Biogeography and Biotic Assembly of Indo-Pacific Corvoid Passerine Birds

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Annual Review of Ecology, Evolution, and Systematics Biogeography and Biotic Assembly of Indo-Paciﬁc Corvoid Passerine Birds Knud Andreas Jønsson,1 Michael Krabbe Borregaard,1 Daniel Wisbech Carstensen,1 Louis A. Hansen,1 Jonathan D. Kennedy,1 Antonin Machac,1 Petter Zahl Marki,1,2 Jon Fjeldsa˚,1 and Carsten Rahbek1,3

1Center for Macroecology, Evolution and Climate, Natural History Museum of Denmark, University of Copenhagen, DK-2100 Copenhagen, Denmark; email: [email protected], [email protected], [email protected] 2Natural History Museum, University of Oslo, 0318 Oslo, Norway 3Department of Life Sciences, Imperial College London, Ascot SL5 7PY, United Kingdom

Annu. Rev. Ecol. Evol. Syst. 2017. 48:231–53 Keywords First published online as a Review in Advance on Corvides, diversity assembly, evolution, island biogeography, Wallacea August 11, 2017

The Annual Review of Ecology, Evolution, and Abstract Systematics is online at ecolsys.annualreviews.org The archipelagos that form the transition between Asia and Australia were https://doi.org/10.1146/annurev-ecolsys-110316- immortalized by Alfred Russel Wallace’s observations on the connections 022813 between geography and animal distributions, which he summarized in Copyright c 2017 by Annual Reviews. what became the ﬁrst major modern biogeographic synthesis. Wallace All rights reserved traveled the island region for eight years, during which he noted the marked

Access provided by Copenhagen University on 11/19/17. For personal use only. faunal discontinuity across what has later become known as Wallace’s Line. Wallace was intrigued by the bewildering diversity and distribution of Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org

ANNUAL life he discovered. But even today we ask ourselves how species formed REVIEWS Further within the region and why they are not evenly distributed. Biogeography, Click here to view this article's online features: phylogeny, dispersal, biotic interactions, and abiotic factors affect the • Download ﬁgures as PPT slides • Navigate linked references assembly of diversity. On the basis of a decade of research on the ecology, • Download citations evolution, and systematics of corvoid passerine birds, we summarize what we • Explore related articles • Search keywords have learned about the biogeography and assembly of island bird diversity. Corvoid passerine birds include nearly 800 species and 2,300 named taxa and thus represent a large, well-described, and globally distributed clade. Understanding the processes inﬂuencing biodiversity in this group is certain to deepen our general understanding of ecology and evolution in the context of biogeography and faunal assembly.

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INTRODUCTION The distributions of organisms within the archipelagos that lie between Asia and Australia have featured prominently in the development of modern biogeography (Wallace 1860, 1863). The island region inspired Alfred Russel Wallace’s (1860, 1863) notion that the geography and geological history of a region shape its current fauna, and provided key evidence for the theory of evolution (Wallace 1876, 1880). During his eight-year exploration of the region, Wallace noted the marked faunal discontinuity across what later became known as Wallace’s Line, which he explained in terms of past movements of landmasses. He speculated about processes that might have influenced the generation, survival, and geographic distributions of insular faunas. Today, the eastern part of the island region bears his name—Wallacea—which, along with Australo- Papua and the Pacific Islands, constitutes the largest archipelagic system in the world, including thousands of islands spanning thousands of kilometers (Figure 1). The Indo-Pacific archipelagos have been a source of, and testing ground for, evolutionary, ecological, and biogeographical theories. The region’s extensive array of islands, which vary in shape, size, isolation, and historical opportunity for colonization, provides multiple replicates for investigating biogeography and community assembly. Rigorous tests of speciation, extinction, and dispersal scenarios using phylogenetically explicit frameworks have contributed to our understanding of accumulation and regulation of diversity (e.g., Rabosky & Glor 2010, Price et al. 2014, Smith et al. 2014). Moreover, plate tectonic reconstructions of the Indo-Pacific island region (e.g., Pigram & Davies 1987; Hall 2002, 2011; Schellart et al. 2006) provide a basis for interpreting the biogeographic history of the regional biota. Much like Wallace, twentieth-century biologists continue to be intrigued by the bewildering diversity of life forms, geographic distributions, and evolutionary innovations discovered in the course of fieldwork throughout the Indo-Pacific islands. MacArthur & Wilson’s (1967) seminal book, The Theory of Island Biogeography, introduced a framework describing island biotas as equilibrial systems in which the number of species reflects a balance between colonization and extinction. Although later modified and extended ( Johnson et al. 2000, Whittaker et al. 2008, Chen & He 2009, Rosindell & Phillimore 2011), this equilibrium model has remained influential. The Indo-Pacific island region also inspired ideas about how communities of coexisting species are structured (e.g., Diamond 1975, Connor & Simberloff 1979). Until recently, little was known about the evolutionary relationships of birds inhabiting the Indo-Pacific archipelagos, including a large clade of songbirds, the Corvides (Cracraft 2014), previously known as the core Corvoidea, which comprise nearly 800 species and 2,300 named taxa

Access provided by Copenhagen University on 11/19/17. For personal use only. (Dickinson & Christidis 2014). Sibley & Ahlquist (1990) suggested that a major clade of songbirds (Corvida, which is partly equivalent to Corvides) had originated in Australasia and later dispersed Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org across the Australasian archipelagos to colonize most other continents. Barker et al. (2002) and Ericson et al. (2002) subsequently showed that, in fact, songbirds (oscines, of which there are approximately 4,500 species and which constitute approximately 45% of all birds) originated in the Australasian region (see also Edwards & Boles 2002). However, taxon sampling has been sparse ( Jønsson & Fjeldsa˚ 2006) and only during the past few years have multigene phylogenies and dense taxon sampling supported a detailed reconstruction of the evolutionary history of the Corvides— one of the most dominant avian clades in the region. Reconstructions of ancestral distributions of songbirds suggest origin and dispersal out of Australia, which match major tectonic events in the Indo-Paciﬁc that facilitated global colonization of both the Passerides and the Corvides (Figure 2).

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Taiwan

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Vanuatu Samoa

Australia Fiji New Caledonia Tonga

New Zealand Access provided by Copenhagen University on 11/19/17. For personal use only. Figure 1

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org Map of the Indo-Pacific archipelagos, excluding Sri Lanka and western Indian Ocean islands as well as eastern Polynesian islands such as Hawaii, French Polynesia, and Cook Islands. Wallace’s (imaginary) line goes between Sundaland ( green) and Wallacea (orange and yellow). In this review, we summarize a decade of work on the biogeography, evolution, and systematics of one of the prominent evolutionary lineages of the Indo-Pacific, the corvoid passerine birds. Although our central focus is on this group and region, we believe that insights derived from this system apply to other taxa and regions as well. Drawing on genetic data that reveal evolutionary relationships, as well as information on life histories, ecology, and distributions from the literature, we discuss how findings provided by the study of Indo-Pacific corvoids relate to the distribution and diversification of species more generally. We refer to classic work on island systems but also discuss more recent issues and future challenges.

www.annualreviews.org • Biogeography and Biotic Assembly 233 ES48CH11-Jonsson ARI 9 October 2017 7:38

Figure 2 World map representation of the present distribution of 789 recognized species of the Corvides in 1◦ × 1◦ grid cells (Rahbek et al. 2012). The bar to the left indicates the number of species per grid cell.

