Tetrapod Zoology

Tetrapod Zoology

Amphibians, reptiles, birds and mammals - living and extinct

Getting a major chapter on birds ALL birds into a major book on dinosaurs


After the long gestation period that’s typical of big, multi-authored volumes, the second edition of The Complete Dinosaur (Brett-Surman et al. 2012) has finally hit the bookstores and I’m now in happy possession of my own copy. The Complete Dinosaur first appeared in 1997 as a compendium of all things dinosaur. Edited by Jim Farlow and Mike Brett-Surman, and including the work of 47 contributing authors, it became well known as a good, non-technical guide to Mesozoic dinosaurs and their world (Farlow & Brett-Surmam 1997). But much has changed in those intervening 15-ish years and, over the past several years, a second, substantially updated edition has been put together.

As you can (hopefully) appreciate from the photos here, The Complete Dinosaur, Second Edition is huge: it’s bigger overall than the first edition at 1112 pp (vs a pathetic 752 pp) and is also larger-format, with a cover size of 288 x 220 mm, vs 175 x 250 mm for the first edition. It’s about 60 mm thick, whereas the first edition is about 30 mm thick. Thomas Holtz is part of the editorial team, meaning that two of the world’s best non-technical volumes on Mesozoic dinosaurs are now associated with his name (the other is his 2007 Dinosaurs: the Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, discussed here on Tet Zoo ver 2).

Some chapters in The Complete Dinosaur, Second Edition are much the same as those in the first edition, but others – especially those on the different dinosaur groups – have been substantially modified, or are totally new (examples: the first edition included a prosauropod chapter by Jacques VanHeerden and a theropod chapter by Phil Currie; the second edition’s chapters on these groups are by Adam Yates and Tom Holtz, respectively). Furthermore, there are new chapters on dinosaur musculature (Dilkes et al.), palaeoneurology (Buchholtz), human history and the fate of dinosaur remains (Chure), and the relevance of dinosaurs for evolutionary theory in general (Padian and Burton). And then there’s the bird chapter. That’s why we’re here.

Picture to text ratio in the bird chapter (Naish 2012). Nice.

Yeah, the bird chapter is written by that Naish guy. The article is broken into the following 21 sections; Introduction; Gerhard Heilmann’s The Origin of Birds; John Ostrom and the Theropod Hypothesis; The “Birds Are Not Dinosaurs” Movement; The Origins of Avialian* Flight; Avialian Anatomy; The Mesozoic Avialian Radiation; The Enantiornithes; Basal Ornithurines; The Rise of Modern Birds, the Neornithines; Assembling the Neornithine Tree; Palaeognaths: Ratites, Tinamous, and Lithornithids; Waterfowl and Gamebirds: the Galloanserae; Neoaves: the Waterbird Assemblage; Shorebirds and Kin, and Grebes and Flamingos; Birds of Prey; Strisores: Swifts, Hummingbirds, and Nightbirds; Pigeons, Cuckoos and Mousebirds; Higher Landbirds: Coraciiformes, Piciforms, Falcons, and Parrots; The Passerine Radiation; and The Future of Neornithine Diversity. That last section is depressing.

* I've bowed to peer-pressure since writing Naish (2012), and now use 'avialan' instead of 'avialian'.

The fossil record gives us continuity between birds and other dinosaurs. Diagrams (corvid skull above, ratite and gamebird pelves below) by Darren Naish, from Naish (2012).

I consider it a minor achievement in the world of dinosaur-bird relations that – unlike most articles that deal with birds and appear in the Mesozoic-themed literature – it covers Cenozoic birds as well as Mesozoic ones (Naish 2012). In fact, about 25 of its 44 pages are devoted to Cenozoic birds, arguably making this the longest contribution on post-Cretaceous dinosaurs to appear in the Mesozoic-themed dinosaur literature.

Why do this? I mean: why bring Cenozoic animals into a book about Mesozoic animals and their world? One argument could be that the continuity between Mesozoic dinosaurs and their post-Cretaceous kin has become an increasingly important theme the more we’ve learnt: remember, ‘dinosaurs’ and birds are not distinct entities – they grade smoothly into one another. So an argument can be made that those interested in Mesozoic dinosaurs will benefit from greater exposure to post-Cretaceous dinosaur biology, diversity and evolution. Conversely, it’s increasingly recognised that our understanding of bird evolution and biology has the potential to be greatly informed by consideration of whatever happened in the non-birds that went before (Prum 2002, Kaiser 2007).

