaragua in the rio San Juan region at Savala. Unreported in northern Nicaragua since Howell (1971) listed it for the Caribbean lowlands near Waspam, RAAN. We have no conclusive evidence for its presence at rio Labu, PNS, but park guards, unprompted by us, described the species in convincing detail and appeared familiar with its natural history. Pending more evidence we regard the species' occurrence as hypothetical.

CENTRAL AMERICAN PYGMY OWL Glaucidium griseiceps

Poorly known in northern Nicaragua. Several recent records from the rio San Juan drainage on the southern Caribbean slope (Martinez-Sanchez & Will 2010). We heard and recorded this owl's distinctive song on 9 March 2013 at Camp La Vaquita, at c.600 m.

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SHORT-TAILED NIGHTHAWK Lurocalis semitorquatus

Widespread species known from only a handful of records in Nicaragua, the earliest a specimen taken by Huber (1932) on the rio Bambana, RAAN. Additional records are given by Martinez-Sanchez & Will (2010). At dusk on 6 and 12 March 2013 we observed singles in the rio Labu drainage PNS. A. Farnsworth (eBird 2013) also reported it there on 27 March 1996.

TODY-MOTMOT Hylomanes momotula

Endemic to southern Mexico and Central America, where uncommon and local throughout. LC-D heard and saw one at 835 m at Cerro el Torito on 18 April 2009.

KEEL-BILLED MOTMOT Electron carinatum

Scarce, local, endemic to the Caribbean slope foothills of Central America, and treated as Vulnerable (IUCN 2013). We obtained sound-recordings at Camp La Vaquita, at c.600 m, in the rio Labu PNS. A. Farnsworth (eBird 2013) also reported the species there on 27 March 1996.

GREAT JACAMAR Jacamerops aureus

Long thought to range no further north than Costa Rica (AOU 1998), this widespread but scarce species was first reported from adjacent eastern Honduras in 2004 (Jones 2004, Vallely et al. 2010). The first records for Nicaragua were in the rio San Juan region (Munera- Roldan et al. 2007). Given decades of habitat destruction, thought to be declining in Central America (e.g. in Costa Rica; Sigel et al. 2005). We heard two and observed another at close range responding to playback on 12 March 2008 in the rio Labu drainage.

CINNAMON WOODPECKER Celeus loricatus

Long known in Nicaragua mainly from the rio San Juan region (AOU 1998). T. R. Howell (in Martinez-Sanchez & Will 2010) accepted its presence in northern Nicaragua based on a specimen taken at 275 m by Huber (1932) at El Eden, RAAN. Howell remarked that 'if its range is continuous it must be rare in most of eastern Nicaragua to have been missed by every other observer and collector, including Richardson'. We found it common in the rio Labu with multiples heard daily and one observed at close range responding strongly to playback. A. Farnsworth (eBird 2013) also reported it on 27 March 1996. ACV & M. Torrez detected a single by voice near Layasiksa in the eastern lowlands of RAAN on 23 August 2009. In northern Nicaragua may be confined to the interior Caribbean foothills, a little-studied region, and perhaps disjunct in relation to the southern Central American population.

WHITE-FRONTED NUNBIRD Monasa morphoeus

This widespread species reaches its northernmost limit in eastern Honduras (AOU 1998, Rasmussen & Collar 2002). It is poorly known in Nicaragua and has perhaps declined in abundance in parts of southern Central America (Sigel et al. 2005). We detected the species once by voice on 4 March 2013 in the rio Labu drainage. A. Farnsworth (eBird 2013) also reported M. morphoeus there on 27 March 1996.

STRIPED WOODHAUNTER Hyloctistes subulatus

Reaches its northernmost limit in eastern Honduras (Vallely et al. 2010) and known in northern Nicaragua from specimens taken at Rio Grande, dpto. Matagalpa, and Penas Blancas, dpto. Jinotega, by W. B. Richardson in 1908-09 (Miller & Griscom 1925). LC-D

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trapped two on 3-4 May 2012 in the rio Labu drainage (Jones & Komar 2013a) and ACV observed a single responding to playback there on 15 March 2013.

SCALE-CRESTED PYGMY TYRANT Lophotriccus pileatus

Long thought to range no further north than Costa Rica (AOU 1998). Marcus (1983) first reported it from eastern Honduras but the species was unknown in Nicaragua until one was closely observed by LC-D at Cerro El Torito on 17-18 April 2009. On 14 May 2009 LC-D & G. Duriaux observed two at Reserva El Jaguar, dpto. Jinotega. On 23 February 2010 G. Duriaux & LC-D observed one at Cerros Las Chachagiias, dpto. Jinotega (Chavarria & Batchelder 2012). We located two by voice near Camp La Vaquita, c.650 m, in the rio Labii drainage on 8-9 March 2013.

BLACK-HEADED TODY-FLYCATCHER Todirostrum nigriceps

Poorly known north of Costa Rica. On 5 March 2013 LC-D observed one in the canopy of tall, humid, broadleaf forest in the rio Labu drainage, discerning the yellow underparts, long, narrow tail and spatulate bill. A. Farnsworth (eBird 2013) also reported the species in the rio Labu drainage on 27 March 1996. These are the northernmost records and represent a range extension of 350 km from the rio San Juan region.

GREY-HEADED PIPRITES Piprites griseiceps

Endemic to Central America and poorly known. T. R. Howell (in Martinez-Sanchez & Will, 2010) knew of just two records in Nicaragua, one collected by Richardson in the rio San Juan region, and a specimen secured by Huber (1932) at El Eden. On 10 March 2013 ACV observed one for several minutes as it foraged with a large mixed-species flock in the midstorey of tall humid broadleaf forest at c.600 m at Camp La Vaquita. The bird perched upright in the open on a large branch before making an abrupt, upward sally to pursue a small lepidopteron. The same day LC-D & G. Duriaux observed a single nearby.

WING-BANDED ANTBIRD Myrmornis torquata

No documented records in Costa Rica (Stiles & Skutch 1989, Sandoval & Sanchez 2012) and only recently reported in eastern Honduras (Vallely et al. 2010), M. torquata is considered Near Threatened (IUCN 2013). In Nicaragua, known from specimens collected hy W. B. Richardson, including several from Penas Blancas, Jinotega. Another was collected in 1922 at El Eden (Huber 1932). Recently found to be locally common at Cerro Musun in dpto. Matagalpa (Chavarria & Duriaux 2011, 2013). LC-D trapped two in the rio Labu drainage on 3 and 5 May 2012. On 5 March 2013 we captured one, and detected others by voice, in the same area. Known from five sites in northern Nicaragua: Penas Blancas, dpto. Jinotega, El Eden, RAAN, Cerro Musun, dpto, Matagalpa, rio Labu drainage, RAAN, and Santo Domingo, dpto. Chontales. Records from southern Nicaragua in the rio San Juan region were presented by Cody (2000) and Munera-Roldan et al. (2007). Some sites listed in the literature are now largely deforested and we assume that the species must be locally extinct, e.g. at Santo Domingo (Salvin 1872) and El Eden (Huber 1932). Our experience suggests that in northern Central America the species prefers tall, humid, broadleaf forest with sparse understorey on steep, well-drained, slopes at 200-1,200 m. Known sites for M. torquata in Nicaragua, as in Honduras (Vallely et al. 2010), feature irregular forest floors with large boulders or karstic outcrops.

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LOVELY COTINGA Cotinga amabilis

Poorly documented in Nicaragua and probably declining in Central America (e.g. in Costa Rica; Sigel et al. 2005), where it is at best uncommon to rare and local throughout (Snow 2004). Kjeldsen (2005) reported it from Alamikangban in the lowlands of eastern RAAN. We did not find the species but A. Farnsworth (eBird 2013) reported it in the rio Labu drainage on 27 March 1996.

THREE-WATTLED BELLBIRD Procnias tricarunculatus

Regarded as Vulnerable (IUCN 2013) and undertakes complex migrations (Powell & Bjork 2004), with those of northern populations incompletely understood. LC-D encountered the species at Cerro El Torito on 17 April 2009. We heard several calling in late March 2013 in the rio Labu drainage, but the species was not common, with a silent female or young male on 6 March 2013 at c.400 m. Present in the north-central Nicaraguan highlands above 1,200 m in February-June (LC-D pers. obs). We consider those we encountered at 425-600 m in PNS as stragglers or late transients.

BLACK-THROATED WREN Pheugopedius atrogularis

Endemic to the Caribbean slope of Central America and poorly known near the northernmost limit of its range. T. R. Howell (in Martinez-Sanchez & Will, 2010) considered it to reach no further north than the rio Escondido. However, it was recently recorded in eastern Honduras, in dpto. Gracias a Dios (Jones 2004, Vallely et al. 2010). LC-D observed one on 17 April 2009 at c.600 m at Cerro El Torito PNS.

AMERICAN DIPPER Cinclus mexicanus

Very rare and local in Central America and unreported in Nicaragua since Richardson collected it at San Rafael del Norte in 1892 (Martinez-Sanchez & Will 2010). During May 2012 and March 2013 we observed singles and pairs on most days in appropriate habitat on the rio Labu and its tributaries. A Farnsworth also reported the species in this region on 27 March 1996 (eBird 2013). First records from Nicaragua in >100 years.

SCARLET-THIGHED DACNIS Dacnis venusta

Long thought to range no further north than Costa Rica (AOU 1998, Isler & Isler 1999), this widespread but scarce species was recently found in Nicaragua in the rio San Juan basin at Bartola (Jones & Komar 2012). On 30 April 2012 LC-D observed a male near Rosa Grande in the buffer zone of PNS. In the rio Lahti drainage, on 5 March 2013, and at Camp La Vaquita, on 7 March 2013, we observed at least two pairs foraging on inflorescences of canopy vines together with Blue Dacnis D. cayana, Shining Honeycreeper Cyanerpes lucidus and Green Honeycreeper Chlorophanes spiza. These are the northernmost records and represent a range extension of 350 km from the rio San Juan records.

Acknowledgements

We thank the following people, each of whom provided valuable assistance during field work. Georges Duriaux kindly helped with logistics and shared his observations. Ranger Atanasio Maldonado assisted us in many practical ways and shared his knowledge of the local avifauna. Marvin Torrez and David Hille also shared their observations. We are grateful to the Ministerio del Ambiente y los Recursos Naturales (MARENA) for granting research permits. ACV is indebted to the staff of the Dept, of Ornithology at the American Museum of Natural History, New York, for permitting access to the collections there as well as for their help in countless other ways. Paul Sweet, John Ascher and Anne Via made useful comments on an early draft. Comments from the referees, Knut Eisermann and Wayne Arendt, as well as the editor, Guy Kirwan, greatly improved the final manuscript.

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References:

American Ornithologists' Union (AOU). 1998. Check-list of North American birds. Seventh edn. American Ornithologists' Union, Washington DC.

Anderson, D. L., Wiedenfeld, D. A., Bechard, M. J. & Novak, S. J. 2004. Avian diversity in the Moskitia region of Honduras. Orn. Neotrop. 15: 447M82.

Chavarria, L. & Duriaux, G. 2011. Captura de Myrmornis torqunta (Wing-banded Antbird) en Reserva Natural Cerro Musun. Bol. Zeledonia 15: 93-97.

Chavarria, L. & Duriaux, G. 2013. Estado del Hormiguero Alifranjeado Myrmornis torqunta en Nicaragua. Cotinga 35: 71-75.

Chavarria, L. & Batchelder, R. L. 2012. Seven new records from Nicaragua and range extensions for two additional species. Cotinga 34: 28-32.

Cody, M. L. 2000. Antbird guilds in the lowland rainforests of southeast Nicaragua. Condor 102: 784-794.

eBird. 2013. eBird: an online database of bird distribution and abundance. Cornell Lab of Orn., Ithaca, NY. http://www.ebird.org (accessed 10 September 2013).

