Open Access

Birds caught in spider webs in Asia

Avian Research20167:16

https://doi.org/10.1186/s40657-016-0051-4

Received: 21 June 2016

Accepted: 20 August 2016

Published: 19 October 2016

Abstract

A recent global review of birds caught in spider webs reported only three Asian cases. Given this surprisingly low number, I made a concerted effort to obtain additional Asian cases from the literature, the internet, and field workers. I present a total of 56 Asian cases which pertain to 33 bird species. As in the global dataset, mostly small bird species were caught in spider webs, with a mean body mass of 17.5 g and a mean wing chord length of 73.1 mm. Consequently, birds with a body mass >30 g were very rarely caught. This Asian review corroborates the global review that smaller birds are more likely to be caught and that Nephila spiders are most likely to be the predators. Continuous monitoring of spider webs is recommended to ascertain the frequency of these events.

Keywords

Spider Predator–prey relationships Asia

Background

Birds may be killed by environmental factors (e.g. weather; Elkins 2004), accidents or parasites (e.g. Jennings 1961), or predators. The most important predators of birds are birds, reptiles and mammals, including humans, but, more rarely, birds are also predated upon by amphibians, fish and insects (Brooks 2012). A presumably rather rare case of death occurs when a bird gets caught in a spider web; in a global review, Brooks (2012) reviewed 68 cases of birds getting trapped and often killed in the webs of large spiders. When a bird flies into a spider web, the bird may either bounce off the web or fly right through it, or it may become entangled; once entangled, the spider may or may not wrap the bird in silk. Entangled birds may then free themselves again, or they die either due to exhaustion or spider predation, while wrapped birds invariably die unless freed by humans (for details, see Brooks 2012).

Birds should therefore always attempt to avoid collision with spider webs, while the interests of spiders may differ depending on the species. Some spider species opportunistically consume trapped birds (especially large Nephila spiders, see below) and may therefore keep their webs inconspicuous to birds. However, other spider species apparently try to avoid collisions and the consequent damage to their webs by making them more visible to birds (Bruce et al. 2005; Walter and Elgar 2011).

As one may expect, Brooks’ (2012) global review documented that it is almost exclusively smaller birds (mean body mass = 10.7 g, mean wing chord length = 61 mm) which get caught in spider webs. Consequently, 88 and 90 % of all caught birds had a body mass ≤15 g and a wing chord length <90 mm, respectively. In the 34 cases in which the spider was identified, 62 % belonged to the genus Nephila, and all were orb weavers except for a single Latrodectus species.

Most of these cases were reported from Africa, Australia, North America, and the Neotropics, but only a few from Europe and Asia (D. Brooks in litt. 2014). Thus, Brooks (2012) only reported three Asian cases: a Spotted Flycatcher (Muscicapa striata) in Iran (Doberski 1973), a juvenile Laughing Dove (Streptopelia senegalensis) in Oman in 2003 (Forsman 2003), and a Dusky Warbler (Phylloscopus fuscatus) in China some time before 2007 (D. Brooks in litt. 2014). Kasambe et al. (2010) presented another four cases from India not mentioned in Brooks (2012). Given that Nephila species are distributed across much of tropical, subtropical and even some parts of temperate Asia (Miyashita et al. 1998; Murphy and Murphy 2000; Lee et al. 2004; Harvey et al. 2007; Su et al. 2007, 2011; http://www.gbif.org/species/2149490), this relative lack of records seemed surprising. Therefore, I made a concerted effort to obtain additional cases of birds being caught in spider webs in Asia using various sources.

Methods

In 2014 and early 2015, I used eight methods to obtain additional cases from the literature, the internet, Asian ornithologists, birdwatchers and birding tour leaders: (1) I emailed all the authors who published in BirdingASIA and Forktail and whose emails I could take from the journals’ websites or the Web of Science. (2) I emailed all the authors of any article published in an ornithological journal listed on the Web of Science which were returned upon using the keywords “bird” and “Asia”. (3) I posted requests on the birding fora of the Birds of Bangalore, Birds of Bombay, Bombay Natural History Society, Hong Kong Bird Watching Society, Hong Kong Wildlife Net, Kerala Birder, Malaysia Birders, Oriental Bird Club, Ornithological Society of the Middle East, and Pengamat Burung Indonesia. (4) I extensively used the web, images and video search functions of Google and Google Scholar using various combinations of the keywords “spider” “catch” “bird” “Asia” and names of Asian countries. (5) Upon any reply, I asked the person to forward my email request to other Asian ornithologists and birdwatchers. (6) I tried to obtain all references given in publications or websites which reported another case. (7) In early 2014, two native Chinese speakers (J.-L. Wu, T.-Y. Wu in litt. 2014) used Google Taiwan to search Taiwanese websites for cases using relevant keywords (see above), and I emailed all Taiwanese ornithologists and birders that I personally knew. (8) In late 2014, two native Japanese speakers (M. Kamioki, M. Mashiko in litt. 2014) used Google Japan to search Japanese websites for cases using relevant keywords (see above). I kindly request that further cases be reported to my email.

