Eudynamys+scolopaceus

Asian Koel //Eudynamys scolopaceus// (Linnaeus, 1758)

toc Source: Jason Thompson, 2013 (Wikimedia Commons) ||= Figure 2: Female //Eudynamys scolopaceus//. Source: Vasu, V.R., 2011 (Wikimedia Commons) ||
 * = [[image:Figure 1 male Asian koel.jpg width="257" height="374" align="center"]] ||= [[image:Figure 2_female Asian koel.jpg width="293" height="383" align="center"]] ||
 * = Figure 1: Male //Eudynamys scolopaceus//.

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=1 Nomenclature =  Binomial name: //Eudynamys scolopaceus// (Linnaeus, 1758) Synonym: //Eudynamys scolopacea// (Sibley & Monroe, 1990)

 Vernacular names: A variety of vernacular names are given, including but not limited to: Asian koel, Eastern koel, Western koel, Pacific koel, Indian koel, Malayan koel, Common koel, Rainbird, Stormbird, Black cuckoo, 噪鹃 (in Chinese), Burung Sewah Tahu (in Malay) [1,2].

 One of the reasons for such a large number of common names is that the //E. scolopaceus// can be found in many regions of the culturally diverse continent, Asia (see Distribution). Another reason is that definition of the species is still under debate (see Variations in taxonomic classification). To minimise confusion, the scientific name //Eudynamys scolopaceus// will be used throughout this article.

1.1 Etymology
The generic name //Eudynamys// was coined by Vigors & Horsfield in 1826, from the derivative Greek word //dunamis//, meaning power or might, with the prefix //eu//-, meaning good, well, or normal [2].

//Scolopaceus// means snipe-like, and is derived by adding the suffix -//aceus// to the Latin word //scolopax//, which in turn is derived from //scolopakos//, Greek for woodcock. Since //Eudynamys scolopaceus// does not possess a snipe-like bill, this could be a reference to the female’s cryptic striped colouration [2].

The common name //koel// is onomatopoeic* in nature, with a number of language variants. It originates from the Tamil word //kuyil// or the Hindi word //koil//, reflecting the male’s distinct call [1,2].

* Onomatopoeic: derived from the associated sound (//cuckoo// is also an onomatopoeic word). = 2 Description = Note: Since there is more than one definition of //Eudynamys scolopaceus//, the description is made in reference to the nominate race //Eudynamys scolopaceus scolopaceus//, which is the first named race of the species. media type="custom" key="26910682" align="center"

<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">//Eudynamys scolopaceus// is a large cuckoo measuring 39–46 cm (from bill tip to tail tip) and weighing 190–327 g [3,4]. It has relatively short wings, but a rather long tail [3]. Its thick, large bill is slightly curved downwards and is pale green, but darker and bluish along the edges of both mandibles [2]. Its irises are crimson, while its legs and feet are pale grey [3]. It is also sexually dimorphic (i.e. males and females look different). Refer to this webpage to learn about the different parts of a bird.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 110%; line-height: 1.5;">2.1 Male
<span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: left;">Males have a glossy bluish-black plumage which may appear dark purple at times (Figure 3) [5]. <span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Figure 3: Male //Eudynamys scolopaceus// with distinctive features. Adapted from Challiyan, 2008 (Wikimedia Commons)

<span style="font-family: 'Times New Roman',Times,serif; font-size: 110%;">2.2 Female
<span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: left;">Females have a dark brown crown with rufous streaks on the head. The back, rump and wing coverts are dark brown with creamy-white or rufous speckles. The underparts are cream to whitish, with fine dark brown bars. The tail is barred dark brown and creamy-white or rufous [3,5].
 * = [[image:Figure 4_female koel front.jpg width="251" height="447"]] ||= [[image:Figure 5_female koel rear.jpg width="250" height="447"]] ||
 * <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Figure 4: Front view of Female //Eudynamys scolopaceus// with distinctive features. Adapted from K. Hari Krishnan, 2014 (Wikimedia Commons) || <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Figure 5: Rear view of Female //Eudynamys scolopaceus// with distinctive features. Adapted from Doug Janson, 2008 (Wikimedia Commons) ||

<span style="font-family: 'Times New Roman',Times,serif; font-size: 110%; line-height: 1.5;">2.3 Juvenile
<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Juveniles are uniformly black after hatching, but moult quickly to resemble adult females. However, their irises are darker (Figure 6) [2].

