new mayfly nymphs (insecta: ephemeroptera) from yixian ...upper jurassic–lower cretaceous of...

ISSN 0031�0301, Paleontological Journal, 2011, Vol. 45, No. 2, pp. 167–173. © Pleiades Publishing, Ltd., 2011Original Russian Text © Jiandong Huang, N.D. Sinitshenkova, Dong Ren, 2011, published in Paleontologicheskii Zhurnal, 2011, No. 2, pp. 46–51.1

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INTRODUCTION

The Jehol Biota is one of the most important fossilfaunas in northern China; it is dated from the latestJurassic to the earliest Cretaceous. It includes repre�sentatives of almost all major groups of terrestrial andfreshwater vertebrates, a wide variety of invertebrates,and a diverse flora, fossils of which are all well�pre�served. Especially as feathered theropod dinosaurs,early birds, putative basal angiosperms, primitivemammals, and pollinating insects have been discov�ered since the early 1990s, the Jehol Biota has been afocus which is drawing paleontologists' attention allover the world. Although the strata that the Jehol Biotacomprises are still contentious, the Dabeigou and Yix�ian formations and the overlying Jiufotang Formationare unquestionably considered as important parts ofthe Jehol Biota (Hong, 1993; Zhou, 2006).

To date, 16 orders of the insect fauna have beenreported in the Yixian Formation, includingEphemeroptera (Hong, 1982; Lin and Huang, 2001;Huang et al., 2007), Odonata (Zhang et al., 2008),Orthoptera (Ren et al., 1995; Zhang et al., 2008), Ple�coptera (Liu et al., 2007), Blattodea (Wang et al.,2007), Phasmatodea (Ren,1997b), Homoptera (Renet al., 1995), Hemiptera (Yao et al., 2008), Raphid�ioptera (Ren, 1997a), Neuroptera (Ren and Engel,2008), Coleoptera (Liu et al., 2008), Mecoptera (Renand Shih, 2005), Diptera (Ren et al., 1995), Tri�choptera (Ren et al., 1995), and Hymenoptera (Gaoand Ren, 2008; Shih et al., 2009). The fauna containsboth primitive and many advances forms.

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Ephemeroptera is an archaic and somewhat aber�rant insect group; at the same time, it is an importantgroup occurring in lacustrine deposits (Kluge and Sin�itshenkova, 2002). Ping (1928) was the first to recog�nize and describe nymphal specimens of Ephemeropsistrisetalis Eichwald, 1864 (Hexagenitidae) in the JeholBiota of the Yixian Formation. Subsequently, nymphsand adults of this species were redescribed by Hong(1982). As Huang et al. (2007) established the genusEpicharmeropsis and the species E. hexavenulosus andE. quadrivenulosus based on some well�preserved adultspecimens from the same locality, it became evidentthat the genus Ephemeropsis is absent in this locality.Another species, Caenephemera shangyuanensis Lin etHuang of the family Hexagenitidae was describedbased on a single nymphal specimen from the YixianFormation (Lin and Huang, 2001). All of the speciesreported are large�sized, with the body at least 25 mmlong.

Recently, ten relatively well�preserved mayflynymph specimens from the Yixian Formation of theHuangbanjigou locality (near the village of Chaomid�ian, Shangyuan Township, Beipiao City, LiaoningProvince, China) were discovered. All of them aresmall�sized, with the body less than 16 mm long, andbelong to three new species. The new mayflies comefrom the same beds as hexagenitids of the genus Epic�harmeropsis. This is the first case of co�occurrence oflarge hexagenitids and other mayflies. In Siberia andMongolia, large hexagenitids (Ephemeropsis, Mongol�ogenites Sinitshenkova, 1986) usually form assem�blages of only one species. The Huangbanjigou local�

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New Mayfly Nymphs (Insecta: Ephemeroptera) from Yixian Formation, China

