Avian species richness in the Indomalayan realm -- Species richness map overlaid by the breeding ranges of 2406 bird species in east Asia. Bird species richness was highest at the base of the Indochina Peninsula and lowest in northern Siberia, the Gobi and Taklimakan deserts and the Tibetan Plateau. Islands had lower bird species richness than adjacent areas in mainland Asia (Ding et al. 2006).

Ding et al. (2006) suggested that primary productivity is the key factor underpinning patterns of bird species richness in east Asia. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.

Map showing Indomalaya avian species richness for native resident species (n = 1544).

Oong et al. (2026) examined the biogeographical patterns of birds within Indomalaya. Their analyses revealed four major biogeographic regions in Indomalaya that were consistent with present knowledge about the geological history of the region. (1) Indian Subcontinent. Analysis consistently indicated that the Indian subcontinent is a distinct biogeographic region in Indomalaya, with evidence of local areas of endemism. For example, Sri Lanka and the Western Ghats have both been noted as regions of high endemism—34 of the 479 bird species on Sri Lanka are endemic to the island, and the Western Ghats has about 20 endemics. Analysis also revealed evidence of habitat-based subdivision in the Indian subcontinent. The central and western portion of the Indian peninsula is more arid and highly seasonal, whereas the southern and eastern portions around the Western and Eastern Ghats are wetter and relatively less seasonal. (2) Continental Southeast Asia (mainland Southeast Asia and southeastern China). Mainland Southeast Asia—including Vietnam, Laos, Cambodia, Thailand, and Myanmar- extends west into the Himalayas. At least 200 bird species have overlapping ranges in this region. Southeastern China includes a hilly coastal strip south of the Yangtze River basin; approximately 50 bird species have overlapping ranges here. (3) Sundaland (Thai-Malay Peninsula, the islands of Sumatra, Borneo, Java, and Palawan, plus several smaller offshore islands). This region is characterized by extensive tropical rainforests that may have developed by the early Miocene and become largely isolated from the rest of Indomalaya with more seasonal habitats to the north and oceans on all other sides. In addition, the region was later subject to repeated sea level fluctuations during the Plio-Pleistocene, altering land cover and connection among the major landmasses. Sundaland is characterised by a high level of endemism, with over 260 endemic land bird species. (4) Oceanic Philippines. The oceanic Philippines is relatively young and did not come into existence until the Eocene at 35 Ma, while its present-day form did not emerge until around 5 Ma (Hall 2012, 2013; Yumul et al. 2003, 2008, 2009). With its history of geological isolation and position at the crossroads between Australasia and the rest of Indomalaya, the Philippines is faunally distinct, with a high level of both species richness and endemism. About 250 species of birds are either endemic or near endemic to the Philippines.

Some Indomalayan groups, e.g., barbets, range as far east as Bali and no further, whereas the range of several Australian birds (e.g., honey-eaters and cockatoos) range as far west as Lombok (35 km from Bali), but not further. This area of rather abrupt change in faunas was first noticed by Alfred Russel Wallace in the mid-1800’s. He observed an expected decline in the proportion of Asian to Australian birds as he traveled west to east through Indonesia, Malaysia, and Brunei, but noted a more substantial shift in that proportion between the islands of Bali and Lombok (islands just 25 km apart) and between Borneo and Sulawesi (or Celebs). The location of the rather abrupt change in species distributions was subsequently called Wallace’s Line. Wallace concluded that the islands west of the line were once connected to each other and to the Asian mainland and the islands east of the line had been similarly connected, but the two areas had always been separate (Wallace 1859). More recent work suggests that the area between Wallace’s Line and Lydekker’s Line (the western boundary of a strictly Australian fauna), called Wallacea, is best considered an area of interchange between Asiatic and Australian floras and faunas, where organisms exhibit a high degree of endemism (Moss and Wilson 1998).

Map of southeast Asia showing the location of Borneo and Sulawesi and Wallace’s Line. East of Java, Wallace’s Line lies between the
islands of Bali and Lombok. Areas of continental shelves are shown in grey (Figure from Moss and Wilson 1998).

The Malay Archipelago, part of the Sundaland biodiversity hotspot, has exceptionally high bird diversity, hosting over 13% of global bird species. The region, including Sumatra, Java, and Borneo, features rich lowland rainforest assemblages with hundreds of species, high rates of endemism—especially in Borneo—and significant populations of hornbills, pittas, and broadbills.

Key Aspects of Bird Diversity in the Malay Archipelago:

• High Species Richness: The region is home to vast numbers of bird species, with Malaysian areas alone recording over 850 species.
• Borneo as a Biodiversity Hotspot: Borneo has the world's highest concentration of many species, including frogmouths, trogons, hornbills, barbets, pittas, broadbills, flowerpeckers, and spiderhunters. In addition, 17 hawk species can be found in Java.
• Endemism and Evolution: The area acts as a center for avian endemism, with many species diverging during the Miocene.
• Unique Habitats: Lowland dipterocarp forests support incredibly dense, diverse bird communities.
• Key Species: The archipelago is famous for harboring species like the Crestless Fireback, Storm's Stork, Wallace's Hawk Eagle, and various kingfishers (e.g., Brown-winged, Collared, and Black-capped).
• Biogeographical Divisions: The region is divided into areas with different levels of endemic bird populations, with Sumatra and the Malay Peninsula sharing the most species, whereas Java has a distinct avifauna due to drier, seasonal lowlands.
• Migratory and Resident Birds: The region hosts a mix of permanent, diverse residents and migratory species that populate mangroves and forests.

Phylogeny of all passerine birds (Order: Passeriformes; n = 1633 species) in the Indo-Australian Archipelago (Figure from: Prasetya 2024).

