ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 16: 1–15 (2017)© by the Fukui Prefectural Dinosaur Museum

VERTEBRATE DIVERSITY OF THE EARLY CRETACEOUS TETORI BIOTA
FROM JAPAN, THE STATE OF THE ART
Shin-ichi SANO
Fukui Prefectural Dinosaur Museum
51-11, Terao, Muroko, Katsuyama, Fukui 911-8601, Japan
ABSTRACTThe comprehensive list of vertebrate skeletal fossils (excluding ichnofossils) from the Early Cretaceous Tetori
Group is provided. Numbers of species of selected vertebrate taxonomic groups of the Tetori Group are comparedwith those from several famous Late Jurassic–Early Cretaceous Lagerstätten to evaluate the vertebrate diversity of
the Tetori Biota. Vertebrate fauna of the Tetori Group is characterized by the presence of a tritylodont synapsid, thatof both diversiﬁed choristoderes and a crocodyliform, remarkably high diversity of the testudines, moderate diversityof mammals, squamates and dinosaurs, and possible low diversity of ﬁsh. Most of the vertebrate taxonomic groups
recognized in other fauna/biotas are present in the Tetori Biota. The vertebrate diversity of the Tetori Biota can beconsidered to be comparable to those of the Late Jurassic–Early Cretaceous Lagerstätten at present, and is expected
to be higher in the near future. Further investigation and comparison of vertebrate diversities in almost coeval strata inseveral regions in East and Southeast Asia probably provide the useful information to reveal the paleo(bio)geographicaland palaeoclimatic reconstruction in Asia, and also the evolution of the Mesozoic terrestrial ecosystem.
Key words : vertebrates, diversity, Tetori Group, Early Cretaceous, Central Japan, East Asia, terrestrial ecosystem,
palaeobiogeography
佐野晋一（2017）手取層群の脊椎動物相はどのぐらい多様なのか．福井県立恐竜博物館紀要 16：1–15．
手取層群の脊椎動物相を概観すべく，論文等で報告された脊椎動物の体化石のリストを作成した．手取層群の脊椎動物相は，トリティロドン類の存在，ワニ類と比較的多様なコリストデラ類の両方の存在，カメ類の顕著な多様性，哺乳類，有鱗類，恐竜類の多様性，魚類の多様性が小さいことによって特徴づけられる．
手取層群の脊椎動物化石の各グループの種数を，後期ジュラ紀～前期白亜紀の幾つかの著名なラガシュテッ
テンと比較した結果，手取生物群には他のラガシュテッテンに知られるほとんど全てのグループが存在しており，かつ，現在知られている種数も他地域のものにほぼ匹敵する規模に達することがわかった．今後，東・東南アジア各地のほぼ同時代の地層から知られる脊椎動物相の多様性を比較することにより，アジアにおける当時の古（生物）地理や古気候の復元，さらには中生代陸上生態系の進化の解明に貢献できるものと期待される．

INTRODUCTION

However, most previous studies mainly focused on the faunasin each area/formation (e.g., Evans et al., 1998; Matsuoka,
2000a; Matsuoka et al., 2002) or on each taxonomic group (e.g.,
Matsumoto et al., 2015; Evans and Matsumoto, 2015; Shibata etal., 2017), and the whole diversity of the Tetori Biota (Matsukawa
et al., 2006) has not been fully understood. In this paper, almost
all available data of the vertebrate skeletal fossils from the Tetori
Group is summarized to evaluate the potential vertebrate diversityof the Tetori Biota in comparison with those of the Jehol Biota
in northeastern China and other Late Jurassic–Early Cretaceous
Lagerstätten.

The presence of rich terrestrial vertebrate fauna has beenrecognized in the Early Cretaceous Tetori Group (Oishi, 1933a,b) in northern Central Japan, and its importance to reveal theevolution of Mesozoic terrestrial ecosystem in East Asia attracts
much attention recently (e.g., Manabe et al., 2000; Matsuoka
et al., 2002; Matsukawa et al., 2006; Sano and Yabe, 2017).
Received June 21, 2017. Accepted October 26, 2017.
Corresponding author ― Shin-ichi SANO
E-mail: ssano＊dinosaur.pref.fukui.jp1

