Page 1

2012 年 4 月

地 层 学 杂 志

第 36 卷 第 2 期

Apr., 2012

JOURNAL OF STRATIGRAPHY

Vol. 36 No. 2

① 国家自然科学基金(No.91114201)和国家重点基础研究发展计划(2012CB821906)资助出版.

文稿接受日期: 2011-08-17; 修改稿收到日期: 2012-03-31.

第一作者简介: 博士, 研究员, 主要从事古脊椎动物和地层研究; E-mail:dzharakuduk@mail.ru

中亚与亚洲中部晚白垩世的陆生脊椎动物组合对比①

A. AVERIANOV 1)

H.-D. SUES2)

1) 俄罗斯科学院动物研究所 俄罗斯圣彼得堡 199034;

2) 史密斯森协会自然历史博物馆古生物学部 美国华盛顿特区 20013-7012

摘 要: 通过对前人建议的 26 个生物地层标志化石存在与否的简约分析, 中亚与亚洲中部晚白垩世的陆生脊椎

动物组合的相对层位得到了更清楚的揭示.此区最古老的组合是乌兹别克斯坦克孜勒库姆沙漠的 Khodzhakul 组

合(早塞诺曼期), 其次为蒙古戈壁沙漠东部 Bayn Shire 组的下部和上部的组合(塞诺曼期至桑顿期).中国内蒙古二

连达布苏动物群与中亚的土伦期—桑顿期动物群属于同一类群, 因为它们均具龟鳖类 Khunnuchelys, 前者时代可

能为桑顿期.三个中亚的组合(Bissekty,Yalovach 和 Bostobe 组合)中有两个地方性的鳄形类(Kansajsuchus 和

Tadzhikosuchus)和一个出现于戈壁沙漠的鳄形类(Shamosuchus)化石.戈壁沙漠的坎潘期至马斯特里赫特期组合与

北美同期动物群为同一类群.Djadokhta 组与 Barun Goyot 组的坎潘期脊椎动物组合具有高度的地方性, 并反应了

半干旱的古环境.产自 Nemegt 组的组合生存于比较潮湿的环境.在组成上,这一组合与其他河流相沉积环境

(Bissekty,二连达布苏以及北美 Judithian 期和 Lancian 期的组合)相似.具顶饰的鸭嘴龙 Saurolophus 的存在, 支持

了 Nemegt 组合为马斯特里赫特期时代.戈壁沙漠的这三个组合(Djadokhta,Barun Goyot 和 Nemegt 组合)被归为

一类, 因为它们共同拥有地方性的龟类 Mongolemys 和兽脚亚目的小驰龙类.亚洲中部和北美的坎潘期至马斯特

里赫特期组合与亚洲更加古老的组合不同在于存有暴龙科,肿头龙亚目和鸭嘴龙科.在中亚, 由于地区性的海侵,

这一时间段内的陆生脊椎动物组合多不清楚.

关 键 词: 生物地层学, 脊椎动物, 晚白垩世, 中亚, 亚洲中部

中图法分类号: P 534.53

文献标识码: A

文章编号: 0253-4959 (2012) 02-0462-24

CORRELATION OF LATE CRETACEOUS CONTINENTAL

VERTEBRATE ASSEMBLAGES IN MIDDLE AND CENTRAL ASIA

A. AVERIANOV 1) and H.-D. SUES 2)

1) Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, Saint Petersburg 199034, Russia,

e-mail:dzharakuduk@mail.ru;

2) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution MRC 121, P.O. Box 37012, Washington,

DC 20013-7012, U.S.A.

Abstract The relative stratigraphic positions of the better-known assemblages of Late Cretaceous continental verte-

brates from Middle and Central Asia are assessed by parsimony analysis of the presence/absence of 26 proposed bio-

stratigraphic marker taxa. The oldest assemblage in the region is Khodzhakul from the Kyzylkum Desert of Uzbekistan

(early Cenomanian). The next stage includes assemblages from the lower and upper parts of the Bayn Shire Formation

of the eastern Gobi Desert, Mongolia (Cenomanian to Santonian). The Iren Dabasu fauna from Inner Mongolia, China,

clusters with the Turonian-Santonian faunas from Middle Asia based on the shared presence of the trionychid turtle

Khunnuchelys and is likely Santonian in age. Three Middle Asian assemblages (Bissekty, Yalovach, and Bostobe) are

endemic in the presence of two crocodyliform taxa (Kansajsuchus and Tadzhikosuchus) but share another crocodyliform

2 期

Averianov A.等:中亚与亚洲中部晚白垩世的陆生脊椎动物组合对比

463

(Shamosuchus) with the Gobi assemblages. The Campanian-Maastrichtian assemblages from the Gobi Desert cluster

with coeval North American faunas. The Campanian vertebrate assemblages from the Djadokhta and Barun Goyot for-

mations are highly endemic, reflecting semi-arid paleoenvironments. The assemblage from the Nemegt Formation,

which existed under more mesic conditions, is similar in composition to those from other fluvial depositional environ-

ments (Bissekty, Iren Dabasu, and North American Judithian and Lancian assemblages). The presence of the crested

hadrosaurine Saurolophus supports a Maastrichtian age for the Nemegt assemblage. Three Gobi assemblages (Djadokhta,

Barun Goyot, and Nemegt) are grouped together based on the shared presence of the endemic turtle Mongolemys and

parvicursorine theropods. The Campanian to Maastrichtian assemblages of Central Asia and North America differ from

the older assemblages in Asia in the presence of derived Tyrannosauridae, Pachycephalosauria, and Hadrosauridae. In

Middle Asia, continental vertebrate assemblages from this time interval remain largely unknown due to regional marine

transgressions.

Key words biostratigraphy, vertebrates, Late Cretaceous, Middle Asia, Central Asia

1 Introduction

Correlation of the Late Cretaceous continental de-

posits of Central Asia to the Standard Global Chro-

nostratigraphic Scale is difficult due to the absence of

biostratigraphically useful fossils. However, the con-

tinental Late Cretaceous strata in Middle Asia are

sometimes intercalated with marine deposits that do

contain such fossils. This permits more precise age

assessments for these units (King, personal communi-

cation). Furthermore, the Late Cretaceous deposits of

Middle Asia contain dinosaurian assemblages similar

to some extent to the better-known diverse assem-

blages from the Gobi Desert and other regions of Cen-

tral Asia (Nessov, 1995, 1997; Averianov, 2007b; Sues

& Averianov, 2009b). This combination of factors

provides a unique opportunity for assessing the ages of

the assemblages of Late Cretaceous continental verte-

brates from Central Asia.

We follow the geographic usage of the term "Mid-

dle Asia" to specifically denote the region within

"Central Asia" that comprises Kazakhstan, Kyrgyzstan,

Tajikistan, Turkmenistan, and Uzbekistan (Dani &

Masson, 1992).

In this paper we review biostratigraphic markers

that are useful for dating the assemblages of Late

Cretaceous continental vertebrates in Asia. Selected

marker taxa are employed for a quantitative (parsi-

mony) biostratigraphic analysis of the principal verte-

brate assemblages. The recovered pattern of correla-

tion between the analyzed assemblages is used to as-

sess the relative temporal succession of the various

faunas of continental vertebrates from the Late Creta-

ceous of Central Asia.

Abbreviations. BM-biostratigraphic marker; PIN-

Paleontological Institute, Russian Academy of Sciences,

Moscow.

2 Methods

There are a number of quantitative approaches for

identifying areas of endemicity in biogeography (see

Simpson, 1960; Cheetham & Hazel, 1969; Raup &

Crick, 1979; Holtz et al., 2004). Notably fewer meth-

ods are available for biostratigraphic analysis. These

methods, reviewed by Makovicky (2008), fall into two

categories. Most are based on quantitative analysis of

either taxon occurrences or their stratigraphic ranges

(e.g., Alroy, 1994, 2000; Fortelius et al., 2006). Other

methods employ the phylogenetic relationships be-

tween taxa (Martinez, 1995). The cladistic biochro-

nologic analysis developed by Makovicky (2008) is a

further elaboration of the latter approach. It is of par-

ticular interest to us because it was applied to the

chronological ordering of the assemblages of Late

Cretaceous continental vertebrates from the Gobi De-

sert of Mongolia and northern China (Makovicky,

2008). Here we employed a different method, based on

the more traditional "stage of evolution" approach to

analyzing evolutionary transformations within selected

groups of vertebrates. This approach has issues that

were reviewed by Makovicky (2008), but it remains

the basic approach for intra- and intercontinental bio-

stratigraphic correlations. There are still significant

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地 层 学 杂 志

36 卷

obstacles to developing a more rigorous approach be-

cause many vertebrate assemblages, such as the Mid-

dle Asian examples discussed here, are documented

for the most part by fragmentary skeletal remains,

greatly limiting their utility for phylogenetic analysis

at lower taxonomic levels.

First, we review the principal groups of the Late

Cretaceous continental vertebrates of Asia and their

potential utility for biostratigraphic purposes. Taxa

selected as biostratigraphic markers should satisfy

four criteria: 1) wide geographic range, being at least

found in more than one locality; 2) at least one trans-

formation event should have occurred within the par-

ticular lineage during the Late Cretaceous; 3) repre-

sented by abundant material; 4) and identifiable even

from fragmentary remains, such as isolated chon-

drichthyan or dinosaurian teeth or parts of turtle shells.

The taxa included in the present study are individual

genera or lower-level clades. We review 12 assem-

blages of Late Cretaceous continental vertebrates in

this paper. The assemblages constitute the rows in the

data matrix, and the columns in the data matrix list the

biostratigraphic markers (Tab. 1). All multi-state bio-

stratigraphic markers were treated as ordered. This

dataset was subjected to parsimony analysis.

The following assemblages of Late Cretaceous

continental vertebrates were selected for this analysis

(Fig. 1).

1) Khodzhakul Formation, southwestern Kyzylkum

Desert, Uzbekistan. Principal localities: Khodz-hakul,

Sheikhdzheili, and Chelpyk.

Tab. 1 Data matrix of 12 Late Cretaceous vertebrate assem-

blages and 26 biostratigraphic markers used (for the par-

simony analysis. For description of biostratigraphic markers see

text)

0

1

2

12345678901234567890123456

outgroup

00000000000000000000000000

Khodzhakul

10101?01100000?0011000?00?

Bissekty

2121201111110?101110001111

Yalovach

22313011111101?0011000???1

Bostobe

2231301111110??0111000???1

Iren Dabasu

21010?11100?0?10111000????

Bayn Shire Lower

??000101100????00?1000????

Bayn Shire Upper

2?00010110010?00021000????

Djadokhta

??010?02100??1?220?0102222

Barun Goyot

??010202100????2?0?0102233

Nemegt

??010?02100?1122?221113???

Judithian

33010300002112201?20113200

Lancian

43010300002112211?21113200

2) Bissekty Formation, central Kyzylkum Desert,

Uzbekistan. Principal locality: Dzharakuduk.

3) Yalovach Formation, Fergana Depression, Taji-

kistan. Principal locality: Kansai.

4) Bostobe Formation, northeastern Aral Sea region,

Kazakhstan. Principal localities: Shakh Shakh, Buro-

inak, Akkurgan, Baibishe, and Egizkara.

5) Iren Dabasu Formation, Gobi Desert, Inner

Mongolia, China. Principal locality: Iren Nor.

6) Lower part of Bayn Shire Formation, Gobi De-

sert, Mongolia. Principal localities: Shine Us Khuduk,

Khara Khutul, and Amtgai.

7) Upper part of Bayn Shire Formation, Gobi Desert,

Mongolia. Principal localities: Bayn Shire, Bayshin

Tsav, and Burkhant.

8) Djadokhta Formation, Gobi Desert, Mongolia

and China. Principal localities: Bayn Dzak, Ukhaa

Tolgod, Baga Tarjach, Alag Teer, Boro Khovil,

Tugrugeen Shire, and Bayan Mandahu.

9) Barun Goyot Formation, Gobi Desert, Mongolia.

Principal localities: Khulsan, Khermeen Tsav (red

beds), Udan Sayr, and Shara Tsav.

10) Nemegt Formation, Gobi Desert, Mongolia.

Principal localities: Nemegt, Altan Ula I-IV, Bugiin

Tsav, Khermeen Tsav (white beds), Gurileen Tsav,

Nogon Tsav, Tsagan Khushu, and Naran Bulak.

11-12) Vertebrate assemblages of the Judithian

and Lancian land-mammal “ages” in North America

(Russell, 1975; Cifelli et al., 2004; Kielan-Jaworowska

et al., 2004).

The outgroup fauna is a hypothetical assemblage

with state '0' for all biostratigraphic markers.

Fig. 1 Approximate geographic locations of the assem-

blages of Late Cretaceous continental vertebrates

in Middle and Central Asia discussed in this paper

1. Khodzhakul; 2. Bissekty; 3. Yalovach; 4. Bostobe; 5. Iren

Dabasu; 6. Lower Bayn Shire; 7. Upper Bayn Shire; 8. Djadokhta;

9. Barun Goyot; 10. Nemegt

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

465

3 Biostratigraphic Markers

3.1 Chondrichthyans

Teeth of sharks and rays are among the most com-

mon elements in Late Cretaceous microvertebrate as-

semblages. Freshwater deposits typically yield few

chondrichthyan species (Estes, 1964; Beavan & Rus-

sell, 1999). Nessov (1997) presented a long list of

shark species for the fluvial Bissekty Formation of

Uzbekistan, but the majority of these identifications

were based on surface-collected specimens derived

from the overlying marine Aitym Formation in the

section at Dzharakuduk (Archibald et al., 1998). In

Middle Asia, two chondrichthyan lineages appear most

suitable for the correlation of the Late Cretaceous

continental deposits: the hybodont shark Hybodus and

the rhinobatoid ray Myledaphus (Mertinene & Nessov,

1985; Nessov & Mertinene, 1986; Nessov et al.,

1994).

BM 1) Hybodus: 0) absent [outgroup only]; 1) H.

hodzhakulensis [Khodzhakul]; 2) H. kansaiensis [Bis-

sekty, Yalovach, Bostobe, Iren Dabasu, Upper Bayn

Shire]; 3) H. montanensis [Judithian]; 4) extinct [Lan-

cian]; ?) microvertebrates not known or not sampled

[Lower Bayn Shire, Djadokhta, Barun Goyot, Nemegt].