<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Young males often have coverts faded blackish-brown with white tips. The unmoulted wing feathers sometimes retain residual white barring, and paler tips to the edges of primaries and wing coverts (Figure 7) [2].
 * = [[image:Figure 6_immature koel 1.jpg width="374" height="353"]] ||= [[image:Figure 7_immature koel 2.jpg width="463" height="353"]] ||
 * <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Figure 6: Immature //Eudynamys scolopaceus//. Note the black crown and darker iris, with newer, moulted feathers resembling those of the adult female. <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Source: J. M. Garg, 2006 (Wikimedia Commons) || <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Figure 7: Immature male //Eudynamys scolopaceus//. Note the darker iris and unmoulted wing feathers retaining residual white barring. <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Source: Amar-Singh H.S.S., 2011 (permission obtained) ||

<span style="font-family: 'Times New Roman',Times,serif; font-size: 110%;">2.4 Diagnosis
<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">While the female //Eudynamys scolopaceus// has diagnostic features, the male is often confused with other species of birds.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Due to the similar size and plumage, it may be mistaken for a crow. In Singapore, there are two species of crows: the house crow (//Corvus splendens//) and the large-billed crow (//Corvus macrorhynchos//). Refer to Figures 3, 8, and 9 to distinguish the male //E. scolopaceus// from //C. splendens// and //C. macrorhynchos// [6].
 * = [[image:Figure 8_koel house crow.jpg width="520" height="343"]] ||= [[image:Figure 9_koel large-billed crow.jpg width="436" height="346"]] ||
 * = <span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Figure 8: House crow (//Corvus splendens//) with features shown. Adapted from Muhammad Mahdi Karim, 2009 (Wikimedia Commons) ||= <span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Figure 9: Large-billed crow (//Corvus macrorhynchos//) with features shown. Adapted from Francisco Restivo, 2010 (Wikimedia Commons) ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">While the Asian glossy starling (//Aplonis panayensis//) has a similar plumage and red irises (Figure 10), it can be distinguished from the male //E. scolopaceus// by its much smaller size (20 cm from bill tip to tail tip) [7]. <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 12pt; text-align: center;">Figure 10: Asian glossy starling (//Aplonis panayensis//) with features shown. Adapted from Yap Lip Kee, 2005 (Wikimedia Commons) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Table 1: Comparison of features among male //Eudynamys scolopaceus//, //Corvus splendens//, //Corvus macrorhynchos//, and //Aplonis panayensis//.
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Species ** ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Male Asian koel **   **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(//Eudynamys scolopaceus//) **  ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">House crow **   **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(//Corvus splendens//) **  ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Large-billed crow **   **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(//Corvus macrorhynchos//) **  ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Asian glossy starling **   **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(//Aplonis panayensis//) **  ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Length (bill tip to tail tip) ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">39–46 cm || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">40–43 cm || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">46–59 cm || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">20 cm ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Colour of iris ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Crimson || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Dark brown or black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Dark brown or black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Red ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Bill ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Thick, large, pale green || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">More slender, black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Thick, large, black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Less curved, black ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Plumage ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Glossy bluish-black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Rich black, with grey-brown neck and breast || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Glossy black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Glossy greenish-/bluish-black overall, matt black on vent ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Colour of legs and feet ** || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Pale grey || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Black || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Black ||

=<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%; line-height: 1.5;">3 Ecology and behaviour =

<span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">3.1 Vocalisation
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">With its loud, distinct calls, //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> is more often heard than seen. The calls can be heard throughout the day, even late at night, and especially early in the morning. It is especially vocal during breeding seasons. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The calls of males and females of this species differ and can be easily distinguished. The male produces a dissyllabic ‘//koo-Ooo’// call, with emphasis on the second syllable. It is usually repeated several times, increasing in pitch and loudness with each successive call, then stopped abruptly [2,5]. This type of call <span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> can be used to proclaim its territory [2]. media type="custom" key="26911364"

