AT Reprinted from: "Collected Papers in Avian Paleontology Honoring the 90th t^ • Birthday of Alexander Wetmore," Storrs L. Olson, editor, 21 May 1976. j#

Oligocene Fossils Bearing on the Origins of the Todidae and the Momotidae (Aves: Coraciiformes)

Sfon-f /,. Off on

Now, a recently discovered fossil and a new inter- pretation of one of the first fossil birds to be de- ABSTRACT scribed, provide us with increased information bearing on the evolution and geographic origins of A new genus and species of tody, f okeofoduf ernyyt, is described from the "middle" Oligocene (Orellan both the Todidae and the Momotidae. land mammal stage) of Wyoming, providing the ACKNOWLEDGMENTS.—I am indebted to Robert J. first record of the modem family Todidae out- Emry for calling the Wyoming specimen to my at- side the West Indies. The fossil bird frofomif tention, permitting me to work on it, and pro- gbrmffwis from the lower Oligocene of Switzer- viding much information and assistance. The land is removed from the Alcedinidae to the manuscript has had the benefit of his comments Momotidae to provide the first occurrence of the latter family outside the New World. The and those of John Farrand, Jr., Alan Feduccia, and Todidae and Momotidae appear to be more closely Pierce Brodkorb. Robert W. Storer (University of related to each other than either is to any other Michigan Museum of Zoology) kindly lent me a family of Coraciiformes. The Momotidae were evi- skeleton of Hylomanes for study, and casts of Swiss dently derived from the Old World. The Todidae fossils were generously lent by Malcolm C. appear to have been derived from a momotid-like ancestor in the Oligocene or earlier. The present McKenna (American Museum of Natural His- distribution of these two families in the New tory). I am grateful to L. B. Isham for his skillful World tropics is relictual. The Coraciiformes ap- illustrations accompanying this paper and to Anne pear to have been one of the prevalent groups of Curtis for rendering Figure 3. small land birds in the Oligocene. An Oligocene Tody from Wyoming Introduction In June of 1972, Dr. Robert J. Emry of the Smithsonian Institution collected several blocks of The five modem species of todies (Todidae), matrix containing great concentrations of bones of endemic to the Greater Antilles, are among the small vertebrates from a deposit of Orellan age most intriguing birds of the West Indies. The Mo- ("middle" Oligocene) in east-central Wyoming. motidae of Central and South America and the Present in these samples are the abundant re- Todidae are the only families of Coraciiformes mains of at least two species of squirrels, various confined to the New World. Apart from late Pleis- smaller rodents, small marsupials, and insectivores. tocene remains of modern species, there has That this great concentration of bone may be at- hitherto been no fossil record of either family. tributed to the work of owls is virtually certain Aon* JL Okm, D*paffm*f*f o/ Kgrt*6nUg Zoofogy, Afaffonaf since the blocks also contain the beautifully pre- Museum of MUwnzf ffbfory, Smfffuonian /njfffWfon, Woj/i- served skeletons of at least four small owls, possibly ingfon, DC. 20W. of two species.