Iconic and Endemic Species of the Indo-Australian Archipelago:

• Maleo (Macrocephalon maleo): An endangered megapode found only on the island of Sulawesi.
• Javan Hawk-eagle (Spizaetus bartelsi): Indonesia's national bird and a highly threatened raptor endemic to Java.
• Rhinoceros Hornbill (Buceros rhinoceros): The national bird of Malaysia, found in the region's rainforests.
• Hylocitrea (Hylocitrea bonensis): A unique, monotypic bird family endemic to Sulawesi.
• Birds of Paradise: Many species, such as the Red Bird-of-Paradise and Wilson's Bird of Paradise, are found in eastern Indonesia.
• Bali Starling (Leucopsar rothschildi): A critically endangered bird found in Bali.
• Bornean Ground Cuckoo (Carpococcyx radiceus): A rare species endemic to Borneo.

Key Avian Families and Groups:

• Megapodes (Megapodiidae): Including the Maleo, Philippine Megapode, and Dusky Megapode.
• Passerines: The most diverse group in the region.
• Birds of Prey: Various species, including the White-bellied Sea Eagle (largest in Indonesia), Crested Serpent Eagle (Spilornis rufipectus), and various Hawk-eagles.
• Pigeons and Doves: Including the Nicobar Pigeon, Green Imperial Pigeon, and numerous Fruit Doves.
• Nightjars: Such as the Malaysian Eared-nightjar (Lyncornis temminckii) and Great Eared-nightjar (Lyncornis macrotis).
• Partridges and Pheasants: Including the Great Argus and various Peacock-Pheasants.

• Australasian

• Australia, New Zealand, Papua New Guinea, eastern Indonesia & several small islands in the area

• generally a dry region, only 4.5% rainforest

• about 70% of Australia is desert & in this area only 17 species of birds occur (3% of Australia's avifauna)

• ~ 1600 species in the Australian region (907 in Australia alone, including 642 breeding birds, 162 vagrants, 77 non-breeding birds, and 26 introduced by humans)

• ~64 families of terrestrial & freshwater birds represented, with several unique:

• Cassowaries (Casuariidae) (Learn more about Cassowaries: Part 1, Part 2, Part 3, Part 4).

• Emus (Drommiceidae)

• Kiwis (Apterygidae)

• (Megapodiidae)

• Owlet-nightjars (Aegothelidae)

• New Zealand wrens (Acanthisittidae)

• Lyrebirds (Menuridae)

• Scrub-birds (Atrichornithidae)

• Birds of Paradise (Paradisaeidae)

• Bowerbirds (Ptilonorhynchidae)

• Honeyeaters (Meliphagidae)

• Corcoracidae (two species, the White-winged Chough and Apostlebird)

Spatial patterns of avian species richness in Australia across three levels of spatial resolution, from 0.25° grid cells to 1° grid cells (Hurlbert and Jetz 2007). The contemporary richness pattern is well explained statistically by actual evapotranspiration (a measure of water–energy balance), operating both directly and indirectly through plant production (Hawkins et al. 2005).

Native bird species richness in Australia at about 50 km x 50 km resolution. Australia has about 830–850
bird species, with approximately 45% being endemic, representing nearly one-tenth of the world's total.
Richness is heavily concentrated in the tropical northeast (Queensland) and regions with high, consistent
moisture (high actual evapotranspiration). In addition, structurally complex habitats, such as those with
diverse foliage profiles or larger tree diameters, are strong predictors of higher bird diversity. Within Australia,
Queensland (Q) has the highest bird diversity, with over 630 species recorded—roughly 60 more than New South
Wales (NSW), the next most species-rich state Key groups include parrots, honeyeaters, and pigeons. WA. Western
Australia; NT, Northern Territory; SA, South Australia; V, Victoria; T, Tasmania (Figure from: McKinney and Kark 2017).

Australia's Eastern Whipbird

Archipelago and island bird colonization times. Circles show the number of species (both extinct and extant) found in each archipelago (at the time of human arrival).
Numbers on the map correspond to numbers to the left of the archipelago name. Numbers to the right of the archipelago name indicate the number of species on
the archipelago. Colonization times plot: grey horizontal lines indicate archipelago ages. Violin plots (blue) show the distribution of times of colonization of bird species
in each archipelago. Thick black lines inside violin plots indicate the interquartile distance; thin black lines indicate the 95% confidence interval; black dots indicate the median.
Archipelagos with no violin plot or dots are cases where no species were present at the time of human arrival. Birds from left to right: Seychelles Sunbird, Seychelles Magpie Robin,
Silvereye, Príncipe Thrush, Laurel Pigeon, Dodo (extinct), Mauritius Fody, Red-moustached Fruit Dove (extinct), Galápagos Warbler, and Norfolk Kaka (extinct).

Valente et al. (2020) compiled a dataset of birds on islands, based on the terrestrial avifaunas of 41 oceanic archipelagos worldwide (including 596 avian taxa), and applied a new analysis method to estimate the sensitivity of island-specific rates of colonization, speciation and extinction to island features (area and isolation). Their model predicts—with high explanatory power—several global relationships, including a decline in colonization with isolation, a decline in extinction with area, and an increase in speciation with area and isolation, i.e., rates of speciation were found to increase through an interactive effect of both island size and distance from the nearest mainland, providing a mechanism that limits radiations to archipelagos that are both large and remote.

Oceanic Islands

• typically support fewer species of land & freshwater birds than equal areas of comparative mainland

• obvious influences on bird diversity on islands are age, topography, size, & distance from nearest source of immigrants

• importance of size & distance was first quantified by MacArthur & Wilson (1967)

• For example, New Zealand (Baker 1991):

• only 65 native species of land & freshwater birds (with an additional 47 species known from fossil remains and 34 species introduced by humans during the past 130 years)

• 2 endemic orders (Apterygiformes or kiwis & Dinornithiformes or moas [extinct]), 3 endemic families (Xenicidae or New Zealand wrens, Callaeatidae or Kokako, huia, & saddleback & Turnagridae or New Zealand thrushes [probably extinct]), and 37 endemic species (57% of native species; 70% if recently-extinct & fossil species are included) 

• avifauna composed largely of elements that dispersed to New Zealand long after the breakup of Gondwanaland

• Australian element, not surprisingly, is dominant - prevailing westerly winds regularly deposit Australian species in New Zealand (e.g., 8 species have successfully colonized New Zealand in the past 150 years)

• other biogeographical elements include Holarctic (just 3 species) & Archaic (endemic taxa with such a long evolutionary history in New Zealand that no close relatives can be identified elsewhere, e.g., kiwis and moas)

• routes of colonization:

• Tasman Sea has been the principle route

• direct route from the Holarctic (e.g., shorebirds & waterfowl)

• Other islands also have (or had) distinct avifaunas, e.g., see Hawaii's Honeycreepers'

 

New Zealand's Kiwis  

The avifauna of the islands of Wallacea includes elements of both Indomalayan and Australian origin (Michaux 2010). Of 513 species of breeding land birds in Wallacea, 300 (58%) are endemics and 213 (42%) are non-endemics that also breed outside of Wallacea. As also found in other island biogeography studies, island geography, especially island area, but also isolation and elevation, largely explain the variation in island species richness and endemism in Wallacea (Dalsgaard et al. 2014).

