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Earth Science Frontiers, Vol. 17, Special Issue, Aug. 2010 ISSN 1005-2321<br />

Froghoppers, Leafhoppers, Planthoppers <strong>and</strong> Their Allies from the<br />

Mesozoic of Northeastern Ch<strong>in</strong>a<br />

(Hemiptera: Cicadomorpha <strong>and</strong> Fulgoromorpha)<br />

250<br />

Shu Li 1 , Y<strong>in</strong>g WANG 2 , Dong Ren 3 , Jacek Szwedo 4 , Hong Pang 1<br />

1. College of Life Sciences, Sun Yat-sen University, 135 X<strong>in</strong>gangxi Road, Haizhu District, Guangzhou 510275, Ch<strong>in</strong>a<br />

(E-mail: shulyla@yeah.net)<br />

2. Beij<strong>in</strong>g Museum of Natural History, 126 Tianqiao South Street, Beij<strong>in</strong>g 100050, Ch<strong>in</strong>a<br />

3. College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beij<strong>in</strong>g 100037, Ch<strong>in</strong>a<br />

4. Museum <strong>and</strong> Institute of Zoology, Polish Academy of Sciences, 64, Wilcza Street, PL00-679 Warszawa, Pol<strong>and</strong><br />

The northeastern Ch<strong>in</strong>a, especially Jiulongshan<br />

Formation of Middle <strong>Jurassic</strong> <strong>and</strong> Yixian Formation of<br />

Late <strong>Jurassic</strong> to Early <strong>Cretaceous</strong>, yield a mass of<br />

notable fossils <strong>in</strong>clud<strong>in</strong>g abounded froghoppers, leaf-<br />

hoppers, planthoppers <strong>and</strong> their relatives (hoppers),<br />

which provide important evidence for study of the<br />

orig<strong>in</strong> <strong>and</strong> early evolution. The published report of<br />

fossil records <strong>in</strong>cluded at least representatives of nearly<br />

50 species placed <strong>in</strong> 20 genera (Wang et al., 2006), but<br />

this number vastly <strong>in</strong>creased s<strong>in</strong>ce then. Representa-<br />

tives of various families are present <strong>in</strong> northeastern<br />

Ch<strong>in</strong>a deposits, both ext<strong>in</strong>ct <strong>and</strong> extant.<br />

The most abundant group present <strong>in</strong> both <strong>Jurassic</strong><br />

<strong>and</strong> <strong>Cretaceous</strong> strata is Procercopidae (Cercopoidea),<br />

which covered about 40% of all specimens collected.<br />

However, the known taxonomic diversity of these<br />

froghoppers must be recognized aga<strong>in</strong> <strong>and</strong> future<br />

systematic revisions are needed.<br />

Tettigarctidae (Cicadoidea) is the most ancient<br />

l<strong>in</strong>eage still present <strong>in</strong> recent fauna, <strong>and</strong> the fossil<br />

records can date back to term<strong>in</strong>al Triassic (Shcher-<br />

bakov, 2009). Ancient tettigarctids are ancestors of<br />

recent s<strong>in</strong>g<strong>in</strong>g cicadas (Cicadidae), known <strong>in</strong> fossil<br />

record s<strong>in</strong>ce the Late <strong>Cretaceous</strong>. Tettigarctidae were<br />

morphologically diverse <strong>and</strong> quite abundant <strong>in</strong> the<br />

<strong>Jurassic</strong> <strong>and</strong> <strong>Cretaceous</strong>, like Cicadidae <strong>in</strong> the Cenozoic<br />

<strong>and</strong> nowadays. The pronotum conceal<strong>in</strong>g major of<br />

mesonotum seems to be an autapomorphy of Tettigarc-<br />

t<strong>in</strong>ae (Wang <strong>and</strong> Zhang, 2009). 23% of the fossil<br />

materials from the northeastern Ch<strong>in</strong>a belong to Tetti-<br />

garctidae, but so far, only 4 def<strong>in</strong>itive species of<br />

Tettigarctidae are known.<br />

Representatives of ext<strong>in</strong>ct l<strong>in</strong>eage Palaeon-<br />

t<strong>in</strong>oidea, i. e. Palaeont<strong>in</strong>idae were huge, moth-like<br />

