Recent dispersal events among Solomon Island bird species reveal differing potential routes of island colonization

By Jason M. Sardell

Abstract Species assemblages on islands are products of colonization and extinction events, with traditional models of island biogeography emphasizing build-up of biodiversity on small islands via colonizations from continents or other large landmasses. However, recent phylogenetic studies suggest that islands can also act as sources of biodiversity, but few such “upstream” colonizations have been directly observed. In this paper, I report four putative examples of recent range expansions among the avifauna of and its satellite islands in the , a region that has recently been subject to extensive anthropogenic habitat disturbance. They include three separate examples of inter-island dispersal, involving Geoffroyus heteroclitus, Cinnyris jugularis, and Rhipidura rufifrons, which together represent three distinct possible patterns of colonization, and one example of probable downslope altitudinal range expansion in Petroica multicolor. Because each of these species is easily detected when present, and because associated localities were visited by several previous expeditions, these records likely represent recent dispersal events rather than previously-overlooked rare taxa. As such, these observations demonstrate that both large landmasses and small islands can act as sources of island biodiversity, while also providing insights into the potential for habitat alteration to facilitate colonizations and range expansions in island systems.

Author E-mail: [email protected]

Pacific Science, vol. 70, no. 2 December 16, 2015 (Early view) Introduction

The hypothesis that species assemblages on islands are dynamic, with inter-island dispersal playing an important role in determining local community composition, is fundamental to the theory of island biogeography (MacArthur and Wilson 1967, Losos and Ricklefs 2009).

Identifying island colonization events can therefore provide important insight into the processes that influence island species assemblages. Traditional models of island biogeography typically portray colonizations as a one-way flow of individuals from larger to smaller landmasses, with small islands acting as sinks of diversity or as stepping-stones for colonization of landmasses further away (MacArthur and Wilson 1967, Bellemain and Ricklefs 2008). However, recent phylogenetic evidence suggests that small islands can also act as sources of biodiversity for species assemblages on continents or larger islands (Filardi and Moyle 2005, Nicholson et al.

2005, Bellemain and Ricklefs 2008, O'Grady and DeSalle 2008, Hutsemékers et al. 2011,

Andersen et al. 2015, Patiño et al. 2015), but very few such “upstream” colonizations have been directly observed in animal systems. Therefore, observations of island colonization events are needed to confirm the potential for small islands to act as sources of biodiversity.

In this paper, I report new locality records for four bird species in Makira-Ulawa Province in the

Solomon Islands, each providing an example of a different possible colonization pattern responsible for generating local species assemblages in island systems. The Solomon Islands avifauna featured prominently in several of the keystone papers in the field of island biogeography (reviewed in Simberloff and Collins 2010), including MacArthur and Wilson’s

(1963, 1967) original formulation of the theory of dynamic equilibrium, as well as Diamond,

Mayr, and Gilpin’s related research (Diamond et al. 1976, Diamond and Mayr 1976, Gilpin and

Diamond 1976); Greenslade’s (1968) empirical work on taxon cycling theory; and Diamond’s

(1975) research on the effect of resource competition on island assemblages . As such, this

2 system is uniquely well-suited for providing important insight into the dynamics of range expansions in island taxa.

Study Site and Methods

Makira-Ulawa province in the Solomon Islands comprises nine major islands. The largest island,

Makira (formerly known as San Cristobal) is 3,188 km2 with a maximum elevation of 1,040m.

Two groups of satellite islands - Ugi along with the small islet Pio, and the three islands comprising the Three Sisters archipelago (Malaupaina, Malaulalo, and Ali’ite) - lie approximately 8 km and 20 km north of Makira, respectively (Fig. 1). In addition, two satellite islands, Santa Ana (Owa raha) and Santa Catalina (Owa riki) lie approximately 7.5-8 km off the eastern point of Makira (Fig. 1). Finally, Ulawa, the second-largest island in the province, is approximately 65 km north of Makira, although its avifaunal assemblage is more similar to that of the island group approximately 40 km to the west (Cain and Galbraith 1956).

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The earliest records of the avifauna of Makira and Ugi were based on specimens collected by

Victorian-era naturalists in the second half of the nineteenth century (reviewed in Ramsay 1883).