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SHIN-ICHI SANOGEOLOGICAL OUTLINE
The Early Cretaceous Tetori Group is sporadically distributedin the Hakusan Region in the western part of the Hida Belt
(Fig. 1), which is considered to be a fragment of continental
block, possibly a part of the North China Block, or the Jiamushi(or Jiamusi) Block in the eastern margin of the Central Asian
Orogenic Belt before the Miocene opening of the Japan Sea (e.g.,
Jin and Ishiwatari, 1997; Kim et al., 2007; Arakawa et al., 2000;
Kunugiza et al., 2010; Zhao et al., 2013; Takahashi et al., 2017).
The Tetori Group is divided into two depositional stages: DS2
and DS3 (Sano, 2015). In DS2 (Berriasian–late Hauterivian),
the brackish environment is dominant with intercalation ofshallow marine environment in some horizons, whereas a ﬂuvial
environment prevailed without any marine/brackish horizons in
DS3 (Barremian–Aptian).
Vertebrate skeletal fossils have been described or figured
mainly from the Itsuki Formation in the Itoshiro area; the Mitarai,
Okurodani and Amagodani formations in the Shokawa area; theTaie Formation in the Hida-Furukawa area; the Kuwajima, Akaiwa
and Kitadani formations in the Shiramine and Takinamigawa
areas; the Kowashimizu Formation in the Asuwa area (Figs. 1 and
2). Depositional stage and age, and stratigraphical relationshipsof vertebrate fossil-bearing formations of the Tetori Group aresummarized in Figure 2 (See Sano (2015) for details). Amongvertebrate fossil localities in the Tetori Group, KO-2 in thelowermost part of the Okurodani Formation corresponds to upper
part of DS2 (possibly assigned to late Hauterivian or earlier, butlater than Berriasian in age), Kasekikabe in the uppermost partof the Kuwajima Formation lower part of DS3 (Barremian), andthe Kitadani Dinosaur Quarry in the lower part of the Kitadani

FIGURE 1. Index map showing the distribution of the Early Cretaceous
Tetori Group in northern Central Japan. The Kuzuryu and Tetori groups
are distributed in the Hakusan Region, though the former shows a narrow
distribution only in the Itoshiro area (modiﬁed from Maeda (1961b) andFujita (2003)).

Formation upper part of DS3 (Aptian) (Sano and Yabe, 2017).
However, since similar vertebrate taxa including same genera havebeen known from KO-2 and Kasekikabe, and the last occurrenceof brackish molluscs, meaning the transition from DS2 to DS3, isrecognized within the Kuwajima and Okurodani formations, it is
supposed that the strata of these two fossil localities are probably
not so different in age (Sano and Yabe, 2017).
The stratigraphy of three other vertebrate skeletal fossil
localities is brieﬂy mentioned here, because it was not discussed
in Sano (2015) and Sano and Yabe (2017). An unidentified
theropod tooth was recovered from a grey mudstone ﬂoat in Eastof Shiramine Village in the Shiramine area (Tanimoto et al., 2009;
Utsunomiya, 2009). Co-occurrence of a bivalve Nippononaia
tetoriensis and a gastropod with a tooth in the float suggests
this float was most probably derived from the Tetori Group.
Yanagidani, the type and only known locality of N. tetoriensis in
the Shiramine area (Maeda, 1962), is located close to the fossil
discovery site. Thus the fossil-bearing float is supposed to beoriginated from the stratum at Yanagidani, or nearby locality/
horizon. Based on the stratigraphical interpretation of the stratumat Yanagidani in Matsuura (2001), original authors considered thedinosaur fossil was derived from the Akaiwa Formation. However,
Sano and Yabe (2017) discussed this fossil-bearing stratum can becorrelated with the lower part of the Kuwajima Formation (upper
part of DS2), and the latter interpretation is accepted here.
Several vertebrate remains including a new squamata
Tedorosaurus asuwaensis have been described or figured fromthe Sakaidera Alternation of the Tetoti Group in the Asuwa area
(Shikama, 1967, 1969; Yasuno, 2004, 2005). Yasuno (2005)
figured freshwater and brackish water bivalves: Nippononaia
tetoriensis, Megasphaerioides okurodaniensis, Myrene sp.,
Tetoria sp., and a shark tooth Hybodus sp. from this stratum,and compared this fossil bivalve assemblage with those of
the Okurodani Formation and the lower part of the Kuwajima
Formation in Yanagidani, and discuss the possible correlationof the fossil-bearing stratum to the upper part of the Itoshiro
Subgroup of the“Tetori Group”(Maeda, 1961b). Yamada et al.
(2008) revised the stratigraphy of the Tetori Group in this area,
redeﬁning the Kowashimizu Formation, which now includes the
Sakaidera Alternation of Maeda (1961a), and assigned it to the
Itoshiro Subgroup. These recent stratigraphical views are accepted
here, and the vertebrate fossil-bearing formation in this area
probably corresponds to the upper part of DS2.
Manabe (1999) described an isolated premaxillary toothof a tyrannosauroid from the Jobu Formation of the“Itoshiro
Subgroup of the Tetori Group”(Yamada et al., 1989) in the
Kamihambara area, in the eastern part of Kuzuryu District.
Sano (2015) proposed the idea that this fossil-bearing stratum isseparated from the Tetori Group, and assigned to the Managawa
Group in the Hida Gaien Belt. Furthermore, the age of the fossilbearing stratum is still controversial. Thus this record is notincluded in the discussion of this paper. Vertebrate fossil records
from the Kuzuryu Group (= DS1 of Sano (2015)) (e.g., a possible
plesiosaur tooth (Board of Education of Toyama Prefecture, 2003)
and ﬁsh scales (Yasuno, 1994, 1995)) are also excluded.

