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'TO S 102

British Ornithologists’ Club

MUSEUM LIBRARY

JUL 2013

PRHSEKjTED _

Volume 133 No. 2
June 2013

‘Uli'cJ •

FORTHCOMING MEETINGS
‘ j fit

Se'e also "BOO webkife: Kttp://wwfw. boc-online.org

BOC MEETINGS are open to all, not just BOC members, and are free.

Evening meetings are held 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_bariey_mow.
html or ask the Chairman for directions.

The cash bar will open 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 about one hour.

It would be very helpful if those who are intending to come would notify the Chairman no later than the day before the
meeting and preferably earlier.

24 September 2013 — 6.30 pm — Dr Roger Safford — Recent advances in the knowledge of Malagasy region
birds

Abstract: The Malagasy region comprises Madagascar, the archipelagos of Seychelles, the Comoros and the
Mascarenes (Mauritius, Reunion and Rodrigues), six more isolated islands or small archipelagos, and the
associated sea areas. It contains one of the most extraordinary and distinctive concentrations of biological
diversity in the world. The last 20 years have seen a major increase in the level of knowledge of, and interest
in, the birds of the region. This talk will draw on research carried out during the preparation of the first
thorough handbook of the region's birds — numbering 487 species — to be compiled since the magnificent
works of the late 19th century. The systematics of most taxa have been assessed using molecular techniques,
revealing numerous surprises and a new family, the Bernieridae (tetrakas), although intriguing questions
remain, not least the relationships of those two most baffling of groups, the Leptosomidae (cuckoo-roller) and
Mesitornithidae (mesites). Current bird work is very patchy, with remarkably little study of 'natural history'
despite the many gaps in our understanding; an interesting exception is the explosion of work of satellite
tracking of seabirds. These and other aspects will be reviewed in a wide-ranging talk.

Biography: Roger Safford has been a frequent visitor to the Malagasy region since 1988, and in 1989-93 he
completed a Ph.D. on the conservation of the endemic passerines of Mauritius, visiting all of the main islands
in the region and developing an intimate knowledge of the region's birds. His subsequent work has always
retained a link to the Malagasy region, with numerous visits and publications resulting, and since 2001 he has
been responsible for supporting the work of the BirdLife International partnership in Madagascar.

19 November— please check the BOC website where details will be posted as soon as they are finalised.

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

BOC Office

P.O. Box 417, Peterborough PE7 3FX, UK
E-mail: boc.office@bou.org.uk. Website: www.boc-online.org
Tel. & Fax: +44 (0)1733 844820.

Club Announcements

Bulletin of the
BRITISH ORNI

Vol. 133 No. 2

TJjSJSJlHiSTORt I BOC- 2013 133<2>

museum ubrahy

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ISTS’ CLUB

Published 14 June 2013

CLUB ANNOUNCEMENTS

Chairman's message

The ninth Conference of the European Ornithologists' Union is being held in the UK at the Univ. of East
Anglia in Norwich on 27-31 August 2013 (Tuesday-Saturday). The BOC will have a stand and I would very
much like to meet any members of the Club who are attending, either at the stand or possibly at an informal
get-together at a bar or other convenient location. I also hope that we will be able to recruit some new
members during the event. If you are coming to the conference it would be very helpful if you could let me
know in advance so that we can make arrangements. Contact me at helen.baker60@tiscali.co.uk.

At the time of writing, we have had our first evening meeting in the very comfortable upstairs room at
the Barley Mow. Do join us on 24 September and 19 November.

We also had a very successful joint meeting with the African Bird Club (ABC) and Natural History
Museum (see below). ABC arranged for Charlie Moores to produce a podcast containing interviews with the
speakers. This is available on the BOC website http://www.boc-online.org/meetings-past-2013.htm. It lasts
for c.l hour and is well worth listening to.

The three supplements to the Bulletin Avian Systematics and Taxonomy (Vol. 112A), Recent Avian
Extinctions (Vol. 126A) and Why Museums Matter (Vol. 123A) should by now be freely available on the
Biodiversity Heritage Library website at www.biodiversitylibrary.org/bibliography/46639.

This is my last message as Chairman and by the time you read it we will have held our AGM and I will
have handed over responsibility to my successor.

Helen Baker

The 972nd meeting of the Club was held on Tuesday 26 February 2013 in the upstairs room at the Barley
Mow, 104 Horseferry Road, Westminster, London SW1P 2EE. Thirteen members and five non-members were
present.

Members attending were: Miss HELEN BAKER (Chairman), Cdr. MICHAEL CASEMENT, RN,
STEPHEN CHAPMAN, MARTIN GAUNTLETT, RICHARD LANGLEY, DAVID MONTIER, Dr ROBERT
PRYS-JONES, ROBIN PRYTHERCH, Dr PETER RUDGE, PAT SELLAR, TONY STATHAM, CHRIS STOREY
and MIKE WALTON.

Non-members attending were: Mrs MARGARET GAUNTLETT, Mrs MARY MONTIER, JAMES
REYNOLDS (Speaker), ROGER WOOLVEN-ALLEN and LINDA WOOLVEN-ALLEN.

Dr James Reynolds (Univ. of Birmingham Centre for Ornithology) spoke on Ascension Island and Sooty
Terns: an ecological disaster or a smorgasbord in the eyes of conservation biologists? Ascension Island in many
ways is an ornithological outpost of the UK. It is one of several UK Overseas Territories and is 'off the beaten
track' because of its remote location midway between West Africa and Brazil in the tropical Atlantic Ocean.
Jim began by explaining the island's historical, geographical and cultural contexts, before describing its
ornithological treasures as a major breeding ground for seabirds within an otherwise landless seascape of
one million square miles. He has worked there since 2008, when he accompanied the Army Ornithological
Society (AOS) as their scientific advisor. The AOS has mounted expeditions to the island since 1990, since
when many seabird species (including Masked Booby Sula dactylatra, Brown Noddy Anous stolidus, Black
Noddy A. minutus, Ascension Frigatebird Fregata aquila and White Tern Gygis alba) have been censused.

However, the focus of Jim's talk was the mainstay of the AOS's and his ongoing research. Sooty Tern
Onychoprion fnscatus breeds on the island and work over the last 23 years has unlocked many of the species'
secrets. For example, he provided empirical evidence from ringing-recapture efforts for its subannual
breeding cycle; he documented the precipitous decline in the population size from as many as 3,000,000
birds as recently as 100 years ago to the current population of 340,000 birds; he described the major predation
pressure from a meso-predator (rat) release event as a result of the eradication of the apex predator (cat) in
2004; and he explored the role of food availability in the tern's population biology. He finished by explaining
how current state-of-the-art tracking technology is revealing movements of the species for the first time,
bridging a major gap in our knowledge about where birds go post-fledging for up to the first seven years of
their lives, about where they forage in the South Atlantic and how they spend the 91% of their lives when
they are not on land.

Club Announcements

Bull. B.O.C. 2013 133(2)

Joint BOC, African Bird Club, Natural History Museum meeting, at the Natural History Museum,
London, 6 April 2013

This was the second of a series of joint meetings held by the BOC and the NHM in conjunction with various
regional bird clubs. This time it was the turn of the African Bird Club. The day's programme comprised
six talks, three each either side of lunch. The morning started with Paul Donald speaking on the Liben
Lark Heteromirafra sidamoensis, possibly the most threatened bird species in Africa. Paul was standing in for
Bruktawit Abdu, who unfortunately had been unable to travel from Ethiopia as originally planned. Paul
reviewed the species' taxonomic history with respect to other forms in the genus elsewhere in the Horn of
Africa, and described research being undertaken to understand the lark's requirements and to conserve its
habitat on the Liben Plain. He also revealed that another population of Liben Larks has been found 600 km
from the Liben Plains, and that sidamoensis is synonymous with the poorly known Archer's Lark H. archeri,
which name has priority (see also Spottiswoode et al. 2013. Rediscovery of a long-lost lark reveals the
conspecificity of endangered Heteromirafra populations in the Horn of Africa. /. Orn. DOI 10.1007/sl0336-013-
0948-1 ). He was followed by a presentation on Lesser Crested Tern Sterna bengalensis in Libya by Abdulmaula
Hamza, who explained that there are now three breeding locations known in Libya and that at each of these
there has been an increase in numbers in recent years. Abdulmaula has carried out detailed research on
the species' feeding requirements and has carefully analysed the fish species taken. He has also ringed a
large number of chicks, c.20 of which have been recovered in different parts of the Mediterranean and West
Africa. The morning session concluded with a travelogue about birds and birdwatching in Rwanda, Africa's
most densely populated country, given by Jason Anderson. Jason described the country's various habitats
and the different species of birds in each. His talk was illustrated with many great photographs and also
accompanied by sound-recordings.

The lunch break was followed by the African Bird Club's AGM and then the talks resumed with a
fascinating account by palaeontologist and artist Julian Hume of his work reconstructing the lost world of the
Dodo and the extinct birds of the Mascarenes, from the fossil record and the scant historical documentation
of the early settlers. Dr Siobhan Cox then spoke about her genetic work reconstructing a phylogeny for the
Zosterops white-eyes of the African mainland. Her results indicate that traditional taxonomy is incorrect in
many respects and that there are more species than currently accepted, with Montane White-eye Z. poliogaster
being especially affected. Nigel Redman concluded the event with an account of his recent visit to the
officially unrecognised Republic of Somaliland, a seldom visited self-governing country that is technically
part of Somalia despite declaring independence in 1991. His talk focused on the endemic and near-endemic
birds of the region, with particular attention paid to the taxonomic status and history of the larks, and was
illustrated by numerous photographs of most species.

Attendance at the meeting was excellent, with well in excess of 100 people present, and there was
enthusiastic support for the planning of future meetings with other regional bird clubs.

David Fisher

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

New and interesting records for the Obi archipelago
(north Maluku, Indonesia), including field
observations and first description of the vocalisation
of Moluccan Woodcock Scolopax rochussenii

by Marc Thibault, Pierre Defos du Ran, Olivier Pineau &

Wesley Pangimangen

Received 6 October 2012

Summary. — The avifauna of the Obi archipelago is rather poorly studied and
current understanding is essentially based on several historic collecting efforts and
few recent visits by modern ornithologists, none of which reached the mountains
above 750 m. Furthermore, the taxonomic position of many bird populations
restricted to the archipelago or shared with nearby Bacan Island remains confused.

We describe the results of a two-week avifaunal survey of Obi in March 2010. We
provide first records since 1982 of the poorly known Moluccan Woodcock Scolopax
rochussenii as well as the first description of its vocalisation and first information on
its habitat, which, contrary to what was previously speculated, includes lowland
forest. We also provide confirmed records of five taxa previously unknown on
the island, including one that possibly represents a new subspecies (Sulawesi
Myzomela Myzomela chloroptera). New elevational information is presented for
34 species. Comments on the taxonomy of several endemic taxa are made on the
basis of new vocalisation or photographic material, suggesting that at least two
deserve biological species status (Northern Golden Bulbul Thapsinillas longirostris
lucasi, Dusky Myzomela Myzomela obscura rubrotincta) and offering further support
to treat Cinnamon-breasted Whistler Pachycephala johni as a species. Finally,
we emphasise the need for taxonomic reappraisal of several other endemic
insular forms (Cinnamon-bellied Imperial Pigeon Dncula basilica obiensis, Violet¬
necked Lory Eos squamata obiensis, Red-cheeked Parrot Geoffroyus geoffroyi obiensis,
Hair-crested Drongo Dicrurus hottentotus guillemardi, Northern Fantail Rhipidura
rufiventris obiensis, Paradise Crow Lycocorax pyrrhopterus obiensis and Island Leaf
Warbler Phylloscopus poliocephalns waterstradti). Although most of our observations
await confirmation, they suggest that endemism on Obi at the species level is
perhaps significantly under-estimated.

Obi and its satellite islands of Bisa, Tapat, Obilatu, Gomumu and Tobala form a remote
oceanic archipelago in Maluku, 33 km south of Bacan, 100 km east of Mangole (Sula Islands)
and 125 km north of Seram (Fig. 1). Total surface area is 3,040 km2 with Obi (2,670 km2)
the largest island, predominantly hilly and rising to at least 1,558 m. Together with the
three large islands of Morotai, Halmahera and Bacan, and several smaller islands, the Obi
archipelago forms part of the Northern Maluku Endemic Bird Area (EBA 171) distinguished
by 32 species and 101 subspecies restricted to this region (following Gill & Donsker 2012).

To date, the resident avifauna of the Obi islands group is known to support 16
Northern Maluku endemic species shared with both Bacan and Halmahera. Only two
endemics currently recognised at species level are confined to Obi (Carunculated Fruit
Dove Ptilinopus granulifrons and Cinnamon-breasted Whistler Pachycephala johni), but the
archipelago also hosts 19 recognised endemic subspecies, e.g. Paradise Crow Lycocorax

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

Figure 1. Location of Obi in eastern Indonesia and localities visited or mentioned in the text.

pyrrhopterus obiensis, while the enigmatic Moluccan Woodcock Scolopax rochussenii and
two subspecies (Chattering Lory Lorius garrulus flavopalliatus and Island Leaf Warbler
Phylloscopus poliocephalus zvaterstradti) are shared only with Bacan and Kasiruta (White &
Bruce 1986). Interestingly, the Obi islands also host endemic subspecies of several residents
that, according to current knowledge, do not occur elsewhere in northern Maluku: Pale
Cicadabird Coracina ceramensis hoogerwerfi, Northern Fantail Rhipidura rufiventris obiensis,
Turquoise Flycatcher Eumyias panayensis obiensis and Hair-crested Drongo Dicrurus
hottentotus guillemardi.

Despite its distinctive avifauna but as a result of its remote situation, Obi has received
little attention from modern ornithologists. Bernstein was the first bird collector to visit the
island in 1862 (Jansen 2008) followed by Guillemard in 1883, Doherty in 1897 (the latter
also visiting Bisa), Lucas in 1898 and Waterstradt in 1902 (Hartert 1903a). Following a gap
of 50 years, an expedition from the Museum Zoologicum Bogoriense was conducted under
the leadership of Wegner in July-November 1953 (Mees 1982) and its results published
by Jany (1954, 1955). Further specimens were collected in 1981-83 by Momou and Tatu,
during expeditions organised by the Smithonian Institution with the assistance of the
Indonesian Nature Conservancy, as part of ethnobiological research conducted by Taylor
(1990). These specimens were deposited at the National Museum for Natural History
(NMNH), Washington DC. In December 1989 Linsley (1995) conducted a bird survey as
part of a collaborative project between Manchester Metropolitan University, the Indonesian
Institute of Sciences (LIPI) and the Directorate-General for Forest Protection and Nature
Conservation (PHPA). He observed nine species not previously reported on Obi and
provided the first ecological information for many taxa and descriptions of the vocalisations
of several endemic forms. A brief exploration of Bisa was conducted on 11 June 1990 by
Bishop (1992). Further ornithological observations were made in February 1992 by Lambert
(1994) during a one-month survey of parrots under the auspices of IUCN; eight species
were recorded on Obi for the first time. While maximum elevations reached by the earlier
collectors are undocumented, Linsley (1995) and Lambert (1994) managed to reach c.500 m
and 730 m, respectively. The basic ecology of a large suite of species remains poorly known
and the rate of new island records hints that the avifauna is still poorly documented. This,
together with the paucity of records for the enigmatic Moluccan Woodcock, encouraged us
to undertake a short expedition to the island with the aim of improving knowledge of its
avifauna.

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

Study area and Methods

On 7-21 March 2010, observations were made at ten sites on Obi. Details of these,
survey dates and habitats are provided in Table 1 and Fig. 1. As many days as possible
were spent above 800 m (10-16 March) and we eventually reached forest between 1,000 and
1,220 m on 13-16 March. Elevations and geographical coordinates were measured using a
Garmin eTrex Vista HCx GPS. Subsequently, elevations and geographical data collected
in the field were compared with satellite data available via Google Earth and some minor
elevation corrections were made. It is of note that the highest elevation of 1,558 m given
for Obi by Google Earth satellite data contradicts both the US Army map SA-52 1:1,000,000
(1968) and that in Coates & Bishop (1997), where 1,611 m is given as the highest elevation.

To support identifications and descriptions, MT took photographs using a Canon
40D digital camera with a 100-400-mm lens. Sound-recordings were made by MT using
an Edirol R09-HR digital recorder with inbuilt microphone. Sonograms were prepared
using Syrinx 2.6h by John Burt (available at www.syrinxpc.com). Recordings of Moluccan
Woodcock have been uploaded to www.xeno-canto.org and further recordings will be
uploaded to this online database.

Recent state of natural habitats on Obi

F. G. Rozendaal in Collar & Andrew (1988), Linsley (1995) and Vetter (2009) already
reported that much of the lower lying forest on Obi was logged during the 1980s and early
1990s, while much of the remainder is under logging concession. According to Vetter (2009),
Obi has suffered some of the largest reductions of lowland forest within north Maluku as
a result of plantation development. In addition, illegal gold mining was reported to be
destroying some areas of forest (F. G. Rozendaal in Collar & Andrew 1988).

Three logging companies, PT Poleko Yurbarson Trad., PT Telaga Bhakti Persada and
PT Pusaka Agro Sejahtera, currently run six large logging licences covering two-thirds of
Obi. These licences generally permit selective logging up to c.100 m3 / ha and for c.20-30
years (N. Brickie pers. comm.). Along the 70-km stretch of coast between Jikotamo and the

TABLE 1

Study site details, survey dates and habitat. See also Fig. 1.

Location (elevation)

Coordinates

Dates

Habitat

Soligi (sea level)

01°39'S, 127°25'E

7-8, 20 March 2010

Village; coconut plantation and other
tree crops; mangrove

PT Telaga Bhakti Persada
logging settlement (sea level)

01°42'S, 127°29'E

19 March 2010

Village; coconut plantation; grassland

Site A (200-300 m)

01°38'S, 127°30'E

19 March 2010

Logged forest

Site B (360-450 m)

01°36'S, 127°33'E

8, 17-19 March 2010

Logged forest

Site C (400-700 m)

01°33'S, 127°34'E

8, 9, 16-17 March 2010

Logged forest

Site D (700-850 m)

01°32'S, 127°35'E

9-13, 16 March 2010

Logged and primary forest

Site E (850-1,220 m)

01°32'S, 127°37'E

13-16 March 2010

Logged and primary forest including
montane forest

Laiwui (sea level)

01°19'S, 127°38'E

20-21 March 2010

Village; coconut plantation; beach

Jikotamo (sea level)

01°20'S, 127°39'E

20-21 March 2010

Village

Site F (50-150 m)

01°21'S, 127°40'E

20-21 March 2010

Logged forest; regrowth forest; coconut
plantation; grassland

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

PT Telaga Bhakti Persada logging settlement south of Soligi, hills were mainly covered by
plantations (including cloves, coconut palm, nutmeg, ambarella and cacao) and fragmented
secondary forest. Hills separating Manis Lake from the west coast were barren as a result
of ongoing open-cast nickel mining. Remnant primary forest was mainly restricted to the
most rugged and inaccessible terrain. Alluvial flats were extensively covered by coconut
plantations, with the notable exception of a relatively large tract of coastal forest bordered
by old-growth mangrove immediately north of Soligi.

Within the PT Telaga Bhakti Persada logging concession south and east of Soligi, forest
was very degraded below 200 m, large areas had already been selectively logged up to 950
m and the lowest patch of primary forest was at 500-700 m, although some large trees there
were marked for logging. At 700-950 m, forest was in better condition, with fragments
of primary forest still covering some of the steeper, less accessible slopes, while the more
accessible areas were mainly covered by selectively logged forest. A small area of tropical
forest explored at 1,100-1,220 m showed no evidence of logging.

Soligi villagers reported that tree-plantation operations were planned by the logging
company and a tree nursery was seen in the concession. Several drill holes excavated
during recent mining exploration were found well inside the logging concession up to 600
m. According to some villagers, these have to date not resulted in any further mining. It
is of note that in the north of the island, a large-scale drilling project was withdrawn in
September 2011. It was initially set to target a large epithermal gold / silver / base metals
system under a license covering 7,700 ha, which is already the subject of extensive artisanal
gold mining (Ashburton Minerals Ltd. 2012).

To our knowledge, Obi lacks any protected area, although a nature reserve of 450 km2
covering highlands in the centre of the island and further protection forests on steep terrain
have been proposed (FAO 1981-82, BirdLife International 2013).

Species accounts

Overall we recorded 74 bird species, of which 66 are resident or presumed so and eight
are Palearctic migrants. Three additional taxa are either unconfirmed or remain unidentified
to species. Tire following details new island records, new elevational or noteworthy records
from a distributional, ecological or taxonomic perspective. Other species encountered
during our survey are listed in Table 2. Taxonomy and nomenclature follow Gill & Donsker
(2012) except that we use Common Golden Whistler Pachycephala pectoralis instead of Black-
chinned Whistler P. mentalis.

MOLUCCAN WOODCOCK Scolopax rochussenii

The following records were obtained at site B at c.420 m, from a logging track bisecting an
extensive area of logged forest on flat and more graded sandstone soils: at c.19.00 h on 17
March 2010, calls reminiscent to those given in display-flight by Bukidnon Woodcock S.
bukidnensis were briefly heard (OP, MT & PDR). Between 02.00 h and 04.00 h on 18 March
2010, similar calls were repeatedly heard by OP and subsequently at 05.55 h and 06.07 h
by the three observers, when sound-recordings (www.xeno-canto.org/109282, www.xeno-
canto.org/109283) and a glimpse of a fast-flying bird were eventually obtained at dawn by
MT. At 07.00-07.15 h (before dusk) on the same day, what was presumably the same bird
was observed three times by MT, when it called in roding (i.e. courtship) display-flight at the
same location. It was flying rather straight, rapidly and low above the canopy, sometimes
<100 m from the observer, and was briefly seen opening its bill when calling. Rock Dove
Columba livia- like size, overall compact stocky shape, with a rounded bulky chest, short tail,
broad pointed wings and a very long straight bill were noted. These characters, together

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

TABLE 2

Species encountered on Obi not mentioned in the main text.

Species name

Dusky Megapode Megapodius fmjcinet
Pacific Reef Heron Egretta sacra sacra
Striated Heron Butorides striata
Eastern Cattle Egret Bubulcus coromandus
Eastern Osprey Pandion cristatus
Pacific Baza Aviceda subcristata rufa
Brahminy Kite Haliastur indus

White-bellied Sea Eagle Haliaeetus leucogaster
Oriental Hobby Falco severus
Spotted Kestrel Falco moluccensis
Common Sandpiper Actitis hypoleucos
Red-necked Phalarope Phalaropus lobatus

Greater Crested Tern Sterna bergii
Pied Imperial Pigeon Ducula bicolor
Great-billed Parrot Tanygnathus megalorhynchos

Barking Owl Ninox connivens rufostrigata
Large-tailed Nightjar Caprimulgus macrurus schlegelii
Glossy Swiftlet Collocalia esculenta
Moustached Treeswift Hemiprocne mystacea confirmata
Common Paradise Kingfisher Tanysiptera galatea obiensis
Blue-and-white Kingfisher Todiramphus diops
Beach Kingfisher Todiramphus saurophagus

Common Kingfisher Alcedo atthis

White-bellied Cuckooshrike Coracina papuensis

Willie Wagtail Rhipidura leucophrys

Torresian Crow Corvus orru

Arctic / Kamchatka Leaf / Japanese Leaf Warbler
Phylloscopus borealis / examinandus / xanthodryas

Gray's Grasshopper Warbler Locustella fasciolata

Moluccan Starling Aplonis mysolensis

Grey-streaked Flycatcher Muscicapa griseistricta

Black-faced Munia Lonchura molucca

Eastern Yellow Wagtail Motacilla tschuschensis ssp.

Grey Wagtail Motacilla cinerea

Comments

1 at c.150 m
1 on the coast
1 on the coast

few seen on the coastal plain

singles seen twice near sea level; presumably resident
1 near sea level

quite common from sea level to 200 m, with additional
records to 1,200 m

4 on coast

1 at Soligi

common in coastal plain
3 near Soligi

common in coastal seas, with 600-1,000 from boat between
Soligi and Jikotamo

c.10 seen inshore between Soligi and Jikotamo

2 in coconut / coastal forest near Soligi

2 near sea level in degraded forest and coconut plantation
near Soligi, 1 heard near Laiwui

1 photographed at site F at c.100 m

seen twice at 400-500 m

from sea level to 940 m

three records at 300 m, 480 m and 600 m.

1 seen at site F

common on coast around Soligi with, e.g. 9 from a boat
along 8 km of coast

1 (A. a. hispidoides) at 480 m; 1 (possibly migrant A. a.
bengalensis) on coast

6 records from sea level to 420 m

1 near sea level near Jikotamo

several encounters from sea level to 1,000 m

3 at 750-840 m

fairly common to 450 m

quite common migrant, from lowlands to 990 m
flock of 30 at 220 m and 2 more at 360 m
1 at site F

5 records

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

with the flight behaviour, are typical of Scolopax. MT also noted that the bird appeared to
be significantly larger than congenerics with which he had previous experience (Eurasian
Woodcock S. rusticola and S. bukidnonensis), but light conditions were too poor to observe
any plumage characters. At 06.02-06.20 h on 19 March 2010, still in poor light conditions, a
bird (probably the same) was seen again once and heard twice more (PDR & MT).

Further observations and sound-recordings (www.xeno-canto.org/109284) were made
at site F, at c.100 m: at c.06.45 h on 20 March 2010, a woodcock was seen in flight and heard
twice roding low over the canopy (MT & WP). The bird was seen in better light conditions
and flew just overhead at a distance of >30 m. Typical Scolopax shape and large size were
noted and WP also noticed yellowish, mainly unstreaked underparts. At 06.02 h the
following morning, what was presumably the same roding bird was seen and heard again
twice at the same location (MT & PDR). The observations were made in a coconut plantation
at the edge of a patch of disturbed, fragmented primary forest, with evidence of ongoing
small-scale logging. The undergrowth included a marshy gully with bare ground bisected
by a low-flowing stream. It was adjacent to a wet grazed pasture and surrounded by steep
hills covered by degraded forest interspersed by scattered clove and nutmeg plantations.

Our observations are the first documented records of Moluccan Woodcock since two
females were collected in August-September 1982 at 'Galala', said to be on Obi (http://
collections.si.edu). The latter locality is considered untraced by BirdLife International
(2011b) and indeed we failed to find a locality of this name on Obi, although a remote
village called Galala exists on the island of Bisa (Fig. 1). The species is known from only six
other specimens, of which three were also collected on Obi (two in the 19th century and
one in 1953, the latter at Wayaloar on the south coast), one on Bacan (in 1902) and two of
unknown origin (ITartert 1903b, BirdLife International 2011b). During their field work on
Obi in 1989 and on Obi and Bacan in 1992, respectively, Linsley (1995) and Lambert (1994)
did not observe the species, but the latter obtained a convincing report from a guide at Kelo,
Obi, who 'occasionally flushed this bird from ridgetops above ca. 500 m'.

Our sound-recordings permit the vocalisation of S. rochussenii to be described. In
display, it gives a loud and distinctive call, comprising a hard, metallic, rattled or staccato
phrase, with 8-11 motifs, each motif given at intervals of 0.04-0.05 seconds (Fig. 2). This
rattle phrase can be transcribed ti'ti'ti'ti'ti'ti'ti'ti'ti. Each motif has a similar structure,
comprising a distinctive pulse (c.4 kHz rising to c.5-5.5 kHz) connecting with a concurrent,
higher piched and slightly variable curled syllable (c.6-6.8 kHz rising to 6. 5-7.3 kHz and
falling to 5.3-6 kHz). The rattle phrase is given at intervals of 1.8-3. 3 seconds. Rattling
calls of Moluccan Woodcock are strikingly similar in structure and frequency to those of S.
bukidnonensis, but the latter have only five motifs and the pulse begins at a lower pitch (c.3
kHz) (Kennedy et al. 2001, www.xeno-canto.org/species/Scolopax-bukidnonensis). Rattles
delivered by S. bukidnonensis are frequently interspersed by lower pitched, growling or
grunting phrases, which were not heard from S. rochussenii, although this might just reflect
the small sample of recordings. The calls of S. rochussenii are very different from those of
New Guinea Woodcock S. rosenbergii (Kennedy et al. 2001; G. Wagner, www.xeno-canto.
org/23274) and from those attributed to Javan Woodcock S. saturata (G. Wagner, www.
xeno-canto.org/42307; F. Ducry & D. Marques, www.xeno-canto.org/56931), but we have
been unable to locate sound-recordings of Sulawesi Woodcock S. celebensis, which are
undescribed, the latter being closest to S. bukidnonensis by morphology. Our results suggest
a close evolutionary relationship between S. bukidnonensis and S. rochussenii.

Based on the very scant information available, Moluccan Woodcock was previously
assumed to be a bird of hill and montane forest (Hartert 1903b, White & Bruce 1986, Coates
& Bishop 1997, Vetter 2009). However, the only traceable locality (Wayaloar, southern

Marc Thibault et al.

Bull. B.O.C. 2013 133(2)

ih|ii]

j|,!

111

nifti .i

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1 2 3 4 5 6 7 8 9 10 11 12

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13 14 15

17 18

) | II | 1 .

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ill,

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Figure 2. Sonograms of the roding display call of Moluccan Woodcock Scolopax rochussenii from (A-C) east of
Soligi, Obi, 18 March 2010 and (D) near Jikotamo, Obi, 21 March 2010 (M. Thibault). X-axis = time (one second
per tick. Fig. 2A; 0.5 seconds per tick. Figs. 2B-D), y-axis = frequency (1 kHz per rick).

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Bull. B.O.C. 2013 133(2)

Obi, August 1953) is on the coast, raising 'the possibility that its optimal (or seasonally
optimal) habitat is to be found in lower-lying regions of islands where the largest, dampest
substrates may perhaps occur' (BirdLife International 2011b). Our records were obtained
at two of three localities visited below 500 m that we surveyed at suitable times, i.e. dusk
and dawn. These records confirm that the species holds territory (and presumably breeds)
in lowland forest, hence supporting the latter hypothesis and that it may not be uncommon
in suitable habitat. Woodcocks were not recorded at the two localities we surveyed at
600-1,200 m despite dedicated searching. Thus, we provide evidence that the species is
not only, if at all, associated with montane forest. Our records also suggest that, to some
extent, Moluccan Woodcock tolerates habitat degradation and fragmentation, and that
wet habitats in or close to forest (i.e. marshes or wet gullies around low-flowing streams)
are perhaps important for the species. However, much remains to be learned concerning
population size and ecological requirements. In the near future, direct habitat destruction
resulting from mining, logging, plantations and agriculture are likely to be extended in
lowland and montane areas on Obi and Bacan. Therefore, its status as Endangered (BirdLife
International 2011b) should be maintained, at least until further data are collected.

METALLIC PIGEON Colwnba vitiensis halmaheira

On 13-15 March 2010, a calling bird was seen and two more heard at site E at c. 1,200 m in
primary forest (MT, PDR & WP). On 14 March 2010 a bird was heard at the same site at
1,100 m (MT). Photographs and sound-recordings were made. No conclusive record was
obtained at lower elevation. The species is widespread in Wallacea. White & Bruce (1986)
and Coates & Bishop (1997) reported its presence on Gomumu, a small satellite island 9 km
south of Obi, but they did not mention it from mainland Obi. These statements contradict
Hartert (1903a), who mentioned that a specimen of 'Columba albigularis' was sent by Lucas
from 'Obi Major'. Furthermore, during a visit to Tring museum in 2011, MT & PDR were
able to check five (three females and two males) C. v. halmaheira collected by Waterstradt
on Obi in January 1903. These specimens might have been collected too late for inclusion in
Hartert's review of the birds of Obi. In any case our investigations clarify the status of this
taxon on mainland Obi and also describe its altitudinal range on this island.

SLENDER-BILLED CUCKOO-DOVE Macropygia amboinensis albiceps
Common in forest and forest edge from sea level to 1,210 m, with photographs made by
MT at c.1,200 m. Our records considerably extend the altitudinal range on Obi as previous
published records are below 500 m (Coates & Bishop 1997) and are in accordance with its
altitudinal range from elsewhere in Wallacea (Coates & Bishop 1997).

SCARLET-BREASTED FRUIT DOVE Ptilinopus bernsteinii micrus

Regularly seen and commonly heard at 150-1,210 m in primary and logged forest. A nest
attended by at least one adult, containing a single egg, was found on 13 March 2010 at 1,210
m in primary forest (WP, PDR & MT). The nest was a flimsy construction on a fern c.80 cm
above the ground, comprising a small moss cup on a larger assemblage of small branches.
P. bernsteini is endemic to North Maluku, with the nominate subspecies found from
Halmahera to Bacan and micrus restricted to Obi (Baptista et al. 1997). Previously known at
180-600 m on Obi (Lambert 1994). Our records include the first description of a nest on Obi,
whose characteristics and situation are similar to those described from Halmahera (Lansley
& Fames 2006).

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Bull. B.O.C. 2013 133(2)

SUPERB FRUIT DOVE Ptilinopus superbus

Common at 150-1,000 m, with further records of two separate birds heard at 1,200 m in
primary montane forest. Our records significantly extend the altitudinal range for the
species on Obi, where it was previously unrecorded above 700 m (Lambert 1994, Coates &
Bishop 1997).

CARUNCULATED FRUIT DOVE Ptilinopus granulifrons

Only recorded once, at site A on 19 March 2010, when at least eight were observed through
a telescope in the upper canopy of a large fruiting Ficus, in logged forest at c.220 m (PDR
& MT). This single record and the species' apparent absence at higher elevations appear to
corroborate that this is an uncommon species closely associated with lowland forest (Coates
& Bishop 1997, BirdLife International 2011a).

SPECTACLED IMPERIAL PIGEON Ducula perspicillata

Commonly heard and occasionally seen from sea level to 1,200 m in forest edge, logged
forest and primary forest. Our records significantly extend the species' altitudinal range on
Obi (Lambert 1994). They also significantly extend the altitudinal range for the species in
the northern Maluku where it was previously recorded to 900 m on Halmahera (Poulsen &
Lambert 2000).

CINNAMON-BELLIED IMPERIAL PIGEON Ducula basilica obiensis

This northern Maluku endemic was fairly commonly seen and heard in forest edge, logged
forest and primary forest at 150-1,210 m (WP, PDR, MT) and it was photographed at c.1,100
m (MT). Our records significantly extend the species' altitudinal range on Obi, where it was
previously unrecorded above 730 m (Lambert 1994) and match the upper limit of 1,230 m
for the nominate subspecies on Halmahera (Poulsen & Lambert 2000).

Sound-recordings of D. b. obiensis were obtained by MT (Fig. 3), enabling comparisons
with vocalisations of D. b. basilica from Halmahera (www.xeno-canto.org/species/Ducula-
basilica). Both forms give similar vocalisations: songs usually comprise a series of 1-2 (less
frequently three) very deep, throaty growls, roozv roooorw-ooow , at 0.4-0. 5 kHz. When the
first call is a short note (lasting 0.2 seconds), the second call is often longer (2. 0-2.5 seconds),
whereas when a first, longer call (0.8-1. 3 seconds) is given, there is no second call or it is
frequently shorter (i.e. 1.3-1. 7 seconds). Longer calls typically rise very slightly and end
abruptly.