For easy comparison, I mirrored Brooks’ (2012) analysis as much as possible. As described in Brooks (2012), I sought data on body mass and wing chord length for each bird species from various data sources (given in Table 1) and, if possible, determined the species of spider (given in Table 1). Unlike Brooks (2012), I added location and date for each record, if possible.
Table 1

Birds entrapped in spider webs in Asia and their respective sizes

Common name

Scientific name

Spider sp.

Mass (g)

Wing (mm)

Location

Date

Source

Glossy Swiftlet

Collocalia esculenta

Np

8.0

950,2,3

Great Nicobar island, Nicobar Islands, India

Before 2010

Manchi and Sankaran (2009)

Edible-nest Swiftlet

Collocalia fuciphaga

S

10.7

1181,1,3

Interview Island, Andaman Islands, India

June 2006

Manchi and Sankaran (2009)

Asian Palm Swift

Cypsiurus balasiensis

Na

9.2

1121,1,0

Doi Kham, Chiang Mai Province, Thailand

6 October 2014

W. Limparungpatthanakij in litt. 2014

Laughing Dove

Streptopelia senegalensis

Na

80.0c

138c

Oman

~ October 2003

Forsman (2003), Brooks (2012), D. Brooks in litt. 2014

Pied Fantail

Rhipidura javanica

S

12.5

821,1,0

Near U Minh Thuong National Park, Kien Giang Province, Vietnam

9 August 2008

M. Le in litt. 2014

Black-naped Monarch

Hypothymis azurea

Np

11.3

69

Sanjay Gandhi National Park, Mumbai, India

October 1996

Andheria (1998, 1999)

Spotted Flycatcher

Muscicapa striata

S

14.0c

80c

Main Kaleh Reserve, Iran

1972

Doberski (1973)

Grey-streaked Flycatcher

Muscicapa griseisticta

S

15.1

83

Taiwan Area

September 2008

http://tinyurl.com/spider-tw2

Grey-streaked Flycatcher

Muscicapa griseisticta

Np

15.1

833,5,8

Iriomote Island, Japan

5 October 2008

http://tinyurl.com/spider-jp6

Asian Brown Flycatcher

Muscicapa dauurica

S

9.9

661,1,0

Thap Lan National Park, Nakhon Ratchhasima Province, Thailand

14 November 1999

P. Round in litt. 2014

Asian Brown Flycatcher

Muscicapa dauurica

S

9.9

661,1,0

Po Toi Island, Hong Kong, China

After 2006

G. Welch in litt. 2014

Asian Brown Flycatcher

Muscicapa dauurica

Np

9.9

661,1,0

Po Toi Island, Hong Kong, China

8 September 2011

M. Hale and G. Welch in litt. 2014

Hill Blue Flycatcher

Cyornis banyumas

Np

14.5

671,1,0

Bukit Larut, Perak State, Malaysia

Unknown

Anonymous (1999b)

Great Tita

Parus major

Np

15.5

591,0,0

Komesu, Itoman City, Okinawa Island, Japan

10 August 2011

http://tinyurl.com/spider-jp1, http://tinyurl.com/spider-jp2

Dusky Crag Martinb

Hirundo concolor

Pf

13.0

981,1,0

Shevaroys (=Servarayan) Hills near Salem, Tamil Nadu

Before 1889

Morris (1889) and Anonymous (1999a)

Light-vented Bulbul

Pycnonotus sinensis

Np

29.7

85

Majia, Pingtung, Taiwan Area

March 2004

http://tinyurl.com/spider-tw3

Styan’s Bulbul

Pycnonotus taivanus

Np

26.2

84

Guangfu, Hualien, Taiwan Area

18 June 2010

http://tinyurl.com/spider-tw4

Yellow-vented Bulbul

Pycnonotus goiavier

Np

27.8

821,4,6

Kledang-Sayong Forest Reserve, Ipoh, Perak, Malaysia

11 February 2014

Amar-Singh (2014a, b), Amar-Singh H. in litt. 2014

Buff-vented Bulbul

Iole olivacea

S

24.5

891,1,0

Near Ban Bang Khram, Khlong Thom District, Krabi (area also known as Khao Nor Chuchi), Thailand

