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Lili Sun, Hongyan Yang, Xiuyuan Lu, Ting Fu, Jia Guo, Sicheng Ren, Waner Liang, Qing Chen, Dongming Li, Theunis Piersma, Nicola Crockford, Yifei Jia, Guangchun Lei. 2025: Shallow water habitats provide high-quality foraging environments for the Spoon-billed Sandpiper at a critical staging site. Avian Research, 16(1): 100233. DOI: 10.1016/j.avrs.2025.100233
Citation: Lili Sun, Hongyan Yang, Xiuyuan Lu, Ting Fu, Jia Guo, Sicheng Ren, Waner Liang, Qing Chen, Dongming Li, Theunis Piersma, Nicola Crockford, Yifei Jia, Guangchun Lei. 2025: Shallow water habitats provide high-quality foraging environments for the Spoon-billed Sandpiper at a critical staging site. Avian Research, 16(1): 100233. DOI: 10.1016/j.avrs.2025.100233
shu

Shallow water habitats provide high-quality foraging environments for the Spoon-billed Sandpiper at a critical staging site

  • a.

    School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China

  • b.

    Center for East Asian-Australasian Flyway Studies, Beijing Forestry University, Beijing 100083, China

  • c.

    Shenzhen Mangrove Wetlands Conservation Foundation, Shenzhen 518000, China

  • d.

    Dongtai Coastal Wetland Tourism and Resort Economic Zone Management Committee, Yancheng 224237, China

  • e.

    Rudi Drent Chair in Global Flyway Ecology at 'BirdEyes - Centre for Global Ecological Change' at the University of Groningen, Groningen, the Netherlands

  • f.

    NIOZ Royal Netherlands Institute for Sea Research, Texel, the Netherlands

  • g.

    RSPB Global Policy, Cambridge, UK

Funds: 

the National Natural Science Foundation of China 42361144873

Yellow Sea Wetland Project HHSDKT202312

the "Saving Spoon-billed Sandpiper" project of Shenzhen Mangrove Wetlands Conservation Foundation (MCF) 

More Information
  • Corresponding author:

    School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China. E-mail address: jiayifei@bjfu.edu.cn (Y. Jia)

    School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China. E-mail address: guangchun.lei@foxmail.com (G. Lei)

  • 1 These authors contributed equally to this work.

  • Received Date: 21 Oct 2024
  • Rev Recd Date: 17 Feb 2025
  • Accepted Date: 23 Feb 2025
  • Available Online: 27 Jun 2025
  • Publish Date: 27 Feb 2025
    Graphical Abstract
    • Abstract

  • Abundant food supply is crucial for the survival of long-distance migratory birds. The continued population decline of the Spoon-billed Sandpiper (Calidris pygmeae), a critically endangered shorebird, is primarily attributed to habitat loss and degradation. However, significant gaps remain in research on their diet and foraging habitat selection, limiting effective conservation and restoration efforts. In this study, we investigated the composition of macrobenthic communities, analyzing habitat and prey selection at the main foraging area of SBS in Tiaozini, Jiangsu Province of eastern China—their most critical staging site during southward migration. Our findings revealed 25 species of macrobenthos in foraging areas, with mobile epibenthos comprising the largest group by biomass, accounting for 73%, and having higher density and biomass nearshore. Observations of foraging Spoon-billed Sandpiper individuals indicated that shallow water habitats were their preferred foraging environments, where mobile epibenthos, which thrive in these habitats after the tide recedes, made up 81% of their biomass intake. We propose that shallow water habitats in intertidal mudflats serve as essential refuges for mobile epibenthos after tidal retreat, thereby providing Spoon-billed Sandpipers with access to high-quality food resources. Habitat protection efforts should prioritize habitats harboring extensive microhabitats with shallow water, especially the nearshore area, and further research is needed to explore the mechanisms underlying the formation of these microhabitats, with the ultimate goal of restoring more high-quality habitats for the Spoon-billed Sandpiper.

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  • 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.

