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Praveen & Lowther: Avian brood in South 103

Avian brood parasitism in Praveen J & Peter Lowther

Praveen, J., & Lowther, P., 2020. Avian brood parasitism in South Asia. Indian 16 (4): 103–119. Praveen J, Villa #5, Embassy Homes, Mudavanmugal, Poojappura P.O. Thiruvananthapuram, Kerala, . E-mail: [email protected] [Corresponding author] Peter Lowther, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL, 60606, USA. E-mail: [email protected] Manuscript received on 14 August 2020.

Abstract Brood parasitism is an evolutionary in which certain , known as brood parasites, have evolved to use other species, known as hosts, to rear their young. While a lot of work to identify species from India and South Asia has been based on collections from the first half of the twentieth century, this has not been followed up with extensive field studies except for certain species. Here we compile information from published literature, online websites, and social media to appraise the current knowledge in this topic, specifically focusing on documentation of hosts feeding parasites. We created an online catalog of 287 instances of host-parasite pairs, covering 15 of the 18 species of brood parasites, across 46 host species. The most commonly observed pair is the -Asian , followed by babblers with Pied , and Common Hawk Cuckoo. We add ten new host-parasite pairs for South Asia, nine of them new to science. Our work did not find any hosts for Chestnut-winged Cuckoo, Whistling Hawk Cuckoo, and Himalayan Cuckoo from South Asia.

We dedicate this work to Vijay Cavale (1961–2020) who inspired Indian photographers to share their work online.

Introduction where the hosts were documented accompanying, or even Brood parasitism is a fascinating feature of avian biology. In feeding, an identifiable chick of a parasite. In that respect, we this evolutionary adaptation, certain species, known as brood trawled online image portals (Macaulay Library, iNaturalist, and parasites, lay their in the of other species, known as Oriental Bird Images), and social media groups (Facebook) to hosts, which then rear the parasitic young, and serve as foster find reliable evidences for host-parasite behavior. In addition, parents. These birds incubate the parasite eggs along with their a short online survey was circulated to collate such instances own, and when the parasite egg hatches, they feed the young, from the public. We downloaded data up to and including June more often than not at the cost of their own reproductive success. 2020, from eBird, for all cuckoo species, filtered out records with In South Asia, brood parasitism is restricted to several species of species comments, and manually went through the comments (Cuculidae: Cuculinae), and one species of for records without relevant media. This altogether resulted in a (Indicatoridae). catalogue of 287 host-parasite entries after removing duplicates Historically, the documentation of avian brood parasitism in [See Supplementary File], most of them (~89%) from the past South Asia primarily followed identification of eggs in clutches decade. We have attempted to include cases where a specimen, (Hume 1890; Baker 1906a,b, 1907a,b, 1934, 1942). However, or a photograph, exists as documentation of the host-chick pair, or this method was not without its flaws as identification of cuckoo at least field notes that are clear. Additionally, where identification species from their eggs is fraught with difficulties. Becking of either the host or the parasite is straightforward, we also (1981) pointed out the dubious of egg identification depend on the reliability of the observer for correct identification. in Baker’s egg collection, and his host-parasite assignments. In all cases of host-parasite pairs, we qualify the observation by Becking, additionally, analysed the eggs using scanning electron the highest evidence type that is available: (S) studies published, microscopy and biochemistry techniques to identify hosts and (M) supporting media and (O) field observations. While general parasites. He also pointed out that not all hosts are viable for the references on this topic cover the species’ entire range (Payne parasite and some eggs would never hatch, and if they indeed 2005; Erritzøe et al. 2012), or are outside South Asia (Yang et do, the chick may not survive. Hence, he suggested that future al. 2012), we list references only if specific documentation was workers should let the cuckoos fledge and study them rather made within South Asia. For common parasite-host pairs, we do than collect the eggs and clutch. not provide exhaustive lists of all references, but limit them to Baker’s ornithological work, most particularly his egg collection, extensive studies. Secondary references like Ali & Ripley (1987) has been a source of controversy that makes it difficult to accept or Billerman et al. (2020) do not augment the information and his collections as the sole evidence for any ornithological discovery hence are not cited specifically. We do not use the list of host/ (see Walters 2006; Rasmussen & Anderton 2012). While Becking victim species from Baker (1942) as definitive but include its taxa (1981) made a substantial contribution in clarifying the air, we only for completeness. take this opportunity to supplement and establish norms and It is worth defining two terms used in brood parasitism. Those exceptions using the best available and latest information. species known to have received a parasite egg in their nests, but are not necessarily known to have reared the parasite young Methodology we term victims. Species that we term hosts are those victims We depart from the traditional methodology of host-parasite known to have reared a parasite young successfully (see also assignment using egg collection, and list only those instances Johnsgard 1997). A brood parasite host table, based solely on 104 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

eggs/clutches, is only capable of identifying victims, though many map is not representative for those areas, however, it is clear victims (unproven hosts) may eventually turn out to be true hosts that the eastern have the highest diversity of brood as more observations and studies accumulate. Victims are easily parasites, followed by the western and central Himalayas, the identified during a single visit to a containing the parasite , and . egg (s) or young; however, identifying hosts require follow up In this paper, we cover seventeen species of cuckoos, and visits to verify successful rearing of the parasite young. It may be one species of honeyguide occurring in our region with tables and mentioned here that Baker’s (1942) is, essentially, a victim list and maps where warranted. The centroids of the districts are overlaid not a host list. Through this work, we attempt to circumvent this on the distribution maps downloaded from State of India’s Birds problem by bringing in cases where hosts were recorded feeding (https://www.stateofindiasbirds.in/). We do not include the the young parasite. Our method too is not flawless, as many of vagrant Horsfield’s Bronze Cuckoo basalis. the host-parasite documentations are made opportunistically, and do not assert the parasite young to have been successfully raised Results by the attending adult, but it is a step in that direction. A caveat worth remembering here are the rare cases where certain non- Chestnut-winged Cuckoo coromandus host species may offer parental care, at least temporarily, masking This is a monotypic cuckoo breeding in the Himalayan and north- the true host from the observer (Sealy & Lorenzana 1997). There eastern Indian foothills, and wintering in Sri Lanka and south- is one such case from our region where a Grey-bellied Cuckoo western India. Laughing-thrushes of the genera Grammatoptila, passerinus raised by (Nectariniidae) was , , and are considered its also actively fed by Common Orthotomus sinensis host species. However, no documentation from our region (Suter 1945). Misguided courtship, by adult cuckoos towards exists of any host species feeding a young Chestnut-winged juveniles, has also been reported in a few species (Kuriakose Cuckoo. The only direct observations of brood parasitism were 2012; Waiker & Sivanandan 2020). of a Chestnut-winged Cuckoo that was ‘noosed’ over a nest of Cuckoos are widely distributed in South Asia but regional a Lesser Necklaced Laughingthrush G. monileger, which it was diversity and abundance varies a lot. Based on eBird data presumably visiting to drop its egg and, that of a bird that was uploaded up to (and including) June 2020, the diversity shot as it left a nest of the same laughingthrush species (Baker (=number of species of brood parasites) and abundance 1907a). Based on Baker’s egg collection, Becking (1981) listed (frequency as percentage of complete lists that has at least one the Greater Necklaced- P. pectoralis and the Lesser Necklaced brood parasite) in South Asia is shown in Fig. 1 at a square with as common hosts, followed by the Striated resolution of 0.5° x 0.5° [c.56x56 km]. Squares with less than Laughingthrush G. striata, Rufous-vented Laughingthrush±1 P. 10 complete lists are omitted for frequency analysis. Admittedly, gularis, Blue-winged Laughingthrush T. squamatum, and Grey- there is insufficient eBird coverage in , , and sided Laughingthrush P. caerulatus. This list is likely to be correct certain parts of east-central and northern India, and hence the based on Becking’s extensive laboratory studies.

