Ornithology NREM/ZOOL 4464 Dr. Tim O’Connell Spring 2015

A Classification of Birds of the World – Laboratory Investigations

As students of Ornithology, a primary objective for you is to develop a working knowledge of the diversity of birds of the world. As you internalize this information, you will develop a deep appreciation for the central role that evolution plays as the driver of biological diversity.

We begin this week in Lab to really explore the diversity of birds through in-depth examination of orders and families. Avian classification is fluid: the ordering and alliance of families and orders differs according to different authorities and new information is continually becoming available that enhances our ability to discern relationships; this results in updated classifications. You are embarking on this voyage during a time of significant taxonomic revision, and much of what you will learn in Lab supersedes classification systems in your field guides.

Different authorities (usually committees) often disagree in their assessment of a taxon. Where one authority might recognize a full species, another might recognize it as a subspecies of some other species. Higher-level assignments are tricky too, e.g., some place the South American rheas in their own order (Rheiformes) whereas others lump it in with other ostrich-like birds in Struthioniformes.

The American Ornithologists’ Union manages two committees (North American and South American Classification Committees) that review information and render decisions on which taxonomic proposals will be recognized. Each July, the AOU publishes updates to its official checklist. That checklist is commonly accessed via a massive book containing information on distribution, evolutionary affinities, etc. for every species, subspecies, hybrid, etc. recognized by the AOU. This book was first authored, and then updated, by the late James Clements:

Clements, J. F., T. S. Schulenberg, M. J. Iliff, D. Roberson, T. A. Fredericks, B. L. Sullivan, and C. L. Wood. 2014. The eBird/Clements checklist of birds of the world: Version 6.9. Downloaded from http://www.birds.cornell.edu/clementschecklist/download/

Clements et al. 2014 is a fascinating body of work. For perspective, the 10,000 or so species of birds are listed one per row in a giant Excel spreadsheet. The distribution of all the subspecies and other taxonomic categories, however, expands that spreadsheet to a whopping 32,000 rows!

As one effort to streamline the list into something more generally manageable, the eBird management team has produced a modified Clements et al. 2014 that lists all of the species and other taxonomic categories that can be entered in eBird. This one is 14,440 rows, and it is the backbone of the classification system we will use: http://www.birds.cornell.edu/clementschecklist/download/.

As you learned in lecture, however, the Avian Phylogenomics Project has recently published a series of papers relying on whole genome analysis (http://avian.genomics.cn/en/index.html), thereby providing unprecedented insights into evolutionary relationships of birds. I would be remiss not to include some big-picture findings of this project into your classification, so I have.

Thus, the official NREM/ZOOL 4464 taxonomic classification blends the eBird scheme (based on Clements’ work) with recent developments from the Avian Phylogenomics Project. This is the only place in the world where you will find this exact taxonomic classification.

1 Now comes the fun part – what you are required to know.

There are 37 orders within the class Aves. You are required to learn 32.

There are 236 families in those orders. You are required to learn 110 of them.

That’s 142 new Greek- and Latin- names for you.

At a minimum, you should concentrate on the following for each family and order:

*unique/characteristic morphology, behavior, or physiology *approximate number of species *primary habitat and foods *life history re: nesting habits, migratory behavior, etc. *biogeography and distribution

My best advice for keeping these straight: Use the information provided to you to make 110 flash cards; bring those cards with you everywhere you go.

Higher order classification

Recall that all taxonomic categories above the species are arbitrarily defined (and “species” is subject to multiple definitions too). Thus, it can be especially confusing to determine what is really a “class” “infraclass,” “subclass”, “superorder”, etc. For example, we learned that birds evolved from dinosaurs. “Dinosauria” is (according to at least one authority) a superorder within an infraclass within a subclass within a class within a series – whatever a series is – of reptiles. By convention, taxonomic classifications among the dinosaurs are not meant to be hierarchical.

For our purposes, we will treat birds traditionally and simply as the Class Aves within the Subphylum Vertebrata of the Phylum Chordata. (If you prefer, you can go full-on dinosaur and consider birds to be in the Class “Reptilia-Dinosauria-Saurischia-Theropoda-Avialae”, but I think “Aves” is a bet less clunky.) There are splits within the Aves that occur at higher than order levels; we will just refer to them generically as “groups” or “clades.”

The Aves first appear in the Jurassic Period of the Mesozoic Era, about 150–200 million years ago. Archaeopteryx lithographica is still one of the earliest known birds (as early as 150 MYA; Anchiornis shows up about 10 million years earlier), but it is not regarded as the primary ancestor of modern birds.

The cladogram (left) illustrates several important splits among early birds. Birds diversified in the Cretaceous Period, and several fossil specimens are described within the Confusiusornithidae, the Enantiornithes, and the loonlike Hesperornithoformes. All modern birds are part of the Neornithes.

Within the Neornithes, the first big split concerns the Paleognathae and Neognathae. These groups are defined by differences in the arrangement and development of the bones that make up the palate in the roof of the mouth: In the more ancestral Palaeognathes, the bones are robust and rigid. In the Neognathes, the bones are more gracile, and actually form a flexible joint within the palate.

