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Edible-nest Swiftlet

(Aerodramus fuciphagus, Thunberg 1812)

Anticlockwise from top:  Edible-nest Swiftlet in flight1  , perched on nest2 , harvested edible nest3 , prepared bird's nest soup.4



Contents:

1. Introduction

    1.1 Description
    1.2 Identification
    1.3 Etymology

2. Range

3. Nest building and human interactions

    3.1 Nest construction
    3.2 Bird's nest soup
    3.3 "Blood nests"
    3.4 House farming

4. Ecology and behaviour

    4.1 Feeding
    4.2 Reproduction
    4.3 Echolocation

5. Taxonomy and phylogenetics

  5.1 Original description
i 5.2 Phylogenetics of swiftlets
i 5.3 Species taxonomy


1. Introduction

1.1 General information

Edible-nest swiftlets are small insectivorous birds from the swift family Apodidae. Weighing in at a mere 8·7–14·8 grams and measuring in at 11·5–12·5 cm in length, this species is known to be an excellent flyer and can spend the long periods foraging in the sky without rest.5  Edible-nest swiftlets may be observed over a wide variety of habitats (even far out at sea) and are widespread throughout South-east Asia today. This bird is well known (and studied) in the region due to its nest being widely collected or farmed as a delicacy known as bird’s nest soup or ‘燕窝’ in Chinese. Natural populations nest in caves, such as the Niah and Gomantong caves in Borneo. However, due to the practice of farming swiftlets in nest houses, many Edible-nest swiftlets have adopted nesting in man-made buildings as early as the 1880s.6

1.2 Identification

A medium sized swiftlet with glossy dark brown upperparts.7  Rump tends to appear paler grey, but variable and can appear uniform. Brown-grey underparts except almost black undertail coverts, significantly forked tail. Naked or lightly feathered tarsi.

Figure 1: Photo composite of Edible-nest Swiftlets with diagnostic characters pointed out. (Composite and photos by Lim Hong Yao, 2018)8
Note that plumage can have subspecies variations.

In Singapore, difficult to distinguish in flight from slightly larger Black-nest Swiftlet (Aerodramus maximus), unless seen exceedingly well enough to determine the Edible-nest Swiftlet's slightly smaller (1-3 cm) size, relatively pointed tail ends with a deeper notch, and less bulky head and body. In hand, identification is possible as Black-nest Swiftlet has more extensive black feathering on tarsi.

1.3 Etymology

The genus Aerodramus stems from two Greek words: ἀερο, "aero" which means 'air', and δρόμος,"dromos" which means racecourse or path.9

The specific epithet fuciphagus originates from Greek as well, with φυκι, "fukos" meaning ‘seaweed,’ and φαγος, "phagos" meaning ‘to eat’. This is likely a result of the (false) historical belief that the swiftlets consume seaweed or substances from sea mist which allows it to produce a gelatinous substance with which it builds its nest with.

2. Range

The Edible-nest Swiftlet has a large range starting from the South of Hainan island spanning southward along the coasts of Vietnam, Myanmar, Thailand and Cambodia, subsequently including Sumatra, Java, Lesser Sundas, Borneo, and West Philippines (Figure 2).5  Inland Peninsula Malaysia is part of its natural range but Edible-nest Swiftlets from house farms are currently inhabiting the area. 8 current subspecies are recognized. For more details, see subspecies taxonomy below.

Range map of Edible-nest SwiftletFigure 2: Range map of the Edible-nest Swiftlet, adapted from HBW (2018)5

3. Nest building and human interactions

3.1 Nest construction

The Edible-nest Swiftlets nest in natural caves, and man-made houses. Breeding has been observed to take place year-round but peak in October and February.10

  • Nest building: Nests are made of salivary excrement that hardens into a cement-like material. Each nesting pair will spend about 25 minutes a day to build the nest, which takes about 45 days to complete.11  Nests are re-used for subsequent nesting by the same pair in the future if not harvested.10  Here is a video of the nest building process:

