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GEODIVERSITAS • 2025 • 47 (15) © Publica ions scien i iques du Muséum na ional d’His oi e na u elle, Pa is. www.geodi e si as.com
KEY WORDS
Thailand,
Sao Khua Fo ma ion,
palynological assemblage,
Valanginian,
humid sub opical
clima e,
Eas e n Te hys P o ince.
Yan ZHANG
Xiao SHI
College o Ea h Sciences, Jilin Uni e si y, Changchun 130061 (China)
[email p o ec ed] (co esponding au ho )
Syl ie BOURQUIN
Uni . Rennes, CNRS, Géosciences Rennes – UMR CNRS 6118, F-35000, Rennes (F ance)
Wenchun GE
College o Ea h Sciences, Jilin Uni e si y, Changchun 130061 (China)
Chuanbiao WAN
Explo a ion and De elopmen Resea ch Ins i u e o Daqing Oil ield Company L d.,
Daqing 163712 (China)
[email p o ec ed] (co esponding au ho )
P adi NULAY
O ice o Mine al Resou ces Region 2, Depa men o Mine al Resou ces,
Minis y o Na u al Resou ces and En i onmen , Khon Kaen (Thailand)
Submi ed on 29 June 2024 | accep ed on 10 No embe 2024 | published on 25 Sep embe 2025
New Ea ly C e aceous palynology o he Sou heas e n
Kho a Pla eau, No heas e n Thailand: palaeobo anical,
palaeoclima ic, and palynophy ogeog aphical implica ions
u n:lsid:zoobank.o g:pub:0AECD667-E640-4D93-AF6A-475951D76EE8
Zhang Y., Shi X., Bou quin S., Ge W., Wan C. & Nulay P. 2025. — New Ea ly C e aceous palynology o he Sou heas e n
Kho a Pla eau, No heas e n Thailand: palaeobo anical, palaeoclima ic, and palynophy ogeog aphical implica ions.
Geodi e si as 47 (15): 641-657. h ps://doi.o g/10.5252/geodi e si as2025 47a15. h p://geodi e si as.com/47/15
ABSTRACT
The Kho a Pla eau, si ua ed in no heas e n Thailand and ex ending pa ially in o Laos and Cambo-
dia, was a h i ing ecosys em du ing he Ea ly C e aceous. I ha bou ed a di e se a ay o e eb a e
auna, including eshwa e hybodon sha k-like ish, ac inop e ygian ishes, u les, c ocodilians,
he opod and sau opod dinosau s, as well as bi al es. Howe e , no many s udies we e ocused on he
lo a o his ecosys em. This s udy ocuses on palynological indings om he Sao Khua Fo ma ion
in he sou heas e n Kho a Pla eau. A o al o 74 species om 43 gene a we e iden i ied, o ming an
assemblage cha ac e ised by Punc a ispo i es-Dichei opollis-Classopollis. Based on ecen adiome ic
da ing o de i al zi con g ains, he Sao Khua Fo ma ion is as assigned o an age no younge han he
mid o la e Valanginian. As he esul , he key axa ound in his o ma ion ha e been calib a ed and
he age o he palynological assemblage is he e o e Valanginian. The assemblage was domina ed by
pollen o Chei olepidiaceae oge he wi h di e si y o p e idophy e spo es, sugges ing a humid sub-
opical clima e wi h pe iodic a id seasons. Compa a i e analysis wi h o he palynological p o inces
places his a ea wi hin he Eas e n Te hys P o ince o he Pan-Te hys Realm.
642 GEODIVERSITAS • 2025 • 47 (15)
Zhang Y. e al.
INTRODUCTION
The Kho a Pla eau is a as pla eau egion si ua ed in no h-
eas e n Thailand. Co e ing an app oxima e a ea o 155 000
squa e kilome es, his expanse is dis inguished by i s la
and a id e ain. The sauce -shaped pla eau is di ided by he
Phu Phan Moun ains in o wo basins: he no he n Sakhon
Nakhon Basin and he sou he n Kho a Basin. (Fig. 1). The
Kho a Basin, one o he la ges and well-exposed con inen al
sedimen a y basins in sou heas e n Asia, spans ac oss eas e n
Thailand, wes e n Laos, and no he n Cambodia. The Ea ly
C e aceous s a a o he Kho a Basin, collec i ely known
as he Kho a G oup, comp ise i e o ma ions (Fig. 1):
hePhu K adung Fo ma ion (Fm), Ph a Wihan Fm, Sao
Khua Fm, Phu Phan Fm and Khok K ua Fm in ascend-
ing o de (Racey& Goodall 2009). The Kho a G oup has
yielded a ple ho a o e eb a e ossils, including selachians,
ac inop e ygians, sa cop e ygians, emnospondyl amphibians,
u les, c ocodyli o mes, p e osau s, non-a ian dinosau s, and
bi ds (Mani koon e al. 2022).