Ill 112 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY

On the surface of one of these blocks, Dr. Emry tional Museum of Natural History, Smithsonian noticed the bill of a small bird and later found a Institution, USNM 205608. Collected in NW 1/4, crushed avian cranium that fit perfectly with the SE 1/4, Sec 27, T32N, R.71W, about 5.6 bilL After the two portion* had been reunited, km SSE of Douglas, Converse County, Wyoming an incomplete avian humerus was found attached (42°42'55"N; 105*21'15"W) on 12 June 1972 by to the lower surface of the skull. There is no rea- Robert J. Emry (Field No. WYO. 72-246) and son to believe that the humerus and skull are not Leroy Glenn. from the same individual. Were they not, the owl HORIZON.—Brule Formation, Orellan land- that cast the pellet containing these remains would mammal stage, "middle" Oligocene. have had to eat two different birds at the same PARATYPE.—Somewhat distorted right humerus lime, since it is rare for elements from a single prey with the shaft crushed and lacking the distal end; item to be found in two successive pellets (Rao same number and data as the holotype. zyriski and Ruprecht, 1974). This would seem un- MEASUREMENTS.—Overall length of skull as pre- likely, particularly in view of the absence of birds served 34.5 mm, length of bill from anterior of nos- other than owls in the remainder of the samples. tril 10.0, length of mandibular symphysis 7.8, width The distinctively shaped, Battened bill of the of mandible at beginning of symphysis 5.6, width fossil was immediately reminiscent of the Todidae, of mandibular ramus 1.7, proximal width of hu- but since bill: of similar shape have evolved inde- merus 6.7. pendently in several groups of birds, many of them ETYMOLOGY.—After Dr. Robert J. Emry, the col- passerines, identification was made cautiously. lector, in recognition of his significant contribu- After careful comparisons, I concluded that this tions to our knowledge of the Oligocene fauna of specimen is indeed referable to the family Todidae. North America. DzsatipnoN.—Bill flat, broad, nearly spatulate, with a broader more rounded tip than in Todtw. Palaeotodus, new genus Intemarial bar long, slender, terete, and somewhat TYPE.—fl@&%60(od%j entry i, new species. heavier than in Todus, continued out the rostrum DIAGNOSIS.—Similar to modem Todua but with as a slightly elevated ridge. Mandible flattened, the different proportions, the wing apparently being symphysis shorter and broader than in Todus; better developed. Bill proportionately shorter and mandibular rami deeper than in Todtw, grooved broader, not as pointed as in Tbdtw. Mandibular anteriorly to form a dorsal shelf. Ventral surface rami not as flattened, the anterior portions of interorbital bridge with three ridges, the middle grooved, so as to form a distinct dorsal shelf. Three one terminating in a pointed process (tip broken ridges on the ventral side of the interorbital bridge off in the type), the outer ones flaring laterally to separate, rather than coalesced as in Todus. form the edges of the huge anterior cranial fenes- ETYMOLOGY.—Greek, pa&was, ancient, plus To- tra. The condition in Todus is essentially similar, dus, the genus of modem todies. See Newton but the three ridges are not as distinct anteriorly (1896:970, footnote) for the etymology of Todtw. and coalesce to form a narrower, deeper interor- bital bridge. The middle process in Todua is a

Palaeotodus emryi, new species Ficuar 1.—Skull (holotype) and humerum (paratype) of Pafwofodw emryf, new genua and apede: (USNM 205608), FIGURE 1 compared with the mame element* of TVxftu MiWWw (USNM 292589): a, dona! view of Anil of P. **wyf; &, HoLoTYPE.