CORVIDES PHYLOGENY: IMPLICATIONS FOR UNDERSTANDING THE ORIGIN OF MODERN DIVERSITY The clade Corvides includes a great diversity of birds (Figure 3), whose body weights vary more than 100-fold (Kennedy et al. 2012). The corvoid clades include some large continental and insular radiations (e.g., Vireonidae and Pachycephalidae), which diversiﬁed over extensive areas with relatively little morphological change; other corvoid clades represent dramatic examples of adaptive radiation with respect to foraging ecology (the Madagascan vangas; Jønsson et al. 2012a) or sexual attraction (birds-of-paradise; Irestedt et al. 2009). Additionally, some small clades (1–3 species) represent ancient relict families restricted to New Guinea or Australia (e.g., Corco- racidae, Eulacestomatidae, Ifritidae, Machaerirhynchidae, Melampittidae, Neosittidae, Oreoici- dae, Paramythiidae, and Rhagologidae).

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Access provided by Copenhagen University on 11/19/17. For personal use only. Figure 3

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org Variation in size and shape among clades of the Corvides, as represented by species from 26 of the 31 families. Top from left to right: Australian magpie (Gymnorhina tibicen, Cracticidae), varied sitella (Daphoenositta chrysoptera, Neosittidae), Australian raven (Corvus coronoides, Corvidae), bay-backed shrike (Lanius vittatus, Laniidae), fork-tailed drongo (Dicrurus adsimilis, Dicruridae). Middle from left to right: white-breasted woodswallow (Artamus leucoryn, Artamidae), yellow-breasted boatbill (Machaerirhynchus flaviventer, Machaerirhynchidae), bristlehead (Pityriasis gymnocephala, Pityriaseidae), common iora (Aegithina tiphia, Aegithinidae), rosy-patched bushshrike (Rhodophoneus cruentus, Malaconotidae), blue vanga (Cyanolanius madagascarinus, Vangidae), yellow-bellied wattle-eye (Dyaphorophyia concreta, Platysteiridae), white-crested helmetshrike (Prionops plumatus, Vangidae), black-eared shrike-babbler (Pteruthius melanotis, Vireonidae), golden whistler (Pachycephala pectoralis, Pachycephalidae), hooded pitohui (Pitohui dichrous, Oriolidae), rufous fantail (Rhipidura rufifrons, Rhipiduridae), Madagascar paradise flycatcher (Terpsiphone mutata, Monarchidae), brown sicklebill (Epimachus meyeri, Paradisaeidae), crested berrypecker (Paramythia montium, Paramythiidae). Bottom from left to right: lesser cuckooshrike (Lalage fimbriata, Campephagidae), Papuan whipbird (Androphobus viridis, Psophodidae), spotted jewel-babbler (Ptilorrhoa leucosticta, Cinclosomatidae), white-winged chough (Corcorax melanorhamphos, Corcoracidae), wattled ploughbill (Eulacestoma nigropectus, Eulacestomidae), blue-capped ifrit (Ifrita kowaldi, Ifritidae), lesser melampitta (Melampitta lugubris, Melampittidae). Watercolor by Jon Fjeldsa˚.

234 Jønsson et al. ES48CH11-Jonsson ARI 9 October 2017 7:38 r i m e r l l u k e i i e e c y t b a a e e n e d l d p o i i m

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m t a a w d r y a r e a o m i t r m a P Paradisaeidae s p a Brown sicklebill B Brown P Paramythiidae r e Epimachus meyeri E s r i a a r t Crested berrypeckerC Crested Paramythia montium Paramythia P t b e i u a p g d i u m t l

t a i l a e p t t i m m

p a r e l a s e m e i t s a l d s a M Melampittidae t e e a e r u a Lesser melampitta L Lesser r M Melampitta lugubris a e d m i

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a e a e e u s e l a Terpsiphone mutata Terpsiphone T r i w u M Madagascar paradise p c d r a o i i p t u t k -

i D Dicruridae r a r k a c c f e r i t I Ifritidae - i k o r i D Dicrurus adsimilis s e f r Fork-tailed drongo F Fork-tailed I Ifrita kowaldi u u l t h s e B Blue-capped ifrit a

t i a s t d i u d u e i v h i

s o k s r o e n u c t t u h a a i l a s i l c c L Laniidae i d i p n n b e i

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e a d y r i o a L Lanius vittatus h f a i o e t a d u r r i g fi d d Bay-backed shrike B Bay-backed n i h g u u O Oriolidae i r a o m o r f

o n t

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i a s t d a p r Pitohui dichrous Pitohui P H Hooded pitohui i s

r u u e p e d m o i l d c f e i o h l b a t u s p l t i s i R Rhipiduridae b t Rufous fantail R Rufous t u e a h a o E Eulacestomidae c s Rhipidura rufifrons Rhipidura R b n i a - Wattled ploughbill W Wattled l l e a e l a u a r k e e r Eulacestoma nigropectus Eulacestoma E i d o i r a e t m l

n h d c t i s s s l e o

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h r h h i e a t l r w p V Vireonidae

u a a e r n e h c e e e - y t p k k d i e P Pteruthius melanotis h l c r c c o a h y l a s s G Golden whistler h Black-eared shrike-babblerB Black-eared Pachycephalidae P t u c t e a a s Pachycephala pectoralis Pachycephala P m e l m o a e h u h l d h i g p

g u d s m n o e p e a t a h o a s h c V Vangidae r

e n d i r o e o d s a c i c y n t e r u - s a e a e r P Prionops plumatus Prionops g n e e n l - r i t o e d n r e i c e i i l c v a h n t r e o m c e a t w

o a o - r n s x W White-crested helmetshrike a a

d e C Corcoracidae o a a i d n r t a w a i r r g i

i e a i o g v o i a y h d a l e c r c

n t d h e a r d s o i a W White-winged chough s i r l a p o u t l C Corvidae v y g

o s C Corcorax melanorhamphos Corcorax t v e m r n

e r a u b l s o a o u e - Australian raven A Australian l s P Platysteiridae u h V Vangidae k Corvus coronoides C i i B Blue vanga u w p r t n o a l h n a l y l s e e e o D Dyaphorophyia concreta Dyaphorophyia h u a Yellow-bellied wattle-eye Y Yellow-bellied n r s d c i a u

a t y s i a b e

o r Cyanolanius madagascarinus Cyanolanius C u h a d n e o p i i d

e o n i t a

c l n h o n a i a c a o h l t n h h p i a a t m i p o h p e t M Malaconotidae r e g - d i m d c a e e y o g t o l a o d s l e Aegithinidae A h i i n e Common iora C Common n o Aegithina tiphia Aegithina A e e Rhodophoneus cruentus R b e r h Rosy-patched bushshrike R Rosy-patched s t v m a e e i a l l a a y t i v d t i b c r o e g s a i