However, the reported provenance of many of these early specimens is dubious. For example, several of the species that Ramsay (1883) lists as being common on Ugi (e.g., Meliarchus sclateri, Myzomela tristrami) are commonly found on Makira but are, at best, rare vagrants to

Ugi (Cain and Galbraith 1956). Later expeditions to this region include visits by A.S. Meek to

Makira in 1908 (Rothschild and Hartert 1908), the Whitney South Seas Expedition to Makira,

Ugi, and Santa Ana for two months in 1927 and one month in 1929 (Mayr 1945), the Crane

Expedition to Ugi for one day in 1929 (Mayr and Camras 1938), the Templeton Crocker expedition to coastal Makira and Ugi in 1933 (Davidson 1934), and the Oxford University

Expedition to Makira (two months) and Ugi (one month) in 1953 (Cain and Galbraith 1956).

3 More recent surveys of the avifauna of Makira and Ugi have been undertaken by J. Diamond in

1974 and 1976 (Diamond 2002), G. Dutson et al. for a total of five weeks in 1990, 1998, 2000, and 2004 (summarised in Dutson 2001, 2011), and the Galathea 3 Expedition project over three weeks in 2006 (Danielsen et al. 2010). In contrast, few recent surveys of the Three Sisters have been undertaken and much of the current knowledge of these islands’ avifauna comes from 13 years of personal observations published by W. French (1957), as well as a 1990 visit by Dutson et. al. (Dutson 2001).

I visited Makira and its satellite islands from 20 June to 18 July 2012, 31 May to 19 July 2013, and 28 July to 10 Sept. 2014 as part of a research project investigating interactions between

Myzomela tristrami and Myzomela cardinalis (Sardell and Uy, in review). While there, I opportunistically observed all other bird species present at my field sites. The location of any noteworthy or unusual sightings were recorded using a Garmin GPSMAP 76CSx global positioning system, with a minimum of ±5m accuracy, and such locations are reported using standard UTM formats for the Southern Hemisphere. All scientific and English names used in this publication are consistent with Gill and Donsker (2014).

Results

I identified four examples of putative recent range expansions of avian species in Makira-Ulawa province, based on observations of multiple individuals outside of their previously known ranges. These observations are described below.

Colonization of satellite island from larger landmass: Song Parrot (Geoffroyus heteroclitus)

The Song Parrot (Geoffroyus heteroclitus) is a medium-sized parrot found primarily in forested habitats throughout the Solomon Islands and Bismarck Archipelago. Although nineteenth-

4 century specimens of G. heteroclitus were labelled as being collected on Ugi (Ramsay 1883), the provenance of these specimens is considered dubious (Cain and Galbraith 1956) and there are no currently accepted records of G. heteroclitus for the Makira satellite islands (Dutson 2011).

I observed a mixed-sex flock of approximately twelve G. heteroclitus in Molehanua village, Ugi

(57L 0798148 8860147) on multiple occasions over two days from 12-13 July 2013. This locality is on the southwest point of Ugi directly across the strait from Makira. However, given the capability of parrots to travel over long distances in search of food, it is unknown whether G. heteroclitus is an established breeder on Ugi or an occasional visitor from the mainland.

Nonetheless, this observation fits the traditional model of populations on large landmasses acting as a source for dispersal to uncolonized islands (MacArthur and Wilson 1967).

Colonization of larger landmass from satellite island: Olive-backed Sunbird (Cinnyris jugularis)

The Olive-backed Sunbird (Cinnyris jugularis) is a wide-ranging nectivorous passerine found in

Asia, Australia, and . It is common, particularly in disturbed habitats, on all major islands in the , with the exception of Makira, Rennell, and Bellona (Dutson

2011). Its absence on Makira is especially striking as it is relatively common on Ugi and the

Three Sisters, particularly near villages, where it co-exists with another nectivorous species,

Myzomela cardinalis (Cain and Galbraith 1956). Relative species distributions and spatial organisation of sunbirds and honeyeaters have often been used to support key ecological theories

(Ripley 1959, Diamond 1970, 1975, Diamond and Marshall 1977, Sanderson et al. 2009,

Carstensen et al. 2011), and it has been hypothesized that C. jugularis may have been unable to undergo range expansion into other Pacific islands due to competitive exclusion by native generalist Myzomela species (Diamond 1970, Diamond and Marshall 1977) .

