Research 75 (2017) 173e178

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Cretaceous Research

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Short communication Qitianniu zhihaoi gen. et sp. nov.: The first cerambycid found in Cretaceous (Coleoptera: )

* Mei-Ying Lin, Ming Bai

Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beichen West Road, Chaoyang Dist., Beijing, 100101, China article info abstract

Article history: A new cerambycid beetle (Qitianniu zhihaoi gen. et. sp. nov.) is described on the basis of a single specimen Received 20 December 2016 embedded in Cretaceous Burmese amber (ca. 99 Ma). Unusual characteristics are hairy antennae and Accepted in revised form 31 March 2017 large lateral eyes, a pronotum with lateral margin, and sinuate protibiae. Based on a phylogenetic Available online 4 April 2017 analysis, the systematic position of Qitianniu is still uncertain and we provisionally place it as Ceram- bycidae incertae sedis. Keywords: © 2017 Elsevier Ltd. All rights reserved. Cerambycidae Qitianniu Burmese amber New

1. Introduction Ma (earliest ) (Shi et al., 2012). The mining locality was at Noije Bum, near Tanai Village (2621033.4100N, 9643011.8800E) Longicorn , including Cerambycidae, Disteniidae, Ves- (Grimaldi et al., 2002; Cruickshank and Ko, 2003, fig. 2; Jałoszynski peridae and Oxypeltidae, are rare in Mesozoic deposits, although et al., 2017, fig. 1), where many important materials were found some taxa have been described as Chrysomeloidea (Yu et al., 2015). including vertebrates and invertebrates (Boucher et al., 2016; To date, only two fossils have been reported, both from the Creta- Oliveira et al., 2016; Xing et al., 2016a, 2016b; Bai et al., 2016a, ceous. One is Cretoprionus liutiaogouensis, from the Lower Creta- 2016b, 2017). These deposits were investigated and dated in detail ceous (ca. 124e122.5 Ma) Yixian Formation of Inner Mongolia, by Cruickshank and Ko (2003) and by Shi et al. (2012). We tentatively China. Its placement in the subfamily is relatively well followed the age (98.8 ± 0.6 Ma) determined by UePb dating of supported (Wang et al., 2014). The second is Sinopraecipuus bilo- zircons from the volcaniclastic matrix of the amber (Shi et al., 2012). batus, from the Lower Cretaceous (approximately 122 Ma) lacus- The type specimen is currently housed in and will eventually be trine deposits of the Yixian Formation in western Liaoning, China, deposited at the Institute of Zoology, Chinese Academy of Sciences whose familial affiliation is less clear but has been treated as (IZAS), the curatorial museum number is IZAS-COL-CERA20160001, Cerambycidae incertae sedis (Yu et al., 2015). Until now, however, no and the specimen will be available for study upon request to cur- Cretaceous Burmese amber species of Cerambycidae have been rent owner, Mr. Zhi-Hao Qi ([email protected]), and the described. In this study, we describe a new extinct beetle species of curators, Dr. Mei-Ying Lin ([email protected]) or Dr. Ming Bai Cerambycidae incertae sedis from the Cretaceous Burmese amber ([email protected]), from Institute of Zoology, Chinese Academy of deposits in the Hukawng Valley of Myanmar. Sciences. The piece of amber containing the specimens was ground and 2. Material and methods polished and examined with a LEICA MZ 12.5 dissecting microscope with a drawing tube attachment. Specimen macrophotography was 2.1. Material and photography conducted using a Visionary Digital photography station, consisting of a Canon EOS 5D DSLR camera equipped with a Canon MP-E A single specimen was found in amber deposits from the 65 mm Macro Photo Lens and tube extensions. An extended Hukawng Valley of Myanmar. The age has been estimated to be ca. 99 depth of field at high magnifications was achieved by combining multiple differently focused images using Helicon Focus 5.1 (Heli- con Soft, Kharkov, Ukraine) software. Final figures were prepared in * Corresponding author. Adobe Photoshop CS5. E-mail address: [email protected] (M. Bai). http://dx.doi.org/10.1016/j.cretres.2017.03.030 0195-6671/© 2017 Elsevier Ltd. All rights reserved. 174 M.-Y. Lin, M. Bai / Cretaceous Research 75 (2017) 173e178