It was recently proposed to split obiensis (which is confined to Obi) from basilica (on
Morotai to Bacan; www.birdlife.org/globally-threatened-bird-forums/2012/) following
application of the Tobias et al. (2010) criteria. Although justification for this taxonomic

nn :r ) m f rmr ; p

■mu

■riMnii'

12 3 4 5 6 7

Time (sec]

Figure 3. Sonogram of Cinnamon-bellied Imperial Pigeon Ducula basilica obiensis, Obi, 12 March 2010 (M.
Thibault). X-axis = time (one second per tick), y-axis = frequency (1 kHz per tick).

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Bull. B.O.C. 2013 133(2)

Figure 4. Cinnamon-bellied Imperial Pigeon Ducula basilica, with (A) basilica (a pair), Halmahera, 3 October
2011 (© F. Steinhauser), (B) obiensis (a pair). Obi, 15 March 2010 (M. Thibault).

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Bull. B.O.C. 2013 133(2)

treatment is unpublished, Hartert (1903a) already emphasised plumage differences
between these two taxa, stating that D. b. obiensis is 'very different' from D. b. basilica, 'the
entire head, throat, foreneck, and breast being much deeper vinous, with a greyish wash;
the hindneck darker grey, separated from the vinous head by a rusty patch; abdomen and
under tail-coverts deep cinnamon, instead of pale cinnamon'. According to Hartert (1903a),
the collector W. Doherty also noted that obiensis has 'iris dark crimson, eyelids vermilion;
feet vermilion; bill nearly black', whereas basilica (from Ternate) has 'iris crimson; feet
pale carmine; bill black'. Although the subtle differences in bare-parts coloration might be
difficult to appreciate in the field, the plumage differences between these taxa are obvious
(Fig. 4). However, given the lack of obvious vocal difference between D. b. basilica and D. b.
obiensis, we suggest that splitting these two taxa might require more investigation.

VIOLET-NECKED LORY Eos squamata obiensis

Recorded regularly from sea level to 700 m in a wide range of habitats including coconut
plantations, primary and logged forest. Less frequently encountered at higher elevations
with two sightings at 850 m and single observations at 1,110 m and 1,210 m (PDR, WP,
MT). Most frequently observed in pairs or trios, although a large flock of c.25 was seen on
11 March at 1,110 m. Previous published records on Obi range up to 730 m. The elevational
range on Obi is therefore significantly wider than previously known, similar to Halmahera
where E. s. riciniata has been found to 1,230 m (Poulsen & Lambert 2000).

No sign of trapping during our visit. E. squamata is widespread on Obi including in
primary forest, logged forest and non-forest habitats. As there was no obvious indication of
a status change compared to Lambert (1993), this species may require quantitative surveys
to detect any, perhaps slight, change in density.

Morphologically, obiensis closely approaches the nominate race (as mentioned by Collar
1997), which is restricted to the West Papuan islands, but is strikingly different from riciniata
on Morotai to Bacan, notably in having the entire head and neck uniform red and greater
wing-coverts and scapulars black (pers. obs.). Based on these observations, we suggest that
the taxonomic position of the taxa currently included in E. squamata should be reassessed
and require a detailed study of morphology, acoustics and DNA.

CHATTERING LORY Lorius garrulus flavopalliatus

Singles and groups of 2-4 recorded almost daily in logged and primary forest, with max.
flock of at least ten foraging birds in selectively logged forest at 440 m on 19 March (WP,
OP, PDR, MT). Regularly recorded to 1,000 m and seen twice at 1,100 m, extending the
altitudinal range of 370 m given previously for Obi (Coates & Bishop 1997). Several captive
E. g. flavopalliatus seen in villages and at a logging settlement, apparently being kept as pets.
As noted by Linsley (1995), this species is the commonest captive parrot on Obi, indicating
that it still faces heavy trapping pressure. However, compared to Lambert (1993), our
observations do not suggest a significant decline. It may require quantitative surveys to
detect any, perhaps slight, change in density.

RED-FLANKED LORIKEET Charmosyna placentis intensior

Very common from sea level to 400 m and less common, albeit regular, to 1,210 m. These
records provide the first data concerning the altitudinal range of C. placentis on Obi and
extend the altitudinal range known elsewhere in northern Maluku, where it had previously
been recorded up to max. 1,010 m on Halmahera (Poulsen & Lambert 2000). C. placentis is
occasionally recorded to 1,600 m in New Guinea (Coates & Peckover 2001).

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Bull. B.O.C. 2013 133(2)

RED- CHEEKED PARROT Geoffroyus geoffroyi obiensis

Total of 21 records of 1-6 birds seen or heard at 200-1,000 m, with 18 records below 600
m, mostly in selectively logged forest. Few were encountered in primary forest, possibly
because of lower density or near-absence at higher altitudes. Our records significantly
extend the altitudinal range of this subspecies endemic to Obi and Bisa, where it had been
reported to 800 m (Coates & Bishop 1997). G. geoffroyi comprises 16 widely recognised
subspecies ranging from Wallacea to New Guinea and north-east Australia (Collar 1997,
Gill & Donsker 2012), of which several differ markedly in plumage, size and vocalisations
(Collar 1997). G. g. obiensis is closer to G. g. cyanicollis from northern Maluku, sharing with
it a long blue collar, although on obiensis this is larger and it has a brownish back which is
lacking on cyanicollis (Collar 1997). The piercing, high-pitched nasal calls of obiensis have
been described by Lambert (1994) and Linsley (1995), who report that they are distinctive
compared to cyanicollis from Bacan, floresianus from Sumba and rhodops from Buru. These
observations suggest that a detailed taxonomic study of G. geoffroyi is required, including
morphological, acoustic and DNA comparisons.

ECLECTUS PARROT Eclectus roratus vosmaeri

Wild birds recorded just twice, both in the lowlands, with three in coastal coconut
plantations near Soligi on 8 March 2010 (all observers) and a female in degraded forest with
cloves and coconut plantations at site F on 20 March (WP, MT). A captive bird apparently
kept as a pet near Soligi. Reportedly very common, at least in the north of the island, in
the 1950s (Lambert 1994), but then declined dramatically due to trapping, so that Linsley
(1995) and Lambert (1994) only obtained one sighting each. Our observations indicate that
E. rotatus remains rare on Obi, where it is primarily confined to lowland and coastal areas.
Despite being protected by Indonesian law, it probably is extensively traded in northern
Maluku (ProFauna Indonesia 2008), although perhaps less so than both lories (Lambert
1993). There is no indication that the species' conservation status on Obi has improved or
even changed since the 1990s.

MOLUCCAN CUCKOO Cacomantis aeruginosas ssp.

Three calling Cacomantis were recorded: a vocalising bird on 11-12 March 2010 at site D at
750 m in primary forest (sound-recorded, MT), another calling bird photographed (Fig. 5)
and sound-recorded (Fig. 6A-E) at site E on 13 March 2010 and heard again on 16 March at

Figure 5. Moluccan Cuckoo Cacomantis aeruginosus ssp. (same individual), Obi, 13 March 2010 (M. Thibault).

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Bull. B.O.C. 2013 133(2)

f , f , , ,i *►*»►*«»

1 2 3 4 5 G 7 8 9 1 0 11 1 2 1 3 1 4 1 5 1 6 1 7 1 8 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47

Time (secj

1 2 3 4 5 6 7

Time (sec)

T— I C:

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25 30 35 40

Time (sec)

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1 2

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Time (sec)

5 6

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1 2

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Figure 6. Sonograms of Moluccan Cuckoo Cacomantis aeruginosus ssp. by M. Thibault, with (A) long, rising
and accelerating song. Obi, 13 March 2010, (B-C) details of A, (D) long level song. Obi, 13 March 2010 (same
individual as in Figs. 6A-C), (E) detail of D, (F) three-note song. Obi, 21 March 2010. X-axis = time (one
second per tick, y-axis = frequency (1 kHz per tick).

870 m in logged forest by MT, with a third briefly seen (WP, PDR, MT) and sound-recorded
(Fig. 6F) on 20-21 March at c.100 m at site F (MT). In addition, a juvenile Cacomantis fed by
a pair of Phylloscopus poliocephalus umterstradti was observed on 13 March 2010 at site E at
1,200 m in primary montane forest (WP, PDR, MT). The latter observation provides the first
documented record of a Cacomantis breeding on Obi.

Taxonomy of resident Cacomantis cuckoos in Maluku is highly confused (Rheindt
& Hutchinson 2007, Tebb et al. 2008, Rheindt 2010, Erritzoe et al. 2012) and the situation
is exacerbated by the seasonal occurrence of migrant Brush Cuckoo C. v. variolosus from
Australia. C. variolosus and C. sepulcralis are sometimes merged into one species (Payne
1997, 2005), but several works maintain the separation of C. v. infaustus and Rusty-breasted
Cuckoo C. sepulcralis aeruginosus in southern Maluku (White & Bruce 1986, Coates & Bishop
1997, Erritzoe et al. 2012). Hartert (1925) separated northern Maluku birds as oblitus, which
is reported to be paler below with longer wings than infaustus, which he considered to be
restricted to southern Maluku. However, most recent works consider oblitus a synonym
of infaustus (White & Bruce 1986, Coates & Bishop 1997, Payne 1997, Erritzoe et al. 2012).

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Bull. B.O.C. 2013 133(2)

Figure 7. From left to right: Cacomantis sepulcrnris virescens (three from Manado, Sulawesi), C. aeruginosus
aeruginosus (one from Buru and one from Ambon), C. variolosus (three from Bacan and two from Ternate) (M.
Thibault / © Tire Natural History Museum, Tring).

Heinrichi is a poorly known taxon described from Bacan and Halmahera by Streseman
(1931) who distinguished it from sympatric C. v. oblitus (= infaustus) on the basis of smaller
size, with proportionately shorter wings and longer tail, more rufous underparts, darker
olive-brown upperparts and yellow legs (instead of yellow-green or brownish). Other
features not mentioned by Streseman (1931) include rufous notches in the outer edges of the
rectrices (Coates & Bishop 1997, Erritzoe et al. 2012), dark grey throat (Payne 1997, Erritzoe
et al. 2012) and eye-ring either undescribed (Coates & Bishop 1997) or greyish (Payne 1997).
Heinrichi is variously treated as a synonym of resident C. v. infaustus (Payne 2005, Erritzoe
et al. 2012) or recognised as an endemic species sympatric with C. v. infaustus and migrant
C. v. variolosus (Heinrich 1956, Payne 1997, White & Bruce 1986, Coates & Bishop 1997).
Alternatively, it was recently suggested, based mainly on vocalisations, that aeruginosus and
heinrichi may belong to the same species (Tebb et al. 2008; J. A. Eaton pers. comm., F. Rheindt
pers. comm.), a view supported by Gill & Donsker (2012), who afforded aeruginosus species
status, including C. a. aeruginosus from southern Maluku and C. a. heinrichi on Halmahera
and Bacan.

A distinct race, C. v. obiensis, was described from Obi by Jany (1955) on the basis of
darker plumage and longer tail compared to resident C. variolosus from northern Maluku,
but he did not compare it with C. aeruginosus aeruginosus. C. v. obiensis is generally treated
as a synonym of C. v. infaustus (White & Bruce 1986, Payne 1997, Erritzoe et al. 2012). White
& Bruce (1986) and Coates & Bishop (1997) mentioned the presence of two taxa on Obi,
namely migrant C. v. variolosus and resident C. v. infaustus, but a calling bird observed and
sound-recorded by F. R. Lambert possibly refers to C. heinrichi (www.xeno-canto.org/67791).

The bird photographed on 13 March 2010 (Fig. 5) was rather long-tailed with dark
upperparts. It had a distinctive pattern on the underside. of some rectrices, with whitish to

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Bull. B.O.C. 2013 133(2)

pale rufous bars extending the full width of the inner web. It also had rich rufous underparts
with greyish feathers restricted to the upper throat and chin, merging with rufous feathers,
yellow eye-ring, brownish tone to the base of the lower mandible and bright yellow legs.
These features most recall C. a. aeruginosus (Fig. 7; http://orientalbirdimages.org/search.
php?p=2&Bird_ID=448&Bird_Family_ID=&pagesize=l) and C. a. heinrichi, based on the
description of Streseman (1931). The bird photographed on Obi also strongly recalls a bird
recently photographed on Halmahera and tentatively identified as Moluccan Cuckoo (Tebb
et al. 2008). It noticeably differs from C. s. virescens from Sulawesi and the Sula Islands,
the latter having greyish feathers extending lower on the breast and shorter whitish
bars restricted to the inner web of the rectrices (Fig. 7). In addition, adults from northern
Maluku referred to C. variolosus in Tring museum all possess paler rufous underparts with
a predominantly greyish throat and breast (Fig. 7).

Most taxa in the C. sepulcralis / variolosus complex give two or more vocalisation
types: (1) trisyllabic call notes repeated at rising frequencies and (2) a series of single calls
usually repeated at level frequency (Coates & Bishop 1997, Rheindt 2010; pers. obs.), albeit
with pronounced inter-racial differences (Coates & Bishop 1997, Payne 2005, Rheindt &
Flutchinson 2007). In addition, distinctive variant vocalisations unambiguously given by C.
a. aeruginosus on Seram, Buru and Ambon (Coates & Bishop 1997, Rheindt & Hutchinson
2007) and presumably by C. a. heinrichi on Halmahera (Tebb et al. 2008) include long series'
of calls repeated very rapidly and delivered on either a level or rising frequency. These
variant vocalisations are unknown in other forms of the sepulcralis / variolosus complex
(Rheindt & Hutchinson 2007, Tebb et al. 2008, Rheindt 2010) and are also lacking in the
many recordings of C. variolosus from the Lesser Sundas, New Guinea and Australia that
we examined in online sound collections (i.e. xeno-canto.org; avocet.zoology.msu.edu/).

Variant calls described above and similar to calls previously recorded on Obi by F. R.
Lambert were given by each of the three birds we recorded. These calls comprised rather
flattened, hook-shaped individual call elements, which are frequently given by various
races of sepulcralis and C. a. aeruginosus, but much less so by C. variolosus. Most interestingly,
we failed to record the typical staple-shaped notes on Obi given by C. variolosus (see Tebb
et al. 2008; F. R. Lambert, www.xeno-canto.org/38144).

Based on plumage and vocalisations, we conclude that our records refer to C. aeruginosus
giving further support to the view that heinrichi and aeruginosus represent a single species
(Tebb et al. 2008; F. Rheindt pers. comm., J. A. Eaton pers. comm.). Furthermore, given that
we failed to record typical variolosus calls, we question the occurrence of a resident variolosus
population (i.e. C. v. infaustus) on Obi and suggest that careful examination of Cacomantis
specimens from Obi identified as variolosus might reveal, if they do not represent the
aeruginosus / heinrichii group, that they are migrants of nominate variolosus from Australia. A
thorough bio-acoustic, morphological and genetic study is required to clarify the taxonomy
of Cacomantis in Australasia and Wallacea. Regarding the resident population on Obi,
further study should clarify whether it belongs to C. a. aeruginosus or C. a. heinrichi, or to the
endemic ohiensis described by Jany (2005), which in our view should not be synonymised
with C. v. infaustus until a thorough assessment is conducted.

MOLUCCAN SCOPS OWL Otus magicus obira

Seen three times and frequently heard at c.100-1,210 m, in forest edge, logged forest
and primary forest. These are the first data on habitats, elevational range, status and
vocalisations of this bird on Obi (White & Bruce 1986, Coates & Bishop 1997). Sound-
recordings were obtained on 9 March 2010 (Fig. 8). The call is a short, harsh, rasping kwok,
repeated at intervals of 5-7 seconds. Calls last 0.33-0.5 seconds, at 0.6-1. 2 khz. When excited

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Bull. B.O.C. 2013 133(2)

Time (sec)

3 -
2

1 -
0 -

Time (sec)

3 -
2 -

1
0

1 1 >4i>nin

Time (sec)

Figure 8. Sonograms of Moluccan Scops Owl Otus magicus obira, Obi, 8 March 2010 (M. Thibault): A: typical
call. B: detail of A. C: excited kivook calls. X-axis = time (five seconds per tick, Fig. 8A; 0.5 seconds per tick.
Fig. 8B; one second per tick. Fig. 8C), y-axis = frequency (1 kHz per tick).

by playback, one bird gave a series of kivook notes slightly longer and deeper than the usual
call and repeated at shorter intervals. These vocalisations were found to be more restricted
in frequency range, but otherwise similar to recordings of O. m. magicus from Burn (F. R.
Lambert, www.xeno-canto.org/68940), O. m. leucospilus from Halmahera (D. Farrow, www.
xeno-canto.org/19771) and O. m. albiventris from Flores (F. R. Lambert, www.xeno-canto.
org/121829).

UNIFORM SWIFTLET Aerodramus vanikorensis

Identified on several occasions among mixed-species groups of Aerodramus and Collocalia
spp. at 200-300 m on 19 March 2010 (MT & PDR). The birds were forced to fly low above
ground by heavy rain, enabling reasonably good views against a forested background.
Identification was based on the combination of larger size in direct comparison to Seram
Swiftlet A. ceramensis and Glossy Swiftlet C. es cidenta, and overall uniform brown plumage.
At least 40 were present. Prior to our observations, only Lambert (1994) had reported this
species on Obi.

SERAM SWIFTLET Aerodramus ceramensis

Also on 19 March 2010, MT & PDR observed at least 12 swiftlets intermediate in size between
A. vanikorensis and C. esculenta, with glossy black upperparts, bold, clear-cut whitish rump
band and paler underparts with dirty white belly and undertail-coverts. Following the split
from Moluccan Swiftlet Aerodramus infuscata (Rheindt & Hutchinson 2007), we identified

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these as Seram Swiftlets, which taxon had been tentatively identified on Obi by Lambert
(1994) and our record suggests that it may be a regular visitor or resident.

WHITE-THROATED NEEDLETAIL Hirundapus caudacutus

On 18 March 2010, several small flocks totalling c.120 of this Palearctic migrant were
recorded at site B at 400 m (OP, WP, PDR, MT). Most were low over the forest canopy,
offering prolonged views in good early morning light. Key identification features including
the extensive white throat were observed, eliminating Purple Needletail H. celebensis
of Sulawesi and the Philippines and Silver-backed Needletail H. cochinchinensis, which
breeds in mainland Asia and winters south-east to Java (Chantler 1999). All were flying
west and might have been migrants. On 19 March 2010, small flocks totalling c.65 were
seen at scattered locations between 320 m and 420 m (PDR, MT). Our records are the first
for Obi and northern Maluku. Previous records from Sangihe, Sulawesi, Taliabu, Burn,
Banda, Lombok, Timor (Coates & Bishop 1997), Atauro (Trainor & Leitao 2007), Tanimbar
(J. A. Eaton pers. comm.) and Peleng (Rheindt et al. 2010) suggest that migrants can occur
anywhere in Wallacea.

BLYTH'S HORNBILL Rhyticeros plicatus

Singles and pairs regularly recorded in logged forest and forest edge from sea level to
400 m, with a single on a forest ridge at 800 m (all observers). No previous information
concerning elevational range on Obi (Coates & Bishop 1997, White & Bruce 1986).

RED-BELLIED PITTA Erythropitta erythrogaster obiensis

Seen twice and fairly commonly heard from c.100 m to 1,050 m. Most frequent in selectively
logged forest and also found in second growth and scrub. Several sound-recordings of
a singing bird obtained on 19 March (Fig. 9). The song comprised two slightly rising
notes, the first note rising more markedly than the second, while the latter is flatter at
the end. It was similar, but not identical, to the vocalisation on Halmahera (R. Drijvers in

3 -
2

n - ■ ■ m \ ' spasms * -

Time (sec)

2 -

1 -

0 -

„ , ; j 1 "i ■* i . . . i -ir- i

• ... ■ [ • .. . . . , 1 , ■ •. •

Time (sec)

Figure 9. Sonograms of Red-bellied Pitta Erythropitta erythrogaster rufiventris, Obi, 19 March 2010 (M. Thibault).
X-axis = time (one second per tick), y-axis = frequency (1 kHz per tick).

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Sharringa 2005, www.xeno-canto.org/89488). However, given that E. erythrogaster presents
considerable individual variation (Rheindt et al. 2010; pers. obs.), we are unable to comment
further on the distinctiveness of the Obi vocalisation. While our records match previous
statements that the species is common on Obi (Lambert 1994), they significantly extend
the upper altitudinal range on the island. E. e. obiensis was described by Hachisuka (1935),
who distinguished it from nifiventris, which occurs in most of northern Maluku, by an
allegedly paler blue breast, despite earlier statement that specimens from Obi did not differ
from nifiventris (Hartert 1903a). However most recent works treat obiensis as a synonym
of nifiventris (White & Bruce 1986, Lambert & Woodcock 1996, Dickinson & Dekker 2000,
Erritzoe 2003). A careful examination of specimens and thorough acoustic comparison are
required to ascertain the taxonomic position of the Obi population.

DUSKY MYZOMELA Myzomela obscura rubrotincta

Common at 200-1,210 m with a single near sea level. Our records extend the altitudinal
range on Obi, where it was previously unknown above 800 m (Coates & Bishop 1997). We
had several opportunities to closely observe and photograph rubrotincta, which is restricted
to Obi and Bisa. It has a bright reddish mantle, wings and tail, and a rosy red wash to the
head-sides and most of the underparts, albeit brighter on the breast-sides and flanks (Fig.
10). MT found the appearance of rubrotincta to be strikingly different from the nominate race
that he has observed in northern Australia, which is an overall greyish-brown bird. We also
found rubrotincta to differ markedly from simplex that we saw on Halmahera immediately
following our visit to Obi, which is a dark greyish bird, with reddish restricted to the outer
edge of the remiges and rectrices, underparts pale greyish brown, with some pinkish-brown
fringes on the breast feathers scarcely visible in the field (see http://orientalbirdimages.org/
search. pbp?Bird_ID=1342).

Figure 10. Dusky Myzomela Myzomela obscura rubrotincta, Obi, 13 March 2010 (M. Thibault).

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M. obscura comprises eight widely recognised races from northern Maluku to New
Guinea and northern Australia (Higgins et al. 2008, Gill & Donsker 2012). Races are
considered to comprise two groups, with obscura (northern Australia), harterti (islands
of Torres Strait and north-east Australia), fumata (New Guinea), nibrobrunnea (Biak) and
aruensis (Aru) comprising the 'nominate group', and mortyana (Morotai), simplex and
rubrotincta the 'simplex group' (Higgins et al. 2008). However, in the nominate group,
which mainly has overall dark grey-brown plumage, rubrobrunnea is remarkable as it has a
brownish-red wash (see www.birdforum.net/opus/images/8/82/4370Dusky_Myzomela.jpg)
and is also reported to have a reddish throat stripe (Higgins et al. 2008). These differences
induced Hartert (1903a) to consider it intermediate between simplex and rubrotincta, and
Mayr et al. (1939) to suggest that rubrobrunnea might constitute a species-level taxon. Based
on important plumage differences, we consider that rubrotincta should be treated as a
species and we suggest that molecular, morphological and perhaps acoustic analyses of
the M. obscura complex should be conducted as they might reveal that non-clinal variation
among island forms elsewhere in the northern Moluccas and on Biak include one or more
additional species-level taxa.

SULAWESI MYZOMELA Myzomela chloroptera (undescribed subspecies?)

On 13 March, an adult male, an immature male and at least one female / juvenile were
observed and photographed at 950 m at site E (MT). On 15 March, two females / juveniles in
the same area (OP, MT). On both occasions, they fed in low trees and dense second growth
bordering a logging track. These are the first confirmed records of M. chloroptera for Obi. It
is of note that a 'mostly red' Myzomela sp. seen in 1992 in the lowlands of Obi could refer to
this species, but the view was too brief for a description (F. R. Lambert pers. comm.).

M. chloroptera currently comprises four widely recognised subspecies: the nominate
from north and central Sulawesi, juga from south Sulawesi, eva from Salayar and
Tanahjampea, and batjanensis from Bacan (Coates & Bishop 1997, Higgins et al. 2008).
Populations recently discovered on Taliabu, Sula Islands (Davidson et al. 1991, Rheindt
2010) and Peleng, Banggai Islands (Rheindt et al. 2010) closely resemble the nominate,
although future research may reveal that they comprise one or two new subspecies.

Figure 11. Male Sulawesi Myzomela Myzomela chloroptera ssp. (presumably the same individual). Obi, 13
March 2010 (M. Thibault). Strongly patterned black-and-red upperparts, all-black scapulars, wing and
tail feathers lacking brownish tinge and pure red on head, throat and breast clearly demarcated from pale
greyish belly and whitish flanks strongly suggest it is a full adult. Underparts coloration strikingly differs
from batjanensis and also shows subtle differences compared to typical adults of the nominate race.

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Figure 12. From left to right: Myzomeln chloroptera batjanensis (three from Bacan), M. c. chloroptera (two from
Minahassa Peninsula, north Sulawesi), M. c. jugn (one from south-west peninsula of Sulawesi), M. c. evn (two
from Salayar and Tanahjampea, respectively) (M. Thibault / © The Natural History Museum, Tring).

The population we discovered on Obi fills a geographical gap between batjanensis and
the populations on Taliabu and Peleng. One of the birds photographed (Fig. 11) was an
adult male based on its strongly patterned black-and-red upperparts, pure red head, throat
and breast lacking any greyish (immature) feathers and clearly demarcated pale greyish
belly and whitish flanks. Preliminary comparisons can be made with other subspecies
of M. chloroptera. Compared to batjanensis (geographically proximate), the adult male
photographed on Obi was strikingly different, having a red (not greyish-olive) breast. The
red below did not extend to the belly and appeared more restricted than on typical adults
of the nominate (pers. obs.), although whether this pattern lies within individual variation
of the latter cannot be fully excluded. It had a whitish abdomen, with no trace of the fawn
grey-brown tinge supposedly distinctive of the nominate (Coates & Bishop 1997; Fig. 12).
Compared to geographically distant eva and juga, no significant plumage difference could
be detected. Compared to the (limited) photographic material available for populations
on Taliabu (Rheindt 2010; F. Rheindt unpubl.) and Peleng (F. Verbelen unpubk), red was
lacking on the upper belly and flanks, suggesting that it possibly represents an undescribed
taxon. Detailed morphological, acoustic and molecular analyses of the M. chloroptera group
are required, as already suggested by Rheindt (2010) and Rheindt et al. (2011).

PALE CICADABIRD Coracina ceramensis hoogerwerfi

Recorded at 300-1,220 m in logged and primary forest and forest edge, seldom in the
lowlands (three records below 700 m) and most commonly above 800 m (ten records).
Endemic to Maluku, the subspecies hoogerwerfi is restricted to Obi. Our observations
provide the first data on the elevational range of hoogerwerfi (Coates & Bishop 1997, White
& Bruce 1986) and add new information concerning its habitat preferences (Linsley 1995).

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RUFOUS-BELLIED TRILLER Lalage aurea

Common in logged forest and forest edge to 420 m. This monotypic species endemic to
northern Maluku is a lowland specialist. Our records slightly extend the altitudinal range
on Obi, where it was previously reported to 300 m (Coates & Bishop 1997).

COMMON GOLDEN WHISTLER Pachycephala pectoralis obiensis

Recorded at 300-1,220 m, most commonly above 700 m. P. pectoralis was reported from the
lowlands of Obi (Linsley 1995), but there were no previous data on the upper elevational
range of P. p. obiensis (White & Bruce 1986, Coates & Bishop 1997). The Common Golden /
Mangrove Whistler P. pectoralis / melanura complex, with 66 named populations, is one of
the most complex examples of avian geographic variation (Jonsson et al. 2008). The IOC
currently treats Obi populations as part of Black-chinned Whistler P. mentalis (northern
Maluku: Gill & Donsker 2012) but it seems preferable to await genetic and vocal analyses of
the complex before accepting this split.

CINNAMON-BREASTED WHISTLER Pachycephala johni

Common in logged forest, primary forest and forest edge, and also seen in regrowth and
scrub, at 300-1,200 m, with a single record near sea level. Our observations significantly
extend the altitudinal range given for this Obi endemic, which was previously recorded at
220-700 m (Lambert 1994).

Most authors (White & Bruce 1986, Coates & Bishop 1997, Boles 2007) placed johni
within P. griseonota, along with five additional subspecies differing significantly in their
plumage and potentially vocalisations (Coates & Bishop 1997, Rheindt et al. 2010), despite
the initial assignment of johni to species level by Hartert (1903a), who argued that it 'has

Figure 13. Male Cinnamon-breasted Whistler Pachycephala johni, Obi, 15 March 2010 (M. Thibault).

Marc Thibault et al. 104 Bull. B.O.C. 2013 133(2)

Figure 14. Sonograms of the songs of (A)
Cinnamon-breasted Whistler Pachycephala johni,
Obi highlands, 16 March 2010 (M. Thibault)
and (B-E) Drab Whistler P. griseonata, with
(B) P. g. griseonata, Kopiboto, Seram (R. O.
Hutchinson), (C) P. g. griseonata, Sawai, Seram (J.
A. Eaton), (D-E) P. g. examinata, Wamlana, Burn
(J. A. Eaton). X-axis = time (one second per tick),
y-axis = frequency (1 kHz per tick).

12 3 4

Time (sec)

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123456789 10 11

Time (sec)

1 -

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123456789 10 11

Time (sec)

6 -

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1 2 3 4 5 6 7 8 9 10 11 12 13 14

Time (sec)

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no very close ally' and that 'its bright rufous-cinnamon underside distinguishes it at once
from all others [Pnchycephala]' . We were impressed by the wholly rich rufous-cinnamon
underparts in male johni (Fig. 13), whereas male P. griseonatn is described as having whitish
(nominate, examinata, lineolata), greyish ( kuehni ) or ashy grey ( cinerascens ) throat, greyish
breast ( examinata , lineolata, cinerascens), tinged ochre (nominate) or brown (knelini) and belly
either rusty ochre (nominate), buffy ochre {kuehni, examinata), white ( lineolata ) or white
with slight ochre tinge ( cinerascens ) (White & Bruce 1986, Boles 2007). Female johni is also
distinctive, being similar to the male, albeit duller and with black streaks on the breast.

The song of johni, previously undescribed, was heard several times and sound-recorded
once. It includes whistling strophes, of 0. 5-2.0 seconds, comprising 3-10 varied notes
delivered at 1. 7-5.5 kHz, which produce a varied tuneful song (Fig. 14A). A preliminary
comparison reveals that it differs markedly from P. g. griseonata on Seram (Fig. 14B-C), P. g.
examinata on Buru (Fig. 14D-E) and P. g. cinerascens on Halmahera (G. Wagner, www.xeno-
canto.org/23102), which mainly utter monotonous phrases. It also differs from the unique
sample from Peleng, possibly attributable to P. g. lineolata (B. Demeuleemester, avocet.
zoology.msu.edu/recordings/12430), and to at least some of the vocalisations described for
kuehni on the Kai Islands (Coates & Bishop 1997). Based on plumage and song, we support
the view (Hartert 1903a, Gill & Donsker 2012) that johni should be regarded as a species and
recommend comprehensive morphological and acoustic studies of the P. griseonata complex
as it might reveal the presence of other species-level taxa.

HAIR-CRESTED DRONGO Dicrurus hottentotus guillemardi

Common in various habitats below 400 m including small patches of degraded forest
mixed with coconut plantations. Much scarcer at higher elevations, with only four records
at 800-1,220 m. Our records significantly extend the altitudinal range on Obi, where
it was previously unrecorded above 800 m (Coates & Bishop 1997). D. h. guillemardi is
endemic to Obi and Bisa (Coates & Bishop 1997). It is one of many allopatric forms of
Dicrurus in the Philippines-Wallacea-New Guinea that are currently lumped in one of two
polytypic species, Hair-crested and Spangled Drongos D. bracteatus. However, important
morphological, behavioural or vocal variation among these numerous forms suggest that
more detailed study might identify 'new' species (Rheindt & Hutchinson 2007, Rocamora
& Yeatman-Berthelot 2009, Rheindt et al. 2010). Future visitors to Obi and Bisa should
concentrate on obtaining good recordings of the vocalisation of D. h. guillemardi as these
would be useful for a reappraisal of the taxonomy of the D. hottentotus / bracteatus complex

NORTHERN FANTAIL Rhipidura rufiventris obiensis

Commonly recorded from sea level to 1,210 m with most records in degraded forest and
edge. Our records significantly extend the upper altitudinal range on Obi as it was not
previously recorded above 550 m (Coates & Bishop 1997). R. r. obiensis is restricted to Obi
and Bisa. R. rufiventris exhibits discrete variation between islands not following clinal
trends; its taxonomy is complicated and unclear, and it has been suggested that DNA
analysis might conclude that many races warrant species status (Rheindt & Hutchinson
2007). R. r. obiensis has a distinctive combination of plumage features not shared by any
neighbouring populations of R. rufiventris in Wallacea, including broad white fringes to the
secondaries and broadly white-tipped outer rectrices (Boles 2006).

RUFOUS FANTAIL Rhipidura rufifrons torrida

Only five singles and one record of a pair, all restricted to a narrow altitudinal zone at
760-1,000 m, at sites D and E (MT & WP). The range of R. r. torrida includes Halmahera,

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Ternate, Bacan and Obi. Our observations apparently constitute the first records on Obi
since one collected by Waterstradt in 1902 (Hartert 1903a) and there were no previous data
on the elevational range on Obi (Coates & Bishop 1997, White & Bruce 1986).

MOLUCCAN MONARCH Symposiachrus bimaculatus diadematus

The diadematus subspecies of this northern Maluku endemic is restricted to Obi and Bisa.
Observed at 260-1,030 nr, with all but one record above 700 m. Widespread in second
growth and forest edge, usually associating with mixed-species flocks. The occurrence of
M. trivirgatus is already well known in the lowlands of Obi, where it was most recently
reported by Linsley (1995). Our survey, however, extends the altitudinal range for the
species, as it is reported to just 850 m on Halmahera (Poulsen & Lambert 2000).

MOLUCCAN FLYCATCHER Myiagra galeata galeata

Recorded in forest edge, secondary forest and scattered trees near coconut plantations,
from sea level to 870 m. Moderately common in the lowlands of Obi, but no published data
concerning its upper altitudinal limit (Lambert 1994, Linsley 1995, Coates & Bishop 1997).

SHINING FLYCATCHER Myiagra aiecto alecto

Seen nine times and heard once. Most records were between sea level and 600 m, but also
seen at 750 m and 850 m. Mostly observed near rivers and forest edge, but one in a dry
gully in secondary forest at 750 m. Although P. alecto is mentioned from the lowlands of Obi
(Linsley 1995), our records provide the first data on its upper elevational limit.

PARADISE CROW Lycocorax pyrrhopterus obiensis

Very common from sea level to 1,220 m in a wide range of habitats including small patches
of degraded lowland forest mixed with coconut plantations and primary montane forest.
Our records significantly extend the altitudinal range on Obi, where it was previously
unknown above 800 m (Lambert 1994, Linsley 1995). L. p. obiensis is very distinct from
both the nominate and morotensis in plumage and measurements (Frith & Beehler 2006)
and has already been considered close to deserving species status (Cracraft 1992). It is also
more manucode-like in appearance (Lambert 1994; pers. obs.) and its vocalisation includes
several frequently delivered distinctive calls not given by the nominate, as previously
described by Lambert (1994). Research on the degree of insular differentiation and on the
evolution of local dialects was recommended by Frith & Beehler (2006).