—Incomplete and partially crushed doraal view of skull of T. fubwfsfw; c, lateral view of mkull skull with most of the anterior portions of the ros- of P. *mfy<; d, ventral view of mandible of P. «myf; «, /, trum intact, including the mandibular symphysis cutaway views of ventral side of interorbital bridge and dor- and parts of both rami; crushed posterior portion sal part of anterior cranial fenestra of T. subulatus; g, h, of cranium with ventral surface of interorbital same views of P. emryi; i, anconal view of humerus of P. emryf; /, proximal end of humen* of P. «mryf, viewed with bridge well preserved; much of the rest of the skull distal portion tilted further upward; k, proximal view of crushed, jumbled, and displaced ventroanteriorly. lame; f, anconal view of humerum of 7*. #i*&**Wuj. (All Ggure: Vertebrate Paleontological Collections of the Na- approximately X 3.) NUMBER 27 113 114 SMITHSONIAN CONTRIBUTIONS TO PALZOBIOLOGY point of attachment for a narrow ligament that bulbous, the capital groove much deeper, and the joins with similar ligaments from the parasphenoid internal tuberosity less perpendicular to the shaft rostrum and the lateral edges of the cranial fenes- than in Pafagofodtw, although these differences are tra to form a tenuous cross holding the anterior considerably less pronounced in the Galbulidae and part of the brain in place. The humerus of PoLwo- Bucconidae. Within the Coraciiformes the humeri fodwj is similar to that of Todwj, but is much of the Upupidae, Phoeniculidae, and Meropidae larger; the internal tuberosity is heavier and much have larger, more triangular deltoid crests than in less lateromedially elongate; and the ligamental PaJaeofodiw, while in the Coraciidae the bicipital furrow and the scar on the external tuberosity are crest is more extensive. The humerus in the both considerably deeper. Alcedinidae has the head more bulbous and situ- REMARKS.—Few nonpasserine landbirds have the ated much higher above the external tuberosity, bill anywhere near as flattened as in Todus and the shaft straighter, the internal tuberosity much Pairofodu*. Those with the most Battened bills arc heavier, and the bicipital surface much less pro- JSkcffon, ffyfomawes, and j4jpo(Aa (Momotidae), duced than in Palaeotodus. The humerus of /acanwzfo/M (Calbulidae), and Myioceyx (Alce- Pa&Mo(odtw is most similar to that of the Todidae dinidae). In all of these forms the bill is deeper and and the Momotidae. In the conformation of the the intemarial bar broader and shorter than in internal tuberosity it is more similar to the Momo- either of the two genera of Todidae. The bony tidae, whereas in the greater excavation of the ex- structure of the bill in passerines, as for example in ternal tuberosity and ligamental furrow it more Todxroffrwyn (Tyrannidae), is quite different from closely resembles the Todidae. that of TWwf and PoZaeofoduj, being deeper and The manner in which the skull was crushed in more arched, with a more troughlike mandible and the type of Pa&wofodwj makes it appear smaller, a shorter, wider, and less terete intemarial bar. while the distortion of the humerus is such as to Pabrofodtw agrees with Todw and differs con make it appear wider and thus larger. Nevertheless, spicuously from the Momotidae, Alcedinidae, Gal- it is quite evident that the proportions of Pa&zeofo- bulidae, and indeed from all of the rest of the dus are different from those of Todus, the wing Coraciiformes and Piciformes, in having the ante- being much larger in relation to the head. rior wall of the cranium and the interorbital sep- This difference in proportions may be due at tum unossiGed. In the Momotidae, Alcedinidae, least in part to the small size of Todwr being aec- and Calbulidae the interorbital septum is partially ondarily derived, since the species of this genus are or very heavily ossified. The anterior wall of the the smallest members of the order Coraciiformes. cranium is ossified in all of these families, whereas In the evolution of vertebrates, body size usually in Todws and Pobfofodur there is a great, open changes at a more rapid rate than head size, so that fenestra. small forms derived from larger ones tend to have The humerus of Pa&zeofodiw is broken and dis- proportionately larger heads, and vice versa. Many torted, with the head crushed down distally relative authors have remarked on the large-headed appear- to the internal and external tuberosities. Although ance of Todtw in life. In the Oligocene, the Todi- in Todwj there is slightly more of a depression be- dae were possibly more diverse than at present and neath the head than in other Coraciiformes, the probably included larger, more actively flying humerus can in no sense be regarded as having a forms with better developed wings than the strictly double fossa, as stated by Bock (1962), and has a sedentary modem todies. single pneumatic opening beneath the internal tuberosity. Pobgofodiw is similar to T"od#w in this An