12

3.2 Bird's nest soup

The Edible-nest Swiftlet’s nests are frequently prepared in a manner known colloquially as ‘birds’ nest soup’, which has long been a Chinese delicacy of immense value since the T’ang Dynasty (618–907 A.D.).13  Like many other Chinese delicacies, birds’ nest soup is not especially tasty, and relies on other culinary additions to appeal to the palate.14  Its value lies in its rumoured nutritional value and health benefits instead; the broth is said to help reduce inflammation, alleviate gastric problems, boost the immune system, metabolism, and mental performance, as well as improve skin complexion.15  As with any claims of these sort, it is crucial to adopt an evidence-based approach in judging if the edible-nests are actually a health product. Thus far, several studies have shown that certain claims may hold some truth to them. The paper Wong (2013) summarises several potential biological effects published in studies, some of which include pro-mitogenic effects, epidermal growth factor-like activity, pro-proliferative effects for lymphocytes and corneal keratocytes, increased anti-body production, anti-oxidative effects, ability to bind to influenza virus to inhibit damage, and potential bone strengthening and skin thickening/enhancing effects.16 17 18  However, it is important to note that all the above-mentioned studies either used ex vitro methods or experimented on mice or rabbits. As such there is no currently no definitive evidence of edible-nests being medicinal in nature after being consumed by humans. Many of the metrics pointing to potential effects may not actually result in desirable effects or appreciable effect sizes; increased femur calcium content and thicker skin does not translate to improved health and beauty. The current state of knowledge regarding the biological effects of edible-nests has not yet been able to prove the numerous claims associated with it, despite the studies pointing to its potential benefits. 

Given the high value of the nests (about USD$20 per nest, nest harvesting naturally became highly popular.15  Despite efforts to keep nesting colonies secret and manage the harvest rates, wild populations of the Edible-nest Swiftlets have plummeted from once immeasurable numbers, and many colonies have now been wiped out, or are critically endangered.19 20  Italy proposed to include the species under Appendix II of CITES to regulate trade in 1994, but opposition from the Southeast Asian nations prevented it, leaving it unlisted and thus unregulated.14

The Black-nest Swiftlet (Aerodramus maximus) also produces nests that are being collected and consumed, but their nests contain inedible feathers that need to be removed. 

3.3 "Blood nests"

Some of the nests develop red stains in them, and were known collectively as “blood nests” under the false belief that the stains composed of swiftlet blood.14  These nests are rarer and used to fetch up to five times the price of a regular nest.15  Instead, the red was due to nitrifying bacteria reacting to ammonia vapour from decaying guano (swiftlet and/or bat excrement).14  Some swiftlet farmers discovered this and induced these “blood nests” by treating the white nests with ammonia. Both artificial and natural red nests are found to have abnormally high levels of nitrite, up to 4400mg/kg, which can increase one’s risk of cancer, and potentially cause food poisoning.21  As a result, China, once the largest importer of swiftlet nests, banned the import of all birds’ nest from Malaysia and Indonesia. This caused a huge shock to the industry as demand of both "blood nests" and regular nests dropped significantly; China currently only imports from several Malaysian firms that meet strict regulations that were subsequently set up after the ban.22

3.4 House farming

Two centuries ago, the Edible-nest Swiftlet nested solely on coastal habitats where caves were available, and there were no records of inland nesting colonies.23  Swiftlets were first reported to nest in houses near cliffs in the village of Sedayu in East Java in 1880, and entrepreneurs eventually began developing methods to attract swiftlets to nest in houses.6  The industry grew as the use of broadcasting swiftlet calls to attract nesting birds became the norm, and a trade in swiftlet eggs began. House nesting swiftlets first appeared in the Malay Peninsula in the 1930s, reportedly first arriving in Singapore from Java, before spreading North across Malaysia.23  The swiftlets have a high degree of nest fidelity, and house farmed swiftlets and their young would return to man-made houses (preferably the same one) to nest again11 , leaving the depleted natural populations and habitats unrestored.23  Furthermore, house nesting swiftlets are suspected to be a hybrid population of A. f. fuciphagus and A. f. inexpectatus or A. f. germani in what could be an ongoing case of domestication; the population of house farmed swiftlets have since spread Northward up to Myanmar, where it may be competing with local subspecies. More extensive regional genetic studies are required to confirm these suspicions.



