identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A187E4FF9CFFB38609D894D8C8FAA8.text	03A187E4FF9CFFB38609D894D8C8FAA8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cinnyris Cuvier 1816	<div><p>CINNYRIS TAXONOMY</p><p>Our results indicate that the olive-backed sunbird represents a superspecies and should be split into at least four species (Supporting Information, Table S10). We have found support for the three-way split suggested by Eaton et al. (2021) while also providing the first genetic, acoustic and morphological evidence that the ‘Wakatobi sunbird’ warrants recognition as a full species. The Wakatobi population exhibits a strikingly non-linear arrangement of population structure, with a range enveloped within that of the more widespread species, and had previously been suggested as a ‘limbo split’ (Rheindt, 2021) based on plumage differences alone (Eaton et al., 2021). Our study has supported these differences with matching patterns in mtDNA and integrative species delimitation or ‘Tobias scoring’ (Tobias et al., 2010). Due to the lower effective population size of mtDNA, along with other factors, differences in mtDNA should be integrated with other forms of evidence in this way (Rubinoff &amp; Holland, 2005). In the light of this new integrative evidence, we recommend that the ‘Wakatobi sunbird  Cinnyris infrenatus ’, originally named by Hartert (1903), be reinstated as a separate species. In addition to its genetic divergence (Fig. 3), the ‘Wakatobi sunbird’ has shorter wings, a shorter bill and longer tarsi than the ‘Sahul sunbird’ (Supporting Information, Fig. S9), as well as exhibiting slower and higher pitched calls over a smaller bandwidth (Supporting Information, Fig. S11). This study is one of several to have remarked upon the distinctness of the Wakatobi avifauna (Kelly et al., 2014; O’Connell et al., 2019a, c), and so we reiterate the recommendation of O’Connell et al. (2020) that the Wakatobi Islands should be protected as an Endemic Bird Area (Stattersfield et al., 1998).</p><p>In addition to the distinctive nature of the Wakatobi lineage, our work supports the splits suggested by Eaton et al. (2021) and one possible additional split in the Philippines (Fig. 5). Under the Eaton et al. (2021) treatment, populations from Sulawesi to the Sahul Shelf and the Solomon Islands were recognized as a species-level taxon, the ‘Sahul sunbird  Cinnyris clementiae ’, the Sunda Shelf populations become ‘ornate sunbird  Cinnyris ornatus ’ and the Philippine birds retain the  Cinnyris jugularis name and take ‘garden sunbird’ as a common name. This is supported by deep genetic divergences (all greater than 5%) between these three putative species, with the Sahul Shelf represented by our new Sulawesi, Australian and PNG sequences along with Solomon Islands birds from Smith &amp; Filardi (2007), the Sunda Shelf by a sequence from Borneo (Boyce et al., 2019) and the Philippines by a number of previously published partial sequences (Supporting Information, Table S3). Although this split has been suggested previously based on limited sampling and incomplete sequences, our more comprehensive sampling of full sequences, taken from both ends of the new ‘Sahul sunbird’ species range, offers stronger support for the division. Our study has also suggested another potential split in this species complex (Fig. 5), outside the geographic range covered in detail by Eaton et al. (2021). ABGD considered the aurora sequences from Busuanga in the western Philippines to represent a distinct species, with a mean genetic distance of 4.5%, from the greater Philippine archipelago (subspecies obscurior and  jugularis). Ornithologists (Rand, 1951; Billerman et al., 2022) have grouped the aurora subspecies separately from these other Philippine subspecies due to its orange breast plumage. The western chain of islands on which the aurora subspecies occurs is geologically and biogeographically distinct from the greater Philippine archipelago (Diamond &amp; Gilpin, 1983). The aurora subspecies displayed a similar level of genetic divergence to that of the more thoroughly sampled ‘Wakatobi sunbird’, but was represented in our analysis by two partial ND2 sequences from Campbell (2013), and so we recommend further sampling of this population.</p><p>The lack of divergence between the Menui population and the wider south-east Sulawesi population in the ‘Sahul sunbird’ confirms that the Menui population belongs to the plateni subspecies. On the Sahul Shelf itself, our ‘Sahul sunbirds’ exhibit a uniform population across PNG and Australia. This is in keeping with the current assignment of these populations to one subspecies,  Cinnyris jugularis frenatus .</p></div>	https://treatment.plazi.org/id/03A187E4FF9CFFB38609D894D8C8FAA8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ó Marcaigh, Fionn;Kelly, David J;O’Connell, Darren P;Analuddin, Kangkuso;Karya, Adi;McCloughan, Jennifer;Tolan, Ellen;Lawless, Naomi;Marples, Nicola M	Ó Marcaigh, Fionn, Kelly, David J, O’Connell, Darren P, Analuddin, Kangkuso, Karya, Adi, McCloughan, Jennifer, Tolan, Ellen, Lawless, Naomi, Marples, Nicola M (2023): Small islands and large biogeographic barriers have driven contrasting speciation patterns in Indo-Pacific sunbirds (Aves: Nectariniidae). Zoological Journal of the Linnean Society 198 (1): 72-92, DOI: 10.1093/zoolinnean/zlac081, URL: https://academic.oup.com/zoolinnean/article/198/1/72/6759114