H. kansaiensis is cited for the Iren Dabasu Forma-

tion based on Currie & Eberth (1993: 136). Shuvalov &

Trusova (1979: 85) listed “H. asiaticus” [nomen nudum]

for the upper part of the Bayn Shire Formation at Bay-

shin Tsav. According to L.S. Glickman (cited by Shu-

valov & Trusova, 1979: 87) this species is "quite

similar to some species of Hybodus known from the

Santonian of Kazakhstan". Currently the material of

Hybodus from the Bostobe Formation of Kazakhstan is

referred to H. kansaiensis (Nessov & Mertinene, 1986;

Nessov, 1988, 1997), so the record for the Upper Bayn

Shire assemblage is here assigned to that species. In

North America Hybodus apparently became extinct

prior to the Lancian (Estes, 1964; Bryant, 1989;

Becker et al., 2004).

BM 2) Myledaphus: 0) absent [outgroup, Khodz-

hakul]; 1) M. tritus (Bissekty, Iren Dabasu]; 2) M.

glickmani [Yalovach, Bostobe]; 3) M. bipartitus [Ju-

dithian, Lancian]; ?) microvertebrates not known or

not sampled [Lower and Upper Bayn Shire, Djadokhta,

Barun Goyot, Nemegt].

Myledaphus is the most common freshwater elas-

mobranch in the Campanian and Maastrichtian conti-

nental strata of North America, where it is represented

by the type species, M. bipartitus (Estes, 1964; Bryant,

1989; Beavan & Russell, 1999; Neuman & Brinkman,

2005). In Middle Asia Myledaphus first appeared in

the uppermost (upper Cenomanian or lower Turonian)

portion of the Khodzhakul Formation (Karachadalysai;

Nessov, 1997; AA, unpublished data). These teeth are

referable to M. tritus, which is also known from the

Turonian Bissekty Formation (Nessov & Udovichenko,

1986; Nessov, 1988, 1997). Currie & Eberth (1993:

136) cited a similar species for the Iren Dabasu For-

mation. In Middle Asia this species is replaced by M.

glickmani in the Santonian Yalovach and Bostobe

formations (Parapalaeobates glickmani of Nessov &

Udovichenko, 1986; generic attribution based on

Cappetta [1992]). The teeth of the three species of

Myledaphus show successive changes in the sculpture

on the occlusal surface and increase in size and may

represent a single evolutionary lineage.

3.2 Osteichthyans

Isolated osteichthyan bones are common in Late

Cretaceous fluvial deposits but are rarely identified

and reported in the literature (Estes, 1964; Nessov,

1985, 1997; Bryant, 1989; Nessov & Panteleeva, 1999;

Peng et al., 2001; Neuman & Brinkman, 2005). The

dominant groups (Lepisosteidae, Amiidae, and Aspi-

dorhynchidae) do not exhibit marked changes during

the entire Late Cretaceous and are represented by very

similar species in Middle Asia and North America. For

example, certain fish scales dubbed “Holostean B”

from the Turonian Bissekty Formation of Uzbekistan

and the Campanian Dinosaur Park Formation of Al-

berta are virtually indistinguishable (Neuman & Brink-

man, 2005; A A, unpublished data). The Cenomanian

Khodzhakul Formation has a larger proportion of

osteichthyans with crushing dentitions, including

pycnodontiforms and the semionotid Lepidotes (Nes-

sov & Golovneva, 1983; Nessov, 1985), which are not

known from the younger, Turonian and Santonian

formations of Middle Asia. However, it is unclear if

this difference is due to local extinctions of these taxa

or reflects greater marine influence during the deposi-

tion of the Khodzhakul Formation. The fish remains

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地 层 学 杂 志

36 卷

from the Gobi Desert have not been studied, although

they are known from the Bayn Shire, Iren Dabasu,

Barun Goyot, and Nemegt formations. The lack of

osteichthyan remains in the Djadokhta Formation,

which include extensive eolian deposits (Dashzeveg et

al., 2005), may reflect unfavorable environmental

conditions for these animals.

3.3 Amphibians

The composition of Late Cretaceous amphibian as-

semblages is quite different between North America

and Middle Asia. In North America, the enigmatic

Albanerpetidae are quite common throughout the Late

Cretaceous (Gardner & Böhme, 2008). In Asia this

group is known only from rare specimens from the

Cenomanian Khodzhakul Formation and possibly a

single specimen from the Turonian Bissekty Forma-

tion (Gardner & Averianov, 1998; Skutschas, 2007).

Salamanders from the Late Cretaceous of North

America comprise Amphiumidae, Batrachosauroididae,

Scapherpetidae and Sirenidae, whereas all Cretaceous

salamanders known to date from Middle Asia are

Cryptobranchidae or Cryptobranchoidea (Skutschas,

2009). In Central Asia, Late Cretaceous salamanders

are unknown; this is either a sampling artifact or an

indication of unfavorable environments. The Middle

Asian cryptobranchid Eoscapherpeton is potentially

important for both biostratigraphy and biogeography

(Nessov, 1997; Skutschas, personal communication).

BM 3) Eoscapherpeton: 0) absent [outgroup, Iren

Dabasu, Lower and Upper Bayn Shire, Djadokhta,

Barun Goyot, Nemegt, Judithian, Lancian]; 1) E.

gracilis [Khodzhakul]; 2) E. asiaticum [Bissekty]; 3)

E. superum [Yalovach, Bostobe].

Late Cretaceous frogs are known mostly from iso-

lated bones in North America and Middle Asia (Estes,

1964; Roček & Nessov, 1993; Sanchiz, 1998; Gardner,

2008; Roček et al., 2010), whereas more complete

anuran skeletal remains have been reported from the

Gobi Desert (Borsuk-Białynicka, 1978; Špinar &

Tatarinov, 1986; Gubin, 1999; Roček, 2008). The dis-

coglossid (or gobiatid) genus Gobiates, which is

known from the Bissekty and Barun Goyot formations

(Roček & Nessov, 1993; Roček, 2008), is potentially

important for interregional correlations, but additional

anuran records from the other formations are needed

for a more comprehensive biostratigraphic assessment

of this group.

3.4 Testudines

Turtles arguably are the most common group of

vertebrates in Late Cretaceous continental deposits.

Even isolated shell elements are usually diagnostic to

the generic level based on characteristic sculpture and

patterns of the keratinous scutes. Furthermore, during

the Late Cretaceous several phyletic lineages of turtles

from different clades underwent considerable evolu-

tionary change in Asia (Sukhanov, 2000). Together

these factors make turtles one of the most important

vertebrate groups for Late Cretaceous continental bio-

stratigraphy.

BM 4) Kizylkumemys: 0) present [outgroup,

Khodzhakul, Lower and Upper Bayn Shire]; 1) absent

[other assemblages].

Kizylkumemys is a pitted-shell turtle (Carettochelyidae)

first described from the Cenomanian Khodzhakul

Formation of the southwestern Kyzylkum Desert of

Uzbekistan (Nessov, 1977). It is also known from the

Cenomanian Dzharakuduk Formation of the central

Kyzylkum Desert (Nessov, 1997) and from the Early

Cretaceous Sao Khua and Khok Kruat formations of

Thailand (Tong et al., 2009). In Middle Asia, Caretto-

chelyidae became extinct during the early Turonian

(Nessov & Golovneva, 1983). In Mongolia Kizylkume-

mys is known only from the Bayn Shire Formation,

from both the lower (Shine Us Khuduk, Khara Khutul)

and upper (Bayshin Tsav) parts (Nessov, 1981; Suk-

hanov, 2000; Sukhanov et al., 2008).

BM 5) Shachemydinae: 0) absent [outgroup, Iren

Dabasu, Lower and Upper Bayn Shire, Djadokhta,

Barun Goyot, Nemegt, Judithian, Lancian]; 1) Fer-

ganemys [Khodzhakul]; 2) Shachemys ancestralis

[Bissekty]; 3) Shachemys baibolatica [Yalovach,

Bostobe].

Shachemydinae is a subfamily of Adocidae, which,

together with Nanhsiungchelyidae, constitutes the

clade Adocusia, which is a sister-group to the clade

comprising the soft-shelled turtles (Trionychia) (Sy-

romyatnikova & Danilov, 2009). Shachemydinae is

endemic to Asia, where it ranges back to the Early

Cretaceous (Lapparent de Broin, 2004). In the Late

Cretaceous this group is known exclusively from

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Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

467

Middle Asia, where it appears to be useful for regional

biostratigraphy (Danilov et al., 2007).

BM 6) Nanhsiungchelyidae: 0) absent [outgroup,

Bissekty, Yalovach, Bostobe]; 1) Hanbogdemys

[Lower and Upper Bayn Shire, Djadokhta]; Zangerlia

[Djadokhta, Barun Goyot]; Basilemys [Judithian, Lan-

cian]; ?) Nanhsiungchelyidae indet. [Khodzhakul, Iren

Dabasu, Nemegt].

The distributional pattern of Nanhsiungchelyidae

mirrors that of Shachemydinae. In Middle Asia

Nanhsiungchelyidae is known only from rather unin-

formative remains from the Cenomanian Khodzhakul

Formation (Danilov & Syromyatnikova, 2008). In

Central Asia and Japan, the group extends back to the

Early Cretaceous and was moderately diverse and

abundant during the Late Cretaceous (Sukhanov, 2000;

Danilov & Syromyatnikova, 2008; Sukhanov et al.,

2008). In North America Nanhsiungchelyidae was

represented only by Basilemys, a common element of

Late Cretaceous continental assemblages starting in

the Coniacian (Hutchison, 2000; Brinkman, 2003).

Makovicky (2008: Fig. 5F) identified Basilemys in the

Iren Dabasu fauna based on an earlier identification in

Currie & Eberth (1993). Currently, however, there are

no indisputable records of Basilemys from Asia, and

earlier identifications should be treated as indetermi-

nate Nanhsiungchelyidae until the material has been

properly evaluated (Sukhanov, 2000; Danilov & Sy-

romyatnikova, 2008).

BM 7) Khunnuchelys: 0) absent [outgroup, Khodz-

hakul, Lower and Upper Bayn Shire, Djadokhta, Ba-

run Goyot, Nemegt, Judithian, Lancian]; 1) present

[Bissekty, Yalovach, Bostobe, Iren Dabasu].

Khunnuchelys is a large soft-shelled turtle (Trio-

nychidae) first reported on the basis of cranial remains

from the Iren Dabasu and Bissekty formations

(Brinkman et al., 1994). Recently a skull of Khunnu-

chelys was described from the Bostobe Formation of

Uzbekistan (Vitek & Danilov, 2010). According to

Vitek & Danilov (2010) this genus is possibly also

represented by shell material in the Yalovach Forma-

tion of Tajikistan. A trionychid taxon of potential im-

portance for intercontinental correlation is Aspidere-

toides, which is known from the Campanian to Maas-

trichtian of North America and the Santonian to Cam-

panian of Middle Asia (Gardner et al., 1995; Vitek &

Danilov, 2010).

BM 8) Lindholmemydidae: 0) absent [outgroup,

Judithian, Lancian]; 1) Lindholmemys [Khodzhakul,

Bissekty, Yalovach, Bostobe, Iren Dabasu, Lower and

Upper Bayn Shire]; 2) Mongolemys [Djadokhta, Barun

Goyot, Nemegt].

Lindholmemydidae is a group of freshwater turtles

known exclusively from the Late Cretaceous and Pa-

leogene of Asia (Sukhanov, 2000). Its monophyly re-

mains to be demonstrated; it possibly represents a

paraphyletic grouping of stem taxa to the crown clade

Testudinoidea. Lindholmemys is known from Middle

Asia and from the Iren Dabasu and Bayn Shire forma-

tions of the Gobi Desert (Danilov & Sukhanov, 2001),

whereas Mongolemys occurs in the younger forma-

tions of the Gobi Desert. A lindholmemydid turtle

from the Khodzhakul Formation of Uzbekistan was

previously identified as Mongolemys (Nessov &

Krassovskaya, 1984) or Khodzhakulemys (Danilov,

1999). It has now been reassigned to Lindholmemys

(Danilov, personal communication). A published re-

cord of Mongolemys from the Lower Cretaceous of

Mongolia (Sukhanov, 2000) is probably derived from

the Upper Cretaceous, possibly from the Nemegt

Formation (Danilov, personal communication).

3.5 Squamates

Snakes apparently originated in Gondwana and are

currently known only from sparse remains from the

Late Cretaceous of Europe and North America (Fox,

1975; Gardner & Cifelli, 1999). There are no records

to date from the Mesozoic of Middle and Central Asia.

Amphisbaenians are predominantly Gondwanan in

distribution but have a modest fossil record in Europe

and North America (Kearney, 2003). Gao & Nessov

(1998) referred the enigmatic Hodzhakulia from the

Khodzhakul Formation of Uzbekistan to Amphisbaenia,

but this assignment was not accepted by later studies

(Alifanov, 2000; Kearney, 2003). Hodzhakulia could

be useful for interregional correlations because it is

also occurs in the Lower Cretaceous, possibly Albian,

of Mongolia (Alifanov, 2000). Isolated lizard bones

are often quite common in the Late Cretaceous fluvial

deposits from Middle Asia and North America (Gao &

Fox, 1996; Nessov, 1997). The Djadokhta and Barun

468

地 层 学 杂 志

36 卷

Goyot formations of the Gobi Desert have yielded a

remarkable diversity of lizards, the majority of which

are represented by articulated skulls and skeletons

(Alifanov, 2000; Gao & Norell, 2000). This diversity,

however, appears to have been overestimated, and

certain taxa are in urgent need of taxonomic revision.

By contrast, the fossil record for lizards from the flu-

vial strata of the Nemegt Formation is much less ex-

tensive (Alifanov, 2000), and only a single lizard bone

has been reported to date from the fluvial deposits of

the Iren Dabasu Formation (Currie & Eberth, 1993).

Lizards could be useful for the biostratigraphic corre-

lations among the Late Cretaceous localities from the

Gobi Desert (Gao & Norell, 2000; Makovicky, 2008),

but considerable endemism of the known assemblages

currently restricts their utility for interregional corre-

lation.

3.6 Choristoderes

The superficially crocodile-like choristoderes were

a common element of continental aquatic ecosystems

in Asia during the Early Cretaceous. They were espe-

cially diverse and successful in ecosystems lacking

crocodyliform archosaurs (Matsumoto & Evans, 2010).

Surprisingly, there is not a single record of choristo-

deres from the Late Cretaceous of Asia, whereas these

reptiles were quite common (if less diverse) in North

America during this time interval and survived the

end-Cretaceous extinction event (Gao & Fox, 1998).

In Asia, choristoderes briefly reappeared during the

Paleocene, possibly by dispersal from Europe (Averi-

anov, 2005).