Howell, T. R. 1971. An ecological study of the birds of the lowland pine savanna and adjacent rainforest in northeastern Nicaragua. Living Bird 10: 185-242.

Huber, W. 1932. Birds collected in northeastern Nicaragua in 1922. Proc. Acad. Nat. Sci. Phil. 84: 205-249.

Isler, M. L. & Isler, P. R. 1999. Tire tanagers. Second edn. Smithsonian Institution Press, Washington DC.

IUCN. 2013. IUCN Red List of threatened species. Version 2013.1. www.iucnredlist.org (accessed 10 September 2013).

Jones, H. L. 2004. The winter season, December 2003 through February 2004: Central America. N. Amer. Birds 58: 290-292.

Jones, H. L. & Komar, O. 2012. The winter season December 2011 through February 2012: Central America. N. Amer. Birds 66: 353-359.

Jones, H. L. & Komar, O. 2013a. Spring migration March through May 2012: Central America. N. Amer. Birds 66: 562-566.

Jones, H. L. & Komar, O. 2013b. The nesting season June through July 2012: Central America. N. Amer. Birds 66: 739-742.

Kaimowitz, D., Faune, A. & Mendoza, R. 2003. Your biosphere is my backyard: the story of Bosawas in Nicaragua. CIFOR Working Paper 25.

Kjeldsen, J. P. 2005. Aves del Municipio Rio Prinzapolka, un inventario de base. Ward 41: 31-64.

Marcus, M. J. 1983. Additions to the avifauna of Honduras. Auk 100: 621-629.

Martinez-Sanchez, J. C. 2007. Lista patron de las aves de Nicaragua; con informacion de nuevos registros, distribucion y localidades donde observar aves. Alianza para las Areas Silvestres, Managua.

Martinez-Sanchez, J. C. & Will, T. (eds.) 2010. Thomas R. Howell's Check-list of the birds of Nicaragua as of 1993. Orn. Monogr. 68.

Miller, W. DeW. & Griscom, L. 1925. Descriptions of new birds from Nicaragua. Amer. Mu s. Novit. 159: 1-9.

Munera-Roldan, C., Cody, M. L., Schiele-Zavala, R. H., Sigel, B., Woltmann, J. S. & Kjeldsen, J. P. 2007. New and noteworthy records of birds from south-eastern Nicaragua. Bull. Brit. Orn. Cl. 127: 152-161.

Powell, G. & Bjork, R. 2004. Habitat linkages and the conservation of tropical biodiversity as indicated by seasonal migrations of Three-wattled Bellbirds. Conserv. Biol. 18: 500-509.

Rasmussen, P. C. & Collar, N. J. 2002. Family Bucconidae (puffbirds). Pp. 102-138 in del Hoyo, J., Elliott, A. & Sargatal, J. (eds.) Handbook of the birds of the world, vol. 7. Lynx Edicions, Barcelona.

Salvin, O. 1872. Notes on the birds of Nicaragua, based upon a collection made at Chontales by Mr. Thomas Belt. Ibis (3)2: 311-323.

Salvin, O. & Godman, F. D. 1896. Biologia Centrali- Americana. Aves, vol. 2. Taylor & Francis, London.

Sandoval, L. & Sanchez, J. E. 2012. Lista de aves de Costa Rica. Union de Ornitologos de Costa Rica, San Jose.

Sigel, B. J., Sherry, T. W. & Young, B. E. 2005. Avian community response to lowland tropical rainforest isolation: 40 years of change at La Selva Biological Station, Costa Rica. Conserv. Biol. 20: 111-121.

Snow, D. W. 2004. Lovely Cotinga Cotinga amabilis. P. 93 in del Hoyo, J., Elliott, A. & Christie, D. A. (eds.) Handbook of the birds of the world, vol. 9. Lynx Edicions, Barcelona.

Stiles, F. G. & Skutch, A. 1989. A guide to the birds of Costa Rica. Cornell Univ. Press, Ithaca, NY.

Stocks, A., McMahan, B. & Taber, P. 2007. Indigenous, colonist, and government impacts on Nicaragua's Bosawas Reserve. Conserv. Biol. 21: 1495-1505.

Vallely, A. C., Gallardo, R. J. & Ascher, J. A. 2010. Notes on the birds of the Rio Platano Biosphere Reserve including four new species for Honduras. Bull. Brit. Orn. Cl. 130: 52-60.

Addresses: Andrew C. Vallely, 84 Riverside Drive, Apt. 2F, New York, NY 10024, USA, e-mail: andrewcvallely@ gmail.com. Liliana Chavarria-Duriaux, Reserva El Jaguar, Jinotega, Nicaragua, e-mail: orion.liliana@ gmail.com

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The avifauna of Mt. Karimui, Chimbu Province, Papua New Guinea, including evidence for long-term population dynamics in undisturbed tropical forest

Ben Freeman & Alexandra M. Class Freeman

Received 27 July 2013

Summary. — We conducted ornithological field work on Mt. Karimui and in the surrounding lowlands in 2011-12, a site first surveyed for birds by J. Diamond in 1965. We report range extensions, elevational records and notes on poorly known species observed during our work. We also present a list with elevational distributions for the 271 species recorded in the Karimui region. Finally, we detail possible changes in species abundance and distribution that have occurred between Diamond's field work and our own. Most prominently, we suggest that Bicolored Mouse-warbler Crateroscelis nigrorufa might recently have colonised Mt. Karimui's north-western ridge, a rare example of distributional change in an avian population inhabiting intact tropical forests.

The island of New Guinea harbours a diverse, largely endemic avifauna (Beehler et al. 1986). However, ornithological studies are hampered by difficulties of access, safety and cost. Consequently, many of its endemic birds remain poorly known, and field workers continue to describe new taxa (Pratt 2000, Beehler et al. 2007), report large range extensions (Freeman et al. 2013) and elucidate natural history (Dumbacher et al. 1992). Of necessity, avifaunal studies are usually based on short-term field work. As a result, population dynamics are poorly known and limited to comparisons of different surveys or differences noticeable over short timescales (Diamond 1971, Mack & Wright 1996).

Here, we report new distributional and ecological observations made during field work on Mt. Karimui, Chimbu Province. Mt. Karimui's avifauna was studied by Jared Diamond in 1965 (Diamond 1972) and we purposely returned to the same ridge he worked. Analysis of elevational changes in Mt. Karimui's avifauna will be presented elsewhere; here, we describe our survey results, including differences from Diamond's historical transect that may reflect avifaunal changes. Tropical bird communities in undisturbed forest are seldom subject to long-term monitoring studies, but populations are thought to be relatively stable (Munn 1985, Greenberg & Gradwohl 1997, Brooks et al. 2005, Martinez & Gomez 2013), albeit with local extinctions and colonisations well documented in fragmented forest (Willis 1974, Robinson 1999, Brook et al. 2003, Sodhi et al. 2004). Finally, we present a comprehensive list of Mt. Karimui's birds, including known elevational ranges and conservation status.

Methods

Study site.— The extinct volcano of Mt. Karimui lies in the southern part of New Guinea's Central Ranges, in Chimbu Province (Fig. 1). Satellite imagery and maps clearly demonstrate Mt. Karimui to be an old volcano with a blown-out caldera. However, when viewed from the Karimui Plateau it appears as a series of discrete peaks. These ridges rise steeply from the relatively level Karimui Plateau (c.1,100 m), and are tallest in the north, where they reach c.2,550 m. Whereas Mt. Karimui is covered in primary forest, a significant part of the fertile Karimui Plateau is under small-scale agriculture, particularly at Karimui Station, a government post c.6 km north-east of Mt. Karimui. For our purposes, we define

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Figure 1. Map of Mt. Karimui, Chimbu Province, Papua New Guinea. The airstrip at Karimui Station, served primarily by small aircraft from Goroka, provides the principal access to the area. Our field work was concentrated on Mt. Karimui's north-west ridge above Yogoromaru village, the same ridge surveyed by Diamond (1972). The transect we surveyed is marked by the black line. See Table 1 for more information on site locations.

the Karimui area as the northern slopes of Mt. Karimui and adjacent Karimui Plateau south and east of the Tua River (see Fig. 1). All field work reported here pertains to this 'Karimui area': the southern ridges, foothills and adjacent lowlands of Mt. Karimui are ornithologically unexplored.

We first visited Mt. Karimui in 2011 during a week-long exploratory visit. We returned for two field seasons in 2012 to survey its birdlife: 13 June to 27 July (June-July) and 12 October to 14 December (October-December). Field work was concentrated on the same ridge studied by Diamond in 1965, Mt. Karimui's north-west ridge located above Yogoromaru village (Diamond 1972, see Fig. 1). In 1965, this ridge was entirely covered by primary forest. Environmental changes in the intervening years have been minor; we found it to be covered with primary forest above c.1,200 m, with the exception of two recently cleared small (c.l ha) patches, the first a subsistence garden at 1,280 m and the second a clearing at the base of a recently constructed cellphone tower at the summit (2,520 m). To facilitate comparisons with previous data, we avoided surveying in the vicinity of non¬ forest habitats. We therefore conducted lower elevation field work (at 1,130-1,330 m) on an entirely forested ridge 0.5 km east of Mt. Karimui's north-west ridge (Camp 3, see Table 1), then surveyed the spine of the ridge from 1,330 m to the summit at 2,520 m (based at Camps 1-2, see Table 1). We also visited several lower elevation sites for short periods (see Table 1 for sites and survey effort).

Field work. — We censused bird communities using mist-net surveys, point counts and ad lib observations (Table 1). Mist-net surveys in June-July were made along the spine of Mt. Karimui's north-west ridge, with a single mist-net survey at Bosiamaru (see Table 1). Mist-nets touched the ground in order to trap terrestrial species. Along Mt. Karimui's north¬ west ridge, we used flagging tape to partition the ridge into sections of 25 vertical m (e.g. 1,400-1,425 m). We measured elevation using the barometric altimeter in a Garmin 62S GPS unit, calibrated at Karimui airstrip (1,112 m per Diamond 1972) and using the average of readings taken on multiple days. We mist-netted along the ridgeline in discrete 'segments' of 24-30 nets (corresponding to 100-175 m elevation), and opened nets from 06.00 h to 13.00 h for two days per segment. Mist-nets were not operated in rain. Upon finishing one segment, we moved nets to higher elevations along the same trail and repeated the

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TABLE 1

Location of field sites surveyed in the Karimui area, with approximate survey effort and brief habitat description. Tire vast majority of field work occurred in the vicinity of our three field camps along

Mt. Karimui's north-west ridge.

	Elevation (m)	Latitude	Longitude	Survey type	Survey effort	Habitat
Mt. Karimui Camp 1	1,420	06°54.123	144°74.263	Mist-nets; point counts	r.45 man-days	Tall primary forest near base of ridgeline
Mt. Karimui Camp 2	1,890	06°54.693	144°75.250	Mist-net; point counts	c.45 man-days	Primary montane forest with epiphyte¬ laden trees, some bamboo tangles
Mt. Karimui Camp 3	1,240	06°52.748	144°74.518	Mist-nets; point counts	20 man-days	Tall primary forest at Mt. Karimui's base
Yogoromaru	c. 1,1 00	06°50.672	144°74.178	Qualitative observations	four man-days	Mostly agricultural landscape
Karimui Station area	c.1,100	06°49.254	144°82.473	Qualitative observations	c.20 man-days	Mostly agricultural landscape
Bosiamaru	1,100-1,150	06°50.689	144=80.149	Mist-nets	six man-days	Second growth, heavily hunted
Sena River	750	06°47.735	144°83.475	Qualitative observations of 'salt lick' site	three man-days	Second growth, heavily hunted
Tua River	570	06°45.176	144°78.164	Qualitative observations	two man-days	Mix of second growth and primary forest

process. Our net-line stretched unbroken from 1,330 m to 2,200 m in this fashion. Difficult terrain in the high-elevation elfin forest permitted only scattered mist-netting above 2,200 m and entirely prevented it above 2,400 m. Finally, we mist-netted in lower elevation forest (1,130-1,330 m) along a parallel ridge (described above. Camp 3, see Table 1). Importantly, this mist-net effort closely matches that of Diamond (1972), who likewise ran a nearly continuous mist-net lane along Mt. Karimui's north-west ridge. Unlike Diamond, we did not collect specimens. Instead, individuals trapped were weighed, measured (wing, tail, culmen, tarsus), scored for moult and photographed. We also took blood samples from the brachial vein of the majority of captured individuals. Finally, we clipped the distal portion of the right three outer rectrices, permitting easy diagnosis of recaptures.

