Canchao YANG, Wei LIANG, Anton ANTONOV, Yan CAI, Bård G. STOKKE, Frode FOSSØY, Arne MOKSNES, Eivin RØSKAFT. 2012: Diversity of parasitic cuckoos and their hosts in China. Avian Research, 3(1): 9-32. DOI: 10.5122/cbirds.2012.0004
Citation: Canchao YANG, Wei LIANG, Anton ANTONOV, Yan CAI, Bård G. STOKKE, Frode FOSSØY, Arne MOKSNES, Eivin RØSKAFT. 2012: Diversity of parasitic cuckoos and their hosts in China. Avian Research, 3(1): 9-32. DOI: 10.5122/cbirds.2012.0004

Diversity of parasitic cuckoos and their hosts in China

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  • Corresponding author:

    Wei LIANG, E-mail: liangwei@hainnu.edu.cn

  • Received Date: 05 Jan 2012
  • Accepted Date: 27 Feb 2012
  • Available Online: 23 Apr 2023
  • In this exposé we provide the first review of host use by brood parasitic cuckoos in a multiple-cuckoo system in China, based on our own long-term field data and a compilation of observations obtained from the literature. In total, we found that 11 species of cuckoos utilized altogether 55 host species. These hosts belong to 15 families, in which Sylviidae, Turdidae and Timaliidae account for 22.6%, 20.8% and 17.0% of parasitism records, respectively. The Common Cuckoo (Cuculus canorus) had the widest range of host species, accounting for 45.5% of the total number of parasitized species (25 in 10 families) of all parasitism records and is the most frequent brood parasite in the country. Cuckoo species differed in their egg coloration and the extent of egg polymorphism with most of them, e.g. the Common Cuckoo, the Lesser Cuckoo (C. poliocephalus) and the Plaintive Cuckoo (Cacomantis merulinus) laying well mimetic eggs with respect to their hosts based on human being's visual observations, while others such as the Large Hawk-cuckoo (C. sparverioides), the Himalayan Cuckoo (C. saturatus) and the Asian Emerald Cuckoo (Chrysococcyx maculatus) usually laid non-mimetic eggs. The use of cuckoo hosts and egg color variation in China are compared with those in other parts of their ranges in Asia.

  • Cuckoo-host systems are textbook examples of co-evolutionary arms races, although most of its research has focused on the Common Cuckoo (Cuculus canorus) and its hosts in Europe, where it is also the only brood parasite in most of the region. On the other hand, there are a number of brood parasitic cuckoo species breeding in sympatry in Asia and many of these have drawn very little attention (Rothstein and Robinson, 1998; Payne, 2005). So far, at least 17 cuckoo species, belonging to six genera, have been recorded in China (MacKinnon and Phillipps, 1999; Zheng, 2011; Appendix 1, see also Fig. 1). For many of these cuckoo species, there is little information on their breeding biology and host use (Payne, 2005).

    Figure  1.  Distribution and richness of breeding cuckoos in China. Numbers near the name of the provinces refer to the cuckoo species breeding in each province. Only residents or summer breeders are included.

    In our investigation we combined our long-term field data, an extensive review of mostly local papers as well as pictures of parasitized nests or cuckoo fledglings attended by hosts, taken by volunteer photographers in order to provide a detailed description of cuckoos and their host use in a multiple-cuckoo system in China. We also provide basic descriptions of egg coloration and egg morphs of the different cuckoos and their hosts.

    Our parasitism records were obtained from 19 provinces in China, including Heilongjiang, Taiwan, Hong Kong and Xizang (Tibet). We employed three approaches to compile the dataset. First, we monitored the occurrence of parasitism in the nests of various passerines at our study area of Kuankuoshui Nature Reserve (28°06′–28°19′N, 107°02′–107°14′E), Guizhou Province, during 1999–2011. Egg color was described based on human eyes whenever possible. Cuckoo species identification was accomplished by both morphology and DNA analysis of blood samples from adult cuckoos and nestlings, by sequencing the Cytb and ND2 genes of cuckoo samples. Second, we supplemented our dataset by including data on parasitism records from all existing published sources in local journals on cuckoos and their hosts in China. Third, we included data from photographs of cuckoo chicks or fledglings attended by their foster parents taken opportunistically by birdwatchers in China (see supplementary figures). For these cases, an important point is host suitability. In this context we differentiated between which host species successfully raised a cuckoo chick and which did not. We also summarized the distribution and richness of breeding parasitic cuckoo species in all provinces of China.

    We obtained 142 parasitism records of 11 cuckoo species (out of all 17 species in China, i.e., 64.7%) from 55 host species, with the hosts of six cuckoo species still unknown in China (Table 1, Appendix 1). Host species were diverse passerines, belonging to 15 families, including Motacillidae, Pycnonotidae, Laniidae, Dicruridae, Sturnidae, Corvidae, Troglodytidae, Turdidae, Muscicapidae, Timaliidae, Paradoxornithidae, Emberizidae, Fringillidae, Cisticolidae and Sylviidae. The most extensively used families were Sylviidae, Turdidae and Timaliidae which, respectively, accounted for 22.6%, 20.8% and 17.0% of the parasitism records. Overall, some of the cuckoo species, such as the Common Cuckoo, the Indian Cuckoo (Cuculus micropterus), the Himalayan Cuckoo (C. saturatus) and the Oriental Cuckoo (C. optanus) use more and taxonomically diverse host species and seem to be generalist parasites, while others, such as the Lesser Cuckoo (C. poliocephalus), the Plaintive Cuckoo (Cacomantis merulinus) and the Asian Emerald Cuckoo (Chrysococcyx maculatus) are specialized in their host use, exploiting a more restricted range of related host species, as well as having a special nest design (domed nests) (Table 2). The Common Cuckoo seems to be the most catholic in its host use since as many as 45.5% (25/55 species, 10 families) of the recorded host species were its hosts (Table 1). However, Common Cuckoo parasitism is also over-represented in our dataset, where it accounts for as much as 40.1% (57/142) of parasitism records.