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The male also produces a continuous excited ‘//kwok-kwok-kwok’// call, which can be described as bubbly. However, this type of call has been observed to accompany aggressive behaviour among rival males [8].

media type="custom" key="26911374" <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The female makes a continuous and shrill //‘kik-kik-kik’// call. media type="custom" key="26910994" align="center"

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Due to //E. scolopaceus// being a brood parasite (link to 3.4 breeding and reproduction), juveniles may learn to imitate the calls of their host parents’ young, though this is not always the case. Click here for a video of a juvenile female calling like a crow.

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3.2 Habitat **
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//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> resides in a variety of natural and man-made habitats, such as light woodland, mangroves, plantations, gardens, parks, and low-density urban areas. It is rarely found in forests, though it can be found around the fringes of secondary forests [9,10].

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3.3 Diet and feeding habits **
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Adults are largely frugivorous, feeding on the fruits of plants like the Ceram palm ( //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Rhopaloblaste ceramica //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">), yellow oleander ( //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Thevetia peruviana //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">), and Indian sandalwood ( //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Santalum album //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">) [11,12]. Occasionally, they feed on insects and eggs of small birds as well [13]. They are shy, arboreal (tree-dwelling) birds, and usually do not venture out of the canopy to feed [14]. media type="custom" key="26910978" align="center"

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3.4.1 Brood parasitism **
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//<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> is a brood parasite*, and parasitises a range of hosts, including the house crow (//Corvus splendens//), Javan myna (// Acridotheres javanicus //), common myna (// Acridotheres tristis //), and long-tailed shrike (// Lanius schach //). The black-naped oriole (// Oriolus chinensis //) is also a possible host [15,16]. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">In Singapore, this brood parasitism behaviour may provide ecological benefits in the form of house crow (//C. splendens//) population management (see Distribution in Singapore).The //C. splendens// population in Singapore had risen rapidly since 1985 and were seen as pests, prompting measures such as culling to curb the population growth [17]. Brood parasitism by //E. scolopaceus// could have lowered the //C. splendens// population further, but there has not been significant correlation between changes in //E. scolopaceus// and //C. splendens// populations [18].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">* Brood parasites manipulate other individuals of the same species or another species (hosts) so that the host raises the young of the parasite. Cuckoos are the most well-known example, though this behaviour can be found in insects and fish as well.

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3.4.2 Breeding season **
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">The breeding season of //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> typically corresponds to that of its usual host. For example, if the usual host is the house crow ( //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Corvus splendens //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">), it will be roughly every three months, throughout the year [19,20].

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3.4.3 Courtship **
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Courtship behaviour in //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> has not been well-studied. However, it is known that the ‘ //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">koo-Ooo //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">’ call is used by males to proclaim their territory, and that rival males confront one another with increasing aggressiveness by flapping their wings and fanning their tails, along with loud ‘//kwok-kwok-kwok//’ calls [8]. These behaviours may better the chances of males securing mates. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Courtship feeding, in which the male feeds the female with fruits or insects, has been observed in India, Australia, and more recently in Singapore [21,22,23,24].

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">3.4.4 Egg laying and parental care **
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<span style="font-family: 'Times New Roman',serif; font-size: 11pt;">The female //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> can lay several eggs per breeding cycle, usually laying an egg in each nest she visits, after the host has laid its first egg [19,25]. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">To allow the female to approach the host’s nest to lay its egg, the male calls loudly close to the nest to advertise its presence, tempting the hosts to give chase. Meanwhile, the female conceals itself nearby. After the hosts are lured far enough, it slips into the nest to lay its egg swiftly [2]. It is unclear whether the hosts’ eggs are destroyed by the female.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The eggs of //E. scolopaceus//, although slightly smaller, resemble crow eggs. They can vary from pale greenish-yellow to greyish-green, with reddish-brown speckles and blotches [26]. Therefore, coevolution* of //E. scolopaceus// and its hosts may have resulted in these eggs matching the eggs of the hosts.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Given that //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">E. scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> eggs have a shorter incubation period than crow eggs (13–14 days compared to 16–20 days), //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">E. scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> tend to hatch first, and the hatchlings could possibly evict host eggs or chicks from the nest [25].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">After a nesting period of 19–28 days, //E. scolopaceus// fledglings continue to be fed by their foster parents for another two to three weeks [25]. Although they may mimic the begging call of the hosts’ young, it is not always the case. Therefore, it is possible that female //E. scolopaceus// do feed their young [25], if there is a lack of parental care by the host parents due to the chicks being unable to mimic such calls. However, this claim has not been supported. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;"> Video of //Corvus splendens// foster parents feeding //Eudynamys scolopaceus// chicks: media type="custom" key="26911004" align="center"

