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

New Haglidae (Insecta: Orthoptera: Hagloidea) from the <strong>Jurassic</strong> of<br />

Ch<strong>in</strong>a <strong>and</strong> Their Implications for the Early Evolution of Acoustic<br />

Communication<br />

Yan Fang, Haichun Zhang, Bo Wang<br />

State Key Laboratory of Palaeobiology <strong>and</strong> <strong>Stratigraphy</strong>, Nanj<strong>in</strong>g Institute of Geology <strong>and</strong> Palaeontology, Ch<strong>in</strong>ese Academy of<br />

Sciences, Nanj<strong>in</strong>g 210008, Ch<strong>in</strong>a (E-mail: fangyan1001@yahoo.cn; yanfang@nigpas.ac.cn)<br />

The orthopteran superfamily Haglioidea is a<br />

crown group of Ensifera which existed from the<br />

Triassic to recent (Sharov, 1968; Gorochov, 1995; Yuan,<br />

2006). The fossil record of this group is very rich <strong>and</strong><br />

diverse dur<strong>in</strong>g <strong>Jurassic</strong> times. It <strong>in</strong>tensively diversified<br />

<strong>in</strong> the Middle <strong>Jurassic</strong> <strong>and</strong> Early <strong>Cretaceous</strong>, <strong>and</strong> then<br />

quickly decl<strong>in</strong>ed. There are two small extant sub-<br />

families: Prophalangops<strong>in</strong>ae, with 5 species <strong>in</strong> 3 genera<br />

from Ch<strong>in</strong>a <strong>and</strong> northeastern India; <strong>and</strong> Cyphoderr<strong>in</strong>ae,<br />

with 4 species <strong>in</strong> 2 genera from western North America<br />

(Zeuner, 1939; Carpenter, 1992; Liu et al., 2009). The<br />

Hagloidea was possibly orig<strong>in</strong>ated from the Carboni-<br />

ferous to Permian Oedischioidea, which is also the<br />

ancestor of two large recent superfamilies: Tetti-<br />

gonioidea <strong>and</strong> Stenopelmatoidea (Gorochov, 1995).<br />

The ext<strong>in</strong>ct family Haglidae is the second largest<br />

group of Hagloidea, <strong>and</strong> conta<strong>in</strong>s about 56 genera from<br />

the Lower Triassic to <strong>Cretaceous</strong> of Europe <strong>and</strong> Asia.<br />

However, most known fossils are just w<strong>in</strong>g impressions,<br />

<strong>and</strong> some of them need re-consideration. So far, few<br />

comprehensive phylogenetic analyses have been at-<br />

tempted. The family Haglidae appears to be a para-<br />

phyletic group, <strong>and</strong> the relationships between major<br />

l<strong>in</strong>eages are still poorly understood (Béthoux, 2002).<br />

The phenomenon of sexual dimorphism is wide-<br />

spread <strong>in</strong> fore w<strong>in</strong>gs of <strong>Jurassic</strong> Haglidae, of which<br />

most males have acoustic mat<strong>in</strong>g system. These <strong>in</strong>sects<br />

produce call<strong>in</strong>g <strong>and</strong> courtship sounds through friction<br />

between specialized ve<strong>in</strong>s of fore w<strong>in</strong>gs, <strong>and</strong> received<br />

these signals primarily via auditory tympana on the<br />

front tibiae of both sexes (Jost <strong>and</strong> Shaw, 2006). This<br />

k<strong>in</strong>d of acoustic structures <strong>and</strong> communication is consi-<br />

dered to be the most primitive of all tegm<strong>in</strong>ally stri-<br />

dulat<strong>in</strong>g Ensifera. Tegm<strong>in</strong>al stridulatory morpho- logy<br />

<strong>and</strong> tibial tympana of Ensifera have evolved more than<br />

once <strong>in</strong> this suborder (Zeuner, 1939; Sharov, 1968; Jost<br />

<strong>and</strong> Shaw, 2006), <strong>and</strong> the Middle <strong>Jurassic</strong> is an im-<br />

portant period of the evolution of stridulatory apparatus<br />

<strong>in</strong> Haglidae. The venation of fore w<strong>in</strong>gs related to<br />

mak<strong>in</strong>g song changed dist<strong>in</strong>ctly <strong>in</strong> the Middle <strong>Jurassic</strong>,<br />