(A) Total species richness on islands in Wallacea. (B) Species richness of regional endemics (only found on islands in Wallacea) in Wallacea.
SUL, Sulawesi (Figure modified a bit from Dalsgaard et al. 2014).

The avifauna of the islands in the tropical Indian Ocean (excluding Madagascar) consists of about 139 species of resident birds (excluding pelagic and introduced species) representing 33 different families of birds. Of these species, 10 species are endemic to the Indian Ocean and 69 are endemic to a single archipelago or island. Most endemic species are in the families Columbidae (pigeons), Psittacidae (parrots), Muscicapidae (thrushes and Old World flycatchers), and Zostertopidae (white-eyes) (Adler 1994). Unfortunately, several of the 139 species of resident birds are now extinct.

The presence of relatively few endemic species of birds on the islands of the tropical Indian Ocean is due to the almost complete absence of any intra-archipelago speciation, likely because the Indian Ocean archipelagos have too few of the large islands needed for such speciation. In addition, some families of birds that are widespread in the Afrotropical and Indomalayan realms, such as Alcedinidae (kingfishers) and Sturnidae (starlings), are either absent or poorly represented on islands in the tropical Indian Ocean. For some families, e.g., kingfishers, this absence may be due the absence of suitable habitat (rivers, streams, or lakes). For other families, reasons for their absence is less clear (Adler 1994).

Avifaunas of islands in the Indian Ocean. The first number below each island’s and group of island’s name is the
number of species of resident breeding birds; the second number is the number of endemic species (Figure modified from Adler 1994).

The islands of the Pacific originated in a variety of ways, including as linear chains of volcanic islands, fragments of continental crust, and obducted portions of adjoining tectonic plates (Neall and Trewick 2008).

Avifaunas of islands in the Pacific Ocean. Numbers below the names of islands indicate, in order, the number of
breeding land birds, number of endemic species, and total number of species including seabirds (Figure modified from Steadman 1995).

The Hawaiian archipelago consists of 45 islands extending across 2450 km of the central Pacific Ocean and is the most remote archipelago on the planet (Juvik and Austring 1979). The Hawaiian Islands once had at least 111 native species of birds, almost all endemic, before humans (Polynesians) arrived about 1000 years ago. Of those 111 species, 55 became extinct after arrival of the Polynesians and are now known only as fossils, and 19 more extinctions occurred after arrival of Europeans (Scott et al. 2001). Of the remaining 37 extant endemic species, 33 are at some risk of extinction. In addition to these endemic species, at least 22 species of seabirds breed on the Hawaiian Islands, and another 54 species of landbirds have been introduced by humans and now breed in Hawaii.

All of Hawaii’s endemic species of birds are thought to have originated from just 20 colonization events, and the many species of honeycreepers (originally more than 50, just 17 remain) are thought to be derived from a single ancestor (Berger 1981, Olson and James 1982). Based on analyses of mitochondrial genomes, Lerner et al. (2011) suggested that Hawaiian honeycreepers are a sister taxon to Eurasian rosefinches (Carpodacus) and that the ancestor of the honeycreepers arrived in Hawaii about 6-7 million years ago. In addition to the honeycreepers, the endemic Hawaiian avifauna includes (or included) several waterfowl, two raptors, a crow (extinct in the wild, but some still live in captivity), and representatives of Old World flycatchers, honeyeaters, and thrushes (Jacobi and Atkinson 1995). The ancestors of these and other endemic species arrived in the Hawaiian Islands beginning about 5 million years ago.

Estimated arrival times (millions of years ago, Mya) of the ancestor of several species and taxa of birds in the Hawaiian Islands
(Price and Clague 2002; except Hawaiian honeycreepers from Lerner et al. 2011).

The islands in the Atlantic Ocean off the coasts of southern Europe and northern Africa are collectively referred to as Macaronesia. This region has 31 principal volcanic islands that are grouped into five archipelagos: Azores, Madeira, Selvagens, Canary Islands, and Cape Verde (Figure below). These archipelagos vary in their distance from the mainland (range = <100 km for the Canary Islands and 1365 km for the Azores), latitude, and age (range = 0.25 – 29 million years) (Hazevoet 1995, Geldmacher et al. 2001, Azevedo and Ferreira 2006). The taxonomic affinities of native bird species suggest that the origin of the extant terrestrial avifauna is largely from the western Palaearctic mainland, but the Cape Verde archipelago, because of its lower latitude and proximity to Africa, has species related to mainland Africa (Hazevoet 1995, Martín and Lorenzo 2001, Clarke et al. 2006). In addition, the avifaunas of the four largest archipelagos (excluding Selvagens) have all reached an equilibrium, with species richness reaching a plateau because rates of colonization and speciation are similar to extinction rates (Valente et al. 2017).

(A) Colonization times of Macaronesian species of landbirds and a map of Macaronesia. The vertical lines to the left indicate the maximum geological ages of the archipelagos.
Filled circles, non-endemic species; unfilled circles, endemic species; unfilled squares, Macaronesian endemic. The Selvagens Islands, although important breeding areas for several species of seabirds,
have only two resident landbirds: Berthelot’s Pipit (Anthus bertheloti) and Common Kestrel (Falco tinnunculus). (B) Estimated n umbers of species over time in each of the archipelagos.
Black lines are estimated median values and the colored areas show the 2.5th–97.5th percentiles. Gray dashed line, pre-human diversity; black dashed line, present diversity (excluding extinct species).
Time before present is in millions of years (Figures from Valente et al. 2017).