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">* Coevolution is used to describe cases where two (or more) species influence each other's evolution. =<span style="font-family: 'Times New Roman',Times,serif; font-size: 16pt;">4 Distribution =

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">4.1 Global **
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//<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> resides in much of Asia, ranging from Pakistan to Indonesia. Some populations are vagrant, migrating to warmer regions during winter. Refer to Figure 11 for the global distribution of //E. scolopaceus// [27]. <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Figure 11: Global distribution of //Eudynamys scolopaceus//. <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 120%; text-align: center;">Source: BirdLife International, 2012. IUCN Red List of Threatened Species Version 2014.3 (in accordance with Terms and Conditions)

<span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">4.2 Singapore
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">The population of //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">in Singapore has not been determined, but the 24th Annual Bird Census conducted by the Nature Society (Singapore) in 2009 found it to be the 20th most numerous avian species in Singapore [28].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The breeding population of //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">E. scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> has become widespread since 1987 [25]. This was perhaps aided by the escalating house crow (//Corvus// //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> splendens //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">) population. After population management strategies implemented in 2003 caused the //C. splendens// population to be greatly reduced, it is believed that //E. scolopaceus// continued to thrive by turning towards other host species, such as the Javan myna (// A //// cridotheres javanicus // <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">), the most common avian species in Singapore [16,28]. In addition, the now more scattered <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">//C. splendens// population results in increased vulnerability to brood parasitism by //E. scolopaceus//, since nesting in groups could lower the occurrence of brood parasitism [16].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The proliferation of //E. scolopaceus// in Singapore has led to some residents’ unhappiness over their loud calls, while others are more tolerant of them. Read about the different opinions at the following links: link 1, link 2, link 3, link 4.

= <span style="display: block; font-family: 'Times New Roman',Times,serif; font-size: 16pt; text-align: left;">5 Conservation status = media type="custom" key="26911140" align="center"

<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Conservation status: least concern (IUCN 3.1) [27] <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Although the population of //Eudynamys scolopaceus// is not known, it occupies a large range globally, and its population trend appears to be stable [27]. Being able to survive in some urban areas (see Habitat), urbanisation does not appear to pose much of a threat to it.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">In Singapore, //E. scolopaceus// is a species protected under the Wild Animals and Birds Act, and is not listed under the Schedule. = <span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">6 Taxonomy and systematics =

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">6.1 Taxonavigation **
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Table 2 provides a hierarchical summary of clade names in the taxonomic classification of //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Table 2: Taxonavigation of //Eudynamys// //scolopaceus//.
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Animalia ||
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Chordata ||
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Aves ||
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Cuculiformes ||
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Cuculidae ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys // ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">E. scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> (Linnaeus, 1758) ||

//<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">E. scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> was originally described by Linnaeus as //Cuculus scolopaceus// based on a specimen he received from the Malabar region [29]. It was later moved to the new genus //Eudynamys// by Vigors and Horsfield in 1827 as it has a relatively more powerful bill and stronger tarsi and feet than true //Cuculus// cuckoos [29].