Another argument is that a review of Cenozoic bird diversity and history is not only interesting but also needed, given that few existing works do this already. If you want a review on Cenozoic bird history, what is there? There’s Feduccia’s (1996) book; I’m not a fan, since I think that his take on bird history and evolution is idiosyncratic and frequently misleading, and I don’t just mean his take on Mesozoic bird history and bird origins (read on). Then there’s Storrs Olson's now very dated (but generally very useful) 1985 book chapter (Olson 1985) and Jiří Mlíkovský’s catalogue of European fossil birds (Mlíkovský 2002). The latter is so quirky (example: hornbills are the sister-group of loons) and contains so many unsubstantiated taxonomic revisions that some ornithologists would prefer it to be ignored (Mourer-Chauviré 2004). More recently, Gerald Mayr has produced an invaluable, wonderfully illustrated technical review of Paleogene birds (Mayr 2009), but (1) it’s only on Paleogene birds, and (2) it’s crazy expensive.

A selection of modern and recently extinct paleognaths. A, Double-wattled or Southern cassowary (Casuarius casuarius). B, Single-wattled cassowary (C. unappendiculatus). C, Brown kiwi Apteryx (the species-level taxonomy of Apteryx is currently unresolved). D, Little spotted kiwi (A. owenii). E, The robust-legged moa Pachyornis elephantopus. F, Head of moa Euryapteryx geranoides. G, Elephant bird Aepyornis maximus. Images by Darren Naish, from Naish (2012).

What I would do differently

My final, finished chapter looks great. In general, I’m happy with it, and I’m certainly very pleased to see it included in such a noteworthy tome. But there are some things that I wish were different. While I put more than enough time into the preparation of the many illustrations (there are 46 different figures, many combining several individual components), I really wish I’d produced even more – when I was writing the text, I didn’t realise it would be published with huge, 70-mm-wide white margins down the outer sides of each page. I could have filled more, most or even all of that margin space with additional diagrams, and it’s annoying now to see several bird groups (including penguins, cranes, trogons, pigeons, kingfishers and kin, woodpeckers, falcons and parrots) lacking illustrated representatives. Oh well, 46 figures seemed plenty enough at the time of submission.

In order, I presume, to save space, some of my illustrations have been published at tiny size. I’m not complaining – I absolutely understand why this had to be done. For those interested in seeing larger versions, they’re included here, and higher-resolution versions will be put on my deviantART page when time allows.

1, Galliformes (gamebirds or fowl); 2, Anseriformes (waterfowl or wildfowl); 3, Strisores (nightbirds, swifts and hummingbirds); 4, Eurypygae (kagus and sunbittern); 5, Columbiformes (pigeons); 6, Phaethontidae (tropicbirds); 7, Phoenicopteriformes (flamingos); 8, Podicipediformes (grebes); 9, Musophagidae (turacos); 10, Otididae (bustards); 11, Cuculiformes (cuckoos); 12, Rallidae (rails); 13, Gruidae (cranes); 14, Gaviiformes (loons or divers); 15, Sphenisciformes (penguins); 16, Procellariiformes (tube-nosed seabirds); 17, Ciconiidae (storks); 18, Pelecanidae (pelicans); 19, Ardeidae (herons); 20, Threskiornithidae (ibises and spoonbills); 21, Fregatidae (frigatebirds); 22, Suloidea (gannets, cormorants, and anhingas); 23, Scolopaci (sandpipers and kin); 24, Turnicidae (buttonquails or hemipodes); 25, Lari (auks, gulls, and kin); 26, Accipitriformes (vultures, eagles and kin); 27, Strigiformes (owls); 28, Coliiformes (mousebirds); 29, Coracii (rollers and kin); 30, Piciformes (woodpeckers and kin); 31, Cariamidae (seriemas); 32, Falconidae (falcons); 33, Psittaciformes (parrots); 34, Suboscines (pittas, broadbills, ovenbirds, and kin); 35, Passerida (finches, thrushes, warblers, and kin); 36, Corvoidea (crows and kin).