The West Indies consist of three groups of islands: (1) the Greater Antilles, including Cuba, Hispaniola, and Puerto Rico, are old fragments of continental crust that originally formed as islands closer to present-day Central America and carried to their present locations by plate movements, (2) the Bahama islands are low-elevation islands located north of the Greater Antilles and southeast of Florida, and (3) the Lesser Antilles were created by a series of volcanoes, some still active, where the South American Plate is being subducted under the Caribbean plate. Bird species richness of the West Indies is low compared to the nearby continents, likely because of the limited dispersal abilities of many avian taxa (Ricklefs and Bermingham 2001). Water barriers reduce the likelihood of colonization by birds that rarely move beyond forest borders like most tropical suboscine passerines and woodpeckers (Terborgh 1973, Terborgh et al. 1978).

 
Birds of Cuba

Of the 243 breeding species in West Indies, 176 (72%) are endemics and 67 (28%) non-endemics (Dalsgaard et al. 2014). The main pathways of immigration into the islands have been from the south (the northern coast of South America and through the Lesser Antilles), from the west (Central America and into the Greater Antilles), and from the north (Florida). Although South America contains a larger species pool of potential immigrants and the Lesser Antilles are close to the coast of South America, more species of birds in the West Indies originated in North and Central America (Bond 1963). However, species that have invaded the West Indies from South America have tended to move further north than species that emigrated from North and Central America. As is typically the case, the larger islands in the West Indies have more landbird species and more endemic species of birds.

A) Total species richness on islands in the West Indies. (B) Species richness of regional endemics, i.e., only found on islands
in the West Indies. FL, Florida; PR, Puerto Rico; SA, South America (Figure modified from Dalsgaard et al. 2014).

Seminar - Challenges and opportunities for Caribbean endemic bird conservation

The islands of the Southern Ocean can be divided into older islands with a continental origin (e.g ., Falkland Islands, South Georgia, Auckland and Campbell Islands), and younger, volcanic islands (e.g., most islands in the southern Indian Ocean). Islands in the Southern Ocean have cool, wet, and windy climates. The islands are covered in tundra vegetation, ranging from scattered dwarf plants to grasslands, herbfields, and bogs. Vegetation in coastal areas includes tussock grasses, graminoids, and herbs; more upland areas are often covered with cushion-forming angiosperms and mosses. Vascular plant species richness ranges from zero (Bounty Islands) to 188 (Auckland Islands), and insect species richness varies from six (McDonald Island) to more than 200 (Auckland Islands) (Bergstrom and Chown 1999).

With the exception of the Falkland and Chatham islands, most islands in the Southern Ocean have few or no breeding landbirds. In addition, the Southern Ocean islands have few endemic species. The few breeding songs generally show affinities to the nearest mainland areas. In contrast to the few breeding landbirds, the Southern Ocean islands are the primary nesting areas for numerous species of seabirds. The productivity of the southern oceans provides abundant food for breeding seabirds.

Islands of the Southern Ocean. Numbers below the names of the islands indicate the number of breeding landbirds, number of endemic species,
and total number of breeding birds. Most birds breeding on these island are penguins, albatrosses, petrels and prions,
shearwaters, storm-petrels, skuas, and gulls and terns (Numbers from Shirihai 2007; Figure modified from Angel et al. 2009).

(A) An artist's impression of H. moorei attacking the extinct New Zealand moa. Evidence of eagle strikes are preserved on skeletons of moa weighing up to 200 kg. (Artwork: John Megahan.) (B) Comparison of the huge claws of H. moorei with those of its close relative  Hieraaetus morphnoides, the “little” eagle. The massive claws of H. moorei could pierce and crush bone up to 6 mm thick under 50 mm of skin and flesh.

Ancient DNA Tells Story of Giant Eagle Evolution -- For reasons not entirely clear, when animals make their way to isolated islands, they tend to evolve relatively quickly toward an outsized or pint-sized version of their mainland counterpart. One avian giant was found in New Zealand:  Haast's eagle (Harpagornis moorei), with a wingspan up to 3 meters. Though Haast's Eagle could fly,  its body mass (10–14 kilograms) pushed the limits for self-propelled flight. As extreme evolutionary examples, large island birds can offer insights into the forces and events shaping evolutionary change. Bunce et al. (2005) compared ancient mitochondrial DNA extracted from Haast's Eagle bones and found that the extinct raptor underwent a rapid evolutionary transformation that belies its kinship to some of the world's smallest eagle species. Their analysis places Haast's Eagle in the same evolutionary lineage as a group of small eagles in the genus Hieraaetus. Surprisingly, the genetic distance separating the giant eagle and its smaller cousins from their last common ancestor is relatively small.       Using  molecular dating techniques, Bunce et al. (2005) proposed a divergence date of  0.7–1.8 million years ago. The increase in body size—by at least an order of magnitude in less than 2 million years—is remarkable, Bunce et al. (2005) argue, because it occurred in a species still capable of flight. The absence of mammalian competitors may have precipitated the rapid morphological shift. Haast's eagle, the authors wrote, “represents an extreme example of how freedom from competition on island ecosystems can rapidly influence morphological adaptation and speciation.” -- Source: (2005) Ancient DNA Tells Story of Giant Eagle Evolution. PLoS Biol 3(1): e20.