One observer (BGF) completed point counts in both 2012 field seasons on Mt. Karimui's north-west ridge. Point counts in June-July were at 1,130-2,520 m (n = 40); in October- December at 1,330-2,520 m (n = 30). Each point count location was at least 150 m distant from neighbouring point count sites. We conducted five-minute audiovisual point counts, repeating counts on each of three separate mornings (06.00-12.00 h, mostly 06.30-09.00 h). We augmented our quantitative surveys with qualitative observations lacking effort information during the course of field work. Survey effort on Mt. Karimui's ridge was approximately equal between Diamond's July-August transect (33 days) and our June-July (38 days) and October-December (34 days) field seasons, facilitating comparisons. Audio¬ recordings will be archived at the Macaulay Library of Natural Sounds at the Cornell Lab of Ornithology, Ithaca, NY, while observational data are archived in the Avian Knowledge Network via eBird.

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Results

Some 271 bird species occur in the Karimui area, with a further four species reported by local informants (Appendix 1). This total sums extensive field work by ourselves and Diamond, and is probably near-complete. Nevertheless, species richness estimated by Diamond's survey and our own differed: we documented 245 species, Diamond 234. These different totals largely reflect geographic differences in survey effort. Diamond (1972) spent more time at low elevations on the Karimui Plateau and employed native hunters to collect specimens, while our field work was concentrated on Mt. Karimui's slopes. However, research effort on Mt. Karimui's north-west ridge was qualitatively similar between historical and modern transects, suggesting that some of the observed differences may reflect changes in species' populations. Diamond's extensive surveys on Mt. Karimui lack quantitative effort data: our mist-net effort summed 3,665 net-hours, during which time we captured 977 individuals of 91 species. Point counts detected 130 species in 2,082 species / point count combinations. We describe our observations of population dynamics, elevational range extensions and ecological notes for 21 species below.

DWARF CASSOWARY Casuarius bennetti / SOUTHERN CASSOWARY C. casuarius

Cassowaries are New Guinea's largest terrestrial animals and highly valued for their meat (Beehler et al. 1986). Informants consistently described them as largely extirpated from the Karimui region, correlating their disappearance to a period of intense hunting in the 1960s and 1970s when metal snares were first used. We never encountered cassowary droppings in the forest— which are frequently encountered where cassowaries are present (BGF pers. obs.) — and saw just one captive bird, a Dwarf Cassowary chick acquired by a Yogoromaru hunter from a remote and seldom-hunted location near the Tua River. The species resident in the Karimui area is Dwarf Cassowary. However, informants described Southern Cassowary as resident in the lowlands south of Mt. Karimui, and reported it to occasionally venture to the Karimui area treated by this manuscript.

COLLARED BRUSHTURKEY Talegalla jobiensis /

BLACK-BILLED BRUSHTURKEY T. fitscirostris

Talegalla are shy forest-dwellers with braying vocalisations. Due to the difficulty in identifying Talegalla vocalisations to species, distributional knowledge is poor. For example. Diamond was unable to identify which Talegalla inhabits Mt. Karimui (Diamond 1972) and it was only recently that Collared Brushturkey was documented south of New Guinea's Central Ranges (Mack & Wright 1996). We frequently heard Talegalla vocalisations below 1,890 m, eventually photographing a Collared Brushturkey at its mound nest at 1,390 m. We suspect Black-billed Brushturkey also occurs at Karimui and replaces Collared Brushturkey at lower elevations: our best local informant described the green-legged Black-billed Brushturkey as a common resident near the Tua River. We consider this informant credible, as he accurately described the leg colours and preferred elevations of the three megapodes we encountered, the montane Collared and Wattled Brushturkeys Aepygpodius arfakianus and widespread Orange-footed Scrubfowl Megapodius reinwardt.

PAPUAN EAGLE Harpyopsis novaeguineae

We recorded this raptor only a few times: one was observed perched in the canopy at 1,300 m in 2011, and vocalising birds were heard c.3 times in 2012 at our 1,420 and 1,890 m camps. While never abundant throughout its range, this eagle is usually easily detected by voice in forested montane environments (Beehler et al. 1986; BGF pers. obs.). Diamond (1972)

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noted the species on Mt. Karimui, but did not assess its relative abundance, which often correlates with hunting levels (K. D. Bishop pers. comm.), and we consider it probable that this species is impacted by hunting at Karimui. For example, one local informant showed us a full set of talons from a bird hunted in early 2012 and several informants told us that hunters frequently target Papuan Eagles.

BUFF-BANDED RAIL Gallirallus philippensis

Widespread throughout Melanesia, colonising even remote Pacific islands (Beehler et al. 1986). We found it relatively common in agricultural areas at Karimui Station. Diamond did not record it. It is unlikely that Diamond overlooked this rail: he surveyed appropriate habitats near Karimui airstrip and collected five Rufous-tailed Bush-hens Amaurornis moluccana (Diamond 1972), a more secretive species (BGF pers. obs.). It is therefore probable that Buff-banded Rail— an excellent coloniser (Diamond & LeCroy 1979) — has recently colonised the area, presumably in response to the large increase in agricultural land since 1965.

RUFESCENT IMPERIAL PIGEON Ducula chalconota

We regularly heard this montane pigeon on Mt. Karimui. Diamond (1972) was familiar with its distinctive vocalisations, but did not find it on Mt. Karimui. Because columbids regularly undertake seasonal movements and are difficult to detect when not vocal (Diamond 1972), this species may have been overlooked by Diamond or was rare or absent at the time of his survey.

ZOE'S IMPERIAL PIGEON Ducula zoeae

We found this lowland species well above its published elevation limit of 1,500 m (Baptista el al. 1997). In June-July, it was one of the most commonly detected species on point counts, vocally abundant to c.1,900 m with some heard up to 2,080 m. It probably undertakes seasonal elevational movements: in October-November, we recorded this species infrequently and only below 1,620 m.

STRIATED LORIKEET Charmosyna multistriata

Formerly considered absent from Papua New Guinea's southern watershed (Beehler el al.. 1986). We identified the species on three occasions in November 2012 at our 1,420 m camp. All observations were of small flocks (2M birds) in flight, identified by their all-green coloration with yellowish-streaked underparts, distinct from the similar Goldie's Lorikeet Psitteuteles goldiei, also present. Lorikeets are difficult to positively identify in flight and our records should be considered provisional. C. multistriata is nomadic, often present at a site for several years before disappearing (K. D. Bishop pers. comm.). Our probable records and recent observations from the Crater Mountain area immediately east of the Karimui Plateau (Mack & Wright 1996) suggest this species' wanderings include much of Papua New Guinea's southern watershed.

LORIKEET SP. Charmosyna sp.

We observed a vocalising Charmosyna in August 2011, when an adult flew by at eye level on the rim of the Karimui Plateau. We judged it to be smaller than Coconut Lorikeet Trichoglossus haematodus, the commonest lorikeet at the site, and provisionally identified it as Josephine's Lorikeet Charmosyna josefinae based on its relatively large size, very long yellow-tipped tail and red rump. However, we could not conclusively eliminate other Charmosyna species (e.g. Papuan Lorikeet C. papou) and Josephine's Lorikeet has not been

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documented east of Mt. Bosavi (Collar 1997), c.175 km west of Mt. Karimui. This was our sole, possible, observation suggesting it may be an occasional visitor to the region and that future field workers should remain alert for Josephine's Lorikeet.

PESQUET'S PARROT Psittrichas fulgidus

Prized for its vermilion and black flight feathers, hunting has extirpated this species in many locations (Beehler et al. 1986, Mack & Wright 1998). We did not record it on the Karimui Plateau, where hunting pressure has been intense for at least 50 years (Wagner 1967). In fact, hunting may have formerly almost extirpated the species from the entire Karimui area: Diamond observed the species just once during several months of field work on the Karimui Plateau and on Mt. Karimui. However, we regularly observed pairs or small groups roosting around our 1,420 m and 1,890 m camps, suggesting that it is currently uncommon on the slopes of Mt. Karimui. Local informants reported the species to be fairly common in parts of the Tua River Valley far from human settlements.

PACIFIC KOEL Eudynamys orientalis

Regularly heard up to 2,120 m, well above its previously known elevational ceiling of 1,500 m (Beehler et al. 1986).

WHITE-CROWNED CUCKOO Cacomantis leucolophus

This lowland species generally occurs below 1,740 m (Coates 1985). We frequently heard it during both 2012 field seasons up to c.2,200 m, with one record from 2,520 m in July 2012. We have also heard the species at 2,200-2,300 at Hogave, Mt. Michael, Eastern Highlands, and it is regularly encountered at other highland locations (e.g. Ambua Lodge near Tari, Papua New Guinea; K. D. Bishop pers. comm.): It is probably widespread in montane forest.

HOOK-BILLED KINGFISHER Melidora macrorrhina

Commonly heard pre-dawn up to 1,870 m. This is a new high-elevation record for this lowland species, which was previously known only to 1,280 m (Woodall 2001).

YELLOWISH-STREAKED HONEYEATER Ptiloprora meekiana This rare and unobtrusive montane species is probably nomadic (K. D. Bishop pers. comm.), and has been recorded only a few times in the Central and Eastern Highlands of Papua New Guinea (Higgins et al. 2008). We did not find it in June-July 2012, nor did Diamond record this species in 1965. However, we observed one in a flowering tree at 1,880 m on three consecutive days in October 2012.

BICOLORED MOUSE-WARBLER Crateroscelis nigrorufa

Patchily distributed throughout montane New Guinea. When presexat, it occupies a narrow elevational band between the closely related lowland Rusty Mouse-warbler C. murina and montane Mountain Mouse-warbler C. robusta. Despite extensive mist-netting effort and numerous mist-netted Rusty and Mountain Mouse-warblers, Diamond did not record Bicolored Mouse-warbler on Mt. Karimui in 1965. In contrast, we mist-netted 16 C. nigrorufa in June-July 2012 (1,620-1,940 m) and regularly observed small parties during both field seasons. Given that Diamond operated a series of mist-nets across the entire elevational zone (J. Diamond pers. comm.), we consider it highly unlikely that this species was overlooked by him. Instead, we suggest that this species was very rare or truly absent on Mt. Karimui's north-west ridge in 1965 and has since become relatively common.

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SCRUBWREN SP. Sericornis sp.

We observed small flocks of an unidentified scrubwren ( Sericornis sp.) on six occasions at 1,280-1,355 m. Flocks comprised 3-8 individuals that foraged 1-12 m above ground. Three were mist-netted at 1,310 m and blood samples taken. They were morphologically similar (if not identical) to Large Scrubwren S. nouhuysi, which was commonly seen and mist-netted above 1,470 (AMCF photographs). However, the only Sericornis we observed at 1,355-1,470 m was the much smaller Grey-green Scrubwren S. arfakianus. Although speculative, these unidentified lower elevation Sericornis could represent a new population of Perplexing Scrubwren S. virgatus, a phenotypically variable low-elevation species confusingly similar to Large Scrubwren. This possibility is bolstered by recent records of Perplexing Scrubwren in southern Papua New Guinea (J. Diamond pers. comm.). Genetic studies are necessary to evaluate the taxonomic status of these unidentified scrubwrens.