One of my diagrams (Fig. 20.19, Naish 2012) depicts a simplified ‘consensus’ tree of affinities within neognath neornithines. Based in part on the topology recovered by Ericson et al. (2006) and Hackett et al. (2008), the tree shows an early split between Galloanserae and Neoaves, and a divergence in Neoaves between a gruiform + Aequornithes clade, and a Charadriiformes + Mirandornithes + ‘landbird’ clade. That last big clade (Charadriiformes + Mirandornithes + ‘landbirds’) corresponds approximately to Terrestrornithes Livezey & Zusi, 2007 (and I’ll use that name from hereon, purely for convenience). ‘Landbirds’ might best be termed Dendrornithes, also as per Livezey & Zusi (2007). The ambiguity about the terms Terrestrornithes and Dendrornithes, incidentally, comes from the fact that they weren’t given specific definitions when named, so it isn’t at all clear (given the marked differences between Livezey & Zusi’s cladograms and those published in other studies) which lineages they have to include before evaporating. Anyway, in the ‘landbird’ (= Dendrornithes?) clade, I showed eagles, owls, mousebirds, rollers and woodpeckers on one branch, and pigeons, Strisores (‘nightbirds’, swifts and hummingbirds), parrots and passerines on another. I didn’t (Naish 2012) show seriemas or Eurypygae (sunbittern and kagu) at all.

Not a muppet, a Large owlet-nightjar (Aegotheles insignis). Image by Darren Naish, from Naish (2012).

As I said in the caption to the tree, showing pigeons and apodiforms as being ‘landbirds’ close to parrots and passerines is very likely not correct. I mostly went for this position because it’s been supported in some morphological analyses (Mayr & Clarke 2003) (I should note here that the hummingbird is meant to be standing in for the whole of Strisores, the neornithine clade that includes ‘nightbirds’ (that is, nightjars, potoos, frogmouths, owlet-nightjars and oilbirds) and swifts as well as hummingbirds). However, Hackett et al. (2008) found Columbiformes, Eurypygae and Strisores to be outside the huge Aequornithes + Terrestrornithes clade. This result – with the inclusion of Columbiformes, Eurypygae and Strisores within Metaves – is also seen in Ericson et al. (2006) and Ericson (2012). If Metaves is identified, I suppose Coronaves must be too (Fain & Houde 2004).

Seeing as I can modify my diagram as and when I choose, the version above is an updated one that accounts for those corrections; it's somewhat different from the version in Naish (2012). It’s still not meant to show all neognath lineages, but here is what I’ve changed: (1) Columbiformes, Eurypygae and Strisores are now outside of the Aequornithes + Terrestrornithes clade, in a clade that mostly matches Metaves; (2) tropicbirds (Phaethontidae) are shown in Metaves as well; (3) Mirandornithes – the flamingo + grebe clade – is shown in two possible positions, one in Metaves, and one close to Charadriiformes; (4) I added turacos (Musophagidae), since molecular datasets seem to agree in showing this group to be close to Aequornithes and/or Gruiformes; finally, (5) seriemas and falcons are now included and shown as close to parrots and passerines. Proposed names have been added where appropriate.

Hugely simplified Metaves section, from diagram shown above. Mirandornithes: is that node-based, or branch-based? In this case, it really is node-based. From Naish (2012).

One problem with my diagram is that the use of big obvious markers for clade names makes it look as if the respective names are specifically node-based or branch-based. That’s not deliberate – I used the markers merely to show as clearly as possible which groups the names applied to, and in most (but not all) cases there are not, in fact, formalised phylogenetic definitions. A specific, node-based definition does exist for Mirandornithes (Sangster 2005), but things aren’t so clear for the other names.

All that new stuff

As usual, many new discoveries have been published since I submitted the final version of the chapter, and I would have referred to them if only I could. Here are some of them.

I included a longish section on the ‘Birds Are Not Dinosaurs’ movement (Naish 2012, pp. 382-383). In part I agree with those who think that the criticisms of Alan Feduccia and colleagues are irrelevant and should be ignored, but I also think that we should take the time to point out that what Feduccia says as goes bird origins is naïve, misleading and erroneous. For those who need a refresher, Feduccia argues that birds are not theropod dinosaurs – they simply can’t be, because (he argues) non-bird dinosaurs are too cursorial, too big, too short-armed etc. – but instead descend from an unidentified cloud of pre-dinosaurian archosauromorphs, ostensibly including Longisquama, Megalancosaurus and anything else small and pointy-snouted (Feduccia 1996, 2012). Unambiguously feathered deinonychosaurs and oviraptorosaurs? They’re not unarguable theropods, absolutely irrelevant to bird origins and only convergently similar to them, as he has long argued (Feduccia 1996), but are actually flightless birds, closer to modern birds than is Archaeopteryx, and any similarity they have with other theropods is the result of the sort of hypothesised convergence (or parallelism) otherwise popular in the 19th and early 20th centuries.

