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https://doi.org/10.24199/j.mmv.1977.38.04 4 April 1977

A NEW OWLET-NIGHTJAR FROM THE EARLY TO MID-MIOCENE OF EASTERN By Pat Vickers Rich and Allan McEvey National Museum of Victoria, Melbourne

Abstract A new owlet-nightjar (Family: Aegothelidae), Quipollornis koniberi gen. et sp. nov., has been recognized from Miocene lacustrine sediments in eastern New South Wales. It thus establishes the occurrence of the aegothelids in Australia during the Miocene. The new form shows distinct and numerous differences from the other two genera in this family, the living Aegotheles from Australia, New Guinea, and the South-west Pacific and its closely related New Zealand counterpart, Megaegotheles, now extinct.

Introduction the coracoidal head; and its short but broad vertebrae. This combination of char- During the past 50 years diatomite deposits cervical to be unique to the Owlet- representing mid-Tertiary freshwater lakes in acters appears within the Caprimulgiformes. eastern Australia have yielded a number of nightjars well-preserved leaves and fish. Among these Quipollornis gen. nov. collections is a single, unique skeleton of a Type Species: Quipollornis koniberi gen. et small bird closely related to the living Owlet- sp. nov. nightjars {Aegotheles) nocturnal birds en- , Age and Distribution: Early to Middle Miocene, demic to, but with wide dispersion in, Aus- eastern New South Wales, Australia. tralia, New Guinea, and the South-west Etymology: Quipolly (Aboriginal), a water- Pacific. The Miocene fossil, Quipollornis hole containing fish; ornis (Greek), Bird. Re- koniberi gen. et sp. nov., is the oldest record ferring to the occurrence of Quipollornis in of the family Aegothelidae, which is repre- lacustrine sediments containing mainly fish. sented by only one other genus, the extinct Diagnosis: Differs from Aegotheles in that the Megaegotheles (Scarlett, 1968; Rich and Scar- jugals are broader and stouter bones, and the lett, 1977) from the Quaternary of New Zea- maxillaries do not expand as much medio- land. This paper describes the new form and laterally towards their posterior termination. compares it with all other caprimulgiform Differs from both Aegotheles and Megaego- families. theles in that the head of coracoid is not as In this study the following forms were avail- inflated dorsally and is decidedly more elongate able to us for comparison: Steatornis caripensis between the coraco-humeral surface and pro- Nyctibius griseus Podargus strigoides (1), (3), coracoid; the scapula appears to lack a marked Aegotheles cristatus (9), Eurostopodus (20), flexure near the posterior end as well as expan- guttatus E. mysticalis (3), and Chordeiles (5), sion of the posterior end present in Megaego- minor ) ( 1 theles] the breadth of the scapula just anterior Systematics to the glenoid facet is somewhat greater than in Aegotheles, similar to that area in Megaego- Class: Aves theles; the deltoid crest is intermediate in its (Ridgway) Brod- Order: Caprimulgiformes dorso-lateral protrusion between Megaegotheles korb, 1971 and Aegotheles; the metacarpal I process is Family: Aegothelidae (Bonaparte) lower and broader, and metacarpal III more robust. Differs from Megaego- Comment: Placement of Quipollornis koniberi markedly theles in that the shaft (viewed ventrolaterally) gen. et sp. nov. in the Aegothelidae is based is not as highly curved. primarily on its broad skull with slender-boned Ratios (see Table 1). The humerus about lower and upper jaws and rounded, inflated twice the length of femur unlike that of braincase; its highly curved internal margin of

247 i

248 PAT VICKERS RICH and ALLAN McEVEY

TABLE I: Measurements (in mm.) of Skull, Shoulder Girdle, and Limbs

of Quipollornis koniberi, n. gen. et n. sp. and Other Caprimulgiforms,

Specimens Podargus Eurostopodui Eurostopodus Aegotheles Steatornis Nyctibiui Quipollornis Megaegotheles Measurements strigoides mystacalis guttatus cristatus caripensis griseui koniberi novaezealandiae AMF49404-5

