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

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).

Continue reading “Two new bird species from the unique and understudied Sulawesi region”

The Wakatobi Flowerpecker: the reclassification of a bird species and why it matters

Wakatobi Flowerpecker - Male

I posted previously about my PhD research studying bird populations from the tropical and biodiversity-rich region of Sulawesi, Indonesia. I am happy to announce that the first paper as part of this research has just been published in the open access journal PLOS ONE. To read the full paper for free, click here. This work is a collaborative effort from staff in the Department of Zoology in Trinity College Dublin and Haluoleo University in Sulawesi. Here, I’d like to discuss the wider importance of the findings of this study.

My current research focuses on bird populations from peninsular South-east Sulawesi and the nearby Wakatobi Islands. The main focus of this paper was to reassess the taxonomic status of a population of birds from the Wakatobi Islands (i.e. whether these birds represent a species or subspecies). The birds in question belong to the flowerpecker family (Dicaeidae); a group of small and colourful, arboreal passerines found from Southeast Asia to Australia. The Wakatobi birds were originally described as a separate species (Dicaeum kuehni) from those on mainland Sulawesi by the renowned avian taxonomist Ernst J. Hartert. However, for reasons that remain unclear in the literature, the Wakatobi birds were later reclassified as a subspecies of the Grey-sided Flowerpecker (Dicaeum celebicum) from mainland Sulawesi. Therefore we decided the Wakatobi populations were deserving of reassessment. From comparisons of plumage and morphology (that is, the measurement of various features such as a bird’s wing and bill), as well as estimates of genetic divergence and phylogenetic relationships between Wakatobi and Sulawesi populations, our results suggest the Wakatobi birds deserve to be recognised as a distinct species. We have therefore recommended the Wakatobi populations be reclassified as Dicaeum kuehni, a species found only on the Wakatobi archipelago and put forward the common name ‘Wakatobi Flowerpecker’.  For more detailed methods and results check out the paper.

“So what?”, you might say. Well, despite centuries of work from naturalists aiming to estimate the number of different species that exist or have existed on Earth (be they animal, plant, fungus, bacteria, etc) and further understand their evolutionary relationships, we still have a lot to learn! Therefore, this research adds another tiny piece to this enormous and incomplete jigsaw. Through a greater understanding of life on Earth we can attempt to answer some of the great philosophical questions, such as ‘Where and how did life start?’; ‘How and why do new species appear?’;  ‘Why has life evolved to become as it is today?’; and ‘How have we, as humans, come to be?’. Anyway, let’s be honest, who doesn’t enjoy learning of a recently discovered species or simply one they haven’t heard of before (be they as cute as the recently discovered olinguito or as frighteningly ugly as the goblin shark)? But the endeavour to discover species and classify and quantify the diversity on life on Earth brings us much more than entertainment and endless fascination, it also has very practical applications. Data on the distribution and conservation status of species are one of the major sources of information used to inform conservation policy. Therefore, as we are in the midst of an extinction crisis, it is vital that these data are accurate.

In order to maximise our understanding biodiversity, particularly in the remote and poorly known Sulawesi region of Indonesia, we require multi-disciplinary research. For example, take a look at Figure 1 below. On the left are a male (above) and a female (below) Grey-sided Flowerpecker from mainland Sulawesi. On the right are a male (above) and a female (below) Wakatobi Flowerpecker. They look very similar, right? This is true. However there are subtle but consistent differences in plumage between the species (again, see the paper for more info on this). Without the collection of detailed morphological data and the generation of genetic sequences, we may have incorrectly concluded that these make up just one species, when in fact they are morphologically distinct, reproductively isolated and genetically very different. This demonstrates the need for modern research, not just in Sulawesi, but globally, to employ integrative research, combining traditional comparisons of colour, size and shape with modern genetic and phylogenetic analyses.