    Table  1.  Parasitism records of cuckoos and their hosts in China
    Cuckoo species Number of host species Number of parasitism nests Number of cuckoo eggs recorded Number of cuckoo nestlings recorded Number of cuckoo egg morphs
    Chestnut-winged Cuckoo (Clamator coromandus) 4 10 3 7 1
    Large Hawk-cuckoo (Cuculus sparverioides) 7 15 15 1 2
    Indian Cuckoo (Cuculus micropterus) 6 9 4 5 1
    Common Cuckoo (Cuculus canorus) 24 57 22 35 10
    Himalayan Cuckoo (Cuculus saturatus) 5 8 6 2 1
    Oriental Cuckoo (Cuculus optatus) 8 9 5 4 2
    Lesser Cuckoo (Cuculus poliocephalus) 4 11 6 5 2
    Plaintive Cuckoo (Cacomantis merulinus) 1 8 3 5 1
    Asian Emerald (Cuckoo Chrysococcyx maculatus) 4 10 2 8 1
    Asian Drongo Cuckoo (Surniculus lugubris) 1 1 1
    Asian Koel (Eudynamys scolopacea) 4 7 2 5 1
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    Table  2.  Host use by parasitic cuckoos in China
    Host name Nest type CWC LHC IC CC HC OC LC PC AEC ADC AK Total
    Rufous-tailed Shrike (Lanius isabellinus) Shrub, open 3 3
    Brown Shrike (Lanius cristatus) Shrub, open 3 3
    Grey-backed Shrike (Lanius tephronotus) Shrub, open 1 1
    Red-billed Blue Magpie (Urocissa erythrorhyncha) Tree, open 2 2
    Azure-winged Magpie (Cyanopica cyanus) Tree, open 3 1 4
    Magpie (Pica pica) Tree, open 1 2 3
    Black Drongo (Dicrurus macrocercus) Tree, open 1 1
    Eurasian Blackbird (Turdus merula) Tree, open 1 1
    Bluethroat (Luscinia svecica) Ground, open 1 1
    Slaty-backed Forktail (Enicurus schistaceus) Ground, semi-open 1 1
    Lesser Shortwing (Brachypteryx leucophrys) Tree, open 1 1
    Black Redstart (Phoenicurus ochruros) Semi-open 2 2
    Daurian Redstart (Phoenicurus auroreus) Hole, semi-open 8 8
    Plumbeous Water Redstart (Rhyacornis fuliginosus) Semi-open 1 1
    White-bellied Redstart (Hodgsonius phaenicuroides) Shrub, open 2 2
    Oriental Magpie Robin (Copsychus saularis) Hole 1 1
    Common Stonechat (Saxicola torquata) Shrub, open 1 1 2
    Grey Bushchat (Saxicola ferreus) Ground, semi-open 2 2
    Black-collared Starling (Sturnus nigricollis) Semi-open 2 2
    Light-vented Bulbul (Pycnonotus sinensis) Shrub, open 1 1
    Collared Finchbill (Spizixos semitorques) Shrub, open 1 1
    Blue-and-white Flycatcher (Cyanoptila cyanomelana) Tree hole 1 1 2
    Brownish-flanked Bush Warbler (Cettia fortipes) Shrub, semi-open 1 9 10
    Aberrant Bush Warbler (Cettia flavolivacea) Shrub, semi-open - 1 1
    Common Tailorbird (Orthotomus sutorius) Tree, open - 8 8
    Black-browed Reed Warbler (Acrocephalus bistrigiceps) Shrub, open 1 1
    Oriental Reed Warbler (Acrocephalus orientalis) Shrub, open 2 4 6
    Buff-throated Warbler (Phylloscopus subaffinis) Ground, domed 1 1
    Humes' Warbler (Phylloscopus humei) Ground, domed 1 1
    Pale-legged Leaf Warbler (Phylloscopus tenellipes) Ground, domed 2 1 3
    Eastern Crowned Warbler (Phylloscopus coronatus) Ground, domed 1 1
    Blyth's Leaf Warbler (Phylloscopus reguloides) Ground, domed 3 1 4
    Bianchi's Warbler (Seicercus valentine) Ground, domed 7 7
    Chestnut-crowned Warbler (Seicercus castaniceps) Ground, semi-open 1 1
    Yellow-bellied Prinia (Prinia flaviventris) Shrub, open 2 2
    Zitting Cisticola (Cisticola juncidis) Shrub, semi-open 1 1
    Masked Laughingthrush (Garrulax perspicillatus) Shrub, open 1 1 2
    Greater Necklaced Laughingthrush (Garrulax pectoralis) Shrub, open 5 5
    Hwamei (Garrulax canorus) Ground, open 2 2 4
    White-browed Laughingthrush (Garrulax sannio) Shrub, open 4 4
    Elliot's Laughingthrush (Garrulax elliotii) Shrub, open 1 1
    Grey-cheeked Fulvetta (Alcippe morrisonia) Shrub, open 1 1
    Spot-breasted Scimitar Babbler (Pomatorhinus erythrocnemis) Ground, semi-open 1 1
    Rufous-capped Babbler (Stachyris ruficeps) Shrub, semi-open 1 1
    Chinese Babax (Babax lanceolatus) Shrub, open 5 5
    Reed Parrotbill (Paradoxornis heudei) Shrub, open 1 1
    Vinous-throated Parrotbill (Paradoxornis webbianus) Shrub, open 2 2
    Ashy-throated Parrotbill (Paradoxornis alphonsianus) Shrub, open 11 11
    Wren (Troglodytes troglodytes) Semi-open 1 1
    White Wagtail (Motacilla alba) Semi-open 2 1 3
    Richard's Pipit (Anthus richardi) Shrub, open 1 1
    Desert Finch (Rhodospiza obsoleta) Tree, open 1 1
    Black-faced Bunting (Emberiza spodocephala) Ground, open 2 2
    Jankowski's Bunting (Emberiza jankowskii) Ground, open 2 2
    Yellow-throated Bunting (Emberiza elegans) Ground, open 2 2
    Total number of species 2 7 5 25 5 8 4 1 4 1 4 NA
    Total number of records 7 15 8 57 8 9 12 8 10 1 7 142
    Note: abbreviation of cuckoo species: CWC, Chestnut-winged Cuckoo; LHC, Large Hawk-cuckoo; IC, Indian Cuckoo; CC, Common Cuckoo; HC, Himalayan Cuckoo; OC, Oriental Cuckoo; LC, Lesser Cuckoo; PC, Plaintive Cuckoo; AEC, Asian Emerald Cuckoo; ADC, Asian Drongo Cuckoo; AK, Asian Koel. Numbers in the table refer to the times of parasitism records.
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    The highest density of cuckoos is found in the southern parts of the country (Fig. 1). The provinces with the highest richness of breeding cuckoos in China are Guangxi (13 species), Yunnan (12 species), Sichuan (12 species), Guizhou (11 species), Jiangsu (11 species) and Hainan (11 species). The northern provinces, such as Qinghai, Xinjiang and Ningxia have the lowest cuckoo richness, with typically one or two breeding cuckoo species. Seven cuckoo species breed in Xizang; however, five of them are breeding in the southeast of Xizang near the borders with Sichuan and Yunnan.