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s c a i d e s B Bristlehead s i d t o n P Pityriaseidae b t a u c e p - y i a l t r h Access provided by Copenhagen University on 11/19/17. For personal use only. u o h s e c y s e r m l a t i a y

n i l w e r o e r l y a Pityriasis gymnocephala Pityriasis P a e r s e e h j h o

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t e c r w t i d e n d t d t a r i i s o o t a i i l r l e e i i s C Cinclosomatidae o l i o h b M Machaerirhynchidae t a r v o e e c

p p P Ptilorrhoa leucosticta a d n Yellow-breasted boatbill Y Yellow-breasted s i n a i Neosittidae N Spotted jewel-babbler S Spotted d y Varied sitella V Varied e u h r d e w Machaerirhynchus flaviventer Machaerirhynchus M o b t o e o w o s

h l o c a l h a n p h u d a p e i a e a p r l o w

u o Daphoenositta chrysoptera D s m b s s r - p a P Psophodidae u d d a t e r t o n Papuan whipbird P Papuan m i o A Androphobus viridis A Artamidae a h t n w woodswallow r e W White-breasted i e A Artamus leucoryn c p i g b e i a a t

d e m a i

k c n e i i n i a r t t a a h i c h r a d l i i s a o a r r g o r n b t C Cracticidae a o s k m h m u y c fi p

Australian magpie A Australian u G Gymnorhina tibicen e e c p

g r a l e m s a a s Lalage fimbriata L Campephagidae C e Lesser cuckooshrike L Lesser

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The first European explorers of Australasia generally assumed that the local birds (and other organisms) belonged to groups with which they were already familiar. New species were simply considered members of known groups. Consequently, species of passerine birds were placed among, for instance, Old World finches (Fringillidae) and flycatchers (Muscicapidae). Skilled nat- uralists such as Wallace realized that several Australasian groups represented very divergent taxa, but many early misclassifications persisted for more than a century. For example, some small insect-eating birds resembled Pachycephala whistlers, which became a repository for unassignable insectivorous birds in the Indo-Pacific. Moreover, much systematic and taxonomic work on Indo- Pacific birds was carried out by Ernst Mayr, whose conservative species-level taxonomy lumped many allopatric island taxa into so-called superspecies complexes. Mayr’s taxonomic legacy lives on to this day, and many of his superspecies will undoubtedly be split following new and genetically informed taxonomic reevaluations. With the advent of DNA systematics, ornithologists began to understand that the Australasian fauna differs in many ways from the avifauna of Europe, and molecular studies published within the past 15 years have resolved most of the deep-time relationships within the Corvides (Aggerbeck et al. 2014, Moyle et al. 2016). Detailed phylogeographic assessments suggest that Indo-Pacific regional diversity is generally underestimated (e.g., Lohman et al. 2010, Irestedt et al. 2013), but also that some polytypic species, such as the golden whistler (Pachycephala pectoralis) complex with 66 named subspecies, actually represent nonmonophyletic populations of birds with a conserved morphology (Andersen et al. 2014, Jønsson et al. 2014). Rapid early diversification has made it difficult to confidently resolve relationships at the base of the corvoid tree. However, a comprehensive, multilocus analysis of the primary (family-level) corvoid lineages established three main corvoid clades (X, Y, and Z of Aggerbeck et al. 2014, Jønsson et al. 2016b), which we refer to as the Orioloidea, Malaconotoidea, and Corvoidea. The Malaconotoidea and Corvoidea together are sister to the Orioloidea. Ongoing analyses of genomic data (see also Moyle et al. 2016) corroborate these three main clades but move the corvoid families Campephagidae, Cinclosomatidae, and Neosittidae to the base of the corvoid tree.

TECTONIC HISTORY AND CLIMATE OF THE INDO-PACIFIC The origin and evolution of the Corvides have taken place against a background of geological change in the configuration of land in the Indo-Pacific island world throughout the Cenozoic (see Figure 4 for part of the region). Extensive rearrangements of landmasses (including emergence, uplift, and disappearance of islands) coupled with periods of global temperature fluctuations (and associated changes of sea levels and habitats) have influenced faunal assembly in the region (e.g., Access provided by Copenhagen University on 11/19/17. For personal use only. Pigram & Davies 1987, Hall 2002, Schellart et al. 2006). Of course, early biogeographers did not

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org know about these changes, although Wallace (1880) pondered the matter. The positions of islands were generally assumed to have been static; only Quaternary sea-level changes were believed to have affected land connections and dispersal opportunities across open sea. Thus, islands and archipelagos were regarded as dead ends, in the sense that the diversity of island life was seen as a consequence of colonization and eventual extinction (e.g., MacArthur & Wilson 1963, 1967; Mayr 1965). More recent studies have demonstrated substantial diversification within archipelagos and even reverse colonization of continents from islands (Filardi & Moyle 2005, Bellemain & Ricklefs 2008, Esselstyn et al. 2009, Jønsson et al. 2011, Fritz et al. 2012, Jønsson & Holt 2015). Consequently, the changing positions and configurations of landmasses over time have had a marked influence on colonization and diversification (e.g., Wallace 1863, Hosner et al. 2013).

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Volcanos Highland Land Carbonate platforms Shallow sea Deep sea

5 Ma 10 Ma Early Pliocene Late Miocene

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15 Ma 20 Ma Middle Miocene Early Miocene

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0°

10°S

20°S 90°E 100°E110°E 120°E 130°E 140°E 150°E 90°E100°E 110°E 120°E 130°E 140°E 150°E

Figure 4 Distribution of land and sea from the mid-Oligocene to the present in the area between Australia and Southeast Asia. Figure courtesy of Robert Hall.

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In the following section we summarize current knowledge about the changes in landmass configuration in the region during the past 30–40 million years. For a more exhaustive overview, we provide key references and note that substantial uncertainties surround geologic and climatic reconstructions. Clearly, however, substantial changes in the distribution of land and sea have influenced the distribution of taxa and the evolutionary diversification of island communities.

Australo-Papua Australia began to separate from Antarctica in the Late Cretaceous approximately 80 Mya (Metcalve 1998). By 40 Mya, seafloor spreading between the continents caused the Australian plate to drift rapidly northward until it collided with terranes of Asian origin approximately 20 Mya (Hall 1998, 2002; van Ufford & Cloos 2005). Through the mid-Tertiary, the northern (Papuan) part of the plate was mostly submerged but intermittent islands were associated with uplift of carbonate platforms. Plate fragments representing the present Bird’s Head (Vogelkop) Peninsula of western New Guinea, the southern Moluccas (with Seram and Buru), and eastern Sulawesi broke apart from the Australian plate in the late Oligocene (the Sula Spur, evidently with some subaerial areas already present at this early stage; Hall & Sevastjanova 2012, Hall 2013) and approached the chain of volcanoes that emerged north of Sulawesi. Such islands may have provided stepping-stones for early dispersal from Australia to the Oriental region. Approximately 10 Mya, plate fragments of Asian and Australian origin had become greatly intermingled, forming distinct groups or chains of islands, with numerous subaerial stepping-stones between the two continents (Hamilton 1979; Hall 1998, 2002). The climate in the northern tropical part of this region was probably stabilized by a warm surface current (Hall 2011). Farther south, however, Australia experienced significant aridification beginning in the Miocene (approximately 15 Mya). This transformed a heavily forested continent to its current arid interior with isolated patches of rain forest and open sclerophyllous forest at the perimeter in the north, east, and extreme southwest (Byrne et al. 2008, 2011).