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The Paleognathae includes 5 extant orders (plus the extinct moas of New Zealand and Elephant Bird of Madagascar). Four of those orders (all lumped at left under the umbrella of the ostrich Struthioniformes) are known as “ratites”: ostriches, rheas, emus/cassowaries, and kiwis. These flightless birds lack a keel on the sternum, and ratite comes from the Greek for “raft”, i.e., a flat-bottomed boat. The 5th order of extant Paleognathes consists of the 49 species of Tinamous from the Neotropics. Tinamous have similar ancestral palate structure, but they differ from the ratites in their ability to fly (barely) and their decent keel on the sternum.

Among the Neognathes, the next big split occurs between the chickens and ducks (orders Galliformes and Anseriformes, grouped together as the “Galloanserae”) and everything else, the Neoaves.

By far, most living birds are classified in the orders and families of the Neoaves. There is a big split in this group between orders that are “pigeony” (Columbea) and “everything else” (Passerea). Passerea includes all of the songbirds (i.e., passerines, order Passeriformes), plus the hawks, shorebirds, pelicans, parrots, etc. Passeriformes is by far the biggest and most diverse order. We recognize three suborders according to the degree of ossification within the syrinx (i.e., avian voice box): the New Zealand wrens (Acanthisitti), the Tyranni (a.k.a. suboscines), and Passeri (oscines). The most accomplished singers with the greatest voice control are oscine passerines; the suboscines and New Zealand wrens have comparatively more simple vocalizations.

All the Birds In the accompanying Excel workbook, you will find three worksheets. These are entitled eBird every species, all orders and families, and for you to know. I hope those titles are self-explanatory, but it’s important for you to examine all three to really understand what we’re doing.

In for you to know, I have first listed orders and families independently. Next you will find a combined list of families within orders, with 1s and 0s indicating the families that you will and will-not need to know for your next practical. Finally, beginning with column J, I have provided you with a list of the specific 110 families in 32 orders you need to know. Your first step should probably be to commit to memorizing the names. For example, if we show you an ibis and ask for the family and order, you will need to reply with Threskiornithidae and Pelecaniformes, respectively, for full credit. “It’s an ibis” or “in the ibis family” would only net you partial credit. Likewise, spelling matters. For full credit, your answers need to be spelled correctly and written legibly.

How will you amass information like this

*unique/characteristic morphology, behavior, or physiology *approximate number of species *primary habitat and foods *life history re: nesting habits, migratory behavior, etc. *biogeography and distribution for 110 families of birds? Easy: go to hbw.com.

When prompted to sign in, use “OKStateOrnithology” and “OrniTax15”. This resource is available to you free of charge for one year, as a gift from your friendly neighborhood Ornithology professor.

3 Your next practical is scheduled for April 1 and 2. The required material will come from:

18/19 Feb: integument and skeleton 25/26 Feb: musculature and organ systems 4/5 Mar: global bird diversity 1 11/12 Mar: global bird diversity 2 25/26 Mar: global bird diversity 3

With the advice that you can begin today to collate all the information you will need to master by April 1&2, here is a breakout of the specific families and orders I expect to cover in our next three Labs.

Global Bird Diversity 1 – 4/5 March

Paleognathae ORDER FAMILY Rheiformes Rheidae (Rheas) Tinamiformes Tinamidae (Tinamous) Casuariiformes Casuariidae (Cassowaries) Apterygiformes Apterygidae (Kiwis)

Neognathae Galloanseres ORDER FAMILY Anseriformes Anatidae (Ducks, Geese, and Waterfowl) Galliformes Cracidae (Guans, Chachalacas, and Curassows) Galliformes Odontophoridae (New World Quail) Galliformes Phasianidae (Pheasants, Grouse, and allies)

Neognathae Neoaves Columbea ORDER FAMILY Podicipediformes Podicipedidae (Grebes) Phoenicopteriformes Phoenicopteridae (Flamingos) Columbiformes Columbidae (Pigeons and Doves)

Neognathae Neoaves Passerea ORDER FAMILY Cuculiformes Cuculidae (Cuckoos) Cuculiformes Musophagidae (Turacos) Otidiformes Otididae (Bustards) Apodiformes Trochilidae (Hummingbirds) Apodiformes Apodidae (Swifts) Caprimulgiformes Caprimulgidae (Nightjars and Allies) Caprimulgiformes Steatornithidae (Oilbird) Opisthocomiformes Opisthocomidae (Hoatzin) Gruiformes Gruidae (Cranes) Gruiformes Rallidae (Rails, Gallinules, and Coots) Gruiformes Aramidae (Limpkin)

