of Mongolia and the former USSR

EVGENII N. KUROCHKIN

Introduction Institutional abbreviations The first Mesozoic avian skeletal remains from IVPP, Institute of Vertebrate Palaeontology and were found in Mongolia by the Polish-Mongolian Palaeoanthropology, Beijing, China; IZASK, Institute Palaeontological Expedition (Elzanowski, 1974, 1976, of Zoology of the Kazakh Academy of Sciences, 1981). from the Late of Alma-Aty, Kazakhstan; JRMPE, Joint Russian- Kazakhstan were described by Bazhanov (1969) and Mongolian Palaeontological Expedition; MGI, Shilin (1977) and numerous feathers from the Geological Institute of the Mongolian Academy of Lower Cretaceous of Mongolia and East Siberia were Sciences, Ulaanbaatar; PIN, Palaeontological Institute collected by palaeoentomologists from PIN at the of the Russian Academy of Sciences, Moscow, Russia; beginning of the 1970s. From the 1970s to the 1990s PO, Collection of the Zoological Institute of the skeletal remains of various avian were recov- Russian Academy of Sciences, St. Petersburg, Russia; ered from the Cretaceous of Mongolia by the Joint TsNIGRI, F.N. Chernyshev Central Museum for Russian-Mongolian Palaeontological Expedition and Geological Exploration, St. Petersburg, Russia; VPM, in the Cretaceous of Middle Asia by the late Lev Volgograd Provincial Museum, Volgograd, Russia; Nesov as a result of his persistent exploration. ZPAL, Institute of Palaeobiology of the Polish Further remains of birds were found in recent years Academy of Sciences, Warsaw, Poland. in Central Asia by the American-Mongolian expedi- tions (see Chapter 12) and it now appears that earlier Systematic description American expeditions in the 1920s also recovered fossil birds, though they were not recognized as such Aves Linnaeus, 1758 until the 1990s (see Chiappe et al., 1996, and refs Subclass Haeckel, 1866 therein). Infraclass Walker, 1981 Mesozoic birds from Mongolia and the former Order Alexornithiformes Brodkorb, 1976 Soviet Union (FSU) are rare and usually fragmentary, Family Alexornithidae Brodkorb, 1976 but they provide some data regarding the early history Contents. Elzanowski, 1974; Alexornis of the group in this territory. In addition, many Brodkorb, 1976; Kizylkumavir Nesov, 1984; Zbyraornis Mesozoic birds have recently been found in China Nesov, 1984 (2 spp.); Nunantius Molnar, 1986 (2 spp.); (Chiappe, 1995). Together, these records provide the Sazavis Nesov and Yarkov, 1989; Chiappe basis for analyses of Mesozoic avian assemblages in and Calvo, 1994; Lenesornir Kurochkin, 1996. Asia (Kurochkin, 199Sa). This chapter presents a Diagnosis. Cranial half of the synsacrum low and summary of Mesozoic birds from the FSU and broad; synsacrum convex dorsally; ischium narrow; Mongolia, primarily discovered by Russian palaeon- acrocoracoid and coracoidal process narrow and tologists. Avian macro- follows Kurochkin tapered; shaft of the strut-like and narrow; (1995~). shaft of the coracoid with a deep and short depression on the dorsal side; scapular facet and scapular glenoid Kizylkumavis cretacea Nesov, 1984 fused; ventral epicondyle of the protrudes Holotypr. TsNIGRI 5111 1915, distal fragment of a distad to a striking degree; wing digits clawless; shaft right humerus. Dzharakhuduk locality, Navoi District, of the tihiotarsus thin; metatarsal 111 straight; metatar- Rukhara Province, Uzbekistan. Outcrop CBI-Sa, sals 11-1V short and gracile. Rissekty Svita (Coniacian). Diagno.ri.r. Same as for . Kizylkumavis Nesov, 1 984 Comments. K cretacea is a very small enantiornithine; Type .rpecie.r. Kky1kumavi.r cretacea Nesov, 1984. the maximum width of the distal end of humerus is Diagnosis. Original diagnosis of Nesov (1984): olecra- 5.1 mm. K. creracea was the first member of the non fossa is narrow and displaced in the direction of Enantiornithes to be described from the Old World the flexor process; flexor process is strongly projected (Nesov, 1984), although in the original description it distally; dorsal condyle very narrow and aligned at an was assigned to Aves inccrrae sedis. In spite of its frag- angle of 65" to the longitudinal axis of the humeral mentary condition, there are no doubts as to the enan- shaft; ventral condyle is short, aligned almost trans- tiornithine relationships of K crctacea since it has no versely to the longitudinal axis of the humeral shaft, fossa for the .&I. brachiali.~,a transverse position of the and only slightly projected in cranial aspect; intercon- dorsal condyle of the humerus, and an inclined posi- dylar furrow runs slightly on the cranial side; brachial tion of the ventral condyle. depression is not developed; a small groove is devel- oped distal to a small ventral supracondylar tubercle Z~yraor?zisNesov, 1984 on the cranial surface of the distal end, and aligned at Type specie.^. Z&yraorni.r kashkarovi Nesov, 1084. an angle of 70" to the longitudinal axis of the shaft. content^: Z. kashkarovi Nesov, 1984; Z. logu?zovi Nesov, Commenr~:As I have not seen the specimen recently, 1002. the original diagnosis, which includes the generic Diag?zosi~:Cranial portion of the synsacrum notice- characters of Kizylkumavis, as well as characters of the ably convex dorsally; cranial end of the synsacrum Enantiornithes and Alexornithiformes is presented remarkahly broad; synsacrum only slightly broadened here. The following characters form an emended diag- across both sacral vertebrae; onty one thoracic verte- nosis for this genus: distal end of the humerus is very bra precedes two sacral vertebrae; caudal half of the wide in the dorsoventral direction; the ventral portion synsacrum long and narrow; the two largest costal of the distal end of the humerus is remarkably processes are inclined caudally; no longitudinal enlarged; the dorsal condyle is broad; the intercondy- groove on the ventral side of the synsacrum. lar furrow is narrow; and the flexor process is strongly Commenr.~.There are four of from projected distally. Dzharakhuduk that are based on synsacra and The humeri of Kizylkumavis and Alexornir of Mexico assigned to the Ichthyornithiformes (Nesov, 1984, (Brodkorb, 1976) are similar in some respects. For 1986, 1990, 1992b, d; Nesov and Yarkov, 1989; example, they share: a remarkable distal projection of Nesov and Psnteleev, 1993). Comparison of these the ventral epicondyle, distal displacement of a remains with the synsacrum of ,lia?zantius valifanovi shallow olecranal fossa, and an abrupt transition from Kurochkin, 1996 showed that they should be the distal end to the shaft. However, they also exhibit assigned to the Enantiornirhes (Kurochkin, 1995c, some differences: the shape of the dorsal condyle 1996). (which is broad in Kizylkumavis and narrow and olive- shaped in Alexornis); a more spacious intercondylar Zhyraornis kashkarovi Nesov, 1984 furrow in Alexornis, and more distal projection of the Holotypc TsNIGRl42/11915, incomplete synsacrum, flexor process in Kizylkumavis. having at least seven coossified vertebrae and lacking the most caudal portion. Dzharakhuduk locality, Mesozoic birds of Mongolia and the former USSK

Navoi District, Bukhara Province, Uzbekistan. 1.ene.ronzi.r Kurochkin, 1906 Outcrop CRI-4, Rissekty Svita (Coniacian). rypespecies. Lene.t.omi.r mcz/t.lhr-~~.kyi(Nesov, 1986). Diagnosis. First on synsacrum expands grad- Diagnn.ri.~: The cranial portion of the synsacrum is ually in the cranial direction; transverse processes on only slightly con\,es dorsally, the cranial articular the second sacral vertebra are slightly marked on the surfice of the synsjcrurn is transversely elongated; the dorsal surface; two pairs of the largest costal processes third and fourth vertehrae of the synsacrurn possess are slender and distinctly inclined length\vise; the the largest costal processes; the costal processes are at synsacrum is generally extended and narrow. a right angle to the sagittal plane; and the ventral comment.^. A thoracic vertebra (TsNIGRI 43/1191.i) groove is uide and shallow. from the Khodzhakulsai locality (lihodzhakul settle- Comments. This skmsacrum was described as ment, Karakalpakia, Uzbekistan; lihodzhakul Svita, Ichtbyov~~ismaltsh~z~skyi I)!. Nesov ( 1 98 6) in the lirnily Cenomanian) (Nesov, 1984, 1992b; Nesov and Borkin, lchthyornithidae (see also Nesov, 1992b, d). However, 1983), regarded as an indeterminate hlesozoic bird by the similarity ol' this hssil to .'n?za?ztius z~al~/2rnnvi Nesov (I 992d), a left ('TsNIGRI 4411 1915) Kurochkin, 1006, and the at~undanceof postcranial and the shaft of a left humerus (TsNIGRI 45/11915) remains of Enantiornithes at the Dzharakhuduk from Dzharakhuduk (Nesor, 1984), were also assigned locality enabled it to be reidentihed as an enantiorni- to Z. kashkarovi. The vertebra has a wide neural canal, thine (Kurochkin, 1996). Because of a noticeable deep lateral excavations on the centrum, and nearly diff'erence from Zbyvanvflis, it was assigned to a separ- flat cranial and ca11da1 articular surfaces. It is compar- ate gent~s. able in size to the vertehrae of the synsacrum of Z. kashkaroz~j.The structure of the glenoid facet on the I,ene.rovui.r nzalt.sbev.rkyi (Nesov, 1 980) scapula and its acrornion show a certain similarity to Holo~ypc. PO 3434, cranial half of the s\ nsacrum. that of the enantiornithines, but this is not sufficient Dzharakhuduk locality, Navoi District. Bukhara evidence for its assignment to Z. kashkavovi 'She Province, Uzbekistan. Outcrop (:U1-14, Uissel\tr S\ it;] humerus shaft cert~linly is not enantiornithine, (Coniacian). because its nutrient foramen is located in the typical Uiagao.ris Same as for genus. position tbr neornithine taxa, whereas in the Enantiornithes the nutrient foramen occurs on tbe Sazavis Nesov, 1080 opposite side of the shaft. 'Iype species. Sazavi.rpvisca Nesov, 108'). Diagno~.is. Small birds; distal end of the tibiotarsus is Z,hyvaov~i.rlogurroz~i Nesov, 1092 wide and its large medial condyle has a rounded dorsal Holo[ype. PO 4600, incomplete synsacrum, having at margin in cranial aspect; intercondylar furrow is dis- least five coossified vertebrae and lacking the most placed !aterally, therefore the lateral condylc is caudal portion. Dzharakhuduk locality, Navoi narrow. l'he diameter of the bone is strongly reduced District, Rukhara Province, Uzbekistan. Outcrop CBI- dorsal to the distal end of the tihiotarsus; the liga~nen- ja, upper member of the Rissekty Svita (Coniacian). tal tubercle on the cranial surface of the tibiotarsus is Diagaosi~: The first vertebra of the synsacrum expands weak and located relati\,el!. distal. abr~lptlyin the cranial direction; the transverse pro- Comme~zt.~:As I ha\,e not seen the specimen recentl!, cesses of the second sacral vertebra are prominently the original di~lgnosis,ivhich includes the generic marked on the dorsal surface; two pairs of the largest characters of Srr-rrz1i.c; as we11 as characters of the costal processes are thick; the costal processes of the Enantiornithes and .-\lesornithiformcs, is presented second sacral vertebra are perpendicular to the sagit- here. It should also be noted that in the original diag- tal plane; and the synsacrum is generally expanded nosis lateral and medial aspects were confused, though and broadened. these are corrected in the diagnosis given above. 'l'he follo\ving characters form an emended diagnosis for a hroad dorsal arch and a flat caudal lamina that is this genus: the transition from the shaft to the distal strongly projected caudally, a low dorsal process, and a end of the tibiotarsus is abrupt; the medial condyle is shallow lateral excavation of the body. l'hese charac- nearly circular in cranial view; and the intercondylar ters allow assignment of PO 3473 to the fossa is somewhat medially displaced. '4lexornithidae (Kurochkin, 1996). 'I'his specimen S~i;nz,i.r was assigned, with some doubt, to the differs from the axis of 3'. valzfitzoziin heing larger: the Alexornithidae (Nesoi. and I'arkov, 1989) or to the distance between tbe cranial and caudal articular sur- Enanriornithes (Nesov, 1992b, d). Assign~nentof S. faces is some 133 Inn] (measured from the figure in prisca to enantiornithine birds is supported hy the Nesov, 1988) but only 6.2 mrn in .N. ual~fiirozli. presence of a bulbous and enlarged medial condyle of Two enantiornithine were described by the til)iotarsus, a small and trans\-crsely compressed Nesov and Panteleev (1993). Specimen PO 4819 is lateral condyle, and a smooth tubercle in rhe centre of represented by the dorsal half of a coracoid with a the ascending process. narrow shaft, slight projection of the lateral margin, and shallow depression on the dorsal shaft, features Sazavispri.sca Nesov, 1989 characteristic for the :llexornithidae. Specimen PO Holo[ype, PO 3472, distal fragment of the right tibio- 4818 is represented by the fragment of a shaft that tarsus. Dzharakhuduk locality, Navoi District, shous a Ixoadened sternal portion and a deep dorsal Rukhara Pro\-ince, Uzbekistan. Outcrop CBI-14, depression (Kurochkin, 1996). Because of its very Rissekty Svita (Coniacian). fragmentary condition the family relationships of this Diap~osis.Same as for genus. hone is uncertain. Commcnt.c S. pi-i.sr~z is another very small enantiorni- thine from the Dzharakhuduk locality. The width of the distal end of the tihiotarsus is 4.5 mm which is Enantiornithidae gen. indet. comparable in size with Kizylkumavis cn7t~zrr~zand a 'The shaft of a right coracoid, TsNlGRl 56/11915, congeneric or conspecific relationship might be sup- some 15 mm long (measured hy the published figure in posed. However, this cannot l)e certainly dernon- Nesov and Rorkin, 1983), was provisionally assigned strated because the remains are non-comparat)le. to birds (Nesov and Rorkin, 1983) and later to the enantiornithines (Nesov and Panteleev, 1903). This Other a1exon~ith~irm.sjpom the Kizylkum Deserl specimen clearly belongs to the Enantiornithidae In various papers Nesov described other fragrnentarv because it shows a distincti~edeep dorsal depression, avian fossils from the Dzl,arakh,lduk localitv in running far dorsallv, and a conlex lateral margin of Bakhara Province, Uzbekisran. Some of these ;,ere the (KurochLinl assigned to the Enantiornithes indet. and some only to Aves indet. 'The remains of alexornithiforms are listed Alexornithifor~nesfamily indet. belo&,. 'IchtLyortzis' ~II~~ZUSCU~USwas based on a single thoracic vertebra, PO 3941 (Nesov, 1990). This specimen Alexornithidae gen. indet shares, with the synsacrum of L. malt.rhmskyi, an ellip- .4 cc)~npleteaxis, PO 3473, was figured as avian I)!- tical protile of the cranial articular surfice and wide Nesob- (1088, fig. 1: 3, 1992h, fig. 2: 0-P) and then cited vertebral foramen. However, because of its fragmen- as similar to Gaviidae (Nesov, 1992d). This specimen tary nature, '1'. mirzuscu1u.r can only be assigned to the shares, with the axis of .\I. valifanovi, a general cranio- Alexornithiformes fam. indet. (Kurochkin, 1996). caudal elongation, poorly developed cranial articular The proximal fragment of a left tarsometatarsus, facets, lateral extensions of the cranial articular facets, PO 3394, was assigned to the Enantiornithidae hy Mesozoic birds of Mongolia and the former USSR