Oligocene Motmot from Switzerland respect. The humerus of Pafoeofodus differs from that of the Passeriformes in the less bulbous head, In 1839, von Meyer called attention to the re- the much lesser development of the bicipital crest, mains of what he thought to be a passerine bird the higher and more pronounced external tuber- from slate deposits (Glamer Fischschiefer) in osity, and the narrower, more ventrally projecting Switzerland, then considered to be of Cretaceous internal tuberosity. In the Piciformes the deltoid age. In a subsequent publication he named this crest is much more expanded, the head more specimen ProforMw g&zrmffww (von Meyer, 1844). NUMBER *7 115

Later (von Meyer, 1856), he emended the name to somewhat shorter than in the modem genera of P. gfaromfnjw and this spelling was in general use Momotidae, being most similar in this respect to until Brodkorb (1971), whom I have followed, re- ffyZomwma;, which genus is generally conceded to vived the original orthography. Lambrecht (1933) be the most primitive of living motmots. maintained Preform** as a genus incertae sedis in In the mandibular articulation of Profoynir the the Passeriformes. At that time the deposits from internal process is a long, thin splint set off from a which the type off, gfamimww was derived were well-developed retroarticular process by a distinct regarded as upper Eocene in age. Subsequent notch, with the actual articulating surface for the studies have shown them to be of lower Oligocene quadrate much reduced. This is exactly the condi- age (Peyer, 1957). tion seen in the Momotidae and Todidae. In the The type of P. gfamifwu consists of a slab con- Alcedinidae the articular cup for the quadrate is taining the bones of all four limbs, the pectoral large and deep, the retroarticular process virtually girdle, a complete mandible, the quadrates, and a absent, and the internal process wide, heavy, and few vertebrae and ribs. These were insufficiently triangular, quite unlike Protornis or the motmots exposed when von Meyer studied them, but Peyer and todies. Bee-eaters, Meropidae, have a fairly (1957) undertook further preparation of the type, long, slender internal process, but it is not set off illustrating his results with numerous photographs from the retroarticular by a notch, and the articu- and x-radiographs. The fossil is slightly distorted lar cup is deep, as in kingfishers. Furthermore, the from stresses imposed on the rock after deposition; bill shape of Profomir is not at all like that of the StOssi (1958) and Baumann (1958) have offered Meropidae. mathematical and optical corrections, respectively, The shape of the hypotarsus in Profomi; is ex- for this distortion. Another less complete specimen actly as in motmots and differs from that of the was referred to Profomif, possibly gfamtfurw, by kingfishers, in which it projects above the cotylae Peyer (1957). I have had access to casts of both in a distinct point. The tarsometatarsus of Pro- these specimens, as well as to Peyer's excellent illus- (omw is only slightly shorter than the middle toe trations. The casts were made by a copper electro- with claw, as in the motmots. In todies the tarsus plate process and appear to be very accurate rep- is longer than the middle toe with claw, whereas in resentations of the original specimens. the kingfishers and bee-eaters the tarsus is squat After his study of the type of P. gbrmeww, and much shorter than the middle toe. The pro- Peyer (1957) concluded that it belonged with the coracoid process appears to be nearly absent in Alcedinidae (kingfishers) and more particularly Protornis, as in motmots and todies, whereas it is that it was nearest to Docefo. I agree with the as- better developed in kingfishers. signment of this form to the Coraciiformes, but From the evidence detailed above it is clear that numerous characters of its skeleton show conclu- Protornw does not belong with the Alcedinidae, sively that Pfofomis cannot be a kingfisher. where Peyer (1957) placed it. The proportions of Profomif gfamiffms is a small bird, slightly the bill and of the hindlimb and toes preclude its smaller than the modem motmot ffyfomwmff assignment to the Todidae. In all of its important momotwfa. As detailed by Peyer, many aspects of features it agrees with the Momotidae. It differs its structure