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">6.2 Phylogeny **
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<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">There has been much uncertainty on the phylogeny of taxa in Aves. Mayr (2008) collated the genetic and morphological information from previous studies to produce the following trees in Figure 12 [30]. The Tree of Life Web Project offers a comprehensive explanation on interpreting tree diagrams.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">Figure 12: Phylogeny of Cuculidae among Neognathous birds derived from (a) a Bayesian analysis of five nuclear genes (β-fibrinogen, c-myc, RAG-1, myoglobin, and ornithine decarboxylase) and (b) a parsimony analysis of 2954 morphological characters. Nodes with posterior probability below 95% were collapsed in (a), while only bootstrap support values above 80% were shown in (b). Source: Mayr, 2008 (permission pending) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Phylogeny within Cuculidae has also been unclear. The Tree of Life Web Project displays the following tree for the subfamily Cuculinae, consisting of Old World parasitic cuckoos (Figure 13) [31]. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 13: Phylogeny within Cuculinae, with position of //Eudynamys// shown. Adapted from Tree of Life Web Project, 2007 (permission pending) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">With respect to 135 osteological characters, Hughes (2000) placed //Eudynamys// as a sister taxon to //Urodynamis// (Figure 14) [32]. This classification was shown to support the evolution of brood parasitism in a single event instead of three events as previously proposed, as all brood parasitic cuckoos formed a clade [32]. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 14: Optimal hypothesis of phylogeny for 33 cuckoo genera, with position of the monophyletic clade containing //Eudynamys// and //Urodynamis// shown. Source: Hughes, 2000 (permission pending)

**<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">6.3 Variations in taxonomic classification **
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//<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Eudynamys scolopaceus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">displays variations across its geographical range, with several island populations established. Some consider the black-billed koel (// E. melanorhynchus //) and Pacific koel (// E. orientalis //) conspecific with //E. scolopaceus//. In this case, the “combined” species //E. scolopaceus// is known as the common koel. Another way of classification is to consider // E. melanorhynchus // <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;"> as a separate species, with <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">//E. scolopaceus// encompassing // E. orientalis // and all its subspecies, with the exception of the Australian koel (// E. cyanocephalus //). Alternatively, the three are considered separate species, owing to differences in plumage, bill colour, and vocalisations. The last view is becoming increasingly accepted [23].

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">According to the 2012 Clements Checklist of Birds of the World, a regularly updated checklist maintained by the Cornell Lab of Ornithology, 18 species and subspecies have been identified [1]. A summary is provided in Table 3.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Table 3: Species and subspecies recognised by the 2012 Clements Checklist, with their respective geographical range.
 * **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Species ** ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Subspecies **  ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Authority **  ||  **<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Geographical range **  ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Linnaeus, 1758) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Tropical southern Asia to south China and Greater Sundas ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus scolopaceus // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Linnaeus, 1758) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Nepal to Pakistan, India, Sri Lanka, Laccadives and Maldives ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus chinensis // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Cabanis & Heine, 1863 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Southern China and Indochina; winters to Borneo ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus harterti // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Ingram, 1912 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Hainan (Southern China) ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus malayanus // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Cabanis & Heine, 1863 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Northeast India to Thailand, Malaya, Sumatra, Borneo and Lesser Sundas ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus simalurensis // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Simeulue, Indonesia (off West Sumatra) ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus mindanensis // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Linnaeus, 1766) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Philippines, Palawan, Sulu Archipelago, Sangihe, Indonesia and Talaud Islands ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus corvinus // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">North Moluccas (Morotai, Halmahera, Ternate, Tidore and Bacan) ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys scolopaceus frater // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Northern Philippines (Calayan and Fuga) ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Linnaeus, 1766) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">South Moluccas (Buru, Manipa, Kelang, Seram, Ambon, Watubela) ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis orientalis // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Linnaeus, 1766) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">South Moluccas (Buru, Manipa, Kelang, Seram, Ambon, Watubela) ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[//rufiventer or minimus//] || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Lesson, 1830) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">New Guinea ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis salvadorii // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Hartert, 1900 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Bismarck Archipelago ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis alberti // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Rothschild & Hartert, 1907 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Solomon Islands ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys orientalis picatus // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Müller, 1843 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Kai Islands and Sumba to Timor and Roma ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys cyanocephalus // ||  ||   || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Australia ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys cyanocephalus cyanocephalus // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(Latham, 1801) || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Northern Queensland to Southern New South Wales; winters to Moluccas ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys cyanocephalus subcyanocephalus // || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Matthews, 1912 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Northwest Australia to Northwest Queensland; winters to South Moluccas ||
 * //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys melanorhynchus // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Müller, 1843 || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Sulawesi, Sula Islands ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys melanorhynchus melanorhynchus // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Sulawesi ||
 * || //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Eudynamys melanorhynchus facialis // ||  || <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Sula Islands ||