    The data suggests little overlap in host use of the different species, since only 12.7% (7/55) of all the host species were hosts of more than one cuckoo species (two in all the cases).

    Based on our visual observations, we conclude that most of the cuckoo species lay more or less mimetic eggs, except for the Large Hawk-cuckoo (Cuculus sparverioides), the Himalayan Cuckoo and the Asian Emerald Cuckoo which lay poorly mimetic or non-mimetic eggs with most of their hosts (Appendix 1). In some cuckoo species, perfect egg mimicry is evident, e.g. the Common Cuckoo and its parrotbill hosts (Yang et al., 2010b). Marked egg mimicry is also the rule for the Lesser Cuckoo and its typically Cettia warbler hosts, with both parties laying uniform chocolate- or red-brown eggs (Yang et al., 2010a). However, these same cuckoo species may lay non-mimetic or poorly mimetic eggs with some other hosts; e.g. Common Cuckoos lay light cyan eggs with brown spots in contrast to the white and brown-spotted eggs of its Black-browed Reed Warbler (Acrocephalus bistrigiceps) host and the Lesser Cuckoo parasitizing Blue-and-white Flycatcher (Cyanoptila cyanomelana) lays light cyan eggs while the host eggs are white (Appendix 1). The Oriental Cuckoo lays highly- or intermediately mimetic eggs with their hosts, mainly leaf warbler hosts. The eggs of the closely related Himalayan Cuckoo appear to be mimetic only to the eggs of some Phylloscopus or Seicercus warblers, which may be its main hosts, but this cuckoo species frequently parasitizes a variety of other host species, differing greatly in egg coloration and hence its mimicry is often very poor.

    The eggs of the Large Hawk-cuckoo are typically non-mimetic, with whitish with brown spots in the nests of Elliot's Laughingthrush (Garrulax elliotii), the Masked Laughingthrush (G. perspicillatus), the Spot-breasted Scimitar Babbler (Pomatorhinus erythrocnemis) and the Eurasian Blackbird (Turdus merula). Of these four hosts, the first one lays light sky-blue eggs with brown spots and streaks, the second one lays light cyan eggs, the third one lays white egg and the last one lays light cyan eggs with brown spots. The Large Hawk-cuckoo lays white eggs in the nest of the Chinese Babax (Babax lanceolatus) which has deep blue eggs (Yang et al., 2012) and also parasitizes Hwamei (Garrulax canorus) in Guangxi Province, laying deep blue eggs as well.

    The existence of distinctive egg morphs is differently pronounced among the cuckoo species. The most polymorphic were the eggs of the Common Cuckoos, of which we could identify 10 egg morphs: 1) turquoise, unspotted, 2) pale blue, unspotted, 3) white, unspotted, 4) dark turquoise, unspotted, 5) turquoise with violet spots, 6) white with grey spots, 7) whitish with brown spots, 8) light cyan with brown spots, 9) pinkish white with sandy brown spots and 10) offwhite with olive spots. Most of these mimic to some extent the eggs of their diverse host species or genera, suggesting the existence of host races (gentes). At the other extreme are the Himalayan Cuckoo and the Asian Emerald Cuckoo which, according to our data, have only one egg morph, white with few tiny brown spots and yellowish-white with brown spots, respectively, regardless of the eggs of the host species. The hosts of the former lay white, chocolate-brown, offwhite with black spots and pink with violet spotted eggs while those of the latter lay white eggs.

    Our study provides the first synthesis of the available data on host use of cuckoos in China. Some cuckoo species appeared to be more generalists in host use, while others were specialized in their host use, parasitizing a few related host species. However, our results must be viewed as preliminary, since the research effort was not equally distributed among the different cuckoo species, their hosts and areas. Some cuckoos also have overlapping ranges of host species, but the data are insufficient to allow reliable estimates of such overlaps and their importance. With several cuckoo species breeding sympatrically, host use segregation as a result of interspecific competition is to be expected as indicated by evidence from Japan (e.g. Higuchi, 1998). Since there are many more species of coexisting cuckoos in China, we expect more pronounced host segregation, a phenomenon to be demonstrated by future studies.

