Aethopyga+siparaja

Crimson sunbird  //Aethopyga siparaja// (Raffles, 1822) Say hello to Singapore's (unofficial) national bird! toc ==

=1. Introduction=

1.1 A brief description
The crimson sunbird is a tiny sunbird that belongs to the family Nectariniidae. It can be found in the tropical regions of southern and south-eastern Asia, ranging from north-western India to eastern Indonesia. Its preferred habitats are forests and plantations. It is generally nectarivorous, but may also feed on insects.

1.2 Singapore's unofficial national bird
In 2002, the crimson sunbird received the most votes to be Singapore's national bird in a poll started by Nature Society Singapore (the white-bellied sea eagle came second). Voters felt that its small size and bright red plumage were good representatives of the country. On 1st November 2015, Dr Shawn Lum, president of Nature Society Singapore, publicly announced that the crimson sunbird had been declared the national bird of Singapore. The following day, however, Dr Lum clarified that this was not official and was still pending an agreement with three government departments. This remains the case till today.

1.3 Not a hummingbird
Crimson sunbirds and other sunbirds are often mistaken for hummingbirds due to their similar appearance in terms of size and colouring. However, this is actually a result of convergent evolution, as both fulfil similar ecological niches by feeding on nectar and therefore play a key role in pollination. Sunbirds and hummingbirds are not closely related at all - hummingbirds' closest cousins are actually swifts and nightjars. In addition, sunbirds and hummingbirds do not coexist anywhere in the world. Sunbirds are found in Asia, Africa and Australia while hummingbirds are found only in North and South America. So the next time you're confused as to whether a bird you've seen is a sunbird or a hummingbird, all you have to do is know which continent you're on!

=2. Etymology=

Etymology describes how a species got its scientific name. For the crimson sunbird, the Latin species epithet //siparaja// was given to it by Sir Stamford Raffles based on the Malay name 'burong sepah raja'. 'Burong' translates to 'bird', 'sepah' refers to the sweet juice of the betel nut, and 'raja' means 'king'.

=3. Biology=

3.1 Habitat
The crimson sunbird occupies a large variety of habitats. It can be found in various forest types (eg. riverine, peatswamp), typically at the forest edge, but also in more open areas such as grasslands, scrub, mangroves, woodlands, parks or gardens. With regards to altitude, it is usually found at heights of below 915m, but can be found at altitudes of 1,690m in Bhutan and 1,800m in Nepal during the summer. Below is a video of a male crimson sunbird in MacRitchie Reservoir:

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3.2 Behaviour
Crimson sunbirds may forage either low or in the canopy; singly, in pairs or in larger family parties. Males often sing from exposed branches in the canopy and can get extremely aggressive with each other as they often establish feeding territories on flowering shrubs or trees. Crimson sunbirds can also reduce their metabolism at night depending on the ambient temperature (reduce by 54% at 5°C, and 62% at 25°C).

3.3 Feeding habits
The crimson sunbird feeds primarily on nectar. It seeks out flowers with high sugar content and pierces the base of the corolla to get to the nectar directly. It plays an important ecological role in pollination, as many native flowering plants rely on sunbirds for this purpose. Many plants have tubular-shaped flowers that actually exclude other pollinators such as bees from feeding on them, just so that the crimson sunbird can do so. The crimson sunbird may also feed on insects and spiders during the breeding season to feed its young.



3.4 Breeding and nesting
The male crimson sunbird is known to perform a mating display, during which it not only sings, but also fluffs out its tertial feathers to display its yellow rump which usually remains unexposed. Watch it in the video below:

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The nest of a crimson sunbird couple is usually attached to palm fronds. It is made of vegetable down, cobweb, grass, roots, and may be found under overhanging banks, such as in Gunung Kinabalu, Sabah.

Egg-laying months are typically March and April. One to three eggs are laid at a time. Eggs are of variable colouring, but are usually pinkish and sporadically marked with dark reddish-brown spots.

3.5 Parasites
The crimson sunbird is a host to //Leucocytozoon sp.//, a group of intracellular blood parasites of birds. //Leucocytozoon sp.// are known to parasitize other sunbirds, as well as at least three other families: Lybiidae (barbets), Picacthartidae (junglefowl) and Pycnonotidae (bulbuls). However, the exact relationship between the birds and the parasites is yet to be determined.