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Figure 15 shows the distribution of four //E. scolopaceus// subspecies: //E. s. chinensis, E. s. malayanus, E. s. mindanensis, and E. s. scolopaceus.//



<span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 15: Geographical distribution of //Eudynamys scolopaceus chinensis//, //E. s. malayanus//, //E. s. mindanensis//, and //E. s. scolopaceus//. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Source: xeno-canto, 2014 (permission pending) = <span style="font-family: 'Times New Roman',Times,serif; font-size: 16pt;">7 Biogeography =

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<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Using the Principle of Parsimony based on primary and secondary “Brooks Parsimony Analysis” (BPA), Posso and Donatelli (2012) proposed that arboreal cuckoos originated in South America, and dispersed to inhabit other continents thereafter (Figure 17) [33]. This is because //Coccyzus//, the most basal genus in Cuculinae, is found in South America [33]. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 16: Strict concensus cladogram with bootstrap support values in parentheses. Spotted lines denote terrestrial/arboreal habits, thick lines denote terrestrial cuckoos, while thin lines denote arboreal cuckoos. Source: Posso & Donatelli, 2012 (permission pending) <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 17: Current distribution of basal cuckoos, with arrows showing the subsequent dispersion of arboreal cuckoos. <span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Source: Posso & Donatelli, 2012 (permission pending) = <span style="font-family: 'Times New Roman',Times,serif; font-size: 16pt;">8 Molecular data =

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<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The database GenBank contains the sequences of genes such as cytochrome oxidase subunit I (COI), cytochrome b (cytb), and NADH dehydrogenase subunit 2 (ND2), although the full genome of //Eudynamys scolopaceus// has not been sequenced.