Haast's Eagle - The King of All Eagles

Avifaunas of the various zoogeographic realms represent a mixture of species of various ages & origins. The mobility of many (but not all) birds, plus their proclivity to migrate, has allowed the movement of some groups between realms - some very long ago, others more recently. And, once a species arrives in a new area, adaptive radiation often generates new species. This combination of avian mobility (or lack thereof) and adaptive radiation has created avifaunas that represent complex mosaics (Gill 1995). Briefly, the avifaunas of the major zoogeographic realms can be described as follows:

• Palearctic

• many genera shared with the Nearctic, Ethiopian, & Oriental regions indicating much dispersal

• geographic 'barriers' between these regions include:

• the Sahara desert (Ethiopian)

• the Himalayas (Oriental) (viewed from space in photo to the right)

• the Bering Sea (Nearctic)

• relatively low species richness, probably due to cold climate & large areas of homogeneous habitat (taiga)

 

Map of species richness among equal area grid cells in the full data set (3,036 taxa) of Palearctic songbirds.
Maximum richness shown in deep rufous and minimum in dark blue. Scores grouped into 32 color-scale classes of
approximately equal frequency. Classes range from 1–3 species per grid cell (blue) to 128–307 (rufous); also a single grid cell
with maximum score (316 species, in northeast Myanmar [Burma]) in red (Roselaar et al. 2007). Most 'hotspot' regions are located
in mountainous regions.

Geographic variation in the richness of bird species in the former USSR. The hatched area in the upper left represents an area classified as the western Palearctic (Hawkins et al. 2003).

Examples of songbirds found in the Palearctic realm. (A) Long-billed Pipit, Anthus similis (Photo by Ian Boustead), (B) Indian Blue Robin, Luscinia brunnea
(Photo by Niranjam Sant), (C) White-throated Redstart, Phoenicurus schisticeps, (D) White-capped Water Redstart, Chaimarrornis leucocephalus,
(E) Black-browed Tit, Aegithalos bonvaloti, (F) Chestnut Thrush, Turdus rubrocanus, (G) Temminck’s Lark, Eremophila bilopha (Photo by Augusto Faustino),
(H) Plain-backed Thrush, Zoothera mollissima (Photos C, D, E, and F by John and Jemi Holmes) (Figure made from photos from Roselaar 2006).

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The Western Palearctic Avifauna (Blondel and Mourer-Chauviré 1998) -- Compared with that of the two other large forested regions of the Northern Hemisphere (eastern North America and eastern Asia), the bird fauna (and particularly forest avifauna) of the western Palearctic is relatively species poor. Controlling as much as possible for the size of areas, Mönkkönen and Viro (1997) have shown that the western Palaearctic (including North Africa) has 50% fewer forest-associated bird species than eastern Asia and 40% fewer than eastern North America. Europe is similar in size to China (9 500 000 km², including Manchuria), but European species richness (about 500 bird spp.) is only about half that reported for China. In the western Palaearctic, less than half of the terrestrial avifauna is associated with forest compared with two-thirds in eastern North America and eastern Asia.

Large and repeated habitat changes during the Pleistocene have undoubtedly accelerated the extinction rates of birds of the western Palearctic, just as they did for plants and large mammals in North America. Comparative analyses using molecular phylogenies of 11 lineages of North American songbirds also indicate a general decrease in rates of species diversification during the Pleistocene. However, molecular evolutionary trees cannot distinguish between reduced speciation rates (e.g. as a consequence of progressive niche filling through evolutionary time) and higher extinction rates as the cause of net changes in diversification rates.

Assuming similar rates of speciation over the whole Northern Hemisphere during the Pleistocene, the relative impoverishment of the western Palearctic bird fauna (compared with the two other regions of the Northern Hemisphere) could be caused by higher extinction rates in the western Palearctic, because there was no connection with the tropical biomes further south. In contrast to those of the western Palearctic, the extant bird faunas of both northeastern North America and eastern Asia share many species in common with southern tropical regions. For example, 76% of passerine species breeding in North America have either conspecific populations or congeneric species that breed in the Neotropics. In eastern Asia, two orders and nine families are tropical, whereas Afro-tropical influence in the fauna of the western Palaearctic is negligible.

The most likely explanation for these differences is that the biota of both eastern North America and eastern Asia remained connected to the tropics over the whole Tertiary–Quaternary, making it possible for a continuous interchange between tropical and temperate avifaunas. In contrast, the massive east–west oriented barriers (mountain ranges, seas and large desert belts) ‘trapped' biota of the western Palaearctic in their southern refugia during glacial periods, preventing them from finding refuge in tropical regions further south, and preventing tropical species from colonizing northern regions. Since the Pliocene, the Sahara desert has never been sufficiently forested to provide a dispersal corridor for forest birds between the Afro-tropics and Eurasia.

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Nearctic

Species richness, compared to most other realms, is relatively low. Contributing factors include:

birds entering from Siberia must pass through areas of tundra & cold coniferous forest (with no access at all during periods of glaciation)

tropic-adapted species from the south are faced with an adverse climatic gradient. For example, few South American birds have colonized the deciduous forests of the eastern United States (one hummingbird - the Ruby-throated Hummingbird - & two tanagers - Summer and Scarlet tanagers).

low species richness in North American deciduous forests because:

they formed a continuous block (restricting opportunities for speciation) and they have been compressed during glacial maxima

Geographic variation in the richness of breeding terrestrial bird species in the Nearctic and northern Neotropics. The heavy dashed lines in southern Mexico distinguish cells classified as Nearctic and cells classified as Neotropical. Numbers represent the number of bird species in each cell (Hawkins et al. 2003).

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Deciduous forest may have barely existed in the southern USA at the Last Glacial Maximum (LGM). Contrary to most of the earlier interpretations, it now seems inappropriate to show any significant area of deciduous forest in the eastern US for the LGM. Current work by S. Jackson (Northern Arizona University), Eric Grimm (Illinois State Museum), and Bill Watts suggests that even in the south there was very little broad-leaved forest at the LGM, with perhaps only isolated pockets of deciduous forest that were surrounded by coniferous vegetation. These deciduous forest pockets at the LGM may have been confined to moister microsites, such as stream gullies (Ritchie 1982). For example, spruce forest has been found to have been growing in Louisiana at the LGM. Tallis (1990) also maps a south-eastern pine forest that extending from about 33°N down to the Gulf coast and northern Florida, and suggests that it would generally have been fairly open in structure. Source: http://www.soton.ac.uk/~tjms/namerica.html

Eastern U.S. - 18,000 years ago Source: http://www.esd.ornl.gov/projects/qen/NAL2215.gif & http://www.esd.ornl.gov/projects/qen/nercNORTHAMERICA.html