FIGURE 2. Depositional stages, stratigraphic distribution of marine and brackish horizons, and the occurrences of characteristic bivalves in the Tetori Group (modiﬁed from Sano and Yabe (2017)).
Stratigraphy and molluscan fossil occurrences in the Tetori Group: Maeda (1952b, 1957) for the Itoshiro area; Maeda (1952a) and Matsukawa and Nakada (1999) for the Shokawa area; Matsukawa et al.
(2007) for the Hida-Furukawa area; Maeda (1958, 1962), Tamura (1990), Isaji (1993) and Sano et al. (2008) for the Shiramine and Takinamigawa areas; Yasuno (2004, 2005) andYamada et al. (2008) for
the Asuwa area; see text and Sano (2015) for details. Ammonoid data: Sato et al. (2003, 2008) for the Mitarai Formation in the Shokawa area; Matsukawa and Fukui (2009) for the Inagoe Formation in the
Hida-Furukawa area. Abbreviation: It–Itsuki Formation in the Itoshiro area; Mi–Mitarai Formation, Ok–Okurodani Formation, Am–Amagodani Formation in the Shokawa area; Ta–Taie Formation, Ig–
Inagoe Formation in the Hida-Furukawa area; Kw–Kuwajima Formation, Ak–Akaiwa Formation, Kt–Kitadani Formation in the Shiramine and Takinamigawa areas; Ko–Kowashimizu Formation in the
Asuwa area. The numerical ages are based on International Commission on Stratigraphy (ICS) (2015).