Nesov and Borkin (1983) or to the Enantiornithes 1976, 1077, 1995) might be features of the (Nesov, 1984, 399211). The specimen shows the proxi- Enantiornithes, though at present these features are mal fusion of the metatarsals and the enlargement of not known in described remains of this group the medial cotyla. 'l'he depth of this proximal end is which include the occipital cranium region of about 4 rnm (measured from the figure in Nesov, Neuque~ior~iisvolans from the -Coniacian of 1992b). The specimen was correctly identified as an Argentina, rostral portions of the and maxil- enantiornithine, and should be assigned to the lary and palatal apparatus of K. valifa~iovifrom the Alexornithiformes on the basis of its very strongly Late of Mongolia, and rostral portions of reduced metatarsal IV (Kurochkin, 1996). the mandible and maxillary and dorsal cranium of embryonic enantiornithines also from the Late Discussion. In total, the Bissekty Svita of the Campanian of Mongolia. Dzharakhuduk locality has produced 13 enantiorni- Thus, there is only circumstantia1 evidence that thines all based on fragmentary remains. Seven of Gobzjtt,ryx belongs to the Enantiornithes, since it does them were described under species status here, but it is not have advanced characters in common with known impossible to compare the other six fragments. enantiornithine birds. In addition Gobzjteryxis charac- Nevertheless, at least three sinall and two middle- terized by some primitive characters of the quadrate, sized enantiornithines might have existed concur- pterygoid and palate which are also unknown in other rentlv in the Coniacian of Uzbekistan. Enantiornithes.

Family lllexornithidae Brodkorb, 1976 Cobipteryx ~ninutaElzanowski, 1974 Cobipteryx Elzanowski, 1974 Holotypc: ZPL4LMgR-1/12, rostral portion of the skull. type specie^: Gobzjteyyx nzinuta Elzanowski, 1074. Khulsan locality, Nemegt Valley, South Gobi Desert, A~ne~ideddiag~rosis. Culmen straight and very thin Mongolia. Baruungoyot Formation (Late Campanian), above the nasal openings; rostral ends of the premax- . illa and mandible flattened with rounded tips; mandi- Rq'22rred material. Rostra1 portion of a skull, ZPAL bula with a low and short symphysis and thin rami; MgR-1/32, from the same locality. contact surfaces on the ventral margin of the premax- Diagnosi.r. Same as for genus. illa and and dorsal margin of the mandible Comment.r. C. ~ninutashows a gracile construction of flat; no distinct grooves on the lateral surface of the the upper and mandible and large rostral choanal rostral portions of the premaxilla and mandible; fenestra bordered caudally by the palatines dorsal mandibular margin distinctly elevated above (Elzanowski, 1995). Following additional preparation the level of the lateral mandibular process; and large of the holotype, Elzanowski (1995) discovered a small choanal fenestra located in the rostral area of the hooked ectopterygoid that has also heen identified in palatal shelf. both the Eichstitt and seventh specimen of Com~nnits.Originally, Gobipteryx was assigned to the (Wellnhofer, 1974; Elzanowski and (Elzanowski, 1976, 1977), but later Wellnhofer, 1996). The ectopterygoid is present in Martin (1083) attributed it to the Enantiornithes. many theropods, and I evaluate this fact as a further There are no reliable arguments for assigning the skull confirmation of the relationship hetween the portions of Gobzjtecyx to the Enantiornithes, since the Enantiornithes and and their assign- skull remains of Gobzjteyyx are not known among ment to a phvlogenetic lineage separate from ornithu- undoubted enantiornithines. However, the bipartite rine birds. 'This and some other cranial mandihular articulation of the quadrate, anterior synapomorphies support a theropodan origin for hihrcation of the pterygoid, the subsidiary palatal Sauriurae, hut not all birds. fenestra, and a hooked ectopterygoid (Elzanowski, E.S. kUKOCIIKIN

.\inttn~rtiu.r Molnar, 1086 side of the mandible; a shallow, broad axial groove on T~prspecies. .Vaimlrti~i.cvn1if;Enozli Kurocll k i n, 1 406. the ventral side of the synsacrurn; a short tibular crest COIJ~CI~~J:.\n?/nutr~is e0.c Molnar, 1980 (ciustral ia, on the tibiotarsus that is approximately five times the ); .I-.i'~/j/~?ll(lvi Kurochkin, 1997 (hlongolia, 1,ate transverse width of the proximal articular surface; Ca~npanian);,YUIIN~~~~I~SS~. (Australia, Alhian). position of the nutrient foramen on the crarrial side of Ding110.ri.1: Maxillary and mandible short, high and the tibiotarsus close to the distal extremity of the stout; culmen very straight and thick above the nasal tibular cresr; subtriangrlilar cross-section and sharp- opening; the humerus has a cur\,ed shaft with a thin ened caudal edge of the proximal shaft of the tibiotar- uiid-portion; the shaft of the radius is bowed; the sus; transversely compressed lateral condyle of the proxi~nalphalanx of the ~najordigit has a rectangular tibiotarsus which proiects markedly craniad in distal top; the tihiotarsus has a long and remarkably thin view; and metatarsal I\- is shorter than n~etatarsalIT. shaft that is bowed laterally; conspicuous intercotylar Commelzts. The Hermiin Tsav locality, situated on the pro~ninenceon the caudal area of the proximal arricu- \vestern border of the 'I'rans-Altai <;obi Desert in lar surfiacc of the rihiorarsus; lateral area of the proxi- Southern Mongolia, is famous for yielding numerous mal articular surfiice of the tibiotarsus slopes distad; fossil of birds, some of which contain e~nbrvos well-developed tibular crcst reaches to the margin of (Elzanowsl:i, 1081; Chatterjee and Kurochkin, 1991; the proximal articular surhcc; tit)ular crest located see helow). 'The nearly complete holorype skeleton of along the craniolateral edge of the shaft of the tibio- N. valfanovi was found on the northern slope of tarsus; proximal origin of the fibular crest and top of Hermiin Tsav, about 1kn1 east of 'Bird's IIill', a local- the cranial cnernial crest united together; elongate ity for fossil avian eggs at the mouth of Hermiin Tsav. depressions run along the cranial and caudal base of Sorne bones were collected in association including: the fibular crcst; medial condyle of the tibiotarsus rostrsl portions of the mandible and premaxillary; a transversel\l elliptical (nor circular) in cranial view; portion of the palatal apparatus; some cervical crte- fihula ver\. short, flat and thin; metatarsals 11-1V of a brae; part of the pelvis and synsacrum; the proximal similar thickness. end of the first phalanx of the major digit with the car- Commej/t.l: In spite of the great geographical and - pornetacarpus; the distal end of the left femur and poral gaps het~veen val~/atzovifrom the ;\lbisn of pelvis; the tarsometatarsus with some pedal pha- htongolia and .\: eos from the Campanian of Australia langes; and other pedal phalanges. Many pieces of (hlolnar, 1086) it was decided to assign the Mongolian eggshell were also found in association with the bone taxon to the salne genus because of the great similarity remains of 1V. valifanovi. The eggshell has a laevisoo- in the derived characters of the tibiotarsus of the two lithid n~icrostructurethus linking this type of eggshell forms. to the Enanriornithes (Kurochkin, 1996; see Chapter 2 8). .la~ralrrtiusz-alifkr~ovi Kurochkin, 1906 N. vnlqanoviis disti~iguishedfrom A:. coo by the rela- Holo

South Gobi .iimag, Mongolia. Middle layers of the shaft, which is rounded in ,V. PUS, a small tubercle at rhe outcrop on the northern slope of tiermiin Tsav, centre of the ascending process which is located

Raruungoyot Formation (1,ate Ca~npanian), Late nearer to the top of this process in h'. EOS, greater Cretaceous. cranial protrudence of the lateral condyle, a slightly U~ng~rosi.~:Shallow longitudinal groove on the ventral compressed proximodistally medial condyle, which is Mesozoic birds of Mongolia and the former USSR *.-

Figure 27.1. Holotype skeleton of Nanantiu~valifanoviKurochkin, 1996 (PIN 4492-1). Scale bar= 20 mrn.

more circular in N. eos, and a deeper and wider cranial fragments of two from the Khulsan locality intercondylar fossa. In additon, the tibiotarsus in N. located some 150 km east of Hermiin Tsav. This taxon valifanovi is about 30% longer than in N. eos. shows some features in common with N. valifanovi in The relationships of N. valifanovi to Gobipteryx the configuration of the maxillary segment and in the minuta Elzanowski, 1974, need to be discussed here, presence of conspicuous rows of nutrient foramina in because they were both found in the Baruungoyot the maxillary and mandible. However, N. ~al~novi Formation of the South Gobi. G. minuta is based on differs from G. minuta in having more robust rostra1 E.N. KUROCHKIX

ing the ability for powerful tlesion ofthe digits. 'l'his is in contrast to the relatively small size of the trochlea of the left metatarsal I, and the phalanges of digit 1. The powerful anterior second and third toes with curved claws and weak hallux also suggest a climbing adapta- tion for the foot. The palatal elements and the rostral portions of the mandible and upper jaw have a very powerful and robust construction. This is, apparently, an adaptation for feeding on tough objects, perhaps fruits or seeds.

Family Brett-Surman and Paul, 1985 Contcy1t.r. Eeay1tiorui.r Walker, 1981; Avi.mtrru.r Brett- Surman and Paul, 1085; Soroaz1i.rnuru.r Chiappe, 1093. Diagy1o.ri.r. Deep fossa on the cranial surface of the scapula; scapular facet and scapular glenoid separated; ischium wide; tibiotarsus with a straight and robust shaft; metatarsal 111 with a strongly convex trans- versely dorsal surface; medial ridge of the trochlea on the metatarsal I11 projects markedly on the plantar side.