CHESTNUT-BACKED JEWEL-BABBLER Ptilorrhoa castanonota /

SPOTTED JEWEL-BABBLER P. leucosticta

Many species in tropical mountains are elevational replacements (closely related species with parapatric elevational distributions: Diamond 1973, Terborgh & Weske 1975, Jankowski et al. 2012, Freeman et al. 2013). Range borders are often very abrupt, but elevational gaps sometimes exist between two replacements (Terborgh & Weske 1975). For example. Diamond noted a substantial gap between the foothill Chestnut-backed Jewel- babbler and montane Spotted Jewel-babbler on Mt. Karimui (Diamond 1972). This gap appears to have been maintained. We expended significant effort determining Ptilorrhoa elevational distributions, and found a gap between the highest Chestnut-backed Jewel- babbler territory at 1,460 m and the lowest Spotted Jewel-babbler territory at 1,510 m.

RUFOUS -N APED WHISTLER Aleadryas rufinucha

Regularly recorded above 1,920 m. Our only record below this was a juvenile mist-netted at 1,300 m, which echoes records of juveniles of other species found well outside their typical elevational distributions in New Guinea, supporting the hypothesis that juveniles disperse beyond regular altitudinal limits (Diamond 1972, Freeman et al. 2013).

SINGING STARLING Aplonis cantoroides

This urban / agricultural species has greatly expanded its distribution in response to urbanisation of New Guinea's landscape. It was not recorded by Diamond at Karimui in 1965. However, we observed a small flock on four occasions in 2012, in a small grove of fig trees adjacent to Karimui airstrip. Local informants declared that the species had arrived within the past decade, but is only seen in the vicinity of the airstrip.

LAWES'S PAROTIA Parotia lawesii

Regularly observed by Diamond (1972), who collected six specimens on Mt. Karimui's north-west ridge. In contrast, despite many weeks of field work at appropriate elevations, our sole observation was a pair at 1,640 m in November 2012. Parotias are vocal and easily detected (BGF pers. obs.) suggesting that the species has declined in abundance along Mt. Karimui's north-west ridge since 1965.

BANDED YELLOW ROBIN Poecilodryas placens

Diamond (1972) found this species near Karimui Station, where he collected two specimens and repeatedly observed lone individuals foraging in the understorey. We did not encounter this easily mist-netted and vocally distinctive understorey species in primary

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forest at Karimui Station, nor did we detect it during opportunistic field work at lower elevations. However, we were unable to mist-net in lower elevation (<1,000 m) forest on the Karimui Plateau. Thus, it is unclear if the species persists, even patchily, in the region or is truly absent.

We also documented minor elevational records (<250 m above previously reported limits) for 12 additional species. STEPHAN'S EMERALD DOVE Chalcophaps stephani : mist-netted at 1,390 m, observed at 1,420 m camp, vs. below 1,200 m (Baptista et al. 1997). RED-CHEEKED PARROT Geoffroyus geoffroyi: to 1,240 m, vs. below 1,113 m (Diamond 1972). VARIABLE DWARF KINGFISHER Ceyx lepidus : mist-netted to 1,385 m, vs. below 1,300 m (Woodall 2001). WHITE-EARED CATBIRD Ailuroediis buccoides: mist- netted to 1,300 m, vs. below 1,200 m (Mack & Wright 1996). TAWNY-BREASTED HONEYEATER Xanthotis flaviventer. to 1,660 m, vs. below 1,500 m (Higgins et al. 2008). RUBY-THROATED MYZOMELA Myzomela eques: to 1,310 m, vs. below 1,200 m (Higgins et al. 2008). GREY -GREEN SCRUBWREN Sericornis arfakianus: to 1,780 m, vs. below 1,700 m (Gregory 2007). YELLOW-BELLIED GERYGONE Gerygone chrysogaster : to 1,030 m, vs. below 800 m (Beehler et al. 1986). GOLDENFACE Pachycare flavogriseum : to 1,920 m, vs. below 1,800 m (Boles 2007). GOLDEN CUCKOOSHRIKE Campochaera sloetii: to 1,240 m, vs. below 1,100 m (Taylor 2005). SOOTY THICKET FANTAIL Rhipidura threnothorax : to 1,240 m, vs. to 1,100 m (Boles 2006). BLACK-FRONTED WHITE-EYE Zosterops atrifrons : to 1,700 m, vs. below 1,460 m (van Balen 2008).

Discussion

Our studies confirm the high avian diversity of Mt. Karimui and the Karimui Plateau: 271 species are documented to occur, a total comparable to other extensively surveyed New Guinean elevational gradients (Freeman et al. 2013), and remarkably high given the absence of lakes, marshes and both low-elevation (<500 m) and upper montane forests (>2,500 m) in the Karimui area. Mt. Karimui's avian diversity includes many species detected during our field work but not by Diamond (1972) in 1965. Conversely, we failed to detect several species reported by Diamond (1972).

Avian community dynamics in the tropics have been seldom studied in undisturbed forests. The sparse data that exist support the hypothesis that tropical bird populations are relatively stable through time, especially among forest-dwelling insectivores (Munn 1985, Brooks 2005, Martinez & Gomez 2013). We lack quantitative data to statistically assess population changes in Mt. Karimui's avifauna. Nevertheless, several species may have undergone substantial population changes during this interval. Most obviously. Bicolored Mouse-warbler was apparently absent on Mt. Karimui's north-west ridge in 1965 but relatively common in 2012. We believe this is the most extreme example of population changes in a resident understorey tropical bird in undisturbed forest. Bicolored Mouse- warbler is patchily distributed across New Guinea and inhabits a narrow elevational zone between two more widespread congeners (Beehler et al. 1986, Freeman et al. 2013), distributional attributes that may predispose this species to local colonisations and extinctions at individual sites (Diamond 1973). Conversely, it seems that the Lawes's Parotia has almost disappeared from Mt. Karimui's north-west ridge since 1965. Examples of local colonisations and extinctions in disturbed tropical habitats are much more common (Diamond 1971), and we documented the probable recent colonisation of agricultural habitats on the Karimui Plateau by Buff-banded Rail and Singing Starling.

Distributional ecology.— Distributional data describing range limits of New Guinean birds have been previously used to test hypotheses of community assembly and

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diversification (Diamond 1973, Diamond 1986, Mack & Dumbacher 2007). We briefly comment on one well-known pattern — the tendency for closely related species to replace one another parapatrically along elevational gradients (Diamond 1986, Freeman et al. 2013). Elevational replacements occur in tropical mountains worldwide in many taxa. Understanding the ecological factors that maintain their parapatric distributions is an active arena of ecological research, focused on answering the question of why elevational gradients contain multiple closely related species that partition elevational space, instead of just one widespread species (Jankowski et al. 2012).

Flowever, the contribution of elevational replacements to tropical montane biodiversity is seldom quantified. We updated Diamond's (1972) list of elevational replacements on Mt. Karimui. Nearly all elevational replacements are congeners with similar body sizes and diets. In total, we identified 24 pairs, five trios and two quartets of elevational replacements on Mt. Karimui (71 species; Appendix 1). Why do 71 species with narrow elevational distributions exist instead of 31 more widespread species? Providing a satisfactory answer to this question is beyond our scope, but these statistics demonstrate that elevational replacements comprise a significant portion of Mt. Karimui's avian diversity. Mt. Karimui contains 238 species of forest-dwelling birds (Appendix 1), a total 20.2% higher than it would be if all 40 'ecologically redundant' elevational replacements were excluded. This coarse analysis is one of the first to explicitly quantify the contribution of elevational replacements to montane biodiversity for a taxonomic group (Terborgh & Weske 1975), strengthening the hypothesis that elucidating the evolution of elevational distributions is a key component of understanding montane biodiversity in New Guinea (Diamond 1973, 1986).

Conservation.— Conserving Mt. Karimui's diverse avifauna is a significant challenge. The principal negative impacts on bird populations result from hunting and, increasingly, forest clearance. The latter will almost certainly be the main cause of avifaunal declines in the Karimui area in the near future. Human population on the Karimui Plateau has quintupled since the early 1960s (Wagner 1967; J. Anuabo pers. comm.), with concomitant habitat loss due to both subsistence and cash-crop (e.g. coffee) agriculture, with forest clearance likely to accelerate via current plans to construct a road to Karimui. Conserving entire elevational gradients of primary forest provides watershed benefits to local communities while conserving the vast majority of montane biodiversity and providing space to accommodate climate change-driven range shifts (Laurance et al. 2011).

Bird populations may also be impacted by hunting. We did not attempt to document the impact of hunting on Mt. Karimui, but did collect several observations consistent with the hypothesis that it affects populations of several species. Subsistence hunting remains common in New Guinean cultures (Wagner 1967). We frequently encountered boys and men hunting birds with slingshots and / or bow-and-arrows, and observed numerous hunting blinds. The latter are used especially frequently (daily or near-daily) during droughts or if located near 'salt licks' where birds, especially columbids, gather to drink water and / or ingest grit or minerals (Diamond et al. 1999, Symes et al. 2006). Informants reported regularly taking large numbers (>10) of birds, principally columbids, on single visits to such sites. Lastly, it is common practice to consume eggs or nestlings on encountering an active nest, even of small (<15 g) passerines. It is probable that hunting has significantly impacted populations of certain species. For example. Dwarf Cassowary is extirpated from accessible parts of the Karimui Plateau and Mt. Karimui. Likewise, Talegalla brushturkeys are absent from forests around Karimui Station, even in large tracts of primary forest. Additionally, Papuan Eagle and Pesquet's Parrot appear to be largely absent from the Karimui Plateau, although the parrot persists on the lower slopes of Mt. Karimui, in rugged terrain near the

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Tua River, and may be increasing. These species are regularly targeted by hunters, and it is probable that their distributions are currently limited by hunting pressure.

Acknowledgements

First and foremost, we thank the many landowners in the Karimui region who enabled all aspects of this research, particularly J. Anuabo of KCRMPI, Gande, S. Banu, D. Goma, J. Buga Tane and W. Paro. Comments from I. Woxvold, B. Benz, J. Mandeville, ]. M. Diamond, K. D. Bishop and G. M. Kirwan improved this manuscript. BGF was supported by the Athena Fund of the Cornell Lab of Ornithology, the Explorer's Club and National Science Foundation GRFP 2011083591. AMCF was supported by National Geographic CRE grant 9117-12.