VERTEBRATE DIVERSITY OF THE TETORI BIOTA FROM JAPAN3

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SHIN-ICHI SANOVERTEBRATE DIVERSITY OF THE TETORI GROUP
Methods
To evaluate the vertebrate diversity of the Tetori Group, only
the skeletal fossils (excluding ichnofossils, such as footprints andeggshells) formally described in scientific papers are surveyed.
Indeterminate material is also included in the list, if at least its
higher level (generally Order) classification is mentioned in
the description. However, in case almost all the information of
selected taxonomic groups from the Tetori Group is summarized
in the review papers (e.g., Hirayama, 2006 for testudines; Shibata
et al., 2017 for dinosaurs), some undescribed taxa mentioned inthese papers are also included in the list with original references(usually abstracts for the scientific meeting). Although acrocodyliform from the Tetori Group has not been formallydescribed yet (Kobayashi, 1998), this record has been frequently
mentioned in the palaeobiogeographical and palaeoclimatic
discussions (e.g., Amiot et al., 2011; Matsumoto et al., 2015; Sanoand Yabe, 2017), and thus is included in the list. Such uncertain
records are shown with asterisk.
Since systematic revision of each taxon is beyond the scopeof this paper, its original taxonomic assignment is generally
followed, but higher taxonomic names are sometimes revised
according to their recent usage (e.g., Pantrionychia Joyce et al.,2004). The change of the names applied for the same specimens
or species are carefully surveyed, and related references are
mentioned in one same row of the list.
Results
The comprehensive list of vertebrate skeletal fossils from the
Tetori Group is provided in Table 1. It should be noted that the
number of species in each taxonomic group is tentative, and canbe changed (probably larger) during the course of future studies.
Two elasmobranch and five neopterygian fish, including two
new taxa: Sinamia kukurihime and Tetoriichthys kuwajimaensis(Yabumoto, 2008, 2014), are recognized. At least two amphibians
including Anura and Caudata are present. One new tritylodontid
synapsid Montirictus kuwajimaensis (Matsuoka et al., 2016),
and seven mammals, including four new taxa: Symmetrolestes
parvus, Hakusanobaatar matsuoi, Tedoribaatar reini and
Hakusanodon archaeus (Tsubamoto et al., 2004; Rougier et al.,2007; Kusuhashi, 2008), were described. Among 13 testudines,
only one new taxon Kappachelys okurai (Hirayama et al., 2013)
was formally described at present. A new lizard Tedorosaurus
asuwaensis is the first vertebrate fossil recovered from the
Tetori Group (Shikama, 1967, 1969), though its identificationis still uncertain (Evans and Matsumoto, 2015). After this
discovery, 10 squamates, including 6 new taxa: Kaganaias
hakusanensis, Kuwajimalla kagaensis, Sakurasaurus shokawensis,
Asagaolacerta tricuspidens, Kuroyuriella mikikoi and Hakuseps
imberis (Evans and Manabe, 1999a, 2008; Evans et al., 2006;
Evans and Matsumoto, 2015), were described, though other taxa
were also probably present (Evans and Manabe, 1999a). The

presence of all three known choristoderan morphotypes (shortnecked longirostrine, short-necked brevirostrine and long-necked
brevirostrine) were recognized in the Tetori Group (Matsumoto
et al., 2015), and one new long-necked taxon Shokawa ikoi
was proposed (Evans and Manabe, 1999b). One crocodyliform,
possibly four pterosaurs, and one avis are present. Dinosaurs
excluding aves represent possibly 10 theropods, two sauropods
and possibly six ornithischians. Among them, two theropods:
Fukuiraptor kitadaniensis and Fukuivenator paradoxus (Azuma
and Currie, 2000; Azuma et al., 2016), one sauropod: Fukuititan
nipponensis (Azuma and Shibata, 2010), and three ornithischians:
Albalophosaurus yamaguchiorum, Fukuisaurus tetoriensis andKoshisaurus katsuyama (Kobayashi and Azuma, 2003; Ohashi
and Barrett, 2009; Shibata and Azuma, 2015) were described as
new taxa. In summary, vertebrate skeletal fossils from the Tetori
Group is composed of about 68 taxa in total, among which 22
taxa are newly described.
COMPARISON WITH VERTEBRATE DIVERSITIESOF THE LATE JURASSIC–EARLY CRETACEOUS
LAGERSTÄTTEN
Methods
Numbers of species of selected vertebrate taxonomic groups
from several famous Late Jurassic–Early Cretaceous Lagerstätten
are compared with those of the almost coeval Tetori Groupto evaluate the vertebrate diversity of the Tetori Biota (Table
2, Fig. 3). Six Lagerstätten: the Yanliao Biota (Middle–LateJurassic, 166–159 Ma) in northeastern China, the Solnhofen
Fauna (Late Jurassic) in southern Germany, the Las Hoyas Fauna(Upper Barremian) in Spain, the Wealden Fauna (Barremian–earliest Aptian) in southern United Kingdom, the Jehol Biota(Early Cretaceous, 131–120 Ma) in northeastern China, and theSantana Fauna (Early Cretaceous) in Brazil are selected, andtheir vertebrate diversities are compared using similar data setspublished in Zhou and Wang (2010, 2017) and Sweetman (2016),
though detailed genera and species lists were published only for
the Wealden and Jehol fauna/biotas. Fish in Table 2 and Figure 3
represents total number of species of an agnathan, elasmobranchs,
and neopterygians. The data of the Solnhofen Fauna is very
different from other faunas, because of its outstanding diversityof ﬁsh, the presence of rich marine reptiles, such as ichthyosaurs,
sphenodonts, crocodyliforms, which have not been recognized inother faunas, and the absence of amphibians and mammals, andthus is not shown in Figure 3.
Results
Most of the vertebrate taxonomic groups recognized in otherfaunas are present in the Tetori Biota, except marine reptiles, suchas ichthyosaurs and plesiosaurs (Table 2). Total number of species
in the Tetori Biota (68 species) is more than that of the Yanliao
Biota (40 species), and comparable to those of the Las Hoyas(63 species) and Santana (69 species) faunas, and less than thoseTABLE 1. The comprehensive list of vertebrate skeletal fossils (excluding ichnofossils) from the Early Cretaceous Tetori Group. Taxon names in bold represent the new taxa from the Tetori Group. The
asterisk denotes the taxa, which has not been described formally. Abbreviation: Ak–Akaiwa Formation; Am–Amagodani Formation; Ta–Taie Formation; It–Itsuki Formation; Ko–Kowashimizu Formation;
Kt–Kitadani Formation; Kw-u–upper part of the Kuwajima Formation; Kw-l–lower part of the Kuwajima Formation; Mi–Mitarai Formation; Ok–Okurodani Formation.