Bird's nest soup prepared with red dates and rock sugar.
(Photo: Steamy Kitchen, 2017)24


Edible-nest Swiftlet "blood nest". 
(Photo: Lim Hong Yao, 2018)25

Swiftlet house farm in Thailand
(Photo: Alexander Heitkamp, 2007)26

4. Ecology and behaviour

4.1 Feeding

Edible-nest Swiftlets are aerial insectivores that catch arthropods on the wing. Diet analyses have been conducted by examining regurgitated food boluses; hymenoptera (bees, wasps and ants), diptera (flies) and ephemeroptera (mayflies) made up majority of the food items, while arachnida (spiders and other arachnids), coleoptera (beetles) and hemiptera (true bugs) made up most of the rest.27

4.2 Reproduction

  • Chick development: Each pair of birds would usually lay two eggs and spend an average of 23 days incubating them. The chicks take about 43 days to develop and fledge, with an approximate 50% survival rate in nest houses when under observation (Nigel Langham 1979). Mortality is mostly attributed to eggs or chicks falling off the nests; it is not known why so many of them fall off the nest.


4.3 Echolocation

As Edible-nest Swiftlets nest in places which are completely dark, they have evolved the ability to echolocate using clicking sounds, along with many other species from the genus Aerodramus.28  This ability, however, is not diagnostic of the genus as the Pygmy Swiftlet (Collocalia troglodytes) has been proven to possess this ability too.

T4254-Aerodramusfuciphagus-snd-1.mp3
Sound recording of an Edible-nest Swiftlet's echolocating clicking calls. (Recorded by Lim Hong Yao, 2018)


5. Taxonomy

5.1 Original description

The Edible-nest Swiftlet was originally described by Carl Peter Thunberg, a well renowned Swedish naturalist from the 19th century. Thunberg collected the specimen in Java and described it as Hirundo fuciphagus in his Remarks about the Swallows that build jelly-like, edible Nests published in Kungl. Svenska vetenskapsakademiens handlingar (1812)29 (Figure 3), which, with help of Google Translate, roughly states that

  • he collected a specimen that appeared similar to Hirundo esculenta, but upon closer examination discovered it was an undescribed species. 
  • H. fuciphagus can be found in caves of the mountainous areas in Java and made jelly like nests that are a precious commodity.
  • The body of H. fuciphagus is black above and immaculate grey below, about four inches long
  • Tail is rounded and black above and below.
  • Wings black, twice the length of the tail and acute. Feet black and short.
  • Similar to H. esculenta but differentiated by the all black tail, chest and abdomen, no spots.

Type information could not be located online as far as my efforts went.

Figure 3: Scans of the original swedish description of H. fuciphagus in Kungl. Svenska vetenskapsakademiens handlingar by Thunberg (1812), obtained from Biodiversity Heritage Library.29

5.2 Phylogenetics of swiftlets

Current accepted classification of the species: Animalia - Chordata - Aves - Apodiformes - Apodidae - Aerodramus - A. fuciphagus 30 Thunberg (1812) named the Edible-nest Swiftlets in Java Hirundo fuciphagus, mistakenly grouping them together with the swallows (Hirundo is a genus of swallows, which are passerines).31  By the 1900s swiftlets were recognized as a separate taxon Collocalini 32 and all its members were initially lumped in the genus Collocalia, then subsequently divided into 3 genera: Hydrochous (Giant Swiftlets) are sister to Aerodramus (medium sized brownish swiftlets)and Collocalia (small glossy plumaged swiftlets). Multiple recent phylogenetic analyses have been conducted to establish that Collocalia are the basal group that are more related to the swifts, followed by Hydrochous and Aerodramus being sister genera (Figure 4).30 33 34 35



Figure 4: Phylogenetic tree of swiftlets obtained using nuclear and mitochondrial DNA analysis, estimated with Bayesian and Maximum Likelihood methods. Node support is denoted as posterior probabilities/bootstrap values. Adapted from Cibois et. al (2018).30