2.2. Micro-CT scanning and 3D reconstruction assigned as the outgroup according to three assumptions: a) Chrysomelidae is a distinct family and Chrysomelinae is a mono- The amber specimen was scanned with a MicroXCT 400 (Carl phyletic subfamily (Reid, 2014); b) it is related to the cerambycoid Zeiss X-ray Microscopy Inc., Pleasanton, USA) at the Institute of assemblage but not too closely related to as Megalopodidae or Zoology, Chinese Academy of Sciences. The scan of the entire beetle Orsodacnidae (Haddad & Mckenna, 2016); and c) Qitianniu is not an (Fig. 1aeb and 1feg) was performed with a beam energy of 60 kV, ingroup of Chrysomelidae. 133 mA, absorption contrast and a spatial resolution of 4.9215 mm. In this study, 51 characters (from the head, mouthparts, pro- The detail scans of the head (Fig. 1ced) and the metatarsi (Fig. 1e) notum, thorax, and legs) were selected (Supplementary Method S2, were performed with 60 kV, 133 mA, absorption contrast, and from Table II of Napp, 1994). The missing data from Napp (1994) spatial resolutions of 1.2762 mm and 1.2762 mm, respectively. On the were changed from “9” to “?”. The characters unavailable in the basis of the obtained image stacks, structures of the specimen were fossil (the hind wing and genitalia) were excluded from the matrix. reconstructed and separated with Amira 5.4 (Visage Imaging, San Some mistakes in Napp (1994) were corrected; for example, char- Diego, USA). The subsequent volumerendering and animations acter 5 for both Saphanus and Atimia were corrected from 9 to 3 were performed with VG Studiomax 2.1 (Volume Graphics, Hei- because both have deeply emarginate eyes, and character 54 for delberg, Germany). The final figures were prepared with Photo- Philus was corrected from 1 to 0 because it has two spurs on the  shopCS5 (Adobe, San Jose, USA). metatibia (Svacha & Lawrence, 2014). The final matrix (Supplementary Method S3) was processed in WinClada and parsimony analyses were conducted with NONA (ratchet, 1000 2.3. Taxon sampling, characters and phylogenetic analysis replicates) and TNT (traditional search; 99,999 random seeds, 1000 replicates) (Goloboff et al., 2003, 2008). Bremer support was The main aim of the phylogenetic analysis in this study was to calculated with TNT. clarify the phylogenetic position of Qitianniu zhihaoi gen. et sp. nov. in the cerambycoid assemblage. We extracted data for 20 genera from the matrix of Napp (1994, table III), containing extant repre- 3. Results sentatives of all four families and eleven subfamilies of Ceram- bycoidea. Additionally, we selected , a representative of 3.1. Systematic paleontology the subfamily , which was missing from Napp (1994), and Chrysomela, a representative of the family Chrysomelidae, as Order Coleoptera Linnaeus, 1758 the outgroup. Together with Qitianniu, 23 genera (Supplementary Superfamily Chrysomeloidea Latreille, 1802 Method S1) were selected for the final matrix. Chrysomela was Family Cerambycidae Latreille, 1802

Fig. 1. 3D reconstructions of Qitianniu zhihaoi gen. et sp. nov., Holotype (IZAS-COL-CERA20160001) based on micro-CT data. AeB The habitus, antennae, mouthparts and legs mostly missing (except mesofemora) due to scan limitations. A dorsal view; B lateral view; CeD head, C in ventral view, D in vento-lateral view, showing the large and coarsely facetted eyes; E metatarsi, showing the bilobed tarsomere III and the short tarsomere IV: arrows indicate the separating of tarsomeres IV and V; F ventrites IV & V in ventral view; G abdomen in ventro-lateral view, showing ventrites. Scale bar in yellow ¼ 1.0 mm, scale bar in red ¼ 0.5 mm. M.-Y. Lin, M. Bai / Cretaceous Research 75 (2017) 173e178 175