Among hem, he Sao Khua Fm is amous o i s abundan
dinosau ossils. Te anu an non-a ian he opods we e ound
in he Sao Khua Fm, including he me iacan hosau Siamo-
y annus Bu e au , Su ee ho n& Tong, 1996, spinosau id
“Siamosau us” Bu e au & Inga a , 1986, a possible second
spinosau id axon “Phu Wiang spinosau id B”, mega ap o an
Phuwiang ena o Sama hi, Chan hasi & Sande , 2019, and
coelu osau ian Vayu ap o Sama hi, Chan hasi & Sande ,
2019, o ni homimid Kinna eemimus Bu e au , Su ee ho n&
Tong, 2009, an unde e mined ca cha odon osau id om
Phu Wang, and an unde e mined e anu an, and a leas wo
sau opods, including he somphospondylan Phuwiangosau us
Ma in, Bu e au & Su ee ho n, 1994, and an unde e mined
MOTS CLÉS
Thaïlande,
Fo ma ion Sao Khua,
assemblage
palynologique,
Valanginien,
clima sub opical
humide,
P o ince de l’es de la
Té hys.
axon (Tucke e al. 2022, and e e ences he ein). The au ho s
also da ed he Sao Khua Fm, om de i al zi con g ains, as
no younge han middle o la e Valanginian.
Mo eo e , palynological ossils ha e been documen ed om
he con inen al sedimen s o he Ea ly C e aceous, helping o
de e mine he palyno lo a o he Kho a G oup o ma ions
(Racey& Goodall 2009). Howe e , only wel e gene a o
spo e and pollen ossils we e ound a he Sao Khua Fm (Racey
e al. 1996; Racey& Goodall 2009). To add ess his gap, a
de ailed palynological s udy was conduc ed on i e samples
om he uppe sec ion o he Sao Khua Fm in Nam Yuen
ci y, no heas e n Thailand (Fig. 1). The new palynological
da a s udied he e p o ide aluable in o ma ion o comple e
he egional palynos a ig aphy and acili a e he econs uc-
ion o he palaeo ege a ion and palaeoclima e o his a ea.
GEOLOGICAL SETTING
The Kho a G oup (Fig. 2) ep esen s he la ges ed-bed
sequence on he Ko a Pla eau, bu he age o each o ma ion
wi hin he g oup emains con o e sial due o he absence o
index ossils. The lowe mos Phu K adung Fm may ex end
in o he Ju assic Pe iod, whe eas he Ph a Wihan, Sao Khua,
Phu Phan, and Khok K ua o ma ions a e da ed as “Ea ly
C e aceous” (Racey e al. 1996; Ca e & Moss 1999; Meesook
2000; Ca e & B is ow 2003; Hasegawa e al. 2010).
The Sao Khua Fm is p ima ily composed o conglome a ic
sands one, sil s one, and clays one, as well as nume ous pal-
aeosol laye s (Tucke e al. 2022). Facies analysis and a chi-
ec u al s udies e eal ha he sedimen a ion o Sao Khua Fm
occu ed wi hin a loodplain se ing ed by la ge meande ing
bedload- ich channels (Tucke e al. 2022). Mul iple ypes
RÉSUMÉ
Nou elles données palynologiques du C é acé in é ieu du sud-es du Pla eau du Kho a , no d-es de la
Thaïlande : implica ions paléobo aniques, paléoclima iques e palynophy ogéog aphiques.