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Addresses: Dept, of Ecology and Evolutionary Biology, Cornell University, W257 Corson Hall, Ithaca, NY, USA; and Cornell Lab of Ornithology, 159 Sapsucker Woods Rd, Ithaca, NY, USA, e-mail: bgf27@ comell.edu

Appendix 1: Complete list of the Karimui area avifauna

Nomenclature follows IOC classification (Gill & Donsker 2013). Conservation status reflects IUCN classification (IUCN 2013), while habitat classifications are based on Beehler et al. (1986) and pers. obs. Elevational replacements (pairs, trios and quartets) are based on references (Diamond 1972, Beehler et al. 1986) and pers. obs. We note those bird species documented by our recent field work and those by Diamond (1972). We also report elevational distributions at Mt. Karimui of most forest-dwelling species. We describe high elevation limits for many species, and low elevation limits for those species whose lower limit lies above c.1,100 m. Finally, we summarise additional information pertaining to our observations as brief notes. Habitats: F = Forest, Ag = Agricultural, Aq = Aquatic. Conservation status: VU = Vulnerable, NT = Near Threatened, DD = Data Deficient.

so

G

UJ

Southern

Cassowary

o> £				\|	S
rz c u 55		J — c C c > O c	Uh y S c	J	\|
_at	7c IS	f— .H 'll at <s> £ 3 g £ 5 at Cl,	< 1 5 -2 CO O	J-H at s	t— I at G- <-■ .	CD a* o
'C cn	X	U (A UJ H	o hJ	D	z
Casuarius	F	VU pair, low				Reported by informants
casuarius						to occur near border of Chimbu and Gulf provinces
Casuarius bennetti	F	NT pair, high				Reported by informants to persist in remote forests, one captive bird in Yogoromaru village

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11	1) B		c			B	B
£ ra c J=, cd ’&b	n 3 C u £ C	n IS	.2 n3 > 1 1	“ nj C C O ft Q J 1> Oh	Ui U S < t5 O CO	T3 £ c — O )_ b a m S	\| Jx ru Cl,	CD Oi o
c w	*G CD	X	L ) Id	s a	5 °	3	z
Wattled	Aepypodius	F			X	X		Heard several times
Brushturkey	arfakianus							around 1,890 m camp
Black-billed	Talegalla	F		pair, low				Reported by informants
Brushturkey	fuscirostris							to occur below c.800 m
Collared	Talegalla jobiensis	F		pair, high	X		1,893
Brushturkey Orange-footed	Megapodius	F			X	X	1,923
Scrubfowl	reinwardt
Brown Quail	Coturnix	Ag						Reported by informants
	ypsilophora							to occur in agricultural land
Salvadori's Teal	Salvadorina	Aq	vu		X	X		Seen at Sena River
	waigiuensis						(730 m) and Tua River (550 m)

Great Egret	Ardea alba	Aq			X			Seen once at Tua River (550 m)
Pied Heron	Egretta picata	Aq				X
Little Black	Phalacrocorax	Aq			X			Seen once at Sena River
Cormorant	sulcirostris						(730 m)
Pacific Baza	Aviceda subcristata	Ag			X	X	1,203
Long-tailed	Henicopernis	F			X	X	2,263
Honey Buzzard	longicauda
Papuan Eagle	Harpyopsis novaeguineae	F	vu		X	X	1,888	Scarce
Pygmy Eagle	Hieraaetus weiskei	F			X			Seen once at 1,300 m
Chestnut¬	Erythrotriorchis	F	DD			X
shouldered Goshawk	buergersi
Doha's Goshawk	Megatriorchis doriae	F				X
Variable Goshawk	Accipiter hiogaster Ag			X	X		Seen 3^4 times around
								Karimui Station
Brown Goshawk	Accipiter fasciatus	Ag				X
Black-mantled	Accipiter	F		pair, high	X	X 1,423	2,143
Goshawk	melanochlamys
Grey-headed	Accipiter	F		pair, low	X	X	1,215
Goshawk	poliocephalus
Collared	Accipiter	Ag			X			Seen once near Karimui
Sparrowhawk	cirrocephalus							Station
Swamp Harrier	Circus	Ag			X	X		Seen once at Karimui
	approximans							airstrip
Brahminy Kite	Haliastur indus	F			X	X	2,383
Nankeen Kestrel	Falco cenchroides	Ag			X	X		Seen twice around Karimui Station
Oriental Hobby	Falco severus	F				X
Brown Falcon	Falco berigora	Ag			X	X		Seen regularly around

Karimui Station

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o>	o> £	C				E	E
E H3 c CD 'ob	n 3 C u GG — •_£ «3 G .tS 0» 45	03 £ c/l e B	-G G 2 a> ! 1 2 is	M-i U S < u-	-T3 G o £ r3	J 11 £	1 Jh 11 Cl.	w 11
c tu	mC * Gn X	O nl U tn	a» Gh S £	o CO	5	o	Cl, D	o Z
Forbes's Forest Rail	Rallicula forbesi F		pair, high	X		1,343	1,763
Red-necked Crake Rallim tricolor F		pair, low		X
Buff-banded Rail	Gallirallus Ag			X				Common around
	philippensis							Karimui Station
Lewin's Rail	Lewinia pectoralis Ag			X				Heard once near Karimui Station
Pale-vented	Amaurornis Ag			X	X			Common around
Bush-hen	moluccana							Karimui Station
Common	Actitis hypoleucos Aq			X				Seen along Tua River
Sandpiper								(550 m)
Australian	Sh/fifl isabella Ag			X	X			Seen once at Karimui
Pratincole							airstrip
Slender-billed	Macropygia F			X	X		1,903	Very common at
Cuckoo-Dove	amboinensis							lower elevations (below c. 1,500 m) and in anthropogenic landscapes
Bar-tailed	Macropygia F		largely replaces	X	X		2,520	Very common at higher
Cuckoo-Dove	nigrirostris		M. amboiensis at					elevations (above
			high elevations, but					c.1,300 m), not recorded
			significant overlap					in anthropogenic areas
Great Cuckoo-	Reinwardtoena F			X	X		2,233
Dove	reinwar dti
Stephan's	Chalcophaps F			X	X		1,423
Emerald Dove	stephani
New Guinea	Henicophaps F				X
Bronzewing	albifrons
Cinnamon	Gallicolumba F		pair, low	X	X		1,288
Ground Dove	rufigula
White-breasted	Gallicolumba F				X
Ground Dove	jobiensis
Bronze Ground	Gallicolumba F		pair, high	X	X	1,363	2,068
Dove	beccarii
Pheasant Pigeon	Otidiphaps nobilis F			X	X		1,693
Southern	Goura F	vu						Reported by elderly
Crowned Pigeon	scheepmakeri							local informants to occur near Tua River, but unclear if still present in the Karimui
								area
Wompoo Fruit	Ptilinopus F			X	X		1,033
Dove	magnificus
Pink-spotted Fruit Ptilinopus perlatus F			X	X			Regular at Sena River
Dove								salt lick (750 m)
Ornate Fruit Dove Ptilinopus ornatus F			X	X		2,520
Superb Fruit Dove Ptilinopus F			X	X		1,273
	superbus
Beautiful Fmit	Ptilinopus F			X	X		1,243
Dove	pulchellus

Ben Freeman & Alexandra M. Class Freeman

43

Bull. B.O.C. 2014 134(1)

<u	E	C				b	b
E 03 c J5 c/5 "Sc	H3 G V X ** 'X C3 c -t: i> X	H3 £ a; t/5 G \|	cc3 rtj C C ■ E S \| S It &-	BGF/AMCF	T3 e O E C33	1 i-H QJ 2	E h, a i O.	C/5 11 o
c uu	CD X	~tr>	E 2	5	o hJ	3	z
White-bibbed Fruit Dove	Ptilinopus rivoli F			X	X	1,243	2,105
Dwarf Fruit Dove	Ptilinopus minus F			X	X			Regular at Sena River salt lick (750 m)
Purple-tailed Imperial Pigeon	Ducula rufigaster F		pair, low	X	X		1,283
Rufescent	Ducula chalconota F		pair, high	X		1,793	2,272
Imperial Pigeon Zoe's Imperial Pigeon	Ducula zoeae F		X	X		2,083


Papuan Mountain Gymnophaps F			X	X		2,333
Pigeon	albertisii
Palm Cockatoo	Probosciger F aterrimus			X	X		1,283	Scarce
Sulphur-crested Cockatoo	Cacatua galerita F			X	X		2,373
Pesquet's Parrot	Psittrichas fulgidus F	vu		X	X		1,903
Red-breasted	Micropsitta F			X		1,453	2,133
Pygmy Parrot	bruijnii
Yellowish-	Chalcopsitta F				X
streaked Lory	scintillata
Dusky Lory	Pseudeos fuscata F			X	X		1,653
Coconut Lorikeet	Trichoglossus F haematodus			X	X		1,423
Goldie's Lorikeet	Psitteuteles goldiei F			X			1,933
Black-capped Lory	Lorius lory F			X	X		1,508
Striated Lorikeet	Charmosyna F			X				Seen at 1,420 m camp
	multistriata							only
Pygmy Lorikeet	Charmosyna F wilhelminae			X			1,933
Red-flanked	Charmosyna F			X	X		1,323
Lorikeet	placentis
Fairy Lorikeet	Charmosyna F pulchella			X	X	1,323	1,961
Josephine's	Charmosyna F			X				Possibly seen once at
Lorikeet	josefime							990 m, but requires confirmation.
Papuan Lorikeet	Charmosyna papou F		replaces other	X	X			Seen twice, at 1,735 and
			Charmosyna at high elevations					1,910 m
Plum-faced	Oreopsittacus F			X	X	1,943	2,520
Lorikeet	arfaki
Yellow-billed	Neopsittacus F				X
Lorikeet	musschenbroekii
Brehm's Tiger Parrot	Psittacella brehmii F			X	X	1,765	2,235
Madarasz's Tiger	Psittacella F			X	X			Seen twice, at 1,820 and
Parrot	madaraszi							1,910 m

Ben Freeman & Alexandra M. Class Freeman

44

Bull. B.O.C. 2014 134(1)

a >	o> a		e				E	E
e t « c 4= CD 'sb	C u 2 c .OS	15	O £ CD g \|	2 QJ O s • S Ol \|S a > Ci*	u s < E O cn	xs c o E	1 J-H a> s	E QJ Cl. Cl,	CD a* Q
c UJ	’0 cn	X	"cD	E 5	5	o hJ	D	z
Red-cheeked	Geoffroyus	F		pair, low	X	X		1,243
Parrot	geoffroyi
Blue-collared Parrot	Geoffroyus simplex	F		pair, high	X	X	1,033	1,953
Eclectus Parrot	Eclectus rorntus	F			X	X		1,393
Papuan King	Alisterus	F			X	X		1,593
Parrot	chloropterus
Orange-breasted	Cyclopsitta	F		pair, low	X	X		1,108
Fig Parrot	gulielmitertii
Double-eyed Fig	Cyclopsitta	F		pair, high	X		1,163	1,243
Parrot	diophthalma
Large Fig Parrot	Psittaculirostris desmarestii	F			X	X		1,243
Ivory-billed	Centropus menbeki F			X	X		1,383
Coucal
Pheasant Coucal	Centropus	Ag			X				One record from near
	phasianinus							Karinuii Station
Dwarf Koel	Microdynamis pawn	F			X	X		1,321
Pacific Koel	Eudynamys orientnlis	F			X	X		2,123
Rufous-throated	Chrysococcyx	F		pair, high	X		1,793	2,520
Bronze Cuckoo	ruficollis
White-eared	Chrysococcyx	F		pair, low	X	X		1,813
Bronze Cuckoo	meyerii
White-crowned	Cacomantis	F			X	X		2,520
Cuckoo	leucolophus
Chestnut-breasted	Cacomantis	F		pair, low	X	X		1,658	Possibly to higher
Cuckoo	castaneiventris								elevations (overlap unclear, Fan-tailed Cuckoo vocally similar)
Fan- tailed Cuckoo	Cacomantis	F		pair, high	X		1,763	2,520	Possibly to lower
	flabelliformis								elevations (overlap unclear, Chestnut¬ breasted Cuckoo vocally similar)
Brush Cuckoo	Cacomantis variolosus	Ag			X	X			Common in gardens
Oriental Cuckoo	Cuculus optatus	F			X				Seen at Tua River (550 m)
Greater Sooty	Tyto tenebricosa	F			X				Heard regularly at
Owl									1,420 m camp
Papuan Boobook	Ninox theomacha	F			X	X		2,520
Marbled	Podargus ocellatus	F			X	X		1,233
Frogmouth Papuan	Podargus	F			X	X		1,233
Frogmouth	papuensis
White-throated	Eurostopodus	Ag				X
Nightjar	mystacalis
Papuan Nightjar	Eurostopodus	F			X				Seen at Sena River
	papuensis								(730 m)