    Most of the cuckoo species showed some degree of adaptation in egg phenotypes to their respective hosts as evidenced by egg mimicry in most cases, except for the Large Hawk-cuckoo and the Asian Emerald Cuckoo. There are some clear cases of very good egg matching, e.g. in the Lesser Cuckoo and their warbler hosts as well as in the Common Cuckoo and parrotbills (see also Yang et al., 2010a, 2010b), suggesting a long history of co-evolutionary interactions and advanced arms races. However, egg color descriptions in our study are based on human vision which differs dramatically from avian vision (e.g. Cherry and Bennett, 2001). Thus, more objective techniques to quantify egg color, such as visual modeling (e.g. Yang et al., 2010b; Antonov et al., 2011), are needed to establish how well the eggs of the different cuckoos match the eggs of their hosts from the perspective of avian vision. What remains to be shown is the existence of host races (gentes) within each cuckoo species. Furthermore, egg recognition and rejection abilities of the different hosts must be tested experimentally to understand the variation in egg mimicry among and within cuckoo species.

    According to our data, the Chestnut-winged Cuckoo (Clamator coromandus) in China parasitizes the Hwamei, a laughingthrush mainly found in southern parts of the country. We did not find any eggs of this cuckoo and have therefore no information on whether they mimic the deep blue eggs of its Hwamei host. In India and Sri Lanka, the Chestnut-winged Cuckoo uses other species of laughingthrush, often the Greater Necklaced Laughingthrush (Garrulax pectoralis) and the Lesser Necklaced Laughingthrush (G. monileger) (Osmaston, 1916; Mackenzie, 1918; Baker, 1942; Becking, 1981). Their eggs are pale blue and unmarked.

    In China, besides three species of common laughingthrushes, the Large Hawk-cuckoo (Cuculus sparverioides) also parasitizes the Chinese Babax (Babax lanceolatus), the Spot-breasted Scimitar Babbler (Pomatorhinus erythrocnemis) and the Eurasian Blackbird (Turdus merula). We recorded three cuckoo egg morphs, 1) unspotted white, 2) unspotted skyblue and 3) whitish brown, spotted. The plain egg white morph has only been described in China so far by Yang et al. (2012). Most of the Large Hawk-cuckoo breeding records outside China are from India and show that they parasitize laughingthrushes, the Himalayan Whistling-thrush (Myiophonus caeruleus), the Nepal Shortwing (Brachypteryx leucophrys), the Hoary Barwing (Actinodura nipalenisis), the Large Spiderhunter (Arachnothera magna) and other songbirds (Hume, 1873; Baker, 1906, 1934; Becking, 1981). Cuckoo eggs in India are of two morphs, 1) light brownish olive, seldom with darker specks and 2) blue, unspotted (Baker, 1906, 1907; Osmaston, 1912, 1916).

    We found that Chinese Common Cuckoos parasitize 3 species of parrotbills, 4 species of redstarts, 2 species of reed warblers, 2 species of buntings, 2 species of chats and 1 species each of bluethroats, cisticolas, flycatchers, magpies, leaf warblers, pipits, shrikes, wagtails and finches. The Common Cuckoo seems to be using all potential insectivorous songbirds suitable as hosts. This species has more than 100 host species in Europe and India, including flycatchers, chats, warblers, wagtails and buntings (Payne, 2005). In Japan, this cuckoo parasitizes at least 20 host species (Higuchi, 1998; Nakamura et al., 1998) and in Korea their hosts are the Daurian Redstart (Phoenicurus auroreus) and the Vinous-throated Parrotbill (Paradoxornis webbianus) (Yoon, 2000). Cuckoo eggs are polymorphic in coloration (6 morphs identified), and typically good mimics of the eggs of their respective host species, indicative of the existence of host-specific races as in Europe where at least 15 cuckoo egg morphs are recognized (Moksnes et al., 1995).

    The Himalayan and Oriental Cuckoos use predominantly warblers of the genera Phylloscopus and Seicercus as hosts (Payne, 2005), which we also found to be the case in China. Eggs are pale blue or white with fine black and brown or reddish stippling (Baker, 1906, 1927, 1934). Furthermore, they use the Yellow-bellied Prinia (Prinia flaviventris) which is widespread in the south of China as well as the White Wagtail (Motacilla alba) in Altai, the northernmost area of Xinjiang Autonomous Region. They also parasitize the Brownish-flanked Bush Warbler (Cettia fortipes) which is the main host of the Lesser Cuckoo. Other host species include the Slaty-backed Forktail (Enicurus schistaceus), the Yellow-throated Bunting (Emberiza elegans), the Collared Finchbill (Spizixos semitorques), the Stonechat (Saxicola torquata), the Lesser Shortwing (Brachypteryx leucophrys) and the Rufous-capped Babbler (Stachyris ruficeps). In China, cuckoo eggs found in these host species are white or white with brown spots.

    In China, Lesser Cuckoos use two species of bush warbles, the Winter Wren (Troglodytes troglodytes) and the Blue-and-white Flycatcher as hosts. Eggs are brown and light cyan, unspotted. In southeast Siberia in Ussuria, Japan and India, the Lesser Cuckoo parasitizes leaf warblers (Phylloscopus ijimae and P. occipitalis), bush warblers (Cettia diphone) and Winter Wrens (Payne, 2005). Cuckoo eggs are immaculately brown (Neufeldt, 1971; Higuchi, 1989, 1998), red or white (Baker, 1906, 1934).

    Indian Cuckoo hosts in China are the Azure-winged Magpie (Cyanopica cyanus), the Oriental Reed Warbler (Acrocephalus orientalis), the Light-vented Bulbul (Pycnonotus sinensis), the Black Drongo (Dicrurus macrocercus), the Eurasian Blackbird (Turdus merula) and the Common Stonechat. We recorded only two cuckoo egg morphs ― pinkish white or offwhite with firebrick markings. Indian Cuckoos in Russia parasitize the Brown Shrike (Lanius cristatus) (Neufeldt, 1966). In south-eastern Asia, they parasitize drongos and lay highly mimetic eggs (Becking 1981; Payne, 2005; Begum et al., 2012). In the Malay Peninsula at Selangor fledglings were observed being fed by a non-passerine bird, the Black-and-yellow Broadbill (Eurylaimus ochromalus) (Wells, 1999). Eggs are either white with reddish-brown markings or unmarked blue (Neufeldt, 1966; Baker, 1906, 1907, 1934, 1942).

