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Early angiosperms from the Lower Cretaceous of Jixi, eastern ...

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G. Sun, D.L. Dilcher / Review <strong>of</strong> Palaeobotany and Palynology 121 (2002) 91^112 95<br />

rations <strong>from</strong> bed 12 yielded species such as Canningia<br />

pistica Helby, Kiokansium polypes (Cooks.<br />

et Eis.) Below, Muderongia testudinaria Burger,<br />

Muderongia tetracantha (Gocht.) Alberti, Oligosphaeridium<br />

complex (White) Davey et Williams,<br />

and Palaeoperidium cretaceum Pocock. Sun et al.<br />

(1992b) suggest that <strong>the</strong> age <strong>of</strong> <strong>the</strong> dino£agellate<br />

assemblage <strong>from</strong> bed 12 is Valanginian^Hauterivian.However,<br />

He and Sun (2000) studied <strong>the</strong><br />

dino£agellates <strong>from</strong> <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> Chengzihe<br />

Formation <strong>of</strong> <strong>Jixi</strong> and reported <strong>the</strong> following<br />

taxa: Odontochitina operculata (O.Wetzel) De£.<br />

et Cooks., Muderongia tetracantha (Gocht.) Alberti,<br />

Batioladinium? exiguum (Alberti) Brideaux,<br />

Gardodinium trabeculosum (Gocht.) Alberti, Palaeoperidium<br />

cretaceum Pocock, Kiokansium sp.,<br />

Escharisphaeridia pocockii (Sarj.) Erkmen et Sarjeant,<br />

Circulodinium asymmetricum (Burger) He et<br />

Sun, Nyktericysta puyangensis Wang et Qiao and<br />

Oligosphaeridium jixiense He et Sun.These two<br />

authors considered <strong>the</strong> age <strong>of</strong> marine bed 12<br />

yielding <strong>the</strong> dino£agellates to be late Hauterivian,<br />

and <strong>the</strong> age <strong>of</strong> <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> Chengzihe<br />

Formation to be late Valanginian^late Hauterivian<br />

(He and Sun, 2000).Therefore, we suggest<br />

that <strong>the</strong> age <strong>of</strong> <strong>the</strong> early <strong>angiosperms</strong> collected<br />

<strong>from</strong> beds 27^47 in <strong>the</strong> Chengzihe Formation is<br />

late Hauterivian or late Hauterivian^early Barremian<br />

(Fig.2).<br />

The Chengzihe Formation is conformably overlain<br />

by <strong>the</strong> coal-bearing Muling Formation.The<br />

two formations correspond to <strong>the</strong> lower part <strong>of</strong><br />

Lipovich Formation and Ussuri Formation, dated<br />

as Barremian^Aptian age.These correlations can<br />

also be extended to <strong>the</strong> Ussuri Formation, which<br />

is <strong>the</strong> equivalent <strong>of</strong> <strong>the</strong> Older Suchan Formation<br />

<strong>from</strong> <strong>the</strong> neighboring Suchan Basin which overlies<br />

<strong>the</strong> Berriasian^Valanginian marine beds (see Fig.<br />

3 for details <strong>of</strong> this correlation).Therefore, <strong>the</strong><br />

Muling Formation is considered middle^late Barremian<br />

to early Aptian, while <strong>the</strong> whole Chengzihe<br />

Formation is considered Valanginian (or late<br />

Berriasian) to late Hauterivian (or early Barremian)<br />

in age (Sun et al., 1992a,b, 2000).<br />

Only about 80 specimens <strong>of</strong> angiosperm megafossils<br />

have been collected <strong>from</strong> <strong>the</strong> Chengzihe<br />

Formation by <strong>the</strong> authors so far, including <strong>the</strong><br />

three angiosperm reproductive organs described<br />

here, while specimens <strong>of</strong> <strong>the</strong> o<strong>the</strong>r plants associated<br />

with <strong>the</strong> <strong>angiosperms</strong> number nearly 1000.<br />

That means <strong>the</strong> early angiosperm megafossils are<br />

rare (about 8%) in <strong>the</strong> assemblage.This is also <strong>the</strong><br />

case when <strong>the</strong> pollen <strong>from</strong> <strong>the</strong>se sediments was<br />

examined (Shang, 1997). Brenner and Bicko¡<br />

(1992) and Brenner (1996) also found that early<br />

angiosperm pollen is present at only very low percentages<br />

in <strong>the</strong> Valanginian^Hauterivian <strong>of</strong> Israel.<br />

Dilcher and Farley (1988) noted in Cenomanian<br />

age sediments that 63^75% <strong>of</strong> <strong>the</strong> megafossils recovered<br />

were <strong>angiosperms</strong> while only 21^23% <strong>of</strong><br />

<strong>the</strong> miospores recovered were <strong>angiosperms</strong>.This<br />

demonstrates that angiosperm diversity and abundance<br />

dominate as megafossils long before <strong>the</strong>y<br />

do so in <strong>the</strong> miospore record.<br />

There are only a few specimens which are compressions<br />

and with cuticle preserved.The preparations<br />

for anatomical study <strong>of</strong> <strong>the</strong> reproductive<br />

axis and <strong>the</strong> cuticles <strong>of</strong> <strong>the</strong> angiosperm leaves<br />

were made with Schultze solution for <strong>the</strong> macerations,<br />

and scanning electron microscopy (SEM)<br />

observations were made with a JAL-3600 SEM.<br />

The surface <strong>of</strong> <strong>the</strong> reproductive axis was examined<br />

with Zeiss epi£uorescence and individual pollen<br />

and clusters <strong>of</strong> pollen grains were studied.<br />

Often extreme oblique light was used to discern<br />

<strong>the</strong> nature <strong>of</strong> <strong>the</strong> leaf margins and <strong>the</strong> venation<br />

while studying <strong>the</strong> specimens with a Wild M 500<br />

dissecting microscope.The described specimens<br />

are housed in Nanjing Institute <strong>of</strong> Geology and<br />

Palaeontology, Academia Sinica, Nanjing, China.<br />

3. Systematics<br />

The following early angiosperm taxa <strong>from</strong> <strong>the</strong><br />

<strong>Early</strong> <strong>Cretaceous</strong> Chengzihe Formation <strong>of</strong> <strong>Jixi</strong>,<br />

China are described in this paper: Asiatifolium<br />

elegans Sun, Guo et Zheng emend.Sun et Dilcher,<br />

<strong>Jixi</strong>a pinnatipartita Guo et Sun emend.Sun et<br />

Dilcher, <strong>Jixi</strong>a chengzihensis Sun et Dilcher sp.<br />

nov., <strong>Jixi</strong>a sp., Shenkuoa caloneura Sun et Guo<br />

emend.Sun et Dilcher, Zhengia chinensis Sun et<br />

Dilcher gen.et sp.nov., Xingxueina heilongjiangensis<br />

Sun et Dilcher, Xingxuephyllum jixiense<br />

Sun et Dilcher gen.et sp.nov., Reproductive organ<br />

A, Reproductive organ B.

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