Le pla eau de Kho a , si ué dans le no d-es de la Thaïlande e s’é endan pa iellemen au Laos e au
Cambodge, é ai un écosys ème p ospè e du an le C é acé in é ieu . Il ab i ai une a ié é impo an e
d’espèces de e éb és, incluan des poissons d’eau douce essemblan à des equins hybodon es, des
poissons ac inop é ygiens, des o ues, des c ocodiliens, des dinosau es hé opodes e sau opodes, e
des bi al es. Cependan , peu d’é udes se son concen ées su la lo e de ce écosys ème. Ce e é ude
se concen e su les décou e es palynologiques de la Fo ma ion Sao Khua dans le sud-es du Pla eau
de Kho a . L’assemblage palynologique, composé de 74 espèces épa ies en 43 gen es, es ca ac é isé
pa Punc a ispo i es-Dichei opollis-Classopollis. Su la base d’une da a ion adiomé ique écen e de
g ains de zi con dé i iques, la Fo ma ion de Sao Khua n’es pas considé ée comme plus jeune que le
Valanginien moyen ou supé ieu . Ainsi, les axons clés ou és dans ce e o ma ion on é é calib és
e l’assemblage palynologique es donc d’âge alanginien. Ce assemblage é ai dominé pa des g ains
de pollen de Chei olepidiaceae, ainsi que pa une di e si é de spo es de ougè es, impliquan un
clima sub opical humide a ec des saisons a ides pé iodiques. L’analyse compa a i e a ec les au es
p o inces palynologiques pe me de considé e que ce e égion se si uai dans la P o ince o ien ale
de la Té hys, au sein du Domaine pan- é hysien.
643
Ea ly C e aceous Palynology om No heas e n Thailand
GEODIVERSITAS • 2025 • 47 (15)
LAOS
KHORAT
PLATEAU
Bangkok
T a
CAMBODIA
Sakhon Nakhon
Ri e
Mekong
Maha Sa akham
Nakhon Ra chasima
Dong ek
Nakhon
Thai
14°
16°
18°N
104°102°E
Khao Ph a Wihan
Phu Phan Upli
Gul o Thailand
Ko Ku
Qua e na y
Maha Sa akham & Phu Thok Fms
Phu Phan & Khok K ua Fms
Ph a Wihan & Sao Khua Fms
P e-C e aceous
100 km
Palynomo ph samples
N
Phu K adung
Range
ig. 1. — Syn he ic geological map o eas e n Thailand (a e Meesook 2011).

644 GEODIVERSITAS • 2025 • 47 (15)
Zhang Y. e al.
o palaeosols we e iden i ied in his o ma ion. Based on he
cha ac e is ics and geochemical da a o hese palaeosols, pal-
aeoclima e condi ions sugges a p edominan ly s able humid
sub opical clima e, wi h a mean annual empe a u e o
app oxima ely 17-20°C and a mean annual p ecipi a ion o
a ound 1000-1400mm pe yea (Tucke e al. 2022). Based
on adiome ic da ing o de i al zi cons ( ia LA-IPC-MS),
hese au ho s conside ed ha he middle pa o he Sa Khua
Fm was deposi ed no la e han 133.8 ± 1.8 Ma (la e Valangin-
ian) and ha he o e lying Phu Phan Fm no la e han 132.4
± 2.0 Ma (lowe Hau e i ian). Consequen ly, he Sao Khua
Fm is conside ed as no la e han mid- o la e Valanginian.
The ou c op om which he samples we e collec ed is
loca ed 11.8 kilome es om Nam Yuen Ci y, nea he bo -
de s o Thailand, Laos, and Cambodia (Fig. 1). In his a ea,
only he yellow-g een sil s one a he op o he Sao Khua
Fm c ops ou in a small alley nea a emple, which loca ed
a 14°21’31”N, 105°12’34”E (Fig. 2). I is o e lain by he
eddish-b own calca eous, conglome a ic sands one a ibu ed
o he base o he Phu Phan Fm, wi h a con o mable con ac
be ween he wo o ma ions (Racey& Goodall 2009; Meesook
2011). Mos o his a ea is a mili a y es ic ed zone because
i lies a he junc ion o Thailand, Myanma , and Cambodia.
MATERIAL AND METHODS
A o al o i e palynological samples we e collec ed om he
yellow-g een sil s one and muds one beds o he Sao Khua
Fm in Nam Yuen Ci y (Figs. 1, 3A). Each sample was spaced
40cm apa and iden i ied as sk21, sk22, sk23, sk24, and
sk25 om bo om o op. Plan mac o- emains (Fig. 3B) can
be obse ed wi hin he yellow-g een muds one laye .