    The only Plaintive Cuckoo host we found in China was the Common Tailorbird (Orthotomus sutorius) but in other parts of its range in Asia, many hosts have been recorded, such as the Yellow-bellied Prina (Prinia flaviventris) and the Black-necked Tailorbird (Orthotomus atrogularis) in the Malay Peninsula (Wells, 1999), the Olive-backed Tailorbird (O. sepium) in Java (Hellebrekers and Hoogerwerf, 1967; Beceking, 1981), prinias and tailorbirds in Borneo (Smythies, 1957, 1999) and the Grey-backed Tailorbird (O. derbianus) in the Philippines (Hornskov, 1995). In Sabah, the Indian Cuckoo is said to parasitize the Common Iora (Aegithina tiphia), the Streaky-breasted Spiderhunter (Arachnothera affinis) and the Yellow-bellied Prinia (Sheldon et al., 2001). Eggs are dull, light buff to green with light olive markings (Baker, 1906, 1907, 1934; Hoogerwerf, 1949; Hellebrekers and Hoogerwerf, 1967; Becking, 1981).

    The major host of the Asian Emerald Cuckoo in Guizhou Province of China is the Bianchi's Warbler (Seicercus valentini). This cuckoo species also parasitizes the Blyth's Leaf Warbler which is also host to the Himalayan Cuckoo. Other host species in China are the Chestnut-crowned Warbler (S. castaniceps) and the Buff-throated Warbler (Phylloscopus subaffinis). The color of the cuckoo eggs we recorded was offwhite with brown spots concentrated at the blunt end. The Asian Emerald Cuckoo hosts in India are the Yellow-backed Sunbird (Aethopyga siparaja), the Little Spiderhunter (A. longirostra) (Baker, 1919) and the Chestnut-crowned Warbler (Seicercus castaneiceps) (Stevens, 1925). The eggs are whitish, blotched with light brown or reddish brown (Baker, 1906, 1942; Harrison, 1969; Becking, 1981).

    The Asian Koel in China uses one species of starling, two species of magpies and one species of laughingthrush as hosts, while in south-eastern Asia it uses crows as common hosts (Begum et al., 2011a, b). From these hosts, the Black-billed Magpie (Pica pica) and the Masked Laughingthrush (Garrulax perspillatus) are also parasitized by the Large Hawk-cuckoo. With regard to the Asian Drongo Cuckoo, only one host species, the Grey-cheeked Fulvetta (Alcippe morrisonia), has been recorded so far in China.