Paciﬁc Archipelagos The Melanesian archipelagos (Bismarck, Solomon, Santa Cruz, Vanuatu, and Fiji Islands) as well as the northern mountain ranges of New Guinea are part of a volcanic island arc system at the subduction zone between the Australian and Paciﬁc plates (Hall 1998, 2002). This arc system was pushed westward in front of the advancing Australo-Papuan plate since at least 25 Mya, causing subduction along the eastern margin of the Eurasian plate and strike-slip faulting along the northern margin of the Australian plate. In this process, several terranes were accreted to the northern margin of the Australian plate, forming the coastal mountain ranges of New Guinea (e.g., the Torricelli and Adelbert Ranges) (Hall 2002). This arc system was nearly continuous from the Access provided by Copenhagen University on 11/19/17. For personal use only. Philippines to Fiji from approximately 25 Mya until approximately 12 Mya, when the opening of

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org the North Fiji Basin began to create a gap between Fiji and Vanuatu (Hall 2002, Schellart et al. 2006). The area of land in this region during different periods is poorly known, but some parts of the Melanesian Arc have undoubtedly been above water during periods of low sea levels, forming partial and intermittent land connections along the chain and potentially facilitating the dispersal of organisms between them. New Caledonia is not part of the Melanesian Arc but belongs to the Norfolk Ridge, which is connected southward to the large continental plateau that includes New Zealand. New Caledonia probably rose above sea level approximately 35 Mya (Cluzel et al. 2001), although some geological evidence suggests that the entire Zealandia plate may have been submerged during the mid- Tertiary; nonetheless, the persistence of some ancient taxa suggests that subaerial land must have remained somewhere in the region at all times (Gibbs 2006). This has some relevance for interpreting the origin of phylogenetically isolated corvoid lineages such as Mohoua and Turnagra.

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Sundaland This area includes the shallow extension of continental Asia, the Malay Peninsula, and the Greater Sunda Islands of Java, Sumatra, and Borneo. The Greater Sunda Islands have increased substan- tially in area since the mid-Miocene because of uplift. These islands were connected to each other and to mainland Asia until the early Pliocene, when rising sea levels caused them to become separate islands, requiring overwater dispersal of land taxa for colonization throughout the Quaternary (Chappell 1987; Hall 1998, 2002; Sheldon et al. 2015). Palawan originated as a distinct microplate near the South Chinese coast in the Eocene, then drifted south but did not become subaerial until the Miocene, and was accreted to the Sunda Shelf during the Pleistocene.

Wallacea During the Miocene, tectonic movements resulted in the formation of the archipelagos between the Asian and Australo-Papuan continents. They formed when the Asian and Australian plates collided, causing plate fragments and oceanic island chains to intermingle in a counterclockwise direction. However, it is difficult to determine when many of these islands rose above sea level (Hall 2009). Many adjacent island groups are separated by deep ocean trenches and have never been connected. The region includes the island of Sulawesi, which is of mixed plate origin and consists of terranes from the Asian, Indo-Australian, and Pacific plates, and volcanic areas that rose from the sea. The Lesser Sunda Islands are oceanic, with the exception of Sumba, which is of Sundaland origin, and Timor, which is of Australian origin, although subaerial only since the Pliocene. Most of the islands in the Moluccas originated from the Australo-Papuan plate, whereas Halmahera was part of the oceanic Philippine–Halmahera Arc system and thus rafted in from far out in the Pacific Ocean (Hamilton 1979, Hall 1998, Moss & Wilson 1998).

Philippine Archipelago The Philippine archipelago is also of mixed geographical origin. The West Philippines, with Luzon, is oceanic and was formed in the mid-Eocene in the Proto-South China Sea northeast of Borneo. The East Philippines, with Mindanao, is also of oceanic origin. It formed in the Oligocene and rafted close by New Guinea’s Bird’s Head in the Miocene, when a very long stepping-stone chain of islands existed between the Philippines and New Guinea via Halmahera. Although the Philippines have been isolated throughout the Tertiary, a Miocene stepping-stone connection between Borneo and Luzon via the Sula Arc is possible, and later, Palawan formed another stepping-stone route (Hall 1998, 2002). More recently, Plio-Pleistocene sea-level changes

Access provided by Copenhagen University on 11/19/17. For personal use only. may have signiﬁcantly altered the amount and extent of land in the Philippines, which likely contributed to the assembly of diversity for some groups (e.g., Hosner et al. 2013). Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org

ORIGIN AND CURRENT DISTRIBUTIONS OF CORVOIDS Corvoid passerine birds are distributed across all continents, except Antarctica, and all zoogeo- graphic regions (Figure 2), but exhibit their highest contemporary diversity in Australasia, specifically on the island of New Guinea, where up to 93 species co-occur within a single 110 × 110 km grid cell (Kennedy et al. 2017). These observation and biogeographic analyses based on molecular phylogenies suggest that the group originated during the Oligocene epoch on islands in the area currently occupied by New Guinea, at the edge of the Australian plate. These proto-Papuan islands may have acted as a springboard for early dispersal and global colonization (Aggerbeck et al. 2014, Jønsson et al. 2011). New Guinea is also home to some small endemic corvoid families [e.g., Eulacestomatidae (monotypic), Ifritidae (monotypic), Paramythiidae (two species, four taxa), and

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Machaerirhynchidae (two monotypic species)]. Thus, New Guinea appears to be both a species pump (cradle) and a museum (graveyard) for corvoids. The formation of the first terrestrial habitats north of Australia, in the Sula Spur area some 30–20 Mya, corresponds to our estimate of the timing of early dispersal of Corvides out of Australo- Papua to the Indo-Pacific islands and subsequently to mainland Asia, Africa, and the Americas ( Jønsson et al. 2011). The Papuan origin hypothesis is corroborated by other animal groups (Oliver 2011, Toussaint et al. 2014). However, critics have argued that taxa previously widespread across a forested Australo-Papuan continent during cooler and wetter times have simply retracted and now occur exclusively as relicts in forest pockets along the perimeter of the Australo-Papuan continent (Schodde & Christidis 2014, Moyle et al. 2016). The Papuan origin hypothesis is also criticized because of the temporal mismatch between the origin of Corvides and the main uplift of New Guinea, which dates back to the late Miocene/Pliocene (Moyle et al. 2016). Although we agree that many corvoid families expanded and diversified at this later time, evidence of subaerial areas already in the late Oligocene (Sula Spur islands; see Hall & Sevastjanova 2012), corresponding to the appearance of the first fossils of oscine birds in the Old World (Manegold 2008), suggests that an early dispersal event may have occurred long before the major period of uplift and diversification in the Indo-Pacific island region. To further assess where corvoid diversity originated and how it became globally distributed over time, we estimated speciation within, and transitions between, regions using ancestral area reconstruction (see Jønsson & Holt 2015 for details). Most diversity is generated within regions. The largest landmass, Australia, exported little of its corvoid diversity, whereas New Guinea was the source area for all other Indo-Pacific regions. The prevalence of Papuan distributions toward the base of the Corvides tree further suggests that the extant Corvides diversity is deeply rooted in New Guinea ( Jønsson et al. 2011, Aggerbeck et al. 2014, Kennedy et al. 2017). This may reflect the retention of ancient Corvides diversity in Papua or the generation of Corvides diversity over millions of years in Papua. Of the 31 Corvides families, 24 occur in Australia or New Guinea. Of these, seven (Corvidae, Monarchidae, Rhipiduridae, Dicruridae, Artamidae, Campephagidae, and Pachycephalidae) have colonized Pacific islands, of which all but two (Dicruridae and Artamidae) extend to remote Pacific islands (Figure 5). To the west of Australo-Papua, the colonization patterns are complex. Some families colonized continental Asia, which may have led to the colonization of the Americas via Beringia (e.g., Vireonidae and Corvidae). Others colonized Africa (e.g., Vangidae, Platysteiridae, and Malaconotidae), but continental Asian groups also back-colonized the Indo-Pacific islands (e.g., Oriolidae, Dicruridae, and Corvidae), and some monotypic families represent relictual forms on large Indo-Pacific islands (e.g., Pityriaseidae on Borneo). Access provided by Copenhagen University on 11/19/17. For personal use only.

