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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The 2020 EcoEvo Hall of Fame

At the start of each year we ask the EcoEvo contributors to share their favourite scientific publications from the past year and why they found them interesting, inspiring, or otherwise worthy of inclusion in the Hall of Fame. Keeping with tradition, here are the EcoEvo Hall of Fame entries for 2020! And if you enjoy reading about our favourite papers from 2020, remember you can also check out our favourites from 2017, 2018 and 2019, too!

Chosen by Andrew Neill

Read the full People and Nature paper here.

I really enjoyed this paper because it tackles a really difficult topic at the intersection of poverty, human rights, development, conservation, and sustainability. It is important to remember that conservation will never meet its objectives without considering how people depend on nature for their needs and livelihoods. The areas of richest biological diversity (and therefore conservation potential) are usually in developing countries with communities experiencing poverty. This paper collects responses from conservation practitioners to examine their viewpoints on poverty in the context of their work. 

F I G U R E 3. Comparison of discourses on five key dimensions of difference. Discourses are compared on a simple ordinal scale, and accordingly should only be interpreted in relative positions to one another (for instance, D3 is more ecocentric than D1).
© 2020 The Authors. People and Nature published by John Wiley & Sons Ltd on behalf of British Ecological Society. The article is distributed under the terms of the CC-BY 4 license.

They found some areas of agreement such as the poorest people should not be expected to shoulder the costs of preserving a global public good (the conservation of biodiversity). However, they also identify differences between responses: Is the focus placed on meeting the needs of people or more closely aligned with the “do no harm” principle? Is poverty a driver of nature’s decline, or is it the over-consumption that drives environmental degradation? This paper was a great opportunity to question my own views on these very complex ideas and to appreciate the wide diversity of thought going on across the world of conservation. 

Fisher, J.A., Dhungana, H., Duffy, J., He, J., Inturias, M., Lehmann, I., Martin, A., Mwayafu, D.M., Rodríguez, I. and Schneider, H. (2020). Conservationists’ perspectives on poverty: An empirical study. People and Nature2 (3), pp.678-692.


Chosen by Fionn Ó Marcaigh

Read the full Nature Communications paper here.

This paper is based on a truly colossal undertaking: to collect their data on dispersal ability, Sheard et al. measured the wings of 10,338 bird species, i.e. 99% of all bird species on Earth. They used the Hand-Wing Index, a measure that correlates with aspect ratio and basically tells you how long and pointed the bird’s wing is. The higher this number (i.e. the pointier the wing), the better the bird will be at dispersing and flying long distances.

a Diagram showing linear measurements used to calculate HWI taken on a standard museum study skin (secondary feathers shown in pale grey; primary feathers in dark grey). Wing length (Lw) is the distance from carpal joint to the tip of the longest primary feather; secondary length (S1) is the distance from carpal joint to the tip of the first secondary feather; Kipp’s distance (DK) is the difference between Lw and S1b Open wing of a passerine bird showing how Lw and S1 are related to the wing’s span and width, and hence to its aspect ratio. c Because it is correlated with the aspect ratio, HWI is in theory positively associated with flight efficiency and key aspects of dispersal ability, including dispersal distance and gap-crossing ability.
© The Author(s) 2020. This article is distributed under the terms of the CC BY 4 license.

This is important for evolution, as the more birds that are able to fly between distant populations the more gene flow there will be and the less likely the populations are to diverge. Sheard et al. found important links between dispersal ability and geography and ecology, as tropical and territorial birds, had lower Hand-Wing Indices and migratory species had higher ones. It’s fascinating to see how these traits affect the ability of a species to move around, which in turn dictates where that species will be found in the world. The authors have made this incredible dataset freely available and it is sure to inform new insights into bird ecology and evolution for years to come.

Sheard C., Neate-Clegg M. H. C., Alioravainen N., Jones S. E. I., Vincent C., MacGregor H. E. A., Bregman T. P., Claramunt  S. & Tobias J. A. (2020) Ecological drivers of global gradients in avian dispersal inferred from wing morphology. Nature Communications, 11 (2463).


Chosen by Sam Ross

Read the full Science paper here.

The COVID-19 pandemic has been extremely challenging for many, so it was great to see some excellent science coming from the ‘natural experiment’ offered by COVID-19 movement restrictions. The authors show that during the COVID-19 restrictions anthropogenic noise (from vehicles etc.) in the San Francisco Bay Area reached a 70-year low, characteristic of the mid-1950s. They use a long-term dataset of White-Crowned Sparrow recordings to show that during the COVID-19 lockdown, when human noise pollution was minimal, Sparrows exploited the emptied acoustic space (usually occupied by human-related noise) by producing higher-performance songs at lower amplitudes, to maximise song distance. The authors highlight the rapidity with which behavioural traits (song characteristics) adapted to changes in human activity, suggesting incredible plasticity and potential resilience to pervasive anthropogenic pressures like noise pollution. To me, this study is a perfect example of nature’s resilience, and also on finding opportunity from tragedy (research made possible by a global pandemic).

Derryberry E.P., Phillips J.N., Derryberry G.E., Blum M.J., Luther D. (2020). Singing in a silent spring: Birds respond to a half-century soundscape reversion during the COVID-19 shutdownScience, 370, 575-579.


Chosen by Jenny Bortoluzzi

Read the full Marine Policy paper here.

