Prosinodendron krelli from the Yixian Formation, China: A missing link among Lucanidae, Diphyllostomatidae and Passalidae (Coleoptera: Scarabaeoidea)
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.
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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).
References (39)
- et al.
High-precision 40Ar/39Ar age for the Jehol Biota
Palaeogeography, Palaeoclimatology, Palaeoecology
(2009) - et al.
Preliminary analysis of the depositional environments of the Lower Cretaceous Yixian Formation in the Sihetun area, western Liaoning, China
Cretaceous Research
(2007) - et al.
Early Cretaceous volcanism and its impact on fossil preservation in Western Liaoning, NE China
Palaeogeography, Palaeoclimatology, Palaeoecology
(2011) - et al.
New evidence for Cretaceous age of the feathered dinosaurs of Liaoning: Zircon U–Pb SHRIMP dating of the Yixian Formation in Sihetun, northeast China
Cretaceous Research
(2007) - et al.
Palaeomagnetic and 40Ar/39Ar dating constraints on the age of the Jehol Biota and the duration of deposition of the Sihetun fossil-bearing lake sediments, northeast China
Cretaceous Research
(2007) - et al.
Oxygen isotopes of East Asian dinosaurs reveal exceptionally cold Early Cretaceous climate
Proceedings of the National Academy of Sciences of the United States of America
(2011) - et al.
A new, well-preserved species of Glaresidae (Coleoptera: Scarabaeoidea) from the Jehol Biota of China
Acta Geologica Sinica (English Edition)
(2010) - et al.
Prophaenognatha, a new aclopines genus from the Yixian Formation, China and its phylogenetic position based on morphological characters (Coleoptera: Scarabaeidae)
Acta Geologica Sinica (English Edition)
(2011) - et al.
Phylogeny of the families of Scarabaeoidea (Coleoptera) based on characters of the hind wing articulation, hind wing base and wing venation
Systematic Entomology
(1995) The phylogenetic significance of ommatidium structure in the compound eyes of polyphagan beetles
Canadian Journal of Zoology
(1986)
New fossil beetles of the family Elateridae from the Jehol biota of China (Coleoptera: Polyphaga)
Acta Geologica Sinica (English Edition)
Distribution and migration of the Jehol Fauna with reference to non-marine Jurassic-Cretaceous boundary in China
Acta Palaeontologica Sinica
An exceptionally well-preserved theropod dinosaur from the Yixian Formation of China
Nature
The Natural Classification of the Families of Coleoptera
The stratigraphic sequence and fossil bearing horizon of the Yixian Formation in western Liaoning, China
Geology and Resources
The systematic position of the genus Diphyllostoma Fall (Coleoptera: Scarabaeoidea)
New Zealand Journal of Science
Phylogeny of Japanese stag beetles (Coleoptera: Lucanidae) inferred from 16S mtrRNA gene sequences, with reference to the evolution of sexual dimorphism of mandibles
Zoological Science
A diapsid skull in a new species of the primitive bird Confuciusornis
Nature
Larval and adult characters of Frickius germain, its relationship to the Geotrupini, and a phylogeny of some major taxa in the Scarabaeoidea (Insecta: Coleoptera)
Canadian Journal of Zoology
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2016, Cretaceous ResearchCitation Excerpt :The study is based on four specimens (including two with parts and counterparts) collected near Liutiaogou Village, Ningcheng County, Chifeng City, from Yixian Formation of Inner Mongolia, China (Fig 1). Yixian Formation, belonging to the famous Jehol Biota (Ren et al., 1997, 2010, 2012), has yielded a number of fossil insects, such as Odonata (Zhang et al., 2006), Chresmodidae (Zhang et al., 2008), Coleoptera (Bai et al., 2012, 2013; Yan et al., 2012a,b; Yu et al., 2013, 2015a,b), Hemiptera (Chen et al., 2014; Yao et al., 2014), Neuroptera (Shi et al., 2015), Hymenoptera (Li et al., 2014; Wang et al., 2015) and Pseudopulicidae (Gao et al., 2012), which provided new information to enhance our understanding of the taxonomy and evolution of the respective groups. The age of Yixian Formation is regarded as Early Cretaceous (latest Barremian to earliest Aptian) and linked to a radioisotope-dating of ca. 125 Ma. (
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2015, Molecular Phylogenetics and EvolutionCitation Excerpt :Hence, we considered this fossil as the oldest crown group lucanid (i.e., Lucanidae sensu stricto; node 1). In addition, the fossil of Prosinodendron krelli Bai et al. was described from the Yixian Formation of Inner Mongolia, China (Bai et al., 2012), which has been dated to 123.9 ± 1.1 MYA of the Lower Cretaceous (Yang et al., 2007). Based on the description and illustrations of the fossil, we were able to associate Prosinodendron with modern lucanids excluding Aesalus, and used it to calibrate the node shared by the Syndesinae, Lampriminae, Lucaninae, and an aesaline genus Nicagus (node 2).