Skull

Width 49.3-55.4(6) 30.0 24.8(1) 22.8 23.6 34.4 43.1-47.2 30.8 26.0-27.0 4 Length 81.5-88.7 52.6 £44.0-48.4 35.4-38.4 60.1 56.8-65.1 40.8 Width/Length 0.58-0.68 0.57 0.51 0.61-0.65 0.57 0.73-0.76 0.75 n 7(6) 1 2(1) 6 3 I Scapula Length 38.2 43.7 32.8 34.4 26.4-27.9 19.0-23.4 46.2 34.9-39.2 >2I2 >23.3 Maximum width, head 9.6- 11.3 5.3- 6.7 4.8- 5.6 4.2- 5.1 8.8 7.4- 7.5 5.3 5.6 n 8 3 3 8 1 I Coracoid Length of head' 9.0- 9.5 6.9- 7.7 6.4 7.0- 9.2 4.( 4.7 Width across 2 head 4.5- 5.5 3.9- 5.7 1.3- 2.0 7.4 3.6- 5.3(2) 2.E 2.1 n 3 2 7 3(2) Humerus

Length 76.0- 81.7(6) 53.2-54.7 43.5-47.2 4 Maximum 59.8-67.3 ©35. 34. 28.3 36.3(20) width proxi mal 3 end 17.4- 21.6 12.9- 18.1 12 9 14.6 8.1 - 9.1 20.9 16.8-20.2 @I0.0 @l 1.2 6.8- 9.0(20) Maximum w idth distal end 13.7- 14.6 9.8- 10.1(6) 7.8 8.4 5.6- 6.2 14.3 10.7- 12.9 @ 7.1 5.3- 7.0 n 7 16) 3 3 3 1 22(20) Ulna Length 81 .7- 96.8 67.4 67.8 57.4 63.0 Maximum 63.4-72.0 est. 44.5 32.5-39.3 diameter, external condyle 6.9- 7.4 5.0- 5.8 4.0- 4.5 2.7- 3.1 5.8- 6.9 3.3- 3.4(3) 3 3 8 3 4(3) Carpometacarpus Length from head to distal end of inter- metacarpal space 32.0-36.6 30.6-31.3 15.2- 16.7 37.6 Minimum width of

metacarpal II 2.0- 2.5 Length of intermeta-

carpal space 19.8-24.1 21.2-21. 19.8 20.6 14.8- 18.6 8 3 3

Carpal Phalanx I Length 8.6- 12.2 5.6

Femur Length 40.2-43.1 28.5 29.2 20.5-22.0 35.1 27.6 >I7.5 23.0-32.0 8 3 2 Tibiotarsus Depth, proximal end 7.8- 8.8 4 5 5.4 3.7- 4 1 3.4- 4.3 7.4 5.0- 6.3 4 6.4- 6.5(2) Length 62.4- 71.2 38.6- 39.4 34.0 38.4 35.4- 38.0 43.5 35.6-42.1 j>22. 3 3 50.0-66.0+ 7(6) 3 Ratios 12(2) Humeral / Ulnar 0.84- 0.93 0.78-0.81 0.740.76 0.78-0.81 0.93-0.96 6 Length/ Length 0.78 0.87-0.92

Humeral / Femoral 1.76- 1.94 1.82- 1.80-1.84 t.26- 1.32 1.97 2.17-2.22 2.0 Lencjth/ Length 1.13- 1.23 Humeral /Skull 0.92- 0.97 1.04 0.930.99 0.72-0.76 2.01 1.02- 1.03 1.14 Length/ Width 1.09- 1.34

Humeral / Proximal 3.70- 4.12 3.33-3.65 3.5 Length,3th/ WidiVidth Humerus

Humeral/ Distal ! Length/ Width 4.92 5.19- 5.34 Humerus

FOOT NOTES:

1. Length of coracoid from dorsalmost part of head to 4 - approximately, @, !$*<{:, slightly greater or slightly less than. ventralmost part of curve between procoracoid and 5. Measurement brachial tuberosity. made by utilizing complete humerus on counterpart to project where proximal end of ulna would occur on other slab. 2. Width from distalmost extension of curve to external 6. Because none of the Megaegotheles margin of shaft. except one where definitely associated, ratios have been computed by considering minimum 3. Width across head to maximum measures expansion of deltoid crest. together and maximum measures likewise. NEW OWLET-NIGHTJAR 249