Euay1tiorni.r Wal ker, 198 1 Type .rpecic:l: Ein~ztttiory1i.rleali Walker, 108 1 Corctr. E Itdi Walker, 1981 (.krgentina, h,Iaastrichtian); E. wnlkeri Nesov and Panteleev, 1003 Figure 2 7.2. Ilcstoration of the skeleton of h'anaiitiu~ (Uzbekistan, Coniacian); 6 rnartitzi Nesov and iwlrf;li~u.jiKurochkin, 1'496. Scale bar= 10 mm. Panteleev, 1993 (,Uzbekistan,Coniacian). Diagy1o.ri.r. Shoulder end of the coracoid robust and portions of the upper jaw and mandible, sharper short; shaft of the coracoid wide; acrocoracoid and rostral ends of the upper jaw and mandible, thin and coracoidal process stout; scapular acromion broad acute contact surfaces of the n~asillaryand mandible, with an obtuse top; fossa or foramen on the cranial deep grooves possessing nutrient foramina both in the surfice of the shoulder end of the scapula; scapular upper jaw and mandible, nutrient foramina only in the facet and glenoid facet of the scapula are separated; anterior half of the maxillary rostrum, nutrient f'ora- dorsal portion of the hulneral head longer than the mina which become larger caudally, and in the pres- ventral portion; distinct cranial fossa in the cranial ence of an axial groove on the ventral side of' the surface of the proximal end of the humerus; deltopec- mandible. toral crest with a thin proximal base; distinct depres- The short tarsometatarsus and relatively short and sion in the caudal surface of the proximal end of the powerful second and third pedal digits with strong and humerus. slightly curved clarvs of N. valifanovi reflect arboreal Co>rrrnctrt.c In the present state of knowledge of the adaptations. The ungual phalanges are sturdy, similar Enantiornithes it is a ditTicult task to develop a generic in size and slightly curi.ed, with symmetrical articular diagnosis for Etratztioriti.r, since almost all authors ana- cotyles and well-developed flexor tubercles, signify- lyzed either the characters of the Enantiornithes or .Lq ut:s[nqy 1e purl(!! alax 'a.81cl pur! (leurs 's%a I!SSSOJ s!q~jo sluaur!pas aqj2 .dc[u!y !clc)f) LII~OS341 LI! 'u!e~ UF!.~E a1s8uola jo sad?(] MI 'la1c7 ,saq]!u1o!]ueug -unoLu 111n uc11y jo q110u 01 ury j 1 J~LIOS'uo!ssa~dap aq] 01 LLI~~Ipaud~sse put: miltr!in .3 se solilqura ,iesL12u!!#!!~ aq] u! pa~cn~~sst qstqfi ',i~qesol aesL

I!SSSOJ aql 11Cpa~!LI8~3al(<~h[)!ySAfiOURZ[g 1JaICl LI!!~!.III~>aql 11: thb 1 U! e!loduoly L~IIIOSLI! gd~qd( put: '(12~661 'g~hl)LI!ILCN .t""w"[qy LLIO.I~ paq!.~s~p aq] ,icl pa~sallosa1a.n 'zno~..r,)ftnzt~~tj!.ri~? '~!~I!IIJ~!ILII:U~ ".a L[s!q.\i 'tih 1 '!~L"uuc"~~ "12U,2211 X,(>~/~ZYO:) a.8~~1e jo sauocl alp~!8Japlnoqs puc r3u!.\\ leJa!\aS 01 soli~cluraasaql pou81sst: aq ,i~!l!cl~:qo.~d11~1 's8.8a FOO(, 1 'UI~qso~11y !non;?zr 1t!zriljl.rn3 al!posc).~s10 a11.1111aq 01 1q8noq1 uanq .ilsno!,ia~dpeq "60 1 'u!y qso1n;y 1!t~llj,z~n3 qs!q.u .\es-~u!!urlaH ruoq s##a aleiiuola llct~rsu1 suo~ at:p!q]!rrJo!]ut:u:{ -J[2JS ~1~.~~!L~l~1~\11~~aC1!12Sap((1861) !yS..MOUt!Z[T c!lo.Zuoly molj solilqwn su!q~!u.~o!~uctrg .LIJILI~:.I~~salau lep!oselos pa~s#uolaa1oLu puc la~\\out!u3q1 liq puL?"yqs ~car!xoldaq] jo du!uapeo~q .CJ!~~LLI~~ILIOS PU~ q1.10~ sr: [lam st: 'snoasoal3 aq~ [CISI~'p!O~t?.~~~Ol~~!.~~llOq~ 'ssaso~dIFp!OSFl03 13.+\01 jo pu.3 J~Ilc t:!~o#uoj,y pa]!qequ! sp!q]!ulo!]ueuJ sS~el -1cu 'az!s lallerus qsnw l!aq] .icl !/WJI g moy paqs!l~;i 1eq1 s.z\oqs put: I?!s\: Iulua:) jo snoaselalg a11!-1 aq] -u!~s!p alc JILZJLIIN~.y p~eZW~IIIRZ .,T .sasoude!p alp LI! AI!SJ~.\!IJ u,nouy l!aq] sasl?alsu! saq]!ulo!lueuy aq] u! pauo!]uarrr salnlsnlls aql u! la,tl!p ,iaq~'la,\afio~{ jo a;l!~e~uasa~dal#rl!,i~ a8~el S!L~I jo ,i~a.ioss!parlLI> ,(sa~n.8ypaqs!lcjnd ulo~jpalnscaru :ssnsold lep!os '21u.11 'y uerl] -ems aq] jo do] aql puc p!tr~elo;rolseail] jo do] nq] 1ari;~e.l.illsu!~yp su~ppu~ ruuu l.r)z d~r!aclsnlamnq aql usa.n]aq ur rLr () 1 .ila~l:[u!xolcidc)az!s .IC[!ILI!S lila,\ I: jo j0 pLI3 IWII!XO.I~ aLllji? LIlp!.M aL[l '[lllq 2dl~1I? S! 11 'LUlOj ale 'paseq ale a,iocl1: pais!l sapads o.\11 aq] rls!rlA\t uodn ,nau s1q1 OI S~rolaqosl~? s11dlc3c1arrrodles pue stupel 'p!oswos aql jo suo!ilod laplnoqs 0.111 aqL :r~usiri1iro3 'e~11n(IF /i[qt:qo~cipue sulaLunrl e jo spus lels!p aq,j, .~no~ss! yerls aqi jo uo!~~od~t:ru!so.~d sq~ pue ,ill" (c(,(,~I'u!yqso~ny) ap!s lelluaa nrp -!pau~-o~alt:~no.11su s! ssaso~dlep!osclo:) :nsouYyg jo aliiue le!uclsoliua,\ aq] uo uo!ssnldrrr! le~uaurt:#!l .(ue!st:!uo3) [?I!.~sAIYSSS!~ 3111 ~u.I;)~III;)I~Ialpp!ru Iwuan -1t:lns~!s daap e sassassod put: a;i.rcl s! snlaulnq '~1-1g:l dols~no'uels!ysqzn 'asu!noq elcrllnH sr11 jo p~aI~:ILI!XC)J~ alp putz'als~acln~ leuua.\ uaureloj '~s!l~s!(~!O.\EN ';i~!lt:su~ ynpnqyeleqza .p!~st:los ~t?!xca~[i ,iq ~)s~t:lojladIOU S! IE~IaSEC] ap!.n '~uoisha.\ 1q3p aql jo 1uaru.8u.q laplnoqs '60r)t 0~1..'d't;o/o~ t: 'pcaq lclns!ut: aql jo suo!~.~od1t:s~op PUB 1~11ua.i COO I '\aala~rr~:dpue hosaN !/~,z~rz7rrrs!/iro,z/u~?uy aql uaa.waq alSue uado pue ap!x nrp '1s9.1s 1c1 -o~sado~lapaql jo asecl lem!xold aql leau uo!ssaldi~r! ~uua~~rc;i!l~es.~op aq~ jo adt:qs ~e~oaql Bu!pnlsu! 's1a1 -seJeqs .y!s~dslsr[lo smoqs osle snlarrrnq aqL .(g'i~ a1n8!,4) 1jt:qs ar11 put: pua laplnoqs aql rraa.niacl qsau ~~~~~~03U!L~.i[qey~t?r~a~ I: pue 'snap!oselos snssas -old aql jo uo!leu!wal lelluaa pslu!od aql 'snlawnq alp jo pua ~m!xo.~dJ~I jo asej~nsIepncs aq~u! rro!s -sa~dap[epnes fiolleqs .i~a~e 'lsals ~elojsado~lapaql jo aseq ys!q~arp 'snlamnq aqi jo pua II:LLI!YO.I~ aql jo ,alp1!.8 laplnoqs aql asejlns IR!UFJS aql jo call: ICLLIIX~J~aq] jo alpp!ru aql jo sauoq pue snlawnq sq] .iq pa~uasa~da.~OSII: sau!ql rr! ~ssojle!uels ,+iolleqs.i~a.~ c 'puoq Jclns!llc lelatunq -!ulo!lueua larpo ruo~jI! qs!n#u!is!p qs!q.\\ J~!~I.IO!/~IIUL~ aql jo suo!llod 1t:slop pue ~UIU~Asq] jo q12ual jo slaneleL[s pasuehpt: aqi isalas UI pa~d~ua~~e1 a iocle ~mba1so1nle aql /iq paz!~a~st:~eqss! !,z~.~.I:)zL 1!~r,(j!;tn3 ua.4 s!sou8e!p aql u1 ,ssu!ql!uJo!]u1:Lra all] uo s~adcd ,uo!lewloLl1.8arua~ (ut:!~qs!l~seel,y ~il~e~-ue!ueclu~eg Iuasal laqlo duew ri;u!pn[su! pue (1861) .~aq[e,&\ a~cljhncraselal:) ole7 oq~OI #1101acl ii~![eso[ ~11l.n du!uu!Saq 'ssclscy~r! s!q~jo sapads ale.redas

NSSn .I~UI.IO~aql put: c!lo;iucr~q+tr spl!cl -,!ozossw E.N. KUROCHKIN

Figure 27.3. Gun'Ipia nezsovi Kurochkin, 1999a, (Late Cretaceous) of Guriliin Tsav, Omnogov' Aimag, Mongolia. Holotype PIN 4499-14, proximal end of the right humerus in (A) cranial and (B) caudal view. Paratype PIN 4499-13, shoulder end of the left coracoid in (C) dorsal and (D) lateral view. Scale bar = 10 mm. the JRMPE. The small type is the same as that embryos (Kurochkin, 1995a, c, 1996). However, after found at Hermiin Tsav (Mikhailov, 1995, 1996), but study of fossil material in the ZPAL collections I now fossil embryos have not yet discovered in eggs from agree that only one taxon is present and that it belongs Khulsan. to the enantionithines as originally proposed by The avian embryos from Hermiin Tsav, now in the Martin (1995a). Differences among the embryos collections of ZPAL and PIN (Figure 27.4), are beauti- (Kurochkin, 1996) represent different age stages, as fully preserved and show even the smallest ossified ele- was first noted by Elzanowski (1981). The structure of ments including, for example, the most proximal wing the eggshell is an important piece of evidence sup- phalanges. This characteristic, as well as the complete porting the identification of the embryos as enantior- ossification of the bones, from the ends across the shafts, nithine. The ratio between the spongy and differs fundamentally from the embryos of recent birds. mammillary layers, the ultrastructure, absence of In the latter case the process of ossification begins from asymmetry, and the greater thickness of the eggshell centres in the ends of bones and in the shaft, so that car- are all characters of enantiornithine eggs (Mikhailov, tilaginous insertions between these ossifications exist 1991, 1996). By contrast, the possible absence of the some weeks after hatching. external layer of vertical crystals distinguishes the In earlier papers I concluded that two groups of Hermiin Tsav eggshell from that of the Ratitae, birds were represented among the Hermiin Tsav , and . amLn suawlsads ohqu~aaql pus xiC~m(tq03 jc) AJFI u1 .sau!ql![rJo!]ueua jo slalseleqs ~e!uels~sodpaz! -~!xtma~daql leql panz~e(1861) !~sMM"uI?~~.soli~qwa -1slsads Jaqlo pus 'slndess pus p!oss~osaql uaamlaq pus xl~nc$qo3~oj alnleaj uowwos d[uo aql s! ssaso~d uo!ls[ns!lJs lagsos pus ssoq aql s~!q!qxa I! asu!s 'auy JB~~~!I.I~U.II~JZUOIF '~a~a~fio~ .saql!uJo!luauz aql ~oj -!uJo!lueua lil~salss! (186 1 '!~SMOU~Z[~)tt/~-$~ili~ s!ls!JalssJeqs aq 01 s~saddspus usslnqx woy nznuzui IL~~J~Zuaw!sads 'usslnqx woy nznuzul xiC~azdtqo3 '3 U! puno] oqt: y qs!qM 'a[q!pusw aql u! ssaso~d 01 pau81ssa aq uss soli~qwa ~ss~~,u!!wlaH aql JRlns!lleoJlaJ sue[ e seq uaru!sads s!ql 'uo!l!ppe laqlaqm o] se sureuraJ [[!IS uo!~sanb aql 'JaAaMoH

