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Tetrapod Zoology

Tetrapod Zoology

Amphibians, reptiles, birds and mammals - living and extinct

More passerines as seen from the peripheries (part III): Great tits!

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Parus major, aka Great tit, photographed in southern England in early 2014. Photo by Darren Naish.

Welcome to another of my articles on passerines from the peripheries. As before, the idea here is that we’re looking at passerine bird groups as seen ‘from the fringes’ – from an obscure, maritime archipelago on the eastern fringes of the North Atlantic, far from the places where these birds underwent most of their evolution and diversification. This time, we once again find ourselves surrounded by tits – Great tits Parus major, no less. If you’re a Tet Zoo regular you’ll recall Great tits being covered here once or twice before, most memorably in the article titled Great tits: still murderous, rapacious, flesh-rending predators! Anyway...

Gosler's 1993 The Great Tit, a fine book on tits.

Here (above) is a photogenic Great tit I encountered recently here in Southampton. The Great tit is an extensively studied bird. My first port of call when needing information on this species is Andrew Gosler’s 1993 book The Great Tit. He starts that book by noting how previous authors have frequently apologised for producing yet another study of what’s been regarded as the most extensively studied of all small birds (Gosler 1993). He also says therein that over 700 published studies have been devoted to this one species, and that was as of 1979, so I wonder what the figure is now. It should be said that tits as a whole are extensively studied: James et al. (2003) noted that at least 100 articles on the group are published every year.

For a tit, P. major is large (up to a gargantuan 20 g in some British birds). Its black ventral stripe, white cheeks and distinctive ‘teecha teecha’ springtime song are all highly familiar. Males and females can be easily distinguished on the basis of how extensive the belly stripe is (its size is also age-dependent), but the birds also exhibit dimorphism in the ultraviolet reflectance of their black crown feathers (Hegyi et al. 2007) – something invisible to us, of course, and something indicating that mutual sexual selection is at play, as it often seems to be in bird species where males and females display similar display patterns.

Great tits (almost) everywhere

Cinereous tit (P. cinereus) in Himachal Pradesh, India. Photo by J. M. Garg, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

The Great tit is insanely widespread, occurring from western Europe eastwards to Japan, across the Mediterranean region of Africa and as far south-east as Sumatra, Java and the Lesser Sundas (though read on). Because it’s so variable in size and plumage across this range, about 30 subspecies have been named, conventionally grouped into four ‘subspecies groups’ or ‘allospecies’. The full story is vastly complex, but the diversity of forms included within the P. m. major group that occur around the Mediterranean suggests that this was an area of early diversification before (some time within the last 8000 years or so) members of this particular group spread north and east. The histories of the other subspecies groups involve the evolution of numerous island-endemic forms, adaptation to high-altitude life on the Himalayan Plateau and elsewhere, and a post-glacial expansion northwards by birds restricted to a south-east Asian refugium during the Pleistocene. [Adjacent photo by J. M. Garg.]

However, those south-east Asian ‘Great tits’ – members of the P. m. cinereus group (they actually occur as far west as Iran) – are increasingly regarded as a distinct species, the Cinereous tit P. cinereus. Molecular analyses indicate that they’ve been distinct from P. major proper for something like 1.5 million years and clearly qualify as ‘distinct species’ when compared to other passerines (Martin et al. 2005).

Distribution map showing range of Great tit (P. major) and related taxa sometimes regarded as part of P. major as well (P. minor is the Japanese tit). Image in the public domain.

European Great tits are extensively predated upon by Eurasian sparrowhawks Accipiter nisus (so much so that some sparrowhawks are able to raise their chicks on Great tit and nothing else), but important predators at the nest site include Great spotted woodpeckers Picus major and weasels Mustela nivalis. Woodpecker predation on nesting birds is really interesting and I must talk about it at length some time. Weasel predation on tits is also interesting both because it involves a substantial amount of arboreal behaviour on the part of the weasels, but also because it’s tightly linked to the fortunes of local rodent populations. [Photo below by Luc Viatour.]