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org DISPERSAL Despite the acceptance of dispersal as a key process in shaping distribution patterns and community assemblages (Ronquist 1997, Zink et al. 2000, de Queiroz 2005, Heaney 2007), we still know little

Figure 5 Indo-Paciﬁc island (extant) species richness maps indicating the number of corvoid species per island larger than 10 km2 (381 corvoid species in total; only islands with more than 2 species are shown). From top left, Campephagidae (65 species), Paradisaeidae (41 species), Corvidae (34 species), Dicruridae (12 species, not including Chaetorhynchus), Oriolidae (22 species, including Pitohui kirhocephalus and Pitohui dichrous), Monarchidae (79 species), Pachycephalidae (51 species), and Rhipiduridae (44 species, including Lamprolia and Chaetorhynchus). Number of species distributed in the tropical Southeast Asian mainland is indicated on each individual mapinthetopleftcorner.

240 Jønsson et al. ES48CH11-Jonsson ARI 9 October 2017 7:38 41 species 22 species 44 species Oriolidae Rhipiduridae Paradisaeidae 1 2 3 4–5 6–33 1 2 3 4 5–6 1 2 3 4–5 6–14 0 6 3 Species richness Species richness Species richness 65 species 12 species 51 species Dicruridae Campephagidae Pachycephalidae 1 2–3 4–5 6–8 9–15 1 2 3 4 5 1 2 3 4–5 6–18 7 1 11 Species richness Species richness Species richness Access provided by Copenhagen University on 11/19/17. For personal use only. Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org s e e a i c d i e v p s r

o 4 Corvoid 3 34 species 79 species C Corvidae 381 species Monarchidae 2–3 4–7 8–12 13–17 18–132 1 2–3 4–5 6 7–8 1 2 3 4–5 6–15 3 14 Species richness Species richness Species richness

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about dispersal and colonization at the individual, population, or community level. One taxon may disperse readily but become established infrequently, whereas another might disperse poorly but establish readily where it does land. Among the Corvides, only four families have radiated extensively within the Pacific archipelagos. Three of these families (Monarchidae, Rhipiduridae, and Pachycephalidae) include small insectivorous species with intermediate hand-wing indexes, suggesting moderate capacity for active dispersal, whereas the Campephagidae have predominately high hand-wing indexes and om- nivorous feeding habits (Kennedy et al. 2016). Assuming the adequacy of wing morphology as a proxy for dispersal, this suggests that active flight alone may not explain the colonization of remote archipelagos or that new island colonizers undergo rapid evolutionary shifts in dispersal ability (Kennedy et al. 2016). The ancestral area analysis across the whole Corvides tree suggests a dynamic exchange of diversity between Indo-Pacific archipelagos. New Guinea has exported diversity to all regions, whereas Australia has been a minor source of diversity. Fifteen colonization events to the west toward Wallacea originated in New Guinea, and two of these extended to the Oriental biogeographic region. Ten colonization events occurred from New Guinea eastward toward Melanesia and Polynesia, with five secondary colonizations of Polynesia from Melanesia. Polynesia itself has not exported any corvoid diversity and is an example of a macroevolutionary sink. In contrast, Micronesia, despite consisting of small, remote islands, has contributed some diversity to other regions. Also noteworthy is that New Guinea appears to have exported 13 lineages to Australia but has received only 2 in return. Inferring dispersal and colonization from phylogenies and contemporary distributions depends on many assumptions. We still know little about the stochastic versus deterministic nature of dispersal (Lowe & McPeek 2015), how dispersal is related to behavioral traits, and how dispersal differs between central and peripheral populations in a species’ range (e.g., Lindstrom¨ et al. 2013).

SPECIATION Some of the most long-standing questions in ecology pertain to why some groups of organisms are more abundant, are more species rich, and have larger ranges than others. In other words, what determines the success of clades? Simpson (1953) suggested that ecological opportunity is an important driver of diversiﬁcation. Across the dynamic Indo-Paciﬁc archipelagos, islands emerge, change in size, are rearranged spatially, and sink or are eroded back into the ocean over time (Whittaker et al. 2008). The highly dynamic nature of these archipelagos likely created rich opportunities for the generation of allopatric populations and, possibly, for speciation. Species richness also might depend simply on time; that is, the longer a clade has existed the Access provided by Copenhagen University on 11/19/17. For personal use only. more time it has had to accumulate species. Previous analyses have reached contrasting conclusions

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org regarding the importance of clade age as a correlate of species richness (Ricklefs 2006, McPeek & Brown 2007, Rabosky 2009, Wiens 2011), and within the Corvides some of the oldest clades have only 1–5 species. Although we cannot know whether such clades always contained few species, the Corvides ( Jønsson et al. 2011, 2014) provide some evidence for an alternative explanation, i.e., that a low species number is a consequence of relictualization following the contraction of clades over time (Ricklefs & Jønsson 2014). Intuitively, it seems reasonable that older clades may at a certain point stop expanding and enter a phase of diversity decline as has been suggested for Caribbean birds (Ricklefs & Bermingham 2008). Diversity decline may be seen in light of population senescence, in which expanding taxa experience an initial release from pathogens, which subsequently catch up and cause the decline of taxa or even whole clades (Ricklefs & Jønsson 2014). This decline would explain the lack of a consistent relationship between clade age and clade diversity and would link the patterns described

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here to the rise and fall of species richness in the fossil record (Raup et al. 1973, Foote 2007). Recent studies suggest that it may be difﬁcult to detect a signal of diversity decline in the absence of a good fossil record, even with complete extant species-level molecular phylogenies (Quental & Marshall 2011). Population persistence is lower, i.e., extinction risk is higher, on small islands than it is on larger islands (Pimm et al. 1988). This is evident from our data on Corvides, which show that large islands harbor older and evolutionarily more distinct species (Figure 6), which have persisted longer than the average lineage life span. Conversely, small islands harbor young taxa, on average.

EXTINCTION Extinctions may play a significant role in structuring island faunas. However, loss of species on remote islands accelerated after the arrival of humans (Steadman 2006). The work of Steadman and others (e.g., Duncan et al. 2002, Boyer 2009) on fossil bird faunas show that the larger flightless landbirds, such as rails and pigeons, which were subjected to greater hunting pressures, have been eradicated. Except on the Hawaiian Islands (e.g., Olson & James 1982, James & Olson 1991), few extinctions of passerine birds have been documented, and among the Corvides only two of these extinctions can be attributed to prehistoric human colonization of Pacific islands. However, with losses of the terrestrial avifauna between 35% and 50% since human colonization (Olson & James 1982, Boyer 2009), extinction could certainly influence the results of analyses assessing the assembly of island communities, even when restricting analyses to a clade such as the Corvides. Although the fossil record for birds is relatively poor, particularly for tropical groups, some Corvides fossils are known from Australia (Boles 1999, Nguyen et al. 2013, Nguyen 2016) and North America (Brodkorb 1972). These fossils represent the only hard evidence for extinctions of Corvides species and provide little insight into whether particular clades, regions, or times have experienced extraordinary rates of extinction. What we do know is that much of the Indo- Pacific island world is associated with active volcanoes (the so-called Pacific Ring of Fire) and that enormous eruptions may have degraded environments on multiple occasions and even wiped out entire island faunas. In a similar way, climate change on the Australian continent caused dramatic habitat shifts from predominantly forest to arid desert (Byrne et al. 2008, 2011). Both volcanic activity across the Indo-Pacific islands and aridification of Australia likely resulted in extinctions and dramatic alterations of species compositions but left little hard evidence behind. The development of extinction models tested with phylogenies of groups with good fossil records has progressed rapidly (Morlon et al. 2011, Etienne et al. 2012), but in many cases results have been counterintuitive and difficult to reconcile with other evidence. However, despite the

Access provided by Copenhagen University on 11/19/17. For personal use only. lack of fossil evidence, we assume signiﬁcant background extinction for island faunas (Marki et al. 2015). An alternative and perhaps more realistic approach would be to search for distributional Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org signatures within clades that are well sampled at the subspecies or population level (e.g., Ricklefs & Bermingham 2008, Jønsson et al. 2014). Here, past extinctions may be inferred from cases of signiﬁcant range disjunctions, as demonstrated by the Pachycephala radiation (described below).