    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 SwiftletCollocalia esculentaNp8.0950, 2, 3Great Nicobar island, Nicobar Islands, IndiaBefore 2010Manchi and Sankaran (2009)
    Edible-nest SwiftletCollocalia fuciphagaS10.71181, 1, 3Interview Island, Andaman Islands, IndiaJune 2006Manchi and Sankaran (2009)
    Asian Palm SwiftCypsiurus balasiensisNa9.21121, 1, 0Doi Kham, Chiang Mai Province, Thailand6 October 2014W. Limparungpatthanakij in litt. 2014
    Laughing DoveStreptopelia senegalensisNa80.0c138cOman~ October 2003Forsman (2003), Brooks (2012), D. Brooks in litt. 2014
    Pied FantailRhipidura javanicaS12.5821, 1, 0Near U Minh Thuong National Park, Kien Giang Province, Vietnam9 August 2008M. Le in litt. 2014
    Black-naped MonarchHypothymis azureaNp11.369Sanjay Gandhi National Park, Mumbai, IndiaOctober 1996Andheria(1998, 1999)
    Spotted FlycatcherMuscicapa striataS14.0c80cMain Kaleh Reserve, Iran1972Doberski (1973)
    Grey-streaked FlycatcherMuscicapa griseistictaS15.183Taiwan AreaSeptember 2008http://tinyurl.com/spider-tw2
    Grey-streaked FlycatcherMuscicapa griseistictaNp15.1833, 5, 8Iriomote Island, Japan5 October 2008http://tinyurl.com/spider-jp6
    Asian Brown FlycatcherMuscicapa dauuricaS9.9661, 1, 0Thap Lan National Park, Nakhon Ratchhasima Province, Thailand14 November 1999P. Round in litt. 2014
    Asian Brown FlycatcherMuscicapa dauuricaS9.9661, 1, 0Po Toi Island, Hong Kong, ChinaAfter 2006G. Welch in litt. 2014
    Asian Brown FlycatcherMuscicapa dauuricaNp9.9661, 1, 0Po Toi Island, Hong Kong, China8 September 2011M. Hale and G. Welch in litt. 2014
    Hill Blue FlycatcherCyornis banyumasNp14.5671, 1, 0Bukit Larut, Perak State, MalaysiaUnknownAnonymous (1999b)
    Great TitaParus majorNp15.5591, 0, 0Komesu, Itoman City, Okinawa Island, Japan10 August 2011http://tinyurl.com/spider-jp1, http://tinyurl.com/spider-jp2
    Dusky Crag MartinbHirundo concolorPf13.0981, 1, 0Shevaroys (=Servarayan) Hills near Salem, Tamil NaduBefore 1889Morris (1889) and Anonymous (1999a)
    Light-vented BulbulPycnonotus sinensisNp29.785Majia, Pingtung, Taiwan AreaMarch 2004http://tinyurl.com/spider-tw3
    Styan's BulbulPycnonotus taivanusNp26.284Guangfu, Hualien, Taiwan Area18 June 2010http://tinyurl.com/spider-tw4
    Yellow-vented BulbulPycnonotus goiavierNp27.8821, 4, 6Kledang-Sayong Forest Reserve, Ipoh, Perak, Malaysia11 February 2014Amar-Singh(2014a, b), Amar-Singh H. in litt. 2014
    Buff-vented BulbulIole olivaceaS24.5891, 1, 0Near Ban Bang Khram, Khlong Thom District, Krabi (area also known as Khao Nor Chuchi), Thailand7 August 2013P. Round in litt. 2014
    Brown-eared BulbulIxos amaurotisNp70.91161, 1, 0Tokunoshima Island, Japan1 August 2010http://tinyurl.com/spider-jp3
    Plain PriniaPrinia inornataNp8.249Tadoba Andhari Tiger Reserve, Chandrapur district, Maharashtra, IndiaOctober 1998Anonymous (1999a)
    Plain PriniaPrinia inornataNp8.249Melghat Tiger Reserve, northern part of Amravati District of Maharashtra State, IndiaBefore 2005Pande et al. (2004)
    Plain PriniaPrinia inornataS8.249Western Ghats, Maharashtra, IndiaUnknownS. Pande in litt. 2015
    Plain PriniaPrinia inornataS8.249Taiwan AreaSeptember 2008http://tinyurl.com/spider-tw2
    Oriental White-eyeZosterops palpebrosusNa8.6511, 1, 5Sungei Buloh Wetland Reserve, Singapore30 April 2012Ong(2012a, b)
    Japanese White-eyeZosterops japonicusNp11.353Mong Tseng Tsuen (near Tsim Bei Tsui), Hong Kong, China22 August 2004So (2005)
    Japanese White-eyeZosterops japonicusNp11.353Keelung, Taiwan Area18 August 2005http://tinyurl.com/spider-tw5
    Japanese White-eyeZosterops japonicusNp11.353Taiwan AreaBefore October 2005http://tinyurl.com/spider-tw7
    Japanese White-eyeZosterops japonicusNp11.353Badouzi, Keelung, Taiwan Island13 August 2011http://tinyurl.com/spider-tw6
    Japanese White-eyeZosterops japonicusNp11.353Okinawa Island, JapanNovember 2012http://tinyurl.com/spider-jp5