Since I wrote the text, Feduccia’s new book – Riddle of the Feathered Dragons: Hidden Birds of China – has appeared (Feduccia 2012). I don’t dislike it because I dislike non-standard phylogenetic hypotheses. I dislike it because many of its arguments are erroneous (e.g., yet again we see the claim that theropods were on some sort of trend toward shortening their forelimbs, as if this is (a) true, and (b) relevant), and also because it involves deliberate attempts at obfuscation (e.g., there’s a lengthy section explaining why Mark Norell and colleagues ended up using the nickname ‘Dave’ for the feathered dromaeosaurid NGMC 91; another lengthy section is a biography of Thomas Huxley that points repeatedly to Huxley’s errors as goes the interpretation of such things as amphioxus. So far as I can tell, these sections are irrelevant to bird origins and dinosaur phylogeny, and only seem included such that they might weaken the authority of the people concerned) (Feduccia 2012). The book is unimpressive in production terms, with highly pixelated diagrams and an over-packed look to the text.

Anyway, my point here is that I would have included some criticism of this book were it out in time. By the way, I’m tired of seeing people defend Feduccia because he’s a decorated and highly respected senior professor. I’m sure that his work and achievements deserve accolade, and there’s no doubt that he’s a brilliant scientist overall, but it doesn’t change the fact that his published arguments on bird and dinosaur evolution fail to account for the data we have, and rely on erroneous arguments.

Enantiornithine nesting colony; image by Julio Laceda.

Various noteworthy contributions or discoveries have been published on Mesozoic birds since my chapter’s text was finalised. Toothless Zhongjianornis – said in my text to be close in the phylogeny to Jeholornis and the omnivoropterygids – has more recently been interpreted as a stem-ornithuromorph, similar in ‘grade’ to Patagopteryx and Vorona (O’Connor & Zhou 2012). Meanwhile, long-tailed, short-armed Dalianraptor – mooted in my text as a possibly flightless relative of Jeholornis – turns out to be "tampered" with, and hence might be a composite. The discovery of an enantiornithine nesting colony (Dyke et al. 2012) would definitely warrant mention, since in a way it makes the biology of these Cretaceous birds more like that of modern ones [adjacent reconstruction of the nesting colony by Julio Laceda: previously covered here on Tet Zoo].

When we come to Cenozoic birds, the remarkable bony-toothed birds, pseudotoothed birds or pelagornithids have been the focus of a fair bit of research interest since the text was written. New records of the group have been described from Africa and Australia; perhaps most importantly, Mayr (2011) has argued that character evidence supposed to link pelagornithids with waterfowl is erroneous (I was fairly kind to this hypothesis), and he argues instead that they might be the sister-group to the whole of Galloanserae (aka Galloanseres). In other Cenozoic news, hoatzins are seemingly now known from Africa (Mayr et al. 2011), and Kurochkin & Dyke (2011) have reviewed the fossil record of owls (they name the new ‘family’ Heterostrigidae for Oligocene owls from Mongolia).

When I was preparing the manuscript, the fluvioviridavids of Eocene Europe and North America were considered possible members of Strisores (the ‘caprimulgiform’ + apodiform clade). Since then, Nesbitt et al. (2011) have argued that Fluvioviridavis is in fact a stem-member of the frogmouth lineage, and hence well nested within Strisores, while Eurofluvioviridavis is not a member of Strisores and remains enigmatic.

A selection of modern anseriforms. A, Northern or Black-necked screamer (Chauna chavaria). B, Whooper swan (Cygnus cygnus). C, Common merganser or Goosander (Mergus merganser). D, Southern or Red shoveler (Anas platalea). E, Tundra bean goose (Anser serrirostris). Images by Darren Naish, from Naish (2012).

While I referred to the idea of a close affinity between falcons, parrots and passerines – I was basing this on Hackett et al. (2008) – this hypothesis has since been bolstered by Suh et al. (2011). Thanks to their study of retroposons – ‘jumping’ genetic sequences that seem to insert themselves randomly in the genome – they were able to provide strong additional evidence for a sister-group relationship between parrots and passerines, with falcons being the next closest relatives of this clade. They termed the parrot + passerine clade Psittacopasserae, and the falcon + (parrot + passerine) clade Eufalconimorphae (Suh et al. 2011). Ericson (2012) also proposed some new names for certain recently recovered clades. Afroaves was suggested for the clade that includes accipitriforms, Leptosomus (the Madagascan cuckoo-roller or Courol), owls, trogons and Picocoraciae, and Australavis for the cariamid + Eufalconimorphae clade. Ericson (2012) also suggested Cantiomimus for the clade named Eufalconimorphae Psittacopasserae by Suh et al. (2011).

A selection of suboscine and oscine passerines. Images by Darren Naish, from Naish (2012).