PHYLOGEOGRAPHY: EVOLUTION ON RECENT TEMPORAL SCALES Although the species-level phylogeny for Indo-Paciﬁc Corvides is near completion ( Jønsson et al. 2016b), more detailed population-level studies are needed, and a review by Beheregaray (2008) demonstrated that phylogeographic studies both for the region (Australasia) and for birds in general are scarce. So far, most studies have analyzed small taxonomic groups or a single archipelago (e.g., Deiner et al. 2011, Kearns et al. 2011, Moltesen et al. 2012, Hosner et al. 2013; but see Jønsson et al. 2014).

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Figure 6 a Taxon cycles predict 25 (a) small islands should not support old or evolutionarily distinct 20 taxa. Large islands hold a combination of old and young taxa. 15 (b)Oldor evolutionarily distinct 10 taxa should inhabit higher elevations. Evolutionary distinctnessEvolutionary Elevational 5 distributions are presented as the mid-elevation for each species. Note that 2 4 6 8 10 12 14 evolutionarily distinct log mean island size (km2) taxa (>15) are absent from the lowest b elevations and 25 evolutionarily indistinct (<2) are absent from the 20 highest elevations. (c)Oldor 15 evolutionarily distinct taxa occur only on a few (large) islands 10 where they are able to persist, whereas young Evolutionary distinctnessEvolutionary species occur on many 5 (but also on few) islands. Evolutionary distinctness (ED) 0 1,000 2,000 3,000 captures the degree of Species mean elevational distribution (m) species isolation on the phylogenetic tree (Isaac et al. 2007) and c was calculated across 25 all species occurring

Access provided by Copenhagen University on 11/19/17. For personal use only. on the examined 20 islands. Islands with

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org high ED scores host faunas that consist of 15 old and isolated phylogenetic lineages (as expected of stage 10 IV of the taxon cycle).

Islands with low ED distinctnessEvolutionary scores host 5 predominantly young and rapidly diversifying species 012345 (stage I of the taxon log number of islands cycle).

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One example that has provided a wealth of information about evolutionary dynamics, as well as detailed information about relationships at the island population level ( Jønsson et al. 2008, 2014; Andersen et al. 2014), comes from the family Pachycephalidae (whistlers). For example, in the Bismarck Archipelago, one whistler species (Pachycephala citreogaster) occurs on the large islands, whereas a different, albeit similar, species (Pachycephala melanura) occurs on the smaller islets and occasionally along the coasts of the larger islands. On the basis of these distributions, Mayr & Diamond (2001) used P. melanura as an example of a supertramp taxon, i.e., a highly dispersive generalist taxon with low competitive ability. Our genetic analyses have first established that P. citreogaster and P. melanura represent distinct taxonomic units. In addition, they revealed that P. citreogaster populations on large islands are approximately three times as old as P. melanura populations on small islands, and that island populations of P. citreogaster are distinct from one another compared with the genetically undifferentiated island populations of P. melanura within the Bismarck islands. Thus, the findings from Bismarck whistler populations support the idea of an early colonization by P. citreogaster leaving sufficient time for establishment on large islands and subsequent between-island differentiation, as well as the disappearance of the species from the smallest islands. P. melanura colonized the Bismarck islands more recently, but this taxon is plausibly still in the highly mobile tramp phase and thus is undifferentiated among islands. Phylogeographic studies may also provide insight into the frequency of long-distance, overwater dispersal. Birds vary greatly in their capacity for long-distance dispersal. Some species are sedentary and stay within vegetation cover; others cross water readily to establish new populations on small newly emerged islands (Diamond 1974, Mayr & Diamond 2001). We may expect that dispersal abilities change, perhaps even rapidly, once a population has become established on an island. Evolution of behavioral flightlessness is documented for several groups of organisms (e.g., birds: Diamond 1981, Moyle et al. 2009; ants: Wilson 1959; butterflies: Holloway 1977). Lamprolia victoriae of Fiji and Chaetorhynchus papuensis of the New Guinean highland present a sister species example of highly sedentary birds confined to primary forest that at some point must have dispersed a long distance over water to reach Fiji, probably via the Melanesian Arc (Irestedt et al. 2008). Even if populations representing this lineage were once widely distributed along the Melanesian island arc, they nonetheless would have required a better dispersal capacity in the past to colonize these islands. Because shifts in dispersal ability and life strategy can be expected of any type of organ- ism, biogeographical patterns should be interpreted with the possibility of change in life-history traits (Moyle et al. 2009). The example of L. victoriae and C. papuensis is at the heart of the debate. Does this range disjunction reflect long-distance dispersal, or does it represent a late Access provided by Copenhagen University on 11/19/17. For personal use only. stage of a taxon cycle in which expansion is followed by range contraction (Wilson 1961,

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org Ricklefs 1970, Ricklefs & Cox 1972)? Will phylogeographic studies provide repeated examples of directional, stepping-stone colonization? Or will we ﬁnd evidence for long-distance dispersal across archipelagos, skipping over whole islands or archipelagos, due to a lack of suitable habitat or resistance from competitors? Population studies may be a powerful tool to answer such questions. Viewing biogeographic patterns across the Wallacean archipelagos, Carstensen et al. (2013) related patterns to stages of taxon cycles, in which species are at ﬁrst widespread across a large number of adjacent islands, generally in coastal or open habitats, but later adapt to closed forest and become disjunctively distributed habitat specialists that in the end persist on single larger islands. Indeed, a large proportion of the birds of tropical archipelagos are single-island endemics, which may persist under stable climates and relatively slow biotic turnover (see also Cronk 1997 for relic island plants).

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SYNTHESIS: WHAT HAVE WE LEARNED FROM STUDIES OF INDO-PACIFIC CORVOID BIRDS? Contemporary distributions and phylogenetic relationships suggest that New Guinea has been a cradle for the diversification of the Corvides. In favor of the Papuan origin hypothesis are the number of ancient lineages and the high contemporary diversity present in New Guinea ( Jønsson et al. 2011, Aggerbeck et al. 2014, Kennedy et al. 2017). Opponents of the Papuan origin hypothesis note that corvoid passerine birds predate the existence of major proto-Papuan land areas, and suggest instead that extensive reduction of forests in Australia shrank corvoid ranges, leaving many of them endemic to New Guinea (Schodde & Christidis 2014). Weighing against this explanation, the Corvides group has no endemic lineages restricted to the mesic tropical forests of Australia, in contrast to several more basal songbird groups (Menurides, Climacterides, and Meliphagides), which are mainly Australian (in a broad range of habitats) but have many terminal subclades in New Guinea (Marki et al. 2017). These issues are currently unresolved. Critics of a Papuan origin of the Corvides have not specifically addressed the possible role of the terranes of the Sula Spur region as sites of early colonization and diversification. Furthermore, New Guinea later served as a center of origin, with subsequent dispersal to the surrounding archipelagos and to Asia. It therefore seems appropriate to consider New Guinea the source for corvoid passerine bird diversity.