5 A single male C. jugularis was observed daily between 28 June -3 July 2013 in gardens behind

Kirakira, Makira (57L 0820279 8842639), the capital and largest settlement in Makira-Ulawa province. A male C. jugularis, potentially the same individual, was also observed in the same location on 20 August 2014. In addition, a male C. jugularis was observed in secondary habitats near the Ravo River road crossing (57L 0824874 8839073), 6.25 km southeast of Kirakira on 4

July 2013. These sightings suggest that C. jugularis recently may be established (but relatively rare) in secondary habitat in lowland northern Makira.

C. jugularis represents the third example of a passerine that likely colonized Makira from populations on Ugi and/or Three Sisters during historically recent times, along with Myzomela cardinalis (Diamond 2002) and Aplonis cantoroides (Dutson 2001). The arrival of these species to Makira coincided with pronounced anthropogenic changes to habitats on the island, beginning with the establishment of missions and government outposts in the late nineteenth century, and more recently, with the expansion of widespread logging throughout the island (Danielsen et al.

2010).

Colonization between satellite islands: (Rhipidura rufifrons ugiensis)

The Rufous Fantail (Rhipidura rufifrons) comprises 18 phenotypically-diverse subspecies that range from Indonesia to Australasia, Melanesia, and Micronesia. Three subspecies are native to

Makira-Ulawa province, including two white-throated races, R. r. russata on Makira and R. r. kuperi on Santa Ana and Santa Catalina, and a black-throated race, R. r. ugiensis, on Ugi.

However, R. rufirons has not previously been reported from the Three Sisters, and was absent from the comprehensive list of native birds that W. French (1957) compiled over 13 years of living and working on these small islands.

6 On 10 July 2012, a R. r. ugiensis was captured during mist-netting operations for other species in secondary habitat on Malaupaina, the southernmost and largest of the Three Sisters. Blood samples were obtained for later study and the bird was banded, photographed, and released. At least one additional R. r. ugiensis was observed nearby while netting on the same day, but not captured. No R. rufifrons subspecies were detected during a later visit to Malaulalo, the middle island of the Three Sisters, from 14-16 July 2013.

Several additional taxa native to both Ugi and Three Sisters either are not currently found on, or are historically-recent colonizers of Makira, including Monarcha castaneiventris ugiensis,

Ptilinopus richardsii, Myzomela cardinalis pulcherrima, and Cinnyris jugularis. Although these populations could be descended from a currently-extinct population on Makira, observation of R. r. ugiensis on Three Sisters suggests that migration between satellite islands likely has an important influence on species assemblages on these islands.

Potential altitudinal range expansion: Pacific Robin (Petroica multicolor polymorpha)

The Pacific Robin (Petroica multicolor) ranges widely throughout islands in the south Pacific.

Many subspecies, including those in , Norfolk Island, Fiji, and Samoa are found in forest and adjacent habitats at all elevations (Boles 2007). However, the four Solomon Island subspecies, P. m. septentrionalis of Bougainville, P. m. kulambangrae of , P. m. dennisi of , and P. m. polymorpha of Makira, are all believed to be restricted to high elevations, none below 800m (Dutson 2011).

On 1-2 Sept. 2014, I twice heard P. multicolor calling from the tree canopy along the upper reaches of the Ravo River upstream from the village of Na’ara, Makira. Both observations were significantly below the previously-reported altitudinal range of this species: one at 132 m (1

Sept. 2014, 57L 0820882 8831651), and one at 40m (2 Sept. 2014, 57L 0820723 8832775).

7 According to Joseph Waihuru, an experienced local naturalist and birding guide, P. multicolor are common along the lengths of the upper Ravo River. However, he has only observed the species nesting at high elevations. It is therefore unclear whether P. m. polymorpha undergoes seasonal altitudinal migration, whether it has been overlooked by previous expeditions, or whether it has undergone recent range expansion. However, this species is quite vocal, and the Ravo River and adjacent river valleys have been surveyed by several previous expeditions (Mayr 1945, Cain and

Galbraith 1956, Dutson 2001, Danielsen et al. 2010), all of which failed to detect this species at low elevations on Makira.