Fig. 1. The number of species and frequency of occurrence (as percentage of complete lists) of brood parasites in South Asia at square size 0.5° x 0.5° [c.56x56 km]; generated from data download from www..org/india. Squares with < 10 complete lists omitted for frequency analysis.

1 Listed under P. delesserti in Becking (1981), and assumed to be P. [d]. gularis, based on range. Praveen & Lowther: Avian brood parasitism in South Asia 105

However, Baker (1942) included many more victims observation may be true considering the large sample size but of the Chestnut-winged Cuckoo, including: Black no observer has come across either of the laughingthrushes Dicrurus macrocercus, Long-tailed Lanius schach, feeding a young Pied Cuckoo. The southern Indian sky-island White-crested Laughingthrush G. leucolophus (also in endemic cachinnans is Baker 1907a), Spot-breasted Laughingthrush G. merulinus, supposedly parasitised by Pied Cuckoos (Hume 1890; Baker Rufous-necked Laughingthrush P. ruficollis, Moustached 1934), however, all evidences relate to cuckoo eggs in nests Laughingthrush Ianthocincla cineracea, Rufous-chinned as reported by collectors, and not to actual young birds seen Laughingthrush I. rufogularis (also in Baker 1907a, 1907b), at a nest, or with this species. The Pied Cuckoo is generally a Laughingthrush T. chrysopterum, Red-faced Liocichla scarce species in the woodlands of the upper Nilgiris, and a Liocichla phoenicea, Rusty-fronted Actinodura confusion of its eggs with that of the Common Hawk Cuckoo egertoni, Blue Whistling Myophonus caeruleus, Slaty- Hierococcyx varius needs investigation. Tooth (1902), and backed Enicurus schistaceus, Orange-headed Thrush Osmaston (1916), reported its eggs from Red-vented ’s Geokichla citrina, and Black-breasted Thrush Turdus dissimilis. Pycnonotus cafer nests, but these remain unconfirmed, as does Though Baker (1907b) listed Spot-breasted the one egg in Baker’s collection (Baker 1934). Reference to the Erythrogenys mcclellandi as one of the victims, it is missing in species parasitizing the erythronotus of the Long- his Baker (1942). Additionally, Baker (1934) cited secondary tailed Shrike in Kashmir (Ali & Ripley 1987) needs verification, information for [=Quaker Thrushes] as the only subspecies listed in Baker (1942) is nigriceps [one poioicephala and (Pycnonotidae) as two unlikely hosts egg]. Baker also recounts shooting this cuckoo from the nest where eggs of this species were found. However, both species of a (Baker 1907). Other hosts listed in Baker are also missing in Baker (1942), suggesting a retraction. Baker (1907b) are , Orange-headed Thrush, (1907a) also got a nest and clutch of an Oriental Magpie Robin and Chestnut-bellied Rock Thrush Monticola rufiventris. Baker Copsychus saularis from a local person that had an egg of this (1942) lists more hosts, presumably all from the region, species. Hume (1890) reported taking an egg from the nest of including several laughingthrushes, namely, White-crested, an Indian Silverbill Euodice malabarica and claimed the cuckoo Rufous-vented, Rufous-necked, Grey-sided, Blue-winged, must have laid the egg elsewhere and pushed it inside the nest Spot-breasted, Striated, and Brown-capped Laughingthrush T. of the Silverbill! austeni, Red-faced Liocichla, P. woodi, Rusty-fronted Barwing, White-tailed Robin Myiomela leucura, Pied Cuckoo Clamator jacobinus , and Blue . Baker (1942) listed This polytypic cuckoo comprises a resident nominate subspecies secondary evidence for the Common Aegithina tiphia (by and a migratory subspecies pica that arrives along with the south- Capt. Salder), and the Oriental Magpie Robin. western monsoon from . Its most preferred host species are the several species of Argya babblers (Table 1). Eudynamys scolopaceus Becking (1981) specifically identified the Streaked The Asian Koel is a widespread and , with a Laughingthrush T. lineatum as one of the ‘Himalayan’ range covering much of the Oriental Region. It has two regionally Laughingthrushes tagged as its popular host, and this is likely occurring subspecies—the widespread nominate, and malayanus; to be correct (See Table 1). However, Baker listed all regionally the latter occupies the extreme south-eastern part of the region. occurring Argya spp., including the Slender-billed Babbler A. This species has the earliest documentation of brood parasitism longirostris [two eggs] (Baker 1942), A. earlei in the world, being mentioned more than 4,000 years ago in the [one egg] (Baker 1907a, 1942 but also Hume 1890), and the Vedas (Ali 1979). It has been documented to use six hosts in the Sri Lankan endemic Orange-billed Babbler A. rufescens (Baker region (Table 2). The favoured host species is the House Crow C. 1907a). A Pied Cuckoo pair was documented attempting to splendens followed by the Large-billed Crow C. macrorhynchus distract an Indian Paradise-flycatcherTerpsiphone paradisi on its (Lamba 1987). Baker’s egg collection has a 112:32 ratio for nest in Gujarat (Bhatt 2019). House Crow:Large-billed Crow. In our cataloging effort, this ratio Baker (1934, 1942) also listed the Lesser Necklaced was even more disproportionately tilted towards the House Laughingthrush [41 eggs] as the most preferred victim, followed Crow (74:3). Baker (1942) also lists one egg from a Common by the Greater Necklaced Laughingthrush [nine eggs]. This ’s nest, which was presumably taken by Inglis and sent to

Table 1. Pied Cuckoo host species No Species Type Reference 1. Argya striata S Hume (1890); Gaston (1976); Nahid et al. (2016a) 2. Yellow-billed Babbler Argya affinis S Ali & Whistler (1936); Johnsingh & Paramanandham (1982); Perera (2009); Sashikumar et al. (2011); Anoop et al. (2016) 3. Argya caudata S Gaston (1976); Seneviratne (1997) 4. Argya malcolmi S Hume (1890); Gaston (1976) 5. Argya subrufa O Sashikumar et al. (2011); Lath (2016); Cherian & Sanuraj (2020) 6. Trochalopteron lineatum O Gaston (1973); Prince (2003) 106 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

Table 2. Asian Koel host species No Species Type Reference 1. Dicrurus macrocercus O Smith (1950) 2. Long-tailed Shrike Lanius schach S Begum et al. (2011a,b); Nahid et al. (2016a) 3. House Crow Corvus splendens S Hume (1890); Lamba (1987); Ali et al. (2007) 4. Large-billed Crow Corvus macrorhynchos S Lamba (1987) 5. Red-vented Bulbul Pycnonotus cafer O Plomp (2019); Mandal (2020) 6. tristis M Choudhury (1998); Ansary (2019); Karthikeyan (2020); Namdeo (2020)