Taxon Cycles: Reconciling Dispersal, Speciation, and Extinction With increasingly detailed phylogenetic data at the subspecies level, we have developed a better understanding of how island avifaunas are assembled through time. New Guinea is both a source and a sink for corvoid lineages. In this context, the idea of the taxon cycle provides a suitable framework for investigating how colonization of the Indo-Paciﬁc archipelagos might have taken place via multiple alternating expansions and contractions of species distributions (Wilson 1961, Greenslade 1968, Ricklefs 1970, Ricklefs & Cox 1972). The so-called stages of the taxon cycle are inferred from geographic distributions and taxonomic or genetic differentiation.

Stage I: expansion. During this stage, populations colonize islands or whole archipelagos from a source area. Some taxa exhibit behaviors (e.g., flocking or highly mobile foraging) that promote overwater dispersal. In some Indo-Pacific archipelagos, widespread species might occur at low densities (Reeve et al. 2015), although this is not a feature of expansion phases in the West Indies (Ricklefs & Cox 1972, Ricklefs & Bermingham 2008). Recent studies have shown that dispersal kernels can be highly dynamic, leading to increased dispersal on the expanding front and Access provided by Copenhagen University on 11/19/17. For personal use only. consequently recurrent pulses of dispersal events over the range edge, at least for continental taxa Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org (Phillips et al. 2008). Alternatively, individuals may be dispersed more passively during storms. This could lead to small birds being carried to remote islands they would not be able to reach via active flight. If dispersal occurs on a larger scale, it may lead to range expansion across large archipelagos, with colonization of numerous islands. In such dispersal events, colonization success will depend on factors such as island size, which relates to the size of a population that an island can sustain; food resources; the likelihood of an available, suitable habitat; and the absence or presence of competitors or predators.

Stage II: differentiation. Following range expansion through dispersal and colonization, populations on different islands begin to differentiate genetically and phenotypically. This may be due to genetic drift, selection, or both, and it might happen rapidly in populations established by a small number of founder individuals.

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Stage III: contraction. Eventually, populations go extinct, beginning with those on the smallest islands (Ricklefs & Bermingham 2008). Loss of landscape complexity due to gradual erosion and geological subsidence (Whittaker et al. 2008) might accelerate this process.

Stage IV: contraction/relictualization. The final stage before lineage extinction is the persistence of a single population (species), generally on one of the larger islands (Carstensen et al. 2012, 2013; Jønsson et al. 2014). New archipelagic range expansions may originate from such relict populations within archipelagos, which might even spread to larger islands or continental regions (Bellemain et al. 2008). Taxon cycles provide some predictions that can be evaluated with empirical data: (a) Young, expanding species inhabit multiple islands without gaps in their distributions; (b) old, relictual species inhabit single islands; (c) young, expanding taxa colonize islands of all sizes; (d ) old, relictual taxa occur only on large islands; and (e) old, relictual taxa are restricted to higher elevations (i.e., farther from the coasts), which presumably support the original habitats of colonists, than younger taxa are, particularly when they co-occur on an island. Recent publications have presented evidence, notably based on work on Pachycephalidae (whistlers) ( Jønsson et al. 2014) and Monarchidae (monarch flycatchers) (Fabre et al. 2014), supporting all five predictions. These studies have documented several expansion events at archipelagic scales, followed by relictualization, as well as intrageneric morphological and elevational segregation between coexisting young and old taxa from the same genus on an island. Similarly, analyses of all Indo-Pacific corvoid passerines reveal patterns that accord with predictions a–d, whereas the pattern of elevational segregation (prediction e)isless prominent, although highly distinct lineages are marginally lacking from the lowest elevations (Figure 6).

THE FUTURE Our knowledge about the processes responsible for the diversification and community assembly of corvoid passerine birds in the Indo-Pacific island world has increased tremendously over the past decade. Most notable is the overall phylogenetic framework and, for some select families, detailed information about the relationships among island populations. We can now test a diverse array of ecological and evolutionary hypotheses pertaining to adaptive radiation ( Jønsson et al. 2012a), taxon cycles ( Jønsson et al. 2014), the evolution of mimicry ( Jønsson et al. 2016a), dispersal (Kennedy et al. 2016), breeding strategies (Marki et al. 2015), and species diversification (Fritz et al. 2012, Kennedy et al. 2017). Access provided by Copenhagen University on 11/19/17. For personal use only. From these studies, it is clear that many processes are simultaneously at play, and that few tax-

Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org onomic groups follow the same evolutionary trajectory. Thus, no single model is likely to explain the current diversity and distribution of corvoid passerine birds in the Indo-Paciﬁc. New technology will likely provide further information about, and insights into, the assembly of insular faunas. With genome sequencing coming of age, it is now feasible to sequence thousands of genes across the genome for hundreds if not thousands of individuals. Applying such technology to population studies will provide information on effective population sizes, dispersal rates, and differentiation across dispersal barriers, which will enable a more informed evaluation of how populations move, differentiate, speciate, and adapt. Moreover, individual birds can now be tracked with satellite transmitters; although costly, this technology brings about the possibility of comparing actual movements with dispersal inferred from genomic data, which will ultimately allow us to assess directly the importance of dispersal in shaping distributions of birds and other organisms ( Jønsson et al. 2016c).

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Although determining the underlying mechanisms that govern distributions in space and time has been a major challenge, another challenge is understanding the relative contribution of each of these mechanisms to contemporary distributions of particular species. Rates of dispersal and differentiation of newly established populations may depend on the life-history strategies of a species. Diamond (1974) discussed changes in dispersal behavior by suggesting how dispersal sweeps were initiated by supertramps, leading to wide distributions across archipelagos (see also Mayr & Dia- mond 2001). This supertramp state was assumed to be followed, quite rapidly, by a shift to more sedentary behavior and genetic divergence of the different island populations. With the large phylogenies now available, we might begin to test the generality of the supertramp hypothesis and determine its association with life-history traits. Using the phylogenetic supermatrix for the Corvides, Marki et al. (2015) documented that dispersal and diversification rates for cooperative breeders were generally low (which appears to be an ancestral state among oscine passerine birds, including the Corvides). Apparently, this breeding system, with long-lasting family bonds, con- strains the tendency to disperse away from the natal environment. However, pair breeding, which evolved independently multiple times, leads to consistently higher rates of dispersal and speciation, suggesting that the tendency to disperse might in some way be associated with specific changes in breeding behavior that allowed flocking and concerted dispersal during the nonbreeding season. To fully understand the causes of high rates of dispersal and speciation in some lineages, other life-history traits should be evaluated. Potential factors could be tendencies to form nonbreeding flocks and to undertake opportunistic exploratory behavior, which may again be linked with flexibility in adapting to new conditions, feeding innovations, and general intelligence ( Jønsson et al. 2012b). However, morphology, including wing shape, does not appear to provide a clear indicator of highly dispersive species (Kennedy et al. 2016). Although long-distance dispersal may require a high aspect ratio of the wing to support rapid flight, this wing morphology is also associated with aerial foraging and might be lost rapidly following colonization of an island and evolution of a more sedentary lifestyle. Thus, combining functional and life-history analyses with genetic information pertinent to dispersal and population differentiation is a priority for understanding the dynamics of distribution and species formation in archipelagic lineages of birds.