I’m sure there are other things in the chapter that are now inaccurate in view of new discoveries and new analyses, and I’m sure there’s also stuff that’s simply wrong and will be shown to be so in time. Of course, it really is incredibly difficult – perhaps impossible – to make material destined for the printed literature to be up-to-the-minute, especially in such a vibrant and fast-moving field as palaeornithology. I tried my best.

So, I’m interested to know what people think of having a lengthy, mostly Cenozoic-themed bird chapter in what is definitely a Mesozoic-themed book. The claim that ‘dinosaur people’ “just don’t know birds” (guess who said this) is wholly unfair and untrue, and the general idea that people either work on animals from the Mesozoic or on animals from the Cenozoic is obsolete and becoming ever more so with time. And, I’ll say it again, reviews of Cenozoic bird diversity and evolution are few and far between. I hope the chapter is considered worthy and interesting, and I even hope that those more interested in Cenozoic animals than Mesozoic ones get something out of it.

I must finish by saying thanks to the editors and everyone else on the editorial and in-house publishing team for steering this enormous project and for getting me on board in the first place. Thanks also to all the colleagues and artists who allowed use of their images: John Conway, Gareth Dyke, Denver Fowler, Bent Lindow, Gerald Mayr, Hanneke Meijer, Greg Paul, Corwin Sullivan, Inge van Noortwijk and Mark Witton. The Complete Dinosaur, Second Edition… another of those ‘must have’ volumes! It's here on amazon, and here on

And - PS - weren't not yet done on big new dinosaur books...

For other Tet Zoo articles relevant to some of the topics discussed here, please see...

Refs - -

Brett-Surman, M. K., Holtz, T. R., Farlow, J. O. 2012. The Complete Dinosaur, Second Edition. Indiana University Press, Bloomington, Indiana.

Dyke, G., Vremir, M., Kaiser, G. & Naish, D. 2012. A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania. Naturwissenschaften 99, 435-442.

Ericson, P. G. P. 2012. Evolution of terrestrial birds in three continents: biogeography and parallel radiations. Journal of Biogeography 39, 813-824.

Fain, M. G. & Houde, P. 2004. Parallel radiations in the primary clades of birds. Evolution 58, 2558-2573.

- ., Anderson, C., Britton, T., Elzanowski, A., Johansson, U., Kallersjo, M., Ohlson, J., Parsons, T., Zuccon, D., & Mayr, G. 2006. Diversification of Neoaves: integration of molecular sequence data and fossils. Biology Letters 2, 543-547.

Farlow, J. O. & Brett-Surman, M. K. 1997. The Complete Dinosaur. Indiana University Press, Bloomington, Indiana.

Feduccia, A. 1996. The Origin and Evolution of Birds. Yale University Press, New Haven & London.

- . 2012. Riddle of the Feathered Dragons: Hidden Birds of China. Yale University Press (New Haven & London).

Hackett, S. J., Kimball, R. T., Reddy, S., Bowie, R. C. K., Braun, E. L., Braun, M. J., Cjojnowski, J. L., Cox, W. A., Han, K.-L., Harshman, J., Huddleston, C. J., Marks, B., Miglia, K. J., Moore, W. S., Sheldon, F. H., Steadman, D. W., Witt, C. C. & Yuri, T. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320, 1763-1768.

Kaiser, G. W. 2007. The Inner Bird: Anatomy and Evolution. UBC Press, Vancouver.

Kurochkin, E. N. & Dyke, G. J. 2011. The first fossil owls (Aves: Strigiformes) from the Paleogene of Asia and a review of the fossil record of Strigiformes. Paleontological Journal 45, 445-458.

Livezey, B. C. & Zusi, R. L. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society 149, 1-95.

Mayr, G. 2009. Paleogene Fossil Birds. Berlin: Springer.

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O’Connor, J. K. & Zhou, Z. 2012. A redescription of Chaoyangia beishanensis (Aves) and a comprehensive phylogeny of Mesozoic birds. Journal of Systematic Palaeontology DOI:10.1080/14772019.2012.690455

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Prum, R. O. 2002. Why ornithologists should care about the theropod origin of birds. Auk 119, 1-17.

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Suh, A., Paus, M., Kiefmann, M., Churakov, G., Franke, F. A., Brosius, J., Kriegs, J. O. & Schmitz, J. 2011. Mesozoic retroposons reveal parrots as the closest living relatives of passerine birds. Nature Communications Aug 23;2:443. doi: 10.1038/ncomms1448.

The views expressed are those of the author and are not necessarily those of Scientific American.

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