him (Inglis 1908); and we found four documentations of this As per Baker (1907a), former reports of eggs taken from host-parasite pair, with three from the eastern part of the Indian the Brownish-flanked Bush WarblerHorornis fortipes, the Brown Subcontinent (Table 2). D’Abreu (1927) reported a Koel’s egg Bush Warbler Locustella luteoventris, the Rufous-capped Babbler in the nest of an Oriolus kundoo but there ruficeps (Hume 1890), and the Grey-bellied are no similar reports and Baker (1934, 1942) do not include it. Tesia cyaniventer were of the ; interestingly, most of these species were listed under Chrysococcyx maculatus by Baker (1942). However, he claimed to have noosed an The Asian Emerald Cuckoo is mainly a resident of the north- Emerald Cuckoo at the nest of a Grey-throated Babbler eastern Indian hills, and is believed to parasitize sunbirds and nigriceps that already had a cuckoo egg (see Becking 1981 for (Nectariniidae). The only incidents of juveniles the argument why the egg is of a Drongo Cuckoo). He also took being fed by hosts were from Garbhanga Reserve , Assam, another egg, possibly of this species, from a Common ’s where the hosts were Crimson Sunbirds siparaja nest. He additionally commented on eggs sent to him by H. A. [79, 80]. All of Baker’s historical reports of eggs and chicks) of Hole, from the nests of Grey-throated Babbler and Buff-chested both, Emerald, and Violet Cuckoos C. xanthorhynchus are so Babbler Cyanoderma ambiguum. Except for the Tailorbird, none muddled with confusion over their identity vis-à-vis that of Lesser of these species figure as victims in Baker (1942)—reported Cuckoo C. poliocephalus, C. merulinus, Banded eggs were from the clutches of Little Arachnothera Bay Cuckoo C. sonneratii, and even Fork-tailed Drongo Cuckoo longirostra [5 eggs], Crimson [5], Mrs Gould’s Sunbird A. dicruroides that only tentative assignments are gouldiae [1], Zitting Cisticola juncidis [3], and Common possible using Baker’s collection (Becking 1981; Payne 2005). Tailorbird [2]. Luku R anjan Nath

80. feeding an Asian Emerald Cuckoo at Garbhanga Reserve Forest, Assam, August 2020.

Violet Cuckoo Chrysococcyx xanthorhynchus The is mainly a resident of the north-eastern Indian Hills, and possibly of the Andaman Islands. It is believed to parasitize sunbirds and spiderhunters. No hosts have been seen feeding a juvenile cuckoo within our region (Ali & Ripley 1987; Payne 2005) until an Olive-backed Sunbird C. jugularis Himanka B. Deka 79. Crimson Sunbird feeding an Asian Emerald Cuckoo at Garbhanga Reserve Forest, Assam, was photographed feeding a juvenile at Shoal Bay, Andaman May 2019. Islands [81, 82]. Praveen & Lowther: Avian brood parasitism in South Asia 107

Baker obtained several eggs from the nests of Nepal C. M. Inglis and A. M. Primrose collected cuckoo eggs from as well as, once, from a Cachar Wedge-billed Babbler S. roberti the nests of Crimson Sunbirds in Cachar, which they thought were (Baker 1907a). He additionally commented that the egg he of this species, which was locally common in their area. Baker received as this species, from H. A. Hole, from Cachar, were likely assigned them, initially, to the Plaintive Cuckoo, but Inglis (1908) those of an Emerald Cuckoo, though the collector himself did refuted that claim citing the rarity of that species in their area. not agree. Later, Baker (1934) included the Blyth’s However, Baker (1908a,b) corrected himself that they belonged Phylloscopus reguloides, and a Cisticola also as victims. Much of to an Emerald Cuckoo, based on a chick that was hatched from this information did not find a place in Baker (1942), wherein a sunbird’s nest by Primrose. The specimen of that chick has he reported eggs from the clutches of the not been located nor been examined by others. Identification of [5 eggs], as well as single eggs with Crimson Sunbird, Zitting juvenile Chrysococcyx is complicated, and hence its real identity Cisticola, , and Yellow-bellied Warbler remains inconclusive (Becking 1981). However, based on the superciliaris. original observations of Inglis and Primrose, this bird could have indeed been a juvenile Violet Cuckoo.

Banded Bay Cuckoo Cacomantis sonneratii A widespread, locally common bird in the hills of the Oriental Region the Banded Bay Cuckoo mostly parasites , (Baker 1907b; Becking 1981; Table 3). While a treatise on its host- parasite relationship is pending, we obtained 24 independent instances of documentation from western India (Fig. 2), where both, James Davidson and T. R. Bell, also reported Common feeding young ones in the late nineteenth century from the district of Uttar Kannada (Baker 1907b). The peak feeding appears to be in October–November, though there are reports from June till February. Though we found reports of five other host species, none of them are backed by media, though all of them are from reliable observers. Baker (1907b) also quoted Davidson to have taken eggs believed to be of this species from Red-whiskered Bulbul, and others who have taken it from the nests of Tawny-bellied Babbler Dumetia hyperythra [but see under Drongo Cuckoo]. Baker (1934) reported a high incidence [33 eggs] from the nests of the Nepal Fulvetta, but these are misidentified Fork-tailed Drongo Cuckoo eggs (Becking 1981). The other sixteen listed victims in his collection, including several small babblers, which could also have been misidentified are (see Becking 1981) [5], Common Tailorbird [3], Red-vented Bulbul [4], Spot-throated Babbler Pellorneum albiventre [4], Puff-throated Babbler P. ruficeps[1], Buff-chested Babbler [2], Grey-throated Babbler [1], Yellow-eyed Babbler Chrysomma sinense [1], Abbott’s Babbler Malacocincla abbotti [2], Chestnut-capped Babbler Timalia pileata [1], Slender-billed Scimitar Babbler Pomatorhinus superciliaris [1], Rufous-bellied Niltava sundara [1], White-browed Rhipidura aureola [1], Pied Bushchat caprata [1], and Tawny-breasted Babbler Both: Moorching Gururaj Spelaeornis longicaudatus [1]—and strangely just one egg from 81 & 82. An Olive-backed Sunbird feeding a Violet Cuckoo at Shoal Bay, Andaman Islands, March 2018. the Common Iora!

Table 3. Banded Bay Cuckoo host species No Species Type Reference 1. Small Minivet Pericrocotus cinnamomeus O Sashikumar et al. (2011) 2. Orange Minivet Pericrocotus flammeus O Philips (1944); Pepper (2008) 3. Black-headed Cuckooshrike Lalage melanoptera O Robson (2010); Shekhawat (2018) 4. Red-whiskered Bulbul Pycnonotus jocosus O Jackson (1971) 5. Common Iora Aegithina tiphia M Davidson (1898), see map (Fig. 2) 6. Common Woodshrike Tephrodornis pondicerianus O Radhakrishnan (2020) 108 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020) Ali Kawser Dany Kawser Ali

Fig. 2. Banded Bay Cuckoo parasitism on Common Iora. Adapted from https://www. stateofindiasbirds.in/ 83. A Plain feeding a Plaintive Cuckoo at Dhaka, , October 2019. Plaintive Cuckoo Cacomantis merulinus The Plaintive Cuckoo is widespread in the lowlands of eastern Grey-bellied Cuckoo Cacomantis passerinus India and known to parasitise Prinia spp., and tailorbirds (Becking The Grey-bellied Cuckoo is a widespread resident in South Asia 1981; Table 4). Not much has been published formally from and is marginally sympatric with the former species in eastern India, and the listing of Striated Prinia P. crinigera, P. India; Grey-bellied Cuckoos are known to parasitise on , socialis, Black-throated Prinia P. atrogularis, Grey-breasted Prinia Common Tailorbird, , and sunbirds (Becking 1981; P. hodgsonii, Plain Prinia P. inornata, Common Tailorbird, and Payne 2005; Table 5; Fig. 3). —as brood parasites in Ali & Ripley (1987), Becking Except for one host in Table 5, the rest have been documented (1981), and Billerman et al. (2020) are based on Baker’s work with photographs. Most of the host-parasite details we know on eggs, and not on observations of actual foster parents feeding of this species have come from R. K. Burnett and others who the cuckoo. As a part of this work, we obtained evidences of reported it commonly parasitizing Ashy Prinias in Hyderabad, and two Prinias and the Common Tailorbird feeding young Plaintive Baker (1907b) reproduced their communication. Suter (1945) Cuckoos. reported a bizarre incident wherein a Grey-bellied Cuckoo fledged Baker (1942) lists several additional species as victims based out of a sunbird’s nest,2 but Common Tailorbirds also participated on his egg collection; some of them may get validated in the in feeding the cuckoo chick while it was in the nest (Suter 1945). future. They are Zitting Cisticola [44 eggs], Striated Prinia [31], Bharucha (1983) also reported an instance of a Black-throated Prinia [25], Common Tailorbird [37], Dark-necked asiaticus bring up a juvenile of this species in in Tailorbird O. atrogularis [5], Grey-breasted Prinia [28], Rufescent the 1960s, but we do not include it here as the observer was Prinia P. rufescens [1], Yellow-bellied Prinia P. flaviventris [1], recounting an incident that had happened 20 years prior, and Yellow-bellied Warbler [1], Buff-chested Babbler [1], Pin-striped hence not considered reliable. Though Becking (1981) suggested Tit Babbler Mixornis gularis [3], Cachar Wedge-billed Babbler it might rather have been a Chrysococcyx, probably because [1], Oriental Magpie-Robin [1], and Indian White-eye Zosterops it selected a sunbird as host, the sizeable number of sunbird palpebrosus [1]; not all of them may have been taken within hosts (mostly Purple-rumped Sunbird) for this cuckoo, in that South Asia. region, indicates the cuckoo must have been identified correctly.