NORTHERN GOLDEN BULBUL Thapsmillas longirostris lucasi

Commonly recorded in primary forest, logged forest and forest edge, at 260-1,210 m,
significantly extending the altitudinal range on Obi, where it was previously unknown
above 800 m (Coates & Bishop 1997). Golden Bulbul is a poorly studied group endemic
to Wallacea, where it is represented by nine recognised taxa. Fishpool & Tobias (2005)
recently split the group into two species, namely Northern Golden Bulbul T. longirostris,
which includes six subspecies in northern Maluku, on Obi, Sangihe, Sula, Banggai and
Togian islands, and Southern Golden Bulbul T. affinis, comprising three subspecies on Burn,
Ambon and Seram. Rheindt & Hutchinson (2007) went further in proposing to split T. affinis
into two species, Buru Golden Bulbul T. mysticalis and Seram Golden Bulbul T. affinis, which
was adopted by Gill & Donsker (2012).

Compared with the subspecies chloris (from Morotai to Bacan), which is geographically
its closest relative, lucasi has very distinctive bright yellowish plumage (whereas chloris is
markedly greener overall) and an extensive bright yellow loral spot ( chloris has a dark loral

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Figure 15. Northern Golden Bulbul Thapsinillas longirostris, with (A) T. I. lucasi, Obi, 10 March 2010 (M.
Thibault) and (B) T. 1. chloris, Halmahera, 23 March 2010 (M. Thibault).

spot extending below the eye) (Fig. 15). The bright yellow loral spot is a feature shared by
no other form of the Northern Golden Bulbul group except platenae, which is restricted to
the geographically distant island of Sangihe (Coates & Bishop 1997). T. 1. lucasi is a fairly
vocal taxon that frequently delivers three different call types, permitting many sound-
recordings to be made. In addition to whistled tweeeuip notes (Fig. 16A) and raucous calls
(Linsley 1995), it also gives a distinctive, piercing call comprising 2-3 notes that can be
transcribed pic pic piie (Fig. 16B). A preliminary comparison was made with incomplete sets
of recordings of the T. affinis / longirostris / mysticalis complex. These included recordings of
platenae on Sangihe (F. Verbelen, avocet.zoology.msu.edu/recordings/3347), aureus on the
Togian Islands (F. R. Lambert, www.xeno-canto.org/90082), hartertii on the Banggai Islands
(F. R. Lambert, www.xeno-canto.org/95775, www.xeno-canto.org/95360, F. Verbelen,
avocet.zoology.msu.edu/recordings/3344), longirostris on Taliabu (D. Verbelen, avocet.

1 1

Time (sec)

Figure 16. Sonograms of Northern Golden Bulbul Thapsinillas longirostris lucasi, with (A) whistled tweeeuip
calls, (B) pic pic piie calls. Obi, March 2010 (M. Thibault). X-axis = time (one second per tick), y-axis = frequency
(1 kHz per tick).

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zoology.msu.edu/recordings/7821), affinis on Seram (F. R. Lambert, avocet.zoology.msu.
edu/recordings/4805, avocet.zoology.msu.edu/recordings/4806) and mysticalis on Burn
(G. Wagner, www.xeno-canto.org/42281, F. R. Lambert, http://avocet.zoology.msu.edu/
recordings/4147). Recording of chloris (from Morotai to Bacan) were unavailable, but its
vocalisation is described as 'a rapid and repetitive chatter of semi-musical, moderately high
pitched notes' of 6-8 seconds duration (Coates & Bishop 1997). From these recordings and
the description of chloris, no vocalisation comparable to those we recorded of lucasi could
be found. Based on our preliminary comparisons, we suggest that lucasi is best regarded as
a species. Further studies may evidence that T. longirostris comprises several other species-
level taxa, as suggested by Rheindt et al. (2010).

UNIDENTIFIED MARTIN Riparia riparia / diluta

On 20 March 2010, a single Riparia sp. was seen at very close range in a flock of Barn
Swallows Hirundo rustica in Laiwui town (PDR & OP). The bird perched for several minutes
on an electric wire just above the observers. It was approximately one-fourth smaller than
the Barn Swallows perched beside it and had dark sandy brown upperparts and top of the
head, and a concolorous collar on its upper breast. The rest of the underparts, including
throat and chin, were creamy white. Its tail appeared relatively short and slightly forked
with the outer tail feathers having a rounded aspect. It took flight after a few minutes and
was not seen again. Despite the prediction by Coates & Bishop (1997) that R. riparia might be
recorded in Wallacea, we have failed to find any record in the literature, with the exception
of a R. riparia on Karakelong, Talaud Islands, on 8 November 2011 (Robson 2012). Our
record is apparently the first for Wallacea of the R. riparia / diluta species group. Based on
several features, including (a) absence of pale grey tone to the upperparts and breast-band
coloration, (b) solid, contrasting and complete dark beige-brown breast-band, (c) absence of
any particular contrast, greyish tinge or paleness on the head, (d) whitish throat contrasting
with dark beige-brown ear-coverts and lores, and (e) slight but clear tail fork, we believe it
was possibly a Sand Martin R. riparia rather than a Pale Sand Martin R. diluta. Overall size
and depth of the tail fork are of limited use on a lone vagrant (Loskot 2006, Schweizer &
Aye 2007). Identification in the held of eastern forms of both species is an unresolved issue
in South-East Asia (http://digdeepl962.wordpress.com/), also because the winter range of
Pale Martin is poorly known. Whereas Pale Martin is regularly recorded in winter only in
Hong Kong (http://digdeepl962.wordpress.com/), eastern Myanmar and eastern Tonkin
(Robson 2008), Sand Martin is more widespread in South-East Asia in the non-breeding
season (Robson 2008) and is a rare but regular winter visitor to Borneo (MacKinnon &
Phillips 1993, Myers 2009) and the Philippines (Kennedy et al. 2000, Allen et al. 2006). It is
also a straggler to Papua New Guinea (Coates & Peckover 2001). In view of this pattern of
occurrence in the Oriental and Australasian regions. Sand Martin is therefore more likely to
be recorded in Wallacea. Elsewhere in Indonesia it has been recorded only in Kalimantan
(http://burung-nusantara.org/birding-indonesia/checklist-birds-of-indonesia/).

BARN SWALLOW Hirundo rustica

Several seen at close range in Laiwui on 20-21 March 2010 (all observers). Easily separated
from Pacific Swallow H. tahitica by the combination of long outer tail-streamers and dark
chest contrasting with very pale creamy-white underparts. This northern migrant was
previously unrecorded on Obi, although its presence is unsurprising given that it is 'likely
to occur anywhere in Wallacea' (Coates & Bishop 1997).

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ISLAND LEAF WARBLER Phylloscopus poliocephalus waterstradti

We recorded this taxon at 360-1,210 m, in primary and logged forest, most commonly above
800 m. P. p. waterstradti is endemic to Bacan and Obi and very little is known of its ecology
except that it was previously recorded above 550 m on Obi (Lambert, 1994) and at 1,500-
2,100 on Bacan (Coates & Bishop 1997). Our records significantly extend the altitudinal
range on Obi and suggest more important range distinctions between Obi and Bacan.

The vocalisation of P. p. waterstradti has not been previously described. Several sound
recordings were obtained (Figs. 17A-B). Songs comprise various trilling strophes lasting
1. 1-2.2 seconds delivered every 1.4-3. 6 seconds. Strophes are most frequently introduced
by a single, lower pitched element at 3. 0-6.5 kFlz. Main trills are level or rising, at 3. 5-9.0
kHz and include repeated inverted-V notes or inverted-V notes alternated with bell-shaped
or more complex notes. These are somewhat reminiscent of the trilling song described for P.
p. ceramensis on Seram (Rheindt & Hutchinson 2007; T. Mark, www.xeno-canto.org/122063),
but the latter also gives more complex phrases alternating between rising and descending
series (M. Catsis, www.xeno-canto.org/38412) that were not heard on Obi. The song of P. p.
waterstradti is also close to Henrietta of Halmahera (Fig. 17C), but distinctly faster and higher
pitched. Our recordings of waterstradti differ markedly from those of all other P. poliocephalus

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Figure 17. Sonograms of songs of (A-B) Island Leaf Warbler Phylloscopus poliocephalus waterstradti, Obi, March
2010 (M. Thibault), (C) P. p. henrietta, Halmahera, October 2011 (R. O. Hutchinson), (D) P. (poliocephalus) taxon
novum, Taliabu, December 2012 (R. O. Hutchinson). X-axis = time (one second per tick), y-axis = frequency
(1 kHz per tick).

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subspecies for which recordings or sonograms are available, i.e. everetti on Burn (Rheindt &
Hutchinson 2007), avicola on Kai (F. R. Lambert, avocet.zoology.msu.edu/recordings/5420),
misoriensis on Biak (B. van Balen, www.xeno-canto.org/75914), giulianettii from central and
south-east New Guinea (I. Woxvold, www.xeno-canto.org/87539) and becki from the eastern
Solomons (D. Gibbs, www.xeno-canto.org/70658). The song of waterstradti also differs
notably from that of 'Taliabu Leaf Warbler' Phylloscopus taxon novum (Fig. 17D), recently
discovered on Taliabu (Davidson et al. 1991, Rheindt 2010) and from 'Peleng Leaf Warbler'
Phylloscopus taxon novum (F. Lambert, www.xeno-canto.org/95794), recently discovered on
Peleng (Rheindt et al. 2010), but is reminiscent in structure to some variant trilling songs of
P. sarasinorum nesophilus from central Sulawesi, although the latter are at a markedly lower
frequency (various recordists, www.xeno-canto.org/species/Phylloscopus-sarasinorum).
This suggests that P. poliocephalus waterstradti might be best treated as a species. However,
detailed taxonomic research into the P. sarasinorum / poliocephalus complex is needed as
it might reveal a number of species-level taxa, as suggested by Rheindt & Hutchinson
(2007) and Rheindt et al. (2010). Furthermore, a comprehensive study should include vocal
comparisons between populations on Obi and Bacan.

CREAM-THROATED WHITE-EYE Zosterops atriceps

This inconspicuous but very vocal white-eye was occasionally seen and commonly heard
at 700-1,200 m in primary and logged forest, forest edge and second growth, with records
down to 400 m (all observers). Our observations significantly extend the altitudinal range on
Obi, where it was previously unknown above 700 m (Coates & Bishop 1997). Surprisingly,
it was first recorded on Obi in 1992 when Lambert (1994) found it uncommon and stated
that 'the birds most closely resemble the nominate subspecies of Bacan with greyish heads
contrasting with the bright green upperparts'. From our field experience, white-eyes on Obi
show striking variation that might be age-related, including in bill size and throat colour,
the latter ranging from creamy white to grey. We failed to notice the greyish tone to the
head mentioned by Lambert, but once obtained photographic evidence of the fuscous-
olive tinge to the crown typical of nominate atriceps (Fig. 18). Vocalisations of birds on
Obi have not been described. The song comprises moderately sweet whistles, strongly
reminiscent of fuscifrons from Halmahera, although apparently more variable in duration
(Coates & Bishop, 1997), with phrases ranging up to eight seconds (Fig. 19). A more detailed
investigation including morphological, vocal and molecular analyses is required to confirm
the taxonomic position of the Obi population.

Figure 18. Cream-throated White-eye Zosterops atriceps (atriceps?),' two different individuals, Obi, 16 March
2010 (M. Thibault); note variation in throat coloration.

Marc Thibault et al.

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Bull. B.O.C. 2013 133(2)

8 -

6 -

4 -

i, I

2 -

0 -i

Time (sec)

Figure 19. Sonogram of song of Cream-throated White-eye Zosterops ntriceps, Obi, 14 March 2010 (M.
Thibault). X-axis = time (one second per tick), Y-axis = frequency (1 kHz per tick).

TURQUOISE FLYCATCHER Eumyias panayensis obiensis

Recorded just twice in logged forest at 800 m at site D and 840 m at site E, with photographs
taken by MT. E. p. obiensis is endemic to Obi (Coates & Bishop 1997). White & Bruce (1986)
traced only two specimens, from 650 m, while Lambert (1994) found it uncommon at 250-
700 m, and Linsley (1995) failed to record it. Our observations slightly extend the altitudinal
range of this taxon and support the view that it is uncommon.

HALMAHERA FLOWERPECKER Dicaeum schistaceiceps

Recorded five times in primary forest, logged forest and edge at 300-1,210 m (all observers).
D. schistaceiceps is a recent split from D. erythrothorax (Rheindt & Hutchinson 2007). Known
elevation is 0-710 m on Halmahera (Poulsen & Lambert 2000), 0-950 m on Bacan (Lambert
1994) and 180M00 m on Obi (Lambert 1994). Our observations significantly extend the
altitudinal range for this species, and suggest that it is fairly common and rather evenly
distributed throughout its elevational range on Obi.

BLACK SUNBIRD Leptocoma sericea

Most commonly recorded from sea level to 500 m, less common at 500-1,000 m with one
record of two birds at 1,200 m (MT). No previous data concerning elevational range on Obi
(White & Bruce 1986, Coates & Bishop 1997).

OLIVE-BACKED SUNBIRD Cinnyris jugularis

We provide the first data concerning elevational range on Obi (Coates & Bishop 1997, White
& Bruce 1986), with sight records from sea level to 420 m.

EURASIAN TREE SPARROW Passer montanns

Common at Laiwui and several at a logging settlement south of Soligi (all observers).
Actively expanding its range in Wallacea (C. Trainor pers. comm.), in Maluku it was first
recorded on Ambon (around 1900) and subsequently on Buru (1980), Ternate (1992) and
Halmahera (1994) (Coates & Bishop 1997). Ours are the first records for Obi and were 40
km distant from each other, suggesting that the species may now be widespread and well
established in the inhabited parts of the island.

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Unconfirmed records

MOLUCCAN MEGAPODE Eulipoa wallacei

On 15 March 2010, two megapodes were flushed in dense, shrubby vegetation adjacent to
a logging track at 940 nr at site E (MT & PDR). Both gave only very brief views in taking-
off but on the first bird, the whitish undertail and overall two-toned plumage with brown
upperparts and dark greyish underparts were seen, suggesting E. umllacei and seemingly
excluding all Megapodius including Dusky Scrubfowl M. freycinet, which is the only
scrubfowl previously known from Obi and which is more uniform blackish grey without
white or brown in the plumage. Moluccan Scrubfowl is near-endemic to Maluku where it
is known from many islands including Halmahera, Meiti, Ternate, Bacan, Buru, Boano,
Seram, Ambon and Haruku (Coates & Bishop 1997). Outside Maluku it occurs only on
Misool Island (West Papua). The lack of previous records from Obi (Coates & Bishop 1997,
White & Bruce 1986) seems remarkable given that the island is central to the species' range.
The elevation of our observation matches information provided by Coates & Bishop (1997),
who mentioned that the species 'inhabits hill and montane forest, generally above c. 750
m'. Tacking previous experience of E. umllacei and as we can only provide an incomplete
description, we stress that our record should be considered provisional. Future observers
should attempt to confirm or refute the species' presence on Obi.

Discussion

We provide the first records since 1982, first data on the vocalisations and basic
information on habitat for a very poorly known species of global conservation concern
( Scolopax rochussenii). These results will be potentially helpful for future research and
conservation action. We also present details of five new bird records for Obi, including one
(Myzomela chloroptera) that potentially involves an undescribed taxon. Three new records
are migrants (including Riparia riparia / diluta, a species group new to Wallacea) and one is
a new colonist whose presence is linked to anthropogenic habitat change ( Passer montanus).
In addition, another new bird record (Eulipoa umllacei) awaits confirmation.

The presence of a population of M. chloroptera in Obi fills a geographical gap between M.
c. batjanensis on Bacan and those populations recently discovered on the Banggai (Rheindt et
al. 2010) and Sula islands (Davidson et al. 1991). This record pertains to a generally montane
species previously unnoticed possibly because most field workers have operated mainly in
the lowlands, although recent colonisation is not fully excluded.

The occurrence of a species not mentioned in recent publications despite evidence
of earlier collectors is confirmed ( Columba vitiensis) and new material supportive of the
presence of a resident population of Cacomantis aeruginosus in Obi is presented.

Our field work failed to reveal the presence on Obi of an unambiguous, distinct
montane bird community, possibly as a result of genuinely limited and fragmented forest
above 800-900 m, as outlined previously for Halmahera (Poulsen & Lambert 2000). While
several resident species were only encountered above 700 m ( Columba vitiensis, Myzomela
chloroptera, Rlupidura rufifrons, Eumyias panayensis), they are not considered strictly montane
elsewhere in their ranges. However further field work focusing on as yet uncovered
elevations (1,200-1,500 m) is needed to confirm our preliminary assessment.

A number of resident landbirds, including two Obi endemics ( Accipiter hiogaster obiensis,
Coracina tenuirostris obiensis), were not encountered by us, suggesting that they are rare or
uncommon and / or that they are restricted to the lowlands, where little time was spent.

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Our field work uncovered substantial new albeit incomplete data pertaining to the
confusing taxonomic placement of several Obi birds. Novel acoustic data are suggestive
of biological species status for Thapsinillas longirostris lucasi and further support calls to
treat Pachycephala johni as a species. Field observations and photographs highlight the
distinctiveness of Myzomela obscura rubrotincta, which we consider deserves species status.
Finally, we highlight the need for a taxonomic reappraisal of several other endemic forms,
some of which were already mentioned by others. These include Ducula basilica obiensis,
Eos squamata obiensis, Geoffroyus geofffoyi obiensis, Dicrurns hottentotus guillemardi, Rhipidura
rufiventris obiensis, Lycocorax pyrrhopterus obiensis and Phylloscopus poliocephalus waterstradti.
Although most of our observations await further taxonomic work, they suggest that the
importance of the Obi avifauna at species level has probably been under-estimated.

Acknowledgements

We are indebted to the MAVA Fondation pour la nature for funding our survey. Thanks are due to the
people of Soligi village for their hospitality and for guiding us to the central highlands of Obi. We are
grateful to H. van Grouw and R. Prys-Jones for affording access to the collections of the Natural History
Museum, Tring, and for permission to publish photographs of specimens. We thank R. O. Hutchinson for
permitting us to publish sonograms of his recordings of Drab Whistler from Seram (Fig. 14), Island Leaf
Warbler from Halmahera and 'Taliabu' Leaf Warbler (Fig. 17). We also thank J. A. Eaton for permitting us to
publish sonograms of some of his recordings of Drab Whistler from Seram and Burn (Fig. 14), as well as the
various recordists who have deposited material in the xeno-canto and AVoCet online sound libraries. MT
thanks F. E. Rheindt and F. Verbelen for sharing their photographs of Sulawesi Myzomela from Taliabu and
Peleng, and for discussing the identification and taxonomy of Moluccan Cuckoo. S. van der Mije provided
photographs of Sulawesi Myzomela specimens from the Netherlands Centre for Biodiversity Naturalis,
Leiden. We thank F. Steinhauser for allowing us to publish his photograph of Cinnamon-bellied Imperial
Pigeon, F. R. Lambert for unpublished data on his 'red' Myzomela sighting on Obi, }. A. Eaton for unpublished
data on White-throated Needletail from the Tanimbars, and Nick Brickie for information on logging licenses
on Obi. Finally, we are most grateful to J. A. Eaton, C. R. Trainor and an anonymous referee who made very
helpful comments that improved the manuscript. G. M. Kirwan is acknowledged for providing substantial
corrections during the editing process.

References:

Allen, D., Espanola, C. P., Broad, G., Oliveros, C. & Gonzalez, J. C. T. 2006. New bird records for the Babuyan
Islands, Philippines, including two first records for the Philippines. Forktail 22: 57-70.

Army Map Service (LU). 1968. Ambon, Indonesia; world (Asia) 1:1,000,000: Ambon; ser. 1301; SA 52, Ambon.

Army Map Service, Corps of Engineers, US Army, Washington DC.

Ashburton Minerals Ltd. 2012. 2012 Annual report, www.ashmin.com.au (accessed 7 January 2013).
Baptista, L. F., Trail, P. W. & Horblit, H. M. 1997. Family Columbidae (pigeons and doves). Pp. 60-243 in del
Hoyo, J., Elliott, A. & Sargatal, J. (eds.) Handbook of the birds of the world, vol. 4. Lynx Edicions, Barcelona.
BirdLife International. 2011a. Species factsheet: P till nopus granulifrons. www.birdlife.org (accessed 25
February 2011).

BirdLife International. 2011b. Species factsheet: Scolopax rochussenii. www.birdlife.org (accessed 4 March

2011).

BirdLife International. 2012. Species factsheet: Lorius garrulus. www.birdlife.org (accessed 5 April 2012).
BirdLife International. 2013. Important Bird Areas factsheet: Pulau Obi. www.birdlife.org (accessed 7
January 2013).

Bishop, K. D. 1992. New and interesting records of birds in Wallacea. Kukila 6: 8-34.

Boles, W. E. 2006. Family Rhipiduridae (fantails). Pp. 200-243 in del Hoyo, J., Elliott, A. & Christie, D. A. (eds.)

Handbook of the birds of the world, vol. 11. Lynx Edicions, Barcelona.

Boles, W. E. 2007. Family Pachycephalidae (whistlers). Pp. 374M37 in del Hoyo, J., Elliott, A. & Christie, D.

A. (eds.) Handbook of the birds of the world, vol. 12. Lynx Edicions, Barcelona.

Chantler, P. 1999. Family Apodidae (swifts). Pp. 388^157 in del Hoyo, J., Elliott, A. & Sargatal, J. (eds.)

Handbook of the birds of the world, vol. 5. Lynx Edicions, Barcelona.

Coates, B. J. & Bishop, K. D. 1997. A guide to the birds of Wallacea - Sulawesi, the Mollucas and Lesser Sunda
islands, Indonesia. Dove Publications, Alderley.

Coates, B. J. & Peckover, W. S. 2001. Birds of New Guinea and the Bismarck archipelago. Dove Publications,
Alderley.

Collar, N. J. 1997. Family Psittacidae (parrots). Pp. 280-477 in del Hoyo, J., Elliott, A. & Sargatal, J. (eds.)
Handbook of the birds of the world, vol. 4. Lynx Edicions, Barcelona.

Marc Thibault et al.

114

Bull. B.O.C. 2013 133(2)

Collar, N. J. & Andrew, P. 1988. Birds to watch: the ICBP world check-list of threatened birds. International Council
for Bird Preservation, Cambridge, UK.

Cracraft, J. 1992. The species of the birds-of-paradise (Paradisaeidae), applying the phylogenetic species
concept to a complex pattern of diversification. Cladistics 8: 1-43.

Davidson, P. J., Lucking, R. S., Stones, A. J., Bean, N. ]., Raharjaningtrah, W. & Banjaransari, H. 1991. Report
on an ornithological survey of Taliabu, Indonesia. Univ. of East Anglia, Norwich.

Dickinson, E. C. & Dekker, R. W. R. J. 2000. Systematic notes on Asian birds. 4. A preliminary review of the
Pittidae. Zool. Verh. Leiden 331: 89-99.

Erritzoe, J. 2003. Family Pittidae (pittas). Pp. 106-161 in del Hoyo, J., Elliott, A. & Christie, D. A. (eds.)
Handbook of the birds of the world, vol. 8. Lynx Edicions, Barcelona.

Erritzoe, ]., Mann, C. F., Brammer, F. P. & Fuller, R. A. 2012. Cuckoos of the world. Christopher Helm, London.

Fishpool, L. D. C. & Tobias, J. A. 2005. Family Pycnonotidae (bulbuls). Pp. 124—251 in del Hoyo, J., Elliott, A.
& Christie, D. A. (eds.) Handbook of the birds of the world, vol. 8. Lynx Edicions, Barcelona.

Food & Agriculture Organisation (FAO). 1981-82. National conservation plan of Indonesia. Eight vols. UNDP /
FAO FO/INS/78/061.

Frith, C. B. & Beehler, B. M. 2006. The birds of paradise: Paradisaeidae. Oxford Univ. Press.

Gill, F. & Donsker, D. (eds.) 2012. IOC world bird names (v. 3.2). www.worldbirdnames.org (accessed 10
January 2013).

Hachisuka, M. 1935. The birds of the Philippine islands with notes on the mammal fauna, vol. 2. London.

Hartert, E. 1903a. The birds of the Obi group. Central Moluccas. Novit. Zool. 10: 1-17.

Hartert, E. 1903b. The birds of Batjan. Novit. Zool. 10: 43-64.

Hartert, E. 1925. Review of the genus Cacomantis Mull. Novit. Zool. 32: 164—174.

Heinrich, G. 1956. Biologische Aufzeichnungen iiber Vogel von Halmahera und Batjan. /. Orn. 97: 31MD.

Higgins, P., Christidis, L. & Ford, H. 2008. Family Meliphagidae (honeyeaters). Pp. 498-691 in del Hoyo, J.,
Elliott, A. & Christie, D. A. (eds.) Handbook of the birds of the world, vol. 13. Lynx Edicions, Barcelona.

Jansen, J. 2008. Pioneer of Asian ornithology - Heinrich Bernstein. BirdingASlA 10: 103-107.

Jany, E. 1954. Nachricht iiber eine Sammelreise nach den Inseln Batjan und Obi. /. Orn. 95: 429.

Jany, E. 1955. Neue Vogel-Formen von den Nord-Molukken. /. Orn. 96: 102-106.

Jonsson, K. A., Bowie, R. C. K., Moyle, R. G., Christidis, L., Filardi, C. E., Norman, J. A. & Fjeldsa, J. 2008.
Molecular phylogenetics and diversification within one of the most geographically variable bird species
complexes Pachycephala pectoralis/melanura. J. Avian Biol. 39: 473-478.

Kennedy, R. S., Gonzales, P. C., Dickinson, E. C., Miranda H. C. & Fisher, T. H. 2000. A guide to the birds of
the Philippines. Oxford Univ. Press.

Kennedy, R. S., Fisher, T. H., Harrap, S. C. B., Diesmos, A. C. & Manamtam, A. S. 2001. A new species of
woodcock (Aves: Scolopacidae) from the Philippines and a re-evaluation of other Asian/Papuasian
woodcock. Forktail 17: 1-12.

Lambert, F. R. 1993. Trade, status and management of three parrots in the North Moluccas, Indonesia: White
Cockatoo Cacatua alba, Chattering Lory Lorius garndus and Violet-eared Lory Eos squamata. Bird Consent.
Intern. 3: 145-168.

Lambert, F. R. 1994. Notes on the avifauna of Bacan, Kasiruta and Obi, North Moluccas. Kukila 7: 1-9.

Lambert, F. R. & Woodcock, M. 1996. Pittas, broadbills and asities. Pica Press, Robertsbridge.

Lansley, P. S. & Fames, R. F. 2006. Nesting of the Scarlet-breasted Fruit Dove Ptilinopus bemsteini. Forktail
22: 139-140.

Linsley, M. D. 1995. Some bird records from Obi, Maluku. Kukila 7: 142-151.

Loskot, V. M. 2006. Systematic notes on Asian birds. 61. New data on taxonomy and nomenclature of the
Common Sand Martin Riparia riparia (Linnaeus, 1758) and the Pale Sand Martin R. diluta (Sharp &
Wyatt, 1893). Zool. Med. Leiden 80: 213-223.

MacKinnon, J. & Phillips, K. 1993. Afield guide to the birds of Borneo, Sumatra, Java and Bali. Oxford Univ. Press.

Mayr, E. & Meyer de Schauensee, R. 1939. Zoological results of the Denison-Crockett expedition to the South
Pacific for the Academy of Natural Sciences of Philadelphia, 1937-1938, Part I. The birds of the island
of Biak. Proc. Acad. Nat. Sci. Phil. 91: 1-37.

Mees, G. F. 1982. Bird records from the Moluccas. Zool. Med. Leiden 56: 91-111.

Myers, S. 2009. Afield guide to the birds of Borneo. New Holland, London.

Payne, R. B. 1997. Family Cuculidae (cuckoos). Pp. 508-607 in del Hoyo, J., Elliott, A. & Sargatal, J. (eds.)
Handbook of the birds of the world, vol. 4. Lynx Edicions, Barcelona.

Payne, R. B. 2005. The cuckoos. Oxford Univ. Press.

Poulsen, M. K. & Lambert, F. R. 2000. Altitudinal distribution and habitat preferences of forest birds on
Halmahera and Buru, Indonesia: implications for conservation of Moluccan avifaunas. Ibis 142: 566-586.

ProFauna Indonesia. 2008. Pirated parrots: ProFauna's investigation of the Indonesian parrot smuggling to
the Philippines. RSPCA, ProFauna International, www.profauna.org

Rheindt, F. E. 2010. New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the
preliminary documentation of two previously undiscovered taxa. Bull. Brit. Orn. Cl. 130: 33-51.

Marc Thibault et al.

115

Bull. B.O.C. 2013 133(2)

Rheindt, F. E. & Hutchinson, R. O. 2007. Taxonomic review. A photoshot odyssey through the confused avian
taxonomy of Seram and Buru (southern Moluccas). BirdingASIA 7: 18-38.

Rheindt, F. E., Verbelen, F., Dadang Dwi Putra, Rahman, A. & Indrawan, M. 2010. New biogeographic
records in the avifauna of Peleng Island (Sulawesi, Indonesia), with taxonomic notes on some endemic
taxa. Bull. Brit. Orn. Cl. 130: 181-207.

Robson, C. 2008. Birds of South-East Asia. New Holland, London.

Robson, C. 2012. From the field. BirdingASIA 17: 123-128.

Rocamora, G. J. & Yeatman-Berthelot, D. 2009. Family Dicruridae (drongos). Pp. 172-227 in del Hoyo, ].,
Elliott, A. & Christie, D. A. (eds.) Handbook of the birds of the world, vol. 14. Lynx Edicions, Barcelona.

Salomonsen, F. 1967. Family Meliphagidae, honeyeaters. Pp. 338-450 in Paynter, R. A. (ed.) Check-list of birds
of the world, vol. 12. Mus. Comp. Zool., Cambridge, MA.

Schweizer, M. & Aye, R. 2007 Identification of the Pale Sand Martin Riparia diluta in Central Asia. Alula 4:
152-158.

Sharringa, J. 2005. Birds of tropical Asia 3.0. DVD-ROM. Bird Songs International, the Netherlands.

Stattersfield, A. J., Crosby, M. J., Long, A. J. & Wege, D. C. 1998. Endemic Bird Areas of the world: priorities for
biodiversity conservation. BirdLife International, Cambridge, UK.

Streseman, E. 1931. Vorlaufiges iiber die ornithologischen Ergebnisse der Expedition Heinrich 1930-1931. V.
Zur Ornithologie von Halmahera und Batjan. Orn. Monatsb. 39: 167-171.

Taylor, P. M. 1990. The folk biology of the Tobelo people: a study in folk classification. Smiths. Contrib.
Anthropol. 34.

Tebb, G., Morris, P. & Los, P. 2008. Spotlight on Sulawesi. New and interesting bird records from Sulawesi
and Halmahera, Indonesia. BirdingASIA 10: 67-76.

Tobias, J. A., Seddon, N., Spottiswoode, C. N., Pilgrim, J. D., Fishpool, L. D. C. & Collar, N. J. 2010.
Quantitative criteria for species delimitation. Ibis 152: 724— 746.

Trainor, C. R. & Leitao, P. J. 2007. Further significant bird records from Atauro Island, Timor-Leste (East
Timor). Forktail 23: 155-158.

Vetter, J. 2009. Impacts of deforestation on the conservation status of endemic birds in the North Maluku
Endemic Bird Area from 1990-2003. M.Sc. thesis. Duke Univ., Durham, NC.

White, C. M. N. & Bruce, M. D. 1986. The birds of Wallacea (Sulawesi, the Moluccas and Lesser Sunda Islands,
Indonesia): an annotated check-list. BOU Check-list no. 7. British Ornithologists' Union, Tring.

Addresses: Marc Thibault, Mas Le Barret, Route de Port-Saint-Louis, 13 200 Arles, France, e-mail: marc.
thibault@freesbee.fr. Pierre Defos du Rau, Chemin de la Garrigue, 30700 Uzes, France, e-mail:
pierre.defosdurau@oncfs.gouv.fr. Olivier Pineau, Le Sambuc, 13200 Arles, France, e-mail: pineau@
tourduvalat.org. Wesley Pangimangen, Lelipang, Tamako 95855, Sangihe, North Sulawesi, Indonesia,
e-mail: wesley_rainbow@yahoo.co.id

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The taxonomic status of the Black-shouldered Nightjar
Caprimulgus nigriscapularis Reichenow, 1893

by H. D. Jackson

Received 13 December 2012

Summary. — The current status of Black-shouldered Nightjar Caprimulgus
nigriscapularis is based on relatively minor differences in voice and plumage from
Fiery-necked Nightjar C. pectoralis. These differences are considered by some
authorities to be no more than geographic or individual variation. Presented here
are mensural data, previously largely overlooked, comparing C. nigriscapularis
with the four races of C. pectoralis. The mean values of five key characters show
clinal variation. Analyses of variance carried out on nine other key characters (in
wing, tail, head and foot) reveal that, except for wing length, the null hypothesis,
linking C. nigriscapularis with C. p. shelleyi, C. p.fervidus, C. p. crepusculans and C. p.
pectoralis, cannot be rejected. Consequently, the conclusion is that the two current
species are conspecific. Black-shouldered Nightjar being the northern race of Fiery-
necked Nightjar.

Black-shouldered Nightjar Caprimulgus nigriscapularis was described as a new species
by Reichenow (1893) and included in his Die Vogel Afiikas (Reichenow 1902-03), which also
recognised South African Nightjar C. pectoralis Cuvier, 1816, and Fiery-necked Nightjar C.
fervidus Sharpe, 1875, as species, while treating Shelley's Nightjar C. shelleyi Bocage, 1879,
as a race of Fiery-necked Nightjar. Sclater (1924) retained C. pectoralis, C. fervidus and C.
nigriscapularis as species in his standard work Systema avium Aethiopicarum but made no
mention of C. shelleyi.

Chapin (1932) noted that 'Pairs of species distinct enough to be granted separate
binomials by most systematists, and which are still mainly separated by the forest belt, at
least in western Africa, include C. nigriscapularis and C. fervidus.' Grant & Mackworth-Praed
(1937) decided that 'As C. nigriscapularis has the same general colour appearance and the
same amount of white in the tail of males as C. fervidus it must be placed as a race of that
species.' Chapin (1939) went further, treating fervidus and nigriscapularis as conspecific with
C. pectoralis, on the basis of similarities in plumage and song. Mackworth-Praed & Grant
(1957, under Addenda and Corrigenda) accepted this arrangement, as did White (1965).

Chappuis (1981) found that nigriscapularis and fervidus have very stereotyped song
phrases throughout their distribution, with significant and constant differences separating
them. He felt that, in the absence of sympatry, the status of superspecies should be
preferred to that of race, since it is possible that the song differences may be insufficient
to prevent hybridisation should they come together again. Fry (1988), in a major review of
the skull morphology, song characteristics and systematics in general of African nightjars,
concluded that C. nigriscapularis is a species, separable from C. pectoralis, based mainly on
vocal differences.

In The birds of Africa Fry & Harwin (1988) distinguished C. nigriscapularis specifically
from C. pectoralis on the basis of evidently constant song differences, but also provided a
suite of plumage characters to separate them in the hand. Louette (1990) tested the colour
criteria proposed in The birds of Africa on Zaire (now Democratic Republic of the Congo =
DRC) material in the Royal Museum for Central Africa (Tervuren, Belgium) and found that

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none holds true. Dowsett & Dowsett-Lemaire (1993), contrary to Chappuis (1981) and Fry &
Harwin (1988), believed that the voices of C. nigriscapularis and C. pectoralis are sufficiently
alike to point to them being conspecific. They felt that Fry and Harwin had overlooked the
natural geographic variation in voice within each form, which is at least as great as that
between the two forms (italics mine). See also below under Discussion.