Ben Freeman & Alexandra M. Class Freeman

45 Bull. B.O.C. 2014 134(1)

<U	E		c				%	£
E rz C	rz e		.2 rz	— c e £	tu U . — i	-a	1	I
-C C/1 "5b	vC c _a»	55 IE	> i-t & § 1	O c •J5 a» 2 is CD Cu	2 < O CD	c o E rz	QJ 2	}— i HI o. 0-1	CD CD ©
e UJ	"c-i c n	X	"55	B E	5	o	3	z
Feline	Aegotheles insignis	F			X	X		1,893	Vocalisations attributed
Owlet-nightjar									to this species heard at 1,420 and 1,910 m
									camps
Wallace's Owlet-nightjar	Aegotheles wallacii	F	DD			X			Aegotheles sp. heard below 1,500 m could
								not be identified to species

Mountain	Aegotheles albertisi F		replaces A. wallacii	X			2,520	Aegotheles sp. heard at
Owlet-nightjar				and A. bennettii at					2,520 m presumed to
				high elevations					be this species based on elevation
Barred	Aegotheles	F				X			Aegotheles sp. heard
Owlet-nightjar	bennettii								below 1,500 m not identified
Moustached	Hemiprocne	F			X	X		1,253
Treeswift	mystacea
Glossy Swiftlet	Collocalia esculenta	F			X	X		2,520
Mountain Swiftlet	Aerodramus	F			X	X			Commonly seen
	hirundinaceus								around Karimui Station
Oriental	Eurystomus	Ag			X	X		1,243
Dollarbird	orientalis
Hook-billed	Melidora	F			X	X		1,873
Kingfisher	macrorrhina
Shovel-billed Kookaburra	Clytoceyx rex	F				X
Rufous-bellied Kookaburra	Dncelo gaudichaud	F			X	X		1,283
Forest Kingfisher	Todirnmphus macleayii	Ag				X
Sacred Kingfisher	Todiramphus	Ag			X	X			Commonly seen
	sanctus								around Karimui Station
Yellow-billed Kingfisher	Syma torotoro	F		pair, low	X	X		1,233
Mountain	Syma	F		pair, high	X	X	1,493	2,158
Kingfisher	megarhyncha
Variable Dwarf	Ceyx lepidus	F			X	X		1,388
Kingfisher Azure Kingfisher	Ceyx azureus	Aq			X	X			Seen once at Tua River (550 m)
Rainbow	Merops ornatus	Ag			X	X			Seen twice near
Bee-eater							Karimui Station
Blyth's Hombill	Rhyticeros plicatus F			X	X			Seen on 3-4 occasions
Red-bellied Pitta	Erythropitta erythrogaster	F			X	X		1,198
White-eared	Ailuroedus	F		pair, low	X	X		1,303
Catbird	buccoides
Spotted Catbird	Ailuroedus melanotis	F		pair, high	X	X	1,363	1,703

Ben Freeman & Alexandra M. Class Freeman

46

Bull. B.O.C. 2014 134(1)

QJ	<u E		G					£	S
£ 03 c on 'ob	c u vB C	IS	O m £ o> CD G	CD G	evational placement	BGF/AMCF	-a c o E	1 J-H a» s	£ a» Gm r\	CD a» Q
c w	"0 CD	X	o u	CD	3 21	5	o hJ	D	z
MacGregor's	Amblyornis	F				X	X	1,693	2,243
Bowerbird	macgregoriae
White-shouldered	Malurus	Ag				X	X		1,253
Fairy wren	alboscapulatus
Orange-crowned	Clytomyias	F				X	X			Family groups seen at
Fairywren	insignis									2,160 m and 2,350 m
Ruby-throated Myzomela	Myzomela eques	F				X	X		1,313
Red Myzomela	Myzomela	F			trio, middle	X	X		1,423	Lower elevation limit
cruentata									unclear (not observed below 1,300 m)

Papuan Black Myzomela	Myzomela nigrita	F			trio, low	X	X		1,243
Red-collared	Myzomela	F			trio, high	X	X	1,273	2,520
Myzomela	rosenbergii
Green-backed Floneyeater	Glycichaera fallax	F				X			1,198
Yellowish-	Ptiloprora	F				X				Seen twice at flowering
streaked Honeyeater	meekiana									tree at 1,880 m
Rufous-backed	Ptiloprora guisei	F				X	X	1,783	2,520
Honeyeater Plain Honeyeater	Pyenopygius	F			pair, low	X	X			Seen once at 1,010 m
	ixoides									near Karimui Station
Marbled	Pyenopygius	F			pair, high	X	X			Seen once at 1,420 m
Honeyeater	cinereus									camp
Spotted	Xanthotis	F				X	X		1,363
Honeyeater	polygrammus
Tawny-breasted	Xanthotis	F				X	X		1,663
Honeyeater	flaviventer
Meyer's Friarbird	Philemon meyeri	F					X
New Guinea	Philemon	F				X	X		1,243
Friarbird	novaeguineae
Long-billed	Melilestes	F				X	X		1,633
Honeyeater	megarhynchus
Common Smoky	Melipotes	F				X	X	1,338	2,520
Honeyeater	fumigatus
Olive Straightbill	Timeliopsis fulvigula	F				X	X	1,633	2,063
Black-throated	Caligavis	F			pair, high	X	X	1,423	2,520
Honeyeater	subfrenata
Obscure Honeyeater	Caligavis obscura	F			pair, low	X	X		1,243
Yellow-browed	Melidectes	F			pair, high	X	X	1,338	2,520
Melidectes	rufocrissalis
Ornate Melidectes	Melidectes torquatus	F			pair, low	X	X	1,333	1,888
Mottle-breasted	Meliphaga	F			pair, low	X	X		1,313
Honeyeater	mimikae
Mountain	Meliphaga	F			pair, high	X	X	1,423	1,883
Honeyeater	orientalis

Ben Freeman & Alexandra M. Class Freeman

Bull. B.O.C. 2014 134(1)

47

Ol	at £		c				B
B PC C -C C/3 15b	re C u vC C	nj 15	.2 ~ c pc e c > o c fc- Ot Ot <*, Ot c 3 > -2	BGF/AMCF	T3 c o B n:	1 1) s	\| J-H at C u p	C/5 at ©
c UJ	"C CD	X	o Jx rr* U « U k<	5	o _J	D	z
Scrub Honeyeater	Meliphaga	Ag		X	X			Common around
	albonotata							Karimui Station
Mimic Honeyeater Meliphaga analoga	F		X	X		1,163
Yellow-gaped	Meliphaga	F			X
Honeyeater	flavirictus
Puff-backed	Meliphaga	F		X	X		1,158
Honeyeater	aruensis
Rusty	Crateroscelis	F	trio, low	X	X		1,773
Mouse-warbler	murina
Bicolored	Crateroscelis	F	trio, middle	X		1,623	1,943
Mouse-warbler	nigrorufa
Mountain	Crateroscelis	F	trio, high	X	X	1,873	2,520
Mouse-warbler	rohusta
Pale-billed	Sericornis	F	quartet, lowest	X	X		1,513
Scrubwren	spilodera
Papuan	Sericornis	F	quartet, highest	X	X	1,943	2,520
Scrubwren	papuensis
scrubwren sp.	Sericornis sp.	F		X				See species account
Large Scrubwren	Sericornis nouhuysi	F		X	X	1,473	2,426
Buff-faced	Sericornis	F	quartet, high	X	X	1,703	2,013
Scrubwren	perspicillatus		middle
Grey-green	Sericornis	F	quartet, low middle X	X	1,378	1,783	One possible record
Scrubwren	arfakianus							from 1,200 m near Bosiamaru
Brown-breasted	Gerygone ruficollis	F		X	X	1,447	2,380
Gerygone Large-billed	Gerygone	F		X				Seen/heard at Sena
Gerygone	magnirostris							River (730 m)
Yellow-bellied	Gerygone	F		X	X		1,033
Gerygone	chrysogaster
Ashy Gerygone	Gerygone cinerea	F		X	X	2,515	2,520
Green-backed	Gerygone	F		X	X		1,383
Gerygone	chloronota
Fairy Gerygone	Gerygone palpebrosa	F		X	X		1,243
Goldenface	Pachycare flavogriseum	F		X	X		1,923
Loria's Satinbird	Cnemophilus loriae F	pair, low	X	X	1,423	2,428
Crested Satinbird	Cnemophilus	F	pair, high	X	X			One male seen at ridgi
	macgregorii							summit (2,520 m)
Yellow-breasted	Loboparadisea	F	NT	X	X	1,433	1,933
Satinbird	sericea
Black Berrypecker Melanocharis	F	pair, low	X	X		1,463
	mgr a
Fan-tailed	Melanocharis	F	pair, high	X	X	1,388	2,520
Berrypecker	versteri
Streaked	Melanocharis	F		X	X	1,453	1,873
Berrypecker	striativentris

Ben Freeman & Alexandra M. Class Freeman

48 Bull. B.O.C. 2014 134(1)

at	at 6		G					B	B
£ n3 c	rs G Ot •LG		O "Ss at CD G		-G G 2 at § i	U-i U s	-a s	1	1
P— CD '5b	G a*	H3 is	CD G	1 Js at Oh	< £5 O CD	o s	i-i at s	H aj &. p	CD at o
c uu	'C CD	X	u	CD	E £	5	o hJ	P	z
Spotted	Rhamphocharis	F				X	X	2,103	2,323
Berry pecker	crassirostris
Dwarf Longbill	Oedistoma iliolophus	F				X	X		1,653
Pygmy Longbill	Oedistoma pygmaeum	F				X	X		1,288
Slaty-headed	Toxorhamphus	F				X	X		1,993
Longbill	poliopterus
Tit Berrypecker	Oreocharis arfaki	F				X	X	1,338	2,520
Crested	Paramythia	F				X	X			Flock seen just below
Berrypecker	montium									summit of ridge (2,49( m)
Spotted	Ptilorrhoa	F			trio, high	X	X	1,508	2,142
Jewel-babbler	leucostida
Blue	Ptilorrhoa	F			trio, middle	X				Heard at Tua River
Jewel-babbler	caerulescens									(550 m)
Chestnut-backed	Ptilorrhoa	F			trio, low	X	X		1,458	Lower elevation limit
Jewel-babbler	castanonota									not determined
Yellow-breasted	Machaerirhynchus	F			pair, low	X	X		1,308
Boatbill	flaviventer
Black-breasted	Machaerirhynchus	F			pair, high	X	X	1,243	2,393
Boatbill	nigripedus
Lowland Peltops	Peltops blainvillii	F			pair, low		X
Mountain Peltops	Peltops montanus	F			pair, high	X	X		2,105
Black Butcherbird	Cracticus quoyi	F				X	X		1,508
Hooded Butcherbird	Cradicus cassicus	F				X	X		1,333
Great	Artamus maximus	F				X	X			Common around
Woodswallow Black-faced	Coracina	Ag					X			Karimui Station
Cuckooshrike	novaehollandiae
Stout-billed	Coracina	F				X	X		J, 593
Cuckooshrike	caeruleogrisea
Boyer's Cuckooshrike	Coracina boyeri	F					X
Common	Coracina	Ag				X				Vocalising bird seen
Cicadabird	tenuirostris									near Karimui Station
Black-shouldered	Coracina incerta	F				X				Single seen on north
Cicadabird										slope of Mt. Karimui (1,250 m)
Grey-headed	Coracina	F			pair, low	X	X		1,363
Cuckooshrike	schisticeps
Black Cicadabird	Coracina melas	F				X	X			Pair seen at Tua River (550 m)
Black-bellied Cuckooshrike	Coracina montana	F			pair, high	X	X	1,338	2,303
Golden	Campochaera	F				X	X		1,243
Cuckooshrike	sloetii
Varied Triller	Lalage leucomela	F				X	X		1,423