Table 4. Plaintive Cuckoo host species No Species Type Reference 1. Common Tailorbird Orthotomus sutorius M Nahid et al. (2016b) 2. Black-throated Prinia Prinia atrogularis M McNAlly (2015); Ryndem (2019) 3. Plain Prinia Prinia inornata M 83

2 This was most likely a Purple-rumped Sunbird zeylonica as mentioned in the text. The reference to Purple Sunbird Cinnyris asiaticus in the title could have been an editorial error since the author probably did not provide the title for his note. Praveen & Lowther: Avian brood parasitism in South Asia 109

Table 5. Grey-bellied Cuckoo Cacomantis passerinus host species No Species Type Reference 1 Black-headed Cuckooshrike Lalage melanoptera O Dharwadkar (2015) 2. Common Tailorbird Orthotomus sutorius M Baker (1906b); Sashikumar et al. (2011) 3. Ashy Prinia Prinia socialis M Baker (1907b); Ali & Whistler (1934) 4. Plain Prinia Prinia inornata M Hume (1890) 5. Zitting Cisticola Cisticola juncidis M 84; Charachar (2018) 6. Purple-rumped Sunbird Leptocoma zeylonica M Bharucha (1983) 7. Passer domesticus M 85

Both Chrysococcyx species do not occur in the Deccan region, even as vagrants. Hume (1890) had identified some additional victims, the latter two as reported by R. Thompson—Red-vented Bulbul from Darjeeling, based on a bird snared near its nest, and young cuckoos seen in the nests of the Long-tailed Shrike and the Yellow-eyed Babbler in Dehra Dun. Thompson’s identity of these young cuckoos, seen in the nest, has not been confirmed. Baker (1942) lists Ashy Prinia [41], Plain Prinia [2], Zitting Cisticola [4], Common Tailorbird [6], P. gracilis [1], Western Crowned Warbler P. occipitalis [1], Dark-fronted Babbler Rhopocichla atriceps [1], and Small Minivet Pericrocotus Shwetha Bharathi Shwetha cinnamomeus [1, probably by Col. Rattray]. The high number of Ashy Prinia eggs in Baker’s collection may have come from Hyderabad through R. K. Burnett as recounted by Baker in Livesey (1936), where these were specifically collected on Baker’s behest. 84. A Grey-bellied Cuckoo about to be fed by a Zitting Cisticola at Mysore, Karnataka, October 2017. Manas Gupta

85. A female House Sparrow feeding a Grey-bellied Cuckoo at Pune, , October 2018.

Himalayan foothills while the Square-tailed Drongo Cuckoo is Fig. 3. Regular hosts of the Grey-bellied Cuckoo. After: https://www.stateofindiasbirds.in/ mostly a resident of eastern India and extending along the base of the Himalayas westwards, until about where both are presumed to be sympatric. Separation of adult birds, based Fork-tailed Drongo Cuckoo Surniculus dicruroides & on morphology itself, is complicated (Gala & Ongole 2018) Square-tailed Drongo Cuckoo Surniculus lugubris and hence assignment of juveniles to either species should be These two resident species of the are treated as tentative, particularly where their ranges overlap. We sometimes considered a single species (Rasmussen & Anderton treat them under one head and indicate likely species based 2012; Dickinson & Remsen 2013). The Fork-tailed Drongo on the location of the observation. They are mostly known to Cuckoo is a resident in Peninsular India extending nearly to the parasitize on small babblers (Payne 2005). 110 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

Table 6. Fork-tailed/Square-tailed Drongo Cuckoo host species No Species Type Reference Species ID (based on range) 1. Yellow-eyed Babbler Chrysomma sinense M 86 & 87 Fork-tailed Drongo Cuckoo 2. Indian White-eye Zosterops palpebrosus M Raiyan (2019) Square-tailed Drongo Cuckoo 3. Tawny-bellied Babbler Dumetia hyperythra O Venkatraman (2019); Sharma (2019) Fork-tailed Drongo Cuckoo 4. Dark-fronted Babbler Rhopocichla atriceps M Becking (1981); Dayananda (2017); Dharwadkar (2018) Fork-tailed Drongo Cuckoo 5. Black-chinned Babbler Cyanoderma pyrrhops M 88 Uncertain

A nestling collected by W. E. Wait, from the nest of a Dark- fronted Babbler in Sri Lanka, has been confirmed from the specimen (NHM #1925.9.5.2) as this species. An egg collected by J. Stewart from Aneichardi, Kerala, from the same host’s nest is also similar to Drongo Cuckoo eggs from elsewhere (Becking 1981). We report four new hosts for this species: Black-chinned Babbler Cyanoderma pyrrhops; Yellow-eyed- and Tawny-bellied Babblers presumed to be parasitizing on Fork-tailed, while Indian White-eye parasitizing on Square-tailed; the species have been assignment based on their range (Table 6). It must be noted that there is a dated, uncorroborated report of Banded Bay Cuckoo eggs (see that species) from Tawny-bellied Babbler nests, and that could have been of Drongo Cuckoos. Baker (1942) had very little to add to this species, presumably having identified many of its eggs as another species. We found no evidence for other victims mentioned in Ali & Ripley (1987): , minivets, ioras, and , as well as additional victims mentioned in

Payne (2005), as from India: Nepal Fulvetta, , and Striated Trilok Singh Bisht Grassbird Megalurus palustris.

88: Black-chinned Babbler feeding a Drongo Cuckoo at Haidakan Road, Uttarakhand, September 2019.