5.3 Species taxonomy

Many studies have attempted to resolve the phylogenetic relationships of the Edible-nest Swiftlet, but A. fuciphagus has often shown up as a paraphyletic clade with one or more individuals of A. fuciphagus are more closely related to A. salangana than other A. fuciphagus.30 34 35
 Even the most recent tree constructed using mitochondrial and nuclear DNA analysis depicts this (Figure 4). The low genetic divergence between closely related taxa is likely due to occasional hybridisation events, which have been reported in Sabah36 , leading to genetic introgression in the form of mitochondrial DNA (mtDNA) sweeps that cause some individuals to appear closer to sister species than its own.37

Furthermore, some authors disagree with classifying all 8 subspecies under a single species A. fuciphagus. The eight recognised subspecies for A. fuciphagus are as follows (descriptions are relative to nominate race unless stated)7 :

  • A. f. amechanus (Oberholser, 1912) – Anambas Is, off SE Peninsular Malaysia.
    • Darker upperparts with green sheen and paler underparts, with greyer rump than germani
  • A. f. germani (Oustalet, 1876) – Coastline from W Hainan S around SE Asia to Malay Peninsula, including Mergui Archipelago (off S Myanmar); coastal N Borneo and W Philippines (Palawan E to Panay and Ticao).
    • Paler underparts and whitish rump
  • A. f. inexpectatus (A. O. Hume, 1873) – Andaman Is and Nicobar Is.
    • Slightly smaller than nominate race
  • A. f. vestitus (Lesson, 1843) – Sumatra, Belitung I and Borneo (except N coasts).
    • Darkest upperparts, lack contrasting rump
  • A. f. perplexus (Riley, 1927) – Maratua I, off E Borneo.
    • Some purple sheen on rectrices and remiges, slight contrasting rump
  • A. f. fuciphagus (Thunberg, 1812) – Java, Kangean Is and Bali to W Lesser Sundas, and Tanahjampea.
    • Dark brown upperparts with slightly paler greyish rump. Underparts brownish-grey.
  • A. f. dammermani (Rensch, 1931) – Flores (EC Lesser Sundas).
    • Slightly paler rump
  • A. f. micans (Stresemann, 1914) – Sumba, Sawu and Timor (C Lesser Sundas).
    • Slightly greyer overall with contrasting rump

Differences amongst subspecies are often subtle and difficult to distinguish in the field due to variations in lighting as well as difficulty in observing constantly fast-moving subjects.

According to the subspecies range, Edible-nest Swiftlets observed in Singapore should be A. f. germani which extends into the Malay Peninsula, but specimens collected appeared identical to the nominate race A. f. fuciphagus, likely because colonies in Malaysia and Singapore are of the house farmed variety (see House Farming below), which is suspected to be of Javan origin (ssp. fuciphagus). To complicate matters, their feeding ranges are likely overlapping.

Several authors believe that this species should be split into two or three. Table 1 on the below presents a summary of the 3 different treatments.

Table 1: Summary of the authors' different species treatments of the Edible-nest Swiftlet complex.

AuthorSpecies

Subspecies

HBW5 & Stresemann (1931)32

A. fuciphagusall 8 listed above

Clement's checklist38

A. germani

germani, amechanus

A. fuciphagus

fuciphagus, vestitus, inexpectatus, perplexus, dammermani, micans

Cranbrook et al. (2013)23

A. fuciphagus

fuciphagus, vestitus, dammermani, micans

A. inexpectatus

inexpectatus, germani, perplexus

A. amechanusamechanus



The Clement's checklist treatment appears to be based on morphology and original descriptions of the subspecies, but does not appear to explicitly explain the treatment in any publication or platform. Despite this, many authors have adopted this treatment.