    Parasitic cuckoos species and their hosts in Chin
    Cuckoo name Host name Nest contents Note contents Cuckoo egg mass (g) and size (mm) Cuckoo egg color Host egg mass (g) and size (mm) Host egg color Source
    Chestnut-winged Cuckoo (Clamator coromandus) Hwamei (Garrulax canorus) Nestling 1C+3H Blue Blue Jia et al., 2007
    Nestling 1C+3H Jia et al., 2007
    Egg Blue La Touche, 1931–34
    Masked Laughingthrush (Garrulax perspicillatus) Egg Blue La Touche, 1931–34
    Greater Necklaced Laughingthrush (Garrulax pectoralis) Nestling 1C Carey et al., 2001
    Nestling 1C Carey et al., 2001
    Nestling 1C Carey et al., 2001
    Nestling 1C Carey et al., 2001
    Nestling 2C+6H Blue Blue This study
    Oriental Magpie Robin (Copsychus saularis) Egg Blue Guangdong Institute of Entomology and Sun Yat-sen University, 1983
    Large Hawk-cuckoo (Cuculus sparverioides) Chinese Babax (Babax lanceolatus) Egg 1C+2H 6.6, 27.1 × 21.4 White 6.0, 27.8 × 20.6 Deep sky blue Yang et al., 2012
    Egg 1C+2H White Deep sky blue This study
    Egg 1C+1H White Deep sky blue This study
    Egg 1C+1H White Deep sky blue This study
    Egg 1C+2H White Deep sky blue This study
    White-browed Laughingthrush (Garrulax sannio) Egg 1C+4H 6.2, 27.0 × 20.6 White 5.7, 27.8 × 19.7 White Jiang et al., 2007
    Egg 1C+2H 7.5, 30.4 × 21.8 White 5.3, 26.1 × 20.0 White This study
    Egg 1C+1H White Blue This study
    Egg 1C+1H White Blue This study
    Elliot's Laughingthrush (Garrulax elliotii) Egg 4.5–5.0, 25.0 × 19.0–27.0 × 19.5 Offwhite with brown spots 5.8, 28.2 × 20.1 Light sky blue with brown spots and steaks Fan et al., 2000
    Masked Laughingthrush (Garrulax perspicillatus) Egg 4.5–5.0, 25.0 × 19.0–27.0 × 19.5 Offwhite with brown spots 5.6, 29.0 × 21.0 Light cyan Fan et al., 2000
    Spot-breasted Scimitar Babbler (Pomatorhinus erythrocnemis) Egg 4.5–5.0, 25.0 × 19.0–27.0 × 19.5 Offwhite with brown spots 6.5, 28.3 × 20.7 White Fan et al., 2000
    Hwamei (Garrulax canorus) Nestling 1C This study
    Egg 1C+2H White Blue This study
    Black-billed Magpie (Pica pica) Egg 2C+4H 4.7, 26.2 × 19.2 Olive grey with brown spots Cheng et al., 1991
    Indian Cuckoo (Cuculus micropterus) Azure-winged Magpie (Cyanopica cyanus) Nestling Yan et al., 1985
    Egg 4.0, 21.0 × 17.5–24.0 × 18.0 6.0, 28.1 × 20.2 Lightcyan with brown and purple spots Bu et al., 1999
    Fledgling This study
    Oriental Reed Warbler (Acrocephalus orientalis) Egg 4.0, 21.0 × 17.5–24.0 × 18.0 3.8, 22.4 × 16.0 Offwhite with olive spots Bu et al., 1999; Hao and Wang, 1992;
    Nestling Yan et al., 1985; Hao and Wang, 1992;
    Light-vented Bulbul (Pycnonotus sinensis) Nestling Yan et al., 1985; Liu and Long, 1986
    Black Drongo (Dicrurus macrocercus) Egg 1C+2H Pinkish white with firebrick spots 4.8, 24.7 × 18.6 Lightpink with brown spots Vaughan and Jones, 1913; Zhang, 2001
    Eurasian Blackbird (Turdus merula) Egg 1C+2H 6.8, 29.5 × 22.7 Offwhite with firebrick spots 7.2, 29.1 × 21.4 Lightcyan with brown spots Zhou et al., 2001
    Common Stonechat (Saxicola torquata) Nestling Cheng et al., 1991
    Common Cuckoo (Cuculus canorus) Ashy-throated Parrotbill (Paradoxornis alphonsianus) Egg 1C+4H 2.6, 20.8 × 16.2 Turquoise 1.3, 16.1 × 12.7 Turquoise Yang et al., 2010b
    Nestling 1C This study
    Nestling 1C This study
    Egg 1C+2H Turquoise Turquoise Yang et al., 2010b
    Egg 1C+3H Turquoise Turquoise Yang et al., 2010b
    Nestling 1C This study
    Nestling 1C This study
    Nestling 1C This study
    Nestling 1C This study
    Egg 1C 3.0, 21.4 × 16.2 Paleblue This study
    Egg 1C+2H 3.1, 21.6 × 16.2 White 1.2, 16.7 × 12.4 Paleblue This study
    Vinous-throated Parrotbill (Paradoxornis webbianus) Fledgling This study
    Egg Turquoise with violet spots Cheng, 1963, 1973
    Reed Parrotbill (Paradoxornis heudei) Egg 1C+2H Offwhite with olive spots Offwhite with olive spots This study
    Daurian Redstart (Phoenicurus auroreus) Nestling 1C This study
    Egg 1C+4H 2.8, 20.8 × 16.3 White with grey spots 1.7, 17.9 × 13.7 Pale turquoise with firebrick spots This study
    Egg 1C+5H White with grey spots White with firebrick spots This study
    Egg 1C+3H This study
    Nestling 1C This study
    Egg 1C+3H White with grey spots White with firebrick spots This study
    Nestling 1C This study
    Nestling 1C Cheng, 1963, 1973
    Black Redstart (Phoenicurus ochruros) Nestling 1C This study
    Fledgling This study
    White-bellied Redstart (Hodgsonius phaenicuroides) Egg 1C+3H 3.8, 25.9 × 17.9 Dark turquoise Dark turquoise This study
    Egg 1C+3H 3.8, 25.1 × 18.0 Dark turquoise Dark turquoise This study
    Plumbeous Water Redstart (Rhyacornis fuliginosus) Nestling 1C This study
    Bluethroat (Luscinia svecica) Nestling Qian and Zhang, 1965
    Common Stonechat (Saxicola torquata) Fledgling This study
    Grey Bushchat (Saxicola ferreus) Egg 1C+3H 2.7, 20.1 × 16.0 Dark turquoise 1.9, 17.6 × 14.4 Medium turquoise This study
    Egg 1C+4H This study
    White Wagtail (Motacilla alba) Fledgling This study
    Nestling 1C Li et al., 1985
    Richard's Pipit (Anthus richardi) Fledgling This study
    Oriental Reed Warbler (Acrocephalus orientalis) Nestling Lu, 1988; Hao and Wang, 1992
    Fledgling This study
    Fledgling This study
    Egg 4.4, 24.0 × 18.0 Offwhite with brown spots 3.8, 22.4 × 16.0 Offwhite with olive spots Tian et al., 1991
    Fledgling This study
    Black-browed Reed Warbler (Acrocephalus bistrigiceps) Egg 1C+4H 3.6, 25.0 × 17.3 Lightcyan with brown spots 1.3, 13.8 × 10.7 Offwhite with brown spots Liu et al., 1984
    Pale-legged Leaf Warbler (Phylloscopus tenellipes) Egg 1.8, 21.5 × 13.3 Pinkish white with sandy brown spots 1.5, 15.6 × 12.3 Pinkish white Gao, 2004