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• avifauna appears to be composed of six elements (Mayr 1946):

• 'Cosmopolitan' - 'mobile' groups with world-wide distributions, including hawks (Accipitridae), swifts (Apodidae), woodpeckers (Picidae), & nightjars (Caprimulgidae)

• 'Panboreal' (or Holarctic) - groups found throughout the Palearctic-Nearctic, including loons (Gaviidae) & some shorebirds

• 'Pan-American' (or Neotropical) - groups that originated in South America, including ,tyrant flycatchers (Tyrannidae), and tanagers (Thraupinae)

• 'Pantropical' - groups with undetermined origins in the southern continents, including parrots (Psittacidae)

• 'Old World'  (or Palearctic):

• early colonizers, including cranes, pigeons, cuckoos, owls, crows, & some thrushes

• later colonizers, including kingfishers, jays, titmice, & some cardueline finches

• recent colonizers, including the barn owl, some swallows, some thrushes, kinglets, & shrikes

• 'Nearctic' - groups that likely originated in North America, including grouse, turkey, wrens, waxwings, & warblers (Parulidae) 

• Pleistocene climatic cycles may have precipitated much speciation, particularly at mid- & high latitudes (Mengel 1964, Selander 1971). This paradigm was challenged based on a review of mtDNA sequence divergences (Klicka and Zink 1997). This review suggested that only 11 of 35 assayed pairs of sister species date to Quaternary separations. Subsequently, Avise and Walker (1998) suggested that divergence of many groups may have started prior to the Pleistocene, but divergence to the species level was promoted by the Pleistocene glaciation.

• a major effect of the southward extensions of the glaciers was to separate many wide-ranging forest/woodland species into eastern & western populations on either side of the prairie grassland belt. Examples of groups within which speciation may have occurred during this time are the:

• flickers (Colaptes spp.; Mayr 1942)

• juncos (Junco spp.; see 'Postglacial range expansion drives the rapid diversification of the genus Junco' & bluebirds (Sialia spp.) (Rand 1948)

• warblers (Setophaga & Vermivora spp.; Mengel 1964)

• mtDNA evidence suggests that other species were likely confined to single small populations during the Pleistocene &, following retreat of the Wisconsin glacier, expanded rapidly throughout much larger ranges, e.g, Downy Woodpecker, Common Grackle, Red-winged Blackbird, Black-capped Chickadee, & Song Sparrow (Avise 2000).

(A) Male Downy Woodpecker (Picoides pubescens) (photograph by Peter de Wit; Wikimedia Commons).
(B) A possible scenario for dynamic range shifts at high latitudes responding to glaciations by contracting (t2) and
then expanding (t1) during more favorable times. (C) Variation in historical population size for Downy Woodpeckers
through time (From Toews 2015).

Few endemic species of birds are found in the Nearctic and the Palearctic. One important reason for this is that climates in
the northern hemisphere have changed dramatically since the Last Glacial Maximum (LGM; 21,000 years ago). (A) Climate-change velocity
(rate of movement of climatic conditions over the Earth’s surface) since the LGM. Velocity is highest in northeastern
North America (Nearctic) and north-central Eurasia (Palearctic), in and near the areas that were glaciated (delimited by black lines).
(B) Proportion of bird species that are endemic throughout the world. Areas where climate-change velocities have been highest are also the
areas with the fewest endemic species, likely due to species extinctions as well as incomplete range filling. Areas with greater
proportions of endemic species are also the areas where climates have exhibited little or no changes since the LGM. Note also
that many of the areas with high proportions of endemic species are islands (Figures from Sandel et al. 2011).

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A, Summer pattern of species richness across North America on the basis of expected number of species over three randomly chosen Breeding Bird Survey routes per 20,000-km2 grid cell. B, Spatial pattern of normalized difference vegetation index (NDVI) in June at 1-km resolution. C, Winter pattern of species richness across North America on the basis of expected number of species over one randomly chosen Christmas Bird Count circle per 20,000-km2 grid cell. D, Spatial pattern of NDVI in December at 1-km resolution (From: Hurlbert and Haskell 2003).

Avian species richness in North America -- The geographic patterns of avian richness in North America vary markedly between seasons and
closely match the seasonal patterns of Normalized Difference Vegetation Index (or NDVI, which is a measure of greenness calculated from reflectance
in the near infrared and red portions of the electromagnetic spectrum). The NDVI has been shown to be correlated with total green biomass, leaf area
index, and percent incident photosynthetically active radiation. During the breeding season, richness is greatest across the eastern U.S.-Canadian border,
along the Appalachian Mountains, and in the northwestern United States, while richness is lowest throughout the Great Plains. During the winter, richness
is greatest at southern latitudes and along the coasts. Latitude alone explains very little of the variation in richness during the breeding season. These data
suggest that birds use their mobility to track spatially and temporally fluctuating resources, even at continental and hemispheric scales. The fact that the
relationship between richness and available energy is similar across seasons despite enormous changes in the geographic pattern of productivity suggests
that bird species distribute themselves in the same way throughout the year in response to resource abundance and the presence of other species without
regard to latitude. Thus, seasonal environmental production appears to determine the number of species that coexist in an area during a given season, and
the rate and timing of that production determine the migratory guild composition of the breeding community (Hurlbert and Haskell 2003).

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• Ethiopian (Afrotropical)

• strong affinities with the Oriental realm; many Palearctic species winter south of the Sahara desert

Source: http://www.rbge.org.uk/research/russcis/rnpsiberia.htm

• similar latitude to Neotropical realm, but fewer than half as many species:

• less tropical forest & generally much drier than the Neotropics

• few mountain ranges

• endemic families generally contain relatively few species, e.g.:

• 1 ostrich (Struthionidae)

• 1 hamerkop (Scopidae)

• 1 shoe-billed stork (Balaenicipitidae)

• 1 secretarybird (Sagittariidae)

• 6 mousebirds (Coliidae)

• 8 woodhoopoes

• 44 helmet shrikes (Prionopidae)

• 23 turacos (Musophagidae)

Species richness for 1599 species of birds endemic to Africa (Figure from: Colwell et al. 2009).
Species richness of endemics (species globally restricted to sub-Saharan Africa) peaks along the
mountains and adjacent lowlands of eastern and southern Africa. Species richness is also relatively
high in the isolated mountains in central and western Africa and the lowlands of the northeastern
Congo Basin (de Klerk et al. 2002).