demonstrate that it does not belong from the modem forms of the family mainly in the with the Passeriformes and the clearly anisodactyl shorter mandibular symphysis and the higher, more feet eliminate the Piciformes from consideration. expanded stemocoracoidal process of the coracoid. The mandibular symphysis is broad, Battened and Profomir gbrmxenfir should, therefore, be assigned somewhat spatulatc, differing from most non- to the family Momotidae. passerine Iandbirds except the Momotidae and A second fossil from the Glarner Fischschiefer, Todidae. The overall conformation of the man- consisting of a slab with both hindlimbs, the right dible is in fact, markedly similar to that of the wing, some ribs, and portions of the pelvis super- motmot genera Ekcfron and ffyfomanas and is imposed on the sternum, was assigned to the genus quite distinct from that of the kingfishers, includ- ProfoTTiw by Peyer (1957), who suggested that it ing the fiat-billed genus Myioceyx (Figure 2). The might, be referable to the species P. gZomgeww. symphysis is broader than in the Todidae and This is plainly impossible, for the second specimen 116 SMTHSONIAN CONTRIBUTIONS TO PA1ZOBIOLOGY

FIGURE 2.—Dorsal views of mandibles: a, Electron platyrhynchum; b, Protornis glarniensis (adapted from Peyer, 1957); c, ffyfomanw fnomofwb; d, Todw* JutuWia; c, Mykc«y* kconW; /, Dacelo novaeguineae. (Not to scale.) is much too large to belong to the same species as, found only in the Trogonidae among modern birds. P. g&zmigfMw; the carpometacarpus, for example, The rest of the skeleton of this specimen is gen- is twice as long. Furthermore, in the right foot of erally similar to that of modern trogons, although this specimen it can be clearly seen that both digits differing in some details. This specimen deserves a I and II are reversed (Figure 3)—a condition great deal more attention since it provides the NUMBER. 2? 117

Discussion

The ten families of the order Coraciiformes fall into several diverse groups whose relationships within and without the order are as yet uncertain. Sibley and Ahlquist (1972:230) maintained that, "no compelling evidence exists to ally any group of the Coraciifdrmes more closely to a non- coraciiform than to other members of the Coracii- formes." On the basis of biochemical analysis of egg-white proteins, Sibley and Ahlquist concluded, as have other taxonomists in the past, that within the Coraciiformes, the Alcedinidae, Todidae, Mo- motidae, and Meropidae appear to form a natural but distantly interrelated group. Feduccia (1975) discovered that these families possess a highly de- rived stapes, which is shared only with the Trogon- idae, and concluded that all five families are closely FIGURE 3.—Diagram of right foot of the so-called second related. Contrary to most earlier opinions, Sibley specimen of Protornis showing the heterodactyl condition and Ahlquist (1972:230) suggested that the Todi- typical of the Tiogonldme. (Adapted from Peyer, 1957, and a dae are more closely allied to the Alcedinidae than cast of the specimen; the distal portion of the fourth toe is to the Momotidae. The osteology of these families present as an impression in the matrix.) does not support this contention, and along with their distributional history strongly indicates that a fairly dose affinity exists between todies and mot- mots and that these families differ considerably earliest evidence of the occurrence of the hetero- from the kingfishers. dactyl foot. It obviously cannot be assigned to Pro- Seven of the ten families of Coraciiformes are fof»ir or the Momotidae and for the present should confined to the Old World. Of approximately 89 be regarded as belonging to the Trogonidae. Four Recent species of Alcedinidae, only six, in two fossil species of trogons are known—all from France genera, are found in the New World. Of these, two (Brodkorb, 1971). Three of these, in the genus are in the genus Ceryle, which also contains two i4rc/w%60ffogom, are from the Phosphorites du Old World species, while the genus CAfofoaaylg, Quercy, which range in age from upper Eocene to which is only weakly differentiated from Ceryk, lower Oligocene, and are thus possibly contempor- contains four species endemic to the New World. aneous with the Swiss specimen. The fourth spe- Clearly the kingfishers are an Old World family