CNEMOPHILINES  Cnemophilidae It is frustrating to write this page. Not only am I 'way out on a limb' in segregating this group as a separate family, but I don't have any photos of actual birds to illustrate the discussion. The best I can do from my slide collection is this shot (below) of the road and habitat on Trauna Ridge, Papua New Guinea, in October 1983. A huge mudslide had blocked our drive to Jimi Valley and in this shot my friends Steve Wilson (left), Brian Finch (pink shirt) and Chris Spooner (upper right; with scope) are crossing it with some difficulty. It was just beyond this spot that we located what was then called Yellow-breasted Bird-of-Paradise, a cnemophiline that I now think is best be considered part of this small endemic New Guinea family.

The Cnemophilidae is a small group of little studied passerines endemic of the uplands of mainland New Guinea. There are just three species, all fabulously illustrated in this plate (left) by William T. Cooper from Frith & Beehler's (1998) The Birds of Paradise. These birds have recently been considered to be birds of paradise; Frith & Beehler (1998) considered them a subfamily [Cnemophilinae] and termed them the "wide-gaped birds of paradise." They are: Yellow-breasted Cnemophilus Loboparadisea sericea (upper right; male between juvenal and female), Crested Cnemophilus Cnemophilus macgregorii (on thick branch in middle; males of two subspecies in center between female and juvenal), and Loria's Cnemophilus C. loriae (lower left; male bottom right flanked by subadult male and female and juvenal higher). Scanning this artwork does not result in the exceptional quality of the published original; I highly recommend that you acquire and study Frith & Beehler (1998) and Cooper & Forshaw (1997) for better reproduction and many more details. These are very poorly known birds. Loria's Cnemophilus was initially described as in its own genus (Loria) but I use Frith & Beehler's generic arrangement here.

This group was initially thought to be related to bowerbirds [Ptilonorhynchidae]; indeed, ornithologists long sought for the bower of Crested Cnemophilus until Ernst Mayr (in Mayr & Gilliard 1954) found anatomical evidence linking it more closely with birds of paradise. Stonor (1937) had already moved Loria's Cnemophilus from the bowerbirds to the birds of paradise. Bock's (1963) osteology studies showed that this group was atypical of both bowerbirds and birds of paradise, although sharing more features of birds of paradise. They had a weak and broad-gaped bill, weak and non-manipulative feet, and an unossified nasal region. They shared some cranial characters, though, with bowerbirds. Bock (1963) and later Gilliard (1969) thought bowerbirds and birds of paradise to be closely related, and to have arisen from the same ancestor. Bock (1963) proposed that the cnemophilines (plus MacGregor's Bird-of-Paradise Macgregoria pulchra) were an ancient group in that lineage that later led to modern bowerbirds and birds of paradise.

The DNA research of Sibley & Ahlquist (1990) showed that bowerbirds and birds of paradise were not closely related. Rather, both the bowerbirds and the cnemophilines are early lineages of the great corvid assemblage that arose in Australasia. Frith & Beehler (1998) acknowledged this, but considered cnemophilines more closely related to birds of paradise than anything else. For one thing, recent information about the nest, eggs, and young of Crested Cnemophilus shows that it is unlike bowerbirds. Yet, Frith & Beehler (1998) admit that "the cnemophilines may, in fact, not be birds of paradise" (at page 174) but "it is certainly convenient to treat these three species" as birds of paradise.

This was the status quo until Cracraft & Feinstein (2000) published molecular evidence that cnemophilines are not closely related to birds of paradise. The biochemical evidence from mitochondrial DNA, as well as some fossil and phylogenetic evidence, show that the cnemophilines, like bowerbirds, are an ancient lineage near the base of the corvoid tree. They are not especially close to bowerbirds, though; the current evidence suggests that their closest relatives may be the cuckoo-shrikes [Campephagidae]. Incidentally, this same paper proved rather convincingly that MacGregor's Bird-of-Paradise is not a bird of paradise, either; it is actually a giant honeyeater [Meliphagidae], most closely related to the genus Melipotes. For those that know the Melipotes in New Guinea, this actually makes a lot of intuitive sense.