Qitianniu gen. nov. carina, apex narrowly rounded; surface irregularly and strongly urn:lsid:zoobank.org:act: C987FBDB-4914-4F56-B49C-9FE7D punctate, covered with semi-erect setae. 4D1E710 Legs short; procoxal cavities posteriorly open; mesocoxal cav- ities open laterally to mesepimeron. Femora clavate; tibiae slightly Description. Sex unknown (probably male), body slender and broadened at apex, protibiae slightly grooved before apex, meso- slightly flattened dorsoventrally. Head declined, prognathous; tibiae without groove. Metatibiae slightly longer than metafemora. vertex with median impression; frons shaped like an inverted Tibial spurs 2-2-2. Tarsi (Fig. 1e) short, tarsomeres II and III both trapezoid, finely punctured; antennal tubercles separated and bilobed; metatarsomere I longer than II or III, but shorter than II elevated; labrum transverse; eyes (Figs. 2d, 1d) very large, strongly plus III; tarsomere IV short and hidden (Fig. 1e); tarsomere V longer convex, coarsely facetted, feebly emarginated and placed laterally, than III, claws simple and widely divergent. narrowly separated from the antennal tubercles. Mandibles quite Underside feebly punctured; prosternum before coxae longer long (Fig. 2d), as long as frons. Antennae (Fig. 2a) eleveneseg- than procoxal cavities, prosternal process lower than trochanter; mented, long, last 5 segments surpass elytral apex; all anten- metasternum much longer than abdominal ventrite I, meta- nomeres fringed with moderately long hairs ventrally and with nepisternum 4 times as long as its anterior width, narrowed dense semi-erect short setae on remaining surface; scape (Fig. 2c, apically. Ventrites I to V subequal in length. 2e) without cicatrix, thickening distad, flattened; antennomere II very short, antennomere III longer than scape, antennomere IV Diagnosis. On the basis of the character combination, including tarsi subequal to III, antennomeres V to VIII subequal to each other, 5-5-5 and tarsomere III bilobed, tarsomere IV short and hidden, antennomeres IX and X shorter than VIII and XI. antennal tubercles present, and long antennae with very short pedicels, Qitianniu gen. nov. can be assigned to Cerambycidae with Pronotum (Fig. 1a) slightly wider than long, constricted basally reasonable certainty (see Discussion for detail). and apically, distinctly narrower than elytral base; sides without Etymology. This name is a combination of the family name of the tubercles or spines and with complete lateral pronotal margins amber owner Mr. Qi and the Chinese spelling tianniu (¼天 (Fig. 2d, 2e); posterior edge more or less straight; disc simple, entire 牛 ¼ longicorn beetle or longhorned beetle). Gender masculine. surface feebly and densely punctured and with hairs. Scutellum transverse, semicircular. Type species: Qitianniu zhihaoi sp. nov. (monotypic). Elytra (Figs. 2a, 1a) fully developed, 2.0 times as long as the Qitianniu zhihaoi sp. nov. combined width and 3.3 times as long as the pronotum, widest at Figs. 1e2 prominent humeri, slightly narrowed around basal 1/3, and urn:lsid:zoobank.org:act: A5AD43BB-7AB1-4A6D-9AEB-513B9F strongly narrowed beyond apical 1/3; elytron without lateral 654674

Fig. 2. Qitianniu zhihaoi gen. et sp. nov., Holotype, IZAS-COL-CERA20160001. AeB Habitus photograph. A in dorso-lateral view; B in ventro-lateral view. C head and prothorax, in antero-lateral view, showing the large and lateral eyes, the scape, the long mandible and the protibia and protarsus; D pronotum in lateral view (left side), arrows indicate the lateral margin; E pronotum in lateral view (right side), arrows indicate the lateral margin; F complete view of the amber piece with a ruler showing the size. Scale bar in red ¼ 1.0 mm. 176 M.-Y. Lin, M. Bai / Cretaceous Research 75 (2017) 173e178

Holotype. Sex unknown but likely a male (see discussion for details), a nearly complete specimen, with most appendages still attached except mesotarsus on the left side and proleg on the right side. The piece of amber (Fig. 2f) containing the specimen was ground and polished to 35.0 17.2 5.2 mm (2.31 g in weight). In the same piece of amber, there are some tiny flies, bugs and other . The type specimen is currently housed in and will eventually be deposited at the Institute of Zoology, Chinese Academy of Sciences (IZAS-COL-CERA20160001). Description. Characters of the genus. Probably male, length 4.6 mm, width 1.5 mm, color uniformly dark, without any markings. Pu- bescence ash-grey, longer setae black-brown. Head with large compound eyes, ca. 1.05 mm wide at maximum, 0.38 mm distance between upper eye lobes (in dorsal view), and 0.46 mm between ventral eye lobes in ventral view. Eyes lateral, coarsely facetted and without interfacetal setae. Mandibles (Fig. 2c) as long as frons, narrowed apically, other details difficult to observe; maxillary palp with last segment longest, not tapered apically; last segment of labial palp not tapered apically. Ratio of antennomere lengths: 5:1:6:6:5:5:5:5:4:4:5. Setae are very dense, with ventral setae nearly twice as long as those on the dorsal side. Pronotum slightly wider than long, ca. 0.9 mm long and 1.0 mm maximum width, constricted basally and apically, basal width ca. 0.67 mm, apical width ca. 0.54 mm. Elytra 3.0 mm long, humeral width 1.48 mm narrowing to 1.35 mm at basal third. Abdomen less than half the length of elytra, ventrites subequal in length, apex of ventrite V rounded (Fig. 1f). Legs moderately long, with dense setae all over, Fig. 3. The first of the three most parsimonious trees from the parsimony analysis of pro- and mesofemora subequal in length, metafemora longer and 21 extant species and 