VERTEBRATE DIVERSITY OF THE TETORI BIOTA FROM JAPAN5·6

VERTEBRATE DIVERSITY OF THE TETORI BIOTA FROM JAPAN

7·8

TABLE 2. Numbers of species of selected vertebrate taxonomic groups of the Tetori Biota are compared with those from several famous Late Jurassic–Early Cretaceous Lagerstätten.

FIGURE 3. Comparison of the species diversity of the Tetori Biota with those of several famous Late Jurassic–Early Cretaceous Lagerstätten.

VERTEBRATE DIVERSITY OF THE TETORI BIOTA FROM JAPANof the Solnhofen (155–190 species), Wealden (110 species) and
Jehol (171 species) fauna/biotas. In general, the diversity of ﬁshshares the signiﬁcant part of the vertebrate diversity of each fauna.
In the tetrapod diversity, only the Wealden (91 species) and Jehol(156 species) faunal biotas exceed the Tetori Biota (61 species).
Thus the vertebrate diversity of the Tetori Biota can be considered
to be comparable to those of the Late Jurassic–Early Cretaceous
Lagerstätten.
Most striking faunal element of the Tetori Biota is a tritylodont
synapsid, which has not been known in other fauna/biotas.
Furthermore, choristoderes have been known only from the Jehol
and Tetori biotas. It should be noted that choristoderes and acrocodyliform coexist only in the Tetori Biota, though these two
faunal groups occur in different formations of the Tetori Group(Matsumoto et al., 2015). Remarkable higher diversity of thetestudines is also another characteristic feature of the Tetori Biota.
Mammals, squamates and dinosaurs are moderately diverse in the
Tetori Biota. It should be noted that the squamate diversity of theTetori Biota is much higher than that of the Jehol Biota, thoughgenerally the diversity of each vertebrate taxonomic group:e.g., mammals, pterosaurs, dinosaurs, and aves, is remarkably
higher in the Jehol Biota than in the Tetori Biota. Low diversityof ﬁsh in comparison with other fauna/biotas may be additional
characteristic feature of the Tetori Biota.
Compositions of dinosaur assemblages are different in almost
coeval Tetori, Jehol and Wealden fauna/biotas. In the Tetori Biota,
the dinosaur fauna is composed of possibly six ornithischians,
two sauropods and possibly 10 theropods. On the other hand,
12 ornithischians, two sauropods and 24 theropods have been
known in the Jehol Biota; 11 ornithischians, 15? sauropods and
11 theropods in the Wealden Fauna (Sweetman, 2016). Striking
differences among three fauna/biotas are the high diversity oftheropods in the Jehol Biota and that of sauropods in the Wealden
Fauna. Besides them, Tetori dinosaur assemblage shows thecomparable diversity with those of the Wealden and Jehol fauna/biotas.
Discussion
Vertebrate diversities of each fauna/biotas was probably
affected by many biases, such as evolutionary history, habitat,
palaeoecology, palaeobiogeography, research history and/or
activities of selected taxa; palaeoclimatic conditions and theirtemporal changes, depositional environments and taphonomical
conditions of the fossil-bearing strata, and so on. For examples,outstanding diversity of ﬁsh and pterosaurs in the Santana Fauna,
those of mammals, pterosaurs in the Yanliao and Jehol biotas,
those of dinosaurs (especially theropods) and aves in the Jehol
Biota are most probably caused by the extensive studies of thesetaxonomic groups in these fauna/biotas (e.g., Kellner and Campos,
1999; Unwin and Martill, 2007; Zhou and Wang, 2010, 2017).
Consequently, total species number of pterosaurs, theropods andaves in the Jehol Biota is over 100, and nearly 60 percent of total
species number of whole vertebrates. In the Wealden Fauna, its
long (almost 200 years) research history probably contributes its