3.7 Crocodyliforms

Isolated crocodyliform teeth and osteoderms are

fairly common in most Late Cretaceous fluvial depos-

its from Asia and North America, but these materials

are often insufficient for lower-level taxonomic iden-

tification. In Asia partial or complete crocodyliform

skulls and partial skeletons are known from the Gobi

Desert (Efimov, 1988; Osmólska et al., 1997; Storrs &

Efimov, 2000; Pol & Norell, 2004a, b; Pol et al., 2009),

but there are also some (mostly as yet unpublished)

cranial remains from sites in Middle Asia. The latter ma-

terial is currently under study by P.P. Skutschas and

A.S. Rezvyi, and we rely on their personal communica-

tions for its identification. The Djadokhta and (to a lesser

extent) Barun Goyot formations have yielded terrestrially

adapted "protosuchian" crocodyliforms (Gobiosuchus,

Zaraasuchus, Zosuchus). Unidentified "protosuchians"

are also known from the Bissekty Formation in Middle

Asia. Neosuchian crocodyliforms are represented by

Shamosuchus in both Central and Middle Asia and

Kansajsuchus in Middle Asia. Late Cretaceous eusu-

chians in Asia are represented to date only by rare

records of Tadzhikosuchus from the Bissekty, Ya-

lovach, and Bostobe formations. This archaic faunal

composition of Late Cretaceous crocodyliform assem-

blages of Asia, with the presence of "protosuchians"

and dominance of non-eusuchian neosuchians, is in

striking contrast to the composition of the Campanian

and Maastrichtian assemblages of North America,

which comprise only eusuchians (Wu, 2005). The fol-

lowing crocodyliform taxa seem to be useful for intra-

and interregional biostratigraphy:

BM 9) Shamosuchus: 0) absent [outgroup, Judithian,

Lancian]; 1) present [all other assemblages].

BM 10) Kansajsuchus; 0) absent [outgroup,

Khodzhakul, Iren Dabasu, Lower and Upper Bayn

Shire, Djadokhta, Barun Goyot, Nemegt, Judithian,

Lancian]; 1) present [Bissekty, Yalovach, Bostobe].

BM 11) Eusuchia: 0) absent [outgroup, Khodzhakul,

Iren Dabasu, Lower and Upper Bayn Shire, Djadokhta,

Barun Goyot, Nemegt]; 1) Tadzhikosuchus [Bissekty,

Yalovach, Bostobe]; 2) Leidyosuchus [Judithian, Lan-

cian].

3.8 Pterosaurs

The diversity of pterodactyloid pterosaurs reached

its peak during the Early Cretaceous and steadily de-

clined during the Late Cretaceous (Butler et al., 2009).

There are three principal groups of Late Cretaceous

pterodactyloids: Pteranodontidae and Nyctosauridae,

restricted to the Western Hemisphere, and Azhdarchi-

dae, almost worldwide in distribution. Remarkably, all

of these last surviving clades of pterosaurs are tooth-

less. But teeth of Ornithocheiridae are still common in

the majority of nearshore marine environments during

the Cenomanian (Barrett et al., 2008). Their presence

in microvertebrate assemblages is usually well docu-

mented by isolated teeth (Wellnhofer & Buffetaut,

1999; Averianov, 2007a; Vullo & Neraudeau, 2009).

BM 12) Pterosauria: 1) Ornithocheiridae + Azhdarchidae

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

469

[outgroup, Khodzhakul]; 2) Azhdarchidae only [Bis-

sekty, Yalovach, Bostobe, Upper Bayn Shire, Judithian,

Lancian]; ?) no pterosaur records [Iren Dabasu, Lower

Bayn Shire, Djadokhta, Barun Goyot, Nemegt].

Pterosaurian bones are known from the Iren Dabasu

Formation but have not yet been identified (Currie &

Eberth, 1993: 136). Azhdarchid remains have recently

been recovered from the Upper Bayn Shire Formation

(Burkhant & Bayshin Tsav; Watabe et al., 2009), and

thus their absence in the Lower Bayn Shire is possibly

a sampling artifact. But the absence of these and any

other pterosaurs in the semi-arid depositional envi-

ronments of the Djadokhta and Barun Goyot formation

is puzzling. Perhaps these settings were not favorable

to pterosaurs. By Nemegt times pterosaurs may have

finally vanished in Asia, as there is no known Maas-

trichtian record of this group for that continent (Bar-

rett et al., 2008; note that these authors mistakenly

placed localities from European Russia in Asia).

3.9 Dinosaurs

An abundance of excellently preserved dinosaurian

specimens has been recovered from Late Cretaceous

sites in North America and Central Asia, whereas

dinosaurian material from Middle Asia typically com-

prises only incomplete, dissociated bones. However,

isolated teeth of a variety of dinosaurs are common at

most of the Late Cretaceous microvertebrate sites and

can be used for biostratigraphic correlations. Some

distinctive dinosaurian dental morphotypes appear to

have a wide stratigraphic and geographic range. For

example, the teeth of enigmatic theropod Richar-

doestesia isosceles from the Campanian and Maas-

trichtian of North America (Sankey et al., 2002;

Sankey, 2008) are virtually identical to those of

"Asiamericana" asiatica from the Turonian of Middle

Asia (Nessov, 1995; Sues & Averianov, unpublished

data). Among the isolated theropod teeth, those of

tyrannosauroids and troodontids are most useful for

correlation purposes.

BM 13) Tyrannosauroidea: 0) basal tyrannosauroids

with labiolingually compressed teeth [outgroup,

Khodzhakul, Bissekty, Yalovach, Bostobe, Iren Da-

basu, Upper Bayn Shire]; 1) more derived tyranno-

saurids with labiolingually thick (incrassate) teeth

[Nemegt, Judithian, Lancian]; ?) unknown [Lower

Bayn Shire, Djadokhta, Barun Goyot].

The biostratigraphic marker is based on character 201

listed by Brusatte et al. (2010). The derived state was

independently acquired by two tyrannosaurid clades of

Campanian to Maastrichtian age: Albertosaurinae and

(Daspletosaurus + (Tarbosaurus + Tyrannosaurus)).

The absence of tyrannosauroids in the Lower Bayn

Shire is possibly a sampling artifact. Large theropods

are known from the Djadokhta Formation at Ukhaa

Tolgod (Dingus et al., 2008), but, at the present time,

it is unknown if they are tyrannosauroids and what

kind of teeth they have.

BM 14) Troodontidae: 0) dental morphotype similar

to Sinornithoides [outgroup, Khodzhakul]; 1) dental

morphotype similar to Saurornithoides [Yalovach,

Djadokhta, Nemegt]; 2) dental morphotype similar to

Troodon [Judithian, Lancian]; ?) unknown [Bissekty,

Bostobe, Iren Dabasu, Lower and Upper Bayn Shire,

Barun Goyot].

The absence of troodontid teeth in the lower and

upper parts of the Bayn Shire Formation and the Barun

Goyot Formation of the Gobi Desert, which have not

been systematically prospected for microvertebrate

remains, possibly represents a sampling artifact. But

their absence in the microvertebrate samples from the

Bostobe Formation of Kazakhstan (Averianov, 2007b)

is puzzling. Troodontids with serrated teeth are not

known from the Cenomanian and Turonian of the cen-

tral Kyzylkum Desert where Urbacodon, a taxon with

unserrated teeth, is present (Averianov & Sues, 2007).

According to Currie & Eberth (1993: 136), in the

Iren Dabasu Formation "troodontid bones are rare, but

include distinctive third metatarsals […] in which the

distal articulation extends onto the posterior surface of

the bone in a broad tongue. A femur (PIN 2549/100,

Kurzanov, 1987) collected by the Sino-Soviet expedi-

tion is probably from a troodontid. These bones are

provisionally referred to Saurornithoides…" Subse-

quently, the presence of Saurornithoides sp. has been

cited for the Iren Dabasu assemblage (e.g., Weisham-

pel et al., 2004). However, Currie & Dong (2001:

1763-1764) later noted that the metatarsal III of the

Iren Dabasu troodontid is more similar to that of

Troodon and other troodontids rather than Saurorni-

thoides. "The Iren Dabasu troodontid, therefore, can-

470

地 层 学 杂 志

36 卷

not be identified further without additional mate-

rial…" (Currie & Dong, 2001: 1764). A femur from

Iren Nor figured by Kurzanov (1987, Fig. 36) is

probably troodontid but cannot be definitely referred

to Saurornithoides. Thus, Saurornithoides should be

removed from the faunal list of the Iren Dabasu For-

mation, and the material from this unit should be listed

as Troodontidae indet.

BM 15) Ornithomimosauria: 0) non-arctometatarsalian

[outgroup, Upper Bayn Shire]; 1) arctometatarsalian, with

ginglymoid distal condyle of metacarpal I [Bissekty, Iren

Dabasu]; 2) arctometatarsalian, with ball-like distal

condyle of metacarpal I [Nemegt, Judithian, Lancian]; ?)

unknown [Khodzhakul, Yalovach, Bostobe, Lower Bayn

Shire, Djadokhta, Barun Goyot].

The biostratigraphic marker is based on apomorphic

characters established by Kobayashi & Lü (2003) and

Kobayashi & Barsbold (2005). Ornithomimosaurs

from the Khodzhakul, Yalovach, Bostobe, Djadokhta,

and Barun Goyot formations are known from incom-

plete specimens that do not permit more precise iden-

tification (Ksepka & Norell, 2004; Alifanov & Averi-

anov, 2006; Averianov, 2007b). The first state charac-

terizes Garudimimus and more basal ornithomimo-

saurs, the second Archaeornithomimus, Sinorni-

thomimus, and the ornithomimosaur from the Bissekty

Formation, and the third characterizes taxa from the

Nemegt Formation (Anserimimus, Gallimimus) and

North America (Ornithomimus, Struthiomimus).

Currie & Eberth (1993: 137) noted that "at least

some of Archaeornithomimus specimens [from Iren

Dabasu Formation] are actually Garudimimus", which

is known from the Upper Bayn Shire Formation

(Bayshin Tsav & Barsbold, 1981; Kobayashi & Bars-

bold, 2005). Identification of Garudimimus in the Iren

Dabasu fauna was based on the putative presence of

the first pedal digit in a single arctometatarsalian

metatarsus. The presence of a first pedal digit was

inferred from a concave facet on the medial side of the

second metatarsal (Smith & Galton, 1990). However, as

was shown by Kobayashi & Barsbold (2005), the pes of

Garudimimus, unlike that of Archaeornithomimus, is not

arctometatarsalian. Furthermore, the contact surface

for the first metatarsal in Garudimimus is located on

the posterior rather than the medial surface of the

second metatarsal (Kobayashi & Barsbold, 2005).

Consequently, Garudimimus should be removed from

the faunal list for the Iren Dabasu Formation.

BM 16) Alvarezsauridae: 0) absent [outgroup,

Khodzhakul, Bissekty, Yalovach, Bostobe, Lower and

Upper Bayn Shire, Judithian]; 1) basal Alvarezsauridae

[Lancian]; 2) Parvicursorinae [Djadokhta, Barun

Goyot, Nemegt].

We have adopted the phylogeny of Alvarezsauridae

by Longrich & Currie (2009), but we follow Xu et al.

(2010a) in using Parvicursorinae for the clade com-

prising the Mongolian alvarezsaurids. Chiappe et al.

(2002: fig. 4.26A, B) referred an isolated fibula from

Iren Dabasu to Alvarezsauridae, but this bone more

likely belongs to Avimimus (Longrich & Currie 2009:

240). In China parvicursorines are known from strata

older than the Djadokhta Formation (Xu et al., 2010a).

BM 17) Oviraptorosauria: 0) absent [outgroup,

Khodzhakul, Yalovach, Lower and Upper Bayn Shire];

1) Oviraptorosauria stem to Oviraptoridae [Bissekty,

Bostobe, Iren Dabasu, Barun Goyot, Nemegt, Judi-

thian, Lancian]; 2) Oviraptoridae (Oviraptorinae +

Ingeniinae) [Djadokhta, Barun Goyot, Nemegt].

The phylogeny of Oviraptorosauria is based on

Longrich et al. (2010). Stem Oviraptorosauria includes

Avimimus, Gigantoraptor, and Caenagnathidae (in-

cluding Elmisauridae).

BM 18) Therizinosauroidea: absent [outgroup,

Djadokhta, Barun Goyot]; Therizinosauroidea basal to

Therizinosauridae [Khodzhakul, Bissekty, Yalovach,

Bostobe, Iren Dabasu, Lower Bayn Shire]; 2) Theriz-

inosauridae [Lower and Upper Bayn Shire, Nemegt]; ?)

unknown [Judithian, Lancian].

We have adopted the phylogeny of Therizinosauroidea

by Zanno (2010). The semiarid environments of the

Djadokhta and Barun Goyot may not have been fa-

vorable for therizinosauroids. The status of alleged

records of therizinosaurs from the Campanian and

Maastrichtian of North America is uncertain (Zanno,

2010).

BM 19) Hadrosauroidea: 0) absent [outgroup]; 1)

basal Hadrosauroidea [Khodzhakul, Bissekty, Ya-

lovach, Bostobe, Iren Dabasu, Upper and Lower Bayn

Shire]; 2) Hadrosauridae [Nemegt, Judithian, Lan-

cian]; ?) unknown [Djadokhta, Barun Goyot].

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

471

The phylogeny of Hadrosauroidea is based on the

analysis by Sues & Averianov (2009b), with Aralosaurus

placed outside Hadrosauridae. Hadrosauroid remains

from the Djadokhta and Barun Goyot formations are

rare, fragmentary, and currently indeterminate. The

single exception to date is an occurrence of several

juvenile hadrosauroid skeletons at Tugrugyin Shireh

(Barsbold & Perle, 1983). Perhaps hadrosauroids used

these semi-arid environments for nesting, but pre-

ferred feeding in more humid, more richly vegetated

settings. Hadrosauroid remains are common in the

fluvial strata of the Bayn Shire, Bissekty, Iren Dabasu,

and Nemegt formations.

BM 20) Saurolophus: 0) absent [outgroup, Khodz-

hakul, Bissekty, Yalovach, Bostobe, Iren Dabasu,

Lower and Upper Bayn Shire, Djadokhta, Barun

Goyot, Judithian]; 1) present [Nemegt, Lancian].

The crested hadrosaurine Saurolophus, from the

Nemegt Formation of Mongolia and from unit 4 of the

Horseshoe Canyon Formation of Alberta, Canada, is

the only known Late Cretaceous dinosaurian genus

definitely shared between Asia and western North

America (Bell, 2011, in press). It provides important

evidence for dating the Nemegt Formation as Maas-

trichtian.

BM 21) Pachycephalosauria: 0) absent [outgroup,

Khodzhakul, Bissekty, Yalovach, Bostobe, Iren Dabasu,

Lower and Upper Bayn Shire]; 1) present [Djadokhta,

Barun Goyot, Nemegt, Judithian, Lancian].