I am currently unaware of further research. From information on the Handbook of the Birds of the World web site, it appears that the cnemophilines will be retained by that project as a subfamily of the birds of paradise. It may be that the evidence presented by Cracraft & Feinstein (2000) will take some time — and probably some replication — to gain much traction against tradition. This may be because the publication is rather obscure. Further, the senior author (Joel Cracraft) has been involved in various controversial topics; he is, for example, a leading proponent of the phylogenetic species concept. His earlier review of all the birds of paradise (Cracraft 1992) recognized some 90 diagnosable phylogenetic species. In contrast (and including the three cnemophilines), the major work by Frith & Beehler (1998) — using a biological species concept — recognizes 42 species (and a host of subspecies). I am of the opinion that the biological species concept is a much better species concept [e.g., see my bird families essay that includes this topic], and therefore I do not agree with Cracraft's opinions on that issue. Yet, this biochemical and phylogenetic work on the cnemophilines seems very persuasive. As they are a well defined small set of similar species that are clearly unrelated to birds of paradise, retaining them in the Paradisaeidae for "convenience" does not seem right. Yes, we don't quite yet know where they will properly be placed among the corvid radiation (and therefore I have them next to the Paradisaeidae "for convenience"), but given that they are as distant from birds of paradise as are cuckoo-shrikes and crows and butcherbirds, it seems best to consider them a separate family.

From the perspective of a world birder, I have wanted to observe at least one representative of each distinctive group of birds; in other words, at least one member of each bird family. Clearly the cnemophilines are a distinctive set of birds and now that it is clear they are not birds of paradise, world birders with my interests should wish to search at least one of them out. Indeed, raising the cnemophilines to family level may serve to accentuate their importance in the broad scheme of things and re-rank priorities on a New Guinea visit, much the same way that raising both the regular berrypeckers [Melanocharitidae] and painted berrypeckers [Paramythiidae] to family status will cause one to revisit priorities. Handbook of the Birds of the World is elevating both those latter groups to family status; it seems eminently logical to do so for the Cnemophilidae as well.

Searching out any of the cnemophilines is not easily done. All are "highly frugivorous montane birds which are rather small, sexually dimorphic, compact with rounded wings, and generally unobtrusive habits" (Beehler et al. 1986). Loria's Cnemophilus may have the broadest range in the central highlands, mostly from 2000-4000m, but is inconspicuous except at fruiting trees. Crested Cnemophilus (once also called the "Sickle-crested Bird-of-Paradise") recalls a MacGregor's Bowerbird Amblyornis macgregoriae (the females are especially similar) and it lives in similar habitats. Coates (1990) calls it "an inconspicuous, compact, sluggish frugivore inhabiting high mountain forest and shrubbery. It is, however, probably the best known of the group since the nest and young have been studied and photographed; e.g., Cliff & Dawn Frith's photos of the nest, with female and young, are published in Coates (1990). Yellow-breasted Cnemophilus is the least known. Almost nothing is known of its biology, and it seems scarce and local within the patches of habitat along the central ranges east to the base of the Huon Peninsula.

While the colorful or exotic males are distinctive, females and young birds are quite drab, making them even more inconspicuous. Alas, although they sometimes move in small groups, most birds encountered seem to such dull-plumaged birds. Fully outfitted males are scarce (Coates 1990, pers. obs.). My experiences in 1983 with two species in cloud forest at about 9000' (2700m) at Murmur Pass, Mt. Hagen, were disappointing. The single Crested was a female-plumaged bird and the single Loria's, although a male, flew across road in front of us as we walked and much better views were desired. Had not Brian Finch been with us, I wonder whether we would have identified it. Finally, while we saw three Yellow-breasted Cnemophilus n roadside forest just past the landslide on Trauna Ridge (6000' elev. = 1800m; top photo, above), they were all immature birds. Thus, while I've ticked all three species, I have yet to have a nice satisfying view of a single male.