Jiandong Huanga, c, N. D. Sinitshenkovab, and Dong RencaAnhui Paleontological Museum, Anhui Province, 230601 China

bBorissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russiae�mail: nina_sin@ mail.ru

cCollege of Life Science, Capital Normal University, Beijing, 100048 Chinae�mail: [email protected]

Received May 20, 2009

Abstract—Three new species of mayfly nymphs, Clavineta excavata sp. nov., Clavineta brevinodia sp. nov.,and Siberiogenites branchicillus sp. nov., from the Yixian Formation (Late Jurassic–Early Cretaceous) inwestern Liaoning Province, China are described. These mature nymphs are small�sized, with the body lessthan 16.0 mm long. The systematic position of the genera Clavineta and Siberiogenites, which are recordedfor the first time in the Yixian Formation, is briefly discussed.

Keywords: Ephemeroptera, nymphs, new species, Yixian Formation, China.

DOI: 10.1134/S0031030111020092

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JIANDONG HUANG et al.1

ity is dominated by nymphs of Epicharmeropsis, whileClavineta and Siberiogenites are much less frequent.

The genus Clavineta Sinitshenkova, 1991 was orig�inally assigned to the family Leptophlebiidae Banks,1900 and considered to be closely similar to FurvonetaSinitshenkova 1989 in the structure of tergaliae (Sinit�shenkova, 1991). Kluge (1993) believed that Clavinetaand Mesoneta Brauer, Redtenbacher et Ganglbauer,1898 are not related to Leptophlebiidae and should bytransferred to the superfamily Siphlonuroidea. Later,based on structural similarity in tergaliae, Sinitshenk�ova (2000) included Clavineta and Furvoneta in thefamily Mesonetidae Tshernova, 1969 (see also Klugeand Sinitshenkova, 2002). At present, Kluge (2004)refers Clavineta to Eusetisura incertae sedis.

To date, all known species of the genus Clavinetahave been described based on nymphs. The presenceof long dense hairs on the inner side of the fore femoraand tibiae, which are positioned close to the head, istheir distinctive morphological feature. Unfortunately,this feature is poorly preserved in C. excavate sp. nov.and C. brevinodia sp. nov. Other characters of theClavineta, such as the position of forelegs near thehead and the presence of ribs on the anterior and pos�terior margins of the tergaliae are well�pronounced.Based on this, we tentatively assign these two new spe�cies to Clavineta until new better preserved and morecomplete material is available.

In China, only one species, Clavineta eximiaZhang, 2006, has been recorded in the JiulongshanFormation of Hebei Province, which is dated MiddleJurassic (Zhang, 2006).

The genus Siberiogenites is reported in China forthe first time. Since each tergalia of Siberiogenites hasa rib on the costal margin and a strong anal rib at a dis�tance from the anal margin, this genus was assigned tothe family Hexagenitidae (Sinitshenkova, 1985).

Kluge (2004) believed that the presence of a longlast tergalia is a unique apomorphy of Hexagenitidae.Siberiogenites is the only genus of this family the sev�enth tergalia of which is not longer than others. Basedon this, Kluge (2004) referred it to as Euplectopteraincertae sedis. In actuality, a larger seventh tergalia isobserved not only in Hexagenitidae but also in oneEarly Cretaceous Australian genus, Promirara Jell etDuncan, 1986, which was initially assigned to the fam�ily Siphlonuridae (Jell and Duncan, 1986).

The Chinese mayfly nymph Shantous Zhang etKluge, 2007 (Zhang and Kluge, 2007) also has anenlarged seventh tergalia, with a well�developedmedian rib. In our opinion, Shantous was mistakenlyreferred to Hexagenitidae, since the structure of itsbody, legs, and tergalia suggest that it is typical mem�ber of Epeoromimidae Tshernova, 1969, with theenlarged last tergalia compared to the others.

An autapomorphy of Hexagenitidae is the presenceof costal and anal ribs on the tergaliae. Thus, theJurassic genus Siberiogenites undoubtedly belongs to

Hexagenitidae, although it is more primitive thanother genera of this family, because its last tergalia isnot longer than others.