Large Hawk Cuckoo Hierococcyx sparverioides The Large Hawk Cuckoo is resident in the Himalayas, sparingly wintering in south-western India apart from the lower foothills. They are known to parasitize mostly on A. magna and leucophris. However, there is one recently documented instance of a juvenile being fed by Chestnut-crowned Laughingthrushes T. erythrocephalum (Sharma 20203; Table 7). There is a debate about the plain blue cuckoo eggs taken by Baker, and listed for laughingthrushes in Baker (1942), that they were from a Common Cuckoo C. canorus (Becking 1981). However, this is despite Colonel Rattray taking a similar blue oviduct egg in Musoorie and noticing similar eggs in the nests of the . Similar eggs were collected by B. B. Osmaston and A. E. Osmaston in Nainital from the nests of Chestnut-crowned Laughingthrush; and on one previous occasion the junior Osmaston having secured a chick of a Large Hawk Cuckoo via his chaprassi [=attendant], which Becking downplayed as he could not trace the skin. Hodgson recounted Chestnut-crowned Laughingthrushes feeding this species, which he claimed was a year old (!) (Hume 1890). Payne (2005) exposed certain limitations in Becking’s analysis, of not reporting the ultrastructure of Heirococcyx sp., and the said eggs may be similar to that of Common Cuckoo’s, but still of this Both: Agnihotri Surendra 86 & 87. A Yellow-eyed Babbler feeding a Drongo Cuckoo at Nagpur, Maharashtra, October 2011. 3 See elsewhere in this issue. Praveen & Lowther: Avian brood parasitism in South Asia 111

species. While the right progenitor of all these blue eggs is still information on its secondary hosts, particularly in regions clouded in mystery, Sharma (2020) establishes that the Large where its primary hosts are missing, is largely unknown. There Hawk Cuckoos indeed use some laughingthrush species to raise is an instance of the Long-tailed Shrike hosting this species in its progeny in India as it has been well established also in Chandrapur District, but we are unable to establish the veracity of (Yang et al. 2012). the same vis-à-vis a Common Cuckoo (Mishra 1989). Contrary to Becking (1981), the ‘brown-type’ eggs attributed Baker (1942) obtained most of his eggs from Jungle to this species in the clutches of Streaked Spiderhunter, Lesser Babbler nests [32 eggs], apart from Large Grey Babbler [7], Shortwing, and other smaller birds need to be challenged. Yellow-bellied Babbler [1], Common Babbler [1], Rufous-necked There are no field observations to back this claim and Becking’s Laughingthrush [4], Moustached Laughingthrush [4], Assam conclusion that these two types of eggs must belong to two Laughingthrush [2], Lesser Necklaced Laughingthrush [1], Rufous- different species may indeed be true, with some of the brown- vented Laughingthrush [1], Grey-sided Laughingthrush [1], Red- type eggs belonging to another cuckoo species. faced Liocichla [2], Rufous-fronted Barwing [2], Indian Paradise- Baker (1942) obtained a large number of ‘brown-type’ eggs flycatcher [1], Rufous-bellied Niltava [1], and Asian Fairy- [98 eggs] from the clutches of Streaked Spiderhunter and Lesser Irena puella [1], and possibly, Orange-billed Babbler in Sri Lanka Shortwing [24] apart from Little Spiderhunter [4], Brownish- (see Lushington 1949). Becking’s (1981) suggestion that many flanked Bush Warbler [4], Buff-chested Babbler [2], Grey-throated of the eggs in Baker’s collection attributed to the Whistling Hawk Babbler [2], Pygmy Blue Flycatcher Ficedula hodgsoni [1], Small Cuckoo may have been of this species. Niltava N. macgrigoriae [1], White-naped Yuhina Yuhina bakeri [1], Y. flavicollis [1], Spot-throated Babbler [2], Puff-throated Babbler [1], Red-vented Bulbul [1], and Long-tailed Shrike [1]; and listed as not in his collection are Nepal Fulvetta and Blue-throated Flycatcher Cyornis rubeculoides. The controversial ‘Blue-type’ eggs are listed for Lesser Necklaced Laughingthrush [12], Greater Necklaced Laughingthrush [6], Moustached Laughingthrush [6], Chestnut-crowned Laughingthrush [2], Grey- sided Laughingthrush [1], Red-faced Liocichla [1], Spot-breasted Scimitar Babbler [2], Blue Whistling Thrush [5], Orange-headed Thrush [2], and Scaly Thrush Zoothera dauma [1]. In addition, two more species listed as, not in Baker’s collection, are Rufous- chinned Laughingthrush and Hoary-throated Barwing Actinodura nipalensis, the latter probably based on Hodgson’s account in Hume 1890. The accounts of Large Hawk Cuckoo building its own nest (Hume 1890), by Rhodes Morgan and Miss Cockburn, from southern India has not found further circulation, and those twig nests may have belonged to an Accipiter (see Baker 1934).

Common Hawk Cuckoo Hierococcyx varius The Common Hawk Cuckoo is a widespread and resident in India, parasitizing Argya babblers (Table 5); mostly on Jungle A. Fig. 4. Common Hawk Cuckoo parasitism on Large Grey Babblers. Adapted from https://www. stateofindiasbirds.in/ striata and Large Grey Babblers A. malcolmi, and Yellow-billed Babblers A. affinis in southern India. Though it is documented as having parasitized laughingthrushes in the Himalayas, evidence Whistling Hawk Cuckoo Hierococcyx nisicolor is lacking. Despite the species being very common, definitive Decidedly the rarest of the three Hawk Cuckoos, and restricted

Table 7. Large Hawk Cuckoo host species No Species Type Reference 1. Chestnut-crowned Laughingthrush Trochalopteron erythrocephalum M Hume (1890), Osmaston (1912); Sharma (2020) 2. Hoary-throated Barwing Actinodura nipalensis O Hume (1890)

Table 8. Common Hawk Cuckoo host species No Species Type Reference 1. Jungle Babbler Argya striata S Lushington (1949); Zacharias & Gaston (1983); Sashikumar et al. (2011) 2 Yellow-billed Babbler Argya affinis M Prasad et al. (2001) 3. Large Grey Babbler Argya malcolmi M See map (Fig. 4) 4. Nilgiri Laughingthrush Montecincla cachinnans O Davison (1883) 112 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