7 August 2013

P. Round in litt. 2014

Brown-eared Bulbul

Ixos amaurotis

Np

70.9

1161,1,0

Tokunoshima Island, Japan

1 August 2010

http://tinyurl.com/spider-jp3

Plain Prinia

Prinia inornata

Np

8.2

49

Tadoba Andhari Tiger Reserve, Chandrapur district, Maharashtra, India

October 1998

Anonymous ( 1999a)

Plain Prinia

Prinia inornata

Np

8.2

49

Melghat Tiger Reserve, northern part of Amravati District of Maharashtra State, India

Before 2005

Pande et al. (2004)

Plain Prinia

Prinia inornata

S

8.2

49

Western Ghats, Maharashtra, India

Unknown

S. Pande in litt. 2015

Plain Prinia

Prinia inornata

S

8.2

49

Taiwan Area

September 2008

http://tinyurl.com/spider-tw2

Oriental White-eye

Zosterops palpebrosus

Na

8.6

511,1,5

Sungei Buloh Wetland Reserve, Singapore

30 April 2012

Ong (2012a, b)

Japanese White-eye

Zosterops japonicus

Np

11.3

53

Mong Tseng Tsuen (near Tsim Bei Tsui), Hong Kong, China

22 August 2004

So (2005)

Japanese White-eye

Zosterops japonicus

Np

11.3

53

Keelung, Taiwan Area

18 August 2005

http://tinyurl.com/spider-tw5

Japanese White-eye

Zosterops japonicus

Np

11.3

53

Taiwan Area

Before October 2005

http://tinyurl.com/spider-tw7

Japanese White-eye

Zosterops japonicus

Np

11.3

53

Badouzi, Keelung, Taiwan Island

13 August 2011

http://tinyurl.com/spider-tw6

Japanese White-eye

Zosterops japonicus

Np

11.3

53

Okinawa Island, Japan

November 2012

http://tinyurl.com/spider-jp5

Lanceolated Warbler

Locustella lanceolata

S

12.9

55

E-Luan-Pi lighthouse, Kenting National Park, Pintung County, Taiwan Area

14 October 1984

TESRI# collection number w672, C.-t. Yao in litt. 2013

Grasshopper Warbler

Locustella naevia

N

14.8

64

Tungareshwar Wildlife Sanctuary, Maharashtra, India

18 November 2006

Kasambe et al. (2010)

Common Tailorbird

Orthotomus sutorius

N

7.5

432,2,5

Mogarkasa Forest, Nagpur, Maharashtra, India

13 November 2008

Kasambe et al. (2010)

Dark-necked Tailorbird

Orthotomus atrogularis

Np

7.7

381,1,0

Kaeng Krachan National Park, Petchaburi Province, Thailand

2012

W. Limparungpatthanakij in litt. 2014

Dusky Warbler

Phylloscopus fuscatus

S

11.0c

57c

Beidahe, Hebei Province, China

Before 2007

D. Zetterström in litt. 2007 (D. Brooks in litt. 2014)

Arctic Warbler

Phylloscopus borealis

Np

10.0

65

Yonaguni Island, Japan

10 September 2008

http://ameblo.jp/attacus/theme2-10004405518.html

Arctic Warbler

Phylloscopus borealis

Np

10.0

65

Bitou Cape, New Taipei City municipality, Taiwan Area

6 September 2011

Y.-P. Chiang in litt. 2013‒2014

Arctic Warbler

Phylloscopus borealis

Np

10.0

65

Pak Sha O, Hong Kong, China

19 September 2015

Geoff Carey in litt. 2015

Greenish Warbler

Phylloscopus trochiloides

N

7.1

60

Bandhavgarh National Park, Madhya Pradesh, India

12 October 2007

Kasambe et al. (2010)

Greenish Warbler

Phylloscopus trochiloides

N

7.1

60

Kanha National Park, Madhya Pradesh, India

22 October 2008

Kasambe et al. (2010)

