No new thing under the sun? Finding sunbird species in Wallacea

A male Wakatobi Sunbird

Our understanding of how species interact and evolve depends on accurate knowledge of the species that exist on Earth. There are still many species to be identified, however, even in evolutionarily significant regions such as Wallacea in central Indonesia, site of Alfred Russel Wallace’s pioneering work. Our new paper, completed jointly with researchers from Universitas Halu Oleo and just published in the Zoological Journal of the Linnean Society, draws on work carried out in Wallacea to identify multiple unrecognised species in the beautiful sunbird family. Made using modern genetic, acoustic, and statistical techniques, these discoveries add to our understanding of how life evolved in this region and reinforce some of Wallace’s original ideas.

The key discovery is the “Wakatobi Sunbird Cinnyris infrenatus”, a species endemic to the small islands of the Wakatobi archipelago, off Southeast Sulawesi in central Indonesia. The Wakatobi Islands have been separated from other landmasses since they first rose out of the sea, and so there has been plenty of time for their populations to evolve in isolation and produce endemic taxa, found nowhere else on Earth. The Wakatobi Islands have been recognised as a Key Biodiversity Area for their importance to the survival of biodiversity. The Wakatobi Sunbird is the latest endemic species to be identified by our research group, following previous work on the Wakatobi Flowerpecker and the double discovery of the Wakatobi White-eye and Wangi-wangi White-eye. It’s important that we know all of the species of Southeast Sulawesi and the Wakatobi Islands, because this region acts as a “natural laboratory” for the study of evolutionary processes such as cryptic sexual dimorphism, the “supertramp strategy”, and the links between behaviour and population divergence. The Wakatobi Sunbird is currently treated as a subspecies of the widespread Olive-backed Sunbird (Cinnyris jugularis), but our findings indicate that the Olive-backed Sunbird is actually made up of at least 4 reproductively isolated species.

The sunbirds (Nectariniidae) fill a similar niche in Africa, Asia, and Australia to the hummingbirds of the Americas. They are small birds with long bills that help them extract nectar from flowers. Like the hummingbirds, many sunbirds (males particularly) exhibit brightly coloured plumage, with beautiful iridescent or “metallic” feathers that reflect the sunlight. In fact, the naturalist William Jardine tells us in his 1843 volume on the sunbirds that sunbirds get their name “from their brightly-tinted dress, appearing in higher splendour when played on by the sun-beams”. For hundreds of years, ornithologists have used the patterns and colours of these feathers to identify sunbird species. Now, however, we can combine multiple forms of data to uncover patterns that weren’t clear from plumage alone. Our paper also looked at the Black Sunbird (Leptocoma aspasia), a species with male plumage that’s hard to examine because it mostly looks jet-black, except when the sun hits it in the right way to reveal other colours. We found a genetic split in this species that had not been suggested by any previous work, probably due to its plumage being less informative.

Finding species like these isn’t just interesting for its own sake. It is also our best evidence to understand how evolution produces new species. This is particularly interesting in a region like Wallacea, which played such a significant role in the development of evolutionary biology. The observations that Wallace made around this region led him to discover evolution by natural selection, work which was published jointly with Charles Darwin in 1858 in the same journal where our sunbird paper has just appeared.

One of the observations that inspired Wallace’s evolutionary thinking was the importance of biogeographic barriers. Wallace noticed that the animals found on Sulawesi are markedly different from those on neighbouring Borneo, evidence that species would evolve on one island and then have difficulty crossing over. This boundary came to be known as Wallace’s Line. We now understand that it represents the beginning of the deep waters of the Wallacea region, which persisted even when sea levels were lower, unlike the shallower waters of the adjoining Sunda Shelf which gave rise to land bridges. A similar barrier to the east came to be known as Lydekker’s Line. As seen in the map below, the range of the Olive-backed Sunbird (in yellow) is currently thought to cross both Wallace’s Line and Lydekker’s Line, while the Black Sunbird (in purple) stops at Wallace’s Line but crosses Lydekker’s Line. It’s quite remarkable to imagine these dainty little birds maintaining gene flow across barriers which block so many other organisms! Our work, however, has indicated that the Olive-backed Sunbird populations on either side of Wallace’s Line actually represent separate species. The same is true of Black Sunbird populations divided by Lydekker’s Line. Modern evidence has actually reinforced Wallace’s original ideas, showing once again that these Lines represent significant biogeographic barriers that block gene flow in most animals.

