New Fossil Elaterid (Coleoptera :Polyphaga:Elateridae)From Yixian Formation of Western Liaoning, China

PROGRESS IN NATURAL SCIENCE Vol .17 , No .10 , October 2007

New fossil elaterid (Coleoptera :Polyphaga :Elateridae)from Yixian Formation of western Liaoning , China＊

Chang Huali, Ren Dong ＊＊ and Shih Chungkun (College of Life Science , Capital Normal University , Beijing 100037 , C hina)

Accepted on April 23 , 2007

Abstract A new species of the elaterid genus Cryptocoelus Dolin and Nel, 2002 from the Late Jurassic to Early C retaceous Yixian Formation in the western Liaoning , China is described .Diagnosis of the genus is revised , and the systematic position of the genus is briefly discussed .Because the elaterids originated in the Early to Mid-Ju rassic , this new material from the Late Jurassic-Early Cretaceous w ill enhance our understanding of the transition betw een ancient elaterids and extan t ones, bridge the gap of cryptic relationships betw een the Mesozoic cupedids and elaterids, and expand our know ledge of their evolutionary history .

Keywords: Elateridae , new species, Yixian Formation, China.

The Elateridae is a large family in the order Karatau[ 7 —11] in Kazakhstan and 18 species in eig ht Coleoptera .The family is composed of more than genera from China[ 11 , 15, 16 , 18, 21 , 23—25] . Elaterids 10000 species , w hich are usually placed in 11 sub- probably originated in the Early to Mid-Jurassic , [ 1] families .Among these , over 800 species have been flourished by the Late Jurassic , and many recent gen- recorded from China[ 2 , 3] .Elaterids are common and era w ere established by the Early Palaeogene[ 4] .Late abundant in all parts of the w orld .Usually , they are Jurassic-Early Cretaceous is just the transitional peri- easily recognized by their ability to jum p into the air od between ex tant and ex tinct elaterids , and the new w hile making a clicking noise and by acute posterio r material from the Yixian Formation w ill enhance our pronotal angles , and possession of well-developed understanding of the transition betw een ancient ela- metacox al plates .The latter feature has m ade it much terids and ex tant ones , and expand our know ledge of easier to identify compressed fossil fo rms of the fami- their evolutionary history . ly[ 4] .Another intrig uing biological feature of this family is having prolonged larval form s and short- Up to date , fossils of one genus and three species lived adult fo rms occupying quite different niches . have been collected from Yixian Formation , w estern [ 11] Adults are nocturnal living on vegetation , while lar- Liaoning , China .The Yixian Fo rmation compris- vae occur in a variety of habitats , including soil , lit- es mainly lacustrine sediments intercalated with vol- ter , and rotten wood . Adult elaterids may be caniclastics[ 29] .The exact age of this fo rmation is still saprophagous , phy tophagous , or predacious, but all contentious .Three different opinions about the age appear to be liquid feeders practicing extraoral diges- (Late Jurassic , Late Jurassic-Early Cretaceous and tion[ 5] .Phytophagous soil-dwelling larvae are com- Early Cretaceous)have been proposed based on both monly know n as wirew orms , w hich have harmful biostratigraphic and radiometric geochronolo- [ 5] economical importance to crops and forests .By gy[ 30 —34] .Recently , by comparing the Yixian biota now , this family contains approximately 165 fossil w ith the Solnhofen biota in Germany , the Purbeck species[ 6—25] and 25 amber species[ 26 —28] that are at- biota in England and Late Jurasic Tero ri-type and tributed to some 72 genera reported from all over the Ryoseki-ty pe floras in Japan , Wang et al.considered w orld.Among these , 108 species in 32 genera are that the age of the Yixian Form ation should be from know n from the Upper Jurassic strata of the Late Tithonian to the Berriasian[ 33, 34] .

＊ Supported by National Natural S cience Foundation of China (Grant Nos.30370184 , 30430100), the Beijing Natural S cience Foundation (Grant No .5032003), Beijing Municipal Commission of Education (Grant No .KZ200410018013)and PH R Project of Beijing Municipal Commission of Edu- cation ＊＊ To w hom correspondence should be addressed .E-mail:rendong @mail.cnu .edu .cn Prog ress in Natural Science Vol.17 No .10 2007 www .tandf .co .uk/ journals 1245

1 Material and methods the mandible to the apex of the abdomen .The body w idth w as measured at the base of the ely tra .Both This study w as the observation o n a new ela- length and width of pronotum w ere measured at the terid , its part and counterpart , collected from the 2nd median line . Bed of Yixian Formation in Huangbanjigou[ 35] , near Chaomidian Village , Shangyuan County , Beipiao 2 Systematic paleontology City , Liaoning Province . Order Coleoptera Linneus , 1758 The specimen was ex amined using a Leica Family Elateridae Leach , 1815 M Z12 .5 dissecting microscope and illustrated with the aid of a drawing tube attachment .The terms of Genus Cryptocoleous Dolin and Nel, 2002 mesoventrite and metaventrite have been used in place of the misapplied terms of mesosternum and metaster- Type species .Cryptocoelus m ajor Dolin and num , follow ing Law rence[ 36] , Beutel and H ass[ 37] Nel , 2002 (Fig .1 (a), (b)), sampled near Chao- and Cleide Costa[ 38] . midian Village , Beipiao City , Liaoning Province , China . The body leng th w as measured from the apex of

