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

Tramps in Transition: Wallacea’s monarch flycatchers and their evolutionary natural experiment

Pale-blue Monarch on the left and Island Monarch on the right

A warm welcome back to all our readers! The new year is now well and truly upon us and we hope you’ve all had a safe and energised return to work. This blog is written by Fionn Ó Marcaigh, summarising his new paper. Congratulations Fionn and we hope our readers enjoy learning about your research as much as we have! So without further ado…

Science is about making observations from the natural world, drawing up hypotheses to explain the patterns you’ve observed, and then testing these hypotheses by experimentation. We tend to imagine scientists in white coats doing experiments in the lab, but our understanding of evolution also owes a lot to work done in “natural laboratories” like islands and other isolated habitats, where evolution has taken place under different conditions. Our new paper, just published Open Access by the International Biogeography Society in their journal Frontiers of Biogeography, has used an important natural laboratory in Southeast Asia to test a classic hypothesis based on a bird called the Island Monarch (Monarcha cinerascens). We’ve made observations that contradict parts of the hypothesis and discovered a possible new species in the process!

Our natural laboratory was a collection of islands around a region known as Wallacea in central Indonesia (see map below). Named after Alfred Russel Wallace, this is where he co-discovered evolution by natural selection while travelling around islands of all shapes and sizes, with the waters around them being so wide and deep that most species have trouble crossing them. Some organisms are better at crossing these barriers than others, with the Island Monarch thought to be particularly adept. As its name suggests, the Island Monarch is one of the kings of small islands. It can be found all the way from the islands off Sulawesi in Wallacea, to the farthest reaches of the Melanesian islands east of Papua New Guinea, but is missing from large islands like Sulawesi and New Guinea themselves.

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Evolution in the understorey

Sulawesi babblers from several islands

What is the first image that comes to mind when you think of evolution? Possibly a line of cartoon primates marching, slouching monkeys at one end and naked men with spears at the other. Or a branching tree diagram where each twig represents an organism, maybe with a tentative “I think” scribbled above it. Alternatively, you may have pictured an illustration of related birds from isolated islands, each showing a dramatically different bill shape adapted to a different diet. Darwin’s Galápagos finches represent a foundational influence in terms of where we tend to look for signs of evolution and what we expect these signs to look like. Our new paper, just published Open Access in Zoologischer Anzeiger: A Journal of Comparative Zoology, provides a contrasting image. We looked at the Sulawesi babbler (Pellorneum celebense), a dull brown bird that spends its time hiding in bushes on less isolated islands in Indonesia, looking pretty similar from one island to the next. Nevertheless, we found that several of its populations are quite different from one another in mitochondrial DNA, in morphology, and in song.

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Science and the English Language

Most people who bother with the matter at all would admit that the English language is in a bad way, but it is generally assumed that we cannot by conscious action do anything about it.”

So George Orwell began his 1946 essay Politics and the English Language, which is still relevant today as both a guide and a warning. Reading it now (the whole thing is available online courtesy of the Orwell Foundation), it strikes me that the decline Orwell saw in the English language might be blamed on science as much as politics. Three of his “five specimens” of poor writing come from academia (one of them written by a prominent zoologist), and many of the specific writing habits he criticises are ones I see regularly in modern papers. One of our recent “NERD Club” discussion sessions was based on Orwell’s essay and related topics, as we looked for the conscious actions that might help us to write clearly and accurately.

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Shy birds hiding evolutionary insights: cryptic sexual dimorphism in the Sulawesi Babbler

A Sulawesi Babbler

The most iconic examples of sexual dimorphism are found in birds. Birds are probably many people’s introduction to this concept in the natural world, when brought to the park as a child and shown the difference between male and female mallards. But sexual dimorphism can be much more subtle than that kind of difference in colour (called dichromatism, and seen also in peacocks, pheasants, sparrows, and many more). Our new paper, just published by the Association for Tropical Biology and Conservation in their journal Biotropica, shows that sexual dimorphism can be missed in some birds even when it is important to their ecology and evolution.

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Boating in Ankobohobo, or The Importance of Bird Areas

Many of Earth’s plant and animal species are at imminent risk of extinction. But with the resources necessary to conserve them so sorely limited, where should efforts be focused? The term “ecological triage” has been coined for such decision-making, after the system invented by French army medics to sort patients by the urgency of their need. This borrowing of metaphor from the Napoleonic and World Wars is not accidental: in envisaging the damage done to nature by humanity, you are to think of cannonballs and scorched earth and mustard gas.

“Ecological triage” can focus attention onto the regions that hold a disproportionately large amount of diversity. Such “biodiversity hotspots” have become key to global conservation, from the Caribbean to Wallacea to New Zealand. But one country stands out as among the “hottest” of hotspots: the great island of Madagascar. I’ve written for EcoEvo before about the field season I spent there, which has just resulted in a new paper published in Scopus: Journal of East African Ornithology. The paper presents findings from a series of boat surveys in the threatened and irreplaceable Ankobohobo Wetland, home to some of the world’s rarest birds.

Header: African Darter (Anhinga rufa) in Ankobohobo, by Jamie Neaves.

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The Bird Life of Wawonii and Muna Islands Part II: mining the literature, mining the hills

Indonesia is a constellation of islands, big and small and many-shaped, born out of volcanism or coral build-up or sheared from the ground of continents, each its own world. The complex geography of this region begat an equally complex biogeography, with evolution working enthusiastically to populate it, and the staggering biodiversity that resulted will surely occupy biologists for centuries to come (as long as it lasts that long). Every nook and cranny is filled with fascinating plants and animals, and our research group recently published a paper on the avian contents of a couple of those nooks and crannies. This post follows one by Darren on the fieldwork stories behind that paper, and the expeditions to remote Muna and Wawonii islands.

Muna and Wawonii in their corner of Indonesia. The numbers mark sites surveyed for the recent paper.
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A Mosaic of Birds in Madagascar

This post is based on the paper ‘The avifauna of the forest mosaic habitats of the Mariarano region, Mahajanga II district, north-west Madagascar’, just published open-access in Bothalia: African Biodiversity and Conservation.

The header image by Jamie Grant-Fraser shows White-faced Whistling Ducks (Dendrocygna viduata) responding to the appearance of a Madagascar Harrier-hawk (Polyboroides radiatus).

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A Happy Réunion at the Island Biology Conference 2019

A beautiful sunset on the southern shore of Réunion ISland, after the ISland Biology 2019 conference

Island biologists often work in beautiful and interesting places. It seems only fitting that when they meet up to discuss their work, they do it somewhere like Réunion, a volcanic oceanic island in the Indian Ocean, administered as a French department and the site of the 2019 Island Biology conference. Armed with my Junior Cert level French, I made the journey to discuss my PhD project on the birds of Sulawesi and the work of the TCD Biogeography Working Group.

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