    Nestling Fu et al., 1984
    Zitting Cisticola (Cisticola juncidis) Egg Turquoise with violet spots Cheng et al., 1991
    Rufous-tailed Shrike (Lanius isabellinus) Fledgling This study
    Fledgling This study
    Fledgling This study
    Brown Shrike (Lanius cristatus) Eggs Qian and Zhang, 1965
    Fledgling This study
    Fledgling This study
    Grey-backed Shrike (Lanius tephronotus) Fledgling This study
    Blue-and-white Flycatcher (Cyanoptila cyanomelana) Nestling Fu et al., 1984
    Black-faced Bunting (Emberiza spodocephala) Fledgling This study
    Nestling Fu et al., 1984
    Jankowski's Bunting (Emberiza jankowskii) Fledgling This study
    Egg 1C+5H Offwhite with brown spots Wang et al., 2011
    Azure-winged Magpie (Cyanopica cyanus) Egg 1C+4H 5.4, 26.0 × 20.0 Lightcyan with brown spots 5.7, 27.4 × 19.3 Lightcyan with brown and purple spots Zhang, 1989
    Desert Finch (Rhodospiza obsoleta) Nestling Qian and Zhang, 1965
    Himalayan Cuckoo Cuculus saturatus) Blyth's Leaf Warbler (Phylloscopus reguloides) Egg 21.5 × 14.0 White with a few brown spots 1.0, 14.9 × 12.2 White La Touche, 1931–34
    Egg 1C+3H 2.1, 20.6 × 13.5 White with a few brown spots 1.1, 14.4 × 11.8 White Yang et al., 2011
    Egg 21.5 × 14.0 White with a few brown spots White La Touche, 1931–34
    Brownish-flanked Bush Warbler (Cettia fortipes) Egg 1C+1H 2.5, 23.4 × 14.4 White with a few brown spots 1.5, 17.7 × 13.0 Chocolate This study
    Slaty-backed Forktail (Enicurus schistaceus) Nestling La Touche, 1931–34
    Yellow-throated Bunting (Emberiza elegans) Egg 1C+2H 2.7, 22.8 × 14.9 White with a few brown spots 1.4, 16.4 × 13.1 Offwhite with black sptos This study
    Nestling Fu et al., 1984
    Collared Finchbill (Spizixos semitorques) Egg 1C+2H 3.0, 21.2 × 16.1 White with a few brown spots 3.3, 24.6 × 17.1 Pink with violet spots This study
    Oriental Cuckoo (Cuculus optatus) Common Stonechat (Saxicola torquata) Nestling La Touche, 1931–34
    Lesser Shortwing (Brachypteryx leucophrys) Egg 21.0 × 14.5 Turquoise with red spots La Touche, 1931–34
    Humes' Warbler (Phylloscopus humei) Egg 1C+3H 2.5, 20.7 × 14.0 White 1.1, 15.0 × 11.1 White Wang et al., 2004
    Pale-legged Leaf Warbler (Phylloscopus tenellipes) Nestling Fu et al., 1984
    Eastern Crowned Warbler (Phylloscopus coronatus) Nestling Gao et al., 1990
    White Wagtail (Motacilla alba) Fledgling This study
    Yellow-bellied Prinia (Prinia flaviventris) Egg 21.0 × 14.7 Pink with dense red spots Pink with dense red spots La Touche, 1931–34
    Egg White with brown spots Pink with dense red spots Zhang, 1980
    Rufous-capped Babbler (Stachyris ruficep) Egg 1C+2H Brown White Lin, 2008
    Lesser Cuckoo (Cuculus poliocephalus) Brownish-flanked Bush Warbler (Cettia fortipes) Egg 1C+3H 2.6, 22.1 × 15.2 Chocolate 1.5, 17.2 × 13.4 Chocolate Yang et al., 2010a
    Nestling Jiang et al., 2006
    Nestling This study
    Nestling This study
    Egg 1C+2H Chocolate Chocolate Yang et al., 2010a
    Egg 1C+3H Chocolate Chocolate Yang et al., 2010a
    Egg 1C+3H Chocolate Chocolate Yang et al., 2010a
    Egg Chocolate Chocolate This study
    Winter Wren (Troglodytes troglodyte) Nestling Li, 1985; Liu et al., 1988
    Blue-and-white Flycatcher (Cyanoptila cyanomelana) Egg 1C+2H Lightcyan 2.3, 20.0 × 15.0 White Zhao, 1985; Zhao and He, 1981
    Aberrant Bush Warbler (Cettia flavolivacea) Fledgling This study
    Plaintive Cuckoo (Cacomantis merulinus) Common Tailorbird (Orthotomus sutorius) Fledgling This study
    Egg 18.3 × 13.5 White with lightrufous spots White with lightrufous spots Vaughan and Jones, 1913
    Fledgling This study
    Nestling This study
    Nestling This study
    Egg 1C+3H Palegreen with lightrufous spots Palegreen with lightrufous spots This study
    Egg 1C+4H White with lightrufous spots White with lightrufous spots This study
    Fledgling This study
    Asian Emerald Cuckoo (Chrysococcyx maculatus) Bianchi's Warbler (Seicercus valentini) Nestling 1C 1.2, 15.8 × 12.0 White Yang et al., 2008
    Egg 1C+1H 1.5, 17.2 × 12.4 White with brown spots 1.3, 15.0 × 12.3 White This study
    Nestling 1C This study
    Nestling 1C This study
    Nestling 1C This study
    Nestling 1C This study
    Nestling 1C This study
    Chestnut-crowned Warbler (Seicercus castaniceps) Egg 1C+4H 1.3, 17.0 × 12.8 White with brown spots 0.9, 15.0 × 11.1 White This study
    Blyth's Leaf Warbler (Phylloscopus reguloides) Nestling 1C White This study
    Buff-throated Warbler (Phylloscopus subaffinis) Nestling White Li, 1985
    Asian Drongo Cuckoo (Surniculus lugubris) Grey-cheeked Fulvetta (Alcippe morrisonia) Fledgling This study
    Asian Koel (Eudynamys scolopacea) Black-collared Starling (Sturnus nigricollis) Egg 33.5 × 22.9 Sage green 29.4 × 21.7 Turquoise Vaughan and Jones, 1913
    Egg Carey et al., 2001
    Red-billed Blue Magpie (Urocissa erythrorhyncha) Nestling 1C+2H Li, 1985
    Nestling Lewthwaite, 1996
    Black-billed Magpie (Pica pica) Nestling 1C Carey et al., 2001
    Nestling 1C Carey et al., 2001
    Masked Laughingthrush (Garrulax perspicillatus) Nestling 1C Carey et al., 2001
    Pied Cuckoo (Clamator jacobinus) Unknown
    Common Hawk-cuckoo (Cuculus varius) Unknown
    Hodgson's Hawk-cuckoo (Cuculus nisicolor) Unknown 22.6 × 16.3 Olive brown with brown spots Ali and Ripley, 1974
    Northern Hawk-cuckoo (Cuculus hyperythrus) Unknown
    Banded Bay Cuckoo (Cacomantis sonneratii) Unknown
    Violet Cuckoo (Chrysococcyx xanthorhynchus) Unknown
     | Show Table
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    Supplementary figures