This paper looked at the human behavioural responses to a blanket ban on thresher shark fisheries in Sri Lanka and fisher’s perceptions of different aspects of the ban. A blanket ban means a complete prohibition on exploitation of a species, and Thresher sharks are considered to be the most vulnerable species of pelagic sharks. A blanket ban might therefore seem like a straightforward and easy conservation measure to protect them. But this study looked at the human impact behind such a drastic policy decision. A ban like this has consequences for the livelihoods of fishers – particularly smaller fishermen who rely highly on thresher shark landings to provide for their families. The study clearly shows the disparity in the impact this conservation policy has had between fishers who rely on these catches to survive and those for whom they are not the primary catch.

The biggest message I took from this paper is how important it is that human lives are taken into account when making conservation decisions; and more importantly that scientists and policymakers need to involve communities early on in the process, communicate better and work together, not against each other if we want conservation to be effective – and supported. This is a message I think more scientists need to hear and integrate into their work and one I hope to take forward in my future career.

Collins C., Letessier T. B., Broderick A., Wijesundara I., Nuno A. (2020). Using perceptions to examine human responses to blanket bans: The case of the thresher shark landing-ban in Sri Lanka. Marine Policy, 121 (104198).

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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The 2019 EcoEvo Hall of Fame

Every year, contributors to the blog look back on their favourite papers of the previous year and tell us what it was about these publications that stuck in their mind so much. With a range of different topics and reasons, it’s always great to see what each of us thinks makes for a great paper! Find out what we elected as our favourite papers in 2018 and 2017, and read on for this year’s entries:


Perrot-Minnot, M. J., Guyonnet, E., Bollache, L., & Lagrue, C. (2019). Differential patterns of definitive host use by two fish acanthocephalans occurring in sympatry: Pomphorhynchus laevis and Pomphorhynchus tereticollis. International Journal for Parasitology: Parasites and Wildlife8, 135-144.

Chosen by Paula Tierney

Read the full International Journal for Parasitology: Parasites and Wildlife paper here

https://ars.els-cdn.com/content/image/1-s2.0-S2213224418300671-fx1_lrg.jpg
Graphical abstract from paper

Sometimes a paper doesn’t have to make huge waves in the broader ecology world to be a great paper and sometimes a paper comes along at just the right time to answer the questions you need answering. Perrot-Minnot et al. 2019 did both for me this year. Since the taxonomic revision of the acanthocephalan parasite Pomphorhychus laevis by Špakulová et al in 2011 resurrected the closely-related species Pomphorhynchus tereticollis, the systematics of the genus in Europe has been something of a taxonomic dumpster fire. It also left one of my PhD chapters with a bit of an identity crisis since,

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Two new bird species from the unique and understudied Sulawesi region

This blog was first published on #theBOUblog. Check it out at https://www.bou.org.uk/blog-oconnell-two-new-white-eye-species-sulawesi/

The Wallacea region has always been known to be home to many unique species, with birds of paradise, giant reptiles and marsupial versions of sloths found among its many islands! The region takes its name from Alfred Russel Wallace, who along with Darwin, developed the theory of evolution from his studies of the species of Wallacea. When I first set my heart on a career as a Zoologist (a decision made with absolute certainty at age 12!) I dreamed of following in the footsteps of these great naturalists. So it is of no surprise that when I finally got around to starting my PhD many years later, I chose to study speciation (the formation of new species during the course of evolution) in the birds of Wallacea, with the hope the region still held mysteries to uncover. Our research focused on South-east Sulawesi, Indonesia. Sulawesi is a weird and wonderful part of the world, and island hopping through that region has provided me with a lifetime of unforgettable memories. It also allowed me to fulfill my dream, as in our recent paper in the Zoological Journal of the Linnean Society, we describe two new bird species from the Wakatobi Islands, an island chain off South-east Sulawesi (Figure 1).

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The 2018 EcoEvo Hall of Fame

As we launch into another year of research, we thought it would be interesting to look back on some of the work that came out in 2018, so we asked the School of Natural Sciences what papers they would like to induct into our second annual Hall of Fame. Read on for the papers we thought were fascinating, notable, or just cool. When you’re finished, you can check out 2017’s Hall of Fame here.

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Kingfisher Evolution in the Wallacea Region

Studying diversification in the Todiramphus kingfishers of Sulawesi often brought to mind the adage, ‘what is rare is beautiful’. Though I certainly also learned that rare beauties can be incredibly frustrating! While I could be guaranteed to catch my other main study taxa, Zosterops white-eyes, by the dozen in the right habitat, Collared Kingfishers (Todiramphus chloris) and Sacred Kingfishers (Todiramphus sanctus) were much more elusive. This made them maddening study species, but ensured that every time I got to grips with one was a special moment. Each bird caught was measured and had a few flank feathers taken before release. Even the most disappointing site could be made worthwhile by catching a kingfisher. In particular, I’ll never forget a particularly fetid swamp in the backwoods of Sulawesi, as I’m pretty sure all the mosquitos in the world lived there. It yielded our worst ever bird catches, but two crucial Collared Kingfishers! These moments of elation have stuck with me, and have ensured that the Todiramphus kingfishers are the study species I’m most fond of.

Darren O’Connell and Adi Karya examine birds, while being examined by passers-by. Photo by Suliman La Ode.

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