cies, farafrogon gafficw, is from lower Miocene that has only recently invaded the Americas. Thus, (Aquitanian) deposits at Langy. the Todidae and Momotidae are the only truly A second species of Profornif, f. oiwmrn, was New World families of modern Coraciiformes. described from the Glarner Fischschiefer in 1865 Recent motmots are neotropical in distribution, by Heer (1876). The type appears to have been ranging from southern Mexico south through Bra- poorly preserved and has not been restudied, its zil. In a classical exercise in zoogeography. Chap- wherealxmts l)eing unknown. Brodkorb (1971) man (1923) analyzed the distribution of the genera, placed this specie*, along with P. gfafmenw, in the species, and subspecies of motmots, concluding that AJcedinidae, but considered its position uncertain. they had originated in Central America, the few From the original illustration one cannot even as- South American forms having been derived from certain that the specimen was avian. In view of the north. Lonnberg (1927), noting that Central this, and since more than one family of birds occurs and North America probably presented a more or in the Glamer deposits, P. Wurmen should be rele- less continuous tropical environment in the Ter- gated to the category of Aves incertae sedis. tiary, felt that the motmots could as easily be con- 118 SMITHSONIAN CONTRIBUTIONS TO PALZOBIOLOGY sidered North American in origin. A North or Momotidae, presently confined to the New World, (Central American origin of the modem members actually had its origins in the Old World. The of the family, as opposed to a South American one, place and time of origin of the Todidae are un- has properly gained general acceptance. Neverthe- certain. The Orellan land-mammal stage repre- less, this leave* unanswered the question of the sents a geologically very short span of time follow- origins of ancestral motmots. ing the much longer Chadronian stage and pre- The modem todies, confined to the West Indies, ceeding the Whitneyan stage, the latter also are also thought of as being North American in representing a very short span of time. The de- origin. This is due in part to their presumed ret posits from which Pa&wofoduj was recovered are lation&hip with the motmots and in part to the about 30 million years old and have traditionally North American derivation of most of the rest of begn regarded as middle Oligocene, although there the Antillean avifauna (Bond, 1966). It might then is as yet no paleontological correlation between the be asked whether the todies evolved their distinc- North American terrestrial deposits of so-called tive characteristics from some momotid-like ances- Oligocene age and those of Europe. By the reduced tor after arriving in the Greater Antilles, or had ossification of the skull, Palaeotodus certainly already assumed these characteristics before colon- seems to be referable to the Todidae rather than izing the islands. the Momotidae, but perhaps with material from Because of the small size and feeble flight of earlier in the Oligocene it would not be possible modem todies, Bond (1966) expressed reservations to distinguish the two families, the family Todidae about their ability to cross even narrow water gaps having assumed its characteristics since that time. and suggested that they might have been rafted to Probably through a combination of climatic the West Indies from Central America. However, change and competition with more advanced land because the five species of modem todies are very birds, the motmot-tody group was entirely sup- similar in plumage and morphology, one must as- planted in the Old World. The deterioration of sume that members of the genus Todus have been tropical conditions in North America in the late able to cross the water barriers between the Greater Tertiary left motmots only in Central America, Antilles within relatively recent geologic time. from whence they have spread into South America Moreover, since there are two species of Todus on since the closing of the Panamanian seaway in the Hispaniola, one must either assume sympatric late Pliocene. Similar factors affected the North speciation or a double invasion of the island. American todies and only the isolated West Indian Bond's own remarks (1974) on the greater similar- relicts of the genus Todus have