    Asian Koel (Eudynamys scolopacea) parasitizes Black-collared Starling (Sturnus nigricollis), Hong Kong (Photo by G.J. Carey)

    Chestnut-winged Cuckoo (Clamator coromandus) parasitizes Hwamei (Garrulax canorus), Guizhou (Photo by Chenxi Jia)

    Chestnut-winged Cuckoo (Clamator coromandus) parasitizes Hwamei (Garrulax canorus), Hubei (Photo by Qiang Ma)

    Large Hawk-cuckoo (Hierococcyx sparverioides) parasitizes White-browed Laughingthrush (Garrulax sannio), Guizhou (Photo by Canchao Yang)

    Large Hawk-cuckoo (Hierococcyx sparverioides) parasitizes Hwamei (Garrulax canorus), Guangxi (Photo by Aiwu Jiang)

    Common Cuckoo (Cuculus canorus) parasitizes Vinous-throated Parrotbill (Paradoxornis webbianus), Sichuan (Photo by Ming Zhang)

    Common Cuckoo (Cuculus canorus) parasitizes Reed Parrotbill (Paradoxornis heudei), Shandong (Photo by Donglai Li)

    Common Cuckoo (Cuculus canorus) parasitizes Daurian Redstart (Phoenicurus auroreus), Guizhou (Photo by Zhumei Li)

    Common Cuckoo (Cuculus canorus) parasitizes Daurian Redstart (Phoenicurus auroreus), Guizhou (Photo by Zhumei Li)

    Common Cuckoo (Cuculus canorus) parasitizes Plumbeous Water Redstart (Rhyacornis fuliginosus), Jiangsu (Photo by A. Lou)

    Common Cuckoo (Cuculus canorus) parasitizes Black Redstart (Phoenicurus ochruros), Gansu (Photo by Shuihua Chen)

    Common Cuckoo (Cuculus canorus) parasitizes White Wagtail (Motacilla alba), Guizhou (Photo by Xiaodong Liao)

    Common Cuckoo (Cuculus saturatus) parasitizes White Wagtail (Motacilla alba), Xinjiang (Photo by Ming Ma)

    Common Cuckoo (Cuculus canorus) parasitizes Oriental Reed Warbler (Acrocephalus orientalis), Heilongjiang (Photo by Suo Tan)

    Common Cuckoo (Cuculus canorus) parasitizes Oriental Reed Warbler (Acrocephalus orientalis), Liaoning (Photo by Xiaojun Wang)

    Common Cuckoo (Cuculus canorus) parasitizes Rufous-tailed Shrike (Lanius isabellinus), Xinjiang (Photo by Damo Gebi)

    Common Cuckoo (Cuculus canorus) parasitizes Rufous-tailed Shrike (Lanius isabellinus), Xinjiang (Photo by Bingchuan Yihao)

    Common Cuckoo (Cuculus canorus) parasitizes Rufous-tailed Shrike (Lanius isabellinus), Shanxi (Photo by Ke Kuan)

    Common Cuckoo (Cuculus canorus) parasitizes Grey-backed Shrike (Lanius tephronotus), Chongqing (Photo by Kacha Chongqing)

    Common Cuckoo (Cuculus canorus) parasitizes Brown Shrike (Lanius cristatus), Beijing (Photo by Sdm)

    Common Cuckoo (Cuculus canorus) parasitizes Brown Shrike (Lanius cristatus), Henan (Photo by Laogao Dongzhai)

    Common Cuckoo (Cuculus canorus) parasitizes Grey Bushchat (Saxicola ferrea), Guizhou (Photo by Canchao Yang)

    Common Cuckoo (Cuculus canorus) parasitizes Richard's Pipit (Anthus richardi), Qinghai (Photo by Suixing Tian)

    Common Cuckoo (Cuculus canorus) parasitizes Black-faced Bunting (Emberiza spodocephala), Qinghai (Photo by Cao Ting)

    Common Cuckoo (Cuculus canorus) parasitizes Jankowski's Bunting (Emberiza jankowskii), Heilongjiang (Photo by Jun Zhao)

    Common Cuckoo (Cuculus canorus) parasitizes Common Stonechat (Saxicola torquata), Guizhou (Photo by Hao Ai)

    Common Cuckoo (Cuculus canorus) parasitizes White-bellied Redstart (Hodgsonius phaenicuroides), Tibet (Photo by Xin Lu)

    Lesser Cuckoo (Cuculus poliocephalus) parasitizes Aberrant Bush Warbler (Cettia flavolivacea), Gansu (Photo by Suixing Tian)

    Plaintive Cuckoo (Cacomantis merulinus) parasitizes Common Tailorbird (Orthotomus sutorius), Guangdong (Photo by Zga)

    Plaintive Cuckoo (Cacomantis merulinus) parasitizes Common Tailorbird (Orthotomus sutorius), Guangdong (Photo by Lool)

    Plaintive Cuckoo (Cacomantis merulinus) parasitizes Common Tailorbird (Orthotomus sutorius), Guangxi (Photo by Cainiao Zishen)

    Drongo Cuckoo (Surniculus lugubris) parasitizesd Grey-cheeked Fulvetta (Alcippe morrisonia), Fujian (Photo by Xiaogang Da)

    We are particularly thankful to all the bird watching enthusiasts who offered photographs of cuckoos and their hosts. We thank Jeremy Wilson, the editor of Ibis, Andrew MacColl, associate editor of Ibis and two anonymous referees of Ibis for providing helpful comments on this manuscript. This work was supported by the National Natural Science Foundation of China (No. 31071938, 31101646), Key Project of Chinese Ministry of Education (No. 212136), the China Postdoctoral Science Foundation funded project (20110490967) and the Program for New Century Excellent Talents in University (NCET-10-0111). We thank the Forestry Department of Guizhou Province and the Kuankuoshui National Nature Reserve for support and permission to carry out this study, as well as X. Guo, L. Wang, X. Xu, N. Wang and T. Su for assistance with the field work.

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