At the end of Jurassic and, particularly, in the EarlyCretaceous, different aquatic insect groups, such as bee�tles (Coptoclavidae), dragonflies (Hemeroscopidae),stoneflies (Platyperlidae), and mayflies (Hexagenitidae,Siphlonuridae, Epeoromimidae) developed adaptationsfor swimming in the water column. The larger last terga�lia developing in mayfly nymphs of different familiescould serve as an oar while swimming (Sinitshenkova,2007).

The specimens were examined using a LeicaMZ12.5 dissecting microscope and illustrated with theaid of a drawing tube attached to the microscope. Linedrawings were produced with CorelDraw 12.0 andPhotoshop CS 10.0. Morphological terms used herefollow those by Kluge (2004). The holotypes andparatypes examined are housed in the Key Lab ofInsect Evolution and Environmental Change, Collegeof Life Sciences, Capital Normal University, Beijing,China.

SYSTEMATIC PALEONTOLOGY

Order Ephemerida (=Ephemeroptera)

Family Mesonetidae Tshernova, 1969

Genus Clavineta Sinitshenkova, 1991

Ty p e s p e c i e s. C. cantabilis Sinitshenkova, 1991,Upper Jurassic–Lower Cretaceous of Mongolia.

S p e c i e s c o m p o s i t i o n. In addition to thetype species, C. transbaikalia Sinitshenkova, 2000 andC. citima Sinitshenkova, 2000 from the Upper Juras�sic–Lower Cretaceous of the Chernovskie Kopi local�ity, Transbaikalia; C eximia Zhang, 2006 from theMiddle Jurassic Jiulongshan Formation of northeast�ern China; and two new species from the Yixian For�mation.

Clavineta excavata Huang, Sinitshenkova et Ren, sp. nov.

Plate 8, figs. 1 and 2

E t y m o l o g y. From the Latin excavatus(incised), because of the concave anterior and poste�rior margins of the pronotum.

H o l o t y p e. CNU�EPH�LN�08001, well�pre�served impression of a nymphal exuviation withoutlegs; Huangbanjigou locality; Upper Jurassic–LowerCretaceous, Yixian Formation.

D e s c r i p t i o n (Figs. 1a, 1b). Mature nymph.The head is relatively small and short, wider than long.Eyes are small, subovate; the frons has a pair of longi�tudinal ridges between the antennae and eyes. Theclypeus and labrum are relatively large. The maxilla ispoorly preserved, only two dentisetae and three�seg�mented maxillary palp are visible; other parts of thehead are not preserved. The pronotum is short, morethan five times as wide as long, with an obviously con�

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NEW MAYFLY NYMPHS (INSECTA: EPHEMEROPTERA) 169

Plate 8

1 3 4

2

E x p l a n a t i o n o f P l a t e 8

Figs. 1 and 2. Clavineta excavata sp. nov.: (1) holotype CNU�EPH�LN�08001; (2) paratype CNU�EPH�LN�08002.Fig. 3. Clavineta brevinodia sp. nov., holotype CNU�EPH�LN�08008.Fig. 4. Siberiogenites branchicillus sp. nov., holotype CNU�EPH�LN�08009.Upper Jurassic–Lower Cretaceous, Yixian Formation, China. Scale bar, 4 mm.

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cave anterior margin and slightly concave posteriormargin. The pterothorax is well developed; the ante�rior pair of wing buds are large, almost triangular, withtwo visible longitudinal veins; their apices reach theposterior margin of the second abdominal segment.The posterior pair of wing buds are completely coveredby anterior ones. The forelegs are positioned close tothe head; the tibia and tarsus are extended forward; thefemur is thickened; the tibia is cylindrical, longer thanthe femur; the tarsus is slightly shorter and thinnerthan the tibia. Hairs on the foreleg are not preserved.The middle legs are similar to the forelegs. The abdo�men is almost three times as long as wide. Each tergumof abdominal segments 1–5 is three times as wide aslong, with the W�shaped anterior margin; the poste�rior margin is straight. Succeeding segments graduallydecrease in size. The posterolateral projections onabdominal segments 1–9 are prominent and extendeddistinctly posteriorly. The tergaliae are well developedon abdominal segment 1–7, about one�third as long asthe width of the abdominal terga; the sixth and seventhpairs of the tergaliae are almost as long as the others;the costal ribs are all longer than anal ribs. The caudaliiare comparatively thin; the paracercus and inner mar�gins of the cerci are covered with long and dense swim�ming hairs at a distance from their base.