Each 50g sample was c ushed in o g ains smalle han 0.5cm
in diame e and subjec ed o ea men wi h HCl (10%) o
24hou s due o he high calca eous con en , ollowed by
HF (36%) o 48hou s, hen ea ed wi h HCl (37.5%) o
24hou s again. To sepa a e he o ganic esidue om mine als
and isola e he palynomo phs, a ZnCl-mixed KI hea y liquid
wi h a densi y o 2.2g/cm³ along wi h an 8µm sie e we e
used. Subsequen ly, he slides con aining he palynomo ph
samples we e examined unde a Leica DM4000B mic oscope.
Image p ocessing was conduc ed using Co elD aw so wa e.
The bo anical a ini ies o spo es and pollen ound in he Sao
Khua Fm we e summa ized acco ding o indings epo ed
in p e ious s udies and lis ed in Table 1.
All samples and slides a e cu en ly housed a he Resea ch
Cen e o Paleon ology and S a ig aphy, Jilin Uni e si y, China.
Sands one
Uncon o mi y
In e bedded clays one, sal , gypsum, anhyd i e and po ash.
Sands one, medium o coa se; conglome a e,
sil s one & clays one, ed-b own.
Sands one, conglome a ic sands one and conglome a e,
Sands one, conglome a e, sil s one and clays one, ed-b own,
silc e es & palaeosols.
Sands one, medium-coa se, conglome a e,
sil s one and clays one, ed-b own.
Sands one, ine-medium g ained, conglome a ic sands one,
whi e-g ey, medium-coa se, c oss-bedded.
conglome a e, whi e-g ey.
? Te .
C e aceous
Ju assic ?
Phu Thok Fm
Maha Sa akham Fm
Khok K ua Fm
Phu Phan Fm
Sao Khua Fm
Ph a Wihan Fm
Phu K adung Fm
1000 m
E apo i es Sands one &
conglome a e Sil s one Clays one Palynomo phs
Kho a G oup
Uppe C e aceous
Lowe C e aceous
ig. 2. — C e aceous s a ig aphic column in he Kho a Pla eau, NE Thailand (a e Meesook 2011).
645
Ea ly C e aceous Palynology om No heas e n Thailand
GEODIVERSITAS • 2025 • 47 (15)
able 1
. — Palynomo phs iden i ied in he Sao Khua Fm, wi h hei bo anical a ini ies and clima e signal. Re e ences: 1, Abbink e al. 2004; 2, Al in 1982;
3, A y e al. 2019; 4, Balme 1957; 5, Balme 1995; 6, Boul e & Windle 1993; 7, Coupe 1958; 8, De mann 1963; 9, De mann 1986; 10, De mann e al. 1992;
11, Fila o 1975; 12, F iis e al. 2004; 13, Guigna d e al. 2009; 14, Ha is 1979; 15, Hubba d & Boul e 1997; 16, Ji 1994; 17, Van Konijnenbu g-Van Ci e 1971;
18, Van Konijnenbu g-Van Ci e 1993; 19, K u zsch 1963; 20, Li 1984; 21, Liu e al. 2024; 22, Mande 2011; 23, Rod íguez-Ba ei o e al. 2024; 24, San os e al.
2022; 25, Song e al. 1986; 26, Song e al. 1999; 27, T a e se 2007; 28, T e isan 1971; 29, Volkheime e al. 2009; 30, Wee akoon e al. 2021; 31, Weyland &
K iege 1953; 32, Wheele e al. 2022; 33, Yang e al. 2007; 34, Ziaja 2006. Symbol: *, g ains less han 10. Abb e ia ion: G ., G ains.
Taxa
sk22 sk23 sk24 sk25 Clima e
signal
Bo anical
a ini yG . G . % G . % G .
