Elsevier

Cretaceous Research

Volume 34, April 2012, Pages 334-339
Cretaceous Research

Prosinodendron krelli from the Yixian Formation, China: A missing link among Lucanidae, Diphyllostomatidae and Passalidae (Coleoptera: Scarabaeoidea)

https://doi.org/10.1016/j.cretres.2011.11.017Get rights and content

Abstract

Prosinodendron krelli Bai, Ren and Yang gen. et sp. nov. (Scarabaeoidea: Lucanidae), an exceptionally well-preserved, new lucanid fossil, is described and illustrated from the Yixian Formation of western Liaoning Province, northeastern China. Based on its unique combination of characters, it is considered to be a missing link among Lucanidae, Diphyllostomatidae and Passalidae. A key to known fossil species of Lucanidae and adjacent families from Mesozoic is given. The close phylogenetic relationship of these three families of the Scarabaeoidea is supported by the fossil evidence.

Introduction

The scarab beetles (Scarabaeoidea or Lamellicornia) are a cosmopolitan monophyletic group of Coleoptera: Polyphaga, comprising about 28,000 described extant species, which are well known for their exceptionally large mandibles (Browne and Scholtz, 1995, Lawrence et al., 2011). The family Lucanidae has long been considered to be one of the most primitive groups in the Scarabaeoidea (Ritcher, 1966, Crowson, 1967, Howden, 1982), and scarabaeoid classifications and evolutionary hypotheses have generally regarded the Lucanidae as basal to all scarabaeoids (Iablokov-Khnzorian, 1977, Howden, 1982, Lawrence and Newton, 1995). However, based on comparison of “primitive” scarabaeoid groups, Scholtz et al. (1994) hypothesized that the family Glaresidae, rather than the Lucanidae, is the most primitive. According to this hypothesis, the Lucanidae is a member of a clade that includes the Passalidae and Diphyllostomatidae.

Prior to the taxonomic elevation of the genus Diphyllostoma to the family Diphyllostomatidae (Holloway, 1972), the Lucanidae was considered as most closely related to the Passalidae (Howden, 1982). Based on shared characters, it is now thought that the Lucanidae is most closely related to the Diphyllostomatidae (Caveney, 1986, Browne and Scholtz, 1995). However these hypotheses have not been supported by the fossil evidence.

So far, 266 fossil scarabaeoid species have been described, of which 231 probably or reliably belong to Scarabaeoidea, 31 are considered doubtful owing to a lack of key characters, and nine belong to other taxa (Krell, 2000, Krell, 2007, Nikolajev, 2007, Bai et al., 2010, Bai et al., 2011). The earliest lucanid species, Juraesalus atavus Nikolajev et al., 2011, was found in Middle Jurassic deposits in Daohugou Village, Chifeng City, Inner Mongolia, China. Hitherto, 20 fossil lucanid species and four passalids have been reported, but no fossil diphyllostomatids have been found and none has been claimed as the missing link of these three families. Prosinodendron krelli Bai, Ren and Yang, gen. et sp. nov., an exceptionally well-preserved scarabaeoid fossil, which is described here, was collected from the Yixian Formation, near Chaomidian Village, Beipiao City, Liaoning Province, China. Based on its unique combination of characters, it is regarded as a missing link among Lucanidae, Diphyllostomatidae and Passalidae. A key to known fossil species from Lucanidae and adjacent families from Mesozoic is given herein.

Section snippets

Geological background

Numerous researchers have discussed the age of the Yixian Formation in light of the biostratigraphical correlations and the radiometric dates (Chen, 1988, Ren et al., 1997, Chen et al., 1998, Wang, 1998, Swisher et al., 1999, Zhou et al., 2003). However it is now generally accepted that it is the Early Cretaceous in age and most likely to be restricted to 129.7–122.1 Ma: Barremian–early Aptian (Yang et al., 2007, Zhu et al., 2007, Chang et al., 2009, Zhou and Wang, 2010).

The strata of Yixian

Material and methods

This study is based on one specimen, registration no. CNU-COL-LB2010607, housed in the fossil insect collection of the Key Lab of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing (CNUB). It was examined using a LEICA MZ 12.5 dissecting microscope and illustrated with the aid of a drawing tube attachment.

Systematic Paleontology

  • Order Coleoptera Linnaeus, 1758

  • Superfamily Scarabaeoidea Latreille, 1802

  • Family Lucanidae Latreille, 1804

  • Genus

Discussion

Key characters among Prosinodendron (Lucanidae), Sinodendron (Lucanidae), Diphyllostomatidae and Passalidae are compared in Table 1 and Fig. 3 and supports the interpretation that Prosinodendron krelli is a missing link among Lucanidae, Diphyllostomatidae and Passalidae, the close phylogenetic relationship of these three scarabaeoid families being supported by this fossil evidence.

Most species of Lucanidae are characterized by pronounced sexual dimorphism owing to the increased development of

Acknowledgments

We are grateful to Dr. Shih Chungkun (College of Life Sciences, CNUB) for improvement of our manuscript. This research was supported by the National Basic Research Program of China (973 Program) (Nos. 2011CB302102, 2012CB821906), the National Natural Science Foundation of China (Nos. 30900144, 31010103913, 31172143, 40872022, 31071964), the Knowledge Innovation Program of Chinese Academy of Sciences (Nos. KSCX2-EW-G-4 and KSCX2-EW-Z-8) and the China Geological Survey (1212011120116).

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