XSS~J~WJO~ aql pus c!qoYr~olqjo sp~!qs!ozosaly similar and thar the latter probably belonged to G. Kholboot is located just 10 kni \vest of the Khurilt nlil~ut~~.Certainl!!, the embryonic specinler. Zl':lL Ulaan Bulag locality where ,~ln~biorr~r.s\\.as found and MgK-1 188 is so~newhatsimilar in its general outlines the sediments at these localities, \vhicli arc currenty to the premasillar)~of G. mi~riit~~,but differs in the assigned to the Biiiintsagaan Corizont (possibly larger and more cranially concave nasal aperture ant1 Neocornian - scc Chapter 14) are of sirnilar age, in the sharper angle between the lateral surfaces of the though Kholhoot may be slightly J8ounger (Sinitsa, prcrnaxillary. The principal tliference between tlieln 1993). is the structure of the rostra1 end of the heak. It is thin The Kholboot specimen consists of portions of the and sharp in Cobipt~~yxfrom Kliulsan, but flat and skull and some shoulder, wing and hind limb bones. rounded in the embryos from Hcrrniin 'Tsav. Initially, it was erroneousl\. identified as 3 pterostlur Moreover, the esterior contact areas het\veen the pre- because of tceth on the ja\vs (Kurochkin, I')Vl), hut nlaxilla and mandible in Gobiptc~ryxare narrow \vhile \vt~s later recognized as avian (Unwin, 1993; they are wide and Hat in cmhryos. 'l'he ernhryos also Kurochkin, 1993; Bakliurina and Unwin, 1905)) show a double-headed quadrate with orbital process; a though suggestions of a relationship with .-1n~biorttr.r ~entralAange on the rostral edge of the mesethmoid, a (Unwin, 1993) arc incorrect. The Lholboot specimen lateral groove on the mandible, a very elongated scap- has clear enautiornithine characters including a Y- ular acroniion with a niedioventral projection, no shaped with a long hypocleideum and Ineta- scapular labruni, a short and wide acrocorncoid, incor- tarsals that are only fused prosinially, and is simil:~r, poration of the sctlpular glenoid and coracoid glenoid for example, in respect of the toothed , to Lower for the glenoid f;icct of tlie humerus, a merged scapu- Cretaceous Chinese enantiornithines, hut distin- locoracoid construction, a proxinlal origin for the dcl- guished from then1 hy the very long tarso~nctatarsus. topectoral crest, a major ~netacarpalthtlt is longer than the minor nietacarpal, a manual phalangeal formula of Sohclass 1-1-0, and a notarium composed of two thoracic Infraclass Odontornithes I:orl)es, 1884 vertebrae. They also exhibit distinctive features such Order Hesperornithiforrnes Fiirbringer, 1888 as a small nasal aperture, a broad and dorsorcntrally L)iaguo.ri.s. See Martin (1084, 11. 147). compressed rostral portion of the premaxiilary, a long C'o~~telzt.~Hesperomithidae hlarsh, 1872; Baptorni- phalanx of the first digit, a very short cranial part thidae .i~ncricanOrnithologists' Union, 1010. of the , a stout fibula which is of siniilar length to Corrrrncwt.r. I lesperornithidae and the genus He.rpt,rnr.lii.r the til~ia,and the absence of ti~sionof the proximal have never really hecn diagnosed. Martin (1084) tarsalia (asrragalo-calcaneus complex) to the tihia. ntte~nptcdto di;~g~,oseHespcrornit11ifi)rmes and to Thus, I consider, in contrast to blartin (19953) and this list one important character can be added: loca- lilzanowski (I995), thar assignment of the emhr\.onic tion of teeth in grooves. specirriens Z1jL4L MgR-1/34, htgK-1/33, and IClgli- 1/88 to Gobipteyy.~or even to G. nziuutn cannot be Family Hesperornirhidae hlarsh, IS72 iustified and they together with the specimens in the Conteuts. H(~.s,f)n.or.~/i.rMarsh, 1872; Parab~:spt~ron~i.r PIN collections (Figure 27.7) must he assigned to a Martin, 1984; .,l.rinbt~sp(.ronzi.rNesov and Prizcmli~i, new taxon (C:harrerjee and Kurochkin, 1994). 1991.

Order Euornirhiformes Kurochkin, I996 He.~croniirMars 11, 1 8 72 I'arnilv no\. Typi~.rpc,cies. I1esperon~i.rri~g~~lis Mars ti, 1 8 72. ,?I he partial ~keletonof a small vertebrate was col- CO?L~(JFL~S.HH~CYOYFL~.~ nznlis hlarsh, 1 8 72; I3t:s/)cr.on~i.r lected by theJRMPE at Kholboot in rhe Eastern urea cm.rsipe.r Marsh, 1 876; He.rpc~ror~~ispcilir Marsh, 1 976; of the Mongolian .ilrai in tlie Bayankhongor Airnag. H~~spwomi.rro.r.ric~zc.s Nesov and Yarkov, 1993. Mesozoic birds of hlongolia and the former USSR

Hespevovr~isrossicus Nesov and Yarkoy 1 993 Holo[ype. VPM 26306/2, proximal portion of the right tarsometatarsus. Right shore of the 'I'zimlyanskoe Reservoir, Ilon river, between Rychkovo and the 278 km Station, Surovikinskii District, in the south-west of \ulgograd Province, Central South Russia. Marine beds, Br~l~~~r~tellocamuxmamillut~rs zone, upper zone of the Early Campanian. Rtft;vvr.d matevial. 'The shaft of a tarsometatarsus, an intermediate phalanx of the fourth pedal digit, a frag- ment of a thoracic vertebra, and a fragment of a cervi- cal vertebra were all recovered from the same 1ocali:y (Nesov and Yarkol: 1003). A fragment of the proximal end of a right tarsometatarsus from the Earlv Campanian of Ivii-Klack, Scone, Southern Sweden has also been assigned to this species (Nesov and Yarkov, 1993). Figure 27.5. Right distal ribiotarsus of a small hesperornithiform from the Nemegt Formation (1,ate Diagr~oszi-.The proximal articular surface of the tarso- Cretaceous) of 'Tsagaan Khushuu, Omniigovl imag, metatarsus has a very large transverse width and small Mongolia. Lateral (A), cranial (B), medial (C), and distal (D) dorsoplantar depth, the diagonal slant is strongly views. .ibbreriations: eg, extensor groove; mc, medial condyle. expressed, the lateral edge of the lateral cotyla Scale bar = 10 mm. exceeds the intercotylar prominence in proximal direction and the rnedial cotyla is located more distal in respect to the lateral cotyla. type of H. rossicus was assigned to sp. Commr.~rts.Ht:rpr~vorni,r is of the masculine gender, thus (Nesov and Yarkov, 1993). This specimen differs from according to the Itztemational Code of Zoological H. vossicusin the more medial location of the intercot- Rrome?zclatuve(1985, article 32d, 33 (II), 31b) the origi- ylar prominence and a larger ridge in the dorsal base nal epithet vossicu (Nesov and Yarkov, 1993) must be of the intercotylar prominence (Nesov and Yarkol, changed to vos.ricus. 1993). H. vossictrs is clearly hesperornithiform, but differs from Hespevovnis vrplis Marsh, 1880, from North Hesperornithidae gen. .qnierica in the larger depth of the proximal articular The distal portion of a tibiotarsus from the Nemegt surface, more proximal projection of the lateral cotyla beds of the 'Tsagaan Khushuu locality in the South and approximately 20% larger size of the tarsometa- Gobi Desert was announced as Buptovnis sp. tarsus. (Kurochkin, 1988). After comparison with all known Because of the proposed presence (see below) of a hesperornithiforms in the collections of the Natural second species (Hfipevovnis sp.) in the same locality, History Museum of Kansas University it was the inclusion of the two vertebrae and the pedal reidentified as representing a bird more closely related phalanx in the referred material of H. vossicus is to Pavabespr.vov~ri~~,on the grounds that it shows few entirely arhitrary. differences between the lateral and medial condyles, no distal projection of the medial condyle, and the Hr.~prrornissp. remarkable medial position of the extensor groove. .I fragment of the proximal end of a left tarsometatar- The transverse width of the distal end of the tibiotar- sus from the same locality and same beds as the holo- sus (Figure 27.5) is 11.7 mm, thus this was a small bird. Subfamily :lsiahesperornithinae Nesov and locality (Ncso~.,1992~; Nesov and Prize~nlin,1991). Prizernlin, 1991 Cow~?tztvat.r.The distal portion of the right tibiotarsus Ari~~b(,.r/)c.,.oi*~ti.iNesov and Prizernli 11, 1901 and the thoracic vertebra previously assigned to .-f. Typt, .rpecic:c Asiabr~.rp(~vovni.rbuzbu?zoai Nesov and buzbatrovi\vere later illustrated as 'hesperornithiforr~~s' Prizen~lin,1901. (see caption to fig. 5 of Nesov and Yarkov, 1993) and, ir~ />iag?zosi.r. Medial condyle of the tit~iotarsusmarkedly the same caption, the proximal portion of the left tar- ~ncdiolaterally cornprcssed, cranial intercondylar sometatarsus was provisionally assigned to another furrow comparatively deep. 'l'he tarsometatarsus is species. comparatively gracile and has parallel lateral and The shah of the tarsometatarsus of .4siabcspri-ovnis medial sides. Both the lateral and medial crest on the bazbunovi exhibits a distinct transverse compression, plantar side of the tllrsometatarsus ha1.e a sharp an acute medial plantar ridge, a strongly reduced facet plantar margin and the flexor groove is shallow. The for n~etatllrsalI, and a markedly craniocaudally com- dorsal facet in the middle c;f the tarsometatarsal shaft pressed distal end of the tibiotarsus. The restored is deep and narrow, and covered hy a high dorsolateral length of the tarsometatarsus is 122 rnm (Nesov and crest and the separate medial facet is de1,eloped on the Prizemlin, 1991). The hesperornithiform frorn distal portion of the shaft. The base of the trochlea of Kushnlurun inhabited the Carnpanian Turgai Inferior the fourth digit is much larger than the base of the Seaway which ran frorn the Polar Ocean to the trochlea of the third digit and the fossa for metatarsal 1 Southern Ocean between Fennoscandia and Eurasia, is very small and short. and was in some ways analogous to the Western Comnzo~ts.'l'he diagnosis gi1,en above is abstracted Inferior Seaway which dilided North r\rnerica in the from the author's original diagnosis and omits some Cainpanian [Nesov and Prizemlin, 1991; Nesov, characters which appear to be characteristic for hespe- 1992a, c). rornithiforms. However, further evaluarion, via direct comparison with remains of Hespci-ovnis is required to 1;amily Baptornithidae American Ornithologists' identitS those characters that are derived for this new Union, 1910 taxon, especially in the contest of some of Nesov's Diagnosis. See Martin and 'l'ate (1976). rernarks that several remains of A.riabespevov?zis can be Co~tte%ts.6laptovni.r hIarsh, 1877 (Coniacian, USA) and assigned to other taxa (see below). The following char- .~udi?aovnis(, Mongolia). acters show that in any case Kushmurun remains Commc~rts.3uditrornisNesov and Borkin, 1083 does not are those of hesperornithiforms: the bones are heavily exhibit any of the characters listed in the diagnosis constructed with well-expressed pachyostosis, a trans- given by Martin and Tate (1976). However, Martin versely coinpressed tarsometatarsus, and a strongly and Tate (1976) noted a small pit lying directly ante- developed fourth trochlea. rior to the diapophysis in the trunk vertebra of B[zptovnis advmus and this is also present in .yudinoi-xis Asiabt~spevov?zirbazbanoi-i Nesov and Prizemlin, 1991 nc~go~ttsaven.ri.rNesov and Borkin, 1983. The flat ventral Holot~pe.IZASK 5/287/86a, shaft of a left tarsometa- side of the body is another character which might also tarsus. Priozernyi Quarry, Kushmurun locality, near be apomorphic for the Baptornithidae and circular the settlement of Kushmurun, Kustanaiskaya pits in the articular surfaces of the centra of the tho- Province, North Kazakhstan. Eginsai S~ita[Latest racic vertebrae are also found in both 6laptovni.r and Santonian-Early Carnpanian). 3udinov~ais. Diagnosis. Same as for rhe genus. Rq'ivved matevial. IZASK 51287186: shaft of a right tar- j7udizovzi.r Nesol and Borkin, 1983 sometatarsus, two thoracic vertebrae and a fragmen- Typr .cpecies. 3udi?zovni.r ilogontsavr~t~.i.rNesov and tary distal portion of the tibiotarsus from the same Borkin, 1983. s! am018 asla.\suell fascj~nsleluels SI! uo als~acln~ q~!~u!ls!psJalscleqs asaqL .papucdxa .4[as~a;\sue~1s! ~su!ls!pc 1111.n pu~padola.\ap1I18uo.11~ SnJawnq aql jo ~jeqs[esJelelaw aq~put! IJOCIS puc Inols s! snslelelaw pua ~w!xcx"~dsq~ jo aSpa le.~luaafu!q~ put. Suol apelq -0s.1t.1 aql 'le~aua8u1 .I Iwuuaw .mj lascj e jo uo!~ ~elndessrycqs aql 01 ~e[ns!puad~adssaso~d plosel -!sod ~w!xo.~daql puc 'calq3o~1puosas aqljo uo!~!sod -oso~dap!.n e Ilaaq e ql!m wnulals !sno(aoso~a~aqaelcl ~LU!XO.KI q8!q aql 'yeqs IRSJRI~I~LU~~J~Iaql jo uo!nas -alJa;\ 1mLAlas isp~!q 8u!iip IIELUS '.r!.rw3~"!p /).)/)tl.~rr~- -ssols pauysu! ue Su!pnlsu! 'sau~loj!ql!u~o~adsa~ ;861 'u!qqsoln~aep!uo!qmy ,il!u~ed aql jo s.~alse.~eqspaapap smoqs uaru!sads s!qLl2 ;86 1 'u!qqso~nxsamloj!llo!qul\: laplo '(9'~za1118!.4) ~-[h++NI~ 'S~S.IFI~~~IIOS.IW ~I[CIUSe jo utr!l~odaB~e1 c jo sls!suos Iepaleul psoj aqL .!qo9 '(q 'USO(,['u!qqso~nn) ado~ng pue e!sy le~luan q~no~u! ;\es~ u!!Scg jo spaq (ue!~qs!~~seew)18au1ap~ jo snoaselalg aql rr! punoj uaaq osle seq dno~8s!ql loj aql uloy pau!elclo se~isnossela.~n ue!loSuo~aql asuap!aa pue aumog put: auasoaled aql u! pale!pel JO SU!Sl" 121k?M lO!lalU! 3qI U! S~UIIO~!~~!UJO~~~S~H aeq~euBoae~edsq11nq1 asuap!Aa poo8 sf a~aql'lrrasald II~ISjo asuaso~d aql jo aJuap!Aa laqund