to north-eastern India till about eastern Nepal, practically nothing Baker obtained the largest number of eggs [24 eggs] from is known about the Whistling Hawk Cuckoo’s forester parents clutches of the Brownish-flanked Bush Warbler, and also had except for the victim list in Baker (1942). eggs from clutches of Blyth’s Leaf Warbler [8], Western Crowned Baker (1942) obtained most of his eggs from the clutches Warbler [4], Yellow-vented Warbler P. cantator [4], Grey- of [23 eggs] and Lesser Shortwing [14]. hooded Warbler P. xanthoschistos [2], Large-billed Leaf Warbler Additional victim species include Blue-throated Flycatcher [7], P. magnirostris [1], Grey-cheeked Warbler P. poliogenys [1], Plumbeous Water fuliginosus [7], Streaked Yellow-bellied Warbler [3], Dark-necked Tailorbird [3], Common Spiderhunter [2], Little Spiderhunter [2], Brownish-flanked Bush Tailorbird [1], Grey-bellied Tesia [4], Chestnut-headed Tesia Warbler [2], Spotted Forktail [2], Snowy-browed Flycatcher F. castaneocoronata [1], Pale-footed Bush Warbler hyperythra [1], thalassinus [1], pallidipes [1], Scaly-breasted Wren Babbler Pnoepyga albiventer Grey-headed Canary-flycatcherCulicicapa ceylonensis [1], [3], Pygmy Wren Babbler P. pusilla [2], B. White-browed Fantail [1], Cachar Wedge-billed Babbler [1], cruralis [1, probably the same mentioned in Osmaston (1904) Grey-bellied Tesia [1], Spotted Forktail [1], Buff-chested Babbler as from Darjeeling], Nepal Fulvetta [1], Indian White-eye [1], and [1], Grey-headed Psittiparus gularis [1], Buff-breasted Streaked Spiderhunter [1]. There is no other available literature Babbler P. tickelli [1], Rufous-vented Yuhina occipitalis [1], and that validated these claims though some of the Phylloscopus Yellow-throated Fulvetta Schoeniparus cinereus [1]. Whether all warblers in this list may be true hosts. these eggs belong to this species, or to the preceding two, is unknown (see Becking 1981). micropterus The Indian Cuckoo is widespread in the Indian Subcontinent, Lesser Cuckoo Cuculus poliocephalus though patchy in distribution, it is known to parasitize mainly This species breeds along the Himalayas and in north-eastern drongos, while it has been suggested that other documented host India and migrates across the Indian Subcontinent, possibly species are dubious (Becking 1981). However, we found host- crossing the Arabian Sea/Indian Ocean, to its non-breeding parasite evidence from only three species (Table 10), including grounds in Africa. As per Becking (1981), they are specialized an oriole, for which Becking (1981) had expressed reservations. in their host requirement, limiting themselves to bush warblers Jones (1941) provided several presumptive evidences for that breed close to the ground; however, the two field reports we Indian Cuckoo parasitizing Black Drongos, though none of them collected were both for Phylloscopus warblers (Table 9). Though were definitive. However, we found nine instances (Table 10; Brooks is believed to have taken an egg from the nest of a Hume’s Fig. 5), apart from a more detailed study by Nahid et al. (2016a) Warbler P. humei in Kashmir (Hume 1890; Baker 1906b), recent in Bangladesh. Ashok Kumar et al. (2001) reported Common studies in Kashmir (Price & Jamdar 1992; Marchetti 1992) found Babblers feeding a juvenile of this species, but this was most likely this warbler to be a ‘rejecter species’ that will not incubate cuckoo a misidentification of Yellow-billed Babblers feeding a Common eggs; at times rejecting their own eggs when artificial eggs were Hawk Cuckoo. Though Baker (1942) listed two types of eggs, placed in its nest (Marchetti 1992). Price & Jamdar (1992) only one amongst the five Drongo-type eggs in his collection studied seven other leaf warblers in Kashmir and found just was a cuckoo egg (Becking 1981); the proven egg believed to one instance of parasitism (Table 9). This is contrary to many of have be taken from the clutch of an Indian Paradise-flycatcher, Baker’s correspondents who found eggs of this species from the presumably taken by Skinner, where the bird was seen sitting on nests of Western Crowned Warbler (Baker 1906b). This same the nest (Baker 1934). Most of the blue eggs of Indian Cuckoo work describes the origins of most of the eggs vis-à-vis the host can be attributed to Colonel Rattray from Murree where he took species that was catalogued in Baker (1942). Apparently, this them from the nests of Streaked Laughingthrush, Indian Blue cuckoo lays multiple variants of its egg—most frequently a ‘red’ Robin brunnea, Saxicola maurus, and a ‘white’ variant. The ‘red’ types are apparently found in nests and White-bellied Redstart phaenicuroides (Baker of bush warblers, and the white in leaf warblers (Rattray 1923; 1934), but the last species was not listed in Baker (1942). Baker 1942). Baker’s (1942) collection had blue eggs from Streaked

Table 9. Lesser Cuckoo host species Balakrishnan Valappil Balakrishnan No Species Type Reference 1. Tickell’s Leaf Warbler Phylloscopus affinis O Price & Jamdar (1992) 2. Grey-hooded Warbler Phylloscopus xanthoschistos M Valappil (2016); 88a 88a. Grey-hooded Warbler feeding Lesser Cuckoo at Devalsari, Uttarakhand, in June 2016.

Table 10. Indian Cuckoo host species No Species Type Reference 1. Black-hooded Oriole Oriolus xanthornus M Eldhose (2000); Jayasekara (2020) 2. Black Drongo Dicrurus macrocercus M O’Donel (1936); Storrs (1944)4; Neelakantan (1952); Prasad & Jayant (1993); Praveen (2009); Sashikumar et al. (2011); Nahid et al. (2016a); Fig. 5 3 Streaked Spiderhunter Arachnothera magna O Gammie (1877)

4 Storrs (1944) not verified, but possibly refers to the same record in O’Donel (1936) Praveen & Lowther: Avian brood parasitism in South Asia 113

Laughingthrush [5], Indian Paradise-flycatcher [1], White-browed Warblers (Baker 1906a). C. L. Wilson found a Himalayan Cuckoo Fantail [1], Grey-winged Blackbird Turdus boulboul [2], Golden hovering around the nest of a Hume’s Warbler P. humei in Bush Robin Tarsiger chrysaeus [1], [1], Kashmir that had a Himalayan Cuckoo-type egg, which he later Siberian Stonechat [1], and Blyth’s Leaf Warbler [1]. He found found in the nests of the Western Crowned Warbler as well drongo-type eggs from D. leucophaeus [3], Black (Baker 1906a). B. B. Osmaston found eggs of this bird in the Drongo [1], and Indian Paradise-flycatcher [1]. nests of Rufous-bellied Niltava in Darjeeling (Baker 1906a). Baker listed a few victim nests from which he had taken eggs (Baker 1907b): Verditer Flycatcher, Rusty-cheeked Scimitar Babbler E. erythrogenys, and Rufous-bellied Niltava, but the first two were not included in Baker (1942). Baker’s (1942) list included a large number of eggs [17 eggs] from the clutches of Blyth’s Leaf Warbler, and records from Chestnut-crowned Warbler P. castaniceps [6], Tickell’s Leaf Warbler P. affinis [5], Western Crowned Warbler [4], Hume’s Warbler [2], Lemon-rumped Warbler P. chloronotus [1], Yellow- vented Warbler [1], Green-crowned Warbler P. burkii [1], Grey- hooded Warbler [1], Cachar Wedge-billed Babbler [2], Tawny- breasted Wren Babbler Spelaeornis longicaudatus [2], Striated Prinia [1], Dark-necked Tailorbird [1], Himalayan Bush Robin Tarsiger rufilatus [1, likely by Rattray, see Baker (1907b)], Rufous-bellied Niltava [1], Rock Emberiza cia [1], and Pygmy Wren Babbler [1]. There is no available literature that has validated these claims though the Phylloscopus warblers in this list may be true hosts.

Common Cuckoo Cuculus canorus The Common Cuckoo is a widespread breeder along the Himalayas and north-eastern India, and is mostly an autumn passage migrant across the Indian Subcontinent. It migrates Fig. 5. Indian Cuckoo parasitism on Black Drongo. Adapted from https://www.stateofindiasbirds.in/ over a broad front, usually spending two to three months before crossing the Indian Ocean to reach its final destination in Africa. Himalayan Cuckoo Cuculus saturates This is, arguably, the most well studied brood parasite in the world Breeding along the Himalayas and north-eastern India, and and has been documented to be catholic in its host choice, with sometimes considered conspecific with the extralimital Oriental more than 100 hosts across the world. Within subspecies, Baker Cuckoo C. opatus, the Himalayan Cuckoo’s hosts are believed (1942) lists 35 victims for ‘Asiatic Cuckoo’ C. c. telephonus, and to be Phylloscopus warblers, specifically the Western Crowned 127 victims of ‘Khasi Hills Cuckoo’ C. c. bakeri. However, the Warbler in the western Himalayas, and Blyth’s Leaf Warbler in the number of actual observations of host-parasite relationship from central and eastern Himalayas (Becking 1981). Unfortunately, India has been meagre (Table 11) and there is concern that recent evidence of cuckoo young being fed by either of these some of the eggs documented for other cuckoo species could species, or in fact by any other species, is lacking. have been of Common Cuckoo (see Becking 1981). Much of Colonel Rattary recounts an incident where a Himalayan the anecdotal notes in Hume (1890) do not refer to specific Cuckoo was harassing a pair of Western Crowned Warblers at species and details of the young are unavailable for assessment. Mussoorie, and the bird, when shot, had an oviduct egg; he found Jones (1941) is said to have recorded a Common Cuckoo laying further eggs of similar kind on the nests of Western Crowned an egg in the nest of a Siberian Stonechat, but does not refer to