= <span style="font-family: 'Times New Roman',Times,serif; font-size: 120%;">9 References = <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[1] Avibase, 2014. //Eudynamys// [//scolopaceus, orientalis, cyanocephalus or melanorhynchus//]. //Avibase//. Retrieved 11 November 2014, from http://avibase.bsc-eoc.org/species.jsp?avibaseid=5D43995351C97BC6 <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[2] Bird Ecology Study Group, 2011. Birds of Singapore: Asian Koel (//Eudynamys scolopacea//). //The Birds of Singapore//. Retrieved 11 November 2014, from http://singaporebirds.net/npassers_01/asian_koel.html <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[3] Oiseaux-Birds, n.d. Asian Koel. //Oiseaux-Birds//. Retrieved 11 November 2014, from http://www.oiseaux-birds.com/card-asian-koel.html <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[4] Dunning, J. B. Jr., 1992. // CRC Handbook of Avian Body Masses // <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">. CRC Press. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">ISBN 978-0-8493-4258-5 <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[5] Encyclopaedia of Life, n.d. Descriptions and Articles about the Common Koel (//Eudynamys scolopaceus//). //Encyclopaedia of Life//. Retrieved 11 November 2014, from http://eol.org/pages/1064881/details#cite_note-payne-10 <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[6] Singapore Birds, 2012. Crows. //Singapore Birds//. Retrieved 11 November 2014, from http://singaporebirds.blogspot.sg/2012/06/crows.html <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[7] Oiseaux-Birds, n.d. Asian Glossy Starling. //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Oiseaux-Birds. //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Retrieved 22 November 2014, from http://www.oiseaux-birds.com/card-asian-glossy-starling.html <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[8] Wee, Y. C., 2007. Koel Confrontation. //Bird Ecology Study Group//. Retrieved 11 November 2014, from http://www.besgroup.org/2007/03/21/koel-confrontation/ <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[9] BirdForum, 2014. Asian Koel. //BirdForum//. Retrieved 11 November 2014, from http://www.birdforum.net/opus/Asian_Koel <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[10] The Birds of NUS, 2012. Asian Koel. //The Birds of NUS//. Retrieved 11 November 2014, from http://nusavifauna.wordpress.com/2012/07/25/asian-koel/ <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[11] <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">Hegde, S. G., Uma Shaanker, R. & Ganeshaiah, K. N., 1991. Evolution of seed size in the bird-dispersed tree //Santalum album// L.: a trade-off between seedling establishment and dispersal efficiency. //Evol. Trends. Plants// 5: 131–135. <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">[12] <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Krishnan, M <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">., 1952. Koels (//Eudynamis scolopaceus//) eating the poisonous fruit of the Yellow Oleander. //J. Bombay Nat. Hist. Soc.// 50: 943–945. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[13] Ali, S. & Ripley, D.S., 1969. //Handbook of the Birds of India and Pakistan.// Vol. 3. Oxford University Press, London. 380 pp. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[14] Roberts, T. J., 1991. //The Birds of Pakistan Vol. 1//. Oxford University Press, Karachi. 666 pp. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[15] Lowther, P. E., 2011. Host List of Avian Brood Parasites. Retrieved 11 November 2014, from http://www.fieldmuseum.org/sites/default/files/Cuckoo-12Sep2011.pdf <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[16] Begum, S., Moksnes, A., Røskaft, E. & Stokke, B. G., 2011. Factors influencing host nest use by the brood parasitic Asian Koel (//Eudynamys scolopacea//). //J. 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Retrieved 11 November 2014, from http://www.ava.gov.sg/NR/rdonlyres/BACEEC21-2D82-40D3-AB42-76F65262D890/27022/Advisoryonasiankoels.pdf <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[21] Tan, S., 2010. Courtship feeding in Asian koel. //Bird Ecology Study Group.// Retrieved 11 November 2014, from http://www.besgroup.org/2010/02/19/courtship-feeding-in-asian-koel/ <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[22] Higgins, P. J. (ed.), 1999. //Handbook of Australian, New Zealand and Antarctic birds. Vol. 4, Parrots to dollarbirds//. Oxford University Press, London. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[23] Lamba, B. S., 1969. The nidification of some common Indian birds – part 12. // J. Bombay Nat. Hist. Soc. //66: 72-80. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[24] Finn, F., 1917. //The Birds of Calcutta//. Thacker, Spink & Co., Calcutta. 136 pp. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[25] Payne, R. B., 2005. //The Cuckoos//. Oxford University Press, New York. 644 pp. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[26] Baker, E. C. S., 1927. //Fauna of British India: Birds. Vol. 4//. Taylor & Francis, London. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[27] BirdLife International, 2012. //Eudynamys scolopaceus//. //The IUCN Red List of Threatened Species. Version 2014.2////IUCN//. Retrieved 11 November 2014, from http://www.iucnredlist.org/details/22684049/0 <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[28] Lim, K. S., 2009. //Report on the 24th Annual Bird Census//. Retrieved 11 November 2014, from http://wildbirdsingapore.nss.org.sg/2009ABC.pdf <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[29] Long, G. (ed.), 1833. //The Penny Cyclopaedia of the Society for the Diffusion of Useful Knowledge. Vol. 8.// Charles Knight, London. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[30] Mayr, G., 2008. Avian higher-level phylogeny: well-supported clades and what we can learn from a phylogenetic analysis of 2954 morphological characters. //J Zool Syst Evol Res// 46: 63–72. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[31] Tree of Life Web Project, 2007. Cuculinae. //Tree of Life Web Project//. Retrieved 11 November 2014, from http://tolweb.org/Cuculinae/103814 <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[32] Hughes, J. M., 2000. Monophyly and phylogeny of cuckoos (Aves, Cuculidae) inferred from osteological characters. //Zool. J. Linn. Soc.// 130: 263–307. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">[33] Posso, S. R. & Donatelli, R. J., 2012. Biogeography on the early distribution of cuckoos (Aves: Cuculiformes). //Zoologia// 29: 187 <span style="color: #252525; font-family: 'Times New Roman',serif; font-size: 12pt;">–194.