Clancey's (1994) review of the two austral races of C. pectoralis found that fervidus was
composite, requiring that three subspecies be recognised, fervidus, shelleyi and a new form,
which he named crepusculans. Cleere (1995) re-examined the entire Fiery-necked Nightjar
group and recognised two species, separated mainly by vocal differences, namely C.
pectoralis, with four subspecies (pectoralis, fervidus, shelleyi and crepusculans), and monotypic
C. nigriscapularis. This arrangement was adopted in both major monographs of the
Caprimulgiformes (Cleere 1998, Holyoak 2001). However, Cleere (1998) noted that some
authorities consider the two species to be conspecific, and Holyoak (2001: 35) explained that
he provisionally treated as species a few controversial forms, such as C. nigriscapularis, in
order to present all relevant data separately from those pertaining to their close allies, rather
than from any conviction that they deserve species rank.

Cleere (1995) suggested that the ranges of the two species may well overlap in parts
of central and southern DRC. However, this was based on a misidentification by Chapin
of a single female specimen from Matadi (Dowsett-Lemaire & Dowsett in litt. 2013). Both
monographs (Cleere 1998, Holyoak 2001) treated their distributions as allopatric. There is a
possible overlap in Rwanda, where Anderson (2012) found and photographed a pectoralis
north-west of a probable nigriscapularis. The latter is a rufous morph and difficult to identify
with certainty.

My examination of the relevant plumage descriptions and / or voice sonograms
published by Fry (1988), Fry & Harwin (1988), Dowsett & Dowsett-Lemaire (1993), Cleere
(1995, 1998) and Holyoak (2001) leads me to question whether the noted differences are
sufficient to warrant specific status for nigriscapularis, or whether they merely represent
geographic and / or individual variation within C. pectoralis. Presented here are mensural
data that I believe help to resolve the problem.

Methods

Key characters (Jackson 2000) were measured on 145 specimens of the five forms
involved: 43 C. pectoralis pectoralis, 42 C. p. crepusculans, six C. p. fervidus, 42 C. p. shelleyi and
12 C. nigriscapularis, listed south to north (for details of specimens see Appendix). Females,
juveniles, feathers in moult and damaged characters were excluded. The data therefore
refer to sound characters on full-grown males alone.

The lengths of the five outer primaries (ppl0-6) were measured by sliding a stopped
ruler under the closed wing until the stop met the bend of the wing, pressing the primaries
flat against the ruler and then reading the five measurements. The position of the wingbar
(an isolated white or buff patch about halfway along the feather) was measured on p9 by
taking the distance from the tip of the primary to the centre of the patch on the inner web.
The extent of the emargination on the leading edge of p9 was measured from the tip of the
primary to the point of flexure in the reverse curve, i. e. the neutral point between the inner
and outer curves (as illustrated in Jackson 1986).

The length of the inner rectrix (rl) was measured from the skin at the base of the
calamus to the tip of the feather. The calamus was exposed by parting the uppertail-coverts
so that the base was located visually, not by feel. The distance from the tip of rl to the tip
of r5 was measured with the tail in the closed position and the difference was then applied

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to the length of rl to derive the length of r5. All rectrix lengths are therefore relative to the
base of rl. The pale apical patches on r4 and r5 were measured by taking the maximum
dimension parallel to the rachis.

The length of the tomium was measured in a straight line from the tip of the bill to the
inside angle of the gape and the width of the gape was measured across the bill from gape
flange to gape flange. Tomium * gape provided an approximate mouth size.

The length of the tarsometatarsus was measured posteriorly from the intertarsal joint to
the base of the last complete scale before the divergence of the toes. The length of the middle
toe (t3) was measured from the base of the last complete scale on the tarsus to the tip of the
pectinated claw. Tarsometatarsus + t3 provided the overall length of the foot.

Characters showing clinal variation were set aside. The remaining characters were
each subjected to an analysis of variance (ANOVA one-way classification) to test, with
a probability of 0.05, the null hypothesis that the five forms represent either a single
population or five populations with equal means.

Results

Clinal variation. — Four characters (one each on wing, tail, head and foot) show clinal
variation, with mensural means decreasing from south to north (Table 1): the ratio of two
primaries in the wing formula (plO / p 7); wing / tail ratio (p9 / rl); relative mouth size
(tomium * gape) and overall foot length (tarsometatarsus + middle toe). Song durations also
appear to decrease clinally south to north (see below).

Analysis of variance. — An ANOVA was carried out on each of nine characters (Table
2); three on the wing, four on the tail, one on the head and one on the foot. The result for
wing length (p9) (F4 13] = 13.89, P = 1.79E-09, Table 2a) was the only one requiring that
the null hypothesis be rejected. It could not be rejected by the results for the other eight
characters: percentage emargination on p9 (F4 n| = 1.79, P = 0.135, Table 2b); position of
wingbar in relation to emargination (F4 = 1.90, P = 0.115, Table 2c); length of inner rectrix

rl (F4 106 = 1.99, P = 0.102, Table 2d); length of outer rectrix r5 (F4 n? = 2.25, P = 0.068, Table
2e); length of patch on outer rectrix (F( |30 = 0.64, P = 0.634, Table 2f); length of patch on r4
(F4 pij = 0.33, P = 0.855, Table 2g); length of tomium (F4 ]32 = 2.06, P = 0.090, Table 2h) and the
ratio tarsometatarsus to middle toe (%) (F4 UQ = 0.63, P = 0.639, Table 2i).

TABLE 1

Mensural characters showing latitudinal clinal variation in five Afrotropical nightjars, with measurement
means decreasing from South African Nightjar Caprimulgus pectoralis pectoralis in the south, through
Clancey's Nightjar C. p. crepusculans, Fiery-necked Nightjar C. p. fervidus and Shelley's Nightjar C. p. shelleyi
to Black-shouldered Nightjar C. nigriscapularis in the north. PlO is the outer primary, rl the inner rectrix.
Mouth = tomium * gape. Foot = tarsometatarsus + middle toe (t3). The sample sizes are shown in brackets

after the means.

Taxon

plO / p7

p9 / rl

Mouth (mm2)

Foot (mm)

pectoralis

1.007 (40)

1.347 (30)

814 (35)

39.79 (43)

crepusculans

1.005 (40)

1.334 (33)

807 (41)

38.17 (42)

fervidus

0.994 (06)

1.329 (06)

756 (06)

37.67 (06)

shelleyi

0.991 (34)

1.328 (29)

753 (39)

37.64 (42)

nigriscapularis

0.954 (09)

1.246 (09)

740 (11)

35.67 (12)

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TABLE 2

Results of ANOVA to test, with probability of 0.05, the null hypothesis that the five nightjars
(as in Table 1) represent either a single population or five populations with equal means.
Degrees of freedom ( df) between groups (upper) and within groups (lower) are shown. F
is the calculated F statistic, F crit the critical value that must be exceeded by F in order to
reject the null hypothesis. F values less than F crit are marked with an asterisk (*).

Taxon (n)

Mean

P-value

F crit

(a) Length of p9 (mm):

pectoralis (41)

163.1

13.894

1.79E-09

2.441

crepusculans (41)

160.2

131

fervidus (6)

165.8

shelleyi (36)

163.6

nigriscapularis (12)

152.8

(b) Emargination on p9 (%):

pectoralis (41)

41.6

1.790*

0.135

2.441

crepusculans (41)

42.4

131

fervidus (6)

42.1

shelleyi (36)

42.2

nigriscapularis (12)

42.0

on inner web of p9 (mm):

pectoralis (41)

4.7

1.898*

0.115

2.440

crepusculans (41)

5.4

132

fervidus (6)

4.2

shelleyi (37)

5.4

nigriscapularis (12)

6.2

(d) Length of inner rectrix rl (mm):

pectoralis (31)

121.6

1.987*

0.102

2.457

crepusculans (34)

120.0

106

fervidus (6)

124.8

shelleyi (31)

123.0

nigriscapularis (9)

122.2

(e) Length of outer rectrix r5 (mm):

pectoralis (37)

120.7

2.249*

0.068

2.449

crepusculans (34)

117.4

117

fervidus (6)

121.3

shelleyi (35)

120.4

nigriscapularis (10)

118.3

(f) Length of apical patch on r5 (mm):

pectoralis (40)

45.0

0.641*

0.634

2.441

crepusculans (39)

43.9

130

fervidus (6)

45.8

shelleyi (38)

44.5

nigriscapularis (12)

45.1

(g) Length of apical patch on r4 (mm):

pectoralis (39)

44.7

0.333*

0.855

2.442

crepusculans (40)

44.0

129

fervidus (6)

44.2

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Taxon (;i)

Mean

P-value F crit

shelleyi (38)

44.5

nigriscapularis (11)

45.4

(h) Length of tomium (nim):

pectoralis (39)

30.7

2.055*

0.090 2.440

crepusculans (41)

30.7

132

fervidus (6)

30.5

shelleyi (40)

30.1

nigriscapularis (11)

29.5

(i) Ratio of tarsometatarsus to middle toe (%):

pectoralis (43)

69.5

0.634*

0.639 2.436

crepusculans (42)

70.8

140

fervidus (6)

75.3

shelleyi (42)

70.0

nigriscapularis (12)

68.3

Discussion

In the absence of published relevant molecular studies, more traditional methods
must be used to re-examine the taxonomic status of Black-shouldered Nightjar, to resolve
whether it is the northern race of Fiery-necked Nightjar, or differs sufficiently to warrant
species status. Its current status as a species is based on vocal and morphological differences
(Fry 1988, Cleere 1995) that are relatively minor.

Plumage differences.— Coloration in nightjars is extremely variable within species,
sometimes geographically and very often individually, and such intraspecific variation is
often greater than differences between species (Jackson 2000). Consequently, most nightjars
are confusingly similar in appearance. For example, Fry (1988: 124) noted that some
specimens of Sombre Nightjar C. fraenatus are 'indistinguishable in the smallest quantitative
detail' from some specimens of the nominate race of C. pectoralis (italics Fry's).

Nightjar plumage patterns have evolved not as species-specific characters, but as
camouflage for the bird at rest; as an adaptation to the general environment and the
particular substrate upon which it roosts and nests. These plumage patterns, which may
not accurately reflect historical relationships, are of minimal diagnostic value in devising
identification keys; the pale patches in the wings and tails of some species do aid species
identification, but are more useful in separating the sexes, males usually having larger,
brighter patches than females (Jackson 2000).

Fry (1988: 105) commented that among African nightjars similarity of plumage is not a
reliable criterion unless accompanied by a suite of derived characters and / or by more or less
parapatric ranges. Fie noted (Fry 1988: 109) that C. nigriscapularis is darker than C. pectoralis
and that in males the white spot on the inner web of plO is vestigial in C. nigriscapularis.
Fry & Harwin (1988) added that C. nigriscapularis differs from C. pectoralis in not having a
small white mark near the corner of the mouth; in lacking a rufous morph, except perhaps
in respect of crown colour (they apparently overlooked Benson & Colebrook-Robjent
1977— see below); in having darker underparts; and colour tones that differ to a greater or
lesser degree over the rest of the plumage. However, as noted above, Louette (1990) found
that none of the plumage criteria proposed by Fry & Harwin (1988) as diagnostic for C.
nigriscapularis is true for DRC material that he examined.

Cleere (1995) noted that C. nigriscapularis tends to have less streaking on the crown than
the four races of C. pectoralis, affording it a rather pale-headed appearance; that the white

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spots on the four outer primaries of the male are generally smaller than on C. pectoralis ; and
that the blackish-brown lesser coverts always contrast markedly with the rest of the coverts,
hence the name Black-shouldered Nightjar. Dowsett-Lemaire & Dowsett (in litt. 2013) noted
that, in non-rufous forms, the uppertail colour is grey in pectoralis, brown in nigriscapularis.

The plumage differences noted above are relatively minor, no greater than those
used by Clancey (1994) to separate the four austral races of C. pectoralis, indicative of their
significance at subspecific level. They do not support specific status for Black-shouldered
Nightjar.

Vocal variation. —Nocturnal and crepuscular birds, such as nightjars, depend less on
plumage patterns and more on voice for conspecific and individual recognition. Nightjars
in song can readily be identified, each species having a highly characteristic song, which
makes it easy to distinguish between species occurring sympatrically in a given area, such
as, for example, southern Africa (Jackson 1986). From the sound-recordings on Ranft &
Cleere (1998) it is evident that neither a musical ear nor a sound spectrograph is necessary
to distinguish the songs of most Afrotropical nightjars, the differences being relatively
substantial.

However, the allopatric Freckled C. tristigma and Nubian Nightjars C. nubicus (C. n.
tamaricus alone: see Jackson 2002a) have remarkably similar songs, as do the allopatric
Abyssinian C. poliocephalus and Montane Nightjars C. ruwenzorii, and the allopatric
Fiery-necked and Black-shouldered Nightjars. Should any of these species-pairs occur
sympatrically, it would be difficult for field workers to distinguish between their songs,
as the differences are relatively minor. A further complication arises because there is
geographic and individual variation in voice, as in any other characteristic, of each species.
The songs of some C. pectoralis males in my Ranelia Farm (Cashel, Zimbabwe) study area
(Jackson 1985) were sufficiently different for me to recognise them individually.

The song durations of C. p. pectoralis (1.7-1. 8 seconds), C. p. fervidus (1.4—1. 6 seconds)
and C. nigriscapularis (1.4 seconds) (Fry 1988, Fry & Harwin 1988) appear to decrease clinally
south to north. Songs are repeated every five seconds ( pectoralis ), four seconds (fervidus ) or
5-8 seconds (nigriscapularis), while the monotonous whooting calls (see Jackson 2002b) are
repeated regularly, for at least 22 seconds, at a rate of 22 per ten seconds (pectoralis), or for
at least 12.5 seconds, at a rate of 41 per ten seconds (nigriscapularis) (Fry 1988, Fry & Harwin
1988). Cleere (1995) noted that the song of C. pectoralis registers within a frequency band
range of 0. 8-2.2 kHz, compared to 1.2-2. 2 kHz for C. nigriscapularis; he also noted that the
whooting calls of the latter are shorter and faster than in C. pectoralis.

Chappuis (1981) noted that significant and constant differences separate the songs of
C. (p.) nigriscapularis and C. (p.) fervidus, but felt it possible that song differences may not be
sufficient to prevent hybridisation should they meet. Dowsett & Dowsett-Lemaire (1993),
after examining a number of sonograms, concluded that song differences between these two
forms are no greater than the geographic variations within each form, and that the voices
of C. pectoralis and C. nigriscapularis are sufficiently alike to suggest they are conspecific.

Measurements.— While song is of no diagnostic value if dealing with a bird in the hand
or museum, measurements of key characters do usually provide reliable diagnoses. My
dichotomous identification keys to the nightjars of Africa were based almost entirely on
mensural characters, since plumage patterns are too variable (Jackson 1984, 2000).

When identifying Afrotropical nightjar species in the hand, the single most useful
character, with a low coefficient of variability, is the percentage emargination on p9
(Jackson 1984). My (Jackson 1993, 2002a, 2003) lack of support for the elevation of both C.
ruwenzorii and C. nigriscapularis to full species status was based on the evidence provided
by this character. The emargination percentages for C. p. poliocephalus (n = 57) and C. p.

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ruwenzorii (n = 39) are 43.3 ± 1.5 (40.1-47.6) and 43.6 ±1.4 (40.0-47.5), respectively (Jackson
2002a), suggesting that they are probably conspecific, a possibility that I am currently
investigating. The mean value of the emargination ratio in nigriscapularis (42.0%) is exactly
halfway between the lowest mean value ( pectoralis 41.6%) and the highest mean value
(crepusculans 42.4%) among races of C. pectoralis (Table 2), which suggests strongly that they
too are probably conspecific.

Clinal variation in some measurements (Table 1) and in song and whooting duration
(see above) tend to isolate C. nigriscapularis in the north, but despite this, the analysis of
variance results (Table 2) show clearly that the null hypothesis linking nigriscapularis to
the races of C. pectoralis cannot be rejected. This conclusion is supported by the evidence
provided by body mass data (Jackson 2003), the mean values (g) of the races being 51.2
(pectoralis), 48.3 (crepusculans), 54.0 (fervidus ), 49.8 (shelleyi) and 50.6 (nigriscapularis). The
mean of nigriscapularis in the north does not differ significantly (t = 0.1150, P > 0.1) from the
nominate race in the south, and matches well with other C. pectoralis races.

Sympatry.— Anderson (2012) presented photographic evidence of possible sympatry in
the south-east corner of Rwanda, with a probable record of C. nigriscapularis near Rusumo
on 13 December 2009, and a positive record of C. pectoralis near Kibungo, c.33 km to the
north-west, on 21 August 2010. It is difficult to identify the former with certainty, despite
the conspicuous blackish shoulders, because it is a rufous morph. According to Fry &
Harwin (1988) C. nigriscapularis does not possess a rufous morph, except perhaps in respect
of crown colour, while C. pectoralis does. However, they overlooked Benson & Colebrook-
Robjent (1977), who noted that erythrism may be frequent (but irregular) in nigriscapularis ;
ten adults in the Natural History Museum, Tring, all have some rufous in the tail, and half
of them, including three from Uganda, have the crown, and / or throat and breast, rufous.
One of the specimens they examined, from Sierra Leone, is illustrated in Anderson (2012).
The rufous plumage of the Rusumo bird does not serve to identify it either way.

Amadon & Short (1992) define sympatric species as taxa that occur in the same area
(range) at least in part and at least during the breeding season. The C. nigriscapularis
breeding season in central Africa appears to cover all months except October-November,
while the breeding season for C. pectoralis is August-December (Holyoak 2001 ). The dates of
the Rwandan photographs appear marginal to the breeding season concerned. No evidence
of breeding was found, and, since the records were eight months and 33 km apart, the
two birds were not actually in the same area in the same breeding season, so it could be
argued that there is no definite sympatry. Southern populations of C. pectoralis are partially
migratory, performing regular post-breeding movements, sometimes in response to high
rainfall (Clancey 1994, Vernon & Dean 2005), so the Kibungo C. pectoralis could have been
an off-season wanderer.

Amadon & Short (1992: 22) stressed the importance of interbreeding between two
taxa in determining that they belong to the same species. That C. nigriscapularis and C.
pectoralis may already have come together, and hybridised, is suggested by four specimens
(from Burundi, Rwanda and DRC) that Louette (1990; and in litt. to J. Anderson 2011— see
Anderson 2012) considered to be intermediate.

Conclusion

Mayr et al. (1953) recommended that it is preferable to treat doubtful allopatric
populations as subspecies. However, Prigogine (1985: 100) wrote 'I believe that it is useful
to give species status to taxa, even when the real taxonomic rank (species or subspecies) is
not fully demonstrated, with the object to draw the attention on a problem that needs more

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investigation.' This appears to be the current practice, and is the one followed by Holyoak
(2001) in his monograph of the Caprimulgiformes.

Prigogine (1985: 93) also stated that 'one has to be careful in giving too much weight
to song differences as a taxonomical criterium . . . and they must be evaluated critically and
compared with other informations of taxonomic signification, especially for allopatric taxa.'
This is the approach that I have followed here by considering not only the differences in
voice and plumage patterns, but also the available mensural data, which appear to have
been largely overlooked by previous authors.

The minor vocal and plumage differences do not appear sufficient to support the
elevation of C. p. nigriscapularis to species. The mensural data, in contrast, especially the
analyses of variance, but also the emargination and body mass data, provide compelling
evidence that C. nigriscapularis is no more than a subspecies of C. pectoralis. My conclusion,
therefore, is that Black-shouldered Nightjar is the northern race of Fiery-necked Nightjar, a
conclusion supported by the possible hybridisation between the two in Burundi, Rwanda
and the DRC (Touette 1990).

Acknowledgements

My thanks go to Johan Ingels for commenting on the original manuscript, and to the editor, Guy Kirwan,
and referees Bob Dowsett, Franchise Dowsett-Lemaire, Michel Louette and Nigel Cleere for constructive
suggestions that improved the paper. The museums listed in the Appendix are thanked for access to relevant
specimen material.

References:

Amadon, D. & Short, L. L. 1992. Taxonomy of lower categories - suggested guidelines. Bull. Brit. Orn. Cl.
112A: 11-38.

Anderson, J. 2012. Fiery-necked Nightjar Caprimulgus pectoralis and Black-shouldered Nightjar C.
nigriscapularis in Rwanda. Bull. Afr. Bird Cl. 29: 194-199.

Benson, C. W. & Colebrook-Robjent, J. F. R. 1977. Erythrism in the Fiery-necked Nightjar Caprimulgus
pectoralis. Bull. Brit. Orn. Cl. 97: 37-39.

Bocage, J. V. B. du 1879. Aves das possessoes portuguezas d' Africa occidental. J. Acad. Sci. Lisboa 24: 260-280.

Chapin, J. P. 1932. The birds of the Belgian Congo. Vol. 1. Bull. Amer. Mus. Nat. Hist. 65: 1-756.

Chapin, J. P. 1939. The birds of the Belgian Congo. Vol. 2. Bull. Amer. Mus. Nat. Hist. 75: 1-632.

Chappuis, C. 1981. Illustration sonore de problemes bioacoustiques poses par les oiseaux de la zone
ethiopienne. Alauda 49: 35-58.

Clancey, P. A. 1994. The austral races of the Afrotropical Fiery-necked Nightjar Caprimulgus pectoralis Cuvier
1816. Bull. Brit. Orn. Cl. 114: 48-55.

Cleere, N. 1995. The identification, taxonomy and distribution of the Mountain Nightjar Caprimulgus
poliocephalus / Fiery-necked Nightjar C. pectoralis complex. Bull. Afr. Bird Cl. 2: 86-97.

Cleere, N. 1998. Nightjars: a guide to nightjars and related nightbirds. Pica Press, Robertsbridge.

Cuvier, G. 1816. Le regne animal, vol. 1. De Lerville, Paris.

Dowsett, R. J. & Dowsett-Lemaire, F. 1993. A contribution to the distribution and taxonomy of Afrotropical and
Malagasy birds. Research Rep. No. 5. Tauraco Press, Liege.

Fry, C. H. 1988. Skulls, songs and systematics of African nightjars. Proc. Pan-Afr. Orn. Congr. 6: 105-131.

Fry, C. H. & Harwin, R. M. 1988. Order Caprimulgiformes. Pp. 155-197 in Fry, C. H„ Keith, S. & Urban, E. K.
(eds.) The birds of Africa, vol. 3. Academic Press, London.

Grant, C. H. B. & Mackworth-Praed, C. W. 1937. On some necessary changes in the status of some eastern
African nightjars. Bull. Brit. Orn. Cl. 58: 34-35.

Holyoak, D., T. 2001. Nightjars and their allies. Oxford Univ. Press.

Jackson, H. D. 1984. Key to the nightjar species of Africa and its islands (Aves: Caprimulgidae). Smithersia
4: 1-55.

Jackson, H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263-276.

Jackson, H. D. 1986. Identifying nightjars in southern Africa. Bokmakierie 38: 41-44.

Jackson, H. D. 1993. The English names of the Afrotropical nightjars (Caprimulgidae). Ostrich 64: 148-159.

Jackson, H. D. 2000. A new key to the identification of the Afrotropical nightjars (Caprimulgidae). Ostrich
71: 371-379.

Jackson, H. D. 2002a. Primary emargination as a guide to Afrotropical nightjar relationships. Ostrich 73:
69-71.

H. D. Jackson

124

Bull. B.O.C. 2013 133(2)

Jackson, H. D. 2002b. Comparison of vocal behaviour in two Afrotropical nightjars: a whistler and a churrer.
Ostrich 73: 173-174.

Jackson, H. D. 2003. On the body mass of the Montane and Pectoral Nightjars. Ostrich 74: 136-138.

Louette, M. 1990. The nightjars of Zaire. Bull. Brit. Orn. Cl. 110: 71-77.

Mackworth-Praed, C. W. & Grant, C. H. B. 1957. African handbook of birds. Birds of eastern and north eastern
Africa, vol. 1. Second edn. Longmans, London.

Mayr, E., Linsley, E. G. & Usinger, R. L. 1953. Methods and principles of systematic zoology. McGraw-Hill, New
York.

Prigogine, A. 1985. Recently recognized bird species in the Afrotropical region - a critical review. Proc. Intern.
Symp. Afr. Vert., Bonn: 91-114.

Ranft, R. & Cleere, N. 1998. A sound guide to nightjars and related nightbirds. Pica Press, Robertsbridge & British
Library, London.

Reichenow, A. 1893. Diagnosen neuer Vogelarten aus Central-Afrika. Orn. Monatsb. 1: 29-32.

Reichenow, A. 1902-03. Die Vogel Afrikas, Bd. 2. Neumann, Neudamm.

Sclater, W. L. 1924. Systema avium Aethiopicarum, vol. 1. British Ornithologists' Union, London.

Sharpe, R. B. 1875. Family Caprimulgidae. Pp. 83-90 in Layard, E. L. The birds of South Africa. New edn.
Quaritch, London.

Vernon, C. J. & Dean, W. R. J. 2005. Fiery-necked Nightjar Caprimulgus pectoralis. Pp. 264-266 in Hockey, P.
A. R., Dean, W. R. J. & Ryan, P. G. (eds.) Roberts— birds of southern Africa. Seventh edn. John Voelcker
Bird Book Fund, Cape Town.

White, C. M. N. 1965. A revised check list of African non-passerine birds. Govt. Printer, Lusaka.

Address: Unit 2, 6 Beer Court, Toowoomba, QLD 4350, Australia, e-mail: des.jackson@tadaust.org.au

Appendix: Museum specimens of Caprimulgus pectoralis analysed.

Museum acronyms: ALBM = Albany Museum, Grahamstown (South Africa); AMNH = American Museum
of Natural History, New York (USA); BMNH = Natural History Museum, Tring (UK); CMNH = Carnegie
Museum of Natural History, Pittsburgh (USA); DURM = Durban Museum (South Africa); ELM = East
London Museum (South Africa); FMNH = Field Museum of Natural History, Chicago (USA); KAFM =
Kaffrarian Museum, King William's Town (South Africa); LACM = Los Angeles County Museum (USA);
MNHN = Museum National d'Histoire Naturelle, Paris (France); NATM = Natal Museum, Pietermaritzburg
(South Africa); NMW = Naturhistorisches Museum, Vienna (Austria); PEM = Port Elizabeth Museum
(South Africa); RMCA = Royal Museum for Central Africa, Tervuren (Belgium); RMNH = Rijksmuseum van
Natuurlijke Historie (Naturalis), Leiden (Netherlands); TMP = Transvaal Museum, Pretoria (South Africa);
USNM = United States National Museum of Natural History, Washington DC (USA).

C. p. pectoralis {n = 43): ALBM no No. a, no No. b, 728, 733, 736, 1249, 2207, 2464, 4092; AMNH 633324,
633325, 633328; BMNH 1933.7.14.178, 52.3.8.6, 74.5.1.187, 90.12.16.58, 90.12.16.59; ELM 1866, 2816, 3222, 7950,
7958, 11352, 12764, 13115; KAFM B785, B786, B2902, B3544; MNHN 1858-393; PEM 1494/76, 1500/93, 1512/15;
RMNH 5, 6; TMP 2662, 2760, 2800, 3268, 7775, 14660, 26278; USNM 163823.

C. p. crepusculans (n =42): BMNH 1905.12.29.203, 1905.12.29.211, 1933.3.1.309-1933.3.1.311, 1933.7.14.179,
98.12.2.424; CMNH 148295; DURM 14502-14505, 16257, 16514, 19374, 20570, 23848, 29427; ELM 8128; FMNH
282662; NATM 2057, 2059, 2060, 2062, 2063, 2066; PEM 1484/31; RMNH 8; TMP 4706, 7794, 9001, 12702, 19065,
19796, 20083-20085, 27959, 27960, 35255; USNM 448384, 457818.

C. p. fervidus (n = 6): BMNH 1910.7.1.243; ELM 9635; FMNH 87253, 87254; LACM 63562; TMP 9213.

C. p. shelleyi (n = 42): AMNH 414149, 633338-633342, 633345, 633352, 633354, 633356-633358, 633360,
633361; BMNH 1932.5.10.730, 1935.10.9.104, 1935.5.11.2, 1939.2.25.484, 88.12.1.5; CMNH 109270, 109277,
147018; FMNH 100090, 205346, 220716, 224172, 224173, 262898; KAFM B2341; LACM 69601; MNHN 1947-
416, 1957-70, 1958-395, 1965-1051; NMW 23510, 35060; RMCA 119359; TMP 28342, 28343; USNM 351690,
448213, 448214.

C. p. nigriscapularis (n = 12): AMNH 158705, 262374-262377, 633329, 633330; BMNH 1926.5.3.305,
1953.1.8; FMNH 194471, 298274; LACM 64152.

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The extinct macaws of the West Indies, with special
reference to Cuban Macaw Ara tricolor

by James W. Wiley & Guy M. Kirwan

Received 7 January 2013

Summary. — The best-known species of West Indian macaw, and the only one for
which skin specimens exist, is Cuban Macaw Ara tricolor, although at least seven
and sometimes as many as 15 different species of Ara and Anodorhynchus have been
speculated to have formerly occurred in the Greater and Lesser Antilles. We review
available historical and prehistorical evidence for the existence of these different
species and conclude that only two or three, St. Croix Macaw Ara autochthones, A.
tricolor and perhaps Montserrat Macaw Ara. sp., and can be definitively admitted,
with reasonably strong evidence for another, Gosse's Macaw Ara gossei of Jamaica.

In addition to reviewing the causes and patterns of decline in West Indian macaws,
we provide a complete overview of available knowledge concerning A. tricolor,
including details of all extant specimens (19). In particular, we draw attention to a
commonly repeated error in much of the ornithological literature over the past six
decades concerning the final demise of Cuban Macaw, as well as to historical data
that suggest the species' range was wider than is often admitted.

Although only a few sketchy passages exist from those who saw live macaws in the
West Indies, those florid descriptions give us an appreciation of the former beauty, now
in large part lost, of those islands' diverse parrot fauna. Columbus described the macaws
of the Antilles as 'by far the most beautiful ornaments of the gloomy forest which covered
the land given up to nature' (Button 1774: 177). Du Tertre (1654: 294) said of West Indian
macaws, 'It is the finest sight in the world to see ten or twelve Macaws in a very green tree;
never are more charming colors displayed.' Gundlach (1893: 152) vividly described Cuban
Macaw Ara tricolor as he lamented the loss of these large, long-tailed birds. Tragically,
Cuban Macaw was just one of many spectacular psittacines lost in the region in historic
times.

Three groups of psittacids inhabited the West Indies at the time of 'discovery' by
Europeans: macaws (Ara and possibly Anodorhynchus), parrots (Amazona) and parakeets
( Aratinga ). Du Tertre (1667: 247) noted that 'These birds are so dissimilar according to the
grounds where they procure their food, that every island has its Parrots, its Aras, and its
Parroquets, different in size of body, in tone of voice, and in the tints of the plumage.'
Williams & Steadman (2001) suggested that the remarkable total of as many as 60 endemic
species of psittacids formerly occupied the West Indies but, as noted by Clark (1905a) and
Forshaw (1978), parrots and their kin are among the first to be exterminated from any given
locality, especially when confined to an insular habitat. Sadly, today only 12 species (three
parakeets, nine parrots) survive in the West Indies, a loss of up to 80% of psittacid species
from the region (Wiley 1991, Williams & Steadman 2001, Wiley et al. 2004).

The original psittacine fauna in the West Indies may have included as many as 15
species of macaws, with most Greater Antillean and several Lesser Antillean islands
supporting one or more indigenous species (Williams & Steadman 2001). Although none
survives today, macaws persisted into historic times on at least Cuba, Isla de Pinos, Jamaica,
Guadeloupe, Dominica and Martinique, but certainly some were lost during prehistoric

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human occupation of islands such as Montserrat. The fossil from the last-named island
has been variously identified as a new species, an unidentified macaw, or the same species
as that formerly present on Guadeloupe (see next section). Rothschild (1905, 1907a,b)
concluded that the region had seven macaw species in historic times and many subsequent
treatments follow his. However, it should be noted that Rothschild completely confounded
much of the evidence associated with insular parrots, for example making significant errors
in describing Mascarene taxa (Hume 2007; J. P Hume in litt. 2013). Most recently, Williams
& Steadman (2001) listed 15 species, with evidence ranging from poor (only hearsay) to
good (specimens or repeated, substantiated observations). Only four are represented by
specimens: three by subfossil skeletal remains and one by entire skins and fossil partial
skeletons (Walters 1995, Olson & Suarez 2008). The paucity of specimens and reliable
reports has led others to tally far fewer species. Olson & Suarez (2008) found no credible
evidence in support of each of the Antilles having one or two indigenous species of macaw;
they noted that the archaeological record, the strongest evidence, suggested perhaps just
two species in the region. Hume & Walters (2012) suggested perhaps only 3-4 species in
the Antilles, noting that 'otherwise the presence of endemic macaws on other West Indian
islands is extremely dubious.' Among the other 11 macaw species Williams & Steadman
(2001) cited, we consider one of good reliability, one of moderate reliability and six of poor
reliability; we present evidence that the three remaining macaws considered by Williams &
Steadman are synonyms of other species or probably did not exist.

Aside from Cuban Macaw and the two or three species evidenced by prehistoric bones,
all information is based on casual accounts by travelers, colonists and the few naturalists who
resided or passed through the islands between the 15th and 19th centuries. Confounding
some early reports are compilations by authors of regional or world faunas (e.g., Buffon),
who cobbled together information in a way that makes identification of species and their
location difficult if not impossible. Nevertheless, sufficient evidence exists to state that
macaws were formerly abundant on several islands. As examples of historical abundance,
Peter Martyr d'Anghera (1457-1525; in MacNutt 1912: 72) proclaimed that parrots and
macaws were 'as numerous in all these islands [West Indies] as sparrows or other small
birds are with us' [Europeans], whereas Hans Sloane (1660-1753; 1725: 297) reported 'the
Small Maccaw' as 'very common in the Woods' of Jamaica, and Bryan Edwards (1743-1800;
1806: 19), who resided on Jamaica most of his life, noted that 'The parrot, and its various
affinities, from the macaw to the parroquet ... are as plentiful in the larger islands of the
West Indies as the rook [Corvus frugilegus] is in Europe.'

Whereas some have argued that many of the macaws described by early explorers were
probably exotics, imported first by Amerindians and subsequently by European settlers,
statements of du Tertre (1654: 296), repeated by Labat (1742: 212), that each island had forms
distinct in plumage suggests that not all of the Antillean macaws were imported species
(Hume & Walters 2012). Further, Hume & Walters (2012) noted that the observations of du
Tertre and Labat on their gentle and docile nature contradicts that of mainland macaws.

Our objective here is to review the relevant historical information concerning the
presence and distribution of macaws in the West Indies, and to present such scant biological
information as is available. We focus on what is known of Cuban Macaw, presenting data
on specimens, and examine possible reasons for the extinction of that species, as well as
other Antillean forms.

Macaws in the West Indies

Unfortunately, so few data are available for most suggested species of West Indian
macaw that they are considered 'hearsay species' (Greenway 1967: 315). Hume &

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Walters (2012) noted that 'no other group of extinct birds has aroused such overzealous
misinterpretation' as Antillean macaws. Nevertheless, much archaeological investigation
remains to be done throughout the West Indies, particularly in the Lesser Antilles.
Compared to many mainland areas, fossil remains are uncommon in the West Indies and
little is known of vertebrate fossils, especially in the Lesser Antilles (Olson 1978, Woods
1989). Here we present additional evidence or clarify certain reports of macaws in the
Antilles.