IV '(95661 'u!?lq""-Jn?l !8861 '9861 'lj"'"~" 3~961 'AOU L[!uJu,~ 'qmg) aJnlsnJlsoJs!m llaqsBBa put JolAeqacl se [lain saw~oj!ql!u~o~adsa~ se 's~alseleqsJelnsaloul pue 1m1Boloqd~o~1sno!leA .4q pal~oddnss! aeqlcu8oaelt.d aql jo .41dqdououl aqL .sru~cj!q~!u~o~adsaqjo s!ls!lalseleqs ale 006 1 '~jus,idaeq1eu8oaeled sselsaled rl-7tq~nsasejlns ~elns!l.~eppne3 PUC le!ueJs paprredxa [(,8 [ 'MCIPI?~ SqI!UlOaN SSt?[3tXjUI .i~asla.\sue~~pue 'slao3o~nald daap AJ~A 'sss.4 -cldodcdLiz le!ueJs rnolleu 'Lpclq aql jo asylns leausa ,]ueloA a1a.n ,iaq~ Iepne3 papuedxa ,ila,\ e s.noqs elqal~aaaqL .sasy.~ns Ieql ,i~!l!q!ssod aql l~oddnsqu~q pu!q acp jo sauoq .1e1ns!1.1eaq1 uaaxlaq q18r1al u! rrrw 1.b~Yu!.~nswu~ paz!~curnaudpalsnllsuo3 .411q8!1 '~~WIIM~~J,.es!Janl\: eJclalJaa 3!se.1oq1 ad,i~aql jo .ipoq [e~clal~aaaql 'pqq q1.10~pue e!lofluol,q jo us!~qs!.~~scel/t.IUIU~U!IU~~ paz!s-a[pp!ul e s! 11 .ppo~p10 a41 u101j paplosal acl o~ aq] n! punoj ale su~~oj!q~!u~o~adsaqI[~!ILIS 'snq~ r~r~cy!q~!u~o~adsaq1SJy aql S! J'!J'~U~U~~UO~IIU~,22~0~,2ptf~ '(3 'cz(,(,[) :IosaN .icl paqsqqnd ,asuap!~a ~alse~cqsLue ]nor11!,n Incl '(el 'el661 Al~uanhasqnsaa~a~'(a~ocle aas) uc~s~lqczeycl1.10~ '9861 hosa~)aep!q~!u~o~deg aq] 01 pallajal st:.% 'as~~!.\o~dedt:qs!eut?~srl~ u! ',i~!lcsol unlnruqsny I! '.1alr?1 '(€861 'u!qx)g put AOSJN)~NJ.I~?J!!J~CJC~:~ fo (uyuedruen .~I.IR~-U~UOIU~S~sa~el) ~I!AS aql 01 pauajal ,illeu!8!~o se,~s~suanvsi~~ollou .r!ir.rorc!l7n~'

!csu18:4 aql u~oy'srrr~oj!q~!u~o~adsaq 11ews jo spJosaJ 't:lrla~~a~~!~c.Io~I alYu!s c uo paseg SIN~LULUOJ laqllng 'VS~aq] puc epeuF3 jo spaq ue!~qs!.~~see~q mua8 loj SR awes :r,rsou.?u!g pue us!uedulen ale1 aql wo1.f sru~oj!ql!u~o~adsaqjo ~snoaselaJ3ale1 'uo!]eur~o.q I~~UI~N sauoq IleLus IclaAas jo 'suo!~sal[oswnasnru ues!laulv .u!l08uol~ ulalpnos "8eru!y ~ofluoqque.ieg '!(lo3 ql~o,y n! 'li~s.\oss!p s!q uo paseq SEM uo!snlsuos !~?IIY-SU~J,I, aql jo eaJe ula]sa.% 'ii~!lcso[,\t?s,l, uoo8o~

"q,~.'(ez(,(,~) ,\oss~-\cl px~~~t?~~"ISJLJEF" w.~oj!q~!u .r.lqallaA 3lse.Ioql a]a1druosu! '(,8{$ ~d 'adfiolo~

-1o~adsaqlue[o.z ,ilq!ssod '~leu~sjo asuals!xa aqLL $86 [ '~!qlogpUP.AOMN J~IS~/,?~~U.L'~UO~'OI~f~.lOzl,2/7tf~ xaplo s!q~jo sa,~!~t:~usssldalu.%orrq woy luaJa.g!p lc1l.liawos oslc Aluo sa!sads adh~urt,~iuo,-j ale sl!s~!~asaqj> .8ew!y (!qoy> 111nos) ,.\oY\?uurg u! .au!l~aluasaql uo ~aq1aBo1 nllqsncln ut!t?dt?s,~,put? at:s~n!!I!Jny> jo spaq 18alua~ asols palesol alt: sas;iqdode8Az Ie!ueJs aq] put: lillep ar11 u! :.ldly~iaq~ .iq pa~xllosa~a~ s~u.~oj!q~!u -nes peolq LJ~AInq a[pp!w aql u! pa.%oueu .4llsu!1 -~o~adsaqI~L:LLIS 8u!luasa~da~ (a[q!pueur t: jo uo!~~od -s!p s! runlluas aq] jo apts 1t:lluaA aq.1, Alas~aasue~~ aq~puc elqalla les!.uas c) su!eruaJ ~aql~njo.nL~ ,w~oj pua~xapuc padcqs-p!ozadcl~ ale elqallaa s!sc~oq~ -!q~!u~o~adsa~lllelus jo uoxel alc~edasc se p~!qs!q~ aq] jo uInJ]uas aq1 jo sasqlns ~elt1s!1~eaqL :~~zso~llvzg Figure 27.6. Left tarsometatarsus of a s~n;~ll,possibly volant hesperorr~itliifi>rmfrom the Nernegr Formation (Late Cretaceous) of Biigiin Tsav localit): i)mniigov' Aimas, blongolia. Medial (A),cranial (B), lateral [C),and plantar (D) views. ;Ibbreviations: tl, facet for mrtatars:ll I; id, infiacotylar depression; tr2, base of trochlea fhrdigit 2. Scale bar= 10 Inm.

short, fixsa-like, and runs dorsoventrally; pneumotri- L. Hou and %. Zhou. I found that Ambiovtu.r and cipital fossa of the humerus not developed. 0togovni.r shared a number of' derived characters Content.r. Ambiovtus Kurochkin, 1982 (Neocomian, including: a thickened, three-edged acrocoracoid with h,fongolia) and 0togorni.r Hou, 1994 (Neocomian, an acute top; Bat, wide humeral articular facet of the China). scapula; ventral position of a small, short, and oval Comme?~rs.0togo~ni.r geng6i.1-i HOU, 1994 is based on articular head of the humerus; and a thin and long associated elements of the and shoulder intermediate phalanx of the major wing digit. These girdle (holotype IVPP V 9607), together with some characters provide evidence for a close relationship flight feathers, from the locality of Chabu Sumu, Otog between A~~rbiortusand Otogovni~,and tor the assign- Qi, Yikezhao-meng, in the Ordos Basin of Inner ment of 0togorni.r to the Ambiortidae (Kurochkin, Mongolia, China. The mudstones which yielded the 1990b). specimen belong to the Yijinhuoluo Formation of the The heterocoelous cervical vertebrae, U-shaped Zidan Group and are earliest Cretaceous or possibly fi~rcula,convex coracoidal cotyla in the scapula, and even Late in age. 0togovni.r was first identified concave scapular cotyla in the coracoid are clear evi- as an enantiornithid (Dong, 1093), but later assigned dence that Ambiortidae belongs in Neornithes. to Aves incevt~zc~edi.r(Hou, 1994). Assignment of this family to Palaeognathae is based In 1995 I had an opportunity to investigate the hol- on the adt.anced condition of the strong ventrocaudal otype of 0togorui.r gengbi.ri through the courtesy of Ur transverse processes of the certical vertebrae; the well Mesozoic birds of Mongolia and the former USSR developed, dorsoventrally compressed scapular acro- mion with the tubercle or prominence on its dorsal side, and the projecting ventral edge of the proximal end of the humerus which bears a remarkable cranial tubercle with a centre pit on its cranial surface.

Ambiortus Kurochkin, 1982 Typespecies. dementjevi Kurochkin, 1982. Diagnosis. Procoracoid process wide and long; scapu- lar acromion long and dorsoventrally compressed; deep groove along lateral side of the scapula; scapular blade narrow; short, fossa-like groove cranial to the tubercle on the projecting ventral edge of the proxi- mal end of the humerus; undivided capital groove in the proximal end of the humerus; metacarpals fused at the proximal end; intermediate phalanx of the major digit dorsoventrally compressed. Comments. Ambiortus represents the earliest known stage in the of neornithine birds. It indicates that early palaeognaths were keeled birds and good fliers. Comparison with the Lithornithiformes and the Ichthyornithiformes does not support the opinion of Martin (1991) and Elzanowski (1995) regarding the close relationships of Ambiortur to the Ichthyornithi- formes. Figure 27.7. Ambiortu~demenqbviKurochkin, 1982. Holotype Ambiortus dementjevi Kurochkin, 1982 PIN 3790-271 + combined with a mould of PIN 3790-272. Holotype. PIN 3790-27 1+, 3790-27 1-, and 3790-272, Scale bar = 10mm. portion of the articulated left shoulder girdle, the left forelimb, and cervical and thoracic vertebrae (Figures 27.7 and 27.8). Khurilt Ulaan Bulag locality, Central and palaeobotanists consider the Khurilt beds as Late Mongolian Altai Mountains, Bayankhongor Aimag. Neocomian (Zherikhin, 1978), Aptian (Krasilov, 1980, Boontsagaan Gorizont, Neocomian, Lower 1982; Dmitriev and Zherikhin, 1988), or just as the Cretaceous. (The age of the Cretaceous shales and youngest insect assemblage among the three Lower sandstones at Khurilt is disputed. According to the Cretaceous assemblages of Central Mongolia latest analysis (Sinitsa, 1993) the Khurilt beds were (Ponomarenko, 1990). Based on geological data, deposited between the Dundargalant Gorizont (Latest Shuvalov (1982, and this volume) assigned the Khurilt Jurassic) and the upper member of the Boijntsagaan and Kholboot beds to the Andaikhudag Formation Gorizont, which is of uncertain age, although, in that he dated as Hauterivian-. Kurochkin places, the Khurilt beds are overlapped by the (1999b) hrther discusses the age of the Khurilt and Khulsangol Svita (Aptian-Albian). The dating of the Kholboot beds and possible correlations with the latter is based on lithofacies data, fossil fish, mollusks, Chinese Jiufotang Formation, aged on the basis of and ostracods, thus the age of the Khurilt beds must be absolute dates. at least older than the Aptian. Palaeoentomologists Diagnosis. Same as for the genus. Figure 27.8. .-It~~biui-tu.~~~l~n~e~~fl;viKn~ocllkin,1082. Holotype PIN 3700-271 + c~)mbinedwith a mold of PIN 3700-272. .\bl)re\iations: ac. ;icrocoracoid; cl, cla\iclc; cm, carpometacarpus; fi, ic;~tlierimprints; 11, humerus, mj, ~najormetacarpal; r, radius; sg, scapular gle~~oid;st, ; tph, terminal (ungual) phalanx of nrajor digit; v8, eighth cervical vertebra; vl0, tenth cervical vcrrehl.a; 111,; ur, ulnare; wph, pruximal and interrr~edi~tephalanges of major digit. Scale I)ar= I0 I~III.