 Stresemann (1931) postulated that populations with a paler rump from A. f. germani and those of a darker rump from A. f. vestitus and germani formed a transition zone over Peninsula Malaysia where intergrades of rump colour can be observed.32  Thus, it was proposed that the populations were interbreeding, and thus the taxa were regarded as subspecies of A. fuciphagus. Medway (1966) also arrived at a similar conclusion.39

Cranbrook et. al (2013) on the other hand, re-examined the specimens used in Stresemann (1931) and postulated that there was not a cline in morphology, but rather an overlap of feeding range in Peninsula Malaysia.23  It was observed that the specimens could be grouped into two main groups: grey-rumped and brown rumped (Figure 5). Given that the rump colour was maintained as a character amongst these two groups with no gradation, it was concluded that these two should be grouped into two species, while amechanus was considered an endemic given its unusual glossy colouration and variable rump band. Grey-rumped swiftlets were grouped under A. inexpectatus while brown-rumped swiftlets remained as A. fuciphagus. Mitochondrial DNA analysis was also conducted in the study with both Maximum Parsimony and Neighbour Joining methods using cyt-b haplotypes but the nodes were all poorly supported and the tree appeared inconclusive regarding the phylogenetic relationships between populations (Figure 6).

However, upon examining the plates, it is apparent that rump colouration can vary significantly within a subspecies; A. inexpectatus germani of plate 3A (Figure 5) clearly shows an individual with a brown rump instead of a whitish rump as described for germani and shown in plate 1A. This raises concerns about using rump colour as a diagnostic trait to treat the species complex (HBW also mentions that the white on the rump is often overstated.7  Even if the morphological breaks separating the populations in different geographical ranges are real, in applying the Biological Species Concept40 , there is insufficient evidence to establish reproductive isolation given that the colonies breed in allopatry. Additionally, Cranbrook et. al (2013) suggested that the house farmed swiftlets could be a hybrid population of fuciphagus and inexpectatus, or fuciphagus and germani, indicating that these taxa may not withstand the test of sympatry when brought together in nest houses. Therefore, A. fuciphagus is currently treated as a single species (encompassing all 8 subspecies) on this page and by several other authors.

In considering the Phylogenetic species concept sensu Wheeler & Platnick (2000)40 , most of these subspecies are likely to be elevated to the rank of species given that the different populations appear to have a unique set of character states in terms of size and plumage differences. However, more work needs to be done as well to adequately sample the different populations to establish the existence of these different character states.

Future taxonomic work is required to concretely establish the relationships between these taxa, both in terms of confirming morphological differences as well as investigating molecular evidence. Goh et. al (2018) recently investigated more house farmed Edible-nest Swiftlets from Peninsula Malaysia and found that they appeared to be closest to A. f. vestitus rather than A. f. fuciphagus (which was first to be recorded in the Malay Peninsula), but the node was not very well supported (bootstrap = 78) and no conclusive statements about their origin could be made.41  Given the many unsuccessful attempts with mtDNA thus far, it is likely that mtDNA is unsuitable as a marker for intraspecific studies as it is only maternally inherited. Coupled with the issues of mtDNA sweeps, it is evident that mtDNA is not an effective  taxonomic indicator for the Edible-nest Swiftlets, and adopting genomic methods using Next Generation Sequencing is likely the way forward to unravel the true relationships between these populations.


Figure 5: Photo plates by Cranbrook et. al (2013) depicting the grey-rumped and brown rumped Edible-nest Swiftlet specimens examined23 .


Figure 6: Maximum Parsimony tree created using cyt-b haplotypes from Edible-nest Swiftlet samples by Cranbrook et. al (2013)23 . Numbers at nodes are the Neighbour Joining/Maximum Parsimony bootstrap values (tree topology was identical for both methods). 

References: 