Bahama Islands. — No evidence of a macaw to date.

Cuba.— One species, Cuban Macaw, Ara tricolor (Bechstein, 1811), survived until the
mid-19th century.

Jamaica.— At least one, perhaps three, native species, but identifications in many
original accounts are muddled. Williams & Steadman (2001) thought it plausible that
multiple species of Ara could have occurred on Jamaica, because it is a large island with
diverse habitats, although it should be noted that the much larger island of Cuba, with
considerably more varied habitats, apparently only ever supported one species of macaw.
Nevertheless, Jamaica is one of only two of the Antilles that supports two endemic
Amazona. Species attributed to Jamaica include Red-headed Green Macaw Ara erythrocephala
Rothschild, 1905, Red-tailed Blue-and-yellow Macaw A. erythrura Rothschild, 1907, and
Gosse's Macaw A. gossei Rothschild, 1905. Evidence is good for at least one species,
although no specimen exists, but poor for the other two species. All Jamaican macaws were
apparently extinct in the wild by the mid-19th century, as William T. March ( 1 804—72; 1863),
a native Jamaican and reliable observer, reported his last information on wild macaws was
in 1849.

Ara erythrocephala: Greenway (1967: 320) portrayed Red-headed Green Macaw as
an 'almost mythical bird,' given the tenuous circumstances upon which Rothschild
(1905) established it as a unique species. Rothschild (1905, 1907a) described the bird as
predominately green, with a red head, based on an account by Richard Hill (1794/5-1872;
in Gosse 1847: 261-262), a resident Jamaican and trustworthy observer. More than a
century earlier, Sloane (1725: 297) wrote that the 'Small Maccaw[s]'... 'are very common
in the Woods, and are eaten as Pigeons, but when young, are tamed, and kept as Parrots.'
Browne (1756: 472) described a 'small green long-tailed Parrot' as native to Jamaica; this was
probably a small macaw, because Browne also listed both native parrots ( Amazona collaria
and A. agilis) and parakeet ( Aratinga nano). But Browne does not describe a macaw similar
to Rothschild's A. erythrocephala.

Hill (in Gosse 1847: 263) noted that a captive Jamaican macaw (A. erythrocephala?)
had been purchased from hog-hunters by a Mr White of Oxford estate (Trelawny parish;
c.18°27'00"N, 77°34'58''W; 200 m). W. Osburn ( d . 1860; 1859) visited Trelawny specifically to
search for macaws, but found none and said the local sportsmen knew nothing of it. Osburn
(1859) referred to a long-term resident of Trelawny who reported 'constantly' seeing
macaws but, when pressed by Osburn, recalling seeing macaws 'about four or five times
in twenty-five years, ... always flying at a great height.' Osburn concluded that macaws
should be listed as 'occasional wanderers' to Jamaica.

Hill (in Gosse 1847: 261) wrote that macaws, 'with unquestionable certainty... are
occasionally, if not constantly, denizens of our mountain forests.' Further, Hill (in Gosse
1847: 262-263) reported macaws were found in 'the Black grounds' [Trelawny parish;
18°15'50"N, 77° 31'58"W; 738 m], 'the never failing resort of these Mexican Macaws [A.
militaris). I have been assured that several birds have been procured there. This is said to be
nearly as far eastward as they have been found.' Hill (ibid.) also noted that young, in 'the

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first year's plumage,' had been procured in the 'neighbourhood of Accompong Maroons'
(18°13'33"N, 77°45'14"W; 450 m), in the hills of St. Elizabeth parish.

Hill (in Gosse 1847: 261-262) thought the birds were Military Macaws A. militaris, which
he believed migrated from the mountain ranges of the continent, where they bred, and did
not return until the first part of the year. During its absence from the mainland. Hill thought
it was a casual visitor in Jamaica, where the macaw inhabited the mountains. Clark (1905b)
argued that it was unlikely that macaws would cross great extents of water, and thought
it more plausible that such individuals represented feral birds. He agreed with Cory (1889:
178) that 'it is not impossible that Ara militaris may have occurred in Cuba and Jamaica,
but it is improbable. The bird recorded as such was perhaps A. tricolor wrongly identified.'
Clark (1905b) suggested that at least some species attributed to the Greater Antilles by early
writers had apparently been brought from Middle or South America to Jamaica, and then
were re-shipped to the Old World as native to the island.

March (1863) observed the partial remains of a macaw shot by a settler near Maroon
Town (18°20'40"N 77°47'43"W; 440 m; St. James parish) in 1834. Although not a perfect
match with A. erythrocephala as described by Rothschild (1905), March's description from
memory ('the head and neck were a bright green with red in the forehead and chin, the tail
blue and red, and the wing blue and green.') is closest to A. erythrocephala among species
attributed to Jamaica by Rothschild (1907a). March's description, however, is much closer
to Military Macaw than to A. erythrocephala.

Snyder et al. (1987: 40) suggested that A. erythrocephala may represent Military Macaw
or Great Green Macaw A. ambiguus, both Middle American species. Lack (1976: 255) thought
the small green macaw described as A. erythrocephala was an endemic species derived from
Military Macaw.

Ara erythrura: Rothschild (1907a) named A. erythrura based on Charles de Rochefort's
(1605-83; 1658: 154) report of blue-and-yellow parrots, with all-red tails. Rothschild (1905)
first named the bird Anodorhynchus coerulens, and said it was from Jamaica, but later (1907a:
54) corrected the name to Ara erythrura nom. nov. and noted it was from 'One of the West
Indian Islands.' However, Greenway (1967: 319) regarded Rothschild's description as not
credible because de Rochefort had not visited Jamaica but, as others (e.g., de la Harpe 1759)
and even de Rochefort noted, took much of his material from du Tertre. Greenway (1967)
suggested that A. erythrura is a synonym of A. martinica [martinicus], a poorly documented
form supposedly from Martinique.

Sloane's (1725: 296) 'Great Maccaw' (' Psittacus Maximus cyanocroceus') is clearly closest
to Ara erythrura among those described for Jamaica by Rothschild, but Sloane's account
differed: 'On each side of the Head was a redish fleshy bare Membrane near the Eyes,
with some few black Feathers growing on it'; also, Sloane (1725) said the tail was blue, not
red as depicted by Rothschild (1907a: PI. 15). In fact, Sloane's bird more closely resembles
Blue-and-yellow Macaw A. ararauna. Browne (1756: 472), who called it 'the blue Macaw
of Edwards,' said he had observed one or two in St. Ann's (St. Ann's parish; c,18°26'25"N,
77°H'23"W), and 'yet keep some of the feathers of one that was killed there by me.'
Although Browne (1756) said it was a native of Jamaica ('tho' seldom catched there'), he
added that 'most of those that are generally seen about gentlemen's houses' had been
'introduced there from the main, where they are more common.' He said they were very
rare in Jamaica, where they 'keep in the most unfrequented inland parts.' Coke (1808: 390)
was probably referring to the same bird when he stated that 'The Blue Mackaw, a more
beautiful bird, though not so gaudy as the red, is a native of Jamaica, but not common, and
is wild in the woods, particularly in the parish of St. Anne.' Revd. Coward (in Gosse 1847:
261) observed macaws in flight about 1842 in 'a plain at the foot of a chain of mountains

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dividing [St. Elizabeth] parish from St fames [parish], and consequently nearly in the medial
line of the island.' Residents of the area familiar with these birds told Coward the macaws
were blue and yellow, which led Gosse (1847) to assume they were A. ararauna. In 1836,
March (1863) was told by Richard Elmas Breary, a resident of St. James, that he had seen
three blue-and-yellow macaws flying high overhead on a mountain road between St. James
and St. Elizabeth parishes.

Lack (1976: 253) thought the blue-and-yellow macaws were introduced, and cited the
fact that no writer since March claimed A. ararauna (A. erythrura ) for Jamaica. Possibly all
were feral A. ararauna, escaped or released after transport to Jamaica as cagebirds.

Ara gossei: Of the three species suggested for Jamaica, the strongest evidence exists for
Gosse's Macaw. Philip Henry Gosse (1810-88; 1847: 260), who resided there from December
1844 to July 1846, referred to it as 'Yellow-headed Macaw? Ara tricolor', noting that it differed
from the Cuban species and was possibly an undescribed species. Like A. tricolor, A. gossei
was predominately red, but its forehead was yellow rather than red. Anthony Robinson ( ob .
1768; in Gosse 1847: 260) examined a stuffed bird (now lost) shot about 1765 by a Mr Odell
in the mountains of Hanover parish, c.10 miles east of Lucea, west of Montego Bay. Gosse
(1847: 260) said it was very rare, but still it is surprising that neither Sloane (1725), who
resided on Jamaica in 1687-88, or Browne (1756) recorded a macaw matching Robinson's
(in Gosse 1847: 260) description of the bird that Rothschild (1905) would designate A. gossei.

Bond (1978) agreed with Lack (1976: 252-254) in rejecting the evidence that an
indigenous Ara (‘A. gossei') inhabited Jamaica. Wetherbee (1985) suggested that A. gossei
was not valid, but was probably a 'tapire' artefact; i.e., a specimen altered in its coloration
by Amerindians.

In addition, non-native red-plumaged macaws were reported from Jamaica by several
writers, including Browne (1756: 472), who wrote that the 'red Mackaw of Edwards ... is
not a native of Jamaica, but they are frequently brought there from neighbouring parts of
the Main, where they are pretty common.' Albin (1738a: 16) also reported a red-plumaged
macaw ('The Maccaw from Jamaica'), considering it the male of a species in which the
female was blue and yellow. A watercolour made in Jamaica by John Lindsay in 1765
depicts a probable Scarlet Macaw A. macao (Turvey 2010). Olson & Maiz Lopez (2008) noted
that the stylised macaw illustrated in Fisher & Warr (2003: 156) from paintings by a Lt.
L. J. Robins entitled The natural history of Jamaica (1765) was similar to Cuban Macaw and
suggested that species had been brought to Jamaica from Cuba.

Hispaniola. — Many accounts of the island's parrots have included a macaw, but
its status is confused. As evidence of a macaw there, writers have referred to reports by
Las Casas (1876: 298-299) and Oviedo y Valdes (1851: 443), as cited by de Armas (1888).
Macaws were said to have been common formerly in Hispaniola, but rare by 1760 (Clark
1905b). Buffon (1793: 160) reported a macaw, based on the statement of resident naturalist
Chevalier Lefebvre Deshayes [ob. 1786] (in lift, to Buffon 1847: 507), who stated that macaws
were once very common in Hispaniola, but had become rarer and confined to tops of
mountains. Olson (2005) dismissed Deshayes' statement noting it had no connection to
Hispaniola. Rothschild (1905) suggested that the Ara on Hispaniola represented a species
other than those already known from Cuba and Jamaica. Williams & Steadman (2001) listed
it as 'Ara tricolor ? or Ara unknown sp.', noting that among the three psittacids reported by
Las Casas (1876: 298) on Hispaniola at the end of the 1400s was a macaw that differed from
those on other islands in having a white forehead, not red like A. tricolor. Most recently,
Olson (2005) examined 16th-century descriptions by Las Casas (1876) and Oviedo y Valdes
(1851) and convincingly concluded that there was 'no credible evidence for the existence of
a macaw on Hispaniola in historic times.' There is a record of a transatlantic shipment of

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'long-tailed parrots [= macaws]' to Spain in 1494, when 60 were brought from Hispaniola to
Cadiz for trading (George 1980: 80).

Based on the de Armas (1888) report, Wetherbee (1985) developed an unfounded
hypothesis that Cuban and Hispaniolan macaws had been confused and extant specimens
represented both species. Further, he applied the name A. tricolor to the Hispaniolan macaw
and renamed the Cuban bird A. cubensis, but Walters (1995) rejected Wetherbee's revision
and returned A. tricolor to the Cuban population, suggesting that the Hispaniolan bird
probably formed a superspecies with the similar Jamaican form, A. gossei.

Puerto Rico. — Olson & Maiz Lopez (2008) reported several bones of one St. Croix
Macaw A. autochthones from an archaeological site in south-central Puerto Rico.

St. Croix, US Virgin Islands. — A tibiotarsus of an adult-sized immature macaw from a
prehistoric archaeological site on St. Croix is the basis for St. Croix Macaw (Wetmore 1937,
Olson 1978). Olson & Maiz Lopez (2008) reported several skeletal elements from Puerto
Rico. Olson (1978), Wing (1989) and Olson & Maiz Lopez (2008), however, cautioned that A.
autochthones was not necessarily indigenous to St. Croix and Puerto Rico because prehistoric
West Indian people were known to trade live psittacids. On the other hand, Williams &
Steadman (2001) argued that St. Croix could have sustained an indigenous macaw.

Montserrat. — An undescribed species, Montserrat Macaw Ara sp., was recently
suggested to have existed on the basis of subfossil remains (University of Florida, Gainesville;
UF; catalogue no. 4416) of an Ara recovered by D. R. Watters at an archaeological site at
Trant's (c,16°45'N, 62°09'W), Saint Georges Parish, (Williams & Steadman 2001). However,
it possibly represents 'A. guadeloupensis' (see above). As skeletons, most large parrots and
macaws are sexually dimorphic in size and exhibit great individual variation (see Hume
2007) and Olson & Lopez have suggested that the single coracoid is not diagnostic at species
level and that more material is needed before any attribution can be made.

Guadeloupe. — Diego Alvarez Chanca (c.1450-1515; in de Ybarra 1907: 428) joined
Columbus on his second voyage (1493-94) and reported taking 'two parrots ['Guacamayos'
or macaws], very large and quite different from the parrots we had before seen' from
the houses of Carib inhabitants of Guadeloupe. Two macaws have been named for the
island: Guadeloupe Macaw A. guadeloupensis Clark, 1905, and Guadeloupe Violet Macaw
Anodorhynchus purpurascens Rothschild, 1905.

Guadeloupe Macaw, a predominately red bird, is based on the account of du Tertre
(1654: 294), who also produced a rudimentary illustration (1667: between pp. 246-247; copied
by Labat 1742: between pp. 216-217). Johann Huttich (c. 1480-1544; 1534: 31) recorded a red
macaw 'present in such numbers as grasshoppers are with us...'. Clark (1905b) considered
A. guadeloupensis as occurring on Guadeloupe, Dominica and Martinique, but later he
(1908, 1934) distinguished the macaw of Dominica (A. atwoodi ) from those on Guadeloupe
and Martinique. Williams & Steadman (2001) found no evidence for Clark's (1905a: 269)
suggestion that A. guadeloupensis also occurred on Dominica and Martinique, but concluded
that it would seem more likely that the Lesser Antillean macaws were endemic to each
island or set of nearby islands based on what is attributed to Jean-Baptiste Labat (1663-1738;
in Clark 1905a: 269).

Hume & Walters (2012) noted the remarkable similarity between the descriptions
of Guadeloupe Macaw by du Tertre (1654: 294) and Labat (1742: 212), and that this may
support the presence of a macaw from the island. That similarity, however, may be the
result of Labat copying du Tertre's work; several illustrations in Labat are re-renderings
of du Tertre's originals. An illustration by D'Aubenton (in Buffon 1765-83: PL 12) closely
resembles the descriptions of du Tertre and Labat, which Hume & Walters (2012) considered
possible evidence that at least one Guadeloupe Macaw reached Europe.

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Because Carib Amerindians were able to inform Columbus the direction of the
mainland, Greenway (1967: 318) suggested that Guadeloupe Macaw could have been an
exotic species, imported through trade with Mesoamerica or Mexico. Still, Williams &
Steadman (2001) argued it was just as likely that macaws were indigenous.

Rothschild (1905) based his description of Guadeloupe Violet Macaw on Martin
Fernandez de Navarrete y Ximendez de Tejada's (1765-1844; cited as 1838: ii: 425 in
Rothschild 1907a: 55, but not found by us) account of Te gros Perroquet de la Guadaloupe',
a very large macaw of an intense purple colour', 'called Onecouli, by the Caraibes, according
to Fernand Columbus' (Rothschild 1907b: 202). Navarrete's narrative was based on Las
Casas' rediscovered abstract of the log Columbus made of his second voyage, not on first¬
hand observations. Martyr (in MacNutt 1912: 72) provided more detailed and convincing
evidence of a purple macaw on Guadeloupe, recounting that Columbus took 'seven parrots
larger than pheasants, and totally unlike any other parrots in colour. Their entire breast
and back are covered with purple plumes, and from their shoulders fall long feathers of the
same colour. ... The other feathers are of various colours, —green, bluish, purple, or yellow.'
Greenway (1967: 320) and Snyder et al. (1987: 40) suggested that Anodorhynchus purpurascens
was based on either a poor description of Guadeloupe Parrot Amazona violacea (now
extinct) or Lear's Macaw Anodorhynchus leari of Brazil, which must have been imported to
Guadeloupe. At best, the evidence for a Guadeloupe Anodorhynchus is weak.

Marie Gal ante.— Williams & Steadman (2001) reported a single ulna of a species
referred to as 'Marie Galante (Guadeloupe?) Macaw Ara cf. guadeloupensis' ; i.e., probably
A. guadeloupensis, but an extinct species for which no specimen exists. Tire specimen was
discovered at an archaeological site at Folle Anse (c.l5°57'N, 61°20'W), Grande Anse, Marie
Galante, and is deposited in the UF archeological collections (Folle Anse 68, Box 68-10).
However, Olson & Maiz Lopez (2008) examined the specimen reported by Williams &
Steadman, and considered it best referred to Imperial Parrot Amazona imperialis.

Dominica.— Dominica Macaw Ara atwoodi Clark, 1908, is a poorly documented species,
based mainly on Thomas Atwood's (oh. 1793; 1791: 29) 18th-century report of a macaw
larger than native Dominican parrots ( Amazona arausiaca, A. imperialis) and was 'in great
plenty, as are also parrots in this island.'

Martinique.— Considerable confusion exists concerning the macaw(s) of Martinique.
Williams & Steadman (2001) listed two species, 'Ara martinica' Rothschild, 1905, and
'Anodorhynchus martinicus' Rothschild, 1905, both called Martinique Macaw. In his three
publications referring to extinct macaws of the West Indies, however, Rothschild does
not mention two species from Martinique. Rothschild (1905) first assigned Anadorhynchus
[Anodorhynchus] martinicus to a blue-and-orange macaw recorded by Pere Jacques Bouton
(1592-1658; 1640: 71). Rothschild used the name Anadorhynchus martinicus, nom. nov. in his
paper (1907b: 202) but, in his Extinct birds (1907a: 53), he referred to it as Ara martinicus,
noting Anadorhynchus martinicus Rothsch. 1905 as a synonym. We suggest only one, if any,
species existed on Martinique, and propose that Ara martinicus Rothschild, 1907, be assigned
to it. Certainly, whether one or two species, it is poorly documented, being based solely
on Bouton's (1640) scanty account. Salvadori (1906) considered it to be Blue-and-yellow
Macaw. Greenway (1967: 319) suggested Bouton described a captive bird and, with Snyder
et al. (1987: 39M0), also thought that the species concerned was Blue-and-yellow Macaw,
which could have been traded to Martinique. Snyder et al. (1987), however, noted that a
distinctive macaw once occurred on Martinique. Regardless of its name, we agree with
Williams & Steadman (2001), who suggested that both supposed species of Anodorhynchus
in the Lesser Antilles require corroboration. As noted by J. P. Hume (in litt. 2013), that an
Anodorhynchus macaw once occurred on the West Indies is, for now, probably best regarded

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as a figment of Rothschild's imagination, given that no modern-day Anodorhynchus occurs
anywhere near the West Indies, and all of the available evidence suggests that this genus
was rare even in historical times making it unlikely that such birds would have arrived
in the Antilles via trade. In contrast, several species of Ara macaws occur on the adjacent
mainland of South and Middle America.

In 1936, the Cuban scientist Mario Sanchez y Roig (1890-1962) thought he had found
a specimen of Ara martinicus. He claimed that it was collected in September 1845 and
mounted in 1846 (in litt. to L. C. Sanford, 21 February 1936). On examining the specimen,
however, J. T. Zimmer determined it to be a composite hoax, using the head, body and
wings of Burrowing Parakeet Cyanoliseus patagonus byroni (now bloxami ) of Chile, and tail
of a dove, apparently an Old World Streptopelia. Zimmer (in litt. American Museum of
Natural History, 27 March 1936) noted 'It has very little resemblance to the Macaws, even
in its made up condition.'

Natural history of Cuban Macaw, with comments on other

West Indian macaws

Earliest records.— Columbus and others, including Jose de Acosta (1540-1600; 1590),
Bartolome de Las Casas (c.1484-1566; 1552-61), Diego Alvarez Chanca (1494; in de Ybarra
1907), and Gonzalo Fernandez de Oviedo y Valdes (1478-1557; 1950: 167) mentioned
macaws in their reports of the exploration of Cuba and other of the Antilles. Juan Ignacio
de Armas (1888: 114-115) summarised these early records. Cuban Macaw's beauty was well
appreciated and illustrated in early treatments based on specimens, including Levaillant
(1801 Vol. 1: PI. 5), D'Aubenton (in Buffon 1765-83: PI. 641: 'Le petit Ara'), Bechstein (1811:
PI. 1) and Brehm (1842: PI. 3). The court painter Leopold Brunner produced a painting of A.
tricolor in the late 18th century (H.-M. Berg pers. comm.).

Relationships. — Speculation on relationships among West Indian macaw species is
presently impossible because no reliable, detailed descriptions or specimens exist other than
for A. tricolor. Williams & Steadman (2001) noted that overall the plumage of Cuban Macaw
suggests that its nearest mainland relative was Scarlet Macaw, because the distribution of
red and blue is similar, as is the presence of a white facial patch that is featherless except for
small crescentic lines of tiny red feathers. A. tricolor is distinct in lacking a yellow shoulder
patch, its all-black bill and substantially smaller size. Greenway (1967: 315) affirmed that
A. tricolor was clearly a representative of A. macao, and believed other forms of what he
considered a superspecies occurred on Jamaica, Hispaniola and Guadeloupe. A. tricolor
and A. macao share a species of feather mite (see Parasites), which further supports their
relationship.

Range and status of Cuban Macaw. — Early writers described the range of A. tricolor
as not only Cuba and Isla de Pinos, but also Haiti and Jamaica (Cory 1892, Gundlach 1893,
Todd 1916), but most recent authors have limited its range to the first two islands (Bond
1978, Olson 2005, AOU 2012).

Clark (1905b) stated that young A. tricolor was largely green, although he presented no
source for this. If the juvenile of A. tricolor was predominantly green, that could account for
early reports of A. militaris on Cuba and Jamaica (Clark 1905b, Williams & Steadman 2001).
On the other hand. Military Macaws were imported to the islands and could have occurred
in a feral state, thereby adding to the confusion (see Trade).

Cuba. — Fossil evidence exists for Ciego Montero, Cienfuegos province (22°20'0"N,
80°24'0"W; Wetmore 1928; Pleistocene); Cueva de Paredones, Caimito, La Habana province
(22°84'0"N, 82°63'0''W; Arredondo 1984; Quaternary); and Casimba en Los Buentes, Mai
Paez, Sagua La Grande, Villa Clara province (22°48'0"N, 80°04'0"W; Olson & Suarez 2008;

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Quaternary) (Fig. 1). Arredondo (1984) said that Cuban Macaw was abundant at the time
of the European arrival, although Barbour (1923: 80) suggested it probably never was
widespread. Moreno (1992) considered it restricted to central and western Cuba, as well as
Isla de Pinos.

The Italian Giovanni Gemelli Careri (1651-1725; Giro del mundo 1699-1700; in de la
Torre 1857) mentioned killing large numbers of parrots and two macaws near La Habana on
20 January 1698. An anonymous resident ('By a physician' 1844) of Cuba listed the macaw
in the 1840s, but by the mid-19th century d'Orbigny (1839) wrote that it was becoming rare.
Most accounts of the macaw's range and numbers at this time are based on Gundlach's
(1810-96) reports from Cienaga de Zapata, a vast swamp comprising about one-third of
the province of Matanzas, on whose southern boundary it is located. Gundlach (1893: 151)
noted that the macaw was still fairly common at the swamp's northern edge (e.g., Hanabana,
where the rio Hanabana flows into the swamp) in 1850, but thereafter it retreated into the
interior. He (1874; 1876: 126) reported it occurring from Hanabana (Hacienda Hanabana
= 'Hato Zarabanda'; 22°24'0"N, 80°58'01"W; 12 m) to the Ensenada (Bahia) de Cochinos
(Bay of Pigs; 22°10'48"N, 81°09'59"W; c.32 km south-west of Hanabana). Gundlach passed
nine months in the Cienaga, from September 1849 to mid 1850, mostly at its north-eastern
edge (Gonzalez Lopez 1990). It was probably during his 1849-50 visit that Gundlach
collected several macaws from one of the last bands, which he said came regularly to
feed at Hanabana (Ramsden 1918). In early 1850, Gundlach traveled to Hato Cabeza de
Toro (22°06'0"N, 80°48'0"W; 10 m; c.38 km south-east of Hacienda Hanabana, and c.39
km east-southeast of Bahia de Cochinos), but left no record of macaws in that part of the
Cienaga. Gundlach made another trip to Cienaga de Zapata in June 1856, his first stop on
an extended journey through Cuba. By 1856, Gundlach (1856) considered the macaw rare in
Cuba. Barely a decade later, Gundlach (1865-66, 1873, 1874, 1876: 126) noted it was confined
to only a few places, with the population consisting of a few pairs. In later years he did not
find it in Cienaga de Zapata, although Gundlach told Cory (1886, 1889: 177-178) that he
believed A. tricolor still survived in Zapata about 1885.

The only report of macaws from western Cuba is that of Barbour (1923: 80), who
was told of its former presence at Guane (western Pinar del Rio province; 22°12'07"N,
84°05'16"W; c.300 km west of Bahia de Cochinos; Fig. 1); he remarked that no macaws were
seen in western Pinar del Rio after the great hurricane of 1844.

In 1867, Antonio Perpiha (1889), a Spanish writer, geographer and priest, traveled
widely through central Cuba, some of the cayos, and as far south-east as Pico Turquino,

Figure 1. Cuba and Isla de Pinos (Isla de la Juventud) with major cities and sites mentioned in text, and
known localities of Cuban Macaw Ara tricolor from observations (dots) and fossil specimens ('x').

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Sierra Maestra (Fig. 1). Perpina described passing through extensive virgin forests in which
he found many animals, and gave a detailed and acceptable description of Cuban Macaw,
which he called the 'king of the parrots' because of its brilliant colour and size. Perpina
(1889: 32) recorded that his entourage shot a macaw, six parrots, four parakeets and 23
other birds during a day of hunting in Valle de San Antonio (c.21°32'0"N, 77°28'0"W; 24
m, near Sierra de Cubitas and c.31 km east-northeast of Camagriey; Fig. 1). Later in his
expedition, Perpina (1889: 242) reported another macaw and many parrots were killed by
his party near Jobabo (20°54'32"N, 77°17'06"W; 82 km south-east of Camagiiey), between
Guaimaro (21°03'0"N, 77°21'0''W) and Las Lomas de Rompe (Sierras de Rompe; 21°02'42"N,
77°10'55"W), west-northwest of Las Tunas (Fig. 1). Further, Perpina (1889: 247) wrote of the
great red macaw ('lorirojo') observed at Las Lomas de Rompe, where he also noted Ivory¬
billed Woodpecker Campephilus principalis bairdii, Cuban Parakeets Aratinga eitops and other
birds. Although Perpina' s observations appear credible, it is surprising that Gundlach did
not encounter macaws in central Cuba, including around Camagriey in May 1859; however,
Gundlach was told of macaws in central Cuba (Gundlach 1893: 151). Although Gundlach
made extensive explorations of the easternmost region of Cuba, he failed to find macaws
and said that no one knew of it there (Gundlach 1893: 151).

Isla de Pinos.— Gundlach explored Isla de Pinos (= Isla de la Juventud) twice, in 1854
(December) and 1855, including six weeks at Santa Fe (21°44'31"N, 82°45'18"W); he also
made a five-week trip early in 1892 (Poey 1866, Gundlach 1854: 426, Dathe & Gonzalez
Lopez 2002; in lift, to F. Ramsden, La Flabana, 26 April 1892; JWW pers. archive). Despite
these extended periods of travel, Gundlach was unable to adequately explore the extensive
Cienaga de Lanier (21°34'0''N, 82°52'0"W; 10 m) because of the distance from his base at
Santa Fe (= La Fe; c.25 km north-east of Lanier) and so passed just one day at the swamp's
northern edge. Although Gundlach (1893: 151) listed the macaw from Isla de Pinos, he
collected none there; he was told that some macaws still survived on the edge of the
Cienaga de Lanier and reported that the macaw and Ivory-billed Woodpecker occurred in
the southern Cienaga (Dathe & Gonzalez Lopez 2002). Walter Reaves Zappey (1878-1914)
traveled extensively through Isla de Pinos, collecting for Rothschild's Tring Museum in
1902, and for E. A. & Outram Bangs in 1904, but did not find maqaws. Zappey was told
that the last pair of macaws known in the island was killed about 1864 at San Francisco
de la Vega (21°37'0"N, 82°46'0"W; Fig. 1), 13 km north of the Cienaga de Lanier; none was
reported thereafter (Bangs & Zappey 1905, Todd 1916). The pair killed at San Francisco de
la Vega apparently was not preserved (Olson & Suarez 2008). The Isla de Pinos macaws
were probably subject to intense harvesting as the Cuban population dwindled. As parrots
became scarcer on Cuba, harvesting became much more intense on Isla de Pinos to supply
demand (Smith 1944); we suspect its macaws also became more vigorously sought for the
same reason.

Because Bangs & Zappey (1905) referred to the locality of the 1864 record merely as
'La Vega,' rather than its complete name of San Francisco de la Vega, many authors (e.g.,
Greenway 1967, Day 1981, Luther 1995, Fuller 2001, Olson & Suarez 2008, Hume & Walters
2012) have wrongly assigned the location to La Vega, Cienfuegos province (22°05'0"N,
80°21'0"W), in Cuba, rather than to the correct Isla de Pinos location. La Vega, Cuba, is c.300
km east-northeast of San Francisco de la Vega, Isla de Pinos. Equally, many authors since
Rothschild (1907a), especially in recent decades, have referred to this record as only having
involved a single individual.

Habitat. — Cuban Macaw occupied open terrain with scattered trees, especially palm
savanna, characteristic of Cienaga de Zapata. Many years after Gundlach, Barbour (1943: 73,
1945: 152) traveled the rio Hanabana to where that river flowed into the swamp, describing

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the morass as bordered by a wide area of open country with scattered clumps of palms
and hardwood cayos or hammocks. The border zone sloped gently toward the swamp and
its extent varied as the rains caused the water levels to rise or fall throughout the entire
Cienaga.

Perpina (1889) described limitless tracts of unbroken forests from north of Camagiiey
to Las Tunas. The natural vegetation of central Cuba was seasonal evergreen, semi-
deciduous (dominated by Swietenia mahagoni) and gallery forests, in stark contrast to the
almost complete conversion of that region to crops and pastures with only remnant forest
and palm savanna today (Borhidi 1991). The habitat where Perpina found macaws around
Lomas de Rompe formerly consisted of extensive rainforest-like gallery forests, but has been
impoverished by human influence, and today only a few degraded stands of forest remain;
natural vegetation has been replaced by treeless marshy meadows and moist savannas
(Borhidi 1991). Also, the area around Guane, Pinar del Rio, is mostly converted habitat,
but the flatlands and gently rolling hill country of the region originally were characterised
by mixed palm-pine woodlands with a loose canopy layer, in which Colpothrinax wrightii,
Acoeloraphe wrightii, Pinus tropicalis and P. caribaea were dominant (Borhidi 1991).

Cienaga de Lanier has not been substantially altered since the mid-19th century and
is today characterised by mangroves at its southern fringe, and a variety of seasonally
inundated habitats including semi-deciduous forest, marshy grasslands, hardwood and
palm hammocks, and palm savanna in the interior.

Buffon (1793: 161), probably speaking of more than one species of red macaw, said
these birds lived in woods and wetlands with palm trees, and fed mainly on fruit of
palms, of which there were immense forests in the flooded savannas. M. de le Borde (in
Buffon 1793: 160) reported that West Indian macaws 'retire to the least frequented places,
and are no more observed to approach the plantations.' Browne (1756: 472) stated that
the 'blue Mackaw of Edwards' generally kept to the most unfrequented inland parts of
Jamaica. Richard Hill (in Gosse 1847: 261) reported Jamaican macaws (Ara sp.) were 'found
exclusively in the central mountains westward of the island, and are observed on the skirt
of the partially cleared country, at an elevation of 2500 or 3000 feet [750-900 m] above the
sea.' March (1863: 283) noted that all records of macaws in Jamaica were from mountainous
and wooded areas, in contrast to the Cuban Macaw, which was mostly found in lowland
savanna, including coastal regions, and low-elevation interior forests.

Habits.— Pew data are available on behaviour of West Indian macaws, but du Tertre
(1654: 294) said 'Their voice is loud and piercing, and they always cry when flying. If one
imitates their cry, they stop short. They have a grave and dignified demeanor'. Labat (1742:
212) likewise noted that the voice of West Indian macaws in general was strong. Atwood
(1791: 29) characterised Dominican Macaw vocalisations as a 'disagreeable, harsh noise,'
and 'loud chattering noise, which at a distance resembles human voices.' Gosse (1847:
263) reported that Jamaican macaws (Ara sp.) flew extremely high, in pairs, giving harsh
screams, before finally alighting on the 'loftiest of the forest trees, in their chosen resting
places.' In Dominica, Atwood (1791: 29) said macaws gathered atop the highest trees, where
they 'feed on the berries in great numbers together.' De Rochefort (1658: 154) said Lesser
Antillean macaws commonly flew in flocks. Gundlach (1874, 1876) reported that Cuban
Macaw lived in pairs or in families, and noted (1893) it had a loud vocalisation like the
macaws of Central America.

Di et.— Gundlach (1874, 1876, 1893) characterised the diet of Cuban Macaw as
including fruits, palm fruit, seeds of Chinaberry tree Melia azedarach, tender shoots and
buds. Chinaberry tree is native to Asia, Australasia, and some Pacific islands, but has been
naturalised widely in warmer parts of the world, including the West Indies. The fruit is a

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slightly fleshy marble-sized (c.15 mm in diameter), nearly round drupe. The oblong seed is
3.5 x 1.6 mm and surrounded by pulp, which was probably the part consumed by macaws,
as it is by Cuban Parrots Amazona leucocephala and Cuban Parakeets (pers. obs.). The fruit
is mildly toxic to humans. Cuba has a diverse palm flora (Leiva Sanchez 1999) and seeds of
several species that occur in Cienaga de Zapata were probably important in the macaw's
diet (Olson & Suarez 2008). Dominant species of palm there include Roystonea regia and
Salmi maritima. Local palms have large fruit upon which many birds, including parrots, feed.
Olson & Suarez (2008) attributed certain features of Cuban Macaw's skull to its probable
diet of very hard seeds, especially those of palms, as is typical of larger mainland macaws.

