The 'Messel rail' Messelornis cristata - a specimen with preserved plumage. Photo from here.
Despite its presentation in years of fieldguides and other popular books, the bird order 'Gruiformes' has in recent times been scattered to the four winds, with analyses both morphological and molecular proclaiming its polyphyly. Nevertheless, molecular analyses such as Hackett
et al. (2008) continue to support a clade roughly corresponding to the suborder Grues as recognised by Cracraft (1973)* containing the cranes and the rails. The morphological analysis of Livezey & Zusi (2007) on the other hand, does not support this clade, but it does support monophyly for each of the two primary divisions within Grues, the ralloid and gruoid lineages.
*
Just to confuse matters, the name "Grues" has been used by different authors for clades of differing inclusivity. Mayr (2009), for instance, uses "Grues" for the Aramus + Gruidae clade, and refers to the larger clade as "core Gruiformes".
The ralloid line contains the living families Rallidae*, the rails, and Heliornithidae, the finfoots (or should that be finfeet?) Cracraft (1973) regarded the Cretaceous
Laornis edvardsianus as a stem ralloid, but no-one else seems to have taken him up on this suggestion. More reliably on the ralloid stem are the Palaeocene to Oligocene Messelornithidae (Mayr, 2009). Messelornithids were medium-sized birds (about the size of a small chicken) best known from
Messelornis cristata for which over 500 specimens are available, some even with preserved feathering.
Messelornis was highly terrestrialised with limited flight capabilities and almost ludicrously long legs (loss or reduction of flight has been a common occurrence among the Grues). Its beak was relatively short and the overall appearance of
Messelornis would probably have not been dissimilar to the modern cariamas.
*
Hackett et al. (2008) resolved the Rallidae as paraphyletic to Heliornithidae, with Sarothrura (the flufftails) closer to Heliornis than to the other two included rails Himantornis and Rallus. A few places, at least online, have suggested recognising Sarothrura as a separate family from the Rallidae as a result, but I'd recommend waiting for a more detailed analysis with greater coverage of the Rallidae. Increased taxonomic coverage may return the flufftails to the other Rallidae, or it may make it more appropriate to treat the finfoots as derived rallids.
The sungrebe Heliornis fulica of tropical South America (I tried to find a picture of one carrying chicks, but no luck). Photo by Jerry Oldenettel.
The finfoots of the Heliornithidae are three species (one in Asia, one in Africa, one in South America) of tropical grebe-like birds, renowned for their reclusiveness. The South American sungrebe
Heliornis fulica is the most distinctive in appearance of the three species (though mitochondrial analysis indicates that it and the Asian
Heliopais personata form a clade to the exclusion of the African
Podica senegalensis - Fain
et al., 2007) and is also very distinct in its nesting behaviour.
Heliopais and
Podica, like most aquatic birds, have chicks that hatch out reasonably well-developed and immediately able to swim after their parents.
Heliornis, in contrast, has altricial chicks that hatch out after only ten to eleven days of incubation. The really amazing bit, though, is what happens
after the chicks hatch. The male sungrebe has a shallow pouch under each wing and he is able to transport the chicks inside this pouch, even flying with them. Whether the chicks remain in the pouches permanently or whether they are only placed in them while the male is travelling remains unknown. Funnily enough, while this chick-carrying behaviour was described by Alvarez del Toro in 1971, it had originally been recorded almost 140 years earlier by Prince Maximilian of Wied. It seems that everyone else had assumed the prince was smoking something.
Grey-winged trumpeters, Psophia crepitans. Photo by A. Vinot.
The gruoid lineage includes
Psophia, the trumpeters,
Aramus guarauna, the limpkin, and Gruidae, the cranes, as well as the fossil taxa
Parvigrus pohli, Geranoididae and Eogruidae. Most recent authors agree that
Aramus and Gruidae form a clade to the exclusion of
Psophia. The chicken-sized Oligocene
Parvigrus was originally described by Mayr (2005) as sister to
Aramus + Gruidae, but he later (Mayr, 2009) revised its position to stem gruoid.
Parvigrus lacked the long beak of limpkins and cranes, as do the Recent trumpeters, three species of similarly chicken-sized birds found in northern South America.