=4. Description and Identification=

From Hails and Jarvis (1987), Jeyarajasingam and Pearson (1999) or Yong, Lim and Lee (2016) unless otherwise stated.

4.1 General Identification
The crimson sunbird is tiny - an adult is only 11cm long on average. It has a thin, down-curved bill of medium length and a brush-tipped tubular tongue, both to facilitate nectar feeding. It also displays sexual dimorphism, with males having the bright crimson colour that gives the bird its vernacular name, and females being more drab. This can be seen from Figures 2a and 2b, which indicate distinctive features of the male and female crimson sunbird for identification purposes. Table 1 below the two figures presents a more detailed description and comparison between the male and the female, and also includes features of the male in eclipse phase - where it has temporary dull plumage after breeding.
 * [[image:crimson_sunbird_male_labelled.jpg width="538" height="433" caption="Figure 3a. Photo of a male Crimson Sunbird with labelled body parts. Photo by Lip Kee (2008) © and edited by Chai Wen Xuan" link="http://www.flickr.com/photos/lipkee"]] || [[image:crimson_sunbird_female_labelled.jpg width="537" height="431" caption="Figure 3b. Photo of a female Crimson Sunbird with labelled body parts. Photo by Lip Kee (2007) © and edited by Chai Wen Xuan" link="http://www.flickr.com/photos/lipkee"]] ||
 * Table 1.** Detailed description of morphology of male, male eclipse and female crimson sunbird

Click [|here] for more information on bird topography.
 * **Body Part(s)** || **Male** || **Male eclipse** || **Female** ||
 * **Forehead** || Dark iridescent green || Dark olive || Dark olive green ||
 * **Head,** **chin, neck** || Bright crimson || Dark olive || Dark olive green ||
 * **Bill** || Medium length, thin, down-curved || Medium length, thin, down-curved || Medium length, thin, down-curved ||
 * **Throat** || Bright crimson || Diffused red || Pale yellow ||
 * **Malar stripes** || Two, dark iridescent violet || Absent || Absent ||
 * **Breast** || Bright crimson || Diffused red || Pale yellow ||
 * **Belly** || Grey || Dark olive || Pale yellow ||
 * **Scapulars** || Bright crimson || Dark olive || Pale yellow ||
 * **Primaries and secondaries** || Brown || Dark olive || Dark olive green ||
 * **Rump** || Yellow || Dark olive || Dark olive green ||
 * **Tail** || Elongate (extends 3cm), iridescent violet || Not elongate, dark olive || Not elongate, dark olive green ||

4.2 Diagnosis
Due to the distinct sexual dimorphism of the crimson sunbird, males and females are morphologically similar to different heterospecifics. Tables 2a and 2b describe the species similar to male and female crimson sunbirds respectively. All species described have overlapping ranges with the crimson sunbird. However, only the brown-throated sunbird, olive-backed sunbird and purple-throated sunbird can be found in Singapore.


 * Table 2a.** Description of morphologically similar species to male crimson sunbirds
 * [[image:black_throated_sunbird.jpg width="300" height="244" caption="Figure 4a. Photo of black-throated sunbird taken by Jason Thompson (2012) ©" link="https://www.flickr.com/photos/79492850@N00/"]] || **Black-throated sunbird (//Aethopyga saturata//)**


 * (NOT FOUND IN SINGAPORE) **
 * Mantle maroon
 * Crown, rump, tail iridescent violet
 * Throat, breast black with violet tinge
 * Belly pale greyish-yellow
 * Same size as crimson sunbird (11cm)
 * Range: Himalayas and southern China through South-east Asia to Malay Peninsula ||
 * [[image:scarlet_sunbird.jpg width="303" height="207" caption="Figure 4b. Photo of scarlet sunbird by Caroline Granycome (2013) ©" link="https://www.flickr.com/photos/cgranycome/"]] || **Scarlet sunbird (//Aethopyga// //temminckii//)**


 * (NOT FOUND IN SINGAPORE) **
 * Morphologically very similar to crimson sunbird
 * Head, breast, mantle scarlet
 * Malar stripe iridescent purple
 * Rump yellow
 * Belly grey
 * Elongate tail 3cm, scarlet
 * Same size as crimson sunbird (11cm)
 * Range: Malay Peninsula to Greater Sundas ||