4 Global Bird Diversity 2 – 11/12 March

Neognathae Neoaves Passerea ORDER FAMILY Charadriiformes Recurvirostridae (Stilts and Avocets) Charadriiformes Haematopodidae (Oystercatchers) Charadriiformes Charadriidae (Plovers and Lapwings) Charadriiformes Jacanidae (Jacanas) Charadriiformes Scolopacidae (Sandpipers and allies) Charadriiformes Alcidae (Auks, Murres, and Puffins) Charadriiformes Laridae (Gulls, Terns, and Skimmers) Gaviiformes Gaviidae (Loons) Sphenisciformes Spheniscidae (Penguins) Procellariiformes Diomedeidae (Albatrosses) Procellariiformes Hydrobatidae (Storm-Petrels) Suliformes Sulidae (Boobies and Gannets) Suliformes Phalacrocoracidae (Cormorants and Shags) Suliformes Anhingidae (Anhingas) Ciconiiformes Ciconiidae (Storks) Pelecaniformes Pelecanidae (Pelicans) Pelecaniformes Balaenicipitidae (Shoebill) Pelecaniformes Ardeidae (Herons, Egrets, and Bitterns) Pelecaniformes Threskiornithidae (Ibises and Spoonbills) Accipitriformes Cathartidae (New World Vultures) Accipitriformes Pandionidae (Osprey) Accipitriformes Accipitridae (Hawks, Eagles, and Kites) Strigiformes Tytonidae (Barn-Owls) Strigiformes Strigidae (Owls) Trogoniformes Trogonidae (Trogons) Galbuliformes Galbulidae (Jacamars) Piciformes Lybiidae (African Barbets) Piciformes Ramphastidae (Toucans) Piciformes Picidae (Woodpeckers) Coraciiformes Todidae (Todies) Coraciiformes Momotidae (Motmots) Coraciiformes Alcedinidae (Kingfishers) Coraciiformes Meropidae (Bee-eaters) Coraciiformes Coraciidae (Rollers) Coraciiformes Upupidae (Hoopoes) Coraciiformes Bucorvidae (Ground-Hornbills) Cariamiformes Cariamidae (Seriemas) Falconiformes Falconidae (Falcons and Caracaras) Psittaciformes Strigopidae (New Zealand Parrots) Psittaciformes Psittacidae (New World and African Parrots)

5 Global Bird Diversity 2 – 25/26 March

Neognathae Neoaves Passerea ORDER FAMILY Passeriformes - suboscine Philepittidae (Asities) Passeriformes - suboscine Pittidae (Pittas) Passeriformes - suboscine Thamnophilidae (Typical Antbirds) Passeriformes - suboscine Furnariidae (Ovenbirds and Woodcreepers) Passeriformes - suboscine Tyrannidae (Tyrant Flycatchers) Passeriformes - suboscine Cotingidae (Cotingas) Passeriformes - suboscine Pipridae (Manakins)

Neognathae Neoaves Passerea ORDER FAMILY Passeriformes – oscine Menuridae (Lyrebirds) Passeriformes – oscine Ptilonorhynchidae (Bowerbirds) Passeriformes – oscine Maluridae (Fairywrens) Passeriformes – oscine Meliphagidae (Honeyeaters) Passeriformes – oscine Callaeidae (Wattlebirds) Passeriformes – oscine Vangidae (Vangas, Helmetshrikes, and allies) Passeriformes – oscine Laniidae (Shrikes) Passeriformes – oscine Vireonidae (Vireos) Passeriformes – oscine Corvidae (Crows, Jays, and Magpies) Passeriformes – oscine Paradisaeidae (Birds-of-Paradise) Passeriformes – oscine Petroicidae (Australasian Robins) Passeriformes – oscine Picathartidae (Rockfowl) Passeriformes – oscine Alaudidae (Larks) Passeriformes – oscine Hirundinidae (Swallows) Passeriformes – oscine Paridae (Chickadees and Tits) Passeriformes – oscine Remizidae (Penduline-Tits) Passeriformes – oscine Sittidae (Nuthatches) Passeriformes – oscine Troglodytidae (Wrens) Passeriformes – oscine Polioptilidae (Gnatcatchers) Passeriformes – oscine Cinclidae (Dippers) Passeriformes – oscine Regulidae (Kinglets) Passeriformes – oscine Sylviidae (Sylviid Warblers) Passeriformes – oscine Muscicapidae (Old World Flycatchers) Passeriformes – oscine Turdidae (Thrushes and allies) Passeriformes – oscine Mimidae (Mockingbirds and Thrashers) Passeriformes – oscine Sturnidae (Starlings) Passeriformes – oscine Dicaeidae (Flowerpeckers) Passeriformes – oscine Nectariniidae (Sunbirds and Spiderhunters) Passeriformes – oscine Motacillidae (Wagtails and Pipits) Passeriformes – oscine Bombycillidae (Waxwings) Passeriformes – oscine Mohoidae (Hawaiian Honeyeaters) Passeriformes – oscine Ptiliogonatidae (Silky-flycatchers) Passeriformes – oscine Calcariidae (Longspurs and Snow Buntings) Passeriformes – oscine Parulidae (New World Warblers) Passeriformes – oscine Thraupidae (Tanagers and allies) Passeriformes – oscine Emberizidae (Buntings and New World Sparrows) Passeriformes – oscine Cardinalidae (Cardinals and Allies) Passeriformes – oscine Icteridae (Troupials and Allies) Passeriformes – oscine Fringillidae (Siskins, Crossbills, and allies) Passeriformes – oscine Passeridae (Old World Sparrows) Passeriformes – oscine Ploceidae (Weavers and Allies)

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