survived up to the ity of the voice of Hiapaniolan 7*. awgur fmwfrw to present. that of Cuban T. multicolor than to that of the Feduccia and Martin (p. 110, herein) have shown other Hispaniolan species, 7*. j%6w&%*wj, suggest a that the predominant order of small land birds of double invasion. Thus, if todies were able to cross the Eocene in North America was the Pidformes. the water barriers between the islands of the It is now becoming evident that the Oligocene was Greater Antilles they might as easily have crossed similarly important in the evolution of the Coracii- from the mainland. Furthermore, the evidence pro- fbrmes. Although the evidence is far from con- vided by PoZofofodua shows that in the Oligocene, clusive, if the Coraciiformes (including the Tro- todies were larger and had proportions suggestive gonidae) were not the predominant perching land of greater powers of Hight. It therefore seems pos- birds of the Oligocene, they were certainly much sible that todies might have colonized the West more prevalent than today. Recently I have ex- Indies over water as easily as, say, kingfishers, and amined a number of fragments of small land birds it is not necessary to invoke rafting to explain their of Chadronian and Orellan age from the western present distribution. United States. All of these appear to be referable Without doubt, the order Coraciiformes, as tra- either to the Coraciiformes or Pic:formes and defi- ditionally conceived, arose in the Old World. The nitely are not passerine. Thus, it would appear that existence of Profonw in the lower Oligocene of the Passeriformes may not have gained a strong Switzerland now provides evidence that the family foothold in North America until the Miocene. NUMBER g? 119

Literature Cited

Baumann, E. Lonnberg, E. 1958. AQine Entzerrug mic einfachen opdachen Mitteln. 1927. Some Speculations on the Origin of the North gtAxLafzarkc/ia PaUkonfofogbcAa ^bhandhmgan, American Ornithic Fauna. JTungWga &/en*Aa Pen 78:17-21, 4 Agnrem. [Separate] fenjAapaAademieru ffandffngar, series 3, 4(6):H(4 Bock, W. J. von Meyer, H. IRK. The Pneumatic Foam of UK Humeru* in the 1839. Bin Vogel im Kreideschlefcr des Kantcms Gfarff Passere*. ^wJk, 79(3):425-443, 2 Ggurem. ATeue* JaArbwcA /Or Mfnerafogfe, Geognnik, Gao! Bond, J. ogfa und Peffe/dAfer^unde^ 1:683-685. 1966. Afhnlties o! the Antillean Avifauna. C*fMi6«an 1844. [Letter.] #eues /oArbucn /Ar Mfnefafogfe, Geog /ournaf o/ SckncP; 6:179-176. »w:a, Geofogfe und Pefre/aAfcnAwnd*, 6:529-540 1974. MnafaanfA g«^km«n* fo (he CAad^fff* a/ Bfrdj 1856. Schildkrote und Vogel aus dem Fischschiefer von of the West Indies (1956). 12 pages. Philadelphia: GImrum. PafaaonfogmpVWca, 4(3):83-96, 2 plates. Academy of Natural Sciences of Philadelphia. Newton, A. Brodkorb. P. 1896. A Dictionary of Birds. 1088 pages. London: Adam 1971. Catalogue of Fossil Bird:, Part 4 (Columbiformes and Charles Black. through Pkdformem). Buffafdn of fha fforkfa ffafa Peyer, B. Muwum, Bfofogfcaf Sckncgf, 15(4): 163-266. 1957. Prof ami* gfaronanw H. v. Meyer Neubeschrelbung Chapman, F. M. des Typusexemplares und eines weiteren Fundes. 1923. The Distribution of the Motmotm of the Genus &cnwefzerfscAen PaMonfofogfjeAen ^IbAandZungan, Momota. Buffatfn of fha idmefiaan Mwawms of 73:1-47, 26 Ggures, 11 plate*. ^afwraf ffWo^, 46:27-59, 4 figures. Feduccia, A. Raczyriski, J„ and A. L. Ruprecht 1975. Morphology of the Bony Stapes (Columella) In the 1974. The Effect of Digestion on the Osteological Com Passeriformes and Related Group*: Evolutionary poddon of Owl Pellets. Xcfa OmffAof(%fca, 14(2): Implications. T&a f/nwarnfy o/ f anja* Afwaum of 1-36. Nafwnaf Hfsfoyy Mwaaffanaou* PubKaaffon, 5:1-54, Sibley, C. G., and J. E. Ahlquist 7 figure;, 16 plate*. 1972. A Comparative Study of the Egg White Proteins Hear, O. of Non-Passerine Birds. Peabody Museum of Nat- 1876. 7*ka Pnmaewaf TPorfd of Sswttarfand. Volume 1. umf g&fory Fafe f/mfoenffy Bwfkffn, 39:1-276, 37 London: Longmans, Green and Co. [English trans- Ggures. ladoa of IX* Uhaaf* der ScAwefz, Vidume 1. ZOrich, Stussi, F. 1856 (not seen).] 1968. Die Entzermng voo FomslUen am Beisplel de* Lambrccht, K. AvfofWs gfaronanjf*. SchaafzarbcAen PaMonfofb- 1933. Handbuch der Palaeornithologie. xix + 1024 pages. gischen Abhandlungen, 73:1-16, 13 figures. [Sepa- Bedln: GebrOder Bomtrager. rate.]