Figure 1. Plumage comparisons-p18pjcggcs1dgo1ulm1sor9s214bc
Figure 1. A comparison of plumage characteristics between male (top row) and female (bottom row) Grey-sided Flowerpeckers (left) and Wakatobi Flowerpeckers (right).

Despite the knowledge that the Sulawesi region is home to a large number of remarkable birds that are found nowhere else in the world, it has remained relatively poorly studied. Furthermore, there has been a lack of integrative ornithological research in the area and very little genetic sampling. Therefore, it is likely that avian species richness for the Sulawesi region is underestimated and that numerous bird species are awaiting description. On top of this, Sulawesi’s biodiversity is facing major threats from a rapidly expanding human population and mass habitat destruction, among other things. Unless we can encourage more multi-disciplinary research within the region, we will likely fail to recognise evolutionarily distinct lineages and run the risk of losing them forever.

Our current findings inspire many further questions. For example, why have the flowerpeckers on the Wakatobi islands become so different to their close relatives on mainland Sulawesi? In other words, what are the evolutionary pressures that have driven the divergence of the Wakatobi Flowerpeckers? By investigating these questions, we hope to learn more about the evolutionary processes of speciation and adaptation to living on islands. As the Wakatobi Flowerpecker is found only on the Wakatobi Islands, the protection status afforded to the islands may require reassessment. Furthermore, considering one unique bird species has evolved on the Wakatobi, could there be more? Watch this space.

Author and Images:  Seán Kelly, kellys17[at]tcd.ie, @seankelly999

Sulawesi Bird Expedition 2013

Beach

Ah the summer, how I miss it! In mid-June I departed (on the horrendously long journey) to the beautifully sunny, tropical islands off the south-eastern coast of Sulawesi, Indonesia. No, I wasn’t on holidays; I am fortunate enough to call this part of the world my study site. During the six week visit, I aimed to gain further behavioural and ecological data on a number of bird species as part of my current PhD project – for more specifics on that see here. I am even more fortunate to be able to carry out this research with the financial and logistical support of Operation Wallacea, an internationally renowned conservation charity that works with researchers from all over the globe, from a variety of different disciplines. As part of this support, I work with students and volunteers in the field, helping them to design effective dissertation projects and field methods. This year I was joined by five students (as opposed to two last year) from a variety of universities in Ireland and the UK.

From the first day it was back to the usual diet (mostly consisting of rice) and routine: up at 4.30am for breakfast and out surveying by 6am. These surveys consisted of walking 1km transects through scrub, farmland and/or forest edge collecting data on my target species’ diets, competitors (via agonistic interactions), social habitats, courtship and breeding, as well as their foraging and flocking behaviours. In the evening we would establish new transects and then get stuck into data entry at night. This routine makes for days that are long and tiring but hugely rewarding. Watching birds so closely allows you to gain intimate insight into their lives and observe some fantastic interactions, such as family groups of Lemon-bellied White-eyes preening each other and pairs reinforcing bonds with gifts of food. You also see how tirelessly and (sometimes) viciously males will fight off other males in order to retain their mates and, therefore, mating privileges, as we saw in the beautifully adorned Olive-backed Sunbird. Spending so much time in the field, you come across a great variety of other wildlife including troops of macaques, the strange bear cuscus, giant monitor lizards, pythons, huge fruit bats and hairy and multicoloured caterpillars that you never touch, to pick out but a few.

An adult Lemon-bellied White-eye returning with food for its chicks
An adult Lemon-bellied White-eye returning with food for its chicks

A beautiful fruit bat relaxing in a banana tree
A beautiful fruit bat relaxing in a banana tree

I’m delighted to say that data collection went exceedingly well for the students and myself – that is, when the weather was on our side (we had a week of non-stop rain while Ireland and the UK were experiencing a heat-wave; typical!). We surveyed five islands in total and got some superb behavioural data on each of our five target species. While managing a large group like this was difficult and tiring at times, it was a great experience and the students were a great bunch really. In the company of the assistants and students on the project, as well as the many other members of staff, students and volunteers from other projects, with their combined wealth of experience and knowledge, it was fantastic to share ideas, brainstorm and discuss current/potential future projects.