9high vertebrate diversity.
Most of vertebrate skeletal fossils of the Tetori Biota have
not yet been intensively studied especially in their systematic
assignment, although some taxa, such as neopterygii, squamatesand choristoderes, have (see Table 1). Thus the detailed
comparison among the Tetori Biota and other fauna/biotas needs
await further taxonomic works. However, some characteristicfeatures of the Tetori Biota are brieﬂy discussed here.
The low diversity of the ﬁsh fauna in the Tetori Biota may be
explained in palaeoenviromental and taphonomical meanings bythat a ﬂuvial environment prevailed in the vertebrate fossil richformations of the Tetori Group (e.g., Sano and Yabe, 2017), since
the ﬁsh diversity is usually higher in large lakes than rivers (e.g.,
Joyce et al., 2005). In addition, the preservation potential of thebiotas is also probably higher in the large lacustrine environments,
where the Yanliao and Jehol biotas are preserved (e.g., Pan et al.,2013; Zhou and Wang, 2017), than the ﬂuvial environments (e.g.,
Allison et al., 2008).
Remarkably high diversity of the testudines in the Tetori
Biota owes mainly to the abundance of the Pantrionychia (e.g.,
Hirayama, 2006), which has been absent in the Wealden Fauna(Sweetman, 2016), and very rare (only one specimen) in the Jehol
Biota (Zhou and Wang, 2017). These data support the hypothesisthat the centre of origin of this taxon group existed in the eastern
margin of East Asia and/or Central Asia in the Early Cretaceous(e.g., Hirayama et al., 2013; Nakajima et al., 2017).
Choristoderes have been known only from the Jehol andTetori biotas among seven fauna/biotas. It should be notedthat all three known choristoderan morphotypes are present
only in the Jehol and Tetori biotas in the world (Matsumoto
et al., 2015). Absence of this group in the Santana Fauna is
not surprising, because choristoderes are considered as theLaurasian taxa (e.g., Matsumoto et al., 2015). Its absence andthe presence of crocodyliforms in the European and Yanliao
fauna/biotas is supposed to be explained by the warmer climaticconditions in these regions than those in the Jehol and Tetori
regions (Amiot et al., 2011; Matsumoto et al., 2015; Sweetman,2016). The faunal change from the choristoderan occurrences
without crocodilyforms to a crocodilyform occurrence withoutchoristoderes in the Tetori Group, indicates the climatic change
from cool to warm during the period (possibly Barremian toAptian) when the Tetori Biota flourished (Yabe et al., 2003;
Amiot et al., 2011; Matsumoto et al., 2015). It is notablethat such an Early Cretaceous climatic change has not beenrecognized in the Jehol Biota (Sano and Yabe, 2017). On thecontrary, the faunal change from the crocodylomorph occurrence
without choristoderes in the Yanliao Biota to the choristoderanoccurrences without crocodylomorphs in the Jehol Biota
occurred in the northeastern China (Zhou and Wang, 2017). It issuggested that crocodylomorphs could expand their distribution
to northeastern China in the Late Mesozoic, and their absencein the Jehol Biota should be explained by some palaeoclimaticor palaeogeographical conditions at that time, or their temporalchanges around the Jurassic–Cretaceous transition (e.g., Xu et al.,2017).