Buff-breasted Babbler

Pellorneum tickelli

S

17.1

611,1,0

Fraser’s Hill, Pahang, Malaysia

5–11 June 2010

S. Pieterse in litt. 2014

Taiwan Yuhina

Yuhina brunneiceps

S

12.2

62

Taiwan Area

Unknown

H.-S. Lin in litt. 2013

Vinous-throated Parrotbill

Paradoxornis webbianus

Np

9.3

52

Mountain Pinglin, Taichung City, Taiping District, Taiwan Area

2007

http://tinyurl.com/spider-tw1

Brown-throated Sunbird

Anthreptes malacensis

S

11.4

661,1,3

Ipoh City, Perak, Malaysia

28 December 2007

Amar-Singh (2014a, b), Amar-Singh H. in litt. 2014

Eurasian Tree Sparrow

Passer montanus

S

23.0

66

Luku, Nantao County, Taiwan Area

1990s

C.-t. Yao in litt. 2013

Eurasian Tree Sparrow

Passer montanus

Np

23.0

66

Taiwan Area

Summer 2004

http://tinyurl.com/spider-tw9

Eurasian Tree Sparrow

Passer montanus

S

23.0

66

Taiwan Area

Before August 2010

http://tinyurl.com/spider-tw8

Eurasian Tree Sparrow

Passer montanus

Np

23.0

66

Jiji, Nantou County, Taiwan Area

13 August 2013

C.-t. Yao in litt. 2013

White-rumped Munia

Lonchura striata

S

11.3

48

Taiwan Area

Unknown

Y.-C. Hsu in litt. 2013

Munia spec.

Np

Bogor Botanical garden, Bogor, Indonesia

Before 1934

Boedijn (1933)

Munia spec.

Np

probably near or in Bogor, Indonesia

Before 1934

Boedijn (1933)

“Small birds”

Np

Thailand

Before 1933

Bristowe (1932)

Unidentified

Np

probably near or in Bogor, Indonesia

Before 1934

Boedijn (1933)

Unidentified

Np

Cheung Sha, Lantau Island, Hong Kong, China

8 October 2006

Anonymous (2006)

Unidentified

Np

Wang Tong River, Mui Wo, Lantau Island, Hong Kong, China

12 October 2009

M. Pearse in litt. 2015

Unidentified

Np

Miyakojima Island, Japan

16 October 2011

http://tinyurl.com/spider-jp7

Common and scientific bird names and taxonomic order follow Inskipp et al. (1996)

Spider species as follows: S = bird was caught by a spider; N = bird was caught by a Nephila species, family Nephilidae, suborder Araneomorphae, order Araneae; Na = bird was caught by Nephila antipodiana; Np = bird was caught by Nephila pilipes (=maculata); Pf = bird was caught by Poecilotheria (=Mygale) fasciata, family Theraphosidae, suborder Mygalomorphae, order Araneae. Body masses were obtained from Glutz von Blotzheim (1966–1996), Dunning (2008), Severinghaus et al. (2010), the Encyclopedia of Life (eol.org) and Wikipedia (en.wikipedia.org). Wing chord lengths were obtained from Glutz von Blotzheim (1966‒1996) and Severinghaus et al. (2010) except when a superscript indicates the number of male, female and unsexed specimens which were measured by P. Capainolo (in litt. 2014) at the American Museum of Natural History, New York, USA, H. van Grouw (in litt. 2014) at the Natural History Museum, Tring, UK, A. Gamauf (in litt. 2014) at the Naturhistorische Museum Wien, Austria, and T. Töpfer (in litt. 2014) at the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany

aAlso classified as Eastern Great Tit (Parus minor)

bThe martin referred to in Morris (1889) must be a Dusky Crag Martin because of the record’s location and the use of a house to build its nest (R. Kasambe, H. Rathore, in litt. 2014)

cI used the body masses and wing chord lengths given for the three Asian cases mentioned in Brooks (2012)

dTESRI refers to Taiwan Endemic Species Research Institute, Jiji, Nantou County, Taiwan Area

Results

In Asia, I was able to document 53 cases in addition to the three cases listed by Brooks (2012) (Table 1). The Asian cases now contain 33 bird species, and together with Brooks’ (2012) global dataset, 84 bird species have been documented so far (Table 2). Three and 12 spider species were identified for Asia and the world, respectively; these are (with the number of Asian cases and cases from other continents in brackets): Aranens trifolium (0/1), Argiope aurantia (0/3), Argiope caphinarium (0/1), Argiope sp. (0/2), Eriophora biapicata (0/1), Latrodectes sp. (0/1), Mastophora sp. (0/1), Neoscona hentzii (0/1), Nephila antipodiana (3/3), Nephila clavipes (0/14), Nephila pilipes (31/32), Nephila sp. (4/8), Nephilengys cruentata (0/2), Poecilotheria fasciata (1/1), and unidentified spiders (17/49) [Brooks (2012) also mentions Nephila inaurata in his text, but it is not listed in his Table 1]. Thus, 38 out of 39 identified cases (97 %) in Asia were Nephila species.
Table 2

