Gathorne Hardy MA PhD PNBS(K) (5 th Earl of Cranbrook) Sarawak Museum (1956 - Yayasan Siswa Lokantara Universiti Malaya Niah cave excavations “Expeditions” (1964 – 92), to Kinabalu, Gn Benom, Vanuatu, Gn Mulu, Belalong (Brunei) Chairman, English Nature (1990-98), Chairman Entrust, Regulator of Environmental bodies under UK Landfill tax regulations (1996)

Two tasks: 1. Sort out swiftlets taxonomy and biology 2. Identify animal remains from Niah excavations.

Diversity peaks World range of swiftlets: Indo-Pacific Mascerenes Eastern Himalayas New Caledonia Hawaii Indo-Malayan region

Swallows share the skies with swiftlets, but are unrelated

Unlike swallows, swiftlets cannot perch. They feed (on aerial arthropods) and drink on the wing. Lim Chan Koon

The world range of swiftlets [nos. of species] (Asia & Indian ocean) [1] [1] [1] [ < 1 > ] [3] [ < [ 8 ] > ] [ 3 ] [ 4 ] [ 7 ] [1] [ 6 ] [1] Migratory

Hot-spot Malaysia 3 genera, 8 species of swiftlets

Waterfall swift (formerly called the “giant” swiftlet Breeding records Sight records

Waterfall swift, Hydrochous gigas

troglodytes esculenta Plain tailed linchi Glossy swiftlets Two species in Malaysia White spots on tail feathers

Speciation among Glossy swiftlets, Collocalia C. esculenta L. 1758 C. linchi H & M 1834 C. troglodytes Gray 1845 All build self-supporting cup-like nests of strands of plant material, fixed with a basal band of edible nest ‘cement’

Collocalia : glossy swiftlets 1. White-bellied swiftlets C. esculenta

Collocalia : glossy swiftlets 1. White-bellied swiftlets C. esculenta cyanoptila

Key character: a single feather tuft on the hind toe (hallux) Collocalia esculenta cyanoptila

Collocalia linchi, Linchi swiftlet : Java, part of Sumatra & Kinabalu, Sabah Green glossed upperparts No tuft on hallux

Black-brown swiftlets Aerodramus Photos: Lim Chan Koon

Speciation among ‘mossy’ nest swiftlets Permanently soft, glutinous salivary nest-cement Lim Chan Koon

A.vanikorensis Q & G 1830 A.salanganus Streubel 1848 * * * = type locality The two allopatric “mossy-nest” swiftlets

There is only one species of swiftlet building the ‘Black’ nests this is Aerodramus maximus Lim Chan Koon

Hevily feathered tarsus = A. maximus Naked (or lightly feathered) tarsus = others Field characters 2.

White nests ( ‘vestitus’) Salai cave, Baram, Sarawak Photo Lim Chan Koon

White nests are highest value These nests are composed almost entirely of edible nest-cement, with no more than a few feathers adhering to or incorporated, Raw nests may be worth RM 7000 per kg of ~100 nests

There are also two species of “white-nest” swiftlets This is the typical dark-rumped form Aerodramus f. fuciphagus of Java

White-nest swiftlets from Sarawak Aerodramus fuciphagus vestitus

White-nest swiftlet from Gomantong cave , Sabah

Topotypes of vestita Lesson from Sumatra

AMNH 634703 f Coll. H.C. Robinson, Koh Pennan (Phangan Is) 13 June 1913. L Wing 112 No moult. Grey rumped, or Germain’s swiftlet Aerodramus germani Coasts and islands from Hainan (China) & Vietnam, Thailand, Malaysia, to Andaman Is Grey rump, with dark shaft streaks

Skins from Satang Kecil (1933)

Young bird from Satang Kecil, Sarawak Turtle Is. Collected in 1957

Two old skins from Selangor (Kelang) 1887 & 1908

Two skins from Juara Bay Tioman island (1907)

P. Condore Type locality Sembilan Is Satang Island Mantanani Is. Berhala Is. Maratua Is. (ssp. perplexus ) Southern and western limits of Grey-rumped swiftlets Aerodramus germani in Malaysia P. Tioman Gomantong Horsborough lighthouse

Two species of white-nest swiftlets Aerodramus germani Aerodramus fuciphagus A. g. germani A. (f.) vestitus ?? A. (g.?) perplexus A. f. vestitus A. f. fuciphagus A. g. hainaensis A. g. inexpectatus A. f. dammermanni A. fuciphagus micans

Topotype of “ Collocalia fuciphaga amechana” Oberholser from Pulau Jimaja, Anamba Is., Indonesia

Glossy swiftlets Collocalia Linchi Black-brown, Aerodramus Giant swiftlet Hydrochous Black nest White nest Mossy nest Bracket vegetable nest Indo-Malayan swiftlets

Six swiftlets of interior Borneo: Comparative wing lengths Scale in mm Chris Stimpson Diagram by C. Stimpson

Karst limestone of interior Borneo Surprisingly, no records of Aerodramus swiftlets in any cave in Msia Semenanjung Caves in limestone and other rocks are the natural nesting sites of swiftlets.