In Mongolia, Pachycephalosauria is represented by

rare but rather complete skeletal remains (Maryańska

et al., 2004). In North America pachycephalosaurs are

mostly known by isolated frontoparietals as well as

teeth from the fluvial strata of Judithian and Lancian age

(Baszio, 1997). Not a single tooth of this morphotype has

been identified to date in any of the extensive microver-

tebrate assemblages from the Late Cretaceous of Middle

Asia. Possibly pachycephalosaurs were not present in

Asia prior to the Campanian.

3.10 Aves

The fossil record of Late Cretaceous birds is gener-

ally sparse but a number of rather complete specimens

have been recovered from the Gobi Desert (El-

żanowski, 1977; Kurochkin, 1996, 2000; Chiappe et

al., 2001; Clarke & Norell, 2002). In Middle Asia,

Enantiornithiformes and taxa of uncertain affinities are

known from isolated bones or bone fragments (Ku-

rochkin, 2000; Chiappe & Walker, 2002). In Mongolia,

both enantiornithiform and ornithuran birds were pre-

sent, whereas ornithurans predominate in the Judithian

and Lancian of North America (Kurochkin, 2000; Hope,

2002; Clarke & Norell, 2004; Longrich, 2009). The

majority of the Late Cretaceous taxa of Enantiornithi-

formes and Ornithurae from Asia are endemic and thus

not useful for interregional correlations. However, one

group, Hesperornithiformes, is potentially important for

biostratigraphy. In North America this group ranges from

the Cenomanian to the Maastrichtian (Tokaryk et al.,

1997; Everhart & Bell, 2009; Wilson et al., 2011). In

Europe Hesperornithiformes are present during the

Campanian (Nessov & Yarkov, 1993; Panteleyev et al.,

2004; Rees & Lindgren, 2005). In Asia all known re-

cords of Hesperornithiformes appear to be Maas-

trichtian in age (Nessov & Borkin, 1983; Nessov &

Prizemlin, 1991; Kurochkin, 2000; Dyke et al., 2006).

BM 22) Hesperornithiformes: 0) absent [outgroup,

Khodzhakul, Bissekty, Yalovach, Bostobe, Iren Da-

basu, Lower and Upper Bayn Shire, Djadokhta, Barun

Goyot]; 1) present [Nemegt, Judithian, Lancian].

3.11 Mammalia

The rapid evolution of many mammalian groups,

the abundance of their fossil remains, and easy identi-

fication of taxa from dental remains were the main

reasons for the development of the land mammal

"ages" or biozones on most continents (e.g., Lindsay,

2003). Three principal land mammal ages have been

proposed for the Late Cretaceous of North America:

Aquilan (late Santonian to early Campanian), Judi-

thian (middle Campanian), and Lancian (late Maas-

trichtian) (Russell, 1975; Cifelli et al., 2004; Kie-

lan-Jaworowska et al., 2004). The mammalian taxa

from these ages are mostly endemic and unknown

outside North America, except for the eutherian

Paranyctoides, which is also known from the Bissekty

and Aitym formations (Turonian and ?Coniacian) in

Middle Asia (Archibald & Averianov, 2001; Averianov

& Archibald, 2003). The mammalian assemblages from

Mongolia are for the most part endemic, but allow cor-

relation among the various localities in the Gobi Desert

(Kielan-Jaworowska, 1974; Kielan-Jaworowska et al.,

472

地 层 学 杂 志

36 卷

2003). One Mongolian multituberculate, Bulganbaatar,

is known also from the early Campanian nearshore

strata with marine fossils in Kazakhstan (Darbasa

Formation; Averianov, 1997), which would indicate a

Campanian date for the Djadokhta Formation (which

is supported by magnetostratigraphic data; Dashzeveg

et al., 2005).

BM 23) Cimolodonta: 0) absent [outgroup only]; 1)

Uzbekbaatar

[Bissekty]; 2) Djadokhtatheroidea

[Djadokhta, Barun Goyot]; Cimolomyidae [Nemegt, Ju-

dithian, Lancian]; ?) unknown [Khodzhakul, Yalovach,

Bostobe, Iren Dabasu, Lower and Upper Bayn Shire].

Multituberculates were the most diverse and abun-

dant group of mammals in the Late Cretaceous of

North America and Central Asia. In the latter region

this group is represented by numerous well-preserved

skulls and skeletons (Kielan-Jaworowska et al., 2004).

On the contrary, multituberculates are absent or very

rare in the eutherian-dominated Late Cretaceous

mammalian assemblages of Middle Asia. In the exten-

sively sampled Bissekty fauna they represent about

1% of all mammalian fossils recovered to date

(Archibald & Averianov, 2005; Averianov & Archibald,

2006). In the latter multituberculates are represented

only by the archaic cimolodontan Uzbekbaatar, the

phylogenetic position of which is unresolved (Kie-

lan-Jaworowska et al., 2004; Averianov & Archibald,

2006). The relationships of Buginbaatar, the only

multituberculate taxon from the Nemegt Formation,

are also uncertain. It has provisionally been referred to

Cimolomyidae (Kielan-Jaworowska et al., 2004), and

is more similar to certain North American taxa than to

any other multituberculate taxon from the Gobi Desert.

BM 24) Deltatheridiidae: 0) absent [outgroup,

Khodzhakul]; 1) Sulestes [Bissekty]; 2) Deltatheridium

or Deltatheroides [Djadokhta, Barun Goyot, Judithian,

Lance]; ?) unknown [Yalovach, Bostobe, Iren Dabasu,

Lower and Upper Bayn Shire, Nemegt].

The metatherian clade Deltatheridiidae is known in

North America from the Early Cretaceous and from a

few records from the Judithian and Lancian faunas

(Fox, 1974; Davis et al., 2003; Kielan-Jaworowska et

al., 2004; Rougier et al., 2004). The latter records

possibly represent dispersals from Asia. In Asia this

group is represented by well-preserved specimens

from Mongolia and more fragmentary material from

Uzbekistan and Kazakhstan (Averianov, 1997; Rougier

et al., 1998, 2004; Averianov et al., 2010). The absence

of this group from other formations in Middle and Cen-

tral Asia possibly represents sampling artifacts.

BM 25) Asioryctitheria: 0) absent [outgroup,

Khodzhakul, Judithian, Lancian]; 1) Daulestes [Bis-

sekty]; 2) Kennalestes [Djadokhta]; 3) Asioryctes

[Barun Goyot]; ?) unknown [Yalovach, Bostobe, Iren

Dabasu, Lower and Upper Bayn Shire, Nemegt].

The endemic Asian eutherian clade Asioryctitheria

is well known from the Bissekty Formation of Middle

Asia and the Djadokhta and Barun Goyot formations

of Central Asia (Archibald & Averianov, 2006). This

group of rapidly evolving mammals has potential for

biostratigraphic correlation between Late Cretaceous

continental vertebrate localities in Asia. Additional

discoveries of asioryctitherians from the fluvial forma-

tions of the Gobi Desert may be helpful in constraining

the stratigraphic positions of these units.

BM 26) Zalambdalestidae: 0) absent [outgroup, Ju-

dithian, Lancian]; 1) Kulbeckia [Bissekty, Yalovach,

Bostobe]; 2) Zalambdalestes [Djadokhta]; 3) Ba-

runlestes [Barun Goyot]; ?) unknown [Khodzhakul,

Iren Dabasu, Lower and Upper Bayn Shire, Nemegt].

This is another clade of Late Cretaceous eutherians

endemic to Asia and also has potential for biostrati-

graphic correlation. In the Khodzhakul Formation

Zalambdalestidae is represented only by a single iso-

lated petrosal to date (Averianov & Archibald, 2005).

Kulbeckia is best known from the Bissekty Formation

of Uzbekistan, but some teeth have also been recov-

ered from the Yalovach Formation of Tajikistan

(Archibald & Averianov, 2003). Beleutinus, a poorly

known zalambdalestid taxon from the Bostobe Forma-

tion of Kazakhstan, is more comparable in its "stage of

evolution" to Kulbeckia than to any of the Mongolian

zalambdalestids (AA, personal observation). Alym-

lestes from the Campanian of Kazakhstan has a de-

rived structure of the lower molars similar to that of

the Mongolian zalambdalestids (Averianov & Nessov,

1995).

4 Parsimony analysis

The data matrix comprises 13 vertebrate assem-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

473

blages and 26 biostratigraphic markers (Table 1) and

was analyzed using three parsimony programs. Im-

plementation of the branch and bound search algo-

rithm of PAUP, version 4.0b10 (Swofford, 2002),

produced five trees, each with a length of 65 steps, a

consistency index of 0.80, and a retention index of

0.83. TNT (Goloboff et al., 2003; new technology

search algorithm) and NONA version 2.0 (Goloboff,

1999; 1,000 repetitions of the parsimony ratchet algo-

rithm) run with the Winclada version 1.00.08 interface

(Nixon, 1999) both produced two trees, each with a

length of 59 steps, a consistency index of 0.88, and

retention index of 0.87. The latter result, requiring

fewer transformations, is accepted here. Both most

parsimonious trees are illustrated in Fig. 2. Only un-

ambiguously optimized apomorphic biostratigraphic

markers are shown in Fig. 2 and discussed here.

The two most parsimonious trees differ in the to-

pology of segment A (Fig. 2), with the North American

faunas either basal (tree 1) or apical (tree 2) to the

assemblage from the Nemegt Formation.

The most basal of the vertebrate assemblages dis-

cussed in this paper is Khodzhakul. Its marker taxa are

Eoscapherpeton gracilis (3[1]) and Ferganemys (5[1]).

The more apical assemblages are united by the ab-

sence of ornithocheirid pterosaurs (12[1]) and pres-

ence of Hybodus kansaiensis (1[2]). The cluster next

branching off unites the two Bayn Shire assemblages

based on the shared presence of Hanbogdemys (6[1]).

The more apical assemblages are united by the ab-

sence of Kizylkumemys (4[1]) and the presence of

Oviraptorosauria stem to Oviraptoridae (17[1]). These

assemblages are divided into two groups, a Middle

Asian one plus Iren Dabasu and a variable segment A.

The first group is supported by the shared presence of

Khunnuchelys (7[1]). The Middle Asiatic assemblages

share the presence of the crocodyliforms Kansajsuchus

(10[1]) and Tadzhikosuchus (11[1]). The Yalovach and

Bostobe assemblages share the presence of

Myledaphus glickmani (2[2]).

The Gobi and North American assemblages are

united either by the presence of tyrannosaurids with

halosauria and Mongolemys (8[2]) (tree 2). On Tree 1

the North American complexes share the absence of

Shamosuchus 9([0]) and the presence of Leidyosuchus

(11[2]) and Troodon (14[2]). The presence of Parvi-

cursorinae (16[2]) unites the Gobi faunas, and thein-

crassate teeth (13[1]), Hadrosauridae (19[2]), and

Pachycephalosauria (21[1]) (tree 1) or by Pachycep-

presence of Djadokhtatheroidea (23[2]) unites the

Fig. 2 Two most parsimonious trees produced by the parsimony ratchet algorithm of NONA version 2.0 using the data

matrix present in Tab. 1

The trees differ in the topology of segment A. Only unambiguous biostratigraphic markers are shown (black circles represent nonhomoplasies and

white circles represent homoplasies). The numbers at the circles are biostratigraphic markers (above) and states (below)

474

地 层 学 杂 志

36 卷

Pachycephalosauria (21[1]) (tree 1) or by Pachycep-

presence of Djadokhtatheroidea (23[2]) unites the

Djadokhta and Barun Goyot assemblages. On Tree 2

the Nemegt and North American faunas are grouped

together by the presence of Hesperornithiformes

(22[1]), and the North American faunas share the ab-

sence of Lindholmemys (8[0]) and Shamosuchus (9[0])

and the presence of Leidyosuchus (11[2]) and Troodon

(14[2]).

5 Succession of the Late Cretaceous ver-

tebrate assemblages in Middle and Cen-

tral Asia

5.1 Middle Asia

In the Kyzylkum Desert there are two principal Late

Cretaceous vertebrate-bearing formations: the early

Cenomanian Khodzhakul Formation in the southwest-

ern Kyzylkum and the middle to late Turonian Bis-

sekty Formation in the central Kyzylkum. Archibald &

Averianov (2005) named the vertebrate assemblages

from these stratigraphic units the Sheikhdzheili and

Bissekty local faunas, respectively. However, the first

fauna is named after the most important locality and

the second name is derived from a stratigraphic unit.

For the sake of consistency, we here use only names

for vertebrate local faunas that are derived from their

respective stratigraphic units rather than the principal

localities. Thus, we now refer to the Sheikhdzheili

local fauna of Archibald & Averianov (2005) as the

Khodzhakul local fauna.

The turtle assemblage of the Khodzhakul Formation

is characterized by the presence of Ferganemys and

Kizylkumemys, which are absent from younger faunal

assemblages in Middle Asia. Ferganemys was origi-

nally described from the Lower Cretaceous of Kyr-

gyzstan (Nessov & Khozatsky, 1977). Syromyatnikova

(2011) has recently questioned the referral of the

Kyzylkum species, F. itemirensis, to the same genus.

Ferganemys itemirensis was first described from the

Cenomanian of the central Kyzylkum Desert (Nessov,

1981). Kizylkumemys and Ferganemys appear to be

useful taxa for correlating pre-Turonian vertebrate

assemblages in Middle Asia.

The dinosaurian assemblages from the Khodzhakul

and Bissekty formations are remarkably similar at the

family level. This is not surprising in view of their

geographic proximity and similar ages (both faunas

are separated by an interval of only five to eight mil-

lion years). The most conspicuous difference between

the two faunas is the presence of the basal neocera-

topsian Asiaceratops in the Khodzhakul assemblage,

where it was one of most common herbivorous dino-

saurs. In the Bissekty local fauna basal neoceratop-

sians are apparently represented by a rare, distinct

taxon. In this fauna there is also a second ceratopsian,

the stem ceratopsid Turanoceratops (Sues & Averi-

anov, 2009a). Turanoceratops is not common in the

Bissekty local fauna and unknown from the Khodz-

hakul local fauna to date. There are some minor mor-

phological differences between the ornithomimids,

therizinosaurids, and hadrosauroids from these faunas.

However, it is difficult to evaluate these differences

because of the fragmentary nature of the available

Khodzhakul dinosaurian remains.

Only eutherians are known from the Khodzhakul

mammalian assemblage: the stem eutherian Bobolestes,

plesiomorphic zhelestids, and a possible zalambdalestid

(Averianov & Archibald, 2005). Zhelestids are repre-

sented by taxa less derived than those from the Bis-

sekty local fauna.