M e a s u r e m e n t s (mm). Holotype: body lengthof nymph, 12.6; body width, 3.0; thorax length, 2.8;abdominal length, 8.8; cercus length, as preserved,3.2; paracercus length, as preserved, 2.8.

Comparison. The new species is similar to Clavin�eta eximia Zhang, 2006 from the Middle Jurassic of theJiulongshan Formation of China and differs in theshape of the pronotum, the anterior pair of wing budsand tergaliae. It clearly differs from other congeners inthe shorter legs, the concave pronotum, and distinctposterolateral denticles n the abdominal segments.

M a t e r i a l. In addition to the holotype, the samelocality has yielded CNU�EPH�LN�08002, exuviationof nymph; CNU�EPH�LN�08003�1 and CNU�EPH�LN�08003�2, part and counterpart of exuviation ofnymph (Fig. 1b; Pl. 8, fig. 2); CNU�EPH�LN�08004�1and CNU�EPH�LN�08004�2, part and counterpart ofmature nymph; CNU�EPH�LN�08005�1 and CNU�EPH�LN�08005�2, CNU�EPH�LN�08006�1 andCNU�EPH�LN�08006�2, CNU�EPH�LN�08007,mature nymphs.

Clavineta brevinodia Huang, Sinitshenkova et Ren, sp. nov.

Plate 8, fig. 3

E t y m o l o g y. From the Latin brevinodius (with ashort abdomen).

H o l o t y p e. CNU�EPH�LN�08008, impressionof a well�preserved nymphal exuviation without head,pronotum, and legs; Huangbanjigou locality; UpperJurassic–Lower Cretaceous, Yixian Formation.

D e s c r i p t i o n (Fig. 1c). Mature nymph. Thesternites of the pterothorax are fused with the sterniteof the first abdominal segment. Abdominal segmentsare wide and short; segments 1–5 are almost 3.8 times

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(a) (b) (c)

Fig. 1. Mayfly nymphs of the genus Clavineta, exuviations: (a, b) C. excavata sp. nov., nymph: (a) holotype CNU�EPH�LN�08001and (b) paratype CNU�EPH�LN�08002; (c) C. brevinodia sp. nov., holotype CNU�EPH�LN�08008. Upper Jurassic–LowerCretaceous, Yixian Formation; China. Scale bar, 2 mm.



as wide as long; the anterior and posterior margins onsternites are straight, while the margins on the tergitesare W�shaped. The posterolateral projections onabdominal segments 1–7 are prominent and extenddistinctly posteriorly, while the posterolateral projec�tions of abdominal segments 8 and 9 are indiscernible.The wing buds are short, do not reach the posteriormargin of the second abdominal segment.

M e a s u r e m e n t s (mm). Body length of pre�served nymph fragment, 10.8, total length, about 13.0;length and width of abdominal segment 3, 0.8 and 3.0,respectively.

C o m p a r i s o n. C. brevinodia sp. nov. clearly dif�fers from all known congeners in the wider and shorterabdominal segments.

R e m a r k s. The wide and short abdominal seg�ments are characteristic of Mesoneta; however, the ter�galiae of this genus lack thickened ribs on the posteriormargin. The new species is similar in the structure oftergaliae to Furvoneta Sinitshenkova, 1989, but in thelatter, posterolateral denticles on the ninth tergite arelong, equal in length to the last tergite. Therefore,although legs in the position characteristic of Clavinetaare not preserved, available features allow the new spe�cies to be assigned to this genus.