Whether Geranoididae and Eogruidae possessed crane-like long beaks is an unknown factor as skull material for both has not been found. Cracraft (1973) placed both outside the crown gruoids, but Clarke
et al. (2005) placed Eogruidae inside the gruoid crown as sister to
Aramus + Gruidae. The Eocene Geranoididae have been described only from leg bones (Wetmore, 1933, assigned some wing bones to
Geranoides jepseni in his original description of this species but did not describe them) so little can be said about them except that they were large and long-legged. Wetmore (1933) commented on the unusually wide spacing of the trochleae (the 'knuckles') at the end of the tarsometatarsus suggesting that
Geranoides had very widely splayed toes, but Cracraft (1969) later attributed to wide spacing to post-mortem distortion. Cracraft (1969, 1973) included a number of Eocene birds in the Geranoididae but admitted a lack of derived characters uniting them; Geranoididae may represent a paraphyletic assemblage of basal gruoids.
Distal ends of tarsometatarsi of the eogruids Proergilornis and Ergilornis, showing reduction of the inner trochlea in Proergilornis and its loss in Ergilornis. Figure from Cracraft (1973).
The Eocene to Pliocene Eogruidae were also decent-sized long-legged birds from central Asia and (in later times) Europe. Earlier authors recognised two families, Eogruidae and Ergilornithidae, but 'ergilornithids' are now recognised as derived eogruids. Eogruids were highly cursorial birds and a humerus attributed to
Ergilornis suggests that it was flightless, though the earlier
Eogrus aeola shows no sign of being so (Clarke
et al., 2005). Originally three-toed, eogruids showed a reduction in the size of the inner toe, and
Ergilornis and
Amphipelargus (the latest of the eogruids) lost it entirely (it is easy to present a progression from flying and three-toed to flightless and two-toed, but be warned that three-toed species survived into the Miocene, well after the appearance of the two-toed forms). The only other birds to reduce the number of toes to two are the ostriches, and a relationship between ostriches and eogruids has been suggested in the past (generally in association with the idea that the ratites do not form a monophyletic group). However, Cracraft (1973) confirmed that eogruids were more similar in their fine morphology to gruoids than ostriches, and modern phylogenetic analyses do not support a close relationship of ostriches and gruoids.
Many people carry the impression that flightlessness in birds is associated with lack of predators. However, eogruids evolved flightlessness in an environment in which predators were no rarity (amongst others, they shared their world with such horrors as hyaenodonts and entelodonts*). Similarly, while the exact circumstances in which they became flightless is unknown, modern ostriches (Africa), emus (Australia) and rheas (South America) all live alongside significant predators or at least did so until recently. Obviously, something other than lack of predators is at play here.
*
I always imagine Roald Dahl's hornswogglers to be something like an entelodont.
REFERENCES
Clarke, J. A., M. Norell & D. Dashzeveg. 2005. New avian remains from the Eocene of Mongolia and the phylogenetic position of the Eogruidae (Aves, Gruoidea).
American Museum Novitates 3494: 1-17.
Cracraft, J. 1969. Systematics and evolution of the Gruiformes (class, Aves). 1, The Eocene family Geranoididae and the early history of the Gruiformes.
American Museum Novitates 2388: 1-41.
Cracraft, J. 1973. Systematics and evolution of the Gruiformes (class Aves). 3, Phylogeny of the suborder Grues.
Bulletin of the American Museum of Natural History 151: 1-127.
Fain, M. G., C. Krajewski & P. Houde. 2007. Phylogeny of "core Gruiformes" (Aves: Grues) and resolution of the limpkin–sungrebe problem.
Molecular Phylogenetics and Evolution 43: 515-529.
Hackett, S. J., R. T. Kimball, S. Reddy, R. C. K. Bowie, E. L. Braun, M. J. Braun, J. L. Chojnowski, W. A. Cox, K.-L. Han, J. Harshman, C. J. Huddleston, B. D. Marks, K. J. Miglia, W. S. Moore, F. H. Sheldon, D. W. Steadman, C. C. Witt & T. Yuri. 2008. A phylogenomic study of birds reveals their evolutionary history.
Science 320: 1763-1768.
Livezey, B. C., & R. L. Zusi. 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): 1-95.
Mayr, G. 2005. A chicken-sized crane precursor from the early Oligocene of France.
Naturwissenschaften 92: 389-393.
Mayr, G. 2009.
Palaeogene Fossil Birds. Springer.
Wetmore, A. 1933. Fossil bird remains from the Eocene of Wyoming.
Condor 35: 115-118.