Footnotes
Ref Notes
1 "Edible-nest Swiftlet in flight," by Lim Hong Yao, 2017. 
2 "Edible-nest Swiftlet on nest," by Lim Hong Yao, 2018. 
3 "Edible birds nest bowl shape" by Ediblebirdsnest. Wikimedia Commons,19 March 2016. URL https://commons.wikimedia.org/wiki/File:Edible-birds-nest-bowl-shape.png (accessed on 19 Nov 2018)
4 "Birds nest soup" by Crue. Blogger, 6 January 2010. URL http://ichikichikehem.blogspot.com/2010/01/thai-hatyai-bird-nest-soup-7ps.html (accessed on 19 Nov 2018)
5 Chantler, P. & Boesman, P. (2018). Edible-nest Swiftlet (Aerodramus fuciphagus). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. (retrieved from https://www.hbw.com/node/55287 on 29 November 2018). [ a b c d ]
6 Lim, C. and Cranbrook, G. (2002). Swiftlets of Borneo. Natural History Publications (Borneo). [ a b ]
7 Chantler, P. & Boesman, P. (2018). Edible-nest Swiftlet (Aerodramus fuciphagus). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. (retrieved from https://www.hbw.com/node/55287 on 29 November 2018). [ a b c ]
8 "Edible-nest Swiftlet identification characters composite," by Lim Hong Yao, 2018. 
9 Jobling, J. (2011). The Helm dictionary of scientific bird names. London: Christopher Helm.
10 Langham, N. (2008). Breeding biology of the edible-nest swiftlet Aerodramus fuciphagus. Ibis, 122(4), pp.447-461. [ a b ]
11 Kang, N., Hails, C. and Sigurdsson, J. (2008). Nest construction and egg-laying in Edible-nest Swiftlets Aerodramus spp. and the implications for harvesting. Ibis, 133(2), pp.170-177. [ a b ]
12 "Salivating for a new nest | Wild Borneo" by National Geographic. Nat Geo WILD YouTube Channel, 3 January 2017. URL: https://www.youtube.com/watch?v=ngPs3kINUXE  (accessed on 30 Nov 2018).
13 Lau, A. and Melville, D. (1994). International trade in swiftlet nests with special reference to Hong Kong. Cambridge, U.K.: Traffic International.
14 Thorburn, C. (2015). The Edible Nest Swiftlet Industry in Southeast Asia: Capitalism Meets Commensalism. Human Ecology, 43(1), pp.179-184. [ a b c d ]
15 Marcone, M. (2005). Characterization of the edible bird’s nest the “Caviar of the East”. Food Research International, 38(10), pp.1125-1134. [ a b c ]
16 Oda, M., Ohta, S., Suga, T. and Aoki, T. (1998). Study on Food Components: The Structure of N-Linked Asialo Carbohydrate from the Edible Bird's Nest Built by Collocalia fuciphaga. Journal of Agricultural and Food Chemistry, 46(8), pp.3047-3053.
17 Roh, K., Lee, J., Kim, Y., Park, J., Kim, J., Lee, J. and Park, D. (2012). Mechanisms of Edible Bird's Nest Extract-Induced Proliferation of Human Adipose-Derived Stem Cells. Evidence-Based Complementary and Alternative Medicine, 2012, pp.1-11.
18 Wong, R. (2013). Edible bird’s nest: Food or medicine?. Chinese Journal of Integrative Medicine, 19(9), pp.643-649.
19 Lau, A. and Melville, D. (1994). International trade in swiftlet nests with special reference to Hong Kong. Cambridge, U.K.: Traffic International.
20 Sankaran, R. (2001). The status and conservation of the Edible-nest Swiftlet (Collocalia fuciphaga) in the Andaman and Nicobar Islands. Biological Conservation, 97(3), pp.283-294.
21 Chibber, A. (2011). China finds nitrite in edible bird's nests from Malaysia. [online] Food navigator-Asia. Available at: https://www.foodnavigator-asia.com/Article/2011/08/25/China-finds-nitrite-in-edible-bird-s-nests-from-Malaysia [Accessed 30 Nov. 2018].
22 News Asiaone. (2013). China lifts ban on bird's nest imports to Malaysia. [online] Available at: http://news.asiaone.com/News/Latest%2BNews/SoShiok/Story/A1Story20130621-431216.html [Accessed 30 Nov. 2018].
23 Cranbrook, E., Goh, W. L., Lim, C. K., and Rahman, M. A. (2013). The species of white-nest swiftlets (Apodidae, Collocaliini) of Malaysia and the origins of house-farm birds: morphometric and genetic evidence. Forktail, 29, pp. 107–119 [ a b c d e f g ]