03A187E4FF9CFFB184BFDC9DDD1AF9E3.text	03A187E4FF9CFFB184BFDC9DDD1AF9E3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Leptocoma Cabanis 1850	<div><p>LEPTOCOMA TAXONOMY</p><p>The movement of species between the genera  Cinnyris,  Leptocoma and  Nectarinia has previously caused confusion for taxonomists working with the black sunbird (LeCroy, 2010). Our analyses suggest that the  Leptocoma genus is not monophyletic (Fig. 2). However, the genus-level branches in our tree had lower support than the species- and subspecies-level branches due to our use of mtDNA. We strongly recommend further sequencing of all species within  Leptocoma to clarify the genus taxonomy.</p><p>Previous research based on plumage differences has not proposed any splits or ‘limbo splits’ within the black sunbird (Eaton et al., 2021). Nonetheless, we found that the black sunbird exhibited genetic divergence consistent with a species-level split between Wallacea and the Sahul Shelf, with a high genetic distance between Sulawesi and PNG (9.1%). This presents a marked contrast to the ‘Sahul sunbird’, which exhibited minimal divergence between those same populations. Further sequencing of birds from intervening areas (such as the Maluku Islands) would help to clarify the boundaries between these potential species. As several intervening populations are absent from our analysis, we cannot rule out the possibility of clinal variation between the Sulawesi and PNG populations (Brumfield, 2005; Cros &amp; Rheindt, 2017). However, the genetic divergence we found was so strong (mean p-distance of 9.1%, higher than any observed in the olive-backed sunbird complex) that it seems more likely the two populations represent two species. The patterns we have observed suggest a division along Lydekker’s Line, and so we predict that sampling of the intervening populations would show a Wallacean species occurring from Sulawesi to the Maluku Islands and a separate species on New Guinea. This would involve the elevation of one of the Wallacean subspecies names to species level, as Lesson &amp; Garnot (1828) named the black sunbird initially for a specimen from Manokwari (Doréry) on the New Guinea mainland. The discovery of cryptic species in the black sunbird highlights the importance of comprehensive sampling of species across their range, even in the absence of obvious plumage differences.</p><p>The structure within the black sunbird contrasted with the patterns shown by the olive-backed sunbird and ‘Sahul sunbird’ in several ways. The black sunbird exhibited a split between Sulawesi and PNG, where the ‘Sahul sunbird’ had a continuous population. The black sunbird also exhibited structure within both of these regions. Most taxonomic treatments (Cheke et al., 2001; Billerman et al., 2022) split the black sunbird into distinct subspecies in PNG’s Western Province (L. a.  aspasia), Northern Province (L. a. vicina Mayr 1936) and the Bismarck Islands (L. a. corinna). However, Gill et al. (2022) merged L. a. vicina into L. a.  aspasia, following Rand (1967). Our genetic work supports L. a.  aspasia and L. a. vicina as distinct subspecies, whereas L. a. corinna appears to warrant species status based on ABGD. However, as this was based on a single sequence, we recommend that the Bismarck populations be examined further before a judgement is made. The Menui black sunbird population was genetically distinct from Sulawesi and the land-bridge islands, with a mean genetic distance (1.72%), a distance greater than that between the two subspecies in PNG (1.48%). This population was also found to be distinct in wing length, bill length, weight and tarsus length (MANOVA, P &lt;0.001 in females). These findings suggest the Menui population of black sunbird may warrant recognition as an endemic subspecies.</p><p>IMPLICATIONS FOR BIOGEOGRAPHY AND EVOLUTION</p><p>The division between the ‘Sahul sunbird’ in Sulawesi and the ‘ornate sunbird’ in neighbouring Borneo (Eaton et al., 2021) reflects one of the most iconic patterns in biogeography. Between these islands runs the boundary between the shallow Sunda Shelf and the deeper waters of Wallacea, which corresponds to the original and best-known version of ‘Wallace’s Line’ (Wallace, 1863; Fig. 1). However, Wallace’s Line, Wallacea and related biogeographic ideas have been treated in an entangled way in the literature, the treatment of Sulawesi and the Philippines being particularly contentious (Ali &amp; Heaney, 2021). The three-way split of the olive-backed sunbird (Eaton et al., 2021), which our work supports, reinforces the more widely used versions of Wallace’s Line (west of Sulawesi, south-east of the Philippines) and Wallacea (excluding the Philippines), as there are separate species in Wallacea (‘Sahul sunbird’), the Philippines (‘garden sunbird’) and the Sunda Shelf (‘ornate sunbird’).