Megaegotheles and Aegotheles where humerus exceeds length on any of cervicals 3-6; in and femur are nearly subequal; the humerus is Chordeiles, width doesn't exceed length in slightly longer than the width of the skull as in cervicals 5-7; in Aegotheles, width and length Megaegotheles, but unlike Aegotheles in which are nearly the same in cervicals 3-7, with a the skull is broader than the length of humerus; resulting shorter neck than in most caprimulgi- humerus broader distally with respect to its forms in this form and in Quipollornis. length than that in Megaegotheles, although Scapula. The breadth of the scapula just quite similar to that of Aegotheles. anterior to the glenoid facet somewhat nar- the narrow Comparisons: Skull. The occipital region of rowed, intermediate between scapula of egotheles and the somewhat cranium is rounded posteriorly (convex pos- A and Chordeiles teriorly) unlike the flattened posterior edge in broader one of Eurostopodus in Podargus, Eurostopodus and Caprimulgus, the slightly but decidedly narrower than that the lateral margin concave edge in Nyctibius, and the slightly Nyctibius, and Steatornis; is highly curved as in Steatornis convex edge in Podargus and Steatornis; the of the shaft not entire scapula is not pre- dorsal surface of cranium lacks the well- and Nyctibius. (The remains for the beginning developed, paired temporal fossae that occur served, but enough of expansion to be evident if present; the left in Podargus and Steatornis. Such fossae are scapula, in fact, appears to be nearly complete.) only slightly developed in Aegotheles, are situated much more ventrally, and thus are not Coracoid. The medial margin of head evident in dorsal view; the premaxillaries, between the brachial tuberosity and the pro- maxillaries, and nasals are very slender as in coracoid is highly curved with procoracoid Aegotheles, but unlike the fully fused elements extending so far dorsally that it leaves only a tuberosity in Podargus and Steatornis that form a com- small gap between it and brachial highly pletely enclosed and continuous upper jaw or as in Aegotheles, and unlike the less the much exanded premaxillae and maxillae curved margins and broader gaps characteristic Chordeiles, Stea- in Eurostopodus or the elongate and very of Podargus, Eurostopodus, slender premaxillae that merge into a broad, tornis, and Nyctibius. nearly continuous palate (except for a slender Humerus. The bone is relatively more

slit along the midline) in Nyctibius; the exter- robust than in other caprimulgiforms; angle nal nasal opening extends further forward than formed between the shaft and proximal margin in all genera examined except Aegotheles; the of bicipital crest is small obtuse angle as in upper jaws, viewed dorsally, rapidly expand Steatornis, Nyctibius, Chordeiles, and Aego- posteriorly, closely resembling Aegotheles, but theles, but differs from Podargus and Eurosto- unlike the gradual expansion that occurs in podus; deltoid crest is not as prominent dorsally Eurostopodus, Caprimulgus, Chordeiles, and but is lower and gently rounded throughout, Steatornis resulting in a narrow V-shaped skull unlike that in Aegotheles, Eurostopodus, and or bill or in Podargus and Nyctibius that have Chordeiles; deltoid crest is also not as broadly a slightly broader V-shape. The skull is very expanded from the shaft as in Steatornis. broad proportionally, decidedly more so than Radius-ulna. The distal end of radius is of in Eurostopodus and Steatornis, but closer to much more closely appressed to distal end that of Podargus, Aegotheles, and Nyctibius, ulna than in Steatornis, Nyctibius, and Chor- A delicate, short anterior nail is formed as in deiles. Aegotheles. Carpometacarpus. Differs from other capri- that the process of metacarpal I Vertebrae (see Table 2). All cervicals pre- mulgiforms in appears to be lower and more rounded; angle served are broad with respect to length. In formed between proximal margin of carpal what are probably cervicals 4 and 7 the breadth trochlea (viewed externo-dorsally) and long across anterior zygopophyses is greater than axis of shaft is more acute; the bone is shorter vertebral length; in Podargus, Eurostopodus, never and much more robust, and metacarpal III is Steatornis, and Nyctibius, anterior width 250 PAT VICKERS RICH and ALLAN McEVEY

TABLE 2: Vertebral Measurements of Quipollornis koniberi, n. gen. »t n. (p.,

and Other Caprlmulgiforms (in millimeters).

Specimen Podargus Eurostopodus Aegothelei Steatornit Nyctlblus Quipollornis Measurement! sir igo ides guttaius cristatus caripensii griseui koinberi NMV B IQI7B NMV 10847 NMV 11033 UCMVZ 141741 UCMVZ 126575 AMF 49404-5

Cervical 4 Anterior width 8-3 5.0 50 7 2 Posterior 7.1 width 7.4 5.6 @ 3.7 7.5 6.5 80 Least width 5,0 5.6 2.0 4 2 63 Left length IL2 6.2 4.7 9.8 Right 7.5 6 7 length 11.2 62 4.7 9.6 7.6 Cervical 7

Anterior width 11.4 6 2 5.1 10.7 Posterior width 9.5 7.4 9.1 46 3.0 8.8 6.2 7.4 Least width 3.9 2 4 2.1 38 Left length 3.0 10 5 7.B 4.2 9.6 7.4 5.4 Right length 9.8 7.7 4.3 * 9.6 8.7 5.4