    Lanceolated WarblerLocustella lanceolataS12.955E-Luan-Pi lighthouse, Kenting National Park, Pintung County, Taiwan Area14 October 1984TESRI# collection number w672, C.-t. Yao in litt. 2013
    Grasshopper WarblerLocustella naeviaN14.864Tungareshwar Wildlife Sanctuary, Maharashtra, India18 November 2006Kasambe et al. (2010)
    Common TailorbirdOrthotomus sutoriusN7.5432, 2, 5Mogarkasa Forest, Nagpur, Maharashtra, India13 November 2008Kasambe et al. (2010)
    Dark-necked TailorbirdOrthotomus atrogularisNp7.7381, 1, 0Kaeng Krachan National Park, Petchaburi Province, Thailand2012W. Limparungpatthanakij in litt. 2014
    Dusky WarblerPhylloscopus fuscatusS11.0c57cBeidahe, Hebei Province, ChinaBefore 2007D. Zetterström in litt. 2007 (D. Brooks in litt. 2014)
    Arctic WarblerPhylloscopus borealisNp10.065Yonaguni Island, Japan10 September 2008http://ameblo.jp/attacus/theme2-10004405518.html
    Arctic WarblerPhylloscopus borealisNp10.065Bitou Cape, New Taipei City municipality, Taiwan Area6 September 2011Y.-P. Chiang in litt. 2013‒2014
    Arctic WarblerPhylloscopus borealisNp10.065Pak Sha O, Hong Kong, China19 September 2015Geoff Carey in litt. 2015
    Greenish WarblerPhylloscopus trochiloidesN7.160Bandhavgarh National Park, Madhya Pradesh, India12 October 2007Kasambe et al. (2010)
    Greenish WarblerPhylloscopus trochiloidesN7.160Kanha National Park, Madhya Pradesh, India22 October 2008Kasambe et al. (2010)
    Buff-breasted BabblerPellorneum tickelliS17.1611, 1, 0Fraser's Hill, Pahang, Malaysia5-11 June 2010S. Pieterse in litt. 2014
    Taiwan YuhinaYuhina brunneicepsS12.262Taiwan AreaUnknownH.-S. Lin in litt. 2013
    Vinous-throated ParrotbillParadoxornis webbianusNp9.352Mountain Pinglin, Taichung City, Taiping District, Taiwan Area2007http://tinyurl.com/spider-tw1
    Brown-throated SunbirdAnthreptes malacensisS11.4661, 1, 3Ipoh City, Perak, Malaysia28 December 2007Amar-Singh(2014a, b), Amar-Singh H. in litt. 2014
    Eurasian Tree SparrowPasser montanusS23.066Luku, Nantao County, Taiwan Area1990sC.-t. Yao in litt. 2013
    Eurasian Tree SparrowPasser montanusNp23.066Taiwan AreaSummer 2004http://tinyurl.com/spider-tw9
    Eurasian Tree SparrowPasser montanusS23.066Taiwan AreaBefore August 2010http://tinyurl.com/spider-tw8
    Eurasian Tree SparrowPasser montanusNp23.066Jiji, Nantou County, Taiwan Area13 August 2013C.-t. Yao in litt. 2013
    White-rumped MuniaLonchura striataS11.348Taiwan AreaUnknownY.-C. Hsu in litt. 2013
    Munia spec.-Np--Bogor Botanical garden, Bogor, IndonesiaBefore 1934Boedijn (1933)
    Munia spec.-Np--probably near or in Bogor, IndonesiaBefore 1934Boedijn (1933)
    "Small birds"-Np--ThailandBefore 1933Bristowe (1932)
    Unidentified-Np--probably near or in Bogor, IndonesiaBefore 1934Boedijn (1933)
    Unidentified-Np--Cheung Sha, Lantau Island, Hong Kong, China8 October 2006Anonymous (2006)
    Unidentified-Np--Wang Tong River, Mui Wo, Lantau Island, Hong Kong, China12 October 2009M. Pearse in litt. 2015
    Unidentified-Np--Miyakojima Island, Japan16 October 2011http://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
     | Show Table
    DownLoad: CSV

    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)315.9 ± 13.7 (7.1‒80.0)68.9 ± 20.9 (38.0‒138.0)
    Species (n = 33)317.5 ± 16.2 (7.1‒80.0)73.1 ± 23.6 (38.0‒138.0)
    World
    Individuals (n = 114)1212.3 ± 10.8 (2.0‒80.0)63.3 ± 20.1 (37.0‒138.0)
    Species (n = 84)1213.5 ± 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
     | Show Table
    DownLoad: CSV

    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.

    Figure  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
    DownLoad: Full-Size Img PowerPoint

    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.

    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.

    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.

    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.

    The author declares that he has no competing interests.

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

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