Fig.1 . Cryptocoelus major Dolin and Nel, 2002 ((a), (b)), and Cryptocoelusbu ffoni Dolin and Nel, 2002 ((c), (d)).(a)and (c) Dorsal view ;(b)and (d)ventral view (Redraw ing from Dolin and Nel[ 11]).

Revised diagnosis:The genus differs from all rial . o ther closely related genera within the same family by the follow ing features:body elongate , posterior angle Cryptocoelus gianteus sp.nov . of prono tum obviously prolonged backw ard , with dis- Etymology:Name derived from Latin “ Gigan- tinct sho rt carina ;antennae short , not reaching pos- teus” , fo r its body is larger than any other fossil ela- terior angle of prono tum , scape robust , pedical much terids that have been previously reported . shorter than scape and antennomere 3 , antennomeres 3 and 4 distinctly elongate ;chin piece moderately o r Holotype : CNU-C-LB2006851-1 , CNU-C- strongly arcuate ;scutellum rounded , semi-oval , o r LB2006851-2 , almost complete part and counterpart subtriangular , never cordate ;ely tra w ith faintly lon- impression of elaterid , housed in Key Lab of Insect gitudinal striae ;mesocoxae open to mesepimeron ; Evolution & Environment Change , Capital Normal metacox al plates obtusely long triangular , evenly nar- University , Beijing , China . rowed laterally ; tarsi w ith five tarsomeres , tarsomeres 1 —4 succeedingly shorter . Horizon and locality :Collected from a site near Chaomidian Village , Beipiao City , Liaoning Remarks :The modified diagnosis of the genus is Province , China ;Upper Jurassic-Low er Cretaceous based on the ty pe species (Fig .1)and the new m ate- Yixian Formation (late Tithonian to Berriasian). 1246 www .tandf .co .uk/ journals Progress in N atural Science V ol.17 No .10 2007

Species diagnosis :Differing from the other two curved, without teeth on inner side ;eyes small , o- species (Fig .1)w ithin the same genus by having a val ;gular sutures parallel ;frons flattened ;labrum longer prosternal process , smaller procoxal cavity , slig htly transverse . different propo rtion of first three segments of antennae , and different proportion of the tarsomeres. Antennae (Fig .3 (c)):short , fail to reach the posterio r angle of prono tum , 11-segmented , serrate Description (Fig .2 (a)):Body quite large , from antennomere 4 ;scape robust , pedicel m uch subcylindrical, smooth , elytra with faintly longitudi- sho rt than scape and antennomere 3 ;antennomere 3 nal striae , w ith short and pointed external ovipositor . to 5 distinctly elongate , 4 approximately 2 .72 times as long as w ide , antennomeres 4 —10 succeedingly Head :oval;mandible robust (Fig .2 (g)), in- narrow er ;antennomere 11 oblong .

Fig.2 . Cryptocoelus gia nteus sp .nov , holotype.(a)Dorsal view ;(b)ventral view ;(c)and (d)hindtibia, dorsal view and ventral view .Ⅰ — Ⅴ , tarsomeres 1 —5 ;(e)pro-mesothorax , 1 prosternal pleural suture;2 procoxae ;3 prosternal process;4 mesoventrite cavity ;5 carinae;(f)6 ovipositor;(g)mandibles. Prog ress in Natural Science Vol.17 No .10 2007 www .tandf .co .uk/ journals 1247

Fig.3 . Cryptocoelus gianteus sp .nov ., holotype .(a)Dorsal view ;(b)ventral view ;(c)antennae, dorsal view ;(d)hindtibia, ventral view ;(e)hindtibia, dorsal view .