24 "Chinese birds nest soup" by Jaden Hair. Steamy Kitchen, 19 June 2017. URL https://steamykitchen.com/44260-chinese-birds-nest-soup-recipe.html. (accessed on 19 Nov 2018)
25 "Swiftlet blood nest," by Lim Hong Yao, 2018. 
26 "Chinese birds nest soup" by Alexander S. Heitkamp. Wikimedia Commons, 2007. URL https://commons.wikimedia.org/wiki/File:Nestinghouse_003.jpg. (accessed on 30 Nov 2018)
27 Lourie, S. and Tompkins, D. (2008). The diets of Malaysian swiftlets. Ibis, 142(4), pp.596-602.
28 Jordan Price, J., P. Johnson, K. and H. Clayton, D. (2004). The evolution of echolocation in swiftlets. Journal of Avian Biology, 35(2), pp.135-143.
29 Thunberg. (1812). Remarks about the Swallows that build jelly-like, edible Nests. Kungl. Svenska vetenskapsakademiens handlingar, Vol. 1, pp.151-156. Accessed from https://www.biodiversitylibrary.org/page/47088095#page/165/mode/1up [ a b ]
30 Cibois, A., Thibault, J., McCormack, G. and Pasquet, E. (2018). Phylogenetic relationships of the Eastern Polynesian swiftlets (Aerodramus, Apodidae) and considerations on other Western Pacific swiftlets. Emu - Austral Ornithology, 118(3), pp.247-257. [ a b c d ]
31 Thunberg. (1812). Remarks about the Swallows that build jelly-like, edible Nests. Kungl. Svenska vetenskapsakademiens handlingar, Vol. 1, pp.51-156. Accessed from https://www.biodiversitylibrary.org/page/47088095#page/165/mode/1up
32 Stresemann, E. (1931). Notes on the systematics and distribution of some swiftlets (Collocalia) of Malaysia and adjacent subregions. Bulletin of the Raffles Museum, 6, pp.83–101. [ a b c ]
33 Päckert, M., Martens, J., Wink, M., Feigl, A. and Tietze, D. (2012). Molecular phylogeny of Old World swifts (Aves: Apodiformes, Apodidae, Apus and Tachymarptis) based on mitochondrial and nuclear markers. Molecular Phylogenetics and Evolution, 63(3), pp.606-616.
34 Price, J., Johnson, K., Bush, S. and Clayton, D. (2005). Phylogenetic relationships of the Papuan Swiftlet Aerodramus papuensis and implications for the evolution of avian echolocation. Ibis, 147(4), pp.790-796. [ a b ]
35 Rheindt, Frank E., Janette A. Norman & Les Christidis. (2014). Extensive diversification across islands in the echolocating Aerodramus swiftlets. Raffles Bulletin of Zoology, 62, pp.89-99. [ a b ]
36 Lee, P., Clayton, D., Griffiths, R. and Page, R. (1996). Does behavior reflect phylogeny in swiftlets (Aves: Apodidae)? A test using cytochrome b mitochondrial DNA sequences. Proceedings of the National Academy of Sciences, 93(14), pp.7091-7096.
37 Rheindt, F. and Edwards, S. (2011). Genetic Introgression: An Integral but neglected component of speciation in birds. The Auk, 128(4), pp.620-632.
38 Clements, J. F., T. S. Schulenberg, M. J. Iliff, D. Roberson, T. A. Fredericks, B. L. Sullivan, and C. L. Wood. 2018. The eBird/Clements checklist of birds of the world: v2018. Downloaded from http://www.birds.cornell.edu/clementschecklist/download/ 
39 Medway, L. (1966). Field characters as a guide to the specific relations of swiftlets. Proceedings of the Linnean Society of London, 177, pp.151–172.
40 Wheeler, Quentin D., and Rudolf Meier, editors. Species Concepts and Phylogenetic Theory: A Debate. Columbia University Press, 2000. JSTOR, www.jstor.org/stable/10.7312/whee10142. [ a b ]
41 Goh, W.L., Siew, W.S., Davies, S.E.W., Ball, S. Ball., Khoo, G., Lim, C.K., Rahman, M. A., and Cranbrook, E. (2018). Genetic diversity among white-nest swiftlets of the genus Aerodramus (Aves: Apodidae: Collocaliini) of house-farms in Malaysia. Raffles Bulletin of Zoology, 66, pp.350–360

This page was authored by Lim Hong Yao

Last curated on 3 December 2018

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