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SHIN-ICHI SANOCONCLUDING REMARKSThe study of the vertebrate fossils from the Tetori Group hasa relatively long history, starting 50 years ago (Shikama, 1967),
but active research have been conducted after the discovery of thedinosaurs about 30 years ago (Shibata et al., 2017, Table 1). As
discussed above, the vertebrate diversity of the Tetori Biota can beconsidered to be comparable to those of the Late Jurassic–Early
Cretaceous Lagerstätten, and provide the unique information of
the vertebrate evolution and climatic change in East Asia.
The present vertebrate diversity of the Tetori Biota basicallyowes to the extensive studies of the vertebrate faunas from thetwo localities: KO-2 in the Shokawa area and Kasekikabe in
the Shiramine area (e.g., Evans et al., 1998; Matsuoka, 2000a;
Matsuoka et al., 2002). The recent studies of mammals and an
avian eggshell from the Kitadani Dinosaur Quarry (Imai and
Azuma, 2015; Miyata et al., 2016) clearly indicate the great
potential of the microvertebrate studies in the Kitadani Formation.
Further studies of ichnofossils, such as footprints and eggshells,
may provide additional information of unrecognized vertebrate
diversity of this biota (e.g., Matsukawa et al., 2005; Isaji andMatsushita, 2005; Isaji et al, 2006; Lee et al., 2010), as with theeggshell study of the Sasayama Group (Tanaka et al., 2016).
Yoshimura and Matsuoka (2010) reported a testudine fossil fromthe Ushimaru Fomation in the Shokawa area, which underlies the
Mitarai Formation. This record represents the earliest vertebrate
record in the Tetori Group, and clearly suggests the lower part ofDS2 of Sano (2015) can be also candidate for the future vertebrate
palaeontological works. Furthermore, since brackish and marine
molluscs were reported from many horizons/localities of DS2 of
the Tetori Group (e.g., Sano, 2015; Koarai and Matsukawa, 2016),
sometimes with shark teeth (e.g., Kitaura et al., 1974; Yamada,1990), future discoveries of marine reptiles from the Tetori
Group also can be supposed, as with the Wealden Fauna, where
the occurrence of a plesiosaur was reported (Sweetman, 2016).
Thus it is expected that more diverse vertebrate fauna should berecognized in the Tetori Biota in near future.
Some similarities and remarkable dissimilarities have beenrecognized in the vertebrate faunal and floral compositionof the coeval Tetori and Jehol biotas, and thus possible
palaeobiogeographical differentiation in East Asia is suggested(Sano and Yabe, 2017). Recently the rich Albian vertebrate fauna
has been recognized from the Sasayama Group in the Tamba–
Sasayama Region, southwestern Japan (Saegusa et al., 2008;
Kusuhashi et al., 2013; Ikeda and Saegusa, 2013; Saegusa and
Ikeda, 2014; Ikeda et al., 2015, 2016; Tanaka et al., 2016; Shibata
et al., 2017), which is possibly correlated with DS4 of Sano (2015)
in the Hakusan Region. Combination of the vertebrate diversitiesof the Tetori Biota and the faunal assemblage of the Sasayama
Group possibly provides the information of the faunal change and/or evolution during the Early Cretaceous in the eastern margin of
East Asia.
Palaeogeographical and palaeoclimatic conditions causing thestrong provincialisms of dinosaurs and other biota in the Early
Cretaceous in East and Southeast Asia attract much attention

recently (e.g., Philippe et al., 2014; Amiot et al., 2015; Matsumoto
et al., 2015; Shibata et al., 2017; Suarez et al., 2017). Further
comparison of vertebrate diversities including microvertebrates
in almost coeval strata in East and Southeast Asia: e.g., theSindong and Hayang groups in South Korea (e.g., Lee et al.,
2001; Yabumoto, et al., 2006), the Xinminbao Group in theMazongshan Region in northwestern China (e.g., Tang et al.,2001), Xinlong Formation in the Napai Region in southern China(Mo et al., 2016), and the Khorat Group in northern Thailand(e.g., Buffetaut et al., 2006; Fernandez et al., 2009), with thoseof northeastern China (the Jehol Biota) and Japan (the Tetori
Biota and the Tamba–Sasayama fauna), probably provides theuseful information to reveal the palaeo(bio)geographical andpalaeoclimatic reconstruction in Asia, and also the evolution of
the Mesozoic terrestrial ecosystem.
ACKNOWLEDGMENTS
The author would like to thank Teppei SONODA for providing
the up-to-date information of the testudine fossils from the Tetori
Group and other strata in East Asia, and Ren HIRAYAMA, Ryoko
MATSUMOTO and Toshihiro YAMADA for providing essential
information on the fossil localities and their stratigraphy, andKiichiro HACHIYA, Steven C. SWEETMAN, and Atsushi YABE
for providing useful references. Thanks are extended to Ren
HIRAYAMA, Junchang LÜ and Hai-lu YOU for reviewing themanuscript, and to Hiroto ICHISHIMA and Soichiro KAWABE
for editorial works. This is a contribution to UNESCO-IUGS
IGCP Project 632 and 608.
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