White cliffs Dizzy heights Twisty tunnels The work of nest collectors is very hazardous

A. (f.) vestitus Salai cave, Baram, Sarawak Rarely can nests be gathered by hand Lim Chan Koon

Access to many caves is arduous Bt Kelam, W. Kalimantan Nest fissure

Climbing Mt Kelam Fixed ladders

Walkways in cave interiors are made of natural materials Panguma cave, Niah, Sarawak Black-nest colony

Nuar at Panguma Fixed climbing aids also use available natural materials

Johari Top of the pole Sarawak Museum

Collecting head at the end of the pole (penyulok ) Huge loss of life – eggs and young birds : Kayan river, Kalimantan Timur George Nawan

HOUSE-FARMING THE LATEST DOMESTICATION Intensive management of housed swiftlets began in Java, Indonesia, in the 1970s

In Java, especially, skills developed in transforming buildings naturally colonised by glossy swiftlets into white-nest houses

Egg transfer

Nest management

House in transition C. linchi > A. fuciphagus Glossy swiftlet nests White nests

New nest house at Bayang Kara, Kaltim. Original wooden house with C. esculenta nests enclosed in concrete shell; eggs imported from W. Java Ventilation holes Cladding, for coolness Javanese technology exported

Spontaneous occupation of buildings by edible/white-nest swiftlets Penang & Butterworth Kuala Terengganu Bintulu Melaka Singapore Java (multiple instances) Banjermasin Since 1890

Spontaneous colonisations of buildings in Pen. Malaysia Penang (1947) K. Terengganu (1974) Kuala Lumpur (1948) Known start dates Melaka (1953) Singapore (1935) Taiping (1950s)

Since 1990s swiftlet-farming has relied on the direct attraction of free-flying birds into specially constructed houses

House-farm buildings are now a familiar part of the street scene Is this the solution to sustainbility ?

Tower block North Sumatra New build (2002)

Vertiginous new structure in Pak Phanang, S. Thailand

In southern Thailand swiftlet houses are a tourist attraction

A young bird at Miri, Sarawak Resembles neither of the wild species of Borneo

An adult from Kuching house

House-farmed white-nest swiftlets of Sarawak do not resemble either of the wild species Aerodramus germani on islands Aerodramus (f.) vestitus of inland caves

Pak Phanang S. Thailand

A bird that flew into a house in Taiping Presumably from a local farm-house

Adult bird no. 2, Penang 29 Oct 08

Adult bird no. 3, Penang 29 Oct 08

Adult house swiftlet at Taiping, 30 Sep 09

Swiftlet at Pusing With unusually dark rump

Adult bird Kota Bahru Oct 08

Neighbour Joining Tree based on partial Cyt b region Sibu (03) Setiawan (SW102) Setiawan (SW01) Setiawan (SW101) Kuantan (KT152) Endau (EN191) Endau (EN193) Rompin (RP61) Selangor (SB32B) Selangor (SB32A) Sumatra ( SM77) Sumatra (SM78) Sumatra (SM77a) Sumatra (SM76) vestitus Goh Wei Lim Baram (LS01) Baram (LS09 ) Kuantan (KT41B ) Setiawan (SW02) Kuantan (KT41A) Kuantan (KT151) Endau (EN192 ) A. maximus 66 95 83 49 59 0.005

1958 sonograph of echo-locating calls of Aerodramus maximus < - - - 2 seconds - - - > Single clicks, very short duration : 0 – 15 kHz ‘white sound’ A||||||||||||||V 20 kHz

Malaysian swiftlets: statistics for wild birds (averages ) Species Ad. Wt. (g) Egg size (mm) Clutch Incub’n (days) Fledge (days) Black-nest 18 25 x 16 1 28 59 White-nest ( vestitus) 12.5 20 x 14 2 25 45 Mossy- nest 12.5 21 x 14 2 23 48 Glossy ( cyanoptila ) 7.5 17 x 11 2 22 42

White-nest : coordinated breeding Lim Chan Koon

White-nest swiftlet: eggs & nestlings Salai cave, Sarawak, 1997-98 1997 1998 Eggs Nestlings Lim Chan Koon

White-nest Swiftlets: % undisturbed, marked nests containing at least one egg = breeding. 3 Sarawak caves Lim Chan Koon

A. (f.) vestitus : ‘Porcupine’ stage

Black-nest swiftlet: Slow nestling growth Body mass (g) Wing length (mm) Adult Kang et al.

Inactive gland Active gland Black nest swiftlet

Commercial nests “ C. brevisrostris” Apus pacificus Apus affinis Human growth hormone

Repackaged as a novel product

Breeding cycle & salivary gland activity Females Males Gonads active Salivary glands inactive Gonads inactive Salivary glands active

Malaysia (2-5 o N) Annual breeding cycle and moult All birds

A. (f.) vestitus : breeding & moult *** 1 st primaries moulted Last primaries moulted *** Lim Chan Koon

Black-nest swiftlet at Niah Annual moult of the primary tract

Black-nest swiftlets at Sarang: moult & breeding % nests with eggs Count of shed feathers Lim Chan Koon

Studies of behaviour at the cave mouth shows that swiftlets feed only during daylight hours, and that weather (especially rain) is an important determinant of the birds’ activity. It is likely that prolonged rainy weather is detrimental to their ability to feed

The influence of weather on behaviour

Analysis of food balls fed to nestlings, compared with passive samples of aerial insects, shows that swiftlets are selective feeders, taking a very wide variety of arthropod prey

The Malaysian government has recognised that the husbandry of swiftlets in houses falls into the remit of the Veterinary Department, including issues such as animal welfare and cruelty

Could managed, selective breeding of house-farmed birds improve the stock, e.g., female fecundity, nest weight or quality, disease resistance ?

In the scenario of this newest domestication, with the backing of sound husbandry and good science , rational planning can ensure the perpetuation and sustainable management of this important biological resource.

Carpometacarpus of a swiftlet Scale in mm Chris Stimpson

Carpometacarpus: Fossils from Niah cave Scale in mm Chris Stimpson

Carpometacarpus: Fossils from Niah cave Scale in mm Chris Stimpson