Ben Freeman & Alexandra M. Class Freeman 49

o»	OJ £		c				E	£
£ rs e JZ c/5 "ab	re c u £ ’£ u	"re IS re	H3 > £ C/5 1 1	1= C “ nj § E 11 U It D-	u* U < b O CO	-a e o £ re	1 L* 1> g	\| hi O) c u p ,
G UJ	’0	X		E S	s	o ■— I	D
Varied Sittella	Daphoenositta chrysoptera	F				X
Mottled Whistler	Rhagologus leucostigma	F			X	X	1,323	2,520
Wattled	Eulacestoma	F			X	X	1,913	2,263
Ploughbill	nigropectus
Rufous-naped	Aleadryas	F			X	X	1,303	2,398
Whistler	rufinucha
Crested Pitohui	Ornorectes cristatus	F			X	X		1,353
Black Pitohui	Melanorectes	F		pair, high (with	X	X	1,573	2,453
	nigrescens			Little Shrikethrush)
Rusty Whistler	Pachycephala hyperythra	F		trio, low	X	X		1,353
Brown-backed	Pachycephala	F			X	X	1,810	2,220
Whistler	modesta
Grey Whistler	Pachycephala simplex	F			X	X		1,463
Sclatehs Whistler	Pachycephala s oror	F		trio, middle	X	X	1,243	1,913
Regent Whistler	Pachycephala schlegelii	F		trio, high	X	X	1,753	2,520
Black-headed	Pachycephala	Ag			X	X
Whistler	monacha
Rusty Pitohui	Pseudorectes ferrugineus	F			X	X		1,143
Little Shrikethmsh Colluricincla	F		pair, low (with	X	X		1,753
	megarhyncha			Black Pitohui)
Long-tailed Shrike Lanius schach	Ag			X
Southern Variable	Pitohui uropygialis F		pair, low	X	X		1,231
Pitohui Hooded Pitohui	Pitohui dichrous	F		pair, high	X	X		1,658
Brown Oriole	Oriolus szalayi	F			X	X		1,443
Pygmy Drongo	Chaetorhynchus papuensis	F			X	X		1,713
Spangled Drongo	Dicrurus bracteatus	F			X	X		1,273
Willie Wagtail	Rhipidura leucophrys	Ag			X	X
Northern Fantail	Rhipidura rufiventris	F			X	X		1,423
Sooty Thicket	Rhipidura	F			X	X		1,243
Fantail	threnothorax
White-bellied	Rhipidura	F			X	X
Thicket Fantail	leucothorax
Black Fantail	Rhipidura atra	F			X	X	1,241	2,520
Chestnut-bellied	Rhipidura	F		pair, low	X	X		1,658
Fantail	hyperythra

Bull. B.O.C. 2014 134(1)

w

u

o

z

Common around Karimui Station

Regular at Karimui airstrip

Lower elevation limit not determined

Common around Karimui Station

Seen once near Karimui Station

Ben Freeman & Alexandra M. Class Freeman

50

Bull. B.O.C. 2014 134(1)

£		c						j	3
R3 c u		o 5>		re C o	c a» s	Ph U s	*T3 c	s	6
	%	OJ	t/1 £		o	<	o
c u	s	J) c	03 >	Q3	up	£ 03	a* £	a» Oh
□	n3	o			D-,	O	o	Oh
c n	X	u	CD	UJ	(U J-H	CO	5		P

Friendly Fantail	Rhipidura albolimbata	F pair, high	X	X	1,633	2,520
Dimorphic Fantail Rhipidura	F	X	X	1,573	2,520
	brachyrhyncha
Rufous-backed	Rhipidura	F		X
Fantail	rufidorsa
Black Monarch	Symposiachrus axillaris	F	X	X	1,188	1,913
Spot-winged	Symposiachrus	F	X	X		1,183
Monarch	guttula
Black-winged Monarch	Monarcha frater	F	X	X		1,623
Golden Monarch	Carterornis chrysomela	F	X	X		1,193
Frilled Monarch	Arses telescopthalmus	F	X	X		1,273
Torrent-lark	Grallina bruijnii	Aq	X	X		1,213
Grey Crow	Corvus tristis	F	X	X		1,233
Lesser Melampitta Melampitta	F	X		1,943	2,520
	lugubris
Blue-capped Ifrit	Ifrit a kowaldi	F	X	X	1,793	2,356
Crinkle-collared	Manucodia	F	X	X		1,433
Manucode	chalybatus
Trumpet	Phonygammus	F	X	X		1,423
Manucode	keraudrenii
Short-tailed	Paradigalla	F	X	X
Paradigalla	brevicauda
Princess	Astrapia	F	X	X	1,713	2,520
Stephanie's Astrapia	stephaniae
Lawes's Parotia	Parotia lawesii	F	X	X
King of Saxony	Pteridophora	F	X	X	1,893 ,	2,520
Bird-of-paradise	alberti
Superb Bird-of-	Lophorina superba	F	X	X	1,283	1,982
paradise Magnificent Riflebird	Ptiloris magnificus	F	X	X		1,473
Black Sicklebill	Epimachus fastosus	F VU	X	X	1,683	2,520
Black-billed	Drepanornis	F	X	X
Sicklebill	albertisi
Magnificent	Diphyllodes	F	X	X		1,709
Bird-of-paradise	magnificus
King Bird-of- paradise	Cicinnurus regius	F	X	X
Raggiana Bird-of-	Paradisaea	F	X	X		1,623
paradise	raggiana
Blue Bird-of-	Paradisaea	F VU		X
paradise	rudolphi

o

Z

Seen once at 1,820 m

Pair seen once at 1,640 m

One mist-netted at 1,710 m

Singles at Sena (730 m) and Tua (550 m) Rivers

Ben Freeman & Alexandra M. Class Freeman

51

o>	i U B		c			B	B
£ n3 c -C err	ci C u vC c		.o ra £ OJ cx	C 2 ai § E o» ■ss ^	u. U s <	T3 E c — O t.	J >-i
Tab		£	o fS	> J2	tu o CD	B ¥ n >	Cl,
c w	KJ CD	X	"So	UJ £	5 2	5-*h P
Banded Yellow	Poecilodnjas	F	NT			X
Robin	placens
Black-throated	Poecilodnjas	F			X	X 1,703	2,520
Robin	albonotata
White- winged	Peneothello	F		quartet, highest	X	X 2,333	2,520
Robin	sigillata
Slaty Robin	Peneothello cyanus	F		quartet, high middle	X	X 1,673	2,398
White-rumped	Peneothello	F		quartet, lowest	X	X	1,283
Robin	bimaculata
White-faced Robin Tregellasia leucops	F			X	X 1,198	1,718
White-eyed Robin Pachycephalopsis	F		quartet, low middle X	X 1,218	1,698
	poliosoma
Torrent Flyrobin	Monachella muelleriana	F			X	X
Canary Flyrobin	Microeca papuana	F		trio, high	X	X 1,763	2,520
Yellow-legged	Microeca	F		trio, middle	X	1,093	1,423
Flyrobin	griseoceps
Olive Flyrobin	Microeca flavovirescens	F		trio, low	X	X	1,313
Garnet Robin	Eugerygone rubra	F			X	X 1,753	2,333
Northern Scrub	Drymodes	F			X	X	1,393
Robin	superciliaris
Lesser Ground	Amalocichla	F			X	X 1,794	2,105
Robin	incerta
Island Leaf	Phylloscopus	F			X	X 1,321	1,961
Warbler	maforensis
Australian Reed	Acrocephalus	Ag			X
Warbler	australis
Papuan Grassbird	Megalurus	Ag			X	X
	macrurus
Black-fronted	Zosterops minor	F			X	X	1,703
White-eye Singing Starling	Aplonis cantoroides	Ag			X
Yellow-faced Myna	Mino dumontii	F			X	X	1,353
Russet-tailed Thrush	Zoothera heinei	F			X	X 1,473	1,643
Pied Bush Chat	Saxicola caprata	Ag				X
Red-capped	Dicaeum	F			X	X	1,723
Flowerpecker	geelvinkianum
Black Sunbird	Leptocoma sericea	Ag				X
Blue-faced Parrotfinch	Erythrura trichroa	F			X	X 1,493	2,313
Streak-headed	Lonchura	F			X
Mannikin	tristissima
Hooded Mannikin Lonchura	Ag			X	X

spectabilis

Bull. B.O.C. 2014 134(1)

z

Seen at Sena (730 m) and Tua (550 m) Rivers

Present around Karimui airstrip

Common around Karimui Station

Flock regular at Karimui airstrip

Seen twice around Karimui Station

Common around Karimui Station

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Bull. B.O.C. 2014 134(1)

On the identification and provenance of some early specimens of grasswrens (Maluridae: Amytornis) and their significance for taxonomy and nomenclature

by Andrew B. Black , Justin J. F. J. Jansen, Steven D. van der Mije &

Clemency T. Fisher

Received l August 2013

Summary. — The identity and provenance of four 19th-century Amytornis grasswren specimens in the Naturalis Biodiversity Center (Leiden) collection are reviewed. Three identified as Thick-billed Grasswren Amytornis modestus inexpectatus enable a revised diagnosis for the extinct subspecies from New South Wales. One of these and one Striated Grasswren A. striatus were acquired from John Gould in, or soon after, December 1840. The other two came via the Frank dealership in 1858 and 1873, but are probably also from Gould's collections. Leiden's A. striatus specimen and another in Philadelphia are identified here as paralectotypes of Dasyornis [ =Amytornis ] striatus Gould, 1840. Evidence is presented that Gould's brother- in-law, Charles Coxen, collected both A. modestus and A. striatus, including type material of the latter, before Gould visited Australia in 1838.

While investigating morphological and genetic diversity within Western Amytornis textilis (Quoy & Gaimard, 1824) and Thick-billed Grasswrens A. modestus (North, 1902) (Black et al. 2010, Black 2011a, Austin et al. 2013) just one specimen of the extinct far eastern (New South Wales) subspecies A. m. inexpectatus (Mathews, 1912) was found in an Australian museum collection and it proved uninformative genetically. Following an enquiry concerning early grasswren specimens in collections outside Australia, it became apparent that four mounted 19th-century specimens were housed in the Naturalis Biodiversity Center (hereafter NBC, formerly Rijksmuseum voor Natuurlijke Historie, RMNH), Leiden, the Netherlands. Three were listed as A. textilis (i.e. either A. textilis or A. modestus, see below) and one as Striated Grasswren A. striatus. Documentation at NBC suggested that at least two of the 'A. textilis' specimens were from New South Wales and might be A. m. inexpectatus. It was also evident that a specimen of each species ('textilis' and striatus) had been sent to RMNH by John Gould, presumably from his own collection. Gould (1840) was the author of A. striatus but, while he collected just one specimen himself, he referred to other specimens from New South Wales (Gould 1848) and claimed to have collected both species on the lower Namoi River, although Schodde (1982) doubted that A. striatus could have been taken there, citing a lack of suitable habitat. The NBC specimens have the potential to contribute to unresolved questions of grasswren taxonomy and nomenclature, including a reappraisal of the phenotype of A. m. inexpectatus and of the type material and type locality of A. striatus.

Methods

Known specimens of A. m. inexpectatus were examined by ABB at the Academy of Natural Sciences, Philadelphia (ANSP) (Gould's collection) on 8 April 2013 and at the American Museum of Natural History, New York (AMNH) (Mathews' collection) on 10 April 2013. Gould's two A. striatus specimens were also examined at ANSP and the four

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NBC specimens were examined at NBC, Leiden, on 16 April 2013. Measurements taken were: bill = length (total culmen to skull insertion) * depth (at level of frontal feathering), wing = flattened chord and tail = central rectrices from tip to base.