M a t e r i a l. Holotype.

Family Hexagenitidae Lameere, 1917

Genus Siberiogenites Sinitshenkova, 1985

Ty p e s p e c i e s. S. angustatus Sinitshenkova,1985; Lower–Middle Jurassic, Ichetuiskaya Forma�tion of western Transbaikalia.

S p e c i e s c o m p o s i t i o n. In addition to thetype species, S. rotundatus Sinitshenkova, 1985 fromthe Lower–Middle Jurassic (Zhargalan Formation) ofwestern Mongolia (Oshin�Boro�Udzur�Ula locality);S. medius Sinitshenkova, 1989 from the Lower Creta�ceous (Tsagantsab Formation) of Mongolia (HaraHutul locality); S. recticostalis Sinitshenkova, 2000from the Upper Jurassic–Lower Cretaceous of theChernovskaya strata of Transbaikalia (ChernovskieKopi locality); S. mongolicus Sinitshenkova, 2002from the Upper Jurassic of Mongolia (Shar Teeg local�ity) (Sinitshenkova, 2002); and S. branchicillus sp. nov.described below.

Siberiogenites branchicillus Huang, Sinitshenkova et Ren, sp. nov.

Plate 8, fig. 4

E t y m o l o g y. From the Greek branchia (gills)and cillus (donkey).

H o l o t y p e. CNU�EPH�LN�08009, part andcounterpart of well�preserved abdominal segments ofa nymph, with tergaliae, caudalii, and a fragment ofwing buds; Huangbanjigou locality; Upper Jurassic–Lower Cretaceous, Yixian Formation.

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D e s c r i p t i o n (Fig. 2). Mature nymph. Thehead and prothorax are not preserved. The anteriorpair of wing buds are large, elongated ovate, andclosely approach the posterior margin of the secondabdominal segment; the posterior pair of wing buds arecompletely covered by anterior ones. The abdomen isalmost three times as long as wide; the first fourabdominal segments are similar in shape, each is about2.8 times as wide as long, with a W�shaped anteriormargin and straight posterior margin. Remaining tergataper gradually terminally; the posterior margin of thetenth tergum is more or less rounded. The posterolat�eral projections on abdominal segments 1–9 areprominent and extend clearly posteriorly. The terga�liae are well developed on abdominal segments 1–7;all tergaliae are single, widely ovate, with a weakrounded tip; tergaliae 1–6 are relatively large, morethan half as long as the width of the abdominal terga.The seventh tergalia is the smallest, less than half aslong as the width of the seventh segment. Each tergaliahas a weak costal rib on the costal margin and a stronganal rib at a distance from the anal margin. The cerciand paracercus are subequal in length; the paracercusand inner margins of the cerci are covered with long anddense swimming hairs at a distance from their base.

Fig. 2. Siberiogenites branchicillus sp. nov., holotypeCNU�EPH�LN�08009, fragment of nymph. UpperJurassic–Lower Cretaceous, Yixian Formation, China.Scale bar, 2 mm.

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M e a s u r e m e n t s in mm. Body length of pre�served nymph fragment, 12.3; total length was about15; length of abdominal segments 1–4, 0.9; theirwidth, 2.5.

C o m p a r i s o n. S. branchicillus sp. nov. differssignificantly from all known species of Siberiogenites inthe weak rib on the costal margin of the tergaliae.

M a t e r i a l. Holotype.

ACKNOWLEDGMENTS

We are grateful to Dr. Shih Chung�kun (College ofLife Sciences, Capital Normal University, Beijing,China) for valuable remarks.

This study was supported by the National NatureScience Foundation of China (projectnos. 408720022, 30811120038), the Nature ScienceFoundation of Beijing (project no. 5082002), Scien�tific Research Key Program and PHR Project ofBeijing Municipal Commission of Education, and theProgram of the Presidium of the Russian Academy ofSciences “Origin and Evolution of the Biosphere.”

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