# a approximately

relatively stouter with respect to metacarpal II. Vertebrae preserved in Quipollornis exhibit The ventral border is slightly more bowed than a distinct constriction at the fore-aft midline. we observed in other caprimulgiforms. If cervicals 3 and 4 are represented in this Femur. When viewed laterally, the bone is series, their morphology differs from that in parallel-sided over much of its length rather Eurostopodus, Steatornis, and Nyctibius, where like all caprimugliforms except Steatornis where no such constriction occurs, but is similar to shaft margins diverge towards either end of that in Aegotheles and Podargus, bone from central part of shaft. Besides the vertebrae, many of the elements Tibiotarsus. See comment, below. discussed in the diagnosis are obscured so that Feather Impressions. Lateral to the region interpretation is often difficult and tentative. of the pectoral girdle and the cervical vertebrae The posterior ends of the scapulae are missing, are the impressions of contour feathers. These, and thus whether the scapula was curved near however, do not show normal feather shape or the end or not is unknown. Likewise, the structure. They preserve the appearance that ventral halves of the coracoids are not visible. can be achieved by wetting soft aegothelid con- The humerii are difficult to interpret because tour feathers so that the barbs in each one cling one cannot be certain of their precise orienta- tion, to the rachis, and the feather then takes on a and slight differences in perspective can narrow, produce apparent slender structure. This process is, of radical differences in shape course, consistent with the mode of preserva- of the deltoid crest and in the prominence of the external tion of the fossil specimen. Viewed in this light, tuberosity along the margin be- tween the the feathers would appear to be typical of, but head of the humerus and the deltoid rather longer than, the soft contour feathers of crest. The proximal end of the carpometa- carpus the living Aegotheles. Because of the lack of is somewhat obscured by the distal end detail of the ulna; similarly, on the fossil feathers, however, it would the femora and proximal be fruitless to carry the comparison further, or parts of the tibiotarsii are either lacking or obscured to claim that the feathers are closer in mor- with bone hash to the extent that few phology to Aegotheles than to a number of qualitative statements can be made concerning other birds. those bones.

Comment: Because only a dorsal view of the Quipollornis koniberi sp. nov. specimen has been preserved, ventral elements (Plate 8, fig. A-B) such as the sternum and more lateral ribs Holotype: Australian Museum (AM) No cannot be discerned. Dorsal elements, how- F49404 and F49405, partial flattened ever, such skeleton as vertebrae are readily visible, at or its impression in diatomite, including part least in part. and counterpart blocks that partly overlap NEW OWLET-NIGHTJAR 251

Representation on these blocks comprises a Discussion and Conclusions complete, but partially obliterated skull, 4-6 Upon first glance, the fossil bird skeleton cervical vertebrae that represent may cervicals preserved in the two diatomite blocks from 4-8 or scapulae, 10, a partial right and possibly Bugaldi appears to retain exquisite detail, but part of the head of the left coracoid, humerii, upon close perusal, it becomes evident that the partial radii ulnae, the left and carpometa- only parts distinct enough for taxonomic use carpus alulu, partial and right and left femora are the skull, the cervical vertebrae, part of the and tibiotarsii, possibly the anterior part of the shoulder girdle and forelimb. Other bones in synsacrum, as well as a number of feather the skeleton do not contradict the assignment impressions that suggest feathers were wet when of this form to the Aegothelidae, but, likewise, bird preserved, with barbs closely adhering to do not provide any qualitative characters that the rachis. can be used to distinguish Quipollornis from Type Locality and Age: Diatomaceous earth other members of the family. deposit at Chalk Mountain, Bugaldi, near Within the Aegothelidae, Aegotheles and , Warrumbungle Mountains, Megaegotheles are very similar to one another eastern New South Wales; Early to Middle (Scarlett, 1968; Rich and Scarlett, 1977) and Miocene. together form a subunit that is distinct from Quipollornis. differences between Measurements: See Tables 1 and 2. The main the two groups are in relative size of the jugals, Etymology: Koniberi (Aboriginal), name of shape of the head of the coracoid, shape of the tribe that once inhabited the geographic area scapular shaft, shape of the metacarpal I from which Quipollornis was recovered. process on the carpometacarpus, and the Diagnosis: as for genus. Same relative lengths of the humerus and femur. Because of the difference, the forelimb seems Stratigraphic Position to have been decidedly more elongate in Diatomite deposits from the Coonabarabran Quipollornis than in other aegothelids (see area in New South Wales have long been Table 1), characteristic of the non-aegothelid known to produce fossil leaves and fish (Kenny, caprimulgiforms, and thus the marked emphasis 1924), the fish being studied some time ago placed on hind limbs in the Owlet-nightjar (Hills, 1946) and found to be close to, if not group had not as yet begun to develop conspecific with, the Murray Cod (Macculloch- significantly.