Birds of Africa

 
Ancient African birds

Lake Nakuru's flamingoes
 

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Turacos are uniquely African, with 23 species recognized by most authorities. The forest-living species are undoubtedly among the most spectacularly colorful of birds, while the savanna-dwellers (known as "go-away birds" because of a well-known call, a harsh "kay-waaay") are predominantly grey in plumage. In South Africa, these birds are better known as louries. All are frugivores, specializing in fruit (particularly figs) and sometimes feeding on leaves, buds and flowers. They are usually gregarious, and frequently associate with parrots, hornbills and barbets, with individuals and groups often returning day after day to the same fruiting tree, until the crop is exhausted.

White-bellied Go Away Bird

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• Oriental (Indomalayan)

• closest affinities to tropical Africa, but linkages to Palearctic & Australian faunas as well. The Himalayas represent a boundary between Oriental & Palearctic regions; water separates the Oriental & Australian regions.

• only one endemic family (Irenidae), but another nearly so (Eurylaimidae); pheasants (Phasianidae) have undergone considerable radiation

• boundary between Oriental & Australasian realms is Wallace's Line; more recent work indicates that the 'line' runs roughly between Celebes & the Moluccas (~600 km east of Wallace's Line & known as Weber's Line

   

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Birds & the Beetles -- Batrachotoxins are neurotoxic steroidal alkaloids first isolated from a Colombian poison-dart frog. In 1992, the toxic principle from feathers and skin of birds in the genus Pitohui, endemic to New Guinea, was isolated and, remarkably, proved to be mainly a batrachotoxin (BTX). BTXs were later found in New Guinea toxic birds of the genus Ifrita. On a weight basis, the batrachotoxins are among the most toxic natural substances known, being 250-fold more toxic than strychnine. It is believed that such toxins provide some protection against the birds' natural enemies, such as parasites and predators, including humans. Neither poison-dart frogs or birds are thought to produce the toxins de novo, but instead they likely sequester them from dietary sources. Dumbacher et al. (2004) described the presence of high levels of batrachotoxins in a little-studied group of beetles, genus Choresine (family Melyridae). These small beetles and their high toxin concentrations suggest that they might provide a toxin source for the New Guinea birds. Stomach content analyses of Pitohui birds revealed Choresine beetles in the diet, as well as numerous other small beetles and arthropods. The family Melyridae is cosmopolitan, and relatives in Colombian rain forests of South America could be the source of the batrachotoxins found in the highly toxic Phyllobates frogs of that region.      The Blue-capped Ifrita (Ifrita kowaldi; shown to the right) is called Nanisani by local villagers. Nanisani is also the word used to describe the tingling and numbing sensation of the lips and face that results from contacting both beetles and bird feathers (Photo Source: Smithsonian National Zoological Park).

A Choresine pulchra beetle with an insert showing  glandular vesicles.

 
Jack Dumbacher and Hooded Piohuis

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• Australasian

• Long history of isolation from other land masses

• 16 families endemic (or nearly so); of 308 passerine species, 80% are 'old endemics' belonging to lineages that likely originated & radiated in Australia & New Guinea

• Some evidence that the parvorder Corvi began an evolutionary radiation in Australia about 55-60 million years ago (Sibley and Ahlquist 1985) & gave rise to:

• Superfamily Menuroidea

• Australian treecreepers (Climacteridae)

• lyrebirds (Menuridae)

• bowerbirds (Ptilonorhynchidae) (check this video about the Vogelkop Bowerbird)

• Superfamily Meliphagoidea

• fairywrens & grasswrens (Maluridae)

• honeyeaters & Australian chats (Meliphagidae)

• Thornbills and allies (Acanthizidae)

• Superfamily Corvoidea

• Australian robins (Petroicidae)

• logrunners (Orthonychidae)

• pseudo-babblers (Pomatostomidae)

• diverse set of other birds:

• crows & jays (Corvidae)

• birds of paradise (Paradisaeidae)

• wood-swallows, Australian magpies, butcherbirds, & currawongs (Artamidae)

• magpie larks (Grallinidae)

• bell magpies (Cracticidae)

• whistlers, shrike-tit, thick heads, & pitohuis (Pachycephalidae)

• This particular scenario is not universally accepted. However, it's apparent that many Australasian species evolved in Australia & nearby areas.

 
Bowerbirds - David Attenborough

Formation of New Guinean avian diversity across barriers and along elevational gradients by Knud Andreas Jonsson

Seminar - Poisonous birds and other stories of New Guinea fieldwork by Jack Dumbacher

 