 * Table 2b.** Description of morphologically similar species to female crimson sunbirds
 * [[image:taxo4254/brown_throated_sunbird_female1.jpg width="308" height="212" caption="Figure 5a. Photo of female brown-throated sunbird by Melvin Yap (2014) ©" link="https://www.flickr.com/photos/mjmyap/"]] || **Brown-throated sunbird (//Anthreptes malaccensis//)**


 * Upperparts olive
 * Underparts bright yellow
 * Larger than crimson sunbird (14cm)
 * Range: Myanmar to southern Indo-China through Malay Peninsula, south to Greater Sundas and Philippines ||
 * [[image:taxo4254/olive_backed_sunbird_female.jpg width="304" height="394" caption="Figure 5b. Photo of female olive-backed sunbird taken by Melvin Yap (2015) ©" link="https://www.flickr.com/photos/mjmyap/"]] || **Olive-backed sunbird (//Nectarinia jugularis//)**


 * Underparts bright yellow
 * Tail very dark brown
 * Same size as crimson sunbird (11cm)
 * Range: Myanmar and southern China through South-east Asia to northern Australia and Solomon Islands ||
 * [[image:taxo4254/purple_throated_sunbird_female.jpg width="309" height="250" caption="Figure 5c. Photo of female purple-throated sunbird taken by Lip Kee (2006) ©" link="https://www.flickr.com/photos/lipkee"]] || **Purple-throated sunbird (//Nectarinia sperata//)**


 * Upperparts dark olive
 * Underparts whitish-yellow
 * Tail very dark brown
 * Smaller than crimson sunbird (10cm)
 * Range: North-east India and Myanmar to southern Vietnam through Malay Peninsula to Greater Sundas and Philippines ||

=4. Vocalization=

The crimson sunbird's call can be described as a high-pitched, sibilant //seep-seep//. Below is a recording of a crimson sunbird's call from the Central Catchment Nature Reserve in Singapore, retrieved from [|xeno-canto]:

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=5. Distribution=

5.1 Global range
The crimson sunbird is exclusively a resident of southern Asia, but has an extremely large range within that area - its estimated extent of occurrence (EOO) is 4,920,000 km 2. Its range covers countries such as India, Bangladesh, China, Myanmar, Thailand, Malaysia, Singapore, the Philippines and Indonesia. It has been described as usually common in all those areas, with the exception of Buton Island in Indonesia, where it is described as uncommon or rare. Figure 4 below shows the range map of the crimson sunbird (demarcated in yellow).



** 5.2 Distribution in Singapore **
The crimson sunbird is commonly found in the Central Catchment Nature Reserve, Bukit Timah Nature Reserve, Sungei Buloh Wetland Reserve and Bukit Batok Nature Park

=7. Conservation Status=

7.1 Global status
The crimson sunbird is evaluated as Least Concern on the IUCN Red List as it does not approach the thresholds for Vulnerable classification under various criteria. It has an extremely large range of almost 5 million km 2 as mentioned in Section 5, and thus does not approach the threshold for Vulnerable under the range size criterion (EOO <20,000km 2 ). It has a stable population trend, and hence does not approach the Vulnerable threshold under the population trend criterion (>30% decline over 10 years or three generations). Its population size, though unquantified, is not estimated to approach the Vulnerable threshold under the population size criterion (<10,000 mature individuals with an estimated continuing decline of >10% in 10 years or three generations). Therefore, the crimson sunbird is evaluated as Least Concern.

7.2 Singapore status
The crimson sunbird is not under any conservation threat in Singapore either. It is listed as 'common' in the 2015 Checklist of the Birds of Singapore compiled by Nature Society Singapore, as well as the Singapore Red Data Book.

=8. Taxonomy and Systematics=


 * ...mostly for the biologists :) **

8.1 Original description
The protonym of the crimson sunbird is //Certhia siparaja//. It was first described by Sir Stamford Raffles in 1822. This description was printed in the Transactions of the Linnean Society of London, and is shown in Figure 7 below.



The type specimen of the crimson sunbird was collected in 1872, and is now located in the Natural History Museum of London. Details of the type specimen are shown in Figure 8 below.



8.2 Taxonomic hierarchy
Below is the taxonomic classification of the crimson sunbird. Taxonomic ranks are not included as they may not provide objective taxonomic information. For more on this, click [|here].

- Animalia -- Chordata --- Aves Passeriformes - Nectariniidae (Vigors, 1825) -- //Aethopyga// (Cabanis, 1850) --- //Aethopyga siparaja// (Raffles, 1822)

**8.3 Subspecies**
The crimson sunbird is a polytypic species. Its 15 subspecies are listed in Table 3 below. //Aethopyga siparaja siparaja// is the only subspecies found in Singapore


 * Table 3.** Description of subspecies of the crimson sunbird and their distributions.
 * **No. ** || **Subspecies ** || **Distribution ** ||
 * 1 || //Aethopyga siparaja siparaja// (Raffles, 1822) (nominate species) || Malay Peninsula (south of Narathiwat), Anamba Islands (east of Peninsular Malaysia), Sumatra (except Aceh) and satellite islands, and Borneo and associated small islands (except Natunas). ||
 * 2 || //A. s. labecula// (Horsfield, 1840) || Bhutan, northeastern India (north West Bengal, Assam, Arunachal Pradesh, Meghalaya, Nagaland, Manipur) and Bangladesh south to Chittagong Hills, east to Myanmar (except in the south), southern China (south Yunnan), northwestern Laos and northwestern Vietnam ||
 * 3 || //A. s. cara// (Hume, 1874) || South Myanmar and Thailand ||
 * 4 || //A. s. nicobarica// (Hume, 1873) || South Nicobar Island (Great and Little Nicobars, Kondal and Meroe Islands) ||
 * 5 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. tonkinensis// (E. J. O. Hartert, 1917) || <span style="font-family: Arial,Helvetica,sans-serif;">Northeastern Vietnam and south China (southeastern Yunnan east to western Guangdong) ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">6 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. owstoni// (Rothschild, 1910) || <span style="font-family: Arial,Helvetica,sans-serif;">Naozhou Island (southwestern Guangdong) in south China ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">7 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. mangini// (Delacour and Jabouille, 1924) || <span style="font-family: Arial,Helvetica,sans-serif;">Southeastern Thailand and central and south Indochina ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">8 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. insularis// (Delacour and Jabouille, 1928) || <span style="font-family: Arial,Helvetica,sans-serif;">Phu Quoc Island, off southern Cambodia ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">9 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. trangensis// (Meyer de Schauensee, 1946) || <span style="font-family: Arial,Helvetica,sans-serif;">Southern Thailand and northern Malay Peninsula ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">10 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. natunae// (Chasen, 1935) || <span style="font-family: Arial,Helvetica,sans-serif;">Natuna Islands ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">11 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. heliogona// (Oberholser, 1923) || <span style="font-family: Arial,Helvetica,sans-serif;">Java ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">12 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. flavostriata// (Wallace, 1865) || <span style="font-family: Arial,Helvetica,sans-serif;">Northern Sulawesi ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">13 || <span style="font-family: Arial,Helvetica,sans-serif;">//A. s. beccarii// (Salvadori, 1875) || <span style="font-family: Arial,Helvetica,sans-serif;">Central, south and southeastern Sulawesi, Kabaena, Muna and Butung ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">14 || <span style="font-family: Arial,Helvetica,sans-serif;">[//A. s. magnifica//] (Sharpe, 1876) (Possibly a separate species) || <span style="font-family: Arial,Helvetica,sans-serif;">West-central Philippine Islands, Marinduque, Tablas, Sibuyan, Panay, Negros, Cebu ||
 * <span style="font-family: Arial,Helvetica,sans-serif;">15 || <span style="font-family: Arial,Helvetica,sans-serif;">[//A. s. seheriae//] (Tickell, 1833) (Possibly a separate species) || <span style="font-family: Arial,Helvetica,sans-serif;">Himalayan foothills in India from W Himachal Pradesh (Kangra) east to Sikkim and Bhutan, south to northern West Bengal, eastern Bihar,eastern Madhya Pradesh and Orissa (possibly northern Andhra Pradesh), and western Bangladesh ||

8.4 Phylogeny
In 2014, Jarvis et al. performed a genome-scale phylogenetic analysis of 48 avian species representing all orders of Neoaves using phylogenomic methods created to handle genome-scale data. The total evidence nucleotide alignment partitioned by data type (introns, UCEs, and first and second exon codon positions) analyzed using Maximum Likelihood analysis under the GTR+GAMMA model of sequence evolution yielded the Total Evidence Nucleotide Tree, a highly-resolved phylogenetic tree that confirmed previously undetermined sister or close relationships.

The order Passeriformes can be split into New Zealand wrens, suboscines and oscines (commonly referred to as songbirds). The crimson sunbird belongs to the oscines. Figure 9 below illustrates the position of the crimson sunbird in the Total Evidence Nucleotide Tree. (In reference to Section 1.3, the position of hummingbirds is illustrated as well, showing how distant the relationship between sunbirds and hummingbirds actually is.)



Within the oscines, there are three superfamilies - Sylvioidea, Muscicapoidea and Passeroidea. Ericson and Johansson (2003) sequenced multiple nuclear and mitochondrial genes and analyzed the data via parsimony jackknifing and Bayesian analysis to obtain four main clades, as shown in Figure 10 below.



Monophyly of this clade is supported in both parsimony jackknifing (94%) and Bayesian analysis (100%), and the maximum likelihood tree in Figure 11 below shows that the //Aethopyga// sunbirds are the first outgroup, together with the flowerpeckers.

Hosner, Nyári and Moyle (2013) then reconstructed the phylogenetic relationships within the //Aethopyga// genus through further mitochondrial and nuclear DNA sequencing. They used maximum likelihood ancestral state biogegraphical reconstructions to examine the ancestral distribution and colonization history of //Aethopyga//. To determine whether the diversification at each node occurred within a continent, across a shallow-water barrier, across a deep-water barrier or within an island, they used statements based on the phylogeny, current distribution of species and bathymetric reconstructions. Ultimately, they derived the position of //Aethopyga siparaja//, as shown in Figure 12 below.



However, the figure also shows that //Aethopyga siparaja// could be treated as a complex of geographically separated species (or allospecies). The most obvious example would be the subspecies //Aethopyga siparaja magnifica// (mentioned as possibly a separate species in Section 8.3 above) which appears to be the sister group of all the //Aethopyga siparaja// subspecies. Prior research on isolation, as well as molecular results , has supported the placement of //A. s. magnifica// as a separate sister species to all other //A. siparaja// subspecies. Further splits within //A. siparaja// might be warranted but similarly rigorous experimentation and DNA sequencing will first have to take place.

8.5 DNA barcoding
Genbank contains many nucleotide sequences for the crimson sunbird, such as the NADH dehydrogenase subunit 2 (ND2) gene, NADH dehydrogenase subunit 3 (ND3) gene, recombination activating protein 1 (RAG1) gene, ornithine decarboxylase (ODC) gene and the cytochrome b (cytb) gene. They can be found [|here]. However, do note that they are not official barcodes as they were not sequenced from the cytochrome c oxidase I (COI) gene.

=9. Glossary=

**nectarivorous:** feeds mainly or exclusively on nectar **convergent evolution:** the process whereby organisms that are not closely related develop similar traits, due to them adapting to similar environments and ecological niches **ecological niche:** the specific role a species plays in its environment **sexual dimorphism:** a distinct difference in size or physical appearance between both sexes of the same species **heterospecific:** of a different species (opposite: conspecific) **sibilant:** characterized by a hissing sound **extent of occurrence**: total surface area over which a species can be found naturally **protonym:** unofficial (usually first) name for a new species **type specimen:** the individual specimen (or set of specimens) on which the description and name of a new species is based **polytypic:** contains more than one taxonomic category of the next lower rank (opposite: monotypic)

=10. Additional information=

Crimson sunbird in MacRitchie Reservoir: https://youtu.be/CWwc8umb7YU Male crimson sunbird mating display: https://youtu.be/uzY0dZ1dfIU More crimson sunbird vocalization recordings: http://www.xeno-canto.org/species/Aethopyga-siparaja

=11. Literature Cited=