Mean body masses and mean wing chord lengths of birds caught in spider webs in Asia (Table 1) and the world (Table 1; Brooks 2012); naturally, cases of unidentified bird species in Table 1 were excluded

Analysis (sample size)

Spider species

Mass (g)

Wing (mm)

Asia

Individuals (n = 49)

3

15.9 ± 13.7 (7.1‒80.0)

68.9 ± 20.9 (38.0‒138.0)

Species (n = 33)

3

17.5 ± 16.2 (7.1‒80.0)

73.1 ± 23.6 (38.0‒138.0)

World

Individuals (n = 114)

12

12.3 ± 10.8 (2.0‒80.0)

63.3 ± 20.1 (37.0‒138.0)

Species (n = 84)

12

13.5 ± 11.8 (2.0‒80.0)

66.4 ± 21.8 (37.0‒138.0)

The analyses were also split into individuals (i.e. all cases) and species (i.e. one case for each bird species). Each entry for body mass and wing chord length gives the mean ± standard deviation and the range in brackets

The mean body mass and mean wing chord length are slightly larger for the Asian than for the global dataset (Table 2). This difference is certainly due to the large number of hummingbirds in Brooks’ (2012) dataset which are all smaller than the smallest Asian species, the Greenish Warbler (Phylloscopus trochiloides; Table 1). Means are also slightly larger for the means calculated across all species than for the means calculated across all individual cases (Table 2). This difference is due to smaller-than-average species caught repeatedly; of the 14 species with more than one case, 11 species had a body mass ≤10 g and 9 species had a wing chord length ≤60 mm (Table 1; Brooks 2012). Among the 49 Asian cases identified to bird species, 71 and 88 % of all caught birds had a body mass ≤15 g and a wing chord length <90 mm, respectively; for the 114 global cases, the respective percentages are 82 and 89 % (Table 1; Brooks 2012). A frequency diagram of all cases shows the great propensity of small-bodied birds being caught (Fig. 1). Cases with a body mass >30 g are exceedingly rare, and the two largest species ever caught, the Laughing Dove (80.0 g) and the Brown-eared Bulbul (Ixos amaurotis, 70.9 g), are anomalies in the general trend.
Fig. 1

Frequency diagram of body mass intervals (in steps of 10 g) of 49 cases (black bars) from Asia (Table 1) and the remaining 65 cases (grey bars) from other continents (Brooks 2012), whereby each case involves one individual bird getting caught in a spider web as described in the text

The oldest case recorded in Asia is the Dusky Crag Martin (Hirundo concolor) reported in Morris (1889) that equals the previous oldest record by McCook (1889) cited in Brooks (2012). Only 11 of the Asian cases are from before 2000 (Table 1); likely reasons are that many records were reported on the internet (Table 1), and that many of the contacted ornithologists and birdwatchers were not active before 2000.

Discussion

Birds are usually predators of spiders or the contents of their webs (e.g. Waide and Hailman 1977; Gunnarsson 2007), but when small birds encounter spider webs of large spiders, the tables can be turned. Overall, this review of Asian cases corroborates the conclusions made by Brooks (2012), namely: (1) the smaller the bird species, the higher the likelihood to be caught in spider webs; and (2) Nephila species are by far the most common spiders to catch birds in their webs.

However, my review of Asian cases suggests that cases of birds getting caught in spider webs may be as common in Asia as in other continents wherever large orb weaver spiders are common. Therefore, the small number of Asian cases in Brooks (2012) represented a biased picture of the Asian situation. Asia covers 30 % of the world’s terrestrial surface, and, due to this review, 46 % (56 out of 121) of all documented cases now come from Asia, thus giving a more representative picture.