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Songbirds Escaped From Australasia, Conquered Rest of World - That male cardinal singing his heart out in your backyard has ancestors that left the neighborhood of Australia 45 million years ago. A comprehensive study of DNA from songbirds and their relatives by Barker et al. (2004) shows that these birds, which account for almost half of all bird species, did not originate in Eurasia, as previously thought. Instead, their ancestors escaped from a relatively small area—Australasia (Australia, New Zealand and nearby islands) and New Guinea—about 45 million years ago and went on to populate every other continent save Antarctica.  The movement of the Australia/New Guinea plate between 60 (a) and 20 (b) million years ago (Mya). When New Zealand  separated from Antarctica ~85 Mya, only South America, Antarctica and Australia/New Guinea remained as the last pieces  of the Gondwanan supercontinent. Around 55 Mya, Australia/New Guinea broke away from Antarctica and moved north. Australia/New Guinea finally collided with southeast Asia ~15 Mya. Possible dispersal routes of birds are indicated by arrows  (From: Heinsohn and Double 2004).   The order Passeriformes, or perching birds, includes all songbirds, such as robins, cardinals, blackbirds, house sparrows, house finches and crows. The group is further divided into birds that must learn their songs “true songbirds” (oscines) and those with the innate ability to sing the “correct” song. True songbirds account for 4,580 of the 6,000 known Passeriformes species. The true songbirds are currently divided into two groups: Passerida (3,477 species, among them many familiar backyard species) and Corvida (1,103 species, including crows and ravens).  The two groups of true songbirds were thought to have separate origins, with Corvida originating in Australasia and the Passerida in Eurasia. The Passerida then supposedly spread from Eurasia to Africa, Australasia and the New World. But in examining the DNA sequences of two genes in all but one family (a closely related group, such as “crows and jays” or “warblers”) of passerine birds, Barker et al. (2004) made a startling discovery. “It was thought that the Passerida arose in Eurasia about 40 million years ago,” said Barker. “But we found that these birds fall into a group within the Corvida. That means all songbirds trace their origins to Australasia and New Guinea.” The Passerida differ from the Corvida because the Passerida somehow made it out of Australasia and New Guinea and onto the Asian mainland long before the Corvida, Barker said. Asia and Australasia are carried on separate plates in the Earth’s crust, and for many millions of years those plates were far apart. Around 45 million years ago, the ancestors of the Passerida dispersed to Asia—over more than 600 miles of open ocean--long before these two plates approached one another (as shown on map with locations of the continents 45 mybp). For some reason, however, ancestors of the Corvida didn’t make it until about 25 million years later, or 20 million years before today. At that time, Asia and Australia were much closer to each other, and island chains that could have allowed the Corvida ancestors to “island hop” to the mainland appeared, Barker said. “There are many endemic Corvida birds on the Indonesian island of Lombok but very few on Bali, the next island to the west,” said Barker. “And, sure enough, the line separating the Asian plate from the Australasian plate runs between Bali and Lombok.

 

 

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• Neotropical

• large number of endemics suggests an ancient & distinctive evolutionary history (Mayr 1964)

• some endemic families have undergone extensive evolutionary radiation & contain many species, e.g.:

• hummingbirds (Trochilidae) - 363 species (The secret life of hummingbirds)

• ovenbirds (Furnariidae) - 242 species

• antbirds, antshrikes, antwrens, antvireos, bushbirds, fire-eyes, & bare-eyes (Thamnophilidae) - 208 species

• cotingas, bellbirds, becards, cocks-of-the-rock (Cotingidae) - 79 species

• antthrushes (Formicariidae) & antpittas (Gralliidae)) - 56 species

• manakins (Pipridae) - 52 species

• woodcreepers (Dendrocolaptidae) - 52 species

• puffbirds, nunlets, and nunbirds (Bucconidae) - 38 species

 
Andean Cock-of-the-rock

 

 

Black-fronted Nunbird

Photo © Arthur Grosset

• tapaculos (Rhinocryptidae) - 30 species

• jacamars (Galbulidae) - 17 species

Rufous-tailed Jacamar

Photograph courtesy Dr. Russell Barrow

 

 

Common Potoo  

• other endemic families have undergone little radiation, e.g.:

• potoos (Nyctibiidae) - 6 species

• seedsnipes (Thinocoridae) - 4 species

• trumpeters (Psophiidae) - 3 species

• plantcutters (Cotingidae) - 3 species

• hoatzin (Opisthocomidae) - 1 species (shown to the right)

• oilbird (Steatornithidae) - 1 species

• in addition to these endemics, several groups are more recent arrivals from the Nearctic: vultures (Cathartidae), pigeons (Columbidae), jays (Corvidae), wrens (Troglodytidae), thrushes (Turdinae), blackbirds (Icterinae), & finches (Fringillidae)

• several other groups have pantropical distributions, including cuckoos (Cuculidae), parrots (Psittacidae), barbets (Capitonidae), & trogons (Trogonidae)

• Neotropical forests are home to nearly 700 genera of birds.

 

Birds of Peru

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Antarctic realm

The continent of Antarctica encompasses about 14.3 million km 2, with most covered by glacial ice. During the Antarctic summer, about 13,700 km 2 of rock and soil is exposed, particularly in the Antarctic Peninsula (Lee et al. 2017). Much of the ice-free area is along the edges of the continent, but other ice-free areas include mountain tops, cliffs, and valleys (Lee et al. 2017). These ice-free areas support breeding populations of 20 species of birds, including five species of penguins.

 

Species of birds that breed in Antarctica (Data from Harris et al. 2015).

Common name	Scientific name	Estimated global population (pairs)
Emperor Penguin	Aptenodytes forsteri	238,000
Gentoo Penguin	Pygoscelis papua	387,000
Adélie Penguin	Pygoscelis adeliae	3,790,000
Chinstrap Penguin	Pygoscelis antarctica	2,650,000
Macaroni Penguin	Eudyptes chrysolophus	6,300,000
Wilson’s Storm-Petrel	Oceanites oceanicus	4 – 10,000,000
Black-belled Storm-Petrel	Fregetta tropica	160,000
Light-mantled Albatross	Phoebetria palpebrata	20,000
Southern Giant Petrel	Macronectes giganteus	50,000
Southern Fulmar	Fulmarus glacialoides	1,000,000
Antarctic Petrel	Thalassoica antarctica	3 - 7,000,000
Cape Petrel	Daption capense	670,000
Snow Petrel	Pagodroma nivea	1,300,000
Antarctic Prion	Pachyptila desolata	16,600,000
Imperial Shag	Phalacrocorax atriceps	13,000
Snowy Sheathbill	Chionis albus	10,000
Kelp Gull	Larus dominicanus	10 – 20,000
Antarctic Tern	Sterna vittata	36,700
South Polar Skua	Catharacta maccormicki	3000 - 7500
Brown Skua	Catharacta antarctica	3000 - 7500

 

Brown and South Polar skuas

 

 

Life cycles of adults in four species of penguins that breed in Antarctica: (a) Emperor Penguins, (b) Adelie Penguins in East Antarctica (EA) and
West Antarctica (WA), (c) Chinstrap Penguins, and (d) Gentoo Penguins. Only Emperor Penguins incubate eggs during Antarctica's winter months (Figure from Handley et al. 2021).

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