Much attention has been given in the literature to the tendency of species ranges to shift upwards toward higher elevations in response to climate change (Wilson et al. 2005, Lenoir et al.

2008, Chen et al. 2009). However, altitudinal range shifts in the opposite direction have also been identified across a wide variety of taxa, and have been hypothesized to be associated with anthropogenic habitat modification (Lenoir et al. 2010). Thus, although inconsistent with the expectation of altitudinal range shifts driven by global warming, further research may provide insight into whether recent widespread logging on Makira is responsible for the downslope expansion of P. multicolor.

Discussion

Three recent examples of native bird species observed outside their known ranges in Makira-

Ulawa province of the Solomon Islands possibly represent three different patterns of inter-island colonization: 1) from larger to smaller landmasses (Geoffroyus heteroclitus on Ugi), 2) from smaller to larger landmasses (Cinnyris jugularis on Makira), and 3) between small islands

(Rhipidura rufifrons ugiensis on Malaupaina). These examples provide some of the first direct evidence that small islands, as well as larger landmasses, may be important sources of island

8 biodiversity. Additionally, recent observations of Petroica multicolor in lowlands on Makira are suggestive of an atypical “downslope” altitudinal range expansion in a species previously known on the island only from much higher elevations.

Compared to other regions, the Solomon Islands have received relatively little focus in terms of ornithological research and avocational birding, so these sightings may potentially represent species that were previously overlooked. However, these species are generally conspicuous and easily observed, and if present, should have been readily detected by several previous expeditions led by distinguished field ornithologists. Accordingly, the observations reported herein likely represent recent arrivals, and possibly true range expansions and incipient colonizations.

Additionally, Makira-Ulawa province has been subject to widespread logging in recent years

(Danielsen et al. 2010). Anthropogenic habit change can be associated with species range expansions on continents (Hill et al. 2001, Kokko and López-Sepulcre 2006, Thomas 2010, Chen et al. 2011). However, while much emphasis has been placed on the ecological consequences of human-assisted species introductions to islands (Howald et al. 2007, Reaser et al. 2007, Sax and

Gaines 2008, Clavero et al. 2009), to my knowledge, no studies have examined the means by which anthropogenic habitat disturbance can facilitate natural colonizations and range expansions in island systems and the resulting potential effects on global biodiversity. Although direct evidence is lacking, such disturbance may have played an important role in the several of these range expansions. For example, during most of the twentieth century, nearly the entirety of the Three Sisters archipelago was covered by commercial coconut plantations (Cain and

Galbraith 1956), and since all records from these islands post-date the beginning of this period of agricultural use, it is possible that associated habitat conversion may previously have driven local extinction of R. rufifrons. Although small scale copra farming still occurs on Malaupaina,

9 secondary vegetation currently covers much of the island (J. Sardell, unpublished data), which could have facilitated a recent successful recolonization by R. rufifrons. Similarly, sunbirds and

Myzomela honeyeaters segregate ecologically elsewhere in Indo-Pacific, with the former found in disturbed open habitats and the latter found in more intact forest (Ripley 1959, Sanderson et al. 2009, Carstensen et al. 2011). Conversion of forested habitat to open habitat on Makira therefore may have facilitated the recent successful colonization of C. jugularis. Future studies of the competition between this species and the two species of Myzomela currently found on

Makira (M. tristrami and M. cardinalis) may provide important insight into the roles of both interspecific competition and anthropogenic habitat disturbance on island species assemblages.

Acknowledgments

I would like to thank the Solomon Islands Ministries of Environment and Education for permission to work in the Solomon Islands, as well the people in the Solomon Islands who made this work possible. In particular, I thank J. and J. Murray of Sanbiz Lodge, J. Tauni of Kirakira, J. and J. Waihuru of Hauta, H. Ha’aina of Molehanua, and the communities of Malaupaina and

Malaulalo. I also thank the local field assistants who provided support. Many thanks to J.A.C. Uy for logistical and financial support, and to D. Baldassarre for providing comments on the manuscript. Funding for this research was provided by the Society of Systematic Biologists,

Kushlan Graduate Research Support Fund, and Jay M. Savage Graduate Research Support Fund.

10 Figure 1: Map of Makira and satellite islands. Inset in upper right denotes location of Makira within the Solomon Islands.

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