Fe ns spo e – 50
31.06
72
19.89
– – –
Fo aminispo is won haggiensis (Cookson &
De mann) De mann, 1963
– 1 0.62 – – – Wa m and we Sphagnaceae (8, 19, 33)
Fo aminispo is asymme icus (Cookson & De mann)
De mann, 1963
– – – 1 0.28 – Wa m and we Sphagnaceae (8, 19, 33)
Densoispo i es sp. – – – 1 0.28 – Wa m and we Pleu omeaceae/Selaginellaceae
(5, 27, 31)
Fo eospo i es sp. – – – 2 0.55 – Wa m and we Selaginellaceae (4, 5)
Neo ais ickia sp. – – – 2 0.55 – Wa m and we Lycopodiaceae/Selaginellaceae
(8, 10)
Ve ucosispo i es g ana um (Bollkh.) Gao & Zhao, 1976 – – – 1 0.28 – Wa m and we Selaginellaceae (27, 33)
Ve ucosispo i es obscu ilaesu a us Pocock, 1962 – – – 1 0.28 – Wa m and we Selaginellaceae (27, 33)
Ve ucosispo i es sci ulus Yu & Zhang, 1982 – – – 1 0.28 – Wa m and we Selaginellaceae (27, 33)
Ve ucosispo i es sp. – 3 1.86 – – – Wa m and we Selaginellaceae (27, 33)
Lep olepidi es c . psa osus No is, 1969 – 1 0.62 – – – Wa m and we Lycopodiaceae (6, 11)
Lep olepidi es e uca us Coupe , 1953 – – – 1 0.28 – Wa m and we Lycopodiaceae (6, 11)
Punc a ispo i es sp. – 19 11.8 12 3.31 – Wa m and we Lepidoca paceae (5)
Calamospo a sp. – 1 0.62 – – – Wa m and we Lycopodiaceae, Equise ales (5,
11)
Apicula ispo i es sp. – 1 0.62 – – – Ho and we Lycopodiaceae, Selaginellaceae
(11)
Bacula ispo i es sp. – – – 1 0.28 – Wa m and we P e idophy a, Osmundaceae (11)
Osmundacidi es sp. – – – 1 0.28 – Wa m and we Osmundaceae (5, 6, 7, 11)
Bi e ispo i es po oniaei Delcou & Sp umon , 1955 – 4 2.48 1 0.28 – Wa m and we Hymenophyllaceae (23)
Bi e ispo i es punc a us Wan & Sun, 2014 – – – 1 0.28 – Wa m and we Hymenophyllaceae (23)
Bi e ispo i es sp. – 2 1.24 1 0.28 – Wa m and we Hymenophyllaceae (23)
Con e ucosispo i es minimus Yu & Miao, 1983 – – – 2 0.55 – Ho and we Dicksoniaceae (5)
Cica icosispo i es c . paucis ia us Han, 1983 – – – 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es c . po omacensis B enne , 1963 – – – 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es c . pseudoau i e (Bolkh.) Li,1959 – – – 2 0.55 – Ho and we Anemiaceae (23)
Cica icosispo i es c . sub o undus B enne , 1963 – – – 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es cunei o mis Pocock, 1964 – – – 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es minu aes ia us (Bolkh.) Pocock, 1964
– 1 0.62 – – – Ho and we Anemiaceae (23)
Cica icosispo i es paucis ia us Han, 1983 – 1 0.62 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es solidus (Pu & Wu) Jia, 1986 – – – 1 0.28 – Ho and we Anemiaceae (23)
Cica icosispo i es sp. – 4 2.48 5 1.38 – Ho and we Anemiaceae (23)
Conca issimispo i es ema cidus Yu,1984 – 2 1.24 – – – Ho and we Lygodiaceae (23)
Conca issimispo i es punc a us (Delcou &
Sp umon ) B enne , 1963
– – – 6 1.66 – Ho and we Lygodiaceae (23)
Conca issimispo i es e ucosus Delcou &
Sp umon ,1955
– – – 1 0.28 – Ho and we Lygodiaceae (23)
Impa decispo a api e uca a (Coupe ) Venka achala,
Ka & Raza, 1969
– – – 4 1.1 – Ho and we Cya heaceae (1, 24)
Impa decispo a c . b e e (Ma yno a) Yu, 1989 – 1 0.62 2 0.55 – Ho and we Cya heaceae (1, 24)
Klukispo i es sp. – 2 1.24 – – – Ho and we Lygodiaceae (23)
Ma onispo i es?– – – 1 0.28 – Ho and we Gleicheniaceae (23)
Schizaeoispo i es sp. – – – 3 0.83 – Ho and d y Schizaeaceae (25, 26)
Todispo i es majo Coupe , 1958 – 1 0.62 – – – Wa m and we Osmundaceae (11, 24)
Todispo i es sp. – 1 0.62 – – – Wa m and we Osmundaceae (11, 24)
T ilobospo i es sp. – – – 1 0.28 – Ho and we Schizaeaceae (3, 11, 24, 29)
P e isispo i es mino Li, 1984 – – – 1 0.28 – Ho and we P e idaceae (20)
P e isispo i es undula us Sung & Zheng in Sung & Li, 1976
– – – 3 0.83 – Ho and we P e idaceae (20)
Cya hidi es aus alis Coupe ,1953 – 3 1.86 1 0.28 – Ho and we Cya heaceae (23)
Cya hidi es mino Coupe , 1953 – 1 0.62 – – – Ho and we Cya heaceae (23)
Del oidospo a i egula is (P lug) Sung & Tsao, 1976 – – – 1 0.28 – Ho and we Dip e idaceae, Ma oniaceae
(5, 6, 7, 13, 18, 22)
Del oidospo a sp. – – – 1 0.28 – Ho and we Dip e idaceae, Ma oniaceae
(5, 6, 7, 13, 18, 22)
Dic yophyllidi es sp. – – – 2 0.55 – Ho and we Dip e idaceae, Ma oniaceae
(6, 7, 11, 13, 18)
Undula ispo i es sp. – – – 1 0.28 – Ho and we Ophioglossaceae (25)
Lae iga ospo i es g acilis Wilson & Webs e , 1946 – – – 2 0.55 – Wa m and we Ophioglossaceae (25)
Tenuicon ac ospo i es c e acius Han, 1983 – 1 0.62 – – – Ho and we Unknown
646 GEODIVERSITAS • 2025 • 47 (15)
Zhang Y. e al.