Comments. New preparation of A. ~/emc.~~tjr;~ishowsthat broken edges of the counterslab (PIN 3790-271) and the cipht and tenth cervical vertebrae have heterocoe- the slab bearing the distal portion of the forelinlb lous not a~nphicoelousarticular surfaces as previously (PIN 3790-272). Thus the carpometacarpus, radius, reported (Kurochkin, 1985a, b). Further investigation and ulna on the main slab (PIN 3790-271 +) extend to also led to the discovery of a contact betwcen the specimen PIN 3790-272. ANI~~OT~UJ.demt~~~rjtmi shows Mesozoic birds of Mongolia and the former USSR some characters that confirm the primitive condition Comments. H. eucretaceu was assigned to the C;ruiforrnes of this bird. The articular head of the humerus is oval scnsu lato, based on some (?) characters of the Rnlli and short, the bicipital crest and intumescence are (Nesov and Borkin, 1983; Nesov, 1992d), but this absent, the pneumatic foramen and fossa are absent; needs to be confirmed. More recently, this taxon was the shaft of the radius is rounded in cross-section, the erroneously assigned to the Enantiornithes by Martin major and minor metacarpals are long, of the same (1995a). Horeznruvis shows such characters of neogna- length, and of similar thickness, the intermediate thous birds as a completely hlsed tarsometatarsus with phalanx of the major digit is long and the major wing dorsal infracotylar depression and an intercotylar digit bears an ungual phalanx. prominence on the proximal articular surface. Itelationships to any extant birds are difficult to estab- Parvclass Pycraft, 1900 lish because of the fragmentary condition of the There are a few neognathous birds from the material. kio~vever,H~)vt~zmavi.c, which was about the Cretaceous of Mongolia, Uzbekistan, and Russia. size of the extant Gallir~ulachloropu.r provides good evi- *?1hey are mainly represented by fragmentary remains dence of the existence of neognathous I~irdsin the and most have not yet been described. However, they latest . provide important data on the distribution of neog- naths in the Cretaceous and on the existence of some Order Anseriformes Bechstein, 1801 extant orders of birds at that time. Family Presbyornithidae Wetmore, 1926 Genus and species nov. Order ? Bonaparte, 1851 Unnamed taxon Family indet. There is a somewhat damaged, but complete tarso- Hor~mnavisNesov, 1983 metatarsus of a presbyornithid from the Baruungoyot '[ljpe species. Horezmavis eocretacca Nesov, 1983. Formation at Uiiden Sair, 6mnijgorr', Bulgan Sum, Diag~~usis.Medial cotyla of the tarsometatarsus Mongolia (Kurochkin, 1988). This very small form, inclined dorsally and located markedly more proximal with a tarsometatarsus length of only 40.3 mm, is the than the lateral cotyla; intercotylar prominence low; only avian from the locality where the maniraptoran dorsal infracotylar fossa deep and elongate; dorsome- and the marsupial Asiathwium were dial margin sharpened; tuberosity for insertion of M. discovered. tibialiscranialisshort, high, and located in the proximal region of the fossa; the large vascular foramen on the Order ? Sharpe, 1891 lateral side and the impression for the ligamental Family ? E'regatidae Garrod, 1891 attachment on the medial side are almost symmetrical Subfamily ? Limnofregatinae Olson, 1977 with respect to the tuberosity; retinacula attachment Volgavh Nesov and Yarkov, 1989 located proximal to the ligamental attachment men- Typt, species. l/olgavi~~~arina Nesov and Ya rkov, 1989. tioned above, close to the dorsomedial margin; plantar Diagtrosir. Tip of the mandible strongly ventrally crest relatively weak (Nesov and Borkin, 1983). deflected. Comments. filgavis was originally assigned, though F~orezmaviseocretacea Nesov, 1983 with some doubt, to the , then later Holotype. PO 3390, proximal end of a left tarsometa- determined as a member of the Limnofregatinae tarsus. Khodzhakul locality, outcrop CS-20, near to which belongs in the Pelecaniformes (Nesov, 19g2d). the north-western end of the Sultan-Uvais mountain This conclusion remains to be confirmed. ridge, Karakalpakia, Uzbekisran. Middle member of the Khodzhakul Svita (Late Albian). Volgavis marina Nesov and Yarkov, 1989 Diugirosis. Same as for the genus. Holotypc. PO 3638, rostra1 portion of the lower jaw \\it11 t)otl~rnnii and a fragment of the surnngular. Family Kuszholiidae Nesov, 1092 h,lal:1\:1 l\:ano\~ka locality, Dul)o\~skii Ilistricr, k;r.izkolin Nesol-, I 992 \i)lgogr:id I'rovince, South Central Russia. Ch~artz- '1,j~pc.ip~~ic:~: Ku.rzkolia i~il.i~xiNcsov, I 092 glauconitc a;lnds of Latest kIaastrichtian or Danian. Corr~nrt.i.Only the t!>pe sllecics. I~ir7g~rosi.1:Same as ti)r the genus. Diagno.ri.i. Synsacrurn wide; transverse process of the Coirrtrtc,nt.r. The loaer jaw fragment of 1'. rnavi~zcr is next to I;~stvertebra on synsacrurn strongl\ developed about 27 Innl, as measused fro111 figure 1-1 ;I of Neso~. and stout; caudal plcurocoels sn~;ill-sized,but deep; and Yarko\ (IOSO), and thus it was a small L~ird.'l'he caudal articular surtllce large, wide :ind dorsoventrally inandih~ilarramus eshil)its the opening of the neuro- cclmprcssed; postzypapophyses of the last vertehra on \,nscular canal on the medial side :~ndsoine neurovas- syns:icrum very large; \,entr:il groove especiall!~deep cul~~rfi~ramina in a shallow grilovc on the lateral side. in articulated areas of the centrll; cenLrum of the third .-\ \.entrally deflected tip of tlie ~nandiblealso suggests vertebra from caudal end heavily dorso\~entrallycom- a strorigl!. hooked end of the upper jaw. pressed (Nesov, 1901). -I > he heds th;lt yielded this fossil, greenish, iluartz- gl~iuconitesaiids were originall!l considered to he Ku.rzkoli~rftlcvzgi Nesov, 1992 Latest Llaastrichtian (Nesol, and Yarkov, 1089; Neso\, Holo(l~p(,.PO 4602, caudal portion of the synsacrum. 100_70), hut arc now thought to Oc Ilanii~n Outcrop CBlL.52, Dzharakhuduk locslity, Na\,oi (PaI;icoct'~ie)(Weso\., 1088, lC)02cj. Ilistricr, Bukhara l'ro\,ince, Uzbekistan. Upper nieln0cr of the Bissekty Svita (Cooiacian). :\\;air/l.(.v/nl' .icadis K(;/i,~~-t,d~~ati~iol. Cranial portion of the synsacrum, T'1~ltnttni~i.rNew\, I002 PO 4623, from the s:ilne outcrop (Fiesol,, lOOZd, plate '1,)pc.i/)c~~i(~,~~. l)/~~tm~li:lis flafrn Nesov, I 992. II; j), and possibly some vertet)rac froin outcrops Dlr/g~to.ri.c Xliddle vertel)rae of siTnsacrulii heavily C:l31-l4~indCI3l-.i7. dorso\~entsallycompressed; vertel)rul foramen \cry Dincq~rosi.r.Same as for the genus. spacious in the middle portion of the synsncruni; CO~>IN~~Z~S.The holot\.pe specimen (1'0 4602) of hl douhle ridge along \~entralside of the synsacrum; tirc~~tyiWJS first figured (fig. 2, ?a-ij, under the n~~rliher pleurocaels \wry Io\v and short; dorsal area of the PO 3486, and identified, in the tigure caption, as the middle \.ertehrae of the s!,nsacrum hroadcnctl iNesov synsacruol of a large iclithyornithid from outcrop 1002). (:El-i2. k: tneflgi\~~sa chicken-sized bird which had a stout synsacruni with an enlarged third pair of trans- I'lrrt~/rtc/i~il.Pram Ne~o\~,I 002 verse processes. Originally placed in the Kuszholiidae Holo!~y)c. 1'0 4601, fragment of the synsacrum consist- itrl,l~vtne sl,rli~., Kuszholi/t was later :issigned to the iug of t\vo or three \,errcbrne. Ilzharakhuduk locality, l'atagopterygiformes (Ncsov and l'anteleev, l99.3), so Naioi Oistrict, Kukl~ara l'rovince, Cjzbekistan. kno\vn only from the Coniacian-Santorii;~r~of O~~rcropCRI-.i;i, lipper rnenll~crof the Bissckty Svita Argentina. 'l'his hvpothesis has yet to he \,erificd, bur it (Coniacian). should LI~noted that the svnsucra of A~iszholinand Dicrg1ro.ri.l: Same as for the genus. Pntnnqo~)tc]SSn laulloj aql u101j slaqleq ~!ssc!j jo plosal pape~apy

.pa,\losa~un su!t:ulaJ uaur!sads s!q~jo d~!luap!~t:lns!lled J~I lnq '~uatuud!ssc ue!.he SI! slloddns '( 100 1 ';rul.j t:.\ounzt:[r>) lidossols!~~UOJl3.312 ~II!UU~?~S.??U!SII 'U?III!SIC~SS!ql 10 uo!1~8!1sa~u!.naN .iut:ld e st: ~!ss(!j s!~ctud!ua s!q] papleBs1 oq.~(~80l) yso~ jo nm!snIsuos aql Ku!~~od -dns pue 'II~!,\CSI? uacc~!~)sdsS!~I jo UO!II?U~!S~Paql Su!

-lsa!al '?no~t~y.~.!ll.io.~uidjo sd!qsuo!lelal aql passnss!p (p~(,o~)AONN xuos (q 11alqnop s! In(] '1aq1t:aj t: jo pJosal umouy lsa!llt:a aql !ilq!ssod s! (8~61'UV!IIIV~~ (h.~za~nB!.d) uau~!sads ay!l-~aqil:?j s!qLl, ~zn.~wwo~~ ,pa~u!od.ipt:als a~t:sc1.1t:cl aql 40 spua !sqq sq~u! sauo ueql ~a2le[alt: ~jeqsaql u! sdes dlnd !a.Zlt:l alt: sdes d1nd sq~:~LIE[C~ [RIUO~!.IC)~ aqi u! paxaH ,i~qeas!lous! laqicaj ~qd!~aql jo ljeqs aql jo uo!~lodlels!p aq] !aut:.~lauu! aql u~q~la.noucu s! sue.? lalno aql !pa~s!.t~~st auea aql jo auald aql !~nojueq~ aloLu IOU s! allacu!lua3 auo u! sqleq 3111 jo Ja(lulnu aql !auea ~I~~~LLIOSB UIJO;~ sq.~t:q isdes d1nd aql ,iq pall9 s! qs!qm '~jeqscl.1~1 aql jo ap!s 1w1opaql 01 .I~:JU palesol saut:,t aql !saut:,t oi ,i~!.~t:[!u~!sawos Bu!,\i\\oqs snql 'pauapeolq put: pauaiit:~sq~t:cl jo sap!s lauu! put: lalno isdt?s d~ndaAt:q sq~cqrsalnqleq olu! dn yealq IOU op daq~asu!s IeH Ala~nlosqcscllecl aql jo &pa

:,nols e jo az!s aql 'p~!qa2.11:l .i[a,it~e[ax.J?SOIL~U?~ ,s!sselnl'a~~?l'e11a~ !t:sueqeleg .ueisqyt?ztq 'asu!,\c)ld luaym!q:) 'li~!lesol (t:y~o

-[!qq!lq) J!~II~'J~C[IWJ 1q.8!~ '~E/(~SZ NI~ .,~d(jo/o~ SL6 [ '~"1nt:x ?i?,OJU+l' S/ILJOdUJd

