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

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only <strong>from</strong> dispersed pollen grains.It is signi¢cant<br />

that we have <strong>the</strong> same pollen type <strong>from</strong> an in£orescence<br />

in situ which is seemingly attached to a<br />

dicotyledonous leaf.Previously it was thought<br />

that <strong>the</strong> sulcus <strong>of</strong> monosulcate angiosperm pollen<br />

were derived <strong>from</strong> a gymnosperm ancestor which<br />

had monosulcate pollen.However, Brenner proposes<br />

a new model for <strong>the</strong> evolution <strong>of</strong> <strong>the</strong> monosulcate<br />

aperture in <strong>angiosperms</strong>.He suggests that<br />

<strong>the</strong> sulcus was developed <strong>from</strong> within <strong>the</strong> angiosperm<br />

lineage and that its presence in <strong>angiosperms</strong><br />

is entirely an angiosperm feature and not<br />

derived <strong>from</strong> a non-angiospermous ancestor.The<br />

inaperturate nature <strong>of</strong> both <strong>the</strong> pollen mention<br />

here and that described by Brenner may be <strong>the</strong><br />

result <strong>of</strong> <strong>the</strong> sometimes cryptic nature <strong>of</strong> apertures<br />

in some <strong>angiosperms</strong>. Endress (1987) indicates<br />

that in some genera <strong>the</strong> apertures are <strong>of</strong>ten hidden<br />

by reticulate exine ornamentation.The occurrence<br />

<strong>of</strong> very similar, small, circular, ‘inaperturate’, reticular,<br />

tectate-columellate pollen which Brenner<br />

found as dispersed pollen in Israel, is known<br />

<strong>from</strong> a megafossil in situ in China.This new material<br />

may provide new characters useful in understanding<br />

<strong>the</strong> nature <strong>of</strong> early <strong>angiosperms</strong>.<br />

The in£orescence consists <strong>of</strong> a clustering <strong>of</strong><br />

small helically arranged £orets borne on a ra<strong>the</strong>r<br />

stout axis.Because <strong>of</strong> <strong>the</strong> carbonaceous nature <strong>of</strong><br />

<strong>the</strong> preservation and <strong>the</strong> crowded nature <strong>of</strong> <strong>the</strong><br />

£orets, it is di⁄cult to discern <strong>the</strong> nature <strong>of</strong><br />

each £oret.The parts seem to be very small, forming<br />

£orets 1.0^1.5 mm in diameter. Since <strong>the</strong> pollen<br />

are derived <strong>from</strong> <strong>the</strong> £orets in situ, we assume<br />

that <strong>the</strong>re were an<strong>the</strong>rs in <strong>the</strong> £orets, but do not<br />

know if <strong>the</strong> £orets were bisexual or unisexual.<br />

However, it seems most probable that since <strong>the</strong><br />

reproductive axis appears to be still attached to<br />

<strong>the</strong> leaf, in which axis it was borne, and because<br />

Plate V.<br />

G. Sun, D.L. Dilcher / Review <strong>of</strong> Palaeobotany and Palynology 121 (2002) 91^112 107<br />

most all <strong>of</strong> <strong>the</strong> £orets are still attached to a common<br />

axis, with no indication <strong>of</strong> fruit formation,<br />

that <strong>the</strong> £orets are young or immature.Because<br />

pollen is present and well formed, perhaps <strong>the</strong><br />

pollen bearing organs matured before <strong>the</strong> ovules.<br />

Never<strong>the</strong>less, until we can collect additional material<br />

and make more preparations, we consider<br />

that <strong>the</strong>se £orets were ei<strong>the</strong>r bisexual or male.<br />

Small bisexual £orets have been described <strong>from</strong><br />

<strong>the</strong> Albian (Friis et al., 1994) and Turonian (Crepet<br />

and Nixon, 1994) sediments in <strong>eastern</strong> North<br />

America.These £orets are small (2.2 mm), pedicellate,<br />

bear monosulcate pollen and are described<br />

as isolated £orets.The only feature similar between<br />

<strong>the</strong> fossil £orets <strong>of</strong> Virginianthus and Xingxueina<br />

is <strong>the</strong> small size <strong>of</strong> <strong>the</strong> £orets.But this is<br />

important because it indicates that several early<br />

<strong>angiosperms</strong> had £owers that were small, many<br />

<strong>of</strong> which were crowded upon an individual axis.<br />

Friis et al.(1994) compares Virginianthus with<br />

Calycanthaceae and notes many similarities between<br />

<strong>the</strong> fossil and several genera <strong>of</strong> that family,<br />

several <strong>of</strong> which have small £orets. Crepet and<br />

Nixon (1994) in <strong>the</strong>ir review <strong>of</strong> <strong>the</strong> fossil £owers<br />

<strong>of</strong> <strong>the</strong> Magnoliidae suggest that <strong>the</strong> angiosperm<br />

ancestral groups probably had small simple £owers<br />

with parts in whorls similar to that known for<br />

<strong>the</strong> Chloranthaceae, Platanaceae, Buxaceae and<br />

Lauraceae (Friis et al., 1991).Also, Dilcher<br />

(1979) suggested that early <strong>angiosperms</strong> may<br />

have had small simple £owers.The reproductive<br />

axis appears to be borne in <strong>the</strong> axil <strong>of</strong> <strong>the</strong> leaf <strong>of</strong><br />

Xingxuephyllum (Plate V, 1B, Fig.4G).There are<br />

two small carbon fragments <strong>of</strong> <strong>the</strong> petiole missing,<br />

but it appears as if <strong>the</strong> petiole continued to <strong>the</strong><br />

slightly enlarged base which is in common with<br />

<strong>the</strong> base <strong>of</strong> <strong>the</strong> reproductive axis.We also considered<br />

a possibility that <strong>the</strong> two fossils just hap-<br />

1A, 3^5. Xingxueina heilongjiangensis Sun et Dilcher.<br />

1A, 3. In£orescence, SC10025. 1A. Bar = 0.5 cm. 3. Bar = 0.2 cm.<br />

4.Pollen grain <strong>from</strong> <strong>the</strong> in£orescence.SEM0775.Bar = 10 Wm.<br />

5.Pollen grains in situ on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> in£orescence as viewed by epi£uorescence microscopy, SC10025.Bar = 80<br />

Wm.<br />

1B, 2. Xingxuephyllum jixiense gen.et sp.nov.Incomplete leaf probably connected organically with <strong>the</strong> in£orescence,<br />

Holotype SC10026. 1B. Bar = 0.5 cm. 2. Bar = 0.6 cm.

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