RESULTS
The palynological eco e y was excellen , ou o he i e
samples (sk22, sk23, sk24, sk25) yielded spo es and pol-
len g ains. No ably, samples sk23 and sk24 showed di e se
palynological ichness, o alling 80 species and 46 gene a
(Table 1, Figs. 4-6). The composi ion o bo h sk23 and sk24
showed s iking simila i y, leading o he ecogni ion o he
Punc a ispo i es-Dichei opollis-Classopollis assemblage. This
assemblage is cha ac e ized by a p edominance o gymno-
spe m pollen (68.32-80.11%), a low p esence o p e idophy e
spo es (19.89-31.68%), and he absence o angiospe m pol-
len. No ewo hy gymnospe m pollen include Dichei opollis
T e isan, 1971 (50.31-59.39%), Classopollis P lug, 1953
(9.12-9.32%), as well as common ypes like A auca iaci es
Cookson ex Coupe , 1953 (1.38-3.73%), Cycadopi es Wode
-
house, 1933 (1.86-3.31%), and Taxodiaceaepolleni es K emp
ex Po onié, 1958 (0.28-2.48%). Addi ionally, spo adically
appea ing ypes include Exesipolleni essp., Jugella allus Yu,
1984, Jugella sp., and Callialaspo i es iloba us (Balme) Sckh
De , 1961. Among p e idophy e spo es, Punc a ispo i es
Po onié& Gelle ich, 1933 a e he mos abundan (3.31-
11.80%), ollowed by Cica icosispo i es Po onié& Gelle ich,
1933 (3.59-3.73%). O he signi ican ypes p esen in he
assemblage encompass T ilobospo i es sp., Impa decispo a c .
b e e (Ma yno a) Yu, 1989, Impa decispo a api e uca a
(Coupe ) Venka achala, Ka & Raza, 1969, Schizaeoispo-
i es sp., Densoispo i es sp., Fo aminispo is won haggiensis
(Cookson& De mann) De mann, 1963, Fo aminispo is
c . Asymme icus (Cookson& De mann) De mann, 1963,
Conca issimispo i es ema cidus Yu, 1984, Conca issimispo i es
punc a us (Delcou & Sp umon ) B enne , 1963, Lep ole-
pidi es c . psa osus No is, 1969, Lep olepidi es e uca us
Coupe , 1953, P e isispo i es mino Li, 1984, P e isispo i es
undula us Sung& Zheng in Sung& Li, 1976, Klukispo-
i essp., Fo eospo i es sp., among o he s.
Taxa
sk22 sk23 sk24 sk25 Clima e
signal
Bo anical
a ini yG . G . % G . % G .