A vertebrate assemblage from the Cenomanian

Dzharakuduk Formation at Itemir in the Itemir-Dzh-

arakuduk Depression of the central Kyzylkum closely

resembles the Khodzhakul local fauna in its composi-

tion. It includes five taxa of dinosaurs: Tyrannosaur-

oidea indet., Urbacodon itemirensis (Troodontidae),

Neosauropoda indet., Hadrosauroidea indet., and Asi-

aceratops salsopaludalis (Neoceratopsia). Skeletal re-

mains of Asiaceratops are common at this locality and

indistinguishable from those from sites in Karakal-

pakistan. The latter have yielded a troodontid with

unserrated teeth, whereas the Khodzhakul fauna has a

troodontid with serrated teeth. A similar taxon with

unserrated teeth, Urbacodon sp., is present in the

younger Bissekty local fauna of the same region (Ave-

rianov & Sues, 2007). Currently it is unclear if this

similarity is due to the close geographic proximity of

the two localities or the younger geological age of the

Itemir local fauna relative to the Khodzhakul local

fauna. The presence of Ferganemys and Kizylkumemys

as well as ornithocheirid pterosaurs in the Dzhara-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

475

kuduk Formation support its Cenomanian age.

The only Cenomanian vertebrate assemblages

known in Middle Asia outside the Kyzylkum Desert

are those from the Sharikhan and other formations in

the Fergana Depression of Kyrgyzstan. These units

have yielded only incomplete dinosaurian remains,

including large Theropoda indet. (Tyrannosauroidea?),

Ornithomimosauria indet., Sauropoda indet., and Hadro-

sauroidea indet. (Nessov, 1995; Averianov, 2006).

In the Itemir-Dzharakuduk Depression the Ceno-

manian and Turonian dinosaur-bearing fluvial deposits

are separated by a thick marine section deposited dur-

ing the early Turonian transgression (Uchkuduk and

Dzheirantau formations; Pyatkov et al., 1967; King,

personal communication). Similarly, the fluvial depos-

its of the Cenomanian Khodzhakul Formation in

Karakalpakistan are overlain by the early Turonian

marine strata of the Beshtyube Formation (Schultz,

1972; King, personal communication). During this

early Turonian transgression a significant faunal turn-

over took place among marine vertebrates and fresh-

water turtles (Nessov & Golovneva, 1983; Nessov,

1997). However, terrestrial vertebrates, including di-

nosaurs, seem to have been little affected by this

turnover. A notable exception is the disappearance of

ornithocheirid pterosaurs in the region after the

Cenomanian (Averianov, 2007a), which may have

been related to changes in the fish fauna.

The vertebrates from the Aitym Formation at

Dzharakuduk (Aitym local fauna) come from near-

shore marine deposits with numerous remains of a

diversity of chondrichthyan and other fishes. This

stratigraphic unit marks a transgression following the

deposition of the fluvial Bissekty Formation, possibly

during the latest Turonian or Coniacian (King, per-

sonal communication). The few dinosaurian teeth re-

covered to date from the Aitym Formation are indis-

tinguishable from those from the underlying Bissekty

Formation. The mammalian fauna is also very similar,

except for the presence of a slightly more derived spe-

cies of the multituberculate Uzbekbaatar, which also

appears to be more common here (Averianov &

Archibald, 2003).

The marine transgression continued in the Ky-

zylkum region during the Santonian and Campanian

and reached its peak in the Maastrichtian (Pyatkov et

al., 1967; King, personal communication). This ex-

plains the rarity of dinosaur occurrences in the

Kyzylkum Desert for this time interval.

The Turonian Zhirkindek Formation in the Aral Sea

region of Kazakhstan is coeval with the Bissekty

Formation of Uzbekistan. Its poorly known dinosaur-

ian assemblage includes Tyrannosauroidea indet., Or-

nithomimosauria indet., Therizinosauroidea indet.,

Dromaeosauridae indet., Sauropoda indet., Hadro-

sauroidea indet., and Neoceratopsia indet. (Nessov,

1995; Kordikova et al., 2001; Averianov, 2007b; Ave-

rianov & Sues, 2009). The hadrosauroid bones from

the Zhirkindek Formation are very similar to those of

Levnesovia from the Bissekty local fauna (Sues &

Averianov, 2009b). The only difference between the

two faunas is the presence of a large basal neoceratop-

sian in the Zhirkindek fauna (Averianov & Sues, 2009).

Other vertebrates from the Zhirkindek Formation in-

clude fishes, turtles, lizards, and crocodyliforms

(Kordikova et al., 2001) but these materials have not

yet been studied in detail.

Younger, Santonian vertebrate faunas are known in

Middle Asia from regions unaffected by the

post-Turonian marine transgression. These are the Ya-

lovach local fauna in the Fergana Depression of Taji-

kistan and the Bostobe local fauna in the Aral Sea re-

gion of Kazakhstan. All common groups of dinosaurs

in the Bissekty local fauna are also present in the

Santonian faunas. The Bostobe hadrosauroid

Aralosaurus is distinctly more derived than the Bis-

sekty hadrosauroid Levnesovia (Sues & Averianov,

2009b), which is consistent with a younger, Santonian

or possibly even early Campanian age for the Bostobe

Formation. Troodontids are currently unknown from

the Bostobe local fauna, but are represented in the

Yalovach local fauna by a taxon with serrated teeth

(Averianov & Sues, 2007), unlike Urbacodon with

unserrated teeth from the Bissekty local fauna. Rich

microvertebrate samples have been recovered from

both the Yalovach and Bostobe formations (Nessov,

1997; Kordikova et al., 2001), but these materials have

not yet been fully documented.

The Turonian-Santonian vertebrate faunas of Mid-

dle Asia are set apart biogeographically from the Cen-

476

地 层 学 杂 志

36 卷

tral Asian Late Cretaceous faunas by the presence of

the endemic crocodyliforms Kansajsuchus and Tadz-

hikosuchus. Eusuchians are unknown in the Late Cre-

taceous of Central Asia, but are present in North

America (Leidyosuchus). The crocodyliform Shamo-

suchus is common in the Late Cretaceous faunas of

Middle and Central Asia, but is unknown in North

America. The other distinct taxon for the Tu-

ronian-Santonian vertebrate assemblages of Middle

Asia is the trionychid turtle Khunnuchelys. In Central

Asia it is known only from the Iren Dabasu fauna

(Brinkman et al., 1994).

Campanian vertebrates in Middle Asia are known

from a microvertebrate sample obtained from near-

shore marine strata of the Darbasa Formation in

southern Kazakhstan. Only the mammal and troodon-

tid teeth from this sample have been described to date

(Averianov & Nessov, 1995; Averianov, 1997; Averi-

anov & Sues, 2007). This fauna is notable for the

presence of the multituberculate mammal Bulgan-

baatar, which was originally reported from the

Djadokhta Formation of Mongolia. The zalambdales-

tid mammal Alymlestes from the Darbasa Formation is

similar in its “stage of evolution” to the Campanian

zalambdalestids from Mongolia and clearly more de-

rived than Kulbeckia from the Turonian Bissekty

fauna.

Only marine vertebrates, mostly sharks and mosa-

saurs, are currently known from the Maastrichtian of

Middle Asia (Nessov, 1997).

5.2 Central Asia

Nessov's pioneering work on the Cretaceous verte-

brates from the Kyzylkum Desert suggested that the

Bissekty fauna most closely resembles that from the

Iren Dabasu Formation of Inner Mongolia, China

(Nessov, 1995, 1997). This correlation led to Nessov

to refer certain dinosaurs from the Bissekty local fauna

to genera previously reported from the Iren Dabasu

fauna: Alectrosaurus sp., Archaeornithomimus (?)

bissektensis, Gilmoreosaurus arkhangelskyi, and Bac-

trosaurus kysylkumensis (Nessov, 1995). The practice

of attribution of fragmentary materials to taxa known

from more complete specimens in a different locality

leads to circular reasoning when the relative age of

localities is considered. Based on our own work we

have reidentified these Bissekty dinosaurs as Tyran-

nosauroidea indet., Ornithomimosauria indet., and

Levnesovia transoxiana or Hadrosauroidea indet.

Nevertheless, the Iren Dabasu local fauna appears to

be quite similar to the Bissekty local fauna in compo-

sition and evolutionary level of the taxa concerned.

The hadrosauroids Levnesovia and Bactrosaurus are

certainly closely related (Sues & Averianov, 2009b).

The principal difference between the dinosaurs from

the two faunas is the presence of different groups of

oviraptorosaurs. In the Bissekty there is the small

caenagnathid Caenagnathasia, whereas the Iren Da-

basu local fauna includes an avimimid and the large

oviraptorosaur Gigantoraptor. Caenagnathidae are

currently known only from the Late Cretaceous of

North America and Middle Asia, but Elmisaurus and

Nomingia from the Nemegt Formation of Gobi Desert

are probably additional representatives of this clade

in Central Asia (Osmólska et al., 2004). This faunal

difference may reflect the different environmental and

paleogeographic positions of the two faunas (coastal

plains of Middle Asia versus inland areas of Central

Asia) rather than a difference in stratigraphic age. The

similarities between the Iren Dabasu and Nemegt bio-

tas in the charophytes and freshwater ostracodes (Van

Itterbeeck et al., 2005) and the shared presence of

Avimimus (or Avimimidae) probably reflect fluvial

depositional settings and mesic climatic conditions

(Jerzykiewicz & Russell, 1991) rather than similarity

in age. This is especially evident when these assem-

blages are compared with those from the Djadokhta

Formation, which represent semi-arid settings

(Jerzykiewicz & Russell, 1991). The presence in the

Iren Dabasu fauna of the turtle Khunnuchelys, which is

otherwise known only from the Turonian to Santonian

of Middle Asia (Brinkman et al., 2004; Vitek &

Danilov, 2010), also suggests a pre-Campanian age for

this assemblage.

The poorly known Shireegiin Gashuun local fauna

of the Gobi Desert of Mongolia with the small basal

neoceratopsian Graciliceratops is similar in its com-

position to the Cenomanian Khodzhakul and Dzhara-

kuduk faunas of Middle Asia, which are dominated by

the small basal neoceratopsian Asiaceratops. However,

the presence of the turtle Lindholmemys in this forma-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

477

tion (Danilov & Sukhanov, 2001) suggests a younger,

possibly Turonian to Santonian age for this unit.

The Bayn Shire Formation in the eastern Gobi De-

sert of Mongolia has been traditionally correlated with

the Iren Dabasu Formation of Inner Mongolia (Currie

& Eberth, 1993; Nessov, 1995, 1997; Hicks et al.,

1999; Averianov, 2002). Based on the stratigraphic

ranges of the turtles, this unit can be divided into a

lower, Cenomanian to early Turonian part with

Kizylkumemys and an upper, late Turonian to Santo-

nian part with Lindholmemys (Nessov, 1997; Sukha-

nov et al., 2008). Based on magnetostratigraphic data,

Hicks et al. (1999) argued that the Bayn Shireh For-

mation was most likely deposited no later than the

latest Santonian. Sukhanov (2000) also reported

Kizylkumemys from the upper part of the Bayn Shire

Formation. Lindholmemys is known also from the Iren

Dabasu Formation but is absent from the younger,

Campanian to Maastrichtian faunas of the Gobi Desert,

where it is replaced by Mongolemys (Currie & Eberth,

1993; Danilov & Sukhanov, 2001). There are no dino-

saurian genera known to be shared between the lower

and upper parts of the Bayn Shire Formation.

Based on the occurrences of turtles and dinosaurs,

Nessov (1997) correlated the lower and upper parts of

the Bayn Shire Formation with the Khodzhakul and

Bissekty formations in Middle Asia, respectively. Al-

though these stratigraphic units share no dinosaurian

genera, this correlation is plausible given the similar

evolutionary levels of the dinosaurs, notably the ther-

izinosauroids and ankylosaurids. The Bissekty theriz-

inosauroid closely resembles Erlikosaurus from the

upper Bayn Shire fauna. The postcranial elements of

Bissektipelta from the Bissekty Formation are almost

identical to those of Talarurus from the upper Bayn

Shire and differ from those of more derived, strati-

graphically younger ankylosaurs.

The remarkably diverse faunas known from the

Djadokhta and Barun Goyot formations document the

next (Campanian) stage of vertebrate evolution in the

Gobi Desert. The dinosaurs from these units differ

considerably from those of the Bissekty fauna in 1) the

rarity of tyrannosauroids, ornithomimids, and hadro-

sauroids; 2) presence of diverse alvarezsaurids and

oviraptorids, which are not known in Middle Asia; 3)

presence of pachycephalosaurs; and 4) abundance of

basal neoceratopsians ("protoceratopsids"), which are

very rare in the Bissekty fauna. The highly diverse

lizard faunas of the Djadokhta and Barun Goyot for-

mations do not have direct parallels in other regions. A

dentary fragment of a lizard from the Bostobe Forma-

tion of Kazakhstan, identified as Slavoia by Kordik-

ova et al. (2001), is best referred to as Scincomorpha

indet. Multituberculate mammals are represented by

an endemic clade Djadokhtatheroidea, which is un-

known in other regions except for Bulganbaatar from

Kazakhstan (Averianov, 1997). The therian mammals

from the Gobi assemblages show more similarities

with the faunas from Middle Asia. Deltatheridiidae,

Asio- ryctitheria, and Zalambdalestidae are present in

both regions, but the Mongolian taxa are clearly more

derived than those from the Turonian of Uzbekistan

(Archibald & Averianov, 2006; Averianov et al.,

2010). However, the most remarkable difference be-

tween the Late Cretaceous mammalian faunas of

Middle and Central Asia is the dominance of Zheles-

tidae in the former and their complete absence in the

latter.

In the Maastrichtian the faunas of the Djadokhta

and Barun Goyot formations were replaced by the

fauna of the fluvial strata of the Nemegt Formation.

The Nemegt dinosaurian assemblage differs markedly

from those of the Djadokhta and Barun Goyot forma-

tions in the abundance of ornithomimids (Gallimimus),

dominance of tyrannosaurids (Tarbosaurus) and

hadrosaurs (Saurolophus), and the absence of "proto-

ceratopsids". In this respect the Nemegt fauna resem-

bles the Bissekty fauna, but this similarity, for the

most part, probably reflects similar floodplain deposi-

tional environments rather than geological age. The

tyrannosaurids, ornithomimids, therizinosaurids, and

hadrosaurids of the Nemegt fauna all are distinctly

more derived than related taxa from the Bissekty fauna.

The Nemegt dinosaurian fauna is also ecologically

similar to the assemblage from the Iren Dabasu For-

mation, which sometimes has led to the mistaken im-

pression that these two faunas are comparable in age.