Great tit photographed in Belgium; photo by Luc Viatour, licensed under Creative Commons Attribution-Share Alike 2.0 Generic license.

Where do tits belong within the passeridan tree?

Tits are passerines, and are undeniably and obviously part of Passerida, the great clade that includes sparrows, finches, thrushes, larks, swallows, Old Word warblers and so on. But where do they belong within this group? That’s been hard to answer: they can’t really be said to have a ‘traditional’ placement, since they really have been put all over the passerine tree in different classification schemes. Since the 1950s, they’ve often been regarded as close relatives of nuthatches, properly termed Sittidae, and for this reason the two are included together in the same books (e.g., Harrap & Quinn 1996). Some phylogenetic studies that feature reasonably good sampling of taxa do actually find these two groups to be sister-taxa on the basis of osteological characters (James et al. 2003)

However, modern molecular studies do not recover any such relationship: nuthatches are part of Certhioidea, a clade that also includes treecreepers and wrens, while tits (Paridae) – together with penduline tits (Remizidae) – belong to a distinct lineage, now termed Paroidea (Johansson et al. 2008, Fjeldså 2013, Alström et al. 2014). Paroidea has generally been of uncertain position within Passerida but might be especially close to Hyliota and Stenostiridae, these three lineages forming a clade that’s close to Sylvioidea according to Alström et al. (2014). The thing that I would have said about tits several months ago – that they’re not especially closely related to any of the other major clades within Passerida – no longer seems to be true.

Substantially simplified depiction of passeridan passerine phylogeny, based mostly on Alström et al. (2014). All images by Darren Naish except the kinglet (by Alpsdake; licensed under Creative Commons Attribution-Share Alike 3.0 Unported license), waxwing (by Randen Pederson; licensed under Creative Commons Attribution-Share Alike 2.0 Generic license), and stenostirid (by Tom Tarrant; licensed under Creative Commons Attribution-Share Alike 3.0 Unported license).

[Kinglet photo above by Alpsdake, waxwing photo by Randen Pederson, stenostirid photo by Tom Tarrant.]

And that’s enough about tits for now. More passerines some time soon! For previous Tet Zoo articles on passerines, see...

Refs - -

Alström, P., Hooper, D. M., Liu, Y., Olsson, U., Mohan, D., Gelang, M., Hung, L. M., Zhao, J., Lei, F. & Price, T. D. 2014. Discovery of a relict lineage and monotypic family of passerine birds. Biology Letters vol. 10 no. 3 20131067.

Fjeldså, J. 2013. The global diversification of songbirds (Oscines) and the build-up of the Sino-Himalayan diversity hotspot. Chinese Birds 4, 132-143.

Gosler, A. G. 1993. The Great Tit. Hamlyn, London.

Harrap, S. & Quinn, D. 1996. Tits, Nuthatches & Treecreepers. Helm/A & C Black, London.

Hegyi, G., Szigeti, B., Török, J. & Eens, M. 2007. Melanin, carotenoid and structural plumage ornaments: Information content and role in great tits Parus major. Journal of Avian Biology 38, 698-708.

James, H. F., Ericson, P. G. P., Slikas, B., Lei, F.-M., Gill, F. B. & Olson, S. L. 2003. Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones. Ibis 145, 185-202.

Johansson, U. S., Fjeldså, J. & Bowie, R. C. K. 2008. Phylogenetic relationships within Passerida (Aves: Passeriformes): a review and a new molecular phylogeny based on three nuclear intron markers. Molecular Phylogenetics and Evolution 48, 858-876.

Martin, P., Martens, J., Eck, S., Nazarenko, A. A., Valchuk, O. P., Petri, B. & Veith, M. 2005. The great tit (Parus major) – a misclassified ring species. Biological Journal of the Linnean Society 86, 153-174.

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

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