Table 11. Common Cuckoo Cuculus canorus host species No Species Type Reference 1. Long-tailed Shrike Lanius schach M Fig. 6 2 Red-billed Leiothrix Leiothrix lutea M Kharkongor et al. (2020)6; Chettri (2015); 90 3. Upland Anthus sylvanus O Hume (1890) 4. Olive-backed Pipit Anthus hodgsoni O Sharma (2020) 5. pectoralis O Osmaston (1925) 6. Blue-fronted Redstart Phoenicurus frontalis M Osmaston (1927); Bates (1949) 7. Phoenicurus fuliginosus M Loke (1952) 8. Pied Bushchat Saxicola caprata O Hume (1890)

5 See elsewhere in this issue 6 See elsewhere in this issue 114 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

any young being raised by that host. It has also been recorded robbing the nest of the Grey Bushchat Saxicola ferreus, possibly prior to parasitism (Sharma 2020) 5. Ali (1968) spotted a Brown Cinclus pallasii feeding a juvenile of the Cuculus sp., at Bringhi Valley, Kashmir, in May 1967. This work also confirms the breeding of this cuckoo in the Indian Subcontinent, away from the main Himalayan range, as has always been suspected (Betham 1910; Ali & Whistler 1940; Bharos 1996, 2005), from Jaipur in , and Jalgaon and Chandrapur in Maharashtra; the last is featured incorrectly in Dhamankar (2004) as a Common Hawk Cuckoo. A bizarre incident of a Common Hoopoe Upupa epops feeding a juvenile Common Cuckoo was reported from Hanle, Ladakh, which would make it the first non- host for any cuckoo in the world (Shivkar 2016, 89). However, observations were not sustained to rule out opportunistic feeding, and hence it is not included here. Prabin Pradhan

90. A Red-billed Leiothrix feeding a Common Cuckoo at Darjeeling, West , October 2010.

Yellow-rumped Honeyguide xanthonotus The only species of honeyguide found in South Asia has a distribution along the central and eastern Himalayas, but nothing is known about its breeding biology. Elsewhere, in Africa parasitize the nests of barbets, kingfishers, and other cavity nesters (Winkler et al. 2020). One case of brood parasitism was reported from Rudraprayag, Uttarkhand, in June 2018, of a young Yellow-rumped Honeyguide being fed by a Green-backed Tit Parus monticolus (Rawat 2018). A honeyguide was seen perched at a hole in a tree that also had a Great Barbet Psilopogon virens nest [91] in May 2015 near Kedarnath Musk Deer Sanctuary, Uttarakhand; whether it was an attempted parasitism is unknown.

Fig. 6. Common Cuckoo parasitism on Long-tailed Shrike. After: https://www.stateofindiasbirds.in/ Anand Vishwanadha Anand Vishwanadha Sahana M Sahana 89. Common Cuckoo just before being fed by a Common Hoopoe at Hanle, Ladakh in August 91. A Yellow-rumped Honeyguide perched at the hole of a tree containing a Great Barbet nest 2016; however an opportunistic feeding cannot be ruled out and the real host could have been near Kedarnath Musk Deer Sanctuary, Uttarakhand, May 2015. While barbets may likely be another species. hosts, the contents of the barbet nest are not known. Praveen & Lowther: Avian brood parasitism in South Asia 115

Table 12. Typical hosts of parasitic cuckoos in comparison with standard literature No Species Payne (2005) Erritzøe et al. (2012) Ali & Ripley (1987) This work 1. Chestnut-winged Laughingthrushes Laughingthrushes Himalayan Laughingthrushes No data Cuckoo 2. Pied Cuckoo Babblers Babblers Babblers, laughingthrushes Jungle- & Yellow-billed Babblers 3. Asian Koel Crows Crows House- & Large-billed Crow House Crow 4. Asian Emerald Crimson Sunbird & Little Spiderhunter Crimson Sunbird & Little Sunbirds (Aethopyga sp.), & Insufficient data Cuckoo Spiderhunter spiderhunters 5. Violet Cuckoo Crimson Sunbird & Little Spiderhunter Crimson Sunbird & Little Sunbirds (Aethopyga sp.), & Insufficient data Spiderhunter spiderhunters 6 Banded Bay Cuckoo Common Iora Common Iora Ioras, bulbuls, small babblers Common Iora 7 Plaintive Cuckoo Warblers with concealed nests, e.g., prinias, Warblers with concealed nests, Common Tailorbird, prinias, Common Tailorbird, prinias tailorbirds, cisticolas e.g., prinias, tailorbirds, & cisticolas cisticolas 8 Grey-bellied Cuckoo Warblers with concealed nests, e.g., prinias, Cisticolas, prinias, tailorbirds Common Tailorbird, prinias, Common Tailorbird, prinias, tailorbirds, cisticolas, as well as sunbirds cisticolas Purple-rumped Sunbird 9 Drongo Cuckoo Small babblers, Nepal Fulvetta, Dark- Shrikes, Dark-fronted Babbler, Iora, Dark-fronted Babbler, Small babblers fronted Babbler, forktails, bulbuls, shrikes, Nepal Fulvetta, forktails, Striated drongos, forktails, minivets Grassbird 10 Large Hawk Cuckoo Streaked Laughingthrush, Chestnut- Lesser Shortwing, Streaked- & Laughingthrushes, Streaked Insufficient data crowned Laughingthrush, Blue Whistling Little Spiderhunter Spiderhunter, Lesser Thrush, Lesser Shortwing, Hoary-throated Shortwing Barwing, Streaked Spiderhunter 11 Whistling Hawk Lesser Shortwing, Small Niltava, and other Lesser Shortwing, Small Niltava, Flycatchers, shortwings Insufficient data Cuckoo flycatchers and babblers and other flycatchers and babblers 12 Common Hawk Babblers Jungle Babbler Babblers, laughingthrushes Jungle-, Yellow-billed & Large Cuckoo Grey Babblers 13 Lesser Cuckoo Tickell’s Leaf Warbler, Blyth’s Leaf Warbler, Leaf Warblers, wren babblers Leaf-Warblers, wren-babblers Insufficient data Brownish-flanked Bush Warbler, Pale- (cupwings) (cupwings), shortwings footed Bush Warbler 14 Indian Cuckoo Black- & Ashy Drongo Black- & Ashy Drongo Drongos Black Drongo 15 Himalayan Cuckoo Seicercus & Phylloscopus warblers Seicercus & Phylloscopus warblers Seicercus & Phylloscopus No data warblers 16 Common Cuckoo Wide range of host species in & Wide range of hosts worldwide, Wide range of hosts Long-tailed Shrike, chats & India, c.100, with a cuckoo egg in the nest c.280 Red-billed Leiothrix' 17 Yellow-rumped Probably barbets Probably barbets Probably barbets Insufficient Data Honeyguide

Table 13. Avian hosts and parasites in South Asia (New host-parasite pairs in bold)

No Parasite Species=> PICU AIKO AECU VICU BBCU PLCU GBCU DRCU LHCU CHCU LECU INCU COCU Total Host Species / No of Hosts => 6 6 1 1 6 3 7 5 2 4 2 3 8 1. Small Minivet Pericrocotus cinnamomeus X 1 2. Orange Minivet Pericrocotus flammeus X 1 3. Black-headed Cuckooshrike Lalage melanoptera X X 2 4. Black-hooded Oriole Oriolus xanthornus X 1 5. Common Woodshrike Tephrodornis pondicerianus X 1 6. Common Iora Aegithina tiphia X 1 7. Black Drongo Dicrurus macrocercus X X 2 8. Long-tailed Shrike Lanius schach X X 2 9. House Crow Corvus splendens X 1 10. Large-billed Crow Corvus macrorhynchos X 1 11. Common Tailorbird Orthotomus sutorius X X 2 116 Indian Birds Vol. 16 No. 4 (Publ. 18 September 2020)