The stratigraphic correlation of the principal Late

Cretaceous vertebrate faunas of Middle and Central

Asia proposed in this paper is illustrated in Fig. 3.

478

地 层 学 杂 志

36 卷

Fig. 3 Attempted stratigraphic correlation of the assemblages of Late Cretaceous continental vertebrates

in Middle and Central Asia discussed in this paper

6 Discussion

Jerzykiewicz & Russell (1991) proposed a succession of

seven Cretaceous Mongolian Land Vertebrate "Ages"

("MOLVAs"), including four for the Late Cretaceous:

Saynshandian, Baynshirenian, Barungoyotian, and Ne-

megtian. All these "ages" were modified versions of the

biostratigraphic horizons used by Soviet geologists (see

Barsbold, 1983). This explains appearance of the earliest

Late Cretaceous Saynshandian "MOLVA" for which no

actual vertebrate fossils were known (Jerzykiewicz &

Russell, 1991) but which is characterized by a distinct

assemblage of freshwater mollusks (Martinson, 1982). In

Inner Mongolia (China), the Iren Dabasu and Bayan

Mandachu formations could be easily referred to the

Baynshirenian and Barungoyotian MOLVAs, respec-

tively (Lucas & Estep, 1998). Less certain is the refer-

ence of the vertebrate assemblage from the Wangshi Se-

ries of Shandong (China) to the Barungoyotian MOLVA

(Lucas & Estep, 2008). The only shared taxon between

Gobi and Shandong dinosaurian assemblages is the an-

kylosaurid Pinacosaurus (Buffetaut, 1995). However,

the lack of cranial material for the Shandong anky-

losaurid renders this identification questionable. In

contrast to the Barungoyotian faunas of Gobi, the Shan-

dong dinosaurian assemblage is dominated by hadro-

sauroids, including the basal hadrosauroid Tanius, the

giant saurolophine Shantungosaurus, and the lam-

beosaurine Tsintaosaurus (Prieto-Márquez, 2010). Cera-

topsia is represented by Sinoceratops, identified as the

most basal centrosaurine ceratopsid, and the leptocera-

topsid Zhuchengceratops (Xu et al., 2010b, c). There is

also a tyrannosaurid Zhuchengtyrannus, which closely

resembles the Nemegtian Tarbosaurus (Hone et al.,

2011). Currently there are no non-dinosaurian vertebrates

recorded from the Wangshi Series, and thus this verte-

brate assemblage was not included into our analysis.

The Wangshi Series is possibly Campanian in age, and

its dinosaurian assemblage resembles contemporane-

ous dinosaurian faunas from North America more

closely than the Gobi faunas (Prieto-Márquez, 2010;

Xu et al., 2010b, c). Lucas & Estep (1998) also re-

ferred to the Nemegtian MOLVA several poorly known

dinosaur assemblages in Xinjiang and Guandong

provinces of China.

The concept "Land Vertebrate Ages" is potentially

useful for correlation of the assemblages of Late Cre-

taceous continental vertebrates in Asia, but "Mongo-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

479

lian Land Vertebrates Ages" ("MOLVAs") should be

transformed to "Asian Land Vertebrate Ages" ("AL-

VAs") and formally defined. The paucity of data con-

cerning non-dinosaurian vertebrates in many East

Asian localities makes such a zonation scheme pre-

mature at present.

Acknowledgments We thank Igor Danilov, Chris King,

Pavel Skutschas, and Elena Syromyatnikova for information

and advice. We gratefully acknowledge financial support of

the National Science Foundation (EAR-9804771 and

EAR-0207004 to J.D. Archibald and H.-D. Sues), the Na-

tional Geographic Society (#5901-97 and #6281-98 to J.D.

Archibald and H.-D. Sues), the Civilian Research and De-

velopment Foundation (RU-G1-2571-ST-04 and RUB1-

2860-ST-07), and the Russian Fund of Basic Research

(07-04- 91110-AFGIRa). Averianov also received research

support from the President's of Russia grant MD 255.

2003.04 and the Russian Fund of Basic Research grants

04-04-49113, 04-04-49637, 07-04-00393, and 10-04-01350,

by the Ministry of Education and Science of Russian Fed-

eration (contract 16.518. 11.7070), and from a Paleon-

tological Museum grant of Saint Petersburg State University.

We thank Benjamin Sames and Spencer Lucas for construc-

tive reviews of the manuscript.

References

Alifanov V R. 2000. The fossil record of Cretaceous lizards

from Mongolia. In: Benton M J, Shishkin M A, Unwin D M

& Kurochkin E N eds. The age of dinosaurs in Russia and

Mongolia. Cambridge: Cambridge University Press. 368-389

Alifanov V R & Averianov A O. 2006. On the finding of orni-

thomimid dinosaurs (Saurischia, Ornithomimosauria) in the

Upper Cretaceous beds of Tajikistan. Paleontological Jour-

nal, 40(1): 103-108

Alroy J. 1994. Appearance event ordination: a new biochro-

nologic method. Paleobiology, 20(2): 191-207

Alroy J. 2000. New methods for quantifying macroevolutionary

patterns and processes. Paleobiology, 26(4): 707-733

Archibald J D & Averianov A O. 2001. Paranyctoides and allies

from the Late Cretaceous of North America and Asia. Acta

Palaeontologica Polonica, 46(4): 533-551

Archibald J D & Averianov A O. 2003. The Late Cretaceous

placental mammal Kulbeckia. Journal of Vertebrate Paleon-

tology, 23(2): 404-419

Archibald J D & Averianov A O. 2005. Mammalian faunal suc-

cession in the Cretaceous of the Kyzylkum Desert. Journal

of Mammalian Evolution, 12(1-2): 9-22

Archibald J D & Averianov A O. 2006. Late Cretaceous

asioryctitherian eutherian mammals from Uzbekistan and

phylogenetic analysis of Asioryctitheria. Acta Palaeon-

tologica Polonica, 51(2): 351-376

Archibald J D, Sues H-D, Averianov A O, King C, Ward D J,

Tsaruk O I, Danilov I G, Rezvyi A S, Veretennikov B G &

Khodjaev A. 1998. Précis of the Cretaceous paleontology,

biostratigraphy and sedimentology at Dzharakuduk (Tu-

ronian?-Santonian), Kyzylkum Desert, Uzbekistan. Bulletin

of the New Mexico Museum of Natural History and Science,

14: 21-28

Averianov A O. 1997. New Late Cretaceous mammals of

southern Kazakhstan. Acta Palaeontologica Polonica, 42(2):

243-256

Averianov A O. 2002. An ankylosaurid (Ornithischia: Anky-

losauria) braincase from the Upper Cretaceous Bissekty

Formation of Uzbekistan. Bulletin de l'Institut Royal des

Sciences Naturelles de Belgique, Sciences de la Terre, 72(1):

97-110

Averianov A O. 2005. The first choristoderes (Diapsida, Cho-

ristodera) from the Paleogene of Asia. Paleontological

Journal, 39(1): 79-84

Averianov A O. 2006. On an ornithomimid dinosaur (Saurischia,

Ornithomimosauria) from the Cenomanian of Fergan. Pale-

ontologicheskii Zhurnal, (3): 88-92

Averianov A O. 2007a. Mid-Cretaceous ornithocheirids (Pter-

osauria, Ornithocheiridae) from Russia and Uzbekistan. Pa-

leontological Journal, 41(1): 79-86

Averianov A O. 2007b. Theropod dinosaurs from the Late Cre-

taceous of North-East Aral Sea area, Kazakhstan. Cretaceous

Research, 28(3): 532-544

Averianov A O & Archibald J D. 2003. Mammals from the Up-

per Cretaceous Aitym Formation, Kyzylkum Desert, Uz-

bekistan. Cretaceous Research, 24(2): 171-191

Averianov A O & Archibald J D. 2005. Mammals from the

mid-Cretaceous Khodzhakul Formation, Kyzylkum Desert,

Uzbekistan. Cretaceous Research, 26(4): 593-608

Averianov A O & Archibald J D. 2006. New specimens of the

multituberculate mammal Uzbekbaatar from the Late Cre-

taceous of Uzbekistan. Acta Palaeontologica Polonica, 51(2):

377-380

Averianov A O, Archibald J D & Ekdale E G. 2010. New mate-

rial of the Late Cretaceous deltatheroidan mammal Sulestes

from Uzbekistan and phylogenetic reassessment of the

metatherian-eutherian dichotomy. Journal of Systematic Pa-

laeontology, 8(3): 301-330

Averianov A O & Nessov L A. 1995. A new Cretaceous mam-

mal from the Campanian of Kazakhstan. Neues Jahrbuch für

Geologie und Paläontologie, Monatshefte (1): 65-74

Averianov A O & Sues H-D. 2007. A new troodontid (Dino-

sauria: Theropoda) from the Cenomanian of Uzbekistan,

with a review of troodontid records from the territories of

the former Soviet Union. Journal of Vertebrate Paleontology,

27(1): 87-98

Averianov A O & Sues H-D. 2009. First record of a basal neo-

ceratopsian dinosaur from the Late Cretaceous of Kazakh-

stan. Acta Palaeontologica Polonica, 54(3): 553-556

480

地 层 学 杂 志

36 卷

Barrett P M, Butler R J, Edwards N P & Milner A R. 2008.

Pterosaur distribution in time and space: an atlas. Zitteliana,

B28: 61-107

Barsbold R. 1981. Toothless carnivorous dinosaurs of Mongolia.

Trudy Sovmestnoi Sovetsko-Mongol'skoi Paleontologicheskoi

Ekspeditsii, 15: 28-39

Barsbold R. 1983. Carnivorous dinosaurs from the Cretaceous

of Mongolia. Trudy Sovmestnoi Sovetsko-Mongol'skoi Pa-

leontologicheskoi Ekspeditsii, 19: 1-120

Barsbold R & Perle A. 1983. On the taphonomy of joint burial of

juvenile dinosaurs and some aspects of their ecology. Trudy

Sovmestnoi Sovetsko-Mongol'skoi Paleontologicheskoi

Ekspeditsii, 24: 121-125

Baszio S. 1997. Systematic palaeontology of isolated dinosaur

teeth from the latest Cretaceous of south Alberta, Canada.

Courier Forschungsinstitut Senckenberg, 196: 33-77

Beavan N R & Russell A P. 1999. An elasmobranch assemblage

from the terrestrial-marine transitional Lethbridge Coal

Zone (Dinosaur Park Formation: upper Campanian), Alberta,

Canada. Journal of Paleontology, 73(3): 494-503

Becker M A, Chamberlain J A & Terry D O Jr. 2004. Chon-

drichthyans from the Fairpoint Member of the Fox Hills

Formation (Maastrichtian), Meade County, South Dakota.

Journal of Vertebrate Paleontology, 24(4): 780-793

Bell P R. 2011. Redescription of the skull of Saurolophus os-

borni Brown 1912 (Ornithischia: Hadrosauridae). Creta-

ceous Research, 32(1): 30-44

Bell P R (in press). Cranial osteology and ontogeny of

Saurolophus angustirostris from the Late Cretaceous of

Mongolia with comments on Saurolophus osborni from

Canada. Acta Palaeontologica Polonica, 56

Borsuk-Białynicka M. 1978. Eopelobates leptocolaptus sp.n. :

the first Upper Cretaceous pelobatid frog from Asia. Palae-

ontologia Polonica, 38: 57-63

Brinkman D B. 2003. A review of nonmarine turtles from the

Late Cretaceous of Alberta. Canadian Journal of Earth Sci-

ences, 40(4): 557-571

Brinkman D B, Nessov L A & Peng J-H. 1994. Khunnuchelys

gen. nov., a new trionychid (Testudines: Trionychidae) from

the Late Cretaceous of Inner Mongolia and Uzbekistan. Ca-

nadian Journal of Earth Sciences, 30(10-11): 2214-2223

Brusatte S L, Norell M A, Carr T D, Erickson G M, Hutchinson

J R, Balanoff A M, Bever G S, Choiniere J N, Makovicky P J

& Xu X. 2010. Tyrannosaur paleobiology: new research on

ancient exemplar organisms. Science, 329(5998): 1481-1485

Bryant L J. 1989. Non-dinosaurian lower vertebrates across the

Cretaceous-Tertiary boundary in northeastern Montana.

University of California Publications in Geological Sciences,

134: 1-107

Buffetaut E. 1995. An ankylosaurid dinosaur from the Upper

Cretaceous of Shandong (China). Geological Magazine,

132(6): 683-692

Butler R J, Barrett P M, Nowbath S & Upchurch P. 2009. Esti-

mating the effects of sampling biases on pterosaur diversity

patterns: implications for hypotheses of bird/pterosaur

competitive replacement. Paleobiology, 35(3): 432-446

Cappetta H. 1992. Nouveaux Rhinobatoidei (Neoselachii,

Rajiformes) à denture specialisée du Maastrichtien du Maroc.

Remarques sur l'évolution dentaire des Rajiformes et des

Myliobatiformes. Neues Jahrbuch für Geologie und

Paläontologie, Abhandlungen, 187(1): 31-52

Cheetham A H & Hazel J E. 1969. Binary (presence-absence)

similarity coefficients. Journal of Paleontology, 43(5):

1130-1136

Chiappe L M, Norell M A & Clark J M. 2001. A new skull of

Gobipteryx minuta (Aves: Enantiornithes) from the Creta-

ceous of the Gobi Desert. American Museum Novitates,

(3346): 1-15

Chiappe L M, Norell M A & Clark J M. 2002. The Cretaceous,

short-armed Alvarezsauridae: Mononykus and its kin. In:

Chiappe L M & Witmer L M eds. Mesozoic birds: above the

heads of dinosaurs. Berkeley: University of California Press.

87-120

Chiappe L M & Walker C A. 2002. Skeletal morphology and

systematics of the Cretaceous Euenantiornithes (Orni-

thothoraces: Enantiornithes). In: Chiappe L M & Witmer L

M eds. Mesozoic birds, above the heads of dinosaurs.

Berkeley: University of California Press. 240-267

Cifelli R L, Eberle J J, Lofgren D L, Lillegraven J A & Clemens

W A. 2004. Mammalian biochronology of the latest Creta-

ceous. In: Woodburne M O ed. Late Cretaceous and Ceno-

zoic Mammals of North America: biostratigraphy and geo-

chronology. New York: Columbia University Press. 21-42

Clarke J A & Norell M A. 2002. The morphology and phyloge-

netic position of Apsaravis ukhaana from the Late Cretaceous

of Mongolia. American Museum Novitates, (3387): 1-46

Clarke J A & Norell M A. 2004. New avialan remains and a

review of the known avifauna from the Late Cretaceous

Nemegt Formation of Mongolia. American Museum Novi-

tates, (3447): 1-12

Currie P J & Dong Z M. 2001. New information on Cretaceous

troodontids (Dinosauria, Theropoda) from the People's Re-

public of China. Canadian Journal of Earth Sciences, 38(12):

1753-1766

Currie P J & Eberth D A. 1993. Palaeontology, sedimentology

and palaeoecology of the Iren Dabasu Formation (Upper

Cretaceous), Inner Mongolia, People's Republic of China.