DISCLOSURE STATEMENT The authors are not aware of any affiliations, memberships, funding, or financial holdings that might be perceived as affecting the objectivity of this review. Access provided by Copenhagen University on 11/19/17. For personal use only. Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org ACKNOWLEDGMENTS We are grateful to Robert Ricklefs, who provided insightful comments on the manuscript. We acknowledge funding from the Danish National Research Foundation to the Center for Macroe- cology, Evolution and Climate (DNRF96). K.A.J. further acknowledges the Carlsberg Foundation CF15–0078 and CF15–0079. M.K.B. is funded by a Marie-Sklodoska Curie Individual Fellow- ship (MSCA IF 2015 no. 707968). D.W.C. is funded by the Danish Council for Independent Research DFF-4181–00309. The authors would like to express their gratitude to numerous mu- seums worldwide and their staff, who have generously provided access to their material for genetic and morphological work over the years. Likewise, K.A.J., J.D.K., P.Z.M., and J.F. are indebted to all field collaborators in the Solomon Islands, Mauritius, Indonesia, and Papua New Guinea, without whom this work would not have been possible.

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Ecological Responses to Habitat Fragmentation Per Se Lenore Fahrig pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp1 Ecological Networks Across Environmental Gradients Jason M. Tylianakis and Rebecca J. Morris ppppppppppppppppppppppppppppppppppppppppppppppppp25 Impacts of Artificial Light at Night on Biological Timings Kevin J. Gaston, Thomas W. Davies, Sophie L. Nedelec, and Lauren A. Holt ppppppppppp49 The Utility of Single Nucleotide Polymorphism (SNP) Data in Phylogenetics Adam D. Leachéand Jamie R. Oaks pppppppppppppppppppppppppppppppppppppppppppppppppppppppp69 The Role of Sexual Selection in Local Adaptation and Speciation Maria R. Servedio and Janette W. Boughman ppppppppppppppppppppppppppppppppppppppppppppp85 The Potential Impacts of Climate Change on Biodiversity in Flowing Freshwater Systems Jason H. Knouft and Darren L. Ficklin ppppppppppppppppppppppppppppppppppppppppppppppppppp111 The Ecology of Mating and Its Evolutionary Consequences in Seed Plants Spencer C.H. Barrett and Lawrence D. Harder pppppppppppppppppppppppppppppppppppppppppp135 Process-Based Models of Phenology for Plants and Animals Isabelle Chuine and Jacques Régnière pppppppppppppppppppppppppppppppppppppppppppppppppppppp159 Evolution of Ecological Niche Breadth

Access provided by Copenhagen University on 11/19/17. For personal use only. Jason P. Sexton, Jorge Montiel, Jackie E. Shay, Molly R. Stephens, and Rachel A. Slatyer pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp183 Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org Analysis of Population Genomic Data from Hybrid Zones Zachariah Gompert, Elizabeth G. Mandeville, and C. Alex Buerkle ppppppppppppppppppp207 Biogeography and Biotic Assembly of Indo-Paciﬁc Corvoid Passerine Birds Knud Andreas Jønsson, Michael Krabbe Borregaard, Daniel Wisbech Carstensen, Louis A. Hansen, Jonathan D. Kennedy, Antonin Machac, Petter Zahl Marki, Jon Fjeldsa˚, and Carsten Rahbek ppppppppppppppppppppppppppppppppp231 Attached Algae: The Cryptic Base of Inverted Trophic Pyramids in Freshwaters Yvonne Vadeboncoeur and Mary E. Power pppppppppppppppppppppppppppppppppppppppppppppppp255

Contents v ES48-FrontMatter ARI 28 September 2017 13:24

Temporal Variation in Trophic Cascades Jonah Piovia-Scott, Louie H. Yang, and Amber N. Wright ppppppppppppppppppppppppppppp281 Anthropogenic Extinction Dominates Holocene Declines of West Indian Mammals Siobhán B. Cooke, Liliana M. Dávalos,Alexis M. Mychajliw, Samuel T. Turvey, and Nathan S. Upham pppppppppppppppppppppppppppppppppppppppppppp301 Spatially Explicit Metrics of Species Diversity, Functional Diversity, and Phylogenetic Diversity: Insights into Plant Community Assembly Processes Thorsten Wiegand, Mar´ıa Uriarte, Nathan J.B. Kraft, Guochun Shen, Xugao Wang, and Fangliang He ppppppppppppppppppppppppppppppppppppppppppppppppppppppp329 Pollinator Diversity: Distribution, Ecological Function, and Conservation Jeff Ollerton ppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp353 Evolution of Animal Neural Systems Benjamin J. Liebeskind, Hans A. Hofmann, David M. Hillis, and Harold H. Zakon pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp377 Variability in Fitness Effects Can Preclude Selection of the Fittest Christopher J. Graves and Daniel M. Weinreich ppppppppppppppppppppppppppppppppppppppppp399 The Ecology of Soil Carbon: Pools, Vulnerabilities, and Biotic and Abiotic Controls Robert B. Jackson, Kate Lajtha, Susan E. Crow, Gustaf Hugelius, Marc G. Kramer, and Gervasio Piñeiro pppppppppppppppppppppppppppppppppppppppppppppppp419 Apparent Competition Robert D. Holt and Michael B. Bonsall pppppppppppppppppppppppppppppppppppppppppppppppppppp447 Marine Infectious Disease Ecology Kevin D. Lafferty ppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp473 Ecosystem Processes and Biogeochemical Cycles in Secondary Tropical Forest Succession Access provided by Copenhagen University on 11/19/17. For personal use only. Jennifer S. Powers and Erika Mar´ın-Spiotta ppppppppppppppppppppppppppppppppppppppppppppp497 Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org Interactions Among Invasive Plants: Lessons from Hawai‘i Carla M. D’Antonio, Rebecca Ostertag, Susan Cordell, and Stephanie Yelenik pppppppp521 Phylogenetics of Allopolyploids Bengt Oxelman, Anne Krag Brysting, Graham R. Jones, Thomas Marcussen, Christoph Oberprieler, and Bernard E. Pfeil pppppppppppppppppppp543 Identifying Causes of Patterns in Ecological Networks: Opportunities and Limitations Carsten F. Dormann, Jochen Fründ,and H. Martin Schaefer pppppppppppppppppppppppppp559

vi Contents ES48-FrontMatter ARI 28 September 2017 13:24

Innate Receiver Bias: Its Role in the Ecology and Evolution of Plant–Animal Interactions Florian P. Schiestl pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp585 Evolutionary Rescue Graham Bell pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp605

Indexes

Cumulative Index of Contributing Authors, Volumes 44–48 ppppppppppppppppppppppppppp629 Cumulative Index of Article Titles, Volumes 44–48 ppppppppppppppppppppppppppppppppppppp633

Errata

An online log of corrections to Annual Review of Ecology, Evolution, and Systematics articles may be found at http://www.annualreviews.org/errata/ecolsys Access provided by Copenhagen University on 11/19/17. For personal use only. Annu. Rev. Ecol. Evol. Syst. 2017.48:231-253. Downloaded from www.annualreviews.org

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