'~"111" HS'LLAO2UJd HL(,I 'ut:!~neasep!ql!uloal:ld S~ILIIE~ 8L()I '~~lnt:~S~UIl?+!ql!UlUat:ld llpJO XLhI 'uE!~"" Salll!UJUJL?Jd SSeIsqnS E.N. KUROCHKIN fossiliferous deposits of the first three localities helong South Mongolia. Nemegt Formation, Upper to the lowermost svitas of the Neocomian. In addition, Cretaceous (hlaastrichtian?). the Lower Cretaceous localities of Baisa, Ust' Kara, Refirred mat~rial.A skull fragment and part of an artic- Pad' Semen, and 'Turga in Transbaikalia, Russia, have ulated postcranial skeleton, MGI 100199; Tiigriigiin yielded about three dozen feathers. lsolated feathers Shiree, South Mongolia; Djadokhta Formation, Upper have been collected from the Late Cretaceous local- Cretaceous (Campanian?). ities of Yantardakh (north of the Krasnoyarskii Krai, 1liagno.ri.r. Edentulous maxilla; pronounced pectoral Khatanga River), Amka (Khabarovskii Krai, Okhotskii crest of humerus; single distal condyle of humerus; Region), Obetzautzii Creek (Magadan Province, ventral tuhercle of humerus pronounced; extremely 1-enkinskii Region) in Russian Siberia, and Taldysai short shafts of ulna and radius; very long olecranon (Dzhezkazgan Province) which yielded Crctaviculus process of ulna; carpometacarpus massive, short, savysuensis based on remains (Bazhanor; 1969; quadrangular with no intermetacarpal space; nianus Shilin and Romanova, 1978) and Tulkeli (Kizylordin digit 1 much larger than digits 2 and 3; claw of manus Province) (Shilin, 1977) in Kazakhstan. digit 1 robust; coracoid not expanded ventrally; Unfortunately, so far, Cretaceous feathers have pro- sternal carina thick; one posterior dorsal vertebra vided little information on birds of the period. biconvex and synsacrum procoelous; zygapophyses, However, the specimens mentioned above are pre- costal fossa, and transverse process of anterior dorsal served in very different conditions. Some represent vertebrae on same level; sharply keeled posterior impressions of the feather structure, while others centra on synsacrum vertebrae; elongate haemal show different kinds of mineralization of the feathers. arches; ischium estremely slender; robust and hori- Most specimens represent small body contour feath- zontally projected antitrochanter; two cnemial crests ers, but they do not show the microstructure of the on tibiotarsus; medial margin of ascending process of distal barbules or microscopic barbicels which are of astragalus excavated by deep notch; metatarsal 111 taxonomic signihcance in modern birds. Some speci- limited to distal third, triangular in cross-section. This mens from Gurvan Ereen represent flight and tail diagnosis is assembled from the original diagnoses in feathers, and some are represented by plumules. One Perle etal., (1993, 1994). specimen from Baisa shows color patterns. Comments. In recent years, there has been much dis- cussion about the relationships of ,lloironykri.r. I do not Fossil avian eggs- - consider it or its relatives to be avian but, because it has been assigned by some to rives, it must be dis- There are a number of fossil avian eggs and eggshell cussed here. remains from the Upper Cretaceous of Mongolia, Cladistic analysis has suggested that Kirgizstan and Uzbekistan (Chapter 28). falls within Metornithes as a sister-group to (Norell et al., 1993; Perle et al., 1993, Non-avians 1994; Chiappe et al., 1996). Art.baeopte~yx is excluded Class Reptilia Linnaeus, 1758 from this evolutionary lineage and assigned to the Marsh, 1881 seirsri Gauthier ( 1986) placing Archaeopteryx as hIaniraptora Gauthier, 1986 a sister-group to taxa including hfaniraptora and tra- Family Parvicursoridae Karhu and Rautian, 1996 ditional Aves, though in a recent paper Chiappe etal. Alorroiryku.r (Perle ctal., 1993) (1996) changed Avialae to Aves for the taxon Type species. Af oirorrykus olecvanus (Perle et al., 1993). Metornithes + Avchaeopteryx. At the same time, the Halotype. MGI 10716, posterior part of skull, most of cladistic approach utilizing the total group concept the precaudal vertebrae, all four limbs, thoracic girdle indicates theropodan relationships for 1210i~onykus and portion of ilium and pubis. Biigiin 'I'sav Locality, (Patterson, 1993a, b). Mesozoic birds of Mongolia and the former USSR

In spire of that, most palaeontologists using a Discussion morpho-phylogenetical concept of homology- analogy have found evidence of a non-avian, theropo- Feather records show a wide distribution of hirds fronl dan nature for .lfononykiis (Ostrom, 1994; Wellnhofer, the earliest Cretaceous in the western part of 1994; Kurochkin, 199.5~;Martin, 199%; Zhou, 1995; Mongolia and in the southern part of Eastern Siberia. Feduccia, 1996). The unambiguous relationships of Late Cretaceous feather records confirm the existence Mononykus to birds was established on five (Perle et al., of birds in Kazakhstan, Northern Siberia, Mongolia, 1993) or six (Chiappe et al., 1996) characters: promi- and the Far East of Russia. Unfortunately, feathers nent ventral processes on cervicodorsal vertebrae, reveal norhing regarding the taxonomic identity of longitudinal and rectangular sternum, ossified sternal these birds, but because of the wide distribution of keel, ischium more than two-thirds of puhic length, downy and fine contour feathers they indicate the carpometacarpus formed from fused distal carpals and extensive distribution of birds with a warm-blooded metacarpals, prominent antitrochanter, and short physiology in the Early Cretaceous. These types of tibula. All these characters can be explained as bipedal feathers may be associated with the origin of neogna- and digging adaptations (Zhou, 1995), and most are thous birds. represented among different velociraptorine or mani- By contrast, the contour for enantiorni- raptoran theropods. Such characters as a single- thine birds is problematic. Chinsamy etal. (1994, 1995) headed quadrate, biconvex posterior dorsal vertebrae reported growth rings in the femora of Argentinian and opisthocoelous thoracic and cervical vertebrae, enantiornithines indicating cyclical growth of bones articulation of the cervical ribs at the same level as the during life and slower growth rates than in extant cranial zygapophyses and transverse processes, an birds. This also provides indirect evidence of their obtuse apex of the scapula that should he directed physiology, thus Chiappe (1995) and Chinsamy ct a/. dorsad in natural articulation to the coracoid (the (1995) have suggested that enantiornithines did not position of the scapula should be vertical, not horizon- have endothermy or ectothermy, hut some kind of tal as it was figured in the original reconstructions intermediate level of metabolism. In any case, as this (Perle et al., 1994 figa. 9 and 20)), a wide coracoid level of physiology is more similar to a reptilian mode lacking an acrocoracoid, a single distal condyle of the of physiology, than an avian one, this may be sufficient humerus, femur longer than the metatarsus, and with a to reject the idea of a good covering of contour feath- fourth trochanter, and unhsed n~etatarsals wirh a ers in the enantiornithines. proximally declined third metatarsal all clearly point Osteological remains are more restricted than to the theropodan nature of .Ifononyku.r (Kurochkin, feathers, both in terms of their geological and geo- 1995~).lndeed Ostrom (1994, p. 1721, concluded that graphical distribution, but they can be attributed to 'Mono~ykuswas not a bird, . . . it clearly was a fleet- particular avian taxa. A small enantiornithine was fossorial theropod.' present in the Early Cretaceous of Mongolia and Recently, Karhu and Rautian (1996) described a during the late Late Cretaceous (Coniacian- new maniraptoran, Paniicuvsov vemotus, from the Late Maastrichtian) small and middle-sized enantiorni- Cretaceous of Mongolia and discussed in detail the thines inhabited hlongolia and Uzbekistan. Their arctometatarsalian type of metatarsus, Parvicuvsov is small size and long, curved pedal claws suggests that extremely similar to Mononykus and both show the they may have been primarily arboreal birds. arctometatarsalian pes with a proximally declined Large, flightless hesperornithiforms are kno~vn third metatarsal. On this basis, Karhu and Rautian from the Campanian of Russia and Kazakhstan, while (1996) argued that arctometatarsalian theropods small and possibly volant hesperornithiforms have are not closely related either to Sauriurae, or to been collected from the Santonian-Campanian beds Ornithurae. of Kazakhstan and from the Maastrichtian beds of L.N. hURO(;tIKlN hlony~lia.'l'his is supported hy data horn North century. Ho~vever,they provide good evidence for a .\merica concerning the existence of previously greater avian diversity at higher taxonomic levels in unknown smal I rcprescntati\,es of hesperornithiforrns the Mesozoic than in the . So, in the in the interior basil) of at the end of Cenozoic, only two evolutionar!r lineages: the Cretaceous. Palaeognathae and Neognathae of the inhaclass 'I'he Palaeognathac is a separate linca~cof 11corni- Ncornithcs survived. During the Cretaceous, at least, thine birds that n.as represented in the Early there were five major phylogenetic lineages: the Cretaceous of hlongolia by dit/biovru.l-. In an adjoining Enantior~~ithcs, Hespcrornithes, Ichthyornithes, rcyion of C;hina, Otogovtiir, a close relative of';Imbiortu.r, Palaeognathae, and Neognathae. has heen reported from the earliest Cretaceous. The Ncognaths fbrnn a fourth group of Cretaceous Acknowledgements hirds. 'l'hey Lvere present in the Early Cretaceous as fossil records from .Asia and show. New data I thank L. Hou and Z.Zhou fix showing Inc thc spcci- on birds tio~nLiaoning in (:hina also contir~nthe cxis- men of 0togort~i.sg.f~,ghiri and fix discussion of this tence of' neopnathol~sI~irds in the tarly Cretaceous fossil, I I. Osmhlska anti K. Sahath for the possibility to (I lou cr ~l.,1000). 111 the Late Cretaceous Nc~nc~twork on specinlens of Cobipte~y.~and the enantiorni- 1:ormation undescrihed remains from Mongolia indi- thine e~nt~ryos,A\.\'. I'antelccr for additional data on cate the existence of charadriifi,r~ns(Graculavidae), birds from Dzharakhuduk, L4.\'. h'lazin of the I'IN anseriforms (I'resl)yornithidae), pe1ecanifi)rms (a cor- lahoratory of photography for the photographs of [norant), and procellariifi~rms (an alt~atross)in this specimens, I-. IvIartin for the idea of moulding speci- region. 'l'his is concordant with discoveries in North men PIN 3700-272 and for its fabrication, and 11. .lmerica and .\ntarctica. 'l'he fifth and last group of Univin and two anonymous rc\,icwcrs for critically .lsian Cretaceous hirds, the Kuszholiidae, of uncertain reading this n~anuscript.'l'his study \\.as supported, relationships, is known only tiom the Late C:retaceo~~s in part, hy the Institute of \'crtebrate Palaeontology of Uzbekistan. and PaIaeoantl1ropology, Beiiing, and hy grants h'lost of these avian fossils are represented hy frag- 95-05-l0919d and 90-04--50822 of the Kussian Fund mentary remains which mainly provide intormation for 1:undarncntal Kesearch. on the distril~utionof hlesozoic hirds. t lo\\yever, some 1. The species of ZL~j~v/o,rt~il-listeti on pp. 533-534 were of the Russian and klongolian Cretaceous hirds, such assigned by Nesov (1984) to tlie Ictithyornithifi)rmes, as .4it/biorrus and .Ymnt~riu.s, pro\,itle very important and subsequently to the Bnantiornitlies by Kurochkir~ e\.idence of the mode of evolutionary processes in (1996, this paper). Ho\\-ever, Lliyraornitliid~~edo not early hirds. .As 1 hypothesized (Kurochkin, 10853, helong to the Enantiornithes, or to any other known 1001), the knolvn record of klesozoic birds is more fossil or living group of birds, as investigation of these restricted than the true diversit\ of hIesozoic hirds. re~nainsin tlie TsNIC;lesbased on later hirds. New fossil data on hlesozoic birds coracoids mentioned on pp. 534-5'4.5 have beer] (Chial~l~c,100.5; Fcclucci~1,100.5; EIou ct ul,, 1005, 1006; described by P~~ntclccv(1008). Kurochkin, 1095~)confirm this assumption and these authors demonstrate a ti~rmerlyunknown Mesozoic diversity of I~irds. References Data on hfesozoic birds has only appeared from ~~kl~,,~i~~~,~\'.y.and ~~~\~i~~, 1),h,, 19~))..\ slIr\,ey Ofptero. Russia and hlongolia during the last Ikw decades: this saurs frorn the Jurassic and Cretaceous of the Forrner is in contrast to North America where Cretaceous Soviet Union and hlongolia. Ifi~iif~~iil.~IBiulo~~ 10: I~irdshave 11een collected and studied fi~rmore than a 197-145. 'hi l-ii [ 7 :O$ J;?.Hl2l.lS qlr/~,yjo ln~~rnogII~!~IIIII<) .eu!q:) jo s~lqndn~s'nldoa,~ '~:!~oSuo~y13~~1 jo UISI:~ sop10 aq] u~oljpl!