Cretaceous Research, 14(2): 127-144

Dani A H & Masson V M. 1992. History of civilizations of

Central Asia. Volume 1. Paris: UNESCO

Danilov I G. 1999. A new lindholmemydid genus (Testudines:

Lindholmemydidae) from the mid-Cretaceous of Uzbekistan.

Russian Journal of Herpetology, 6(2): 63-71

Danilov I G & Sukhanov V B. 2001. New data on lindholme-

myid turtle Lindholmemys from the Late Cretaceous of

Mongolia. Acta Palaeontologica Polonica, 46(1): 125-131

Danilov I G & Syromyatnikova E V. 2008. New materials on

turtles of the family Nanhsiungchelyidae from the Creta-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

481

ceous of Uzbekistan and Mongolia, with a review of the

nanhsiungchelyid record in Asia. Proceedings of the Zoo-

logical Institute of the Russian Academy of Sciences,

312(1/2): 3-25

Danilov I G., Syromyatnikova E V & Sukhanov V B. 2007.

Turtles of the genus Shachemys from the Upper Cretaceous

of Asia. In: Rozanov A Y, Lopatin A V & Parkhaev P Y eds.

Modern paleontology: classical and new methods. Moscow:

Paleontological Institute of the Russian Academy of Sci-

ences. 59-72

Dashzeveg D, Dingus L, Loope D B, Swisher C C III, Dulam T

& Sweeney M R. 2005. New stratigraphic subdivision, de-

positional environment, and age estimate for the Upper Cre-

taceous Djadokhta Formation, southern Ulan Nur Basin,

Mongolia. American Museum Novitates, (3498): 1-31

Davis B M, Cifelli R L & Kielan-Jaworowska Z. 2008. Earliest

evidence of Deltatheroida (Mammalia: Metatheria) from the

Early Cretaceous of North America. In: Sargis E J & Da-

gosto M eds. Mammalian evolutionary morphology. A Trib-

ute to Frederick S. Szalay. Berlin: Springer. 3-24

Dingus L, Loope D B, Dashzeveg D, Swisher C C III, Minjin C,

Novacek M J & Norell M A. 2008. The geology of Ukhaa

Tolgod (Djadokhta Formation, Upper Cretaceous, Nemegt

Basin, Mongolia). American Museum Novitates, (3616):

1-40

Dyke G J, Malakhov D V & Chiappe L M. 2006. A re-analysis

of the marine bird Asiahesperornis from northern Kazakh-

stan. Cretaceous Research, 27(6): 947-953

Efimov M B. 1988. On the fossil crocodiles of Mongolia and

the USSR. Trudy Sovmestnoi Sovetsko-Mongol'skoi Pale-

ontologicheskoi Ekspeditsii, 34: 81-90

Elżanowski A. 1977. Skulls of Gobipteryx (Aves) from the

Upper Cretaceous of Mongolia. Palaeontologia Polonica, 37:

153-165

Estes R. 1964. Fossil vertebrates from the Late Cretaceous

Lance Formation, eastern Wyoming. University of Califor-

nia Publications in Geological Sciences, 49: 1-180

Everhart M J & Bell A. 2009. A hesperornithiform limb bone

from the basal Greenhorn Formation (Late Cretaceous; mid-

dle Cenomanian) of North Central Kansas. Journal of Verte-

brate Paleontology, 29(3): 952-956

Fortelius M, Gionis A, Jernvall J & Mannila H. 2006. Spectral

ordering and biochronology of European fossil mammals.

Paleobiology, 32(2): 206-214

Fox R C. 1974. Deltatheroides-like mammals from the Upper

Cretaceous of North America. Nature, 249(5455): 392

Fox R C. 1975. Fossil snakes from the upper Milk River For-

mation (Upper Cretaceous), Alberta. Canadian Journal of

Earth Sciences, 12(9): 1557-1563

Gao K. & Fox R C. 1996. Taxonomy and evolution of Late

Cretaceous lizards (Reptilia: Squamata) from western Can-

ada. Bulletin of Carnegie Museum of Natural History, 33:

1-107

Gao K & Fox R C. 1998. New choristoderes (Reptilia: Diapsida)

from the Upper Cretaceous and Palaeocene, Alberta and

Saskatchewan, Canada, and phylogenetic relationships of

Choristodera. Zoological Journal of the Linnean Society,

124(4): 303-353

Gao K & Nessov L A. 1998. Early Cretaceous squamates from

the Kyzylkum Desert, Uzbekistan. Neues Jahrbuch für

Geologie und Paläontologie, Abhandlungen 207: 289-309

Gao K & Norell M A. 2000. Taxonomic composition and sys-

tematics of Late Cretaceous lizard assemblages from Ukhaa

Tolgod and adjacent localities, Mongolian Gobi Desert.

Bulletin of the American Museum of Natural History, 249:

1-118

Gardner J D. 2008. New information on frogs (Lissamphibia:

Anura) from the Lance Formation (Late Maastrichtian) and

Bug Creek Anthills (late Maastrichtian and early Paleocene),

Hell Creek Formation, USA. In: Sankey J T & Baszio S eds.

Vertebrate microfossil assemblages: their role in paleoecol-

ogy and paleobiogeography. Bloomington: Indiana Univer-

sity Press. 219-249

Gardner J D & Averianov A O. 1998. Albanerpetontid am-

phibians from the Upper Cretaceous of Middle Asia. Acta

Palaeontologica Polonica, 43(3): 453-467

Gardner J D & Böhme M. 2008. Review of Albanerpetontidae

(Lissamphibia), with comments on the paleoecological pref-

erences of European Tertiary albanerpetontids. Vertebrate

Microfossil Assemblages. In: Sankey J T & Baszio S

eds.Their Role in Paleoecology and Paleobiogeography.

Bloomington: Indiana University Press. 178-218

Gardner J D & Cifelli R L. 1999. A primitive snake from the

Cretaceous of Utah. Special Papers in Palaeontology, 60:

87-100

Gardner J D, Russell A P & Brinkman D B. 1995. Systematics

and taxonomy of soft-shelled turtles (Family Trionychidae)

from the Judith River Group (mid-Campanian) of North

America. Canadian Journal of Earth Sciences, 32(5):

631-643

Goloboff P. 1999. NONA (ver. 1.9). Software published by the

author, San Miguel de Tucumán, Argentina. Available online

at www.cladistics.org

Goloboff P, Farris J S & Nixon K C. 2003. Tree analysis using

new technology. Program and documentation available from

the authors (and online at www.zmuc. dk/public/phylogeny)

Gubin Y M. 1999. Gobiatids (Anura) from the Upper Creta-

ceous locality Khermeen Tsav (Gobi Desert, Mongolia).

Paleontologicheskii Zhurnal, (1): 76-87

Hicks J F, Brinkman D L, Nichols D L & Watabe M. 1999.

Paleomagnetic and palynologic analyses of Albian to Santo-

nian strata at Bayn Shireh, Burkhant, and Khuren Dukh,

eastern Gobi Desert, Mongolia. Cretaceous Research, 20(6):

829-850

Holtz T R, Chapman R E Jr & Lamanna M C. 2004. Mesozoic

biogeography of dinosauria. In: Weishampel D B, Dodson P

& Osmólska H eds. The dinosauria (Second edition). Berke-

ley: University of California Press. 627-642

482

地 层 学 杂 志

36 卷

Hone D W E, Wang K, Sullivan C, Zhao X, Chen S, Li D, Ji S,

Ji Q, & Xu X. 2011. A new, large tyrannosaurine theropod

from the Upper Cretaceous of China. Cretaceous Research,

32(4): 495-503

Hope S. 2002. The Mesozoic radiation of Neornithes. In:

Chiappe L M & Witmer L M eds. Mesozoic birds, above the

heads of dinosaurs. Berkeley: University of California Press.

339-388

Hutchison J H. 2000. Diversity of Cretaceous turtle faunas of

eastern Asia and their contribution to the turtle faunas of

North America. Paleontological Society of Korea Special

Publication, 4: 27-38

Jerzykiewicz T & Russell D A. 1991. Late Mesozoic stratigra-

phy and vertebrates of the Gobi Basin. Cretaceous Research,

12(4): 345-377

Kielan-Jaworowska Z. 1974. Multituberculate succession in the

Late Cretaceous of the Gobi Desert (Mongolia). Palaeon-

tologia Polonica, 30: 23-44

Kielan-Jaworowska Z, Cifelli R L & Luo Z-X. 2004. Mammals

from the age of dinosaurs: origins, evolution, and structure.

New York:Columbia University Press. 1-630

Kielan-Jaworowska Z, Hurum J H & Badamgarav D. 2003. An

extended range of multituberculate Kryptobaatar and dis-

tribution of mammals in the Upper Cretaceous of the Gobi

Desert. Acta Palaeontologica Polonica, 48(2): 273-278

Kearney M. 2003. Systematics of the Amphisbaenia (Lepi-

dosauria: Squamata) based on morphological evidence from

recent and fossil forms. Herpetological Monographs, 17(1):

1-74

Kobayashi Y & Barsbold R. 2005. Reexamination of a primitive

ornithomimosaur, Garudimimus brevipes Barsbold, 1981

(Dinosauria: Theropoda), from the Late Cretaceous of

Mongolia. Canadian Journal of Earth Sciences, 42(9):

1501-1521

Kobayashi Y & Lü J-C. 2003. A new ornithomimid dinosaur

with gregarious habits from the Late Cretaceous of China.

Acta Palaeontologica Polonica, 48(2): 235-259

Kordikova E G, Polly D P, Alifanov V R, Roček Z, Gunnell G F

& Averianov A O. 2001. Small vertebrates from the Late

Cretaceous and Early Tertiary of the northeastern Aral Sea

Region, Kazakhstan. Journal of Paleontology, 75(2):

390-400

Ksepka D T & Norell M A. 2004. Ornithomimosaur cranial

material from Ukhaa Tolgod (Omnogov, Mongolia). Ameri-

can Museum Novitates, (3448): 1-4

Kurochkin E N. 1996. A new enantiornithid of the Mongolian

Late Cretaceous, and a general appraisal of the infraclass

Enantiornithes (Aves). Moscow, Paleontological Institute,

Special Issue. 60

Kurochkin E N. 2000. Mesozoic birds of Mongolia and the

former USSR. In: Benton M J, Shishkin M A, Unwin D M &

Kurochkin E N eds. The age of dinosaurs in Russia and

Mongolia. Cambridge: Cambridge University Press. 533-559

Kurzanov S M. 1987. Avimimidae and the problem of the origin

of birds. Trudy Sovmestnoi Sovetsko-Mongol'skoi Paleon-

tologicheskoi Ekspeditsii, 31: 1-92

Lapparent de Broin F de. 2004. A new Shachemydinae (Chelo-

nii, Cryptodira) from the Lower Cretaceous of Laos: pre-

liminary data. Comptes Rendus Palevol, 3(5): 387-396

Lindsay E H. 2003. Chronostratigraphy, biochronology, datum

events, land mammal ages, stage of evolution, and appear-

ance event ordination. Bulletin of the American Museum of

Natural History, 279: 212-230

Longrich N R. 2009. An ornithurine-dominated avifauna from

the Belly River Group (Campanian, Upper Cretaceous) of

Alberta, Canada. Cretaceous Research, 30(1): 161-177

Longrich N R & Currie P J. 2009. Albertonykus borealis, a new

alvarezsaur (Dinosauria: Theropoda) from the early Maas-

trichtian of Alberta, Canada: implications for the systematics

and ecology of the Alvarezsauridae. Cretaceous Research,

30(1): 239-252

Longrich N R, Currie P J & Dong Z-M. 2010. A new oviraptorid

(Dinosauria: Theropoda) from the Upper Cretaceous of Bayan

Mandahu, Inner Mongolia. Palaeontology, 53(5): 945-960

Lucas S G. & Estep J W. 1998. Vertebrate biostratigraphy and

biochronology of the Cretaceous of China. New Mexico

Museum of Natural History and Science Bulletin, 14: 1-20

Makovicky P J. 2008. Telling time from fossils: a phylog-

eny-based approach to chronological ordering of paleobiotas.

Cladistics, 24(3): 350-371

Martinez J N. 1995. Biochronology and parsimony methods.

Bulletin de la Société géologique de France, 166(5): 517-526

Martinson G G. 1982. Late Cretaceous mollusks of Mongolia.

Trudy Sovmestnoi Sovetsko-Mongol'skoi Paleontologicheskoi

Ekspeditsii, 17: 1-84

Maryańska T, Chapman R E & Weishampel D B. 2004.

Pachycephalosauria. In: Weishampel D B, Dodson P &

Osmólska H eds. The dinosauria (Second edition). Berkeley:

University of California Press. 464-477

Matsumoto R & Evans S E. 2010. Choristoderes and the fresh-

water assemblages of Laurasia. Journal of Iberian Geology,

36(2): 253-274

Mertinene R A & Nessov L A. 1985. Hybodont sharks from the

Cretaceous of Middle Asia. Doklady Akademii Nauk Tadz-

hikskoi SSR, 28(1): 588-592

Nessov L A. 1977. A new genus of two-clawed turtle from the

Upper Cretaceous of Karakalpakia. Paleontologicheskii

Zhurnal, (1): 103-114

Nessov L A. 1981. On a turtle of the family of the Dermatemy-

didae from the Cretaceous of the Amur River Basin, and

some other rare finds of ancient turtle remains of Asia. Her-

petological Investigations in Siberia and Far East. L Y

Borkin. Leningrad: Zoologicheskyi Institut AN SSSR. 69-73

Nessov L A. 1985. Rare bony fishes, terrestrial lizards and

mammals from the zone of estuaries and coastal plains of the

Cretaceous of Kizylkum. Ezhegodnik Vsesoyuznogo Pale-

ontologicheskogo Obshchestva, 28: 199-219

Nessov L A. 1988. Assemblages of late Mesozoic and Paleo-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

483

cene vertebrates of Middle Asia. In: Bogdanova T N &

Oshurkova M V eds. Formation and evolution of the conti-

nental biotas. Leningrad: Izdatelstvo Nauka. 93-101

Nessov L A. 1995. Dinosaurs of Northern Eurasia: new data

about assemblages, ecology and paleobiogeography. Saint

Petersburg: Izdatelstvo Sankt-Peterburgskogo Universiteta.