Naturally, reporting bias is likely to be considerable for rare natural history events like these, and Brooks (2012) therefore emphasized “the importance of reporting interesting natural history notes and keeping good field records.” An example of positive reporting bias is likely to be Taiwan. At 35,883 km2, Taiwan has only 0.08 and 0.02 % of the terrestrial surface of Asia and the Earth, respectively. However, the 16 cases reported from Taiwan (Table 1) represent 29 % of all Asian and 13 % of all global cases. One reason may be that large spiders are certainly common in Taiwan Island, and especially in somewhat disturbed or semi-open habitats with many small gaps and openings suitable for building webs, such as the coastal forests at Bitou Cape (cf. Table 1) where a large spider web can be seen approximately every 10 m. Accordingly, Brooks et al. (2008) and Brooks (2012) hypothesized that disturbed habitats, e.g. forests disturbed by severe storms, may see an increase in the number of large spiders in the lower strata, as possible attachment sites for webs were destroyed in the upper strata, and Taiwan is regularly subjected to devastating typhoons. Furthermore, Taiwan has a very active bird-watching community and widespread internet use, evidenced by the fact that 10 of the 16 Taiwanese cases were reported on the internet (Table 1). The internet and citizen-science can thus play an increasing role in gathering and disseminating natural history information (e.g. Sullivan et al. 2014; Lin et al. 2014).

Certainly, a bird being caught in a spider web remains a rather rare event. I never encountered such a case in several years of birdwatching in tropical and subtropical regions, and 58 out of 68 people (85 %) who replied to my request for information also never encountered such a case. The remaining people had only encountered one case in their entire life except for Amar-Singh H., S. Pande, P. Round, G. Welch, and C.-t. Yao who each had encountered two (this does not include the multiple cases reported in the publications of Boedijn 1933; Manchi and Sankaran 2009; Kasambe et al. 2010). For any small bird, it is nevertheless a considerable risk because it carries the highest fitness cost, i.e. death. Combined with the facts that some spider species attempt to make their webs more visible to birds (Bruce et al. 2005; Walter and Elgar 2011), presumably to avoid collisions and the consequent damage to their webs, and that small bats are also at risk of spider predation (Nyffeler and Knörnschild 2013), the risks of collision, entanglement or death are probably high enough to facilitate the evolution of some avoidance behaviour in small birds. Even for larger bird species, there may be fitness costs; a 142 g Hooded Butcherbird (Cracticus cassicus) had to spend several minutes to preen itself after a collision with a spider web (Brooks 2012). To even begin to evaluate the magnitude of this risk, continuous video monitoring of spider webs would be required to establish collision frequencies, or captive birds could be used in experimental settings with spider webs.

Conclusions

This study adds to the previously presented evidence (Brooks 2012) that small birds face a risk of injury or death wherever large spiders build large spider webs. Although we can assume that these events are relatively rare compared to other risks of death (e.g. predation by hawks, snakes, or humans), what remains unknown is the frequency of these events, and thus the evolutionary pressure for the evolution of countermeasures in birds. Future studies should also elucidate if spiders carry a cost (damaged web) or a benefit (additional prey) from these events, and if their web building strategies have accordingly become adapted to account for these presumably rare events.

Declarations

Acknowledgements

I acknowledge the great help I received from my sources and translators, namely Amar-Singh H.S.S., Stephen Awoyemi, Anthony Bain, Daniel Brooks, Geoff Carey, Yun-Peng Chiang, Yu-Wen Emily Dai, Martin Hale, Yu-Cheng Hsu, Masayoshi Kamioki, Raju Kasambe, Manh Hung Le, Yong Ding Li, Wich’yanan Limparungpatthanakij, Hui-Shan Lin, Ruey-Shing Lin, Miyuki Mashiko, Satish Pande, Merrin Pearse, Sander Pieterse, Himanshu Rathore, Philip Round, Richard Thomas, Bas van Balen, Geoff Welch, Martin Williams, Jian-Long Wu, Tsai-Yu Wu, Cheng-te Yao, and Barure Nirmala of the Bombay Natural History Society library, all of whom I thank profusely. I also greatly thank Nancy Greig, Mark Harvey, Peter Jäger, Matjaž Kuntner and Adalberto Santos for spider identifications, and Peter Capainolo, Anita Gamauf, Paul Sweet, Till Töpfer, Tom Trombone, and Hein van Grouw for helping to obtain measurements from bird specimens. I also thank two anonymous reviewers for insightful comments.

Competing interests

The author declares that he has no competing interests.

Funding

I acknowledge financial support from Taipei Medical University through a SEED Grant.

Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.

Authors’ Affiliations

(1)
Master Program in Global Health and Development, College of Public Health, Taipei Medical University

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