.c y nc.y :.+\o.)solys1.~.7.rri! .#I no!ln/oc,l acp

~IIIJs!.r~,f.l .?!~IIII,I.~II!~ X~III,~.~II~.~.(,) '[.pa) . !).k 'oqua.~cLLI~OU~)~ (I! s I z-$0; .d(l -sa~ua.1.1n22opa]clniu[l.>2c at11 jo pot11a111.icl elep at11 uo sa!l!u~t!j IJJSLI! LII .i~!sl.>,\ip 40 saduslp a11.1~1.$$(,I .,\.,\ '11!1{q1~a117~LII! .II.~;I ',\~!JI!CCI~ .ssa~duopualc1:j

:plOjX() 'vjc,llllYl/O 1 /"l,)i/,'; l~~JIJl.l~.~~~s.~~~NcN~~/s~~~'[ 70,/

.S~OI/UAJ,<1, /;I IIOIJ~I~I/IS~~I/:)p//wLi(,7Jo/liyi/ ~y 1, '(,pa) .f 1y

'uo~uaclUI .lid 'sp~~cl. .lo sap"> ~o'lc~uaq.~ '~~61-

'(,(,$-$s$ :Z I l20/0lql&l//ld '".Ilq jo uo!]osy!s~a\!p ,il.i~:a pur: u!d!.io aqLl, ,986 [ .[ '1jc.12e.13

'tLS-[')i :I ~l~"l"!l/~~~//~~'sLIo!IL!.~~[~Lu! IC.>!~~I[O!S~<~~

:a.1113311.13S0.1.711Ii 3LIOCl UCI.\C. .3IOZOS31\i. '

:h I-')(, 1 :89$.%l//L/l~v '">.l!q 2!0705'3]$ U! s;iu!l q~"<~.if)'t~f, [ .,I 'LIO+)O(I t~~rjy'l'acldc!q:) '.I- '.<~uesu~qn

' :$ $-L 5 1 :($ ) (j5 llln,l.rnl\' ~l/111/.1~//.1.11/~

.ql Jo .u~IIIIL.)~~,'IJasaCI !clot) jo snoasc1a.13 31s-1

all] luoy (aep!lnnzalcal\- :sa.\y) sn,pfituno~~~$0 uo!] -!sod 2!lairadol,iild '966 1 'J\T'['~.IL?[~) pUl? 'F'ly '113~)~'-

YjtZ-c Z i :+I ~t';z0]01/10,1/17[]

J jo /nr~,(noL' .(cu!~uaYly) u!uo31:1ed lo snons1:1s.13 .~add,qaql u~oy(sa\\-) saq~!ulo!~ueu:d nau c '.r/tu/on s.!/~rort,~ndtc,~~\- '0.1 '0.11~3 PUG - - :8Lf ,?,l"Uy 'UO!lIlIO.\a uc!,~~:jo sll:a,i uo!ll!m ~8 ]sly aqL ,5661 .~y'l'adde!qg y(); :.[~icl11s (5)+ 1 ,t~o~olllo'I]u[/

.E L-L y :L lXop,~q~,~~u~/OJ srto!/nq,1,1/no3 nn!/rosrl/,~lic~.(ac11?u!.1e3 :sa.<\-) saculog!s!d pue sa~u.~oj!!~elo-~Japlo aql 01 lellsasue 'il~ua

.~ccldc'p.r!q snoasclal:] I: jo .ila,\oss!a ,9161 'c1~ojpo~g '8 saaua!>S jo ,iurapcsy c!u.r

,jo rto!~tzlo.~~.?yl pnu .rp,(!g -/i, ~I,Z?Z,(~r>y '('pa) 'y 'us!psd u! 7~-isdd '11ld!u LIC!.ZC jo u!S!.~o 1salo(l1e aL1.1- .o$(,[- 'tS-hi .~~~,l.l~//O~)//>.l~~O/~i~J!l~,/() ~~llO~lfl/l,~,~Jli] IIJf 1 .~>'/l~/?~l,l.l~,l~

'S3l1lll~l!.. . .3l!llIl?)[ .l>llJt>I,\2 . 1l!llll!.I2 . 211.1, '$96 l .f,q'1208 .pe~du!r~a-~

'9-5 :U.!/~~,l//.Jl/.iq()~l.~~ll~.~:"/.l!~~]ll~~~ll,~/~l~ l)~~/f~/Z~b.l',l.~,/!!LC15 1.1, .?O/?II].YII(/ A:t.!r.;/, ['assn 2111 u! suua2claln a111 u! d~lI.\!lIllI!lLl2J t>.llil "I" P.lo.72.l 2ql "01 '6961 'S:\ '.\0~11?1[~1?~

XSSn L~UI.I(!J arp pus e1l03uo1.q jo cp~!cl~!OZ~SJ~T E.N. KUKOCHKIK

101-199 in Il'ichev, \:D. and Gavrilo? V.h.1. (eds.), Cretaceous. Couviev bbv~~chungritistitutSt,rirkcilbevg 18 1: '4cta XClll (,'oi~giz,ssusIntenzcctiona1i.r Ovnithologic.i, v. 1. 23-36. hloscow: Nau ka. -1OOib. The relationship of Mnnoi/ykus to ornitho~nirnid -19Xib. :\ true carinate bird from Lower Cretaceous . .~ounzccl of' firtchvute Puleo~ztolo~g 15 deposits in Mongolia and other evidence of early (3)Suppl.: 43'4. Cretaceous birds in Asia. Cvctucc.ous Keseavcb 6: -and Tate, J., Jr. 1976. 'l'he skeleton of H~ptovnisudvenrrs 271-278. (Aves: 1lesperornithiformes). Smithsoftimv Cnntributions -1088. (:retaceous hirds of Mongolia and their to P~ilt:obiolo~2 7: 35-66. significance ior study of the phylogeny of class ;\ves.] hlikhailov, K.E. 1091. Classilication of fossil eggshells of Tvudy So~~tnesttioiSovrtsko-:.Wnngolikoi Pa1mntulogichr.rkoi amniotic vertebrates. Palaenntologia Polor/irtr 36: Ek.rpedit.rii 34: 33-42. 193-238. -1 09 1. [Pvoroccvir,:171zbiorttr.r and other palaeornithological -1995. Systematic, faunistic and stratigraphic diversity of rarities.] Pvivodu 12: 43-53, Cretaceous eggs in Mongolia: comparison with 1993.['I%epvinriptristrzg~,~' in r~olutionof class.he.r.1 Thesis China, pp. 16.5-~168in Sun, A. and Wang, Y. (eds.), on the scientific degree of the Doctor in Biological Si.& Symposium o~iMesozoic 'I hve.rtria1 Eco.rystems and Sciences. kloscow: Palaeontological Institute, RAS. Hinta, Shovt Paprvs. Beiiinp: China Ocean Press. XIS, 64 pp. -1096. [Birds eggs in the Late Cretaceous of hlongolia.] -1995a. The assemblage of the Cretaceous hirds in .4sia, Palt~o~itologicheskii%birvi~~/ 1 : 1 1 9-1 21 . pp. 203-208 in Sun, A. and Wang, Y. (eds.), Sixth Molnar, K.L. IYSh.12n enantiornithine bird fioni the Lower Sl~nyosiunion :Clesozoic 'fivvestvial E[v.y.rrc.tn.r and Riotli. Cretaceous of Queensland, ;\ustralia. Natuve 322: Sbovt Pape~r.Beijing: China Ocean Press. 736-738. -1995b. Xlorphological differentiation of the Nesov, L.A. 1984. [Upper Cretaceous pterosaurs and hirds 1':llaeognathous and the Neognathous birds. Cuuviev from central hia.1 Pul~o~ito/~giCbe~kiiZhuvnal 1:47-c7. t.br.sr.hung~in.rtitutSntcketlbevg 18 1: 79-88. -1986. \The first record of the Late Cretaceous bird -1995~. Sy~~opsisof Mesozoic hirds and Early Evolution Icbt4yorni.r in the Old World and some other bird of class .4ves. .-lvchaeopteyyx 13:47-66. bones from the Cretaceous and Paleogene of Soviet -1006. il Icc>rn t'ir~i~tiov~~ithidof the Mor~golianLate Cvetaccous Middle Asia.] fiuky Zoologirbeskogo Imtitutu, A.V SSSR ~ilda ge~~evulrrppvazsal of the infiacb.r.r b;nantiov~zitbr,s 147: 31-38. (Awed. Special Issue. 50 pp. hlosco\r: Palaeontological -1088. [New Cretaceous and Paleogene birds of Soviet Institute. hliddle Asia and Kazakhstan and their environinents.] -1099a. [A new large enantiornithid from the Upper Tvudy %onlogirAn1.kagoIn.rtituta, AN .CSSR 1 8 2: 1 1 6 12 3. Cretaceous of hlongolia (.4ves, Enantiornithes).] -1990. [Small Ichtbyovnis and others findings of the bird 7 kudy Znolagicbe.rkogo Institutu. K:1N 2 77: 1 30-1 41. hones from the Uissekty Svita (Upper Cretaceous) of -19991>. The relationships of the Early Cretaceous central Kyzylkurn Desert.] 'li.udy Zoologirhe.skop ilnibiovtus and Otogovnts (.lves: .4n1I)iortiformes). I~~sritutu.AN SSSK 2 10: .5%62. Smithsoniccn Contrib~rtioizsto P~leobiolo~g89: 2 75-2 84 1902a. [Flightless birds of meridional Late Cretaceous hlartin, L.11. 1983. The origin and early radiation of birds, sea straits of North .\rnerica, Scandinavia, liussia, and pp. 291-338 in Rrusl~,All. and Clark, (;.A. Jr. (eds.), Kazakhstan as indicators of features of oceanic circu- Pei-.rp[.c.tive.r if? ovnithology. Cam bridge: Cam bridge lation.] Byulleten' Moskovskogo Obshchcstva I.rpytatclti University Press. Pvivody, Otdel (;eolnL~irl~tskii6 7 (5): 78-8 3. -1984. :\ new hesperornithid and the relationships of the -1992b. Mesozoic and Paleogene Birds of the USSR and h'fesozoic birds. Tvttn.qcir.tions of the Kansas Acrmhn!y oj' their paleoenvironments. Natuval Ilistory Museum, Los Science87 (34): 141-1 50. Angc1c.r (hnty,Science Sevies no. 36: 465478. -1991. hiesozoic birds and the origin of hirds, pp. -1992~. liussia, p. l 3 in Mourer-Chauvirk, C. (ed.), Society 18.5-540 in Schultze, 13.-P. and Trueb, I-. (eds.), oJ' dilirli~pale onto lo^ arid Evolution, In@vmation I,ett~v,6, Origins of the highcvgvoups of tc~tvr~pods,contvuyy aad cow November 1992. .qcnsu.r. Ithaca: Cornell University Press. 1992d. [Record of the localities of the Mesozoic and 1995a. The Enantiornithes: terrestrial birds of the Paleogene with avian remains of the USSR, and the '62-1 :SOT f S,li"I"!tlU~\r UIn,?.l'nl4' iLtl.l~A21N~'e!l08uoly

jo snoasclaln ale-I aq] cuo.~j(acle!.tv :t?p~do.~aq.l.)J.ritr .fY6-850 v-r.7~p~nqrli~o1ru~l1jo ddoloqdloru [e]a[aqs '+hf,[ '~'l\i :Z[I .ynl-, ;pl!cl I: ~'v,ptiioiroj~.s[ 'chhr .sqBuoq% 'noqz '1p.10~pu': .~y.['7.1~13 ''8 'ploqueg '.~\i.l'adde!r[3'-

'86 1- 1 :591 y,y~~,\.F, c~~in~~i.s~q ouXuqs,~y~!~o~oit~o,~~nd 'YZY-f 5') :Z9f.~-"'lvI\l

~tj~n-tl,[.saxald~uos cuncj s!ozouan puc sntrasolsl3 ,e!loduoly jo snoasclal3 aq] moy pl!q ssal1q8!1~4 aql jo ~uat~raseldalpuu uo!lcrulo,11 [ ;\:\ 'u!~[y!laq% 'f 661 .[\i.f'y~":) pUE '1q.1 'add~:!q:) '.v'\~'[[J~oN "\: 'allad 'SOL-hh; :6If '815 3'1,171d ~~~l,lU,l~.l~.i,l/J ,J~1il~/JlJ.lfJ/ 2fJ .SYIpLL,la S,?~l~li/O,')'SpJ!q :991 ,IAniv,y .sa!ldal uoslallud .saurcu Sn!u~e~'q~661- lo uo!~nlu,\a.il1ca pueu!B!.~o aq] uo elep xaK .i-661- '5i-1; :59f,i~niv~"\; jlnesou!p~o p~!g'cthh 1 '3'uoslal~ed '08 1-58 .( I -f :S f q.r~d.Ah-.rs~~(/sqy

:S 8 1y Irrn/iazyl,. '~3!y(/v~Soi1ro,in~1~d.xL;LJ~(/o,~I~~~A~ 7U2Utl'/7 !!$~".7~.7!~0~01i!1U()Ilq.rJlIH ' [~llnqlhdxI EJ1113:) uo.\ ~cldruaxal~alaql;allu!g sca 'b~61a 'lajoqu[[a~ lo snoaselal3 laddn ILI~~(saq~!ulo!~ueu~ :sa.\v) 'fc-15 :z6~.1,1ni1l.y.m!Jatu\. [[]nos jo snoasela.1.j sau!q]!ulo!lueua jo sa!.7ads zl\a~],866 1 ;,\.v ',taa[a~ued aql tuoy spl!q jo sselsql\s nnax '[#()I '\-3 '.~a~lch\ 'ssald aassauuaLLlo .;\Fun

.11e~pug LI~NI~C~~):uopuq ~JAO.I,I~~~SS~!~A/./, :a]1 !.IXOU>I 'L"!~I"U

't[ 1-91) 1 :t1"IlAn'lZ oXoq.rnq47!Bo~oo~tlj~i~~~ ['sanv sse13 jo uo!]nloAa pue rrg.r,~

.sasua!ss jo imapcsy '60 1-80 1 19 1 [ flSS ,&I,' '%iYi2!iJIL/ o~o,~s,~~.~!~o/ou;I~(/I~l-l~ ass,? a111 jo qsuelH ulalscg 1e.q :qo~s~o.z!peln [TITa1pp11y la!nos pue t?![o8uol\l jo snoase1aJ:J aq~ ')/SLY,1 ,?<71 40 S'?iU.J.l8Z,lAJ ~ll~/l.l7L~l/if~~)'('\la 1 'v;!l ',\01!~1?1~ lllolj SaUOq pl!Xl j0 sRn!puy XJN] 'f 86 I .f1 '~!710gPUG

"1 80 [-SO I .

~ISS~JauLJoj aql put: r!~oSr[olyjo sp~!cls!ozosajq