The biggest highlight for me was catching up with the elusive ‘Wangi-wangi’ White-eye, a bird we know very little about. It was touch and go for a while, and I was getting quite worried to be honest, but eventually we got excellent data on their flocking and feeding behaviour and who they compete with, directly and indirectly. Between us, the group racked up a number of new bird records for the islands and saw some spectacular species such as the Great-billed Kingfisher, Rainbow Bee-eater, Yellow-eyed Imperial Pigeon, Great-billed Parrot, Yellow-billed Malkoha and Red-knobbed Hornbill. Phwoar!

The stunning Yellow-billed Malkoha, a Sulawesi endemic
The stunning Yellow-billed Malkoha, a Sulawesi endemic 

Sadly, this summer was the last of my field trips to Indonesia as part of my PhD project. I enjoyed it immensely and, for certain, I will be back. I am grateful to Operation Wallacea for allowing me to be involved in such a programme and I hope that they will continue to expand into more areas of this highly unique and understudied part of the world that is full of discoveries yet to be made.

Author and Photo credits:

Seán Kelly: kellys17[at]tcd.ie, @seankelly999

Sulawesi field report

A Lemon-bellied White-eye Zosterops chloris with nesting material in its bill, Tomia Island

July and August of this year saw members of the Behavioural and Evolutionary Ecology research group embark on another field season studying the birds of tropical south-east Sulawesi, Indonesia. Principal investigators Dr. Nicola Marples and Dr. Dave Kelly were joined this year by PhD student Seán Kelly, as well as a number of undergraduates from the university. This year’s expedition, carried out in collaboration with Operation Wallacea, consisted of two teams: the mist netting team, led by Dr. Marples and Dr. Kelly, and the behavioural team led by Seán Kelly.

The netting team trapped birds using mist nets at various locations on Buton island, mainland south-east Sulawesi and Wangi-wangi island (of the Wakatobi archipelago). While small passerines such as white-eyes, sunbirds and flowerpeckers were the target species, individuals from a total of 35 species were caught. The season proved to be a great success with over 300 birds trapped and processed. Data on plumage, morphology, age, sex and breeding condition were collected from each bird, which was colour-ringed and released unharmed. A small number of body feathers were also plucked from each bird for later genetic and stable isotope analyses.

The behavioural team spent the season on various islands of the Wakatobi archipelago collecting detailed behavioural ecology data on the white-eye, sunbird and flowerpecker species present. This included information on their diets, competitors, preferred habitats, social habits, courtship and breeding, as well as their foraging and flocking behaviours. Data collection took place in the early morning and evening, walking 1 km transects through scrub, farmland or forest edge habitats. This resulted in some fantastic insights into the behaviour and ecology of these poorly studied species.

From analysis of the plumage, morphometric and genetic data we have found a number of significant differences between bird populations on the Wakatobi archipelago and mainland Sulawesi, as well as between populations within the Wakatobi. It is hoped that the behavioural data gathered this season will help us to understand the selective pressures driving this divergence, giving us further insight into the evolution of this region’s fascinating avifauna.

Author

Seán Kelly: kellys17[at]tcd.ie

Photo credit
Seán Kelly

Is island life easier?

Lemon-bellied White-eye (Zosterops chloris)

For over 10 years we have been making regular visits to islands in the Sulawesi region of Indonesia. We trap birds on these islands, collecting morphometric data. Each bird we trap is measured, marked with a plastic ring and released. As our dataset grows we gain more insight into the lives of the birds on these islands.

In 2007 and 2010 we visited the island of Kaledupa in the Wakatobi archipelago. In 2010 we made a point of revisiting all of the sites we had trapped at in 2007. This gave us an opportunity to look for the birds we had originally caught in 2007. Continue reading “Is island life easier?”