In Jamaica, Browne (1756: 250) noted that the seeds of a Sloanea ('the large oval-leafed
Sloanea, or brake-axe tree'; probably S. jamaicensis) were 'much coveted by the mackaw and
parrots,' further stating that these were the only birds that could 'break thro' those thick and
lignous seed-vessels.' Further, Browne (1756: 343-344) listed two Jamaican palms— 'The
Mackaw Tree' and 'The Great Mackaw Tree' — but without reference to the bird. He noted
that their fruits were large and rich in palm-oil. Liman (1814: 468) said the Macaw-Tree was
so called 'from a large bird that feeds upon the fruit of this tree, which is of the palm kind.
There are two sorts of them...': 'great macaw-tree' and 'small macaw tree.' There may be
confusion as to the true origin of the names of these palms, because the vulgar names may
relate to the Portuguese name of macauba for a kind of palm tree.

Oviedo y Valdes (1535: Libro Septimo, Capitulo ii: lxxii) reported West Indian macaws
fed on manchineel Hippomane mancinella, which is common in the West Indies but is
a powerful caustic poison to man and other animals. Du Tertre (1654: 296) said West
Indian macaws fed 'on seeds and several fruits of trees, but mainly on the apples of the
manchineel.' Later, du Tertre (1667: 249) elaborated, saying macaws only ate manchineel
in times of necessity, and cautioned that meat of macaws that had eaten manchineel was
unhealthy and even poisonous to man.

Breeding. — Unfortunately, Gundlach did not observe the breeding behaviour of A.
tricolor as he did not visit Hanabana at the appropriate season (Gundlach 1865-66, 1873,
1874, 1876). He described the nest as a hollow in a palm based on sites shown to him by local
residents (Gundlach 1876). The egg has not been preserved or described.

Atwood (1791: 29) noted that Dominican Macaws 'breed on the tops of the highest
trees.' Du Tertre (1667: 249) stated 'The male and the female [West Indian macaw] are
inseparable companions, and it is rare that one is seen singly', further noting, 'When they
wish to breed (which they do once or twice a year) they make a hole with their beaks in the
trunk of a large tree, and construct a nest with feathers from their own bodies. They lay two
eggs, the size of those of a pigeon, marked like those of partridge.'

Parasites. — Du Tertre (1667: 249-250) said West Indian macaw nestlings were infested
with two worms in the nostrils, with another in a small tumour on top its head, and noted
that the maggots died once the chicks were feathered. These worms were probably warble
fly Philornis pici larvae, which commonly parasitise psittacine nestlings in the West Indies
(Snyder et al. 1987; pers. obs.). The maggots pupate within their host, and so may appear
as though the larvae have died. Mey (2005) reported a new species of chewing louse
Psittacobrosus bechsteini, now probably extinct, from a Cuban Macaw collected by Gundlach
c.1849. Cuervo Pineda & Perez Ortiz (2009) found a feather mite Genoprotolichus eurycnemis
on a Cuban Macaw collected by Gundlach in 1849. This species is also known from Scarlet
Macaw. Cuervo Pineda & Perez Ortiz (2010) also found a new species of mite, Distigmesikya
extincta, on the skin of a Cuban Macaw.

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Population decline and extinction of West Indian macaws

At the time of Columbus, parrots and macaws were abundant in the islands (Huttich
1534: 31, Sloane 1725: 297, Atwood 1791: 29, Deshayes in Buffon 1793: 160, Edwards 1806:
19, Martyr in MacNutt 1912: 72), but by the 18th century macaws were rare in the Lesser
Antilles (and presumably elsewhere) (Clark 1905a). M. de le Borde (in Brisson 1760: 183)
stated 'In all these [West Indies] the Macaws have become very rare'. Seventy years earlier,
John Taylor (fl. 1687; 2008: 142-143) noted that Jamaican macaws were 'held as a great
rarity, there being but a few of them on this island and those hard to be come at, because
they are noewhere to be found but in the woody mountaines.' Further observers of macaws
in Jamaica also reported declining populations. Browne (1756: 472) said the 'blue mackaw of
Edwards' was very rare, Stewart (1822: 79) noted that 'The maccaw is become very scarce',
and Gosse (1847: 260) reported they were very rare during his residence (December 1844-
July 1846). March (1863) stated that he had not heard of any reports on Jamaica since 1849,
the last dependable report for the island.

Several dates have been proposed for the extinction of Cuban Macaw, including c.1850
(Bond 1940, Adams et al. 2003), c.1864 (Bond 1959, Balat & Gonzalez 1982, Olson 2005),
c.1884 (Vincent 1966-71, Moreno 1992), c.1885 (Greenway 1967, Prestwich 1970), between
1864 and 1885 (Lantermann 1984), and end of the 19th century (Garrido & Garcia Montana
1975, Moreno 1992). The last reliable reports, however, were Gundlach's from Cienaga de
Zapata in the early 1850s, Perpena's in central and south-west Cuba in 1867, and possibly
Zappey's second-hand record from Isla de Pinos in 1864.

Causes of extinction

Hunting. — Hunting for psittacids was a part of subsistence and traditional cultural
activities of Amerindians before the arrival of Europeans. In Mexico, Hernando Cortes
(Diaz del Castillo 1956) saw parrots for sale as food in the Aztec capital Tenochtitlan in
1520. Antillean Amerindians included parrots and macaws in their diets, and du Tertre
(1667: 249) said that macaws were eaten regularly on some islands. The presence of macaw
remains in St. Croix, Puerto Rico and Montserrat kitchen middens supports this. On
Dominica, Columbus' men were shocked to find 'popingiayes' (macaws) along with geese,
ducks and human flesh in earthen vessels, presumably being food larders, in houses of
natives (Huttich 1534: 31, Arber 1885: 30).

West Indian macaws were easily killed, even using bow and arrows (Labat 1742: 81).
Martyr noted the ease at which macaws could be captured by hand or noose, and the
curious fact that they were not alarmed by gunshots (in Buffon 1793: 162, MacNutt 1912:
409). Du Tertre (1654: 294) said West Indian macaws were 'far from being alarmed by many
shots fired under a tree where they are perched, they gaze at their companions who fall
dead to the ground without being disturbed at all, so that one may fire five or six times into
the same tree without their appearing to be frightened.' de Rochefort (1658: 154) considered
Lesser Antillean macaws 'bold and resolute, for they are not alarmed by the report of fire¬
arms, and if none are wounded at the first discharge, they await a second without moving
from the place where they are; but there are many who attribute this boldness to their
natural stupidity rather than to their courage.'

Du Tertre (1667: 248-249) described how Amerindians in the Antilles captured macaws
alive. The hunters waited until they descended to the ground to feed on fallen fruit,
at which point the men stealthily approached, surrounding the birds, then burst forth,
shouting and clapping, which startled the birds so that they did not fly, but fell on their
backs and defended themselves with bills and claws. The Indians shoved sticks at the birds.

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which they grasped with claws and beak, whereupon the birds were tied up with vines.
In Cuba captive parrots were used as decoys. A native equipped with a captive parrot and
a noose would climb a tree, where he hid in a straw 'hide.' When the trapper touched the
decoy parrot's head, it cried out and attracted other parrots. The noose was slipped over the
head of any inquisitive parrots, its neck wrung and let fall to the ground (Krieger 1929: 44).

Live macaws and parrots were kept captive until such time as they became meals for
Amerindians. Huttich (1534: 31), speaking of Ara guadeloupensis, noted 'they feed some of
them so that they are better to eat.' Martyr (in MacNutt 1912: 72) reported that macaws and
parrots were kept in captivity by Amerindians, 'and just as we keep magpies, thrushes, and
similar birds to fatten them, so do these islanders also keep birds to eat, though their forests
are full of parrots.'

Parrots, including macaws, were hunted as food by 15th and 16th-century explorers
(Cooper & Armitage 2013), though they were probably eaten by the Europeans far less than
by native peoples, and then only in desperation (Boehrer 2004, Cooper & Armitage 2013).
Still, Labat (1742: 215) noted that while in Guadeloupe he ate more parrots than partridges
when in Europe. Long (1774: 865, 951) noted that parrots were eaten by Amerindians
but, although he listed parrots and parakeets, he did not include macaws among birds
commonly eaten by Jamaican colonists. The reason for macaws not being eaten by
Europeans may relate to the quality of their flesh. Du Tertre (1654: 294) said of West Indian
macaws, 'The flesh of this bird is very tough, and considered by many unwholesome, and
even poisonous.' Atwood (1791: 29-30) described the flesh of the Dominican Macaw as
'...being very fat, it wastes in roasting, and eats dry and insipid; for which reason, they
are chiefly used to make soup of, which is accounted very nutritive.' Conversely, Taylor
(2008: 142-143) noted that the flesh of Jamaican macaw ( Ara sp.) was 'very good but they
are seldom eaten.' Sloane (1725: 297) said 'the Small Maccaw' was 'eaten as Pigeons',
and further noted that other native parrots were eaten, including the 'common parrot of
Jamaica' (Yellow-billed Parrot Amazona collaria) which he recorded as being 'bak'd in Pyes'
and tasting of pigeon. This is more or less seconded by Coke (1808: 391), who stated that
'There are two species of parrots in the woods, natives of the island, both green; but one
sort has a yellow bill. They are neither so beautiful, nor such good talkers, as the gray and
green of other countries; they are, however, esteemed as delicate food, and are served up at
some tables in preference to pigeons.' Browne (1756: 473) noted that parrots were 'generally
reckoned very delicate meat, and eat not unlike pigeons; ... and frequently served up at
gentlemen's tables in all the country parts of the island [Jamaica].' M. de le Borde (in Brisson
1760: 183) stated 'In all these [West Indies] ... the inhabitants destroy [macaws] for food.'
The price commanded by parrots as game is suggested by Taylor (2008: 142), who wrote
that 'you may bie of them dead for two royalls a duzen... at Jamaica.'

Several writers have pointed to hunting as contributing to Cuban Macaw's extinction
(Rothschild in Lonnberg 1928, Barbour 1943, Arredondo 1984). Greenway (1967) noted
that hunting was the only known reason for its extinction. Apparently, as reported for
other West Indian macaws, Cuban Macaws were easily taken; Barbour (1943) was told
that adults were stupid and slow to take flight when approached. Gemelli Careri (1699;
in Perez de la Riva & Berthe 1971) commented on the tasty flesh of Cuban Parrots, but
diplomatically remarked that the Cuban Macaw should be conserved for the beauty of its
feathers. Gundlach (1865-66, 1873, 1874, 1876) was more direct and noted that macaw flesh
was undesirable, being tough and having an especially unpleasant odour (Gundlach 1893).
Nevertheless, it was killed for food (Greenway 1967) and Todd (1916) noted it was even
valued for its meat.

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Trade. — Harvesting activities began even before the European arrival, with substantial
numbers of animals (including macaws) captured by Amerindians for pets or trade
among islands (Olson 1982, Wing 1989). Olson & Maiz Lopez (2008) suggested that some
populations of West Indian macaws may have been so precious in trade that every possible
nest was sought to obtain young birds, and that vigorous harvesting of such large and
comparatively uncommon species might conceivably reduce populations to extinction over
the nearly 2,000 years that some islands had been occupied. They further suggested that
Amerindians may have maintained the last Hispaniolan macaws in captivity, even after
their extinction in the wild but, with the collapse of their cultures soon after the European
arrival (Rouse 1992), the macaws became extinct.

The earliest explorers' accounts strongly suggest macaws were kept commonly as
pets and perhaps bred in captivity by Amerindians (Greenway 1967, Dunn & Kelley 1989).
Although adults were taken alive, nestlings were preferred as companion birds, but du
Tertre (1667: 249) noted that adults could be tamed even when caught by such traumatic
methods as described above. In northern Mexico and the American south-west, where they
do not occur naturally, macaws (mainly Ara macao) were an important item of commerce
and ritual among Amerindians, and large captive-breeding facilities were used to sustain
ritual sacrifices (Hargrave 1970, Minnis et al. 1993, Creel & McKusick 1994). Bernal Diaz
del Castillo (1492-1584; 1956: 212) described Hernando Cortes' entrance into Mexico City,
where he found a royal zoo containing birds of all sorts, including 'parrots of many different
colours, and ... so many of them that I forget their names.' Cortes (in Boehrer 2004: 56)
described the Aztec zoo as superior to any in Spain. Smith (1937) noted that Columbus was
'astonished at the swarms of tame birds at liberty in the villages' of Cuban Amerindians.
Chanca (in de Ybarra 1907) reported Columbus' men took two macaws from the houses
of Caribs on Guadeloupe in November 1493. Being the largest and most colourful of West
Indian parrots, macaws were probably of great prestige and value, and would have been
traded afar.

In addition to native species, Amerindians may have imported macaws from the
mainland. The husbandry and trade in live psittacines by West Indian people has raised
concern among archaeologists and biogeographers who analyse material excavated from
cultural sites (Wing 1989, Williams & Steadman 2001). Prehistoric peoples transported
animals for trade and sustenance on their passages among islands, thereby confounding
the determination of the former number and original distribution of macaws and other
psittacids (Oviedo 1959, Wilson 1990, Williams & Steadman 2001). Thevet (1971) reported
that native peoples carried macaws with them during war against enemies, '...for to
eate them, and other things.' Olson (1982) presented evidence of bird transportation and
extinctions caused by early man, noting that a lively trade in macaws existed between
Indians in tropical Mexico and those in south-west North America, and suggested that trade
in macaws probably also occurred in the West Indies.

Besides using macaws and other psittacines as barter and as pets, they were vigorously
sought by Amerindians for their feathers, to decorate ceremonial dress and otherwise
enhance the body (Labat 1742: 85, Navarrete 1828: 277, 280, Krieger 1929). An account of
Francisco de Orellana's 1542 exploration of the Amazon describes local Indians keeping
macaws as a source of ceremonial feathers (Heaton 1934: 415). The value native people placed
on parrot feathers was recorded by Amerigo Vespucci who noted that those peoples' riches
consisted in part 'of variegated birds' feathers,' and one tribe presented him with 'feathers
of very great value, ... and ... numberless parrots of different colors' (Waldseemiiller 1907:
98, 109). Du Tertre (1654: 294) noted that 'The natives [of Guadeloupe] hold the feathers of
the tail [of macaws] in great esteem; they stick them in their hair, and pass them through the

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lobe of the ear and the septum of the nose to serve as mustaches, and consider themselves
then much more genteel and worthy of admiration of Europeans.'

Macaws and parrots were among the valued items traded by Amerindians for the
trinkets brought to the Antilles by the earliest European explorers and, in turn, were
displayed among the treasures as those explorers returned to Europe. Columbus traded
red caps, bells and glass beads to natives, who in turn gave him such things as they valued
most, including gold and parrots (Navarrete 1828: 42, 45, 186, Dunn & Kelley 1989: 65, 71,
223, 259, 271). In fact, Columbus was eager to obtain macaws and requested these from
Amerindians who readily supplied them (Navarrete 1828: 186). In Columbus' triumphant
march to Barcelona, he paraded his most cherished and representative treasures, leading
his entourage on horses and mules, with captive Indians adorned with feathers and various
bright ornamental apparel, their ears decorated with gold pendants, their arms with
bracelets and wearing bead necklaces. Red-and-green parrots perched on their shoulders.
All of these treasures Columbus presented to the king and queen (Collis 1977: 108, Davidson
1997: 285). Oviedo y Valdes (1535: Libro Segundo, Capitulo vii: ix) recorded that Columbus
arrived in Barcelona with riches including gold, native Indians and many macaws. Martyr
(in MacNutt 1912: 65) related that Columbus returned from his first voyage with 'some forty
parrots, some green, others yellow, and some having vermilion collars like the parrakeets of
India...; and all of them have the most brilliant plumage. Their wings are green or yellow,
but mixed with bluish or purple feathers, presenting a variety which enchants the eye.'
Morison (1970: 435) recounted that Antonio de Torres, returning from Hispaniola to Cadiz,
in March 1494, brought 'sixty parrots of different colors, eight of them being big as falcons
and the fairest species of fowl that fly in the air.'

Throughout the Age of Exploration of the New World, travelers brought specimens
of exotic animals to Europe: live, to be exhibited in royal collections, dead, for museum
cabinets (George 1980). As travel to the New World increased with accelerating exploration
and colonisation, so did trade. Much as did the Amerindians, Europeans valued macaws
and carried them among islands and to the mainland from the beginning of commerce in
the New World. The Italian naturalist, Ulisse Aldrovandi (1522-1605) saw his first macaw
at Mantua, Italy in 1572, and remarked that they were then of great admiration and were
highly esteemed, and that the nobility gave them to one another as rare and valuable
presents (in Buffon 1793: 157). Although parrots were considered novelties during this
era, before long the entrepreneurial nature of adventurers and travelers recognised the
trade value of parrots in the Old World. In fact, so many were brought back to Europe
that a mere 34 years after Columbus' first voyage, Oviedo y Valdes (1950: 167) wrote 'there
are many parrots, and of so many kinds and diversity, that it would take a long time to
describe them, but because so many are brout to Spain, I will not waste time talking of
them.' Similarly, Labat (1742: 211) stated that West Indian psittacines were too well known
for him to waste time portraying them. Long (1774: 896), writing for a European audience,
said of Jamaica's five species of psittacines, including 'Blue Mackaw', 'These are all native,
and too generally known to need any description.' Nicolaus Joseph Freiherr von Jacquin's
(1727-1817) expeditions to the Caribbean islands and elsewhere in the New World, 1754-59,
resulted in large collections of natural history objects, but Jacquin (1784) did not include Ara
tricolor among those species reported from his exploration of Cuba (1759) and, in fact, he was
given instructions by Emperor Francis I not to collect parrots (C. Riedl-Dorn pers. comm.),
again, probably because these birds were already so well known in Europe.

As Europeans rapidly became familiar with New World parrots, their desire to own
these exotic status symbols continued to increase. The number of species described from
captive birds attests to the multitude of psittacine species and numbers that were kept as

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pets in Europe in the 17th and 18th centuries, particularly among the upper classes and
especially the nobility (e.g., Albin 1738a, 1740, Edwards 1751). Edwards (1751: PI. 161)
stated that 'the large Green Parrots, [are] commonly brought to us from the West-Indies,'
and elsewhere (1751: PI. 161) said the 'Great Green Parrot, from the West-Indies' was
'pretty common in London.' Edwards (1751: PI. 168), speaking of the 'Little Green Parrot,'
said '...I take it to be from the West-Indies, from whence most of the Green Parrots we
have in London are brought.' Further, authors of descriptive works on parrots frequently
mentioned several London and European dealers specialising in foreign birds, including
macaws. Apparently even stuffed parrots found a market in Europe, as Edwards (1751: PI.
173) mentioned buying a 'beautiful and rare little Parrot ... exposed to Sale (stuffed, and set
on a Perch) in a Toy-Shop Window, in London'.

The great incentive to harvest parrots for European trade can be appreciated from the
high prices commanded by macaws and others of their tribe there. Even in their native
islands, psittacines were worth a considerable sum of money. For example, of Amazona
parrots, Taylor (2008) recorded that '...a young one alive out of the nest will cost six royalls
[reales] at Jamaica.' But captive parrots were valued far more in Europe. As examples, Albin
(1738b: PI. 13) wrote that a 'Laurey from the Brasils' [Purple-naped Lory Lorius domicella]
sold for 'twenty Guineas,' the equivalent in today's money of c.£1725; Blue-and-yellow
Macaw was 'commonly sold for ten Guineas' (c.£860 today) (Albin 1840: PI. 10); and the
'Maccaw from Jamaica' [likely Ara gossei, but perhaps A. macao], which was 'commonly
brought from Jamaica, and other parts of the West-Indies,' were 'commonly sold for ten
Guineas' (Albin 1738a: PI. 16). About 1680, the cost of a parrot in Amsterdam was recorded
as 'roughly sixty guilders' (Margoesy 2010), when a guilder was a 10.61 g, 90% pure silver
coin, demonstrating the high value of pet birds. Such prices initially made owning a macaw
or parrot accessible only to the affluent and thus a substantial status symbol in Europe.
But as more parrots were imported, classes other than the nobility and aristocrats were
able to acquire them. Thus wealthy merchants and affluent citizens sought the parrots
that were becoming staples in the growing number of bird dealers' shops. Thomas (1983)
wrote that commercial bird dealers first appeared in England 'in Tudor times and in the
late seventeenth century there was a large London market in singing-birds, some caught
at home by professional bird-catchers, others exotics imported from the tropics.' Trade in
parrots continued to increase into the 19th century, with parrot popularity as household
animals reaching its peak in the 1800s (Boehrer (2004). This high point in the importation of
psittacines coincides with what is probably the period when the last West Indian macaws
became extinct.

Macaws were kept as companions by colonists in the islands. Du Tertre (1667) included
an illustration depicting what appears to be a macaw perched at a 17th-century plantation.
Taylor (2008) noted that macaws were 'kept tame as a curiosity' in Jamaica, and Gosse
(1847: 263) reported that a captive Jamaican macaw kept by a Mr White was 'for some time
the admiration and talk of the country round.'

Trade in all Cuban psittacines has long been a cause of concern (Gundlach 1893, Bangs
& Zappey 1905, Todd 1916, Noegel 1979) and is vigorous today, even under national and
international regulation, bringing about modern declines. The enormity of the trade in
recent times is particularly well documented on Isla de Pinos, where parrots and parakeets
were formerly numerous (Gundlach 1893, Gonzalez Alonso et al. 2012). Gundlach (1893)
predicted that if rates of capture for the pet trade were maintained, the parakeet would be
exterminated on Isla de Pinos in a few years. Sadly, his prediction was fulfilled; the parakeet
was extirpated early in the 20th century (Bangs & Zappey 1905). The parrot, however, has
survived there, albeit in much-reduced numbers.

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Although we have found no data to appreciate how many macaws and other psittacids
were being exported during the 16th through 19th centuries, the few statistics of the
enormous parrot trade in Cuba during the early 20th century provide an appreciation
of the international demand in that period. During the first decade of the 20th century,
some 3,000-5,000 pairs of parrots were exported from Isla de Pinos to the USA annually
(Anon. 1910a). Because of their depredation of crops, parrots were unprotected until 1909.
Numbers increased rapidly thereafter, but protests by planters soon led to the protective
legislation being repealed and within a few years large numbers were again being exported;
e.g., one large shipment of 2,500 parrots in 1910 (Anon. 1910b, c) and c. 1,000 birds imported
by one US dealer in July 1914 (Anon. 1914). It is certainly within reason to assume that the
Cuban Macaw was even more desirable than parrots and parakeets (Olson 1982).

Cuban Macaws were kept locally and also exported to private collectors and royal
and public zoos overseas, leading several writers to suggest that harvest for the pet and
commercial trade was in large part responsible for its extinction (Gundlach 1874, Lonnberg
1928, Moreno 1992, Dathe & Gonzalez Lopez 2002). Local interest in Cuban Macaws as pets
had a long and consistent history, until the time of its extinction. Gundlach (1865-66, 1873,
1874, 1876) reported that the macaw was a much sought-after cagebird, but noted (1874)
that captives damaged furniture and other items using their powerful bills. Yet, despite this,
we suspect that the macaw was a more desirable pet in Cuba than Cuban Parrot, which
continues to be vigorously sought by island residents and, despite strong legal protection,
is common in households.

Cuban Macaws were equally popular overseas. Garcia Montana (1980) said that they
were given to the kings of Spain, and that over the centuries thousands were exported
to Europe. Barbour (1943) supported that claim and Finsch (1863) stated that it was well
known in captivity in Europe, but even so was rare. Ldnnberg (1928) suggested that at one
time Cuban Macaw had been very popular and that it was kept as an ornamental bird in
Sweden in the 1700s.

The ever-increasing local and international demand for parrots certainly had a
devastating effect. Lonnberg (1928) recognised the problem of such harvesting of young
macaws in that natural reproduction was greatly reduced. As populations of native
macaws diminished and it became more difficult to supply local demand, exotic species
were imported to meet it. Exotics from other islands and beyond were imported to supply
cagebirds for local markets in the face of dwindling native populations. Importing macaws
and parrots during the colonial period is particularly well recorded in Jamaica, where, as
early as the 17th century, Taylor (2008) wrote that the 'Affrican or Guinea parrat' (Grey
Parrot Psittacus erithacus) was 'brought to Jamaica and sold at reassonable rates.' Regarding
Grey Parrot, Edwards (1751: PI. 163) wrote 'I am well assured that what we have are brought
from Africa, generally by the Way of the West-Indies, by our Guinea Traders, that supply
our Sugar Islands with Negroes.' Sloane (1725: 297) also reported that Grey Parrot was
'brought to the Island of Jamaica in great Quantities from Guinea' [Africa]. Further, Sloane
(1725) said 'Psittacus viridis alarum costa superna rubente’ [Red-shouldered Macaw Diopsittaca
nobilis or Yellow-crowned Parrot Amazoua ochrocephala ; neither from the West Indies] was
brought to Jamaica 'from the Spanish Main, or Continent of America frequently hither'. He
also noted that Cuban Parrots were 'brought from Cuba to Jamaica frequently.' In addition,
Sloane (1725: 296) recorded that 'the Great Maccaw' 'Psittacus Maximus cyanocroceus’ ( Ara
ararauna of South America) was kept in captivity. Browne (1756: 472) noted that most of
the 'blue Mackaw[s] of Edwards' were introduced to Jamaica from the mainland. Further,
Browne (1756: 472) said the non-native 'red Mackaw of Edwards Psittacus maximus Jons' was
frequently brought to Jamaica from the neighboring parts of the main'.

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Further evidence of extensive trade in exotic macaws in Jamaica comes from a painting
by the Revd. John Lindsay dated 1765 and part of a series of volumes of watercolour
illustrations and manuscript descriptions of Jamaican flora and fauna in the collection of
the Bristol City Museum and Art Gallery. The macaw appears to be Scarlet Macaw, and
probably represents a captive individual or introduced population (Crane 1981, Fisher &
Warr 2003, Turvey 2010). Scarlet Macaw was one of the most popular of parrots among
18th-century Parisian nobility (Robbins 2002) and was widely traded in the Americas by
pre-Columbian Amerindians (Olson & Maiz Lopez 2008). Thus, the species could have been
imported to Jamaica via trade.

One of the attractions of psittacines as pets is their ability to 'speak', and that capacity
was often noted by early writers. Taylor (2008) reported that Jamaican macaws 'spake much
plainer and lowder than any parrat whatsoever.' Labat (1742: 212) noted that West Indian
macaws could be tamed and, especially if taken young, spoke very well, their voices being
strong and distinct. Sloane (1725: 296) said a Jamaican macaw, probably A. erythrura, was
'more Articulate than any Bird I ever heard.' In contrast, accounts gave Cuban Macaw
poor marks regarding speaking ability. Gemelli Careri (in de la Torre 1857) remarked that
Cuban Macaws could not speak. Gundlach (1865-66, 1873, 1874, 1876) said that although it
imitated some words, Cuban Macaw did not have the skills of Cuban Parrot. It is probable,
however, that ability to talk took second place to the size and showiness of macaws.
Several other observers noted their poor speaking abilities. Of Lesser Antillean macaws, de
Rochefort (1658: 154) commented that the tongue was 'too thick to enable them to speak as
well as parrots and the smaller parakeets', and Atwood (1791: 29) said macaws 'cannot be
taught to articulate words.'

West Indian macaws were often kept in royal menageries and public zoos. Taylor
(2008) said that he saw one Jamaican macaw ('marcough' Ara sp.) 'amongst His Majesty's
birds at Saint James's Park [London].' Moreno (1992) noted that many Cuban Macaws
were exported to Europe where they were sought for exhibit in zoological gardens. Cuban
Macaw was probably not rare in zoological collections in Europe judging from the number
of specimens preserved after dying in zoos. In total, six specimens are known to have been
kept previously in zoos, while an additional four were probably held as captives (Table
1). Additional Cuban Macaws were also reported in zoos, but are not known to have been
deposited in museums after death; e.g., Bolle (1856) reported one in the London Zoo on 2
January 1856 (perhaps BMNH 1858.5.13.1?).

Habitat destruction. — D'Orbigny (1839: 120-121) noted that Cuban Macaw was
becoming increasingly rare as natural lands were cultivated in association with human
population growth. Later writers also considered habitat destruction as probably
contributing to the macaw's extinction (e.g., Garcia Montana 1980, Moreno 1992, Dathe &
Gonzalez Lopez 2002). Of particular harm was the way in which nestlings were obtained
for trade. Gundlach (1893) described how adults were observed to find the nest tree, then
the harvester would wait until the chicks were well grown before felling the tree to obtain
the nestlings. Some chicks undoubtedly died as a result, but a more serious and long-
lasting effect was the cumulative loss of the best nesting habitat; i.e., palms with cavities of
sufficient dimensions and security. After many years of such selective destruction, macaws
may have been left with few, and possibly only suboptimal, nest sites. This method of
harvesting psittacine chicks in Cuba has continued to the present, to the point that such
habitat destruction has greatly affected populations of the two surviving species (de las
Pozas & Gonzalez Alonso 1984, Kirkconnell & Wiley in prep.).

This practice was also common on other islands and may have affected their macaws
as well. Taylor (2008) wrote that parrots and parakeets in Jamaica 'commonly breed in the

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TABLE 1

Museum specimens of Cuban Macaw Am tricolor known or suspected
to have originated as zoo captives or cagebirds.

Museum1 and catalogue no.

Zoological institution where
formerly held

Comments

BMNH 1858.5.13.1

Presumably a captive bird

Obtained from the Zoological Society (Knox &
Walters 1994).

WML D645

Knowsley Park aviaries of

13th Earl of Derby

Died March 1846; arrived in museum 1851.

MNHN 2000-725

Jardin de Plantes, Paris

Donated to museum in 1842 after its death
(Jouanin 1962).

NMW 50.796

Imperial Menagerie, Schonbrunn

Died 1835 (Fitzinger 1853) or > 1806 (Moreno

1992).

SMNG A03466a

Probably a cagebird (C. Diicker pers. comm.).

NRM 569592

Thought to have been a cagebird in Stockholm;
wing feathers clipped (U. Johansson pers.
comm.).

NRM 523094

Amsterdam Zoo

Died 1858

SMTD 3466

Probably formerly a captive, by its frayed
plumage (M. Packert pers. comm.).

MCZ 72526

Obtained from Lafresneye Collection, France;
with one clipped wing,

AMNH 205178

Zoological garden of Berlin

Presented to museum by Barbour, 1923.

'Abbreviations: BMNH = Natural History Museum, Tring; WML = World Museum, Liverpool; MNHN = Museum
National d'Histoire Naturelle, Paris; NMW = Naturhistorisches Museum, Wien; SMNG = Senckenberg Museum fur
Naturkunde, Gorlitz; NRM = Naturhistoriska Riksmuseet, Stockholm; SMTD = Staatliches Museum fiir Tierkunde,
Dresden; MCZ = Museum of Comparative Zoology, Cambridge, Massachusetts; AMNH = American Museum of
Natural History, New York.

hollow trunck of cabadge trees up in the mountains soe that they are hard to come by, and
you must fell the tree to come at 'em.'

Crop pest. — Gund lach (1865-66, 1873) mentioned that Cuban Macaws may have caused
damage to fruit trees, but noted (1874, 1876) that, because they lived far from dwellings, the
damage was not great (but added, also, that the bird provided no benefit). Hill (in Gosse
1847: 261) noted that macaws in Jamaica fed in 'small companies' on the 'full-eared maize,
while the grain was soft, milky, and sweet, and the very husk was sugary.' There is no
reason not to believe that Cuban Macaw also depredated crops and, as a consequence, was
persecuted. Today, Cuban Parrots and, especially, Cuban Parakeets are soundly punished
for such infractions!

Hurricanes. — Barbour (1923) reported that Cuban Macaw disappeared from western
Pinar del Rio after the great hurricane of 1844 (4-5 October). Even if that storm did not
eliminate the last macaws in western Cuba, the devastating hurricanes of 1846 (the 'Great
Havana Hurricane'; 10 October; Category 5) and 1856 (late August; Category 3) could have
further decimated remnant and fragmented habitat, and scattered populations beyond
recovery. Similarly, a strong tropical storm ravaged Cienaga de Zapata on 21 August 1851.
When original forests are extensive, hurricanes have positive effects, producing habitat for
cavity-nesting birds. However, once macaw populations were restricted to a few, relatively
small areas of fragmented habitat, a direct hit by a powerful storm could destroy sufficient
critical habitat to cause extirpation (Wiley & Wunderle 1993). Some of the small, disjunct
populations of the Critically Endangered Puerto Rican Parrot Amazona vittata were probably
destroyed by such events (Snyder et al. 1987). Not only did those populations suffer the

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direct effects of the storm, but habitat critical to their survival was severely damaged.
Further, in the aftermath of hurricanes, parrots were more vulnerable as they moved to
hostile areas seeking food, meeting humans also desperately seeking to survive.

Disease. — Imported pets and domestic fowl may have carried exotic diseases, which
could have spread to wild indigenous parrots, and against which they had little or no
resistance. Du Tertre (1654: 294) mentioned that although Guadeloupe Macaw was long-
lived ('live longer than a man'), they were 'almost all subject to a falling sickness [epilepsy].'
Labat (1724: 217) also remarked on the long-lived psittacids of the West Indies, and their
susceptibility to the 'falling sickness.' De Rochefort (1658: 154) noted that Lesser Antillean
macaws were 'so sensitive to cold that it is difficult to bring them across the sea.' Williams
& Steadman (2001) suggested that a disease outbreak, combined with hunting pressure,
caused the extinction of Guadeloupe Macaw.

Specimens of Ara tricolor

No modern skeletal specimens of A. tricolor exist (Williams & Steadman 2001) but we
can account for 19 skins or mounted specimens in 15 collections (Table 2). All were collected
in Cuba, or from an unknown locality. When and from where and whom the museums
obtained these specimens is unresolved in many cases.

Gundlach's specimens.— Gundlach collected several of the
last group of Cuban Macaws that came regularly to feed at
a site in Cienaga de Zapata in 1849-50 (Barbour 1923). These
may have been the only macaws Gundlach collected, because
he does not mention additional specimens taken during later
visits to the Cienaga. If true, Gundlach must have kept several
specimens for years, eventually giving all but one to friends
and institutions. Before he left Germany for Cuba in November
1838, an organisation was formed to furnish support funds by
the sale of stock, to be repaid by Gundlach with specimens
collected during the expedition, which was originally planned
to continue to Surinam. Although Gundlach made it only as
far as Cuba, he set to work collecting and shipping materials
to those who had bought stock, not ceasing until he paid all his
debts. Even after meeting his financial obligations, he continued
to send specimens to Germany, including his precious macaws.

Cory (1889) mentioned that Gundlach had several macaw
specimens, by which time Gundlach had dispersed a good
portion of his collection to foreign institutions. Although
Gundlach did not record how many macaws he collected, it was
at least four (see below). By the time of his death in 1896, only
one remained at the Institute de Segunda Ensenanza, where
Gundlach's collection was maintained (Ragues 1914).

During the mid-19th century, it became fashionable
among wealthy Cubans to adorn their drawing-rooms with
natural history specimens, and Gundlach gifted friends and
acquaintances specimens he had prepared. He thus became known to the most influential
citizens of that day, who in turn provided him with assistance (Ramsden 1918). Gundlach
may have included one or more macaw specimens among those given to Cuban friends and
patrons (see below).

TABLE 2
Distribution of known skin
or mounted specimens of
Cuban Macaw Ara tricolor
among 15 collections
worldwide.

Collection

No.

IES

BMNH

WML

MNHN

NMW

ZMB

SMF

SMNG

NRM

SMTD

EHM

RMNH

MCZ

AMNH

USNM

'Stolen in 2007.