1-156

Nessov L A. 1997. Cretaceous nonmarine vertebrates of northern

Eurasia. In: Golovneva L B & Averianov A O eds. Saint Pe-

tersburg: Izdatel'stvo Sankt-Peterburgskogo Universiteta. 1-218

Nessov L A & Borkin L Y. 1983. New findings of bird bones

from the Cretaceous of Mongolia and Middle Asia. Trudy

Zoologicheskogo Instituta AN SSSR, 116: 108-110

Nessov L A & Golovneva L B. 1983. Changes of vertebrate

complexes of Cenomanian-Santonian (Late Cretaceous) of

Kyzylkum. In: Oleinikov A N ed. Paleontology and evolu-

tion of biosphere. Transactions of XXV Session of the

All-Union Paleontological Society. Leningrad: Nauka.

126-134

Nessov L A & Khozatsky L I. 1977. Freshwater turtle from the

Early Cretaceous of Fergana. Ezhegodnik Vsesoyuznogo Pale-

ontologicheskogo Obshchestva, 20: 248-262

Nessov L A & Krassovskaya T B. 1984. [Transformations in the

composition of turtle assemblages of the Late Cretaceous of

Middle Asia]. Vestnik Leningradskogo Universiteta (Seriya

Biologiya), 1(3): 15-25

Nessov L A & Mertinene R A. 1986. Remains of chondrich-

thyan fishes from the Cretaceous of Middle Asia and Ka-

zakhstan as a source of information on the age and origin of

the deposits. In: Krymgoltz G Y & Belenkova V S eds. Pa-

leontology and detailed stratigraphic correlation. Leningrad:

Nauka. 35-41

Nessov L A & Panteleeva T M. 1999. Gar Atractosteus tura-

nensis sp. nov. (Lepisosteidae) from the Upper Cretaceous of

Kyzylkum Desert. Trudy Zoologicheskogo Instituta RAN,

277: 104-118

Nessov L A & Prizemlin B V. 1991. A large advanced flightless

marine bird of the order Hesperornithiformes of the Late

Senonian of Turgai Strait: the first find of the group in the

USSR. Trudy Zoologicheskogo Instituta AN SSSR, 239:

85-107

Nessov L A, Sigogneau-Russell D & Russell D E. 1994. A sur-

vey of Cretaceous tribosphenic mammals from Middle Asia

(Uzbekistan, Kazakhstan and Tajikistan), of their geological

setting, age and faunal environment. Palaeovertebrata, 23:

51-92

Nessov L A & Udovichenko N I. 1986. [New findings of re-

mains of Cretaceous and Paleogene vertebrates of Middle

Asia]. Voprosy Paleontologii, 9: 129-136

Nessov L A & Yarkov A A. 1993. [Hesperornithiforms in Rus-

sia]. Russkii Ornitologicheskii Zhurnal, 2(1): 37-54

Neuman A G & Brinkman D B. 2005. Fishes of the fluvial beds.

In: Currie P J & Koppelhus E B eds. Dinosaur Provincial Park:

a spectacular ancient ecosystem revealed. Bloomington:

Indiana University Press. 167-185

Nixon K C. 1999. Winclada (Beta) version 0.9.9. Software

published by the author, Ithaca, NY. Available online at

www.cladistics.org

Osmólska H, Hua S & Buffetaut E. 1997. Gobiosuchus kielanae

(Protosuchia) from the Late Cretaceous of Mongolia: anat-

omy and relationships. Acta Palaeontologica Polonica, 42(2):

257-289

Osmólska H, Currie P J & Barsbold R. 2004. Oviraptorosauria.

In: Weishampel D B, Dodson P & Osmólska H eds. The di-

nosauria (Second edition). Berkeley: University of Califor-

nia Press. 165-183

Panteleyev A V, Popov E V & Averianov A O. 2004. New re-

cord of Hesperornis rossicus (Aves, Hesperornithiformes) in

the Campanian of Saratov Province, Russia. Paleontological

Research, 8(2): 115-122

Peng J H, Russell A P & Brinkman D B. 2001. Vertebrate mi-

crosite assemblages (exclusive of mammals) from the

Foremost and Oldman formations of the Judith River Group

(Campanian) of southeastern Alberta: an illustrated guide.

Provincial Museum of Alberta, Natural History Occasional

Paper, 25: 1-54

Pol D & Norell M A. 2004a. A new crocodyliform from Zos

Canyon, Mongolia. American Museum Novitates, (3445):

1-36

Pol D & Norell M A. 2004b. A new gobiosuchid crocodyliform

taxon from the Cretaceous of Mongolia. American Museum

Novitates, (3458): 1-31

Pol D, Turner A H & Norell M A. 2009. Morphology of the Late

Cretaceous crocodylomorph Shamosuchus djadochtaensis

and a discussion of neosuchian phylogeny as related to the

origin of Eusuchia. Bulletin of the American Museum of

Natural History, 324: 1-103

Prieto-Márquez A. 2010. Global historical biogeography of

hadrosaurid dinosaurs. Zoological Journal of the Linnean

Society, 159(2): 503-525

Pyatkov K K, Pyanovskaya I A, Bukharin A K & Bykovskii Y K.

1967. Geological structure of Central Kyzylkum. Tashkent:

Fan. 1-177

Raup D M & Crick R E. 1979. Measurement of faunal similar-

ity in paleontology. Journal of Paleontology, 53(5):

1213-1227

Rees J & Lindgren J. 2005. Aquatic birds from the Upper Cre-

taceous (lower Campanian) of Sweden and the biology and

distribution of hesperornithiforms. Palaeontology, 48(6):

1321-1329

Roček Z. 2008. The Late Cretaceous frog Gobiates from Cen-

tral Asia: its evolutionary status and possible phylogenetic

relationships. Cretaceous Research, 29(4): 577-591

Roček Z & Nessov L A. 1993. Cretaceous anurans from Central

Asia. Palaeontographica A, 226: 1-54

Roček Z, Eaton J G, Gardner J D & Přikryl T. 2010. Evolution

of anuran assemblages in the Late Cretaceous of Utah, USA.

Palaeobiodiversity and Palaeoenvironments, 90(4): 341-393

484

地 层 学 杂 志

36 卷

Rougier G W, Wible J R & Novacek M J. 1998. Implications of

Deltatheridium specimens for early marsupial history. Na-

ture, 396(6710): 459-463

Rougier G W, Wible J R & Novacek M J. 2004. New specimen

of Deltatheroides cretacicus (Metatheria, Deltatheroida)

from the Late Cretaceous of Mongolia. Bulletin of Carnegie

Museum of Natural History, 36: 245-266

Russell L S. 1975. Mammalian faunal succession in the Creta-

ceous System of western North America. In: Caldwell W G

E ed. The Cretaceous System of the Western Interior. The

Geological Association of Canada, Special Paper Number 13:

137-161

Sanchiz B. 1998. Salientia. Handbuch der Paläoherpetologie.

Teil 4. Munich: Verlag Dr. Friedrich Pfeil. 1-275

Sankey J T. 2008. Diversity of latest Cretaceous (late Maas-

trichtian) small theropods and birds: teeth from Lance and

Hell Creek formations, USA. Sankey J T & Baszio S

eds.Vertebrate microfossil assemblages. their role in pa-

leoecology and paleobiogeography. Bloomington: Indiana

University Press. 117-134

Sankey J T, Brinkman D B, Guenther M & Currie P J. 2002.

Small theropod and bird teeth from the Late Cretaceous (late

Campanian) Judith River Group, Alberta. Journal of Pale-

ontology, 76(4): 751-763

Schultz S S Jr. 1972. [Geological structure of the Junction Zone

of Urals and Tian-Shan]. Moscow: Nedra. 1-208

Simpson G G. 1960. Notes on the measurement of faunal re-

semblance. American Journal of Science, 258-A: 300-311

Shuvalov V F & Trusova E K. 1979. [Late Cretaceous conchostracans

of Mongolia]. Trudy Sovmestnoi Sovetsko-Mongol'skoi Pa-

leontologicheskoi Ekspeditsii,8: 83-93

Skutschas P P. 2007. New specimens of albanerpetontid am-

phibians from the Upper Cretaceous of Uzbekistan. Acta

Palaeontologica Polonica, 52(4): 819-821

Skutschas P P. 2009. Re-evaluation of Mynbulakia Nesov, 1981

(Lissamphibia: Caudata) and description of a new salaman-

der genus from the Late Cretaceous of Uzbekistan. Journal

of Vertebrate Paleontology, 29(3): 659-664

Smith D & Galton P M. 1990. Osteology of Archaeornithomimus

asiaticus (Upper Cretaceous, Iren Dabasu Formation, Peo-

ple's Republic of China). Journal of Vertebrate Paleontology,

10(2): 255-265

Špinar Z V & Tatarinov L P. 1986. A new genus and species of

discoglossid frog from the Upper Cretaceous of the Gobi

Desert. Journal of Vertebrate Paleontology, 6(2): 113-122

Storrs G W & Efimov M B. 2000. Mesozoic crocodyliforms of

north-central Eurasia. In: Benton M J, Shishkin M A, Unwin

D M & Kurochkin E N eds. The age of dinosaurs in Russia

and Mongolia. Cambridge: Cambridge University Press.

402-419

Sues H-D & Averianov A O. 2009a. Turanoceratops tardabilis

—the first ceratopsid dinosaur from Asia. Naturwissenschaften,

96(5): 645-652

Sues H-D & Averianov A O. 2009b. A new basal hadrosauroid

dinosaur from the Late Cretaceous of Uzbekistan and the

early radiation of duck-billed dinosaurs. Proceedings of the

Royal Society B, 276(1667): 2549-2555

Sukhanov V B. 2000. Mesozoic turtles of Middle and Central

Asia. In: Benton M J, Shishkin M A, Unwin D M &

Kurochkin E N eds. The age of dinosaurs in Russia and

Mongolia. Cambridge: Cambridge University Press. 309-

367

Sukhanov V B, Danilov I G & Syromyatnikova E V. 2008. The

description and phylogenetic position of a new nanhsiung-

chelyid turtle from the Late Cretaceous of Mongolia. Acta

Palaeontologica Polonica, 53(4): 601-614

Swofford D L. 2002. PAUP*. Phylogenetic analysis using Par-

simony (*and other methods). Version 4.0. Sunderland,

Sinauer Associates

Syromyatnikova E V. 2011. Turtles of the genus Ferganemys

Nessov et Khosatzky, 1977 (Adocidae): shell morphology

and phylogenetic position. Proceedings of the Zoological

Institute of the Russian Academy of Sciences, 315(1): 38-52

Syromyatnikova E V & Danilov I G. 2009. New material and a

revision of turtles of the genus Adocus (Adocidae) from the

Late Cretaceous of Middle Asia and Kazakhstan. Proceed-

ings of the Zoological Institute of the Russian Academy of

Sciences, 313(1): 74-94

Tokaryk T T, Cumbaa SL & Storer J E. 1997. Early Late Creta-

ceous birds from Saskatchewan, Canada: the oldest diverse

avifauna known from North America. Journal of Vertebrate

Paleontology, 17(1): 172-176

Tong H, Claude J, Suteethorn V, Naksri W & Buffetaut E. 2009.

Turtle assemblages of the Khorat Group (Late Jurassic-Early

Cretaceous) of NE Thailand and their palaeobiogeographical

significance. In: Buffetaut E, Cuny G, Le Loeuff J & Sutee-

thorn V eds. Late Palaeozoic and Mesozoic Ecosystems in

SE Asia. London: Geological Society of London, Special

Publications 315: 141-152

Van Itterbeeck J, Horne D J, Bultynck P & Vandenberghe N.

2005. Stratigraphy and palaeoenvironment of the dino-

saur-bearing Upper Cretaceous Iren Dabasu Formation, In-

ner Mongolia, People's Republic of China. Cretaceous Re-

search, 26(4): 699-725

Vitek N S & Danilov I G. 2010. New material and a reassess-

ment of soft-shelled turtles (Trionychidae) from the Late

Cretaceous of Middle Asia and Kazakhstan. Journal of Ver-

tebrate Paleontology, 30(2): 383-393

Vullo R & Neraudeau D. 2009. Pterosaur remains from the Ceno-

manian (Late Cretaceous) paralic deposits of Charentes, west-

ern France. Journal of Vertebrate Paleontology, 29(1): 277-282

Watabe M, Tsuihiji T, Suzuki D & Tsogtbaatar K. 2009. The

first discovery of pterosaurs from the Upper Cretaceous of

Mongolia. Acta Palaeontologica Polonica, 54(2): 231-242

Weishampel D B, Barrett P M, Coria R A, Le Loeuff J, Xu X,

Zhao X J, Sahni A, Gomani E M & Noto C R. 2004. Dino-

saur distribution. In: Weishampel D B, Dodson P & Osmól-

ska H eds. The dinosauria (Second edition). Berkeley: Uni-

2 期

Averianov A.等: 晚白垩世中亚与亚洲中部的陆生脊椎动物组合对比

485

versity of California Press. 517-606

Wellnhofer P & Buffetaut E. 1999. Pterosaur remains from the

Cretaceous of Morocco. Paläontologische Zeitschrift,

73(1/2): 133-142

Wilson L E, Chin K, Cumbaa S L & Dyke G J. 2011. A high

latitude hesperornithiform (Aves) from Devon Island: pa-

laeobiogeography and size distribution of North American

hesperornithiforms. Journal of Systematic Palaeontology,

9(1): 9-23

Wu X C. 2005. Crocodylians. In: P J Currie & E B eds. Dinosaur

provincial park: a spectacular ancient ecosystem revealed.

Koppelhus. Bloomington: Indiana University Press. 277-291

Xu X, Wang D Y, Sullivan C, Hone D W E, Han F, Yan R H &

Du F M. 2010a. A basal parvicursorine (Theropoda: Alva-

rezsauridae) from the Upper Cretaceous of China. Zootaxa,

2413: 1-19

Xu X, Wang K, Zhao X & Li D. 2010b. First ceratopsid dino-

saur from China and its biogeographical implications. Chi-

nese Science Bulletin,55(16): 1631-1635

Xu X, Wang K, Zhao X, Sullivan C & Chen S. 2010c. A new

leptoceratopsid (Ornithischia: Ceratopsia) from the Upper

Cretaceous of Shandong, China and its implications for

neoceratopsian evolution. PLoS ONE, 5(11): e13835

Zanno L E. 2010. A taxonomic and phylogenetic re-evaluation

of Therizinosauria (Dinosauria: Maniraptora). Journal of

Systematic Palaeontology, 8(4): 503-543