Gymnospe ms pollen – 111
68.94
290
80.11
– – –
A auca iaci es aus alis Cookson, 1947 – 6 3.73 2 0.55 – Wa m, semia id-
semihumid
A auca iaceae (5, 9, 17)
A auca iaci es sp. – – – 3 0.83 – Wa m, semia id-
semihumid
A auca iaceae (5, 9, 17)
Callialaspo i es dampie i (Balme) Sukh De , 1961 – 1 0.62 – – – Wa m, semia id-
semihumid
A auca iaceae (5, 6, 11, 15, 17)
Callialaspo i es iloba us (Balme) Sckh De , 1961 – – – 1 0.28 – Wa m, semia id-
semihumid
A auca iaceae (5, 6, 11, 15, 17)
Psophosphae a g andis Bolkho i ina, 1956 – – – 1 0.28 – Wa m, semia id-
semihumid
A auca iaceae (16)
Psophosphae a unda a (Bolkho i ina) Zhang,1978 – – – 2 0.55 – Wa m, semia id-
semihumid
A auca iaceae (16)
Psophosphae a sp. – – – 1 0.28 – Wa m, semia id-
semihumid
A auca iaceae (16)
Classopollis sp. *15 9.32 33 9.12 *Ho and a id Chei olepidiaceae (7, 14, 34)
Dichei opollis e uscus T e isan, 1971 – 81
50.31
215
59.39
*Ho and a id Chei olepidiaceae (28)
Concen ispo i es lep os Yu & Zhang, 1982 – – – 2 0.55 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Concen ispo i es opimus Yu & Zhang, 1982 – – – 2 0.55 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Concen ispo i es sp. – – – 3 0.83 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Pe inopolleni es lima us Lu & Wang, 1983 – – – 2 0.55 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Exesipolleni es sp. – – – 1 0.28 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Taxodiaceaepolleni es sp. – 5 3.11 1 0.28 – Cool and
semihumid
Cup essaceae (1, 2, 24)
Pinuspolleni es minu us (Zakl.) Sung & Zheng,1978 – – – 1 0.28 – Cool and semid y Pinaceae (5)
Pinuspolleni es sp. – – – 1 0.28 – Cool and semid y Pinaceae (5)
Podcca pidi es sp. – – – 1 0.28 – Cool and humid Podoca paceae (25, 30)
Cycadopi es elonga us (Balkh.) Zhang, 1978 – – – 1 0.28 – Ho and semia id Cycadales, Benne i ales,
Ginkgoales (5, 11)
Cycadopi es sp. – 3 1.86 11 3.04 – Ho and semia id Cycadales, Benne i ales,
Ginkgoales (5, 11)
Eucommiidi es sp. – – – 4 1.1 – Ho and semia id Cycadales, Gne ales (32)
Jugella allus Yu, 1984 – – – 1 0.28 – A id Ephed ales, Gne ales (12, 21)
Jugella sp. – – – 2 0.55 – A id Ephed ales, Gne ales (12, 21)
Cay onipolleni es sp. – – – 1 0.28 – Wa m and we Cay oniaceae (25)
To al * 161 – 362 100 *
Table 1. — Con inua ion.
647
Ea ly C e aceous Palynology om No heas e n Thailand
GEODIVERSITAS • 2025 • 47 (15)
DISCUSSION
Age o he pAlynoAssemblAge And sAo KhuA m
The age o he Sao Khua Fm has been discussed o se e al
decades (Tucke e al. 2022, and e e ences he ein). Recen ly,
U/Pb da ings on de i al zi cons eco e ed om he middle
pa o his o ma ion and he o e lying Phu Phan Fm allowed
assigning he Sao Khua Fm o a mid o la e Valanginian age
(Tucke e al. 2022). This well-de ined ch onos a ig aphy
es ablished wi h adiome ic me hods allows o calib a e he
bios a ig aphical anges o some key palynological axa o
he Ea ly C e aceous.
The Punc a ispo i es-Dichei opollis-Classopollis assemblage
con ains a a ie y o axa wi h s a ig aphical signi icance,
such as Cica icosispo i es Po onié& Gelle ich, 1933, T ilo-
bospo i es Po onié, 1956, Impa decispo a Venka achala, Ka &
Raza, 1969, Schizaeoispo i es Po onié ex Delcou & Sp u-
mon , 1955, Densoispo i es De mann, 1963, Fo aminispo is
K u zsch, 1956, Conca issimispo i es Delcou & Sp umon ,
1955, Lep olepidi es Coupe , 1953, P e isispo i es Sun& Zheng,
1976, Klukispo i es Coupe , 1958, Fo eospo i es Balme, 1957,
Exesipolleni es Balme, 1957, and Jugella Mchedlish ili&
Shakhmundes, 1973. Howe e , many o hem ha e long-
anging s a ig aphical occu ences.