Pronotum:smoo th , subsquare , wider than Ventral surface (Fig .3 (b)):chin piece nor- head , anterior margin nearly straight , lateral sides mally arcuate ;prosternal sutures single ;procoxal arcuate in front of hind ang les , basal margin moder- cavity open behind , rounded ;prosternal process ex- ately bisinuate , width equal to leng th if leng th mea- tending far behind cox ae ;mesoventral cavity appar- sured at median line , 1 .14 times as lo ng as w ide if ently open anteriorly , V-shaped ;metaventrite sm all, leng th is measured from anterior angle to posterio r w ithout transverse sutures ; mesocox ae open to ang le disc slig htly convex ;hind angles acute , project- mesepimeron ;mesoventrite and metaventrite separat- ing backw ardly , w ith distinct short carinae (Fig .2 ed by distinct suture ;metaventrite with longitudinal (e)). suture ;metepisternum narrow ;metaco xal plates obtusely long triangular , evenly narrowed laterally ;ab- Scutellum:small, subtriang ular . domen with 5 visible ventrites , superimposed each Elytron:approximately 1 .14 times as wide as others, narrow ed from the base of fifth visible ven- protho rax , 4 times as long as w ide , lateral sides near- trite , first visible ventrite much longer than others, ly parallel at middle , flattened on disc , w ith 5 faintly 1 .23 times as long as the previous one ;ovipositor visible striae , intervals flat , epipleural rim narrow , conifo rm (Fig .2 (f)). w ith rounded humeral angle , apex of ely tra slightly Legs:tibia and tarsi setosed ;procox a rounded ; obtuse , not completely cover the last visible ventrite protibiae more o r less straight , slightly swollen api- (Fig .3 (a)). 1248 www .tandf .co .uk/ journals Progress in N atural Science V ol.17 No .10 2007 cally , w ith one short spur at tip ;mesocox a rounded , ly , in a w idely accepted taxonomic framew ork of the big ger than procoxa , meso trochanter oval ;metacox a Elateridae by Stibick , the subfamily Agrypninae is transverse , metatrochanter oval, metatarsi with 5 degraded to a tribe status—Ag rypnini , subo rdinate to [ 1] tarsomeres , tarsomere 1 much longer than others , Py rophorinae .Acco rding to this system , the genus tarsomeres 1 —4 succeeding ly sho rter , the fourth one in question has a closer relationship with the Py- is the sho rtest , subcordate ;claw s simple , falciform rophorinae as indicated by the following characteris- (Fig .3 (d), (e)). tics :chin piece normally arcuate ;prosternal process no rmally elongate ;mesocox ae open to mesepimeron ; Measurement (mm ):body length 27 ;body scutellum never co rdate . w idth 8 ;elytron length 16 . (4)We also do not support Dolin and Nel' s as- 3 Discussion and conclusions sig nment of Cryptocoelus to the fossil tribe Crypto- (1)The genitalia of beetles is largely internal in cardiini[ 10 , 11] .Although this genus has high resem- both sexes , and the ovipositor is highly reduced as a blance to Cryptocardius (another genus of Crypto- result .This feature is probably related to the general cardiini know n from Karatau[ 10, 11] ), as the scutellum body structure of beetles because the delicate genitalia of Cryptocoelus never cordate , this feature contra- can be protected internally w hen the beetles w edge dicts the tribe diag nostic feature of Cry ptocardiini into tight space or in soil[ 4] .But the new species and Dolin , 1980 ;thus , it seems inconsequent to attribute Cry ptocoelus buf foni Dolin and Nel , 2002 obviously this genus to Cryptocardiini as suggested by Dolin and preserved with short , pointed , ex ternal oviposito r , so Nel[ 11] . w e may deduce that their oviposito r m ay be more o r less sclerotized , and during their lifetime they should (5)We still canno t assign this genus to any oth- lay their egg s into the soil just like the ex tant species er know n subfamily or tribe due to the absence of the do , fo r the larvae of most ex tant species live under- characteristics currently used to diag nose the fami- [ 1] g round[ 2 , 3] . ly , for exam ple , the position of the mouthparts and whether w ith basal setae on claw s or no t .It is (2)One distinct feature of the new species is its still inadvisable to set a new subfamily or tribe just by possession of robust mandibles, different from extant one single genus, so we temporarily make its subfam- elaterids , but similar to the beetles of the Jehol ento- ily and tribe status uncertain , and look forward to [ 39] mofauna , cupedids .Now adays , cupedids are relict more exquisite-preserved fossil elaterids of this genus beetles , with cry ptic habits , but they are one group to be collected and studied in the future . of primitive beetles .From the Triassic to the Early Cretaceous, cupedids were quite diverse , but in the Acknowledgements T he authors thank D r.Jakub P ro kop (Department of P alaeonto logy , Charles University , Czech Re- M id to Late Cretaceous the diversity of the group de- public), Prof .Jiang Shihong (Applied Biological Engineering creased .Some authors deduced that the extinction of Department, Shenzhen Poly technic), and edito r of V estinik some Cupedomorpha might have been caused by com- Zoologii from U kraine , for providing us with the literature ma- petition w ith modern w ood-eating forms , such as Ela- terials that are indispensable to this research.W e are particular- [ 40 , 41] teridae or Buprestidae , especially elaterids, fo r ly g rateful to the colleagues of our lab fo r their detailed com- they have obvious advantage in escaping their natural ments and fruitful sug gestions. enemy' s hunting .The new elaterid from the Liaon- References ing beds may further attest to this deduction . 1 Stibike JNL .Classification of the Elateridae. 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