Taking away the three cnemophilines and MacGregor's Bird-of-Paradise (now a honeyeater) from the Paradisaeidae leaves just 38 biological species in that family, according to Frith & Beehler (1989). They may be a bit conservative in their taxonomy but wisely they do not include either Lesser Melampitta Melampitta lugubris or Greater Melampitta Melampitta gigantea in the birds-of-paradise family. Some have placed the melampittas among logrunners and allies [Orthonychidae; e.g., Beehler et al. 1986] but the evidence is growing that these two mystery birds of New Guinea are neither logrunners, nor birds-of-paradise, nor even closely related to each other. Lesser Melampitta could be a babbler (see Frith & Beehler 1990) or in some other group (even its own family?) at the base of the corvoid tree (e.g., Sibley & Ahlquist 1990). Greater Melampitta may be related to pitohuis [in the Whistler family, Pachycephalidae] because it, like some pitohuis, seems to be poisonous to eat (e.g., see Frith & Beehler 1990). While I now elevate cnemophilines to family status, it is possible that there are other endemic New Guinea bird families yet to be defined.

Photos: The photo of Trauna Ridge, Papua New Guinea, and the habitat of Yellow-breasted Cnemophilus was taken on 27 Oct 1983. As noted above, the artwork of the cnemophilines is © William T. Cooper from Frith & Beehler (1998), and is used pursuant to "fair use" doctrine. All photos © D. Roberson.

Bibliographic notes

Family book: Rating HHHHN
Frith, C. B., and B. M. Beehler. 1998. The Birds of Paradise. Oxford Univ. Press, Oxford.

This is a superb family book by two of the world's leading experts on the family. As discussed above, they (and all previous family books on BOPs) include the cnemophilines among the birds of paradise. This book appeared two years before the new biochemical evidence showing cnemophilines to be unrelated to BOPs. That aside, this is a wonderful book. Both authors studied BOPs in the field extensively, and this book is up-to-the-minute on what is known about behavior and ecology (as to 1998). It is in the Oxford Press's Bird Families of the World series which is becoming far and away the best series of family books. This entry is a 600 page tome and outstanding in every way except one, which is why I give it 4 1/2 stars instead of 5: the plates (beautifully done by William Cooper) are painted  in "field guide" style (a few have wonderfully mossy branches on which to perch). Perhaps it is just me, but I would much prefer individual full-page plates of each bird in situ, showing habitat and behavior, rather than static field guide poses on a white background. In that sense Cooper & Forshaw has better art. But this text is full of maps, sonograms, and line drawings of behavior, almost making up for this one small deficiency. Their taxonomic conclusions are a bit more conservative than many recent authors (e.g., they do not split the riflebirds into quite as many species as most), but the text does contain detailed justifications for their decisions.
Family book: Rating HHHN
Cooper, W. T., and J. M. Forshaw. 1977. The Birds of Paradise and Bower Birds. Collins, Sydney.
For its time, this was a great book. I have treasured it, and each of the spectacular full-page paintings of the birds and their habitat is etched in my memory. The text was about as good as one could get at the time, relying heavily on Gilliard (1969), but it is quite dated. Now one enjoys this book primarily for the artwork.
Literature cited:
Beehler, B.M., T.K. Pratt, and D.A. Zimmerman. 1986. Birds of New Guinea. Princeton Univ. Press, Princeton, N.J.

Bock, W.J. 1963. Relationship between the birds of paradise and the bowerbirds. Condor 65:91-125.

Coates, B.J. 1990. The Birds of Papua New Guinea. Part II. Dove Publ., Ltd., Alderley, Australia.

Cracraft, J. 1992. The species of the birds-of-paradise (Paradisaeidae): applying the phylogenetic species concept to a complex pattern of diversification. Cladistics 8:1-43.

Cracraft, J., and J. Feinstein. 2000. What is not a bird of paradise? Molecular and morphological evidence places Macgregoria in the Meliphagidae and the Cnemophilinae near the base of the corvoid tree. Proc. R. Soc. London B. 267:233-241.

Gilliard, E.T. 1969. Birds of Paradise and Bowerbirds. London: Weindenfeld & Nicholson.

Mayr, E., and E.T. Gilliard. 1954. Birds of central New Guinea. Bull. Amer. Mus. Nat. Hist. 103:311-374.

Sibley, C.G., and J.E. Ahlquist. 1990. Phylogeny and Classification of Birds: A Study in Molecular Evolution. Yale Univ. Press, New Haven, CT.

Stonor, C.R. 1937. On the systematic position of the Prilonorhynchidae. Proc. Zool. Soc. London, 107, ser. B:475-490.

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