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Specimens in Cuban ami foreign institutions

Instituto de Ecologia y Sistematica (IES), La Habana, Cuba One: CZACC-6. 59937. No date
or precise locality; coll. Juan Gundlach. Adult. Mount. Specimen no. 165 from the Gundlach
collection formerly in the Instituto de Segunda Ensenanza, it was stolen from IES in 2007.

Natural History Museum, Tring (BMNH) Two: BMNH 1851.7.21.5. No date (received
no later than 1859), locality or collector. Adult. Skin, ex-mount. Purchased from Lt. A.
Smith, but R. Prys-Jones (pers. comm.) thinks tins may be in error, and all that can be said
safely is that it was purchased. Knox & Walters (1994) noted that the correct registration
number should be 1851.7.24.5. BMNH 1858.5.13.1 No date, locality or collector. Adult. Skin,
ex-mount. Source given as Zoological Society and presumed to have been a captive bird
(Knox & Walters 1994).

World Museum, Liverpool (WML) One: D645. No date or collector. Locality given as
'Mexico'. $. Skin. Obtained from Knowsley Park aviaries of 13th Earl of Derby, where it
died March 1846. Acquired in 1851, and maintained today in Derby Collection.

Museum National d'Histoire Naturelle, Paris (MNHN) Two: MNHN 2000-725. No date,
locality or collector. Adult. Mount. Maintained in captivity in Jardin de Plantes, Paris, from
30 May 1840 and, after its death, on 6 October 1842, presented to MNHN by L. F. E. Rousseau
(1788-1868), an assistant naturalist at MNHN (1834-60) (from Catalogue des Animaux morts
a la menagerie 1840-1853, where noted 11 October 1842; A. Previato pers. comm.). MNHN
CG 2000-726. No date, locality or collector. Adult. Mount. Type. Jouanin (1962) noted it as
from 18th century. M. Portas (pers. comm.) believes it was from old collection of Jardin du
roi (museum before Revolution = pre-1789). It formed part of the 'Exposition permanente
des Produits de l'Algerie et des Colonies,' displayed in the Palais del'Industrie in Paris in
1860, and curated by Aubry-Lecomte, but no information as to duration of exhibit or when
transferred to MNHN (A. Previato pers. comm.).

Naturhistorisches Museum, Wien (NMW) One: NMW 50.796. No date, locality or
collector. $. Mount. Obtained from Imperial Menagerie of Emperor Franz II (Franz I of
Austria; 1768-1835) (de Germiny 1937; G. Heindl pers. comm., H.-M. Berg pers. comm.).
Acquired by NMW in 1832, the same year it died at the menagerie (Fitzinger 1853: 166; H.-M.
Berg pers. comm.), although Moreno (1992) set date of acquisition as 1806. Fitzinger (1853:
86) listed two A. tricolor, a male and female, in the menagerie as from 'Brasilien. 1760,' but G.
Heindl (pers. comm.) noted that one of these birds was probably A. macao. G. Heindl (pers.
comm.) suggested the two specimens, including the Cuban Macaw, from 1760 were collected
during Nicolas Joseph Jacquin's expeditions to the Caribbean islands in 1754-59, including
Cuba in 1759, but Jacquin (1784) does not include A. tricolor among species reported by his
expedition (C. Riedl-Dom pers. comm.).

Museum fur Naturkunde, Zentralinstitut der Humboldt-Universitat, Berlin (ZMB) One:
ZMB 24886. No date or locality; coll. Gundlach, probably in 1849 [1850?] at Hanabana,
Cienaga de Zapata (Stresemann 1954). $. Adult. Skin. Sent by Gundlach to ZMB in
November 1880 (letter from Gundlach to Peters, 16 November 1880; S. Frahnert pers.
comm.). Received by ZMB in December 1880.

Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt (SMF) One: SMF 17345.
No date, locality or collector. Ex-mount. From Hartert Coll. (No. 2399) (Mertens &
Steinbacher 1955).

Senckenberg Museum fur Naturkunde, Gorlitz (SMNG) One: SMNG A03466a. No date,
locality or collector. $. Mount. Possibly a cagebird (C. Diieker pers. comm.). Donated
(without year) to 'Naturforschende Gesellschaft zu Gorlitz' by Dr H. Bdttcher (1834-1904)

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from village of Quolsdorf. Bottcher bought the macaw from G. Schneider (1834-1900), a
dealer in natural history objects and corresponding member of the Goerlitz Society, Basle,
Switzerland. Bottcher, a renowned bird collector, donated more than 100 birds to the Gorlitz
museum in 1861; the macaw might have been included in that gift (C. Diieker pers. comm.).

Naturhistoriska Riksmuseet, Stockholm (NRM) Two: NRM 569592. No date, locality or
collector. Mount. Very old specimen from private collection of A. U. Grill (1752-92). Grill's
collection was donated to NRM in 1829. No date when acquired by Grill, but probably
c.1791-92 (U. Johansson pers. comm.). Locality given as 'Stockholm?', which U. Johansson
(pers. comm.) interprets as it having been a cagebird in Stockholm. NRM 523094. No date,
locality or collector. Mount. Purchased 1858 from G. A. Frank, a dealer in natural history
objects in Amsterdam. Frank bought 80 birds from T. G. van Lidth de Jeude (1788-1863)
and U. Johansson (pers. comm.) believes the NRM macaw came via this route, although it
is unclear whether there is any documentation in favour of this supposition. The van Lidth
de Jeude collection originated in the early 18th century and was assumed to include many
old specimens (Boeseman 1970). U. Johansson (pers. comm.), however, notes that if a bird
died in Amsterdam Zoo in that period, the NRM specimen may originate from there (see
Lonnberg 1928). Moreno (1992) noted that a Cuban Macaw died in the Amsterdam Zoo in
1858, and presumed it was the NRM specimen.

Staatliches Museum fur Tierkunde, Dresden (SMTD) One: SMTD 3466. No date, locality
or collector. Mount. Moreno (1992) said it was part of an old collection from mid-19th
century. Formerly a captive, by its frayed plumage (M. Packert pers. comm.).

Eichsfelder Heimatmuseum, Heilbad Heiligenstadt (EHM) One: EFIM IA1/310. No date,
locality or collector (fide V. Merten). Mount. From among 750 bird and other natural history
specimens accumulated by K. W. J. Strecker (1818-87) of Dingelstadt. Strecker donated, via
his will, his collection to EHM in 1885.

Nationaal Natuurhistorisch Museum, Leiden (RMNH) One: RMNH.Aves. 110095. No
date or collector. Locality given as 'Cuba'. Mount. No further specimen or acquisition
information (S. van der Mije pers. comm.).

Museum of Comparative Zoology, Harvard University, Cambridge, MA (MCZ) One:
MCZ 72526. No date, locality or collector. Original label missing. Ex-mount. From
Lafresnaye Collection. Fine condition, but one wing clipped, suggesting a cagebird.

American Museum of Natural History, New York (AMNH) One: AMNH 205178. No
date, locality or collector. Mount. Lacks tail. Died Berlin Zoo, date unknown. T. Barbour
presented it to AMNH in 1923. Original label inscribed 'Psittacus tricolor Bechst. 1881 West
Indies, 10189,' as well as a note in ink: 'Zool. Garden, Berlin.' Also: AMNH FARB 6199 ?Ara
tricolor, proximal end of carpometacarpus Pleistocene, Banos de Ciego Montero; coll. B.
Brown, 1918.

United States National Museum of Natural History, Smithsonian Institution, Washington
DC (USNM) Two: USNM 135137. No date or locality; coll. Gundlach. Skin. Acquired by
AMNH from Gundlach 13 December 1894. USNM 17167. No date, locality or collector.
Mount. On exhibit in Hall 13 'Birds of the World.' History of this specimen involved and
partially unsolved. Probably collected in Cienaga de Zapata by Gundlach and carried
with him, along with other birds, to Puerto Rico during Cuba's Ten Years War (1868-78).
Barbour (1923) said Gundlach used the specimens, which also included two Ivory-billed
Woodpeckers, to repay favours afforded by his friends, probably the apothecary and
naturalist Tomas Blanco y Gonzalez (1840-92) and naturalist Agustin Stahl (1842-1917).

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With the encouragement of Blanco and Stahl, in 1866 the Padres Jesuitas de Puerto
Rico invited Gundlach to build a natural history collection for the Colegio Seminario they
had established in San Juan in 1865. Gundlach eventually made two extended trips to
Puerto Rico to collect specimens for the Instituto Civil de Segunda Ensenanza de San Juan,
established the same year as his first expedition (1873; Lopez Yustos 1991). Stahl became a
professor of natural history there, and Gundlach apparently provided the woodpecker and
macaw specimens. Stahl and Blanco maintained the specimens at the Instituto, along with
others collected by Gundlach during his 17 months in Puerto Rico. But, after the Spanish-
American War (1898) and the Treaty of Paris, the Instituto was closed in 1899. Blanco had
died in 1892 and Stahl was deported from Puerto Rico by the Spanish government in 1898,
so the Instituto collection had lost its main advocates. Barbour (1923) thought Gundlach's
Ivory-billed Woodpeckers were later acquired by AMNH, which matches at least one of the
three specimens, AMNH 144873, a male, which has two labels, one of which reads, 'This
bird was collected by Gundlach in Cuba and sold by him to Porto Rico High School.' The
second label says 'Purchased from High School collection at St. Juan Porto Rico in 1901.' The
AMNH catalogue adds 'collected in Cuba and sold to the Porto Rico High School where it
was found' (P. Sweet in litt. 2012).

The macaw was apparently found in the Instituto by Major W. A. Glassford, US Army,
Chief Signal Officer, stationed in San Juan, Puerto Rico during the post-Spanish- American
War occupation by the US military. C. W. Richmond (1868-1932), who was in Puerto Rico
collecting birds and herptiles with L. H. Stejneger (1851-1943) from 12 February to 19 April
1900, received the specimen from Glassford, whereupon Richmond deposited the macaw at
USNM (accession date 16 May 1900).

Hypothetical or lost specimens. — Several rumours exist of additional specimens of Cuban
Macaw in collections within Cuba, and some may have been extant quite recently. One
rumour involves a macaw among the natural history exhibits at the Escuela 'Rafael
Maria Mendive' (= Colegio Dolores or Convento de Dolores) at Santiago de Cuba. G. G.
Hechavarria and L. O. Melian Hernandez (pers. comm.) recalled that the exhibits contained
a large collection of mounted birds including two Ivory-billed Woodpeckers and a Cuban
Macaw. By the time JWW was permitted to visit the school in May 2006, all that remained
were four badly damaged mounts, none of them a macaw. The school's director informed
JWW that all of the other specimens had been destroyed during building reconstruction
over the previous two years. The director said that, until the reconstruction, the natural
history materials, including a macaw, had been well conserved and the most valuable
specimens were in a case away from the main school activities.

That such specimens may have existed in the Escuela 'Rafael Maria Mendive' is
supported by Leyva (1922), who noted a specimen of Ivory-billed Woodpecker and
collection of butterflies made by Gundlach in the Museo Municipal de Santiago de Cuba,
part of which may have found its way to the Escuela. O. H. Garrido (pers. comm.) recalled
that he and F. Garcia Montana obtained a mounted specimen of the near-extinct Cuban Kite
Chondrohierax uncinatus wilsoni from the same collection. Further, documents in the Archivo
Nacional de Cuba include records that establish the collection of birds in the Escuela came
from the Brooks family, which may link that collection to Gundlach. In 1884-85, Gundlach
was guest of his friend Theodoro Brooks at Cafetal 'Jaguey' in the mountains of Yateras and,
as characteristic of Gundlach, he may have given Brooks bird and other specimens to repay
the family for their hospitality.

Barbour (1945) recounted that other macaw specimens had been in Cuban collections,
including one in the cabinet of the Habana Academy of Sciences, but that disappeared
and was rumoured to have gone to a notorious private collector abroad. Fuller (2001:

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236) suggested that collector was Walter Rothschild. Barbour (1945) reported that another
specimen was said to have been in a museum at Cardenas, but thought it had been
destroyed by insects; JWW found no record of it at Museo 'Oscar Maria de Rojas/ the
museum presently holding natural history specimens in Cardenas. Barbour (1945) claimed
that yet another specimen was formerly in the Matanzas Institute.

Among Cuban Macaw specimens not currently accounted for in museum collections is
one that was part of the collection of Prince Massena d'Essling, Duke of Rivoli (1799-1863)
(Souance 1856: 57). Massena accumulated a collection of 12,500 avian specimens, which
he sold to Dr T. B. Wilson in 1846. Wilson gave the collection to the Academy of Natural
Sciences of Philadelphia, but that collection does not contain a Cuban Macaw.

Discussion and Conclusions

Additional sources of information on West Indian macaws. — It is reasonable to hope
that further paleontological and archaeological investigations will yield data to clarify the
numbers of species and their distribution in the West Indies. Current work by W. Suarez
& S. L. Olson in Cuba and their planned investigations of fossil and subfossil materials in
Hispaniola may provide the type of information that will enable a better understanding
of macaw populations. Investigation of Amerindian sites also may produce further
information on macaw distribution and the interactions of native peoples with indigenous
and introduced macaws.

Additional research of published and manuscript accounts by early travelers to the
Antilles could yield confirmation of known reports of macaws. Also, correspondence
archived in natural history museums may provide records of additional macaw specimens,
now lost. We also hope that further surveys of museums, large and small, may reveal
additional macaw specimens.

Another potential source of valuable information on West Indian macaws may lie
in art museums, especially in Europe. Many parrot owners included macaws within
formal portraits that hang on gallery walls. Macaws were considered prestige symbols,
especially in the Renaissance, and often appear in the background or alongside their
owners. Although often stylised, some depictions approximate species suggested for the
West Indies. As Cooper & Armitage (2013) have suggested, examination of such art could
prove a fertile field for learning more of macaw distribution and description. Greenway
(1967) noted that a macaw in an early 17th century painting by Roeland Savery matches
descriptions of some West Indian macaws. Macaws feature prominently in other paintings,
including Bartholomeus van Bassen's (1590-1652) 'Renaissance interior with banqueters'.
Hume & Walters (2012), however, urged caution in intrepreting such evidence as found in
paintings, noting that artistic license may alter an artist's rendition of a bird, as well as the
fact that the painting may have been made from fading memory, plagiarised or modified
to suit composition.

Extinction of West Indian macaws.— Any one of the many problems that faced
the Cuban and other West Indian macaws could have caused their extinctions, but the
combination of environmental changes and human-related pressures most certainly sealed
the fate of all Antillean macaws. Killing of adult macaws for subsistence or to protect
crops probably had a substantial effect; although such persecution had been underway for
some two millennia, the arrival of Europeans with their advanced guns must have vastly
increased the efficiency with which macaws could be killed, thereby accelerating population
declines. Similarly, harvest and trapping of wild macaws had its origin in Amerindian
culture, but it was not until the opening of many markets, far and wide, that populations
suffered to the point of extirpation. As Europeans colonised the Antilles, they increased the

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speed and extent of habitat modification, through improved tools and ever-growing human
populations. Particularly devastating was the practice of felling macaw nest trees to obtain
young birds for personal use or trade. With the high demand for macaws, domestically and
abroad, the harvest of nestlings and associated destruction of nesting habitat must have
had a profound effect. The effects of other factors, including hurricanes and disease, are
less easily predicted, but nonetheless potentially contributed to declines and extinctions.
Disease, perhaps introduced via domestic poultry or exotic parrots bought to islands in
trade, and to fill the demand for pets as local species became rare or disappeared, could
have had an immediate, fateful effect on populations reduced to small numbers and
restricted to remnant habitat. Island populations of birds are particularly susceptible to
introduced diseases, often against which no defences exist. Whereas hurricanes have always
been powerful forces of change in the West Indies, even the strongest storm probably had
little effect on healthy populations of macaws when original habitats were extensive. Once
habitat was fragmented by human activities and macaws had been reduced and confined to
a few remnant habitats, a direct hit by a powerful storm would have destroyed forest critical
to a population's survival, as well as perhaps killing birds outright. No matter whether one
or a combination of factors led to the extinction of the Antillean macaws, certainly birds so
rare, so easy to kill, so much in demand, and so very beautiful had little hope for survival.

Number of spiecies and distribution in the Antilles. — Macaws certainly occurred
over a wide area of the Caribbean, and on several islands, but the number of species is
controversial. At this point, with available evidence, just how many species and on which
islands depends in large part on how much credence is given to the vague accounts of early
explorers, travelers and colonists. If all records unsupported by skins, fossil or subfossil

TABLE 3

Records of extinct Ara and Anodorhynchus in the West Indies, with level of confidence of their former

existence based on source of evidence.

Island

Cuba

Isla de Pinos
Jamaica

Hispaniola
Puerto Rico
St. Croix
Montserrat
Guadeloupe

Marie Galante

Dominica

Martinique

Species

Ara tricolor

Ara erythrura

Ara erythrocephala

Ara gossei [Ara tricolor ?]

Ara tricolor ? or Ara unknown species
Ara autochthones
Ara autochthones
Ara undescribed sp.

Ara guadeloupensis
Anodorhynchus purpurascens

Ara cf. guadeloupensis

Ara atwoodi
Ara martinica

Anodorhynchus martinicus

Evidence

19 skins; fossil materials; last recorded
1850s

Multiple accounts; last recorded in 1864

Multiple accounts; possible introduced
species

Multiple accounts

Skeletal material from kitchen midden
Skeletal material from kitchen midden
Skeletal material from archaeological site
Multiple accounts & possible illustration
Multiple accounts; possible introduced
species

Skeletal material from archaeological site;
= Amazona imperialist
Single account

Single account; possible introduced
species

Synonym of Ara martinica

Confidence

Specimens

Good, based on
competent reporter

Poor

Good, based on
competent observers

Poor

Specimen

Moderate

Poor

Specimen

Poor

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remains are rejected, the number of species may be just three (Cuban Macaw, St. Croix
Macaw, Montserrat Macaw) from five islands (Cuba, Isla de Pinos, Puerto Rico, St. Croix
and Montserrat) (Table 3). Even among those species and islands with specimens, questions
persist as to whether at least some species were native or transported there. If the list of
probable species is extended to accept the most reasonable of those species supported only
by written accounts, the number of islands and species is increased by only one: Gosse's
Macaw from Jamaica. Yet even that species, the best described of all of the other purported
macaws in the region, could have been based on a feral population of Scarlet Macaw
introduced via trading and pet-keeping activities in the colonial period.

Among other suggested species, we consider the possibility of Guadeloupe Macaw
moderately plausible. But, again, existence of Guadeloupe Macaw as endemic is confounded
by possible introduced or transported species.

We consider all other species of macaws noted by early writers and formalised by
Rothschild and others as poorly supported by current evidence. To date, no reliable
verification is available of a macaw on Hispaniola. The existence of Dominica Macaw
rests solely on Atwood's account, and macaws on that island may have their origin as
an introduced or transported species. One named species, Anodorhynchus martinicus, is a
synonym of Ara martinicus. Guadeloupe Violet Macaw was fairly well described as unique
by early explorers, but there is a reasonable likelihood that this was a South American
species transported to the island by Amerindians or colonists. Similarly, two, and perhaps
all, of the three macaws listed by some for Jamaica were possibly feral birds resulting from
releases or escapes of imported birds. Certainly, a vigorous trade in exotic parrots was
established at the time accounts of macaws on Jamaica were recorded and their descriptions
could relate to species known to have been traded on the island.

What we know of psittacid speciation and distribution in the Antilles supports the
possibility, and even probability, of multiple endemic species of macaws, so there is
no reason why each of the macaws described by explorer-naturalists could not have
existed. Nevertheless, although the early accounts are alluring, without further proof, we
recommend a conservative stance. We hope that future information, perhaps best searched
for in the fossil and subfossil record, will provide solid proof that the primordial forests of
most Antilles were alive with the squawks and brilliant plumages of macaws.

Acknowledgements

We are grateful to the many museum and archive curators who provided information on macaw specimens
and records. Among these we particularly thank Robert Prys-Jones and Mark Adams, Natural History
Museum (Tring); Veronika Merten, Museum Director, Eichsfelder Heimatmuseum; Christa Riedl-Dom,
Archiv fur Wissenschaftsgeschichte, Naturhistorisches Museum, Wien; Hans-Martin Berg, Naturhistorisches
Museum, Wien; Marie Portas and Anne Previato, Museum National d'Histoire Naturelle, Paris; Ulf Johansson,
Naturhistoriska Riksmuseet, Stockholm; Craig Ludwig and James Dean, United States National Museum of
Natural History, Washington DC; Tony Parker, National Museums Liverpool; Sylke Frahnert, Museum
fur Naturkunde, Humboldt-Universitat, Berlin; Christian Duker, Senckenberg Museum fur Naturkunde,
Gorlitz; Steven van der Mije, Nationaal Natuurhistorisch Museum, Leiden; Paul Sweet and Carl Mehling,
American Museum of Natural History, New York; Clare Flemming, Ewell Sale Stewart Library and Archives,
Academy of Natural Sciences, Philadelphia; Gerhard Heindl, Schonbrunner Tiergarten Gmbh, Wien; Nancy
Machado Lorenzo, Biblioteca Nacional 'Jose Marti', La Habana; Barbara Danzie and Marta Casal, Archivo
Nacional de Cuba, La Habana; Aurea Roman Fandino, Maria Herminia Pandolfi Camara, Rossio Boronat,
Regia Hernandez, Magalis Reyes and Graciela Guevara, Museo Nacional de Historia de las Ciencias 'Carlos
J. Finlay', La Habana; Rafaela Aguiler Roman and Naomi Cuervo Pineda, Instituto de Ecologia y Sistematica,
La Habana; and Colin Banfield, St. James' Park, London. Ariel Ruiz Urquiola, Centro de Investigaciones
Marinas, Universidad de La Habana, and Nils Navarro, provided information on historic habitat. Lynda
Garrett, Patuxent Wildlife Research Center, and Lloyd F. Kiff and Travis Rosenberry, Peregrine Fund library,
generously helped in obtaining obscure literature. For their stimulating discussions on extinct macaws, we
thank Osvaldo Jimenez, Gabinete de Arqueologia de la Oficina del Historiador de Ciudad de La Habana,
Cuba; Catherine Levy, Windsor Research Centre, Jamaica; and William Suarez, Museo Nacional de Historia

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Natural de Cuba, La Habana. Frank Steinheimer provided some additional references. We thank Errol Fuller

and Julian Hume for their comments on the submitted version of the manuscript.

References:

de Acosta, J. 1590. Historic natural y moral de las Indias. Casa de Iuan de Leon, Sevilla.

Adams, M. P., Cooper, J. H. & Collar, N. J. 2003. Extinct and endangered ('E&E') birds: a proposed list for
collection catalogues. Bull. Brit. Orn. Cl. 123A: 338-354.

Albin, E. 1738a. A natural history of birds, vol. 2. Royal Society, London.

Albin, E. 1738b. A natural history of birds, vol. 1. Royal Society, London.

Albin, E. 1740. A supplement to the natural history of birds, vol. 3. Royal Society, London.

American Ornithologists' Union (AOU). 2012. Check-list of North American birds, www.aou.org/
committees/nacc/ (accessed 10 December 2012).

Anon. 1910a. Parrot question is a serious one. Isle Pines News 2(34): 1.

Anon. 1910b. 2,500 parrots for the United States. Isle Pines News 2: 1.

Anon. 1910c. Parrots may be exported. Isle Pines Nezvs 2: 1.

Anon. 1914. Parrot crop ahead of all anticipation. Isle Pines News 6(33): 1.

Arber, E. (ed.) 1885. The first three English books on America. [? 1511J-1555 A.D. Being chiefly translations,
compilations, &c., by Richard Eden, from the writings, maps, &c. of Pietro Martire, of Anghiera (1455-1526),
Apostolical Protonotary, and Counciller to the Emperor Charles V.; Sebastian Munster, the Cosmographer
(1489-1552), Professor of Hebrew, &c., at the University of Basle; Sebastian Cabot, of Bristol (1474-1557), Grand
Pilot of England: with extracts, &c., from the works of other Spanish, Italian, and German writers of the time.
Turnbull & Spears, Edinburgh & Birmingham.

de Armas, J. I. 1888. La zoologia de Colon y de los primeros exploradores de America. Establecimiento Tipografico,
La Habana.

Arredondo, O. 1984. Sinopsis de las aves halladas en depositos fosiliferos pleisto-holocenicos de Cuba. Rep.
Invest. Inst. Zool. 17: 1-35.

Atwood, T. 1791. The history of the island of Dominica. Frank Cass & Co., London.

Balat, F. & Gonzalez, H. 1982. Concrete data on the breeding of Cuban birds. Acta Sci. Nat. Acad. Sci.
Bohemoslovacae Brno 16(8): 1-46.

Bangs, O. & Zappey, W. R. 1905. Birds of the Isle of Pines. Amer. Natur. 39: 179-215.

Barbour, T. 1923. The birds of Cuba. Mem. Nuttall Orn. Cl. No. 6, Cambridge, MA.

Barbour, T. 1943. Cuban ornithology . Mem. Nuttall Orn. Cl. No. 9. Cambridge, MA.

Barbour, T. 1945. A naturalist in Cuba. Little, Brown & Co., Boston.

Bechstein, J. M. 1811. [Description of Ara tricolor], Johann Lathams allgemeine Ubersicht der Vogel 4: 64.

Boehrer, B. T. 2004. Parrot culture. Our 2,500-year-long fascination with the world's most talkative bird. Univ. of
Pennsylvania Press, Philadelphia.

Boeseman, M. 1970. The vicissitudes and dispersal of Albertus Seba's zoological specimens. Zool. Meded. 44:
177-206.

Bolle, C. 1856. Verzeichniss lebender Vogel der zoologischen Garten in London. J. Orn. 4: 163-171.

Bond, J. 1940. Check-list of birds of the West Indies. Acad. Nat. Sci., Philadelphia.

Bond, J. 1959. Fourth supplement to the Check-list of birds of the West Indies (1956). Acad. Nat. Sci., Philadelphia.

Bond, J. 1978. Twenty-second supplement to the Check-list of birds of the West Indies (1956). Acad. Nat. Sci.,
Philadelphia.

Borhidi, A. 1991. Phytogeography and vegetation ecology of Cuba. Akademiai Kiado, Budapest.

Bouton, J. 1640. Relation de I'establissement des Franqois depuis Pan 1635. En Pisle de la Martinique, Pune des
Antilles de PAmerique. Des moeurs des Sauuages, de la situation, & des autres singularitez de Lisle. Chez
Sebastien Cramoisy, Paris.

Brehm, C. L. 1842. Monographic der papageien oder vollstandige Naturgeschichte aller bis jetzt bekannten Pap'ageien
mit getreuen und ausgemalten Abbildungen. Jena August Schmid, Paris.

Brisson, M.-J. 1760. Ornithologie, vol. 4. Ad Ripam Augustinorum, Paris.

Browne, P. 1756. The civil and natural history of Jamaica. B. White & Son, London.

Buffon, G. L. L. 1765-83. Histoire naturelle des oiseaux, avec la description du Cabinet du Roi. 10 vols. LTmprimerie
Royale, Paris.

Buffon, G. L. L. 1774 [1775]. Histoire naturelle des oiseaux, vol. 6. LTmprimerie Royale, Paris.

Buffon, G. L. L. 1793. The natural history of birds, vol. 6. A. Straham & T. Cadell, London.

Buffon, G. L. L. 1847. Obras completas de Buffon, con las clasificaciones comparadas de Cuvier, y la continuacion hasta
el dia, de Mr. Lesson, miembro del Instituto de Francia. Tradicoda al Castellano de la ultima edicion francesa, vol.
10(3). Mellado, Madrid.

By a physician. 1844. Notes on Cuba, containing an account of its discovery and early history; a description of the
face of the country, its population, resources, and weather; its institutions, and the manners and customs of its
inhabitants. With directions to travellers visiting the island. James Munroe & Co., Boston.

Clark, A. H. 1905a. The Lesser Antillean macaws. Auk 22: 266-273.

Clark, A. H. 1905b. The Greater Antillean macaws. Auk 22: 345-348.,

James W. Wiley & Guy M. Kirwan

153

Bull. B.O.C. 2013 133(2)

Clark, A. H. 1908. The macaw of Dominica. Auk 25: 309-311.

Clark, A. H. 1934. Note on the Guadeloupe Macaw (Ara guadeloupensis). Auk 51: 377.

Collis, J. S. 1977. Christopher Columbus. Stein & Day, New York.

Coke, T. 1808. A history of the West Indies, containing the natural, civil, and ecclesiastical history of each island:
with an account of the missions instituted in those islands, from the commencement of their civilization; but more
especially of the missions which have been established in that archipelago by the society late in connexion with the
Rev. John Wesley, vol. 1. Nuttall, Fisher & Dixon, Liverpool.

Cooper, J. H. & Armitage, P. L. 2013. A parrot of the Caribbean? A remarkable find from a 17th-century
Spanish shipwreck. Bull. Brit. Orn. Cl. 133: 52-58.

Cory, C. B. 1886. The birds of the West Indies, including the Bahama Islands, the Greater and the Lesser
Antilles, excepting the islands of Tobago and Trinidad. Auk 3: 454-472.

Cory, C. B. 1889. The birds of the West Indies. Estes & Lauriat, Boston.

Cory, C. B. 1892. Catalogue of West Indian birds. Alfred Mudge & Son, Boston.

Crane, M. D. 1981. Arthur Broughton, a late eighteenth century botanist in Bristol and Jamaica. Archiv. Nat.
Hist. 10: 317-330.

Creel, D. & McKusick, C. 1994. Prehistoric macaws and parrots in the Mimbres area. New Mexico. Amer.
Antiquity 59: 510-524.

Cuervo Pineda, N. & Perez Ortiz, T. M. 2009. Primer registro de Genoprotolichus eurycnemis (Acari:
Pterolichidae) para Cuba con la description de sus estadios inmaduros. Poeyana 497: 9-13.

Cuervo Pineda, N. & Perez Ortiz, T. M. 2010. Nueva especie del genero Distigmesikya Atyeo, Gaud & Perez,
1984 (Acari, Pterolichidae), del Guacamayo Cubano, Ara tricolor (Aves, Psittacidae). Poeyana 499: 13-18.

Dathe, W. & Gonzalez Lopez, R. M. 2002. Johann Christoph Gundlach (1810-1896). Un naturalista en Cuba /
Naturforscher aufKuba. Basilisken Presse, Marburg an der Lahn.

Davidson, M. H. 1997. Columbus then and now: a life reexamined. Univ. of Oklahoma Press, Norman.

Day, D. 1981. Vanished species. Gallery Books, New York.

Diaz del Castillo, B. 1956. The discovery and conquest of Mexico 1517-1521. Farrar, Straus & Cudahy, New York.

D'Orbigny, A. D. 1839. Mamiferos y aves [de la Isla de Cuba], In Sagra, R. de la (ed.) Historia fisica politico y
natural de la isla de Cuba. Segunda parte. Historia natural, vol. 3. Arthus Bertrand, Paris.

Dunn, O. & Kelley, J. E. 1989. The diario of Christopher Columbus's first voyage to America 1492-1493. Abstracted
by Fray Bartolome de las Casas. Univ. of Oklahoma Press, Norman.

Edwards, B. 1806. The history, civil and commercial, of the British colonies in the West Indies, vol. 1. James
Humphreys, Philadelphia.

Edwards, G. 1751. Histoire naturelle de divers oiseaux, pt. 4. Royal College of Medicine, London.

Finsch, 0. 1863. Naamlijst der in de Diergaarde levende Papegaaijen, ten dienste der bezoekers van den Turn
ingerigt. Nederlandsch Tijdschrift voor Dierkunde 1: 6-24.

Fisher, C. T. & Warr, F. E. 2003. Museums on paper: library & manuscript resources. Bull. Brit. Orn. Cl. 123A:
136-164.

Fitzinger, L. J. 1853. Versuch einer Geschichte der Menagerien des osterreichisch-kaiserlichen Hofes mit besonderer
Beriicksichtigurig der Menagerie zu Schonbrunn nebst einer Aufzahlung der in denselben Gehaltenen Thiere von
der Altesten bis auf die neueste zeit. Kaiserlich-Koniglichen Hof- und Staatsdruckerei, Wien.

Forshaw, J. M. 1978. Parrots of the world. Second [rev.] edn. Lansdowne Editions, Melbourne.

Fuller, E. 2001. Extinct birds. Rev. edn. Comstock Publishing, Ithaca, NY.

Garcia Montana, F. 1980. Las aves de Cuba: especies endemicas, vol. 1. Ed. Gente Nueva, La Habana.

Garrido, O. H. & Garcia Montana, F. 1975. Catdlogo de las aves de Cuba. Acad. Cienc. Cuba, La Habana.

Gemelli Careri, G. F. 1699. Giro del mondo del dottor d. Gio. Francesco Gemelli Careri, vol. 2. Naples.

Gemelli Careri, G. F. 1700. Giro del Mondo. Parte sesta contenente le cose piit ragguardevoli vedute Nellanvova
Spagna. Naples.

George, W. 1980. Sources and background to discoveries of new animals in the sixteenth and seventeenth
centuries. Hist. Sci. 18: 79-104.

de Germiny, G. 1937. Quelques oiseaux eteints du Museum de Vienne. Oiseau R.F.O. 7: 376-377 .

Gonzalez Alonso, H., Rodriguez Schettino, L., Rodriguez, A., Mancina, C. A. & Ramos Garcia, I. (eds.) 2012.
Libro Rojo de los vertebrados de Cuba. Ed. Academia, La Habana.

Gonzalez Lopez, R. M. 1990. Juan Cristobal Gundlach. Apuntes biograficos. Ed. Academia, La Habana.

Gosse, P. H. 1847. The birds of Jamaica. John Van Voorst, London.

Greenway, J. C. 1967. Extinct and vanishing birds of the world. Second rev. edn. Dover Publications, New York.

Gundlach, J. 1854. [List of birds, reptiles, insects, and terrestrial molluscs from Isla de Pinos]. Pp. 426-428 in
Poey, F. (ed.) Apuntes sobre la fauna de la Isla de Pinos. Memorias sobre la historia natural de la isla de Cuba,
vol. 1. Imprenta de Barcina, La Habana.

Gundlach, J. 1856. Beitrage zur omithologie Cuba's. ]. Orn. 4: 97-112.

Gundlach, J. 1865-66. Revista y catalogo de las aves Cubanas. Pp. 281-302 in Poey, F. (ed.) Repertorio fisico
natural de la isla de Cuba, vol. 1. Imprenta del Gobierno y Capitania General, La Habana.

Gundlach, J. 1873. Catalogo de las aves Cubanas. An. Soc. Espahola Hist. Nat., Madrid 2: 81-191.

James W. Wiley & Guy M. Kirwan

154

Bull. B.O.C. 2013 133(2)

Gundlach, J. 1874. Neue Beitrage zur Ornithologie Cubas. Nach eigenen 30 jahrigen Beobachtungen
zusammengestellt. J. Orn. 22: 113-166.

Gundlach, J. 1876. Contribution a la ornitologia cubana. Imprenta 'La Antilla', La Habana.

Gundlach, J. 1893. Ornithologia cubana 6 catdlogo descriptivo de todas las especies de aves tanto indigenas como de
paso anual o accidental observadas en 53 anos. Ed. Imprenta 'La Moderna', La Habana.

Hargrave, L. L. 1970. Mexican macaws. Comparative osteology and survey of remains from the southwest.
Anthrop. Pap. Univ. Arizona 20: 1-67.

