Stilts and Avocets

Visit a healthy wetland in many parts of the world and you may be able to see boldly patterned, lightly built birds with remarkably long legs and bills wading through the shallows. These are the members of the Recurvirostridae, commonly known as the stilts and avocets.

American avocets Recurvirostra americana, from here.


About a dozen species of recurvirostrid are currently recognised, depending on the exact classification scheme in play. They are divided between three genera with the avocets forming the genus Recurvirostra and the stilts divided between Himantopus and Cladorhynchus. The most obvious distinction between the two subgroups is in the shape of the bill: that of stilts is straight but avocets have a distinct upwards curve towards the end of theirs. A fourth genus has often been included in the Recurvirostridae for the ibisbill Ibidorhyncha struthersii, a striking-looking inhabitant of the upland rivers of the Himalayan plateau, but uncertainty about this bird's phylogenetic position has led most recent authors to exclude it from the family.

The recurvirostrids feed mostly on small aquatic invertebrates such as brine shrimp or insect larvae. Their long legs, among the longest relative to body size of any bird, allow them to wade in deeply in search of prey. Stilts actively probe the waters and underlying sediment whereas avocets tend to forage by sweeping their bill through the water side to side. Avocets and the banded stilt Cladorhynchus leucocephalus of Australia prefer brackish waters such as lagoons and estuaries, with the banded stilt congegrating around the great salt lakes of inland Australia. Breeding is conducted by monogamous pairs that share the duty of incubating their simple nest on the ground near water. These nests may be gathered into loose colonies; the banded stilt forms particularly large colonies in which the chicks are herded into communal creches of several hundred.

Pied stilt Himantopus leucocephalus, copyright JJ Harrison.


The majority of recurvirostrids are patterned with black or dark brown and white. The red-necked avocet Recurvirostra novaehollandiae has the head and neck coloured reddish-brown as does the American avocet R. americana during the breeding season. The banded stilt has a broad reddish-brown band across the top of the breast. There is also the black stilt Himantopus novaezelandiae of New Zealand, which is somewhat self-explanatory. Beaks are black in all species; the legs are grey in avocets and red in stilts.

Four geographically distinct species of avocet occupy the modern world: the American avocet in North America, the red-necked avocet in Australia, the pied avocet Recurvirostra avosetta in Eurasia and Africa, and the Andean avocet R. andina in South America. The Andean avocet is a bird of high altitudes, occupying shallow, alkaline lakes in the upper Andes. Cladorhynchus includes only the banded stilt. The most varied taxonomy concerns the genus Himantopus. Historically, all the black-and-white stilts (and sometimes also the black stilt) have been recognised as a single near-cosmopolitan species. In more recent years, the trend has been towards recognition of five or six distinct species in the genus. Most of these species are well separated geographically except for in New Zealand where the black stilt shares its range with the pied stilt Himantopus leucocephalus, a more recent immigrant from Australia. The breeding range of the black stilt is currently restricted to a relatively small area of New Zealand's South Island, and the species is considered endangered due to factors such as habitat alteration and the threat of hybridisation with the more abundant pied stilt*.

*It's worth spending some thought on the role of hybridisation as a conservation risk. Some observers may express concern that regarding hybridisation as a threat per se carries uncomfortable intonations of "racial purity", and that limiting the available gene pool may do more harm than good. After all, it's not as if the black stilt heritage of hybrid individuals is just gone (hybrids between the two species are, I believe, fully fertile and able to produce offspring of their own). The question is, I suppose, do the black stilt genes actually persist in the mixed population? Or does selection and/or drift winnow them out over time? This would be a difficult question to answer, and not without risk to find out.

Banded stilts Cladorhynchus leucocephalus and red-necked avocets Recurvirostra novaehollandiae, copyright Ed Dunens.


Phylogenetically, it is reasonably well established that recurvirostrids form a clade with the ibisbill and oystercatchers. This clade is in turn closely related to the plovers of the Charadriidae; indeed, many recent phylogenies have indicated that the recurvirostrid-oystercatcher clade may even be nested within the plovers as generally recognised. Considering the relatively small number of species in each clade, it might seem reasonable to suggest the recurvirostrids be reduced to a subfamily of the Charadriidae, but bird taxonomists being bird taxonomists, there seems to be more of a push to divide the Charadriidae up instead.

The fossil record of the Recurvirostridae is limited. A handful of species have been assigned to this family from the Eocene, but all are known from limited remains and their position is questionable. Coltonia recurvirostra is known from part of a wing from Utah; it was a relatively large bird, appearing to be more than one-and-a-half times the size of any living recurvirostrid. Fluviatilavis antunesi was described from a femur, humerus and radius from Portugal but was described as exhibiting some primitive features not found in modern recurvirostrids. It is also worth noting that its original description (Harrison 1983) compared it most favourably with the ibisbill, so if that species is not to be regarded as a recurvirostrid, probably neither is Fluviatilavis.

REFERENCE

Harrison, C. J. O. 1983. A new wader, Recurvirostridae (Charadriiformes), from the early Eocene of Portugal. Ciências da Terra 7: 9–16.

4 comments:

  1. A very striking group of shorebirds! It's a shame the charadriiform fossil record is generally so poor. Indeed, "Coltonia" is now considered a species of the anseriform Presbyornis, though the study that made this revision (Ericson, 2000) doesn't seem to be available online.

    Ericson, P.G.P. 2000. Systematic revision, skeletal anatomy, and paleoecology of the New World early Tertiary Presbyornithidae (Aves: Anseriformes). PaleoBios 20: 1-23.

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    Replies
    1. The article you refer to is available at https://ucmp.berkeley.edu/research/paleobios/paleobios-back-issues/.

      It seems odd that there are so few charadriiform fossils. One might think that their association with aquatic habitats would make them prime fossilisation candidates. Are their bones too delicate? Are they too difficult to recognise as Charadriiformes? Or is something else going on?

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    2. Oh, that's good to know! Last I checked, I wasn't able to find much of an online presence for older PaleoBios articles; it's good to see that they've been made digitally available after all.

      The dearth of Paleogene charadriiform fossils is a curious puzzle indeed. I wonder if they simply didn't become diverse/numerous until the Neogene, at which point their fossil record does become somewhat better represented.

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    3. One thing I forgot to mention earlier along those lines is that, if there are no true Palaeogene recurvirostrids, that could be interesting in light of the suggestion from molecular phylogenies that the 'family' is more phylogenetically nested than previously appreciated.

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