The main key species o he Punc a ispo i es-Dichei opollis-
Classopollis assemblage s udied he e is Dichei opollis e uscus
T e isan, 1971. Fi s iden i ied by T e isan (1971) in he
Be iasian-Ba emian Biancone Fm in Tuscany, I aly, Dichei-
opollis e uscus has been ound in a ious egions ac oss
A ica (Libya, Egyp , Mo occo, Senegal, I o y Coas , Gabon,
Came oon, Congo, Angola, Sudan, Chad, and Sou h Sudan),
Sou h Ame ica (Venezuela, B azil), and Asia (Yemen, Thai-
land, Cambodia, and China). I s p edominan p esence in
he Be iasian-lowe Ba emian wo ldwide designa es i as a
aluable ma ke species o he Be iasian-lowe Ba emian
s age in he no he n Gondwana egion (Fig. 7).
Ini ially disco e ed in Ba emian deposi s in no he n Mo occo
(Hoculi 1981), subsequen indings ha e also e ealed he p es-
ence o Dichei opollis e uscus du ing he Be iasian-ea ly Ap ian
age in e al, co-occu ing wi h angiospe ms such as Cla a ipol-
leni es hughesii, S ella opollis sp., Re imonocolpi es sp., A opollis
zona us Doyle, Ja diné& Doe enkamp, 1982, and A opollis
ope cula us Doyle, Ja diné& Doe enkamp, 1982, and A opollis
ope cula us (Gübeli, 1984). In he lowe pa o he Abu Gab a
Fm in he Muglad Basin, Sou h Sudan, Dichei opollis e uscus
has been iden i ied wi h angiospe m pollen species like A opol-
lis zona us, Re imonocolpi es a iplica us Sch ank& Mahmoud,
1998, and S ella opollis densio na us (Lima) Wa d, 1986. The
s udies by Eisawi e al. (2012) and Cole e al. (2017) espec-
i ely de e mined he age o his palynological assemblage o
be Be iasian-Ba emian and Be iasian-lowe Ba emian. The
e olu iona y de elopmen s age o angiospe m pollen in his
assemblage aligns closely wi h ha o he angiospe m pollen
ound in conjunc ion wi h Dichei opollis e uscus in no he n
Mo occo (Gübeli e al. 1984), indica ing ha i s geological
age likely ex ends o he lowe Ap ian. No ably, Dichei opollis
e uscus has been obse ed oge he wi h dino lagella e cys s
in he uppe Hau e i ian-lowe Ba emian ma ine deposi s in
Libya and Egyp (Uwins& Ba en 1988; Dea e al. 2016),
and in Yemen (Racey& Goodall 2009), i s p esence in ma ine
deposi s has been ela ed o a ime ame anging om uppe
Valanginian-lowe Ba emian, as de e mined by dino lagella e
cys s and ac i a chs. In summa y, while Dichei opollis e uscus
may ha e o igina ed as ea ly as he La e Ju assic and con inued
o he lowe Ap ian o he Ea ly C e aceous, i s p ima y p e a-
lence aligns wi h he Be iasian-lowe Ba emian.
AB
Phu Phan Fm
Phu Phan Fm
Sao Khua Fm
Sao Khua Fm
ig. 3. — A, Ou c op o he upmos o he Sao Khua Fm in Nam Yuen Ci y; B, Cup essinocladus Sewa d, 1919 ( he pen is 15 cm in leng h). Pho os: Xiao Shi.
654 GEODIVERSITAS • 2025 • 47 (15)
Zhang Y. e al.
cha ac e ized by a no able p esence o Dichei opollis e uscus,
alongside a ious species o Classopollis, and ypical pollen
axa, such as A auca iaci es, Inape u opolleni es, Ephed ipi es,
and Eucommiidi es, among o he s. Bisacca e coni e pollen
is a e, wi h a low pe cen age o spo es, which is consis en
wi h he ypical lo a o he No he n Gondwana p o ince
(o Dichei opollis e uscus/A opollis P o ince). Consequen ly,
his assemblage ep esen s a signi ican componen o he lo a
wi hin he Pan-Te hys Realm, si ua ed in he sou he nmos
pa o he Eas e n Te hys P o ince.
Acknowledgemen s
We hank P o . Jianguo Li o help ul discussion and
cons uc i e sugges ions o he manusc ip . The au ho s
hank D . S. Mullin o p oo eading he English con en .
This pape is unded by he China Schola ship Council.
We a e e y g a e ul o he edi o Emmanuel Cô ez, and
wo anonymous e iewe s o hei cons uc i e commen s
and sugges ions.
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