</p><p>Unlike Wallace’s Line, a similar barrier in the east has not prevented gene flow in this species, as the ‘Sahul sunbird’ appears to cross Lydekker’s Line with only shallow divergence between populations on either side. It is remarkable that the ‘Sahul sunbird’ appears to maintain a range all the way from Sulawesi to Australia, while the ‘Wakatobi sunbird’ seems to have diverged in a small archipelago nested in this range. Buton is only 27 km from the Wakatobi Islands, while the stretch of islands between Sulawesi and New Guinea contains deep-water barriers up to 100 km wide (this largest barrier is that between Misool and Seram). This striking pattern appears to conform with the ‘taxon cycle’ (Wilson, 1959, 1961; Ricklefs &amp; Bermingham, 2002), whereby taxa become less dispersive as they age. In this framework, the ‘Sahul sunbird’ would occupy an early stage in the taxon cycle, leading to a dispersive lifestyle and a relatively recent colonization of the Sahul Shelf (or perhaps a back-colonization of Sulawesi) across Lydekker’s Line. This would be in keeping with previous hypotheses that the olive-backed sunbird colonized the Sahul Shelf from Wallacea quite recently (Schodde, 1977). The ‘Wakatobi sunbird’ would occupy a later stage of the taxon cycle, maintaining a sedentary lifestyle in a small range. The ‘Wakatobi sunbird’ may thus represent a biological species, at it would have maintained its genetic and phenotypic divergence during this recent colonization wave. In other bird families that are widespread in this region, populations on small oceanic islands have been subject to selection for reduced dispersal ability, leading to a pattern of widespread continental species and isolated island relatives (Pedersen et al., 2018; Le Pepke et al., 2019). We have found evidence that this might also be the case in sunbirds, as the ‘Wakatobi sunbird’ exhibited a significantly lower dispersal index than the ‘Sahul sunbird’ populations of Sulawesi, its land-bridge islands and Menui. Since the earliest days of biogeography (Wallace, 1880), small, isolated ‘oceanic’ islands like the Wakatobi Islands have been known to harbour disproportionate numbers of endemic species. Birds on smaller islands are subject to different evolutionary processes than those on continents, exhibiting lower effective population sizes and losing fewer deleterious mutations to natural selection (Leroy et al., 2021).</p><p>This study also demonstrates the somewhat stochastic nature of island colonization, as we have found that the sunbirds of Runduma colonized this island (located over 50 km east of the main Wakatobi Archipelago) from an entirely different direction than the white-eyes of the island. These are among the only two small passerines on Runduma that are not island specialists or human commensals. The ‘Wakatobi sunbird’ arrived on this tiny island from the Wakatobi Islands: CJ_W03 was the only haplotype found on Runduma, and is shared with Hoga, Tomia, Binongko and Lintea (Fig. 3). Runduma’s white-eye population, on the other hand, is most closely related to that of mainland south-east Sulawesi (O’Connell et al., 2019c). Despite their evolutionary importance, many small islands in the Indo-Pacific have received little ornithological attention until recently (Monkhouse et al., 2018; O’Connell et al., 2020; Sin et al., 2021).</p><p>The black sunbird was already known to be subject to Wallace’s Line, as Sulawesi represents the western extent of its range. Cheke et al. (2001) hypothesized that the black sunbird might be part of a superspecies, with its sister species on the other side of Wallace’s Line. Unlike the ‘Sahul sunbird’, our work indicates that  Leptocoma sunbirds have speciated across Lydekker’s Line, based on the strength of divergence between populations in Sulawesi and PNG. Where the ‘Sahul sunbird’ exhibits a similar population throughout Australia and PNG, the black sunbird exhibits noticeable structure on this scale. That the Bismarck Islands hold an endemic taxon illustrates once again the importance of small and isolated islands in generating biodiversity.</p></div>	https://treatment.plazi.org/id/03A187E4FF9CFFB184BFDC9DDD1AF9E3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ó Marcaigh, Fionn;Kelly, David J;O’Connell, Darren P;Analuddin, Kangkuso;Karya, Adi;McCloughan, Jennifer;Tolan, Ellen;Lawless, Naomi;Marples, Nicola M	Ó Marcaigh, Fionn, Kelly, David J, O’Connell, Darren P, Analuddin, Kangkuso, Karya, Adi, McCloughan, Jennifer, Tolan, Ellen, Lawless, Naomi, Marples, Nicola M (2023): Small islands and large biogeographic barriers have driven contrasting speciation patterns in Indo-Pacific sunbirds (Aves: Nectariniidae). Zoological Journal of the Linnean Society 198 (1): 72-92, DOI: 10.1093/zoolinnean/zlac081, URL: https://academic.oup.com/zoolinnean/article/198/1/72/6759114