<strong>and</strong> two opposite trends occur <strong>in</strong> the evolution of<br />

stridulatory apparatus of this group (Gorochov, 2003).<br />

The detailed research on the <strong>Jurassic</strong> Haglidae is very<br />

important not only for underst<strong>and</strong><strong>in</strong>g the early evolu-<br />

tion of acoustic communication, but also for <strong>in</strong>vesti-<br />

gat<strong>in</strong>g the phylogenetic relationships among recent<br />

l<strong>in</strong>eages of Tettigonioidea <strong>and</strong> Stenoplmatoidea.<br />

Very recently, abundant Haglidae were discovered<br />

from the Middle <strong>Jurassic</strong> of Ch<strong>in</strong>a (Fang et al., 2007,<br />

2009; Gu et al., 2009). A new genus with two species of<br />

Haglidae, was identified based on well-preserved<br />

fossils from the Middle <strong>Jurassic</strong> Daohugou Biota (Inner<br />

Mongolia, Ch<strong>in</strong>a). The diagnostic characters <strong>and</strong> syste-<br />

matic position of genera Mongolovoliopus (Gorochov,<br />

1986), Bacharovololiopus (Gorochov, 1986) Bachara-<br />

boilus (Gorochov, 1988) <strong>and</strong> Tettaboilus (Gorochov,<br />

1988) are discussed <strong>in</strong> detail. The term<strong>in</strong>al diagnostic<br />

characters of Haglidae are discussed, <strong>and</strong> subfamilies,<br />

genera, <strong>and</strong> species are keyed. The newly described<br />

genus is characteristic by the long arched ScA, oblique<br />

ve<strong>in</strong> between RP <strong>and</strong> MA, <strong>and</strong> stridulatory on fore<br />

w<strong>in</strong>gs fairly strong <strong>and</strong> plumbed to posterior marg<strong>in</strong>.<br />

The new genera shares some features with basal<br />

Haglids. The absence of the oblique ve<strong>in</strong> between RP<br />

<strong>and</strong> MA <strong>and</strong> strengthen of CuP <strong>in</strong>dicate the transform of<br />

orientation of stridulatory ve<strong>in</strong> <strong>and</strong> reduction of middle<br />

part of fore w<strong>in</strong>gs. These characters reflect the im-<br />

provement of stridulatory apparatus of tegm<strong>in</strong>a pro-<br />

bably due to the appearance of new predatory verte-<br />

brates with good visual analyser <strong>in</strong> the <strong>Jurassic</strong>.<br />

Key words: Haglidae; Orthoptera; <strong>Jurassic</strong>; Ch<strong>in</strong>a<br />

References:<br />

Béthoux O., Nel A. Venational pattern <strong>and</strong> revision of<br />

Orthoptera sensu n. <strong>and</strong> sister group. Phylogeny of<br />

Orthoptera sensu n. Zootaxa, 2002, 96: 1-88.<br />

Charpenter F.M. Treatise on Invertebrate Palaeon-<br />

tology. Part R. Arthropoda 4, 3. Superclass Hexa-<br />

poda. The Geology Society of America, Boulder,<br />

Colorado, <strong>and</strong> University of Kansas, Lawrence,<br />

Kansas, 1992: 655.<br />

Fang Y. Zhang H.C. Wang B., et al. New taxa of Aboi-<br />

l<strong>in</strong>ae (Insecta, Orthoptera, Prophalangopsidae)<br />

from the Middle <strong>Jurassic</strong> of Daohugou, Inner<br />

Mongolia, Ch<strong>in</strong>a. Zootaxa, 2007, 1637: 55-62.<br />

Fang Y. Zhang H.C. Wang B. A new species of Aboilus<br />

(Insecta, Orhtoptera, Prophalangopsidae) from the<br />

Middle <strong>Jurassic</strong> of Daohugou, Inner Mongolia,<br />

Ch<strong>in</strong>a. Zootaxa, 2009, 2249: 63-68.<br />

Gorochov A.V. System <strong>and</strong> evolution of the suborder<br />

Ensifera (Orthoptera) (<strong>in</strong> 2 parts). Proceed<strong>in</strong>gs of<br />

the Zoological Institute, Russian Academy of<br />

Sciences, 1995, 260 (1/2): 1-224, 1-213 (<strong>in</strong> Rus-<br />

169

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