Heaton, H. C. (ed.) 1934. The discovery of the Amazon according to the account of Friar Gaspar de Carvajal and other
documents. Amer. Geogr. Soc., New York.

Hume J. P. 2007. Reappraisal of the parrots (Aves: Psittacidae) from the Mascarene Islands, with comments
on their ecology, morphology and affinities. Zootaxa 1513: 1-76.

Hume, J. P. & Walters, M. 2012. Extinct birds. T. & A. D. Poyser, London.

Huttich, J. 1534. Die New Welt, der landschafften unnd Insulen, so bis hie her alien Altbeschrybern unbekant, Jungst
aber von den Portugalesern unnd Hispaniern im Nidergenglichen Meer herfunden. Georgen Ulricher von
Andla, Strasbourg.

Jacquin, J. F. 1784. Beytraege zu Geschichte der Voegel. C. F. Wappler, Wien.

Jouanin, C. 1962. Inventaire des oiseaux eteints ou en voie d' extinction conserves au Museum de Paris. Terre
et Vie 109: 257-301.

Kirkconnell, A. & Wiley, J. W. In prep. Zapata Peninsula: important breeding sites for Cuban endemic birds
are endangered!

Knox, A. G. & Walters, M. P. 1994. Extinct and endangered birds in the collection of the Natural History Museum.
Brit. Orn. Cl. Occ. Publ. No. 1. Brit. Orn. CL, Tring.

Krieger, H. W. 1929. Archeological and historical investigations in Samana, Dominican Republic. US Natl.
Mus. Bull. 147: 1-90.

Labat, J. B. 1742. Nouveau voyage aux Isles de I'Amerique, contenant I'histoire naturelle de ces pays, Torigine, les
moeurs, la religion & le gouvernement des habitans anciens & modernes, vol. 2. Paris.

Lack, D. 1976. Island biology, illustrated by the land birds of Jamaica. Univ. of California, Berkeley.

de La Harpe, J.-F. 1759. Abrege de I'histoire generate des voyages. Paris.

Lantermann, W. 1984. Aras. Freileben, Haltung gefangenschafts-vermehrung Verhalten, Arten. Enzyklopddie der
Papageien und Sittiche. Horst Miiller-Verlag, Walsrode.

Las Casas, B. de. 1552-61. Historia de las Indias. Madrid.

Las Casas, B. de. 1876. Historia de las Indias, vol. 5. Miguel Ginesta, Madrid.

Leiva Sanchez, A. T. 1999. Las palmas en Cuba. Ed. Cientifico-Tecnica, La Habana.

Levaillant, F. 1801. Histoire naturelle des perroquets, vol. 1. Levrault, Schoell & Cie, Paris.

Leyva, A. 1922. Museo. Imprenta 'Arroyo Hermanos,' Estrada Palma, baja, numero 13, Santiago de Cuba. [Caja 25,
no. 58: Archivo Nacional]

Long, E. 1774. The history of Jamaica or, a general survey of the ancient and modern state of the island. 3 vols. T.
Lowndes, London.

Lonnberg, E. 1928. Kubas Arapapegoja, Ara tricolor Bechst. Fauna och Flora (Stockholm) 1: 203-208.

Lopez Yustos, A. 1991. Historia de la education en Puerto Rico. Publicaciones Puertorriquenas, San Juan.

Lunan, J. 1814. Hortus Jamaicensis, or a botanical description, vol. 1. St. Jago de la Vega Gazette, Jamaica.

Luther, D. 1995. Die ausgestorbenen Vogel der Welt 4: unverdnderte Auflage. Die Neue Brehm-Biicherei 424.
Westarp- W issenschaften, Magdeburg.

MacNutt, F. A. 1912. De orbe novo, the eight Decades of Peter Martyr d'Anghera, vol. 1. G. P. Putnam's Sons,
New York.

March, W. T. 1863. Notes on the birds of Jamaica. II. Proc. Acad. Nat. Sci. Philadelphia 15: 283-304.

Margoesy, D. 2010. 'Refer to folio and number': encyclopedias, the exchange of curiosities and practices of
identification before Linnaeus. J. Hist. Ideas 71: 63-89.

Mertens, R. & Steinbacher, J. 1955. Die im Senckenberg-Museum vorhandenen Arten ausgestorbener,
aussterbender oder seltener Vogel. Senckenbergiana Biol. 36: 241-265.

Mey, E. 2005. Psittacobrosus bechsteini : eine neuer ausgestorbener Federling (Insecta, Phthiraptera, Amblycera)
vom Dreifarbenara Ara tricolor (Psittaciiformes), nebst einer annotierten Ubersicht liber fossile un rezent
ausgestorbene Tierlause. Anz. Verh. Thiiring. Orn. 5: 201-217.

Minnis, P. E., Whalen, M. E., Kelley, J. H. & Stewart, J. D. 1993. Prehistoric macaw breeding in the North
American Southwest. Amer. Antiquity 58: 270-276.

Moreno, A. 1992. Uber den am Ende des 19. Jahrhunderts ausgestorbenen Kuba-Ara (Ara cubensis). Bongo
20: 65-68.

Morison, S. E. 1970. Admiral of the ocean sea. A life of Christopher Columbus. MJF Books, New York.

Navarrete, M. F. de. 1828. Relations des quatre voyages entrepris par Christophe Colomb pour la decouverte du
Nouveau-monde de 1492 d 1504. Treuttel & Wiirtz, Paris.

Noegel, R. P. 1979. The Isle of Pines Amazon Amazona leucocephala palmarum. Avicult. Mag. 85: 85-88.

Olson, S. L. 1978. A paleontological perspective of West Indian birds and mammals. Pp. 99-117 in Gill, F. B.
(ed.) Zoogeography in the Caribbean. Spec. Publ. No. 13. Acad. Nat. Sci., Philadelphia.

James W. Wiley & Guy M. Kirwan

155

Bull. B.O.C. 2013 133(2)

Olson, S. L. 1982. Biological archeology in the West Indies. Fla. Anthropologist 35: 162-168.

Olson, S. L. 2005. Refutation of the historical evidence for a Hispaniolan macaw (Aves: Psittacidae: Am).
Carib. J. Sci. 41: 319-323.

Olson, S. L. & Maiz Lopez, E. j. 2008. New evidence of Am autochthones from an archeological site in Puerto
Rico: a valid species of West Indian macaw of unknown geographical origin (Aves: Psittacidae). Carib.
J. Sci. 44: 215-222.

Olson, S. L. & Suarez, W. 2008. A fossil cranium of the Cuban Macaw Am tricolor (Aves: Psittacidae) from
Villa Clara Province, Cuba. Carib. J. Sci. 44: 287-290.

Osburn, W. 1859. Notes on the bats and birds of Jamaica. Zoologist 17: 6587-6594.

Oviedo y Valdes, G. F. 1535. La historia general de las Indias. Juam Cromberger, Sevilla.

Oviedo y Valdes, G. F. 1851. Historia general y natural de las Indias, islas y tierra-firme del mar oceano. Real Acad.
Hist., Madrid.

Oviedo y Valdes. G. F. 1950. Sumario de la natural historia de las Indias. Fondo de Cultura Economica, Mexico.
Oviedo, G. F. 1959. Natural history of the West Indies. Univ. of North Carolina Press, Chapel Hill.

Perez de la Riva, J. & Berthe, J. P. 1971. La Habana de fines del siglo XVII vista por un italiano. Rev. Biblioteca
Nac. Jose Marti 62: 63-85.

Perpina, A. 1889. El Camagiiey. Viajes pintorescos por el interior de Cuba y por sus costas con descripciones del pais.
J. A. Bastinos, L. Niubo, Barcelona.

Poey, F. (ed.) 1866. Repertorio fisico natural de la isla de Cuba, vol. 1. Imprenta del Gobierno y Capitania General,
La Habana.

de las Pozas, G. & Gonzalez Alonso, H. J. 1984. Disminucion de los sitios de nidificacion de Cotorra y Catey
(Aves: Psittacidae) por la tala de palmas en Cienaga de Zapata, Cuba. Misc. Zool., Inst. Zool. Acad. Cienc.
Cuba 18: 4.

Prestwich, A. A. 1970. Extinct, vanishing, and hypothetical parrots. Avicult. Mag. 76: 198-204.

Ragues, P. V. 1914. Museo Cubano 'Gundlach' catalogo general. Imprenta 'Cuba Intelectual', La Habana.
Ramsden, C. T. 1918. Vida y exploraciones zoologicas del Dr. Juan Gundlach en Cuba (1839-1896). Soc. Ed. Cuba
Contemporanea, La Habana.

Robbins, L. E. 2002. Elephant slaves and pampered parrots: exotic animals in eighteenth-century Paris. Johns
Hopkins Univ., Baltimore.

[de Rochefort, C.] 1658. Histoire naturelle et morale des iles Antilles de V Amerique: enrichie des plusieurs belles
figures des raretez les plus considerables qui y sont d'ecrites: avec un vocabulaire Cara'tbe. Rotterdam.
Rothschild, W. 1905. [Extinct parrots of the West Indies]. Bull. Brit. Orn. Cl. 16: 13-15.

Rothschild, W. 1907a. Extinct birds. Hutchinson & Co., London.

Rothschild, W. 1907b. On extinct and vanishing birds. Pp. 191-217 in Hartert, E., Bonhote, J. O. & Lewis, J.

(eds.) Proc. Fourth Intern. Orn. Congr., London. Dulan & Co., London.

Rouse, 1. 1992. The Tainos. Rise and decline of the people who greeted Columbus. Yale Univ. Press, New Haven, CT.
Salvadori, T. 1906. Notes on the parrots. (Part V). Ibis (8)6: 451-465.

Sloane, H. 1725. A voyage to the islands Madera, Barbadoes, Nieves, St. Christophers, and Jamaica; with the natural
history of the herbs and trees, four-footed beasts, fishes, birds, insects, reptiles, &c. of the last of those islands, vol.
2. London.

Smith, H. L. 1937. The cage birds of Cuba. Avicult. 37(2-3): 29-32.

Snyder, N. F. R., Wiley, J. W. & Kepler, C. B. 1987. The parrots ofLuquillo: natural history and conservation of the
Puerto Rican Parrot. West. Found. Vert. Zool., Los Angeles.

Souance, C. 1856. Catalogue des perroquets de la collection du prince Massena d'Essling, due de Rivoli, et
observations sur quelques especes nouvelles ou peu connues de psittacides. Rev. Mag. Zool. (2)8: 56-64,
152-158, 208-226.

Stewart, J. 1822. A view of the past and present state of the island of Jamaica. Oliver & Boyd, Edinburgh.
Stresemann, E. 1954. Ausgestorbene und aussterbende Vogelarten vertreten im Zoologischen Museum zu
Berlin. Mitteilungen aus dem Museum fur Naturkunde in Berlin. Zool. Mus. u. Inst. Spez. Zool. (Berlin)
30: 38-53.

Taylor, J. 2008. Jamaica in 1687: the Taylor manuscript at the National Library of Jamaica. Univ. of West Indies
Press, Mona.

du Tertre, J.-B. 1654. Historic generate, des isles des Christophe, de la Guadeloupe, de la Martinique, et autres dans
V Amerique. J. & E. Langlois, Paris.

du Tertre, J.-B. 1667. Histoire generate des Antilles habitees par les Frangois, vol. 2. Paris.

Thevet, A. 1971 [1568]. The new found world, or Antarcticke. Da Capo Press, Amsterdam.

Thomas, K. 1983. Man and the natural world: changing attitudes in England 1500-1800. Oxford Univ. Press.
Todd, W. E. C. 1916. The birds of the Isle of Pines. Incorporating the substance of field-notes by Gustav A.
Link. Ann. Carnegie Mus. 10: 146-296.

de la Torre, J. M. 1857. Lo que fuimos y lo que somos o La Habana antigua y moderna. Imprenta de Spencer y
Compania, La Habana.

Turvey, S. T. 2010. A new historical record of macaws on Jamaica. Archiv. Nat. Hist. 37: 348-351.

Vincent, J. 1966-1971. Red data book. 2. Aves. IUCN, Morges.

James W. Wiley & Guy M. Kirwan

156

Bull. B.O.C. 2013 133(2)

Waldseemiiller, M. 1907. The Cosmographiae introductio of Martin Waldseemiiller in facsimile followed by the four
voyages of Amerigo Vespucci, with their translation into English. US Catholic Historical Soc., New York.

Walters, M. 1995. On the status of Ara tricolor Bechstein. Bull. Brit. Orn. Cl. 115: 168-170.

Wetherbee, D. K. 1985. The extinct Cuban and Hispaniolan macaws (Ara, Psittacidae), and description of a
new species, Ara cubensis. Carib. j. Sci. 21: 169-175.

Wetmore, A. 1928. Bones of birds from the Ciego Montero deposit of Cuba. Amer. Mus. Novit. 301: 1-5.

Wetmore, A. 1937. Ancient records of birds from the island of St. Croix with observations on extinct and
living birds of Puerto Rico. /. Agric., Univ. Puerto Rico 21: 5-15.

Wiley, J. W. 1991. Status and conservation of parrots and parakeets in the Greater Antilles, Bahama Islands,
and Cayman Islands. Bird Conserv. Intern. 1: 187-214.

Wiley, J. W. & Wunderle, J. M. 1993. The effects of hurricanes on birds, with special reference to Caribbean
islands. Bird Conserv. Intern. 3: 319-349.

Wiley, J. W., Gnam, R. S., Koenig, S. E., Dornelly, A., Galvez, X., Bradley, P. E., White, T., Zamore, M., Reillo,
P. R. & Anthony, D. 2004. Status and conservation of the family Psittacidae in the West Indies. /. Carib.
Orn. 17: 94-154.

Williams, M. 1. & Steadman, D. W. 2001. The historic and prehistoric distribution of parrots (Psittacidae) in
the West Indies. Pp. 175-198 in Woods, C. A. & Sergile, F. E. (eds.) Biogeography of the West Indies. Second
edn. CRC Press, Boca Raton, FL.

Wilson, S. M. 1990. Hispaniola: Caribbean chiefdoms in the age of Columbus. Univ. of Alabama Press, Tuscaloosa.

Wing, E. S. 1989. Human exploitation of animal resources in the Caribbean. Pp. 137-152 in Woods, C. A. (ed.)
Biogeography of the West Indies, past, present and future. Sandhill Crane Press, Gainesville, FL.

Woods, C. A. 1989. Biogeography of the West Indian rodents. Pp. 741-798 in Woods, C. A. (ed.) Biogeography
of the West Indies, past, present and future. Sandhill Crane Press, Gainesville, FL.

de Ybarra, A. M. F. 1907. The letter of Dr. Diego Alvarez Chanca, dated 1494, relating to the second voyage
of Columbus to America (being the first written document on the natural history, ethnography, and
ethnology of America). Smithsonian Misc. Coll. 48: 428-457.

Addresses: James W. Wiley, P.O. Box 64, Marion Station, MD 21838, USA. Guy M. Kirwan, Research Associate,
Field Museum of Natural History, 1400 South Lakeshore Drive, Chicago, IL 60605, USA.

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Extent of overlap between two Common Fiscal
Lanius collaris subspecies in Lesotho

by Grzegorz Kopij

Received 11 January 2013

Common Fiscal Lanius collaris Linnaeus, 1766, is widespread throughout the Afrotropics,
and is generally considered to be represented by up to ten subspecies (Harris & Franklin
2000). In southern Africa, four subspecies are currently accepted: L. c. collaris, L. c.
subcoronatus A. Smith, 1841, L. c. capelli (Bocage, 1879) and L. c. aridicolus Clancey, 1955
(Dean 2005). The entire region encompassed by the Republic of South Africa, Lesotho and
Swaziland is inhabited by the nominate and L. c. subcoronatus (Dean 2005). However, the
boundaries between these two subspecies in southern African are not so clearly delimited
as the maps in Hockey et al. (2005) suggest. In fact, L. c. collaris and L. c. subcoronatus appear
to be almost randomly scattered across the region. The latter subspecies differs from the
nominate one in its distinct white supercilium, while the rump and uppertail-coverts are
greyer (less white) in the nominate. The two subspecies are easily separated in the field,
making it possible to precisely delimit their geographical ranges.

Maclean & Maclean (1976) found evidence of a clinal change in the percentage of the
races L. c. subcoronatus and L. c. collaris: between Winburg (28°31'S, 27°01'E) and Brandfort,
Free State (28°42'S, 26°28'E) 100% of examined birds were L. c. collaris (n = 5), Brandfort-
Kimberley 35% : 65% (n = 20), Kimberley-Kuruman 44% : 56% (n = 48), Kuruman-Van
Zylsrus 77% : 23% (n = 9) and between Van Zylsrus and Askham, Northern Cape (26°12'S,
18°35'E) 100% were L. c. subcoronatus (n = 2). In the most recently published maps of the
distribution of these subspecies, the nominate is shown as occurring throughout Lesotho,
except for a narrow strip in the northern lowlands, where the nominate and L. c. subcoronatus
co-occur (Hockey et al. 2005, Chittenden et al. 2012).

Lesotho is an enclave within the Republic of South Africa. Three-quarters of the country,
the highlands (or Maloti) lie above 2,200 m. Lowlands are those areas below 1,700 m while
the foothills are between 1,700 and 2,200 m. Lowlands (total surface area 6,051 km2) form a
strip of land lying along the enclave's north-east to south-west frontier and extending east
to the Cave Sandstone Foothills. The foothills (2,964 km2) form a narrow band bordering
the highlands to the east and lowlands to the west, constituting an intermediate region.
The Senqu Valley lies below 1,800 m and occupies 3,398 km2 (Ambrose et al. 2000). Natural
vegetation in the lowlands is Highveld Grassland, while in the foothills and highlands it is
Afromontane Grassland (Acocks 1988).

During the years 1998-2002, which I spent in Lesotho studying birds (Kopij 2011),
I attempted to distinguish the two subspecies of L. collaris. As shown in Table 1, both
subspecies occur together throughout Lesotho. The extent of this overlap differs regionally
and by altitude. Clinal change is evident in the lowlands: the proportion of collaris to
subcoronatus was 1 : 0 in the north, but 0.5 : 0.5 in the centre and 0.6 : 0.4 in the south.
Both in the foothills and Senqu Valley, the proportion was 0.3 : 0.7. In the highlands, it
was everywhere 0.4 : 0.6, without any apparent regional (clinal) change. Soobramoney et
al. (2005) found that patterns of variation in morphology and anatomy were significantly
correlated with 11 climatic trends along an alititudinal gradient. Common Fiscals were
largest in cooler, less humid, more arid areas, and smallest in warmer, wetter, more humid
localities, in accordance with Bergmann's Rule.

Grzegorz Kopij

158

Bull. B.O.C. 2013 133(2)

TABLE 1

Extent of overlap between two Common Fiscal Lanins collaris subspecies in Lesotho during the years

1998-2002.

Site

Coordinates

Number of birds

LOWLANDS

subcoronatus

(29)

collaris

(35)

Northern (north of Maputsoe)

(0)

(10)

Leribe Plateau

28°07'S, 29°05'E

Central

(12)

(13)

Berea

29°17'S, 27°38'E

Ha Baroana: Lipiring Gorge

29°21'S, 27°47'E

Thaba-Bosiu

29°22'S, 27°37'E

St. Michael

29°26'S, 27°40'E

Maseru

29°20'S, 27°12'E

Southern (south of Maseru)

(17)

(12)

Morija

29°38'S, 27°31'E

Matsieng

29°37'S, 27°35'E

Mantsebo: Qeme Plateau

29°29'S, 27°31'E

FOOTHILLS (central)

(16)

(35)

Nyakosoba

29°31'S, 27°46'E

Moitssupeli

29°34'S, 27°45'E

Popanyana

29°31'S, 27°44'E

Thaba Telle

29°45'S, 27°40'E

Dikolobeng

29°29'S, 27°55'E

SENQUE VALLEY

(5)

(15)

Masitise-Villa Maria

30°24'S, 27°38'E

Quthing

30°25'S, 27°43'E

Qacha's Nek

30°07'S, 28°42'E

HIGHLANDS

(21)

(29)

Northern

(12)

(15)

Mokhotlong

29°18'S, 29°03'E

Mapholaneng

29°12'S, 28°52'E

Central

(7)

(11)

St. Theresa

29°37'S, 28°48'E

Thaba Tseka

29°31'S, 28°36'E

Marakabei

29°33'S, 28°07'E

Southern

(2)

(3)

Semonkong

29°51'S, 28°03'E

3 .

TOTALS

114

Overall in Lesotho, the nominate was almost twice as common as L. c. subcoronatus. The
extent of overlap varied spatially and probably also temporally. Although both subspecies
are common and widespread in Lesotho, hybridisation is apparently rather rare. Only
four (2.3%) intermediate individuals (the white supercilium being less distinct, shorter
and narrower, and the rump dark grey) were recorded during my work (at Matsebo: 28
September 1999, Matsekua: 13 May 2000, Morija: 3 July 2001, and Matsieng: 3 July 2001 ), but
others could have been missed. In the Roma Valley, where the species was closely studied
during the breeding season, in 1999-2002 26-36 pairs bred on the 82-ha National University
of Lesotho campus, with c.10% of 31 pairs in the 2000/2001 breeding season mixed, while in
all others both sexes were of the nominate subspecies (Kopij 2002, 2004, 2006). Because there

Grzegorz Kopij

159

Bull. B.O.C. 2013 133(2)

Figure 1. Numbers of two subspecies of Common Fiscal Lanius collaris recorded in Lesotho: subcoronatus
(first) and collaris (second).

is only slight sexual dimorphism in Common Fiscal (Dean 2005), no attempt was made to
detect any sexual bias towards one subspecies.

A low level of interbreeding between L. c. collaris and L. c. subcoronatus recorded
in Lesotho perhaps indicates that some L. c. subcoronatus migrate from the highlands to
the lowlands in the dry season, i.e. the non-breeding season, which possibility was also
suggested by Clancey (1980). However, in this study most records were made in the wet
(breeding) season. The two forms, therefore, occur in Lesotho sympatrically. Furthermore,
at least in the Roma Valley, both the level of hybridisation is low and the number of hybrids
even lower. It is, therefore, plausible that the two forms might represent separate species
within a superspecies. This premise merits further investigation.

Acknowledgements

Prof. Coleen Downs and Bob Dowsett are gratefully acknowledged for their suggestions, corrections and
comments on an earlier draft of this note.

References:

Acocks, J. P. H. 1988. Veld types of South Africa. Mem. Bot. Sur. S. Afr. 57: 1-146.

Ambrose, D., Talukdar, S. & Pomela, E. M. 2000. Biological diversity in Lesotho: a country study. National
Environment Secretariat, Maseru.

Chittenden, H., Allan, D. & Weiersbye, I. 2012. Roberts' geographic variation of southern African birds. John
Voelcker Bird Book Fund, Cape Town.

Clancey, P. A. 1976. Intergradation between two subspecies of the Fiscal Shrike. Ostrich 47: 145.

Clancey, P. A. (ed.) 1980. S.A.O.S. checklist of southern African birds. S. Afr. Orn. Soc., Houghton.

Dean, W. R. J. 2005. Common Fiscal Lanius collaris. Pp. 752-753 in Hockey, P. A. R., Dean, W. R. J. & Ryan, P.

G. (eds.) Roberts' birds of southern Africa. John Voelcker Bird Book Fund, Cape Town.

Harris, T. & Franklin, K. 2000. Shrikes and bush-shrikes. Christopher Helm, London.

Grzegorz Kopij

160

Bull. B.O.C. 2013 133(2)

Hockey P. A. R., Dean, W. R. J. & Ryan, P. G. (eds.) Roberts' birds of southern Africa. John Voelcker Bird Book
Fund, Cape Town.

Kopij G. 2002. Temporal variation in territoriality of the fiscal shrike. Vogelwelt 42: 40.

Kopij, G. 2004. Breeding density of Fiscal Shrike (Lanius collaris). Biol. Lett. 41: 77-85.

Kopij, G. 2006. Breeding biology of the Fiscal Shrike, Lanius collaris (Laniidae), in a peri-urban environment
in Roma (Lesotho). Vestnik Zool. 40(6): 3SM15.

Kopij, G. 2011. Avian diversity in ruderal and urbanized habitats in Lesotho. Berkut 20(1/2): 22-28.

Maclean, G. L. & Maclean, C. M. 1976. Extent of overlap in two races of the Fiscal Shrike. Ostrich 47: 66.

Soobramoney, S., Downs, C. T. & Adams, N. J. 2005. Morphological variation in the Common Fiscal Lanius
collaris along an altitudinal gradient in southern Africa. Ostrich 76: 130-141.

Address: Dept, of Wildlife Management, Univ. of Namibia, Katima Mulilo Campus, Private Bag 1096, Winela
Road, Katima Mulilo, Namibia, e-mail: gkopij@unam.na

Clive F. Mann

161

Bull. B.O.C. 2013 133(2)

First record of Oriental Cuckoo Cuculus saturatus optatus

in Africa

by Clive F. Mann

Received 25 January 2013

While examining specimens at the Natural History Museum, Tring, for a monograph
on cuckoos (Erritzoe et al. 2012) I discovered a hepatic female Oriental Cuckoo Cuculus
saturatus optatus specimen (BMNH 1911.12.23.966), collected by Boyd Alexander in Zambia
(near the confluence of the Zambezi and Kafue Rivers, close to the modern border with
Zimbabwe) on 27 December 1898. He did not label it to species, but subsequently it was
labelled C. canorus (Common Cuckoo) and placed with the African Cuckoos C. gularis.
However, he did identify it as canorus in a paper (Alexander 1900) that was subsequently
mentioned by Benson (1956).

The most noticeable features are the heavy dark barring on the deep rufous-chestnut
rump and upper tail-coverts, with no pale feather tips, and the broader dark barring on the
ventral surface. Hepatic female Common Cuckoo has little or no barring on the rump and
uppertail-coverts. Hepatic juveniles of the latter are barred, but each feather has a narrow
whitish tip, and the rufous-chestnut is generally paler. Hepatic juvenile African Cuckoo is
much less rich, often with some grey admixed. The ventral barring of the Alexander skin is
broader than that of Common or African Cuckoos. The two subspecies of Oriental Cuckoo
(nominate and optatus) are extremely similar (Figs. 1-2), being reliably separated only by
measurements or perhaps vocalisations. The wing length of Alexander's specimen (224
mm) places it in optatus rather than the nominate (Table 1).

Madagascar Cuckoo C. rochii and Lesser Cuckoo C. poliocephalus can be eliminated
on size. Red-chested Cuckoo C. solitarius, Black Cuckoo C. clamosus, African Cuckoo C.
gularis, and the two subspecies of Common Cuckoo, C. canorus bakeri and C. c. bangsi, can
be eliminated by plumage.

Characters that indicate optatus. — The rump of the Alexander skin lacks pale tips to
the dark feathers. Hepatic adult female Common Cuckoo has almost completely unbarred
rump / uppertail-coverts; hepatic juvenile has dark bars, but the feathers have small whitish
or grey tips. Both subspecies of Oriental Cuckoo (nominate and optatus) lack these tips. The
ratio of the width of dark to pale bars on the ventral surface of Alexander's specimen is
0.751 (versus C. c. canorus 0.321-0.573, n = 30; C. c. subtelephonus 0.316, 0.355, 0.465, n = 3; C.
c. bakeri 0.483, 0.654, n = 2; C. s. optatus 0.605-1.21, n = 16). Five dark and five pale bars were
measured in the mid-chest area, and the means calculated, which were then used to calculate
the ratios (from hepatic female / juvenile specimens at the Natural History Museum, Tring).
Furthermore, the Alexander specimen matches the wing formula for saturatus given by
Payne (2005). However, the wing formulae of canorus and saturatus are very similar, and I
found them to be unreliable for separating the two species over long series.

Distribution (from Erritzoe et al. 2012, and references therein). — Oriental Cuckoo C. s.
saturatus Blyth, 1843. Southern Himalayas, north and east Burma, Thailand to China south
of 32°N, including Taiwan and Hainan. Winters in South and South-East Asia, Greater
Sundas, Philippines and western New Guinea. Some vagrancy. C. s. optatus Gould, 1845.
European Russia east of 45°E to Pacific coast, south to Kazakhstan and Mongolia to north of
32°N in China. Winters and migrates through South-East Asia, Sundas and Philippines east
to south-west Pacific and Australia. Much vagrancy, with two records in Israel, in August

Figures 1-2. Dorsal and ventral views of, left to right, Alexander's specimen (BMNH 1911.12.23.966); Oriental
Cuckoo Cuculus saturatus optatus ; C. s. saturatus ; Common Cuckoo C. canorus canorus ; C. c. subtelephonus ; and
African Cuckoo C. gularis (Harry Taylor © Natural History Museum)

Clive F. Mann

163

Bull. B.O.C. 2013 133(2)

TABLE 1

Mensural data (wing, tail and bill) from Cuculus specimens at the Natural History Museum, Tring,
including Alexander's specimen (BMNH 1911.12.23.966) collected close to the modern-day Zambia /

Zimbabwe border.

wing length (mm)
(max. chord)

range

mean

n =

Alexander's specimen

224

Cuculus saturatus saturatus
male

174-194

183.4

±5.8

C. s. saturatus female

163-190

175.4

±8.2

C. s. optatus male

214-223

218.4

±1.9

C. s. optatus female

213-222

216.9

±2.3

C. canorus canorus male

213-230

221

±4.3

C. c. canorus female

204-216

210

±3.7

C. c. subtelephonus male

201-249

221.9

±15.2

C. c. subtelephonus female

180-223

200.5

±13.1

C. c. bakeri male

198-232

214.3

±9.9

C. c. bakeri female
(no hepatic morph)

195-215

205.1

±6.8

C. c. bangsi male

C. c. bangsi female

199-215

206.7

±6.2

(no hepatic morph)

194-207

200.6

±5.4

C. gularis male

C. gularis female

204-226

215.3

±8.4

(no hepatic morph)

197-209

205.3

±5.1

Mensural data

tail length (mm)

bill to skull (mm)

range

mean

n =

range

mean

n =

167

22.9

113-152

136.4

±11.7

17.9-22.1

20.5

±1.1

113-152

136.6

±10.6

18.5-23.0

±1.4

128-173

156.7

±11.8

19.7-24.6

±1.7

128-168

142.6

±11.8

17.9-23.7

20.9

±1.7

170-186

177

±4.1

25.5-31.2

27.7

±1.5

158-177

167

± 5.6

25.2-28.6

26.8

±1.0

25.2, 26.2

24.7-27.0

26.2

±1.1

24.5-29.5

26.9

±1.0

25.9-28.2

27.2

±1.2

143-166

152.9

±10.8

20.7-23.6

26.1

±0.5

141-156

147

±5.5

19.0-22.4

20.7

±1.2

1985 (Shirihai 1996) and April 2008 (www.
israbirdig.com/israelbirdsforum), being
the closest to Africa.

Common Cuckoo C. c. canorus
Linnaeus, 1758. Palearctic from Ireland to
Kamchatka and Japan. Winters equatorial
Africa, India and South-East Asia. C.
c. bangsi Oberholser, 1919. Iberia and
Maghreb. Probably winters Africa south
of equator. C. c. subtelephonus Zarudny,
1914. Transcaspia to west and north
China, Iran and Afghanistan. Winters
Middle East, India and Africa south of
Sahara. C. c. bakeri E. Hartert, 1912. North¬
east Indian Subcontinent, Assam, Nepal,
Bhutan, northern South-East Asia, south-
South-East Asia.

TABLE 2

Wing formulae data for relevant Cuculus taxa. The
formulae for saturatus, optatus, canorus and gularis are
from Payne (2005). However, in long series I found
them to be an unreliable discriminant.

Wing formulae

(except Alexander's specimen,

from Payne 2005)

Alexander's specimen

P 8>9>7>6>5>4>1 0>3>2> 1

C. saturatus saturatus

P 8>9>7>6>5>4>10>3>2>1

C. saturatus optatus

P 8>9>7>6>5>4> 1 0>3>2> 1

C. canorus canorus

P 8>7>9>6>5>4>1 0>3>2> 1

C. gularis

P 8>7>9>6>5>4> 1 0>3>2> 1

east Tibet and south China. Winters India and

Clive F. Mann

164

Bull. B.O.C. 2013 133(2)

Conclusion

Although C. s. optatus has an easterly migration route, it is prone to vagrancy and
has been recorded twice in Israel (Shirihai 1996; www.israbirdig.com/israelbirdsforum).
Some Cuculus are notoriously difficult to distinguish in the field unless vocalising. Birds
of this genus are infrequently trapped in mist-nets, and birds that die in Africa are usually
scavenged very quickly. Thus species such as Oriental Cuckoo may occasionally occur
in Africa undetected. The specimen in question, although quite distinct, could easily be
passed off as Common Cuckoo. Field observers would not normally consider identifying a
cuckoo in Africa as Oriental, due to the lack of precedent, but I urge them to consider this
possibility in the future and attempt to capture any Cuculus that does not match the known
local species. Collections may also benefit from closer scrutiny.

Acknowledgements

I thank Robert J. Dowsett for his encouragement and advice, and with Robert A. Cheke, for refereeing this
communication. Gratitude is expressed to Robert Prys-Jones and Mark Adams for access to specimens at the
Natural History Museum, Tring, and to Harry Taylor, also of the Natural History Museum, for providing
the photographs.

References:

Alexander, B. 1900. An ornithological expedition to the Zambezi River. Ibis (7)6: 70-109.

Benson C. W. 1956. A contribution to the ornithology of Northern Rhodesia. Occ. Pap. Nat. Mus. S. Rhod.
21B: 1-51.

Erritzoe, J., Mann, C. F., Brammer, F. P. & Fuller, R. A. 2012. Cuckoos of the world. Christopher Helm, London.
Payne, R. B. 2005. The cuckoos. Oxford Univ. Press.

Shirihai, H. 1996. 77ie birds of Israel. Academic Press, London.

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Registered Charity No. 279583

Bulletin of the British Ornithologists' Club
ISSN 0007-1595
Edited by Guy M. Kirwan
Associate Editor: Frank D. Steinheimer

Volume 133 Number 2, pages 81-164

CONTENTS

Club Announcements . 81

THIBAULT, M., DEFOS DU RAU, P„ PINEAU, O. & PANGIMANGEN, W. New and interesting
records for the Obi archipelago (north Maluku, Indonesia), including field observations and first
description of the vocalisation of Moluccan Woodcock Scolopax rochussenii . 83

JACKSON, H. D. The taxonomic status of the Black-shouldered Nightjar Caprimulgus nigriscapularis

Reichenow, 1893 . 116

WILEY, J. W. & KIRWAN, G. M. The extinct macaws of the West Indies, with special reference to

Cuban Macaw Am tricolor . 125

KOPIJ, G. Extent of overlap between two Common Fiscal Lanius collaris subspecies in Lesotho . 157

MANN, C. F. First record of Oriental Cuckoo Cuculus saturatus optatus in Africa . 161

EDITORIAL BOARD

Murray Bruce, R. T. Chesser, Edward C. Dickinson, Franchise Dowsett-Lemaire, Steven M. S. Gregory, Jose
Fernando Pacheco, Robert B. Payne, Pamela C. Rasmussen, Cees Roselaar, Thomas S. Schulenberg, Lars
Svensson

Registered Charity No. 279583

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Printed on acid-free paper.

Published by the British Ornithologists' Club
Typeset by Alcedo Publishing of Arizona, USA, and printed by Latimer Trend, UK

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