Table 13. Avian hosts and parasites in South Asia (New host-parasite pairs in bold)

No Parasite Species=> PICU AIKO AECU VICU BBCU PLCU GBCU DRCU LHCU CHCU LECU INCU COCU Total 12. Black-throated Prinia Prinia atrogularis X 1

13. Plain Prinia Prinia inornata X X 2

14. Ashy Prinia Prinia socialis X 1

15. Zitting Cisticola Cisticola juncidis X 1

16 Red-vented Bulbul Pycnonotus cafer X 1 17. Red-whiskered Bulbul Pycnonotus jocosus X 1

18. Tickell's Leaf Warbler Phylloscopus affinis X 1

19. Grey-hooded Warbler Phylloscopus xanthoschistos X 1

20. Yellow-eyed Babbler Chrysomma sinense X 1

21. Indian White-eye Zosterops palpebrosus X 1

22. Tawny-bellied Babbler Dumetia hyperythra X 1 23. Dark-fronted Babbler Rhopocichla atriceps X 1

24. Black-chinned Babbler Cyanoderma pyrrhops X 1 25. Jungle Babbler Argya striata X X 2

26. Yellow-billed Babbler Argya affinis X X 2

27. Common Babbler Argya caudata X 1

28. Large Grey Babbler Argya malcolmi X X 2

29. Rufous Babbler Argya subrufa X 1

30. Streaked Laughingthrush Trochalopteron lineatum X 1

31. Chestnut-crowned Laughingthrush X 1 Trochalopteron erythrocephalum 32. Nilgiri Laughingthrush Montecincla cachinnans X 1

33. Hoary-throated Barwing Actinodura nipalensis X 1

34. Common Myna Acridotheres tristis X 1

35. Red-billed Leiothrix Leiothrix lutea X 1

36. Himalayan Rubythroat Calliope pectoralis X 1

37. Blue-fronted Redstart Phoenicurus frontalis X 1

38. Plumbeous Water Redstart Phoenicurus fuliginosus X 1

39. Pied Bushchat Saxicola caprata X 1

40. Purple-rumped Sunbird Leptocoma zeylonica X 1

41. Olive-backed Sunbird Cinnyris jugularis X 1 42. Crimson Sunbird Aethopyga siparaja X 1

43. Streaked Spiderhunter Arachnothera magna X 1

44. House Sparrow Passer domesticus X 1 45. Anthus sylvanus X 1

46. Olive-backed Pipit Anthus hodgsoni X 1

No of hosts => 6 6 1 1 6 3 7 5 2 4 2 3 8

Abbreviations: AECU=Asian Emerald Cuckoo; AIKO=Asian Koel; BBCU=Banded Bay Cuckoo; CHCU=Common Hawk Cuckoo; COCU=Common Cuckoo; DRCU=Drongo Cuckoo; GBCU=Grey-bellied Cuckoo; INCU=Indian Cuckoo; LECU=Lesser Cuckoo; LHCU=Large Hawk Cuckoo; PICU=Pied Cuckoo; PLCU=Plaintive Cuckoo; VICU=Violet Cuckoo Praveen & Lowther: Avian brood parasitism in South Asia 117

Table 14. Number of incidents of brood parasitism in Argya babblers excluding detailed studies No Species Pied Cuckoo Clamator jacobinus Common Hawk Cuckoo Hierococcyx varius 1. Jungle Babbler Argya striata 26 32 2. Yellow-billed Babbler Argya affinis 32 26 3. Large Grey Babbler Argya malcolmi 1 27 4. Rufous Babbler Argya subrufa 3 0

Discussion Conclusion There is a large disparity between field knowledge and information This work concludes that much needs to be done in understanding of victims obtained by oologists on parasitizm (Table 12) that the brood parasitism of South Asian cuckoos. There is an absolute triggered this fresh catalogue (Table 13). Whether it is a reflection dearth of basic host-parasite information for most cuckoos that of the ‘mistakes’ that cuckoos make in laying their eggs in victims’ breed in the Himalayas or north-eastern India; most peninsular nests, or the challenges in identifying eggs visually, is unclear, and Indian cuckoos are slightly better off in this regard. The only cases could be a manifestation of both. While the intent of the work where detailed investigation has been done are of House Crow- was a catalogue, there are also certain patterns on the brood Asian Koel pairs, and Pied Cuckoo-Argya babblers, with limited parasitism of South Asian cuckoos that seem to emerge. work in the cases of Long-tailed Shrike-Asian Koel, and Common South Asian cuckoos parasitize 20 passerine families and 32 Hawk Cuckoo-Jungle Babbler. We know almost nothing about passerine genera. Most cuckoos are specialized, with 40 of 46 the breeding of the Yellow-rumped Honeyguide. Hopefully, this host species (~87%) having only one parasite species to worry catalog can serve as a launch pad for further studies and may about within their range. After the well-known Asian Koel-House encourage birdwatchers and photographers to report new host- Crow pair, Argya babblers seem to be the most parasitized species parasite pairs—some may even by lying in their field notebooks (Table 13), with both, the Pied Cuckoo, and the Common Hawk or media collections. Argya babblers parasitised by two widely Cuckoo parasitizing Jungle and Yellow-billed Babblers almost distributed cuckoos in peninsular India are perfect subjects for equally. The Large Grey Babbler is preferentially parasitised by further exploration into strategies of cuckoo brood parasitism in the Common Hawk Cuckoo with just one documented case South Asia. of being cuckolded by a Pied Cuckoo (Selvaganesh 2020). Interestingly, we could not find any instance of brood parasitism Acknowledgments in Common Babbler, unlike what Gaston (1976) reported for This work would not have been possibly without the hundreds of birdwatchers Pied Cuckoos. While we got several cases of Long-tailed Shrikes from South Asia, who uploaded their observations and media in www.ebird.org feeding young Common Cuckoos, we were not able to collect or in various Facebook groups, and their editors and administrators who kept a high bar on documentation by insisting on adding minimum details, which made any evidence of Asian parasitizing this species, as reported this compilation possible and also fun. Twenty-seven birders responded to the by Begum et al. (2011a,b), and Nahid et al. (2016a) from online survey and an unknown number of supporters cross-posted the request and Dhaka, Bangladesh. helped in publicity. We retrieved a published literature list for South Asia from the Most evidence of parasitized cuckoo chicks being fed is online ‘Bibliography of South Asian ’ (Pittie 2020). Our thanks to C. during July–November, peaking in September–October when Sashikumar for scans of relevant literature, Puja Sharma, Sayam Chowdhury, Tarun much of the peninsula has received ample monsoon rains and Menon, Sanjay Molur, and P. O. Nameer for links to papers, and to Ramit Singal for reviewing an early draft. Puja Sharma, Ashwin Viswanathan, Shashank Dalvi, and are abundant (Fig. 7). Despite so much having being Pritam Baruah helped with some difficult identifications. Sayam Chowdhury, and claimed to be known about cuckoos and their brood parasitism Deepal Warakadoga for chasing details of some reports from their countries. Our (Baker 1942), field workers post 2000 added ten new host- thanks to photographers Himanka B. Deka, Luku Ranjan Nath, Gururaj Moorching, parasite pairs from India (Table 13), nine of which are first reports Anand Vishwanadha, Sahana M., Prabin Pradhan, Trilok Singh Bisht, Manas Gupta, worldwide (Payne 2000). Shwetha Bharathi, Surendra Agnihotri, Balakrishnan Valappil, and Ali Kawser Dany who kindly provided the media. The data compiled as a part of this work is freely accessible online and is in principle owned by the birdwatcher , and future workers are encouraged to use the same and build on it.

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