Results

NBC specimens. — RMNH.AVES. 172018. Label: 'Amytis striatus [crossed out] Malum s [crossed out] textilis Gould auct pi Uranie Zool. pi 67 f 2. Australie.' Under the socle (base of stand): 'Amytis textilis Gould auct Malurus textilis Uranie Zool. PI 67 f 2 Australie.' On examination, an A. modestus mount in good condition, female with bright flank patches; slightly convex lower mandible, rather dark and heavily streaked on throat and upper breast for modestus. Bill 12.5 x 5.1 mm, wing 57 mm, tail 80.2 mm (Fig. 1). RMNH.AVES.172019. Label: 'Amytis textilis S (Quoy & Gaim) Frank 1873 N. S. Wales.' Under the socle: 'mas Frank 1873 N. S. Wales.' On examination, a male A. modestus mount showing moderate disturbance to plumage, similarly dark and relatively streaked below for modestus, biconvex bill profile. Bill 12.4 x 5.9 mm, wing 64 mm, tail 84.8 mm (Fig. 2). RMNH.AVES. 172020. No label. Under the socle 'Frank 1858.' On examination, a male A. modestus mount showing slight disturbance to plumage; similar to RMNH.AVES.172018 and 172019, moderately convex lower mandible. Bill 12.3 * 5.4 mm, wing 58 mm, tail 80.2 mm (Fig. 3). RMNH. AVES. 172021. Label: 'Amytis striata (Gould) Avant 1850 Australie.' Under socle: 'Amytis striatus Gould auct pi textilis pi 67 f 2 Lesson. Australie.' On examination, an A. striatus mount in good condition with rufous underparts merging laterally into brighter flanks, and thus female. Bill 12.8 x 4.2 mm, wing 63 mm, tail 89.4 mm (Fig. 4).

ANSP and AMNH specimens of A. m. inexpectatus. — NNSV 16887. Label: 'Gould Coll S Amytis textilis (Quoy et Gaim.) New South Wales T. B. Wilson.' Male, bill 13.6 x 5.4 mm, wing 62 mm, tail 82.5 mm. ANSP 16888. Label: 'Gould Coll $ Amytornis textilis (macrourus [szc]) New South Wales Thos. B. Wilson.' Female, bill 12.2 x 5.4 mm, wing 61 mm, tail 87.2 mm. ANSP 16889. Label: 'Rivoli [sic] Gould Coll Thos. B. Wilson.' Male, bill 13.6 x 5.6 mm, wing 62 mm, tail 79.5 mm. AMNH 598073. Labels. 'Diaphorillas textilis inexpectatus S New South Wales TYPE Mathews 1912 etc.' Male, bill 12.6 x 5.0 mm, wing 63 mm, tail 80.3 mm. AMNH 598072. Labels. 'Diaphorillas — ROTH Exc GMM juvenile S New South Wales.' Male, bill 13.5 x 5.4 mm, wing 63 mm, tail 91.2 mm. All the specimens above are relatively dark for the species and possess moderate to heavy underparts streaking.

ANSP specimens of A. striatus.— ANSP 16890. Labels. 'Gould Coll TYPE <$ Amytis striatus (Gould) New South Wales T. B. Wilson. 379 S N S Wales 16890 Dasyornis striatus Gld. PZS. 1839. P 143 etc.' Male, bill 12.3 x 4.9 mm, wing 60 mm, tail missing. ANSP 16891. Labels. 'Gould Coll TYPE $ Amytis striatus (Gould) New South Wales T. B. Wilson. This has no type sig.' Female, bill 13.0 x 4.9 mm, wing 63 mm, tail 88.4 mm.

Discussion

Identification and provenance of the 'A. textilis' specimens. — RMNH.AVES.172018 appears to have first been identified as Amytis striatus and later as A. textilis. While these two species are very distinct, their identification was not made consistently by all early workers (see below). RMNH. AVES. 172018-020 are all identified here as Amytornis modestus. Until the latter was described (North 1902), all such specimens were generally included under A. textilis. Failure to recognise A. textilis and A. modestus as separate (but sister) species commenced with Gould himself (1848, 1865) who believed that grasswrens he collected in north-eastern New South Wales were the same species, A. textilis, as described by Quoy and Gaimard (1824) from Shark Bay, Western Australia, on the opposite side of the

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Bull. B.O.C. 2014 134(1)

l&CO .

-Si *

Figure 1. RMNH Aves 172018: female Amytornis modestus inexpectatus, from the collection of John Gould 1840-41, the plains bordering the lower Namoi, northern New South Wales (Justin J. F. J. Jansen)

Figure 2. RMNF1 Aves 172019: male Amytornis modestus inexpectatus, from Frank, dealers, 1873, New South Wales (Justin J. F. J. Jansen)

Figure 3. RMNH Aves 172020: male, probable Amytornis modestus inexpectatus, from Frank, dealers, 1858, provenance uncertain (Justin J. F. J. Jansen)

Figure 4. RMNH Aves 172021: female Amytornis striatus striatus, from the collection of John Gould 1840-41, probably collected by Charles Coxen; paralectotype of Dasyornis striatus Gould, 1840, Liverpool Plains, northern New South Wales (Justin J. F. J. Jansen)

continent. North (1902) subsequently described Amytis modesta from central Australia and included Gould's New South Wales birds in his new species (see Fig. 5).

Despite much instability in the taxonomy of this genus in ensuing years, our present understanding (Black et al. 2010) is as follows. Briefly, A. textilis is darker, more heavily streaked and has a longer tail and more slender bill profile than A. modestus but there is variation in all of these characters among populations of both species (Black 201 la, b).

As noted above, the three NBC specimens are relatively dark and heavily streaked for A. modestus and closely resemble the ANSP and AMNH specimens of the New South Wales subspecies A. m. inexpectatus. Mathews (1912) first listed New South Wales birds, including Gould's specimens, as Diaphorillas textilis inexpectatus, but subsequently (Mathews 1922-

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0 modestus -f- O mdulkanna o obscurior O raglessi o curnamona O 'eyre'

Q inexpectatus f

Figure 5. Map showing distributional records of Western Grasswren Amytornis textilis and Thick-billed Grasswren A. modestus. Subspecies are indicated by individual colours; t signifies extinct subspecies. 'Q&G 1824' represents the type locality otMolurus textilis, Shark Bay, Western Australia (WA); 'G 1840 *' shows an approximate type locality of Dnsyornis striatus, Liverpool Plains, New South Wales (NSW), east of the lower Namoi, where Gould collected 'A. textilis'; 'N 1902' represents type localities of Amytis modesta in central Australia (CA); 'M 1912' represents the probable type locality of Diaphorillas textilis inexpectatus; ‘A. t. myall' indicates the population of Western Grasswren occurring on the Eyre Peninsula and in the Gawler Ranges, South Australia (SA) (modified from Austin et al. 2013).

1923) elevated eastern populations to species status, distinguishing them from the much paler isolate in north-western New South Wales, which he included in A. modestus. He also included birds from the Gawler Ranges, South Australia, in D. inexpectatus under the trinomial D. i. myall (= A. textilis myall) (Fig. 5). By combining inexpectatus with a subspecies of Western Grasswren A. textilis, Mathews drew attention to its relatively dark and heavily streaked plumage compared to more typical A. modestus. A further attribute of the NBC specimens is that each has the tail >80 mm, thus within the range of A. m. inexpectatus, as shown, but above that for all other A. modestus subspecies apart from some specimens of the extinct nominate subspecies from central Australia (Black 2011a). Given that these three specimens are also relatively dark and heavily streaked for A. modestus an alternative is that they might be A. textilis. However, RMNH.AVES. 172019 is documented as being from New South Wales and RMNH.AVES. 172018 was acquired from Gould (see below) and therefore by implication is also from New South Wales. Furthermore, no specimens of A. textilis are known to have been taken in Western Australia between Quoy and Gaimard's visit and the late 1890s (Black 2011b) and, while some were collected in the Gawler Ranges, South Australia, in the 1870s (ABB unpubl.), this is too late to account for RMNH.AVES.172020.

Gould (1848) stated that he had 'killed and dissected many examples' of 'A. textilis' from New South Wales. Three of his specimens are in ANSP (as above); one went to the British Museum (BMNH 41.2.1496), where it was mounted (Sharpe 1883) but has not survived; another is identified here (RMNH.AVES. 172018) and it is possible that RMNH. AVES. 172019-020 also came from Gould, since he disposed of many specimens via the

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Amsterdam-based (later London-based) Frank family business. Documents reveal that many Australian bird specimens entered the Leiden collection between c.1831 and 1876 via the Frank agency, including known specimens from Gould's collections as well as the 1858 grasswren (RMNH.AVES. 172020) (JJFJJ pers. data), but details of their origins and collectors are imprecise or lacking, and we have been unable to locate any documents that show from where the Frank dealership acquired them.

Diagnosis of A. modestus inexpectatus.— This subspecies from eastern New South Wales is distinguished by its heavily streaked underparts from the central Australian nominate, whose underparts are barely streaked, and from all other subspecies of A. modestus by tail length (range 79.5-91.2 mm, compared to 63.8-78.8 mm in other subspecies and 74.1-83.6 mm in the nominate).

Identification and provenance of the A. striatus specimen.— The fourth Leiden specimen (RMNH.AVES.172021) is certainly A. striatus. Its label proves that it was received in Leiden before 1850 and evidence for its provenance and that of RMNH.AVES. 172018 is held in the NBC archives. These include a list of Australian specimens sent by Gould to C. J. Temminck, inaugural Director of RMNH, which accompanied two copies of Part 1 of Gould's The birds of Australia, published in December 1840. The list includes skins and skeletons of mammals, and specimens, skeletons and eggs of birds; 114 species are listed among the birds, including one Ann/tis textilis and one A. striatus. From the list of then recently named and yet-to-be-named species in Gould's list, as well as the inclusion of Gould's The birds of Australia Part 1, it is evident that these specimens were sent to Temminck in December 1840 or early 1841. With little doubt, Gould had returned from Australia with them but was able to dispose of any material surplus to his immediate needs.

Documentation accompanying both RMNH.AVES.172018 ('A. textilis') and RMNH. AVES. 172021 (A. striatus) refers to 'pi 67 f 2' [i.e. Plate 67 figure 2 of Lesson (1831)]. Reference to 'Uranie Zool' (RMNH.AVES.172018) acknowledges Quoy & Gaimard's (1824) zoological account in Voyage autour du monde sur les corvettes de I'Uranie et la Physicienne, including the description of A. textilis, which they figured on PL 23. Pl. 67 fig. 2 of Lesson (1831) also illustrated A. textilis but Gould did not believe that this was the case. He observed (Gould 1848, 1865) that 'the bird figured in the "Voyage de I'Uranie" doubtless represents the present species ['Textile Wren Ann/tis textilis'], while that figured by Lesson in the Atlas to his "Traite d'Ornithologie," ... as clearly belongs to A. striatus.’ Even in his initial description of ‘Dasyornis [=Amytornis] striatus’ Gould (1840) wrote that it is 'nearly allied to the Ann/tis textilis of Lesson' (but by inference not to Malurus ( =Amytis ) textilis of Quoy and Gaimard). Gould was mistaken; while Lesson's figure shows a more rufous-plumaged bird than is depicted in Quoy & Gaimard (a distinctive feature to which Gould correctly attributed significance), it is still recognisably A. textilis, since it lacks the distinctive white throat and black moustachial stripe of the other species.

Possible alternative early specimen sources of A. modestus and A. striatus in New South Wales. — While it is almost certain that Gould supplied two of the NBC specimens and possibly all four, potential alternative 19th-century suppliers must be considered. Sharpe (1883) listed among specimens of 'A. textilis’ in the British Museum a mount supplied by Governor George Grey of South Australia,