ella macquariensis) . The leaves, placed in the Despite such differences from the Pleistocene genus Cinnamomum (David, 1950), suggest and living aegothelids, Quipollornis clearly the presence nearby of rainforest, but its extent belongs to this family, and establishes that is unknown. many of the cranial specializations of the Conveniently, the Warrumbungle volcanics family had already developed some 20 million also occur in this area and are known to over- years ago. However, as mentioned above, lie the diatomite deposits producing the fish emphasis on the hind limb at the expense of and aegothelid (Dulhunty and McDougall, the wing had not begun, suggesting a primitive 1966; David, 1950). These volcanics have condition for Quipollornis within the aego- been dated using the potassium-argon tech- thelids. Such lack of emphasis on the hind nique and range in age from 13*5 to 17 million limb further suggests that Quipollornis was an years B. P. (Dulhunty and McDougall, 1966; aerial 'insectivore' like most caprimulgiforms Wellman and McDougall, 1974). No vul- rather than primarily a terrestrial forager, as canism is known in eastern New South Wales are the Owlet-nightjars. younger than 10 million years ago, so a mini- Acknowledgements mum date of mid-Miocene can be assigned to the Chalk Mountain deposits and their included We are greatly indebted to Dr Alex Ritchie fossils. and the Australian Museum () for 252 PAT VICKERS RICH and ALLAN McEVEY graciously loaning us the specimen for study; Literature Cited Dr Ned K. Johnson (Museum of Vertebrate Brodkorb, P., 1971. Catalogue of fossil birds: Pt. 4 Zoology, University of California, Berkeley) (Columbiformes through Piciformes). Bull. for the loan of other modern caprimulgiforms Florida St. Mus. 15 (4): 163-266. David, T. W. E„ 1950. The geology of the Common- used in comparison with the fossil form. Photo- wealth of Australia. London, Edward Arnold graphs are by Frank Coffa (National Museum and Co. Dulhunty, J. A. and I. McDougall, of Victoria). The material was originally 1966. Potas- sium-argon dating of basalts in the Coona- donated by W. Burton in R 1961, and Harold barabran- District, New South Wales. Fletcher first recognized the avian affinities of Aust.Journ. Sci. 28: 393-394. Hills, E. S., 1946. Fossil Murray Cod (Macculloch- the specimen. We are most indebted to ella macquariensis) from diatomaceous earths the late Dr R. A. Stirton (formerly of the in New South Wales. Rec. Aust. Mus. 21: 380- Department of Paleontology, University of 382. Kenny, E. J., 1924. Diatomite, siliceous earth and California, Berkeley) for bringing this speci- sands. N.S.W. Mines Dept. Bull. \5\ 9-10. to men our attention. Mrs Lyn Anderson typed Rich, P. V. and R. J. Scarlett, 1977 (in press). Another the manuscript. Mary Lee Vickers typed the look at Megaegotheles, a large owlet- nightjar from Zealand. tables. New Emu. 11 (1). Scarlett, R. J., 1968. An owlet-nightjar from New Zealand. Notornis. 15 (14): 254-266. Wellman, P. and I. McDougall, 1974. Potassium- argon ages on the Cainozoic volcanic rocks of New South Wales. Jour. Geol, Soc. Aust. 21 (3) 247-272. NEW OWLET-NIGHTJAR 253

Explanation of Plate 8 coracoid; rF, right femur; rH, right humerus; rR, right radius; rS, right scapula; rTi, right tibiotarsus; Quipollornis koniberi gen. et sp. nov. Abbreviations rU, right ulna; ?Sy, possible synsacrum; Fig. A, include: A, alula; C, cranium; CV, cervical vertebrae; AM F49404, block containing skull opposite counter- part; Fig. B, lCd, left coracoid; lCp, left carpometacarpus; IF, left AM F49405, counterpart block, see scale for size. Fig. femur; 1H, left humerus; 1R, left radius; IS, left A printed at slightly smaller scale than Fig. B. scapula; ITi, left tibiotarsus; 1U, left ulna; rCd, right MEM. NAT. MUS. VICT. 38 PLATE 8

A