Map showing populations sampled for our new paper. We found evidence that the Olive-backed Sunbird is actually composed of separate species in the Philippines (“Garden Sunbird”). the Sunda Shelf (“Ornate Sunbird”), the Wakatobi Islands (“Wakatobi Sunbird”), and the islands from Sulawesi to the Sahul Shelf (“Sahul Sunbird”). We also found that Black Sunbirds in New Guinea are strongly genetically divergent from those in Sulawesi. These findings reinforce the importance of biogeographic barriers like Wallace’s Line and Lydekker’s Line to evolution.

Both evolutionary biologists and ecologists are gaining new insights from large datasets on the traits and genomics of species. However, as these datasets are organised by species, they rely on our species lists being accurate in the first place. Data from  “species” like the Olive-backed Sunbird or Black Sunbird might prove misleading, as each of these actually represent multiple species. Meanwhile, a small population like that of the Wakatobi Sunbird may not be included in such a dataset at all if it isn’t recognised as a species.

To quote an 1863 paper by Wallace, the world’s species represent “the individual letters which go to make up one of the volumes of our earth’s history and, as a few lost letters may make a sentence unintelligible, so the extinction of the numerous forms of life which the progress of cultivation invariably entails will necessarily render obscure this invaluable record of the past”. It is therefore an irrevocable loss to the world, to humanity, and to science when a species goes extinct while still unrecognised, “uncared for and unknown”.

Hoga, one of the Wakatobi Islands where sunbirds were sampled for this study. It has been a privilege to see such beautiful places and animals for this research.

While the Wakatobi Islands are biogeographically “remote” due to their small size and the permanent water barriers that surround them, it is worth noting that they are not the stereotypical “desert islands” Western readers may imagine. The islands have been part of important shipping lanes since at least the 14th century, and the people of the Wakatobi are known for their maritime traditions and unique language. As Dr David Kelly, the second author on the recent paper remarked: “The identification of the Wakatobi Sunbird serves to remind us that biodiversity is everywhere. This bird wasn’t found in a remote rainforest, but along the scrubby margins of busy towns and villages. Let us hope the children of the Wakatobi will be able to enjoy these special birds for generations to come.”

A relief from Borobudur Temple on Java, built over 1000 years ago. The birds in the yellow boxes were identified as Olive-backed Sunbirds by Ashari et al. (2021). Their fascinating paper is available at https://li01.tci-thaijo.org/index.php/tnh/article/view/253401

The beauty of the sunbirds has attracted scientists and artists for many years. Elsewhere in Indonesia, Java’s Borobudur (the largest Buddhist temple in the world, constructed in the 8th or 9th century CE) displays carvings of Olive-backed Sunbirds drinking nectar on its walls. The researchers who identified these carvings hypothesise that this symbolises the Buddhist ideal of enlightenment. Over a thousand years later, sunbirds are still enlightening us on the origin of species.

Female (top) and male (bottom) Olive-backed Sunbirds, from Shelley’s monograph, published 1876-1880 and available from the Biodiversity Heritage Library at https://doi.org/10.5962/bhl.title.53516. Our new paper has found that the Olive-backed Sunbird is actually made up of at least four separate species.

To find out more, read our paper in the Zoological Journal of the Linnean Society here: https://doi.org/10.1093/zoolinnean/zlac081

Battle of the sexes – Niche contraction in females but not males in high density island populations

olive-backed sunbird sexual dimorphism and competition in Sulawesi, Indonesia

This blog was first published on #theBOUblog. Check it out at https://www.bou.org.uk/blog-oconnell-olive-backed-sunbird-sexual-dimorphism/

Going right back to the time of Darwin, competition has always been seen as a driver of evolution. When we think of natural selection we often think of hyaenas and lions fighting it out on the plains of Africa, or a pack of wolves hassling a bear over a carcass. Resource competition like this is a daily part of survival for most animals, with a delicate balance of power between competitors which can change in different environments. However competition for resources takes place not only between species (interspecific competition), but within them (intraspecific competition), and often this is where competition is at its most fierce. Sexually dimorphic species present a special case when studying competition in different environments. They are species where the males and females are physically different, often in body size, but also in colouration and other characteristics. Males and females in sexually dimorphic species often occupy different but interlinked ecological niches. If intraspecific competition becomes more intense within a population of a sexually dimorphic species then one sex may suffer more than the other due to being smaller. In our recent paper in Emu – Austral Ornithology we report on one such case with a niche contraction in female Olive-backed Sunbirds (Cinnyris jugularis), but not males, in a highly competitive environment on small islands. This work was carried out on the remote and understudied islands of South-east Sulawesi, Indonesia (Figure 1), where a joint team of scientists from Trinity College Dublin (TCD) and Halu Oleo University (UHO) have been carrying out research into bird evolution since 1999.

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