<strong>in</strong>sects, resembl<strong>in</strong>g a little, but not related to s<strong>in</strong>g<strong>in</strong>g<br />

cicadas, which witnessed the rise <strong>and</strong> fall of d<strong>in</strong>osaurs.<br />

Their w<strong>in</strong>g span ranges 10-36 mm; w<strong>in</strong>gs are very<br />

often disruptive patterned (Wang <strong>and</strong> Ren, 2009), <strong>and</strong><br />

body is often covered with scales or dense hairs. A<br />

comb<strong>in</strong>ation of cryptic coloration <strong>and</strong> disruptive<br />

mark<strong>in</strong>g enhanced their camouflage. Palaeont<strong>in</strong>ids<br />

existed from the Triassic to the Mid <strong>Cretaceous</strong>. They<br />

make up 14% of fossil specimens from northeastern<br />

Ch<strong>in</strong>a.<br />

Hylicellidea, the common ancestors of the liv<strong>in</strong>g<br />

Cicadomorphora, evolved from Prosboloidea by the<br />

Middle Triassic. The ext<strong>in</strong>ct family Hylicellidae had<br />

the first appearance <strong>in</strong> the Middle Triassic (Anisian) of<br />

Vosges, France. It was abundant <strong>in</strong> the <strong>Jurassic</strong> <strong>and</strong><br />

became ext<strong>in</strong>ct by the Mid-<strong>Cretaceous</strong> (Shcherbakov<br />

<strong>and</strong> Popov, 2002). Hylicellidae are quite common <strong>in</strong><br />

the <strong>in</strong>sectiferous deposits from the northeastern Ch<strong>in</strong>a,<br />

often very well preserved <strong>and</strong> present<strong>in</strong>g dist<strong>in</strong>ct<br />

morphological disparity. They constitute at least 7% of<br />

the known specimens, but their taxonomic diversity is<br />

still weakly recognised <strong>and</strong> needs further <strong>in</strong>vesti-<br />

gations.<br />

The knowledge on the other groups is virtually<br />

rear; a few weakly preserved specimens could be<br />

ascribed to Karajassidae or their descendants Cicadel-<br />

lidae. Some others seem to be related to Archijassidae,<br />

however the taxonomic status of this family is still<br />

under debate.<br />

The 2% rema<strong>in</strong><strong>in</strong>g <strong>in</strong>clude some uncerta<strong>in</strong> taxa<br />

<strong>and</strong> only a few Fulgoromorpha. The study of fossil<br />

Fulgoroidea from Ch<strong>in</strong>a is still at the debut stage, but<br />

the number of new outcrops <strong>and</strong> specimens is <strong>in</strong>-<br />

creas<strong>in</strong>g. S<strong>in</strong>gle specimen ascribed to the paraphyletic<br />

‘Fulgoridiidae’ was identified <strong>in</strong> the late Middle<br />

<strong>Jurassic</strong> deposits of Daohugou. However, it clearly<br />

differs from all forms known from Europe <strong>and</strong> Asia.<br />

Another phanthoppers group of taxa <strong>in</strong>clud<strong>in</strong>g<br />

genera Lapicixius Ren, Y<strong>in</strong> et Dou, 1998, Y<strong>and</strong>ucixius<br />

Ren, Lu et Ji, 1995 <strong>and</strong> another not yet described genus<br />

superficially resembles Cixiidae, but <strong>in</strong> details of vena-<br />

tion pattern, head capsule <strong>and</strong> leg structures clearly<br />

differs from it <strong>and</strong> deserve familial placement.<br />

The <strong>in</strong>crement of number of specimens collected<br />

<strong>and</strong> exam<strong>in</strong>ed is br<strong>in</strong>g<strong>in</strong>g a number of new taxa<br />

described. The known taxa are under revisionary<br />

studies as well. These allow further comparisons. By<br />

analyz<strong>in</strong>g the record of animals <strong>and</strong> associated plants<br />

fossil, palaeoclimatic data <strong>and</strong> palaeoecological data<br />

the processes of evolution of Cicadomorpha <strong>and</strong> Ful-<br />

goromorpha could be traced <strong>and</strong> analyzed. Concurrent<br />

changes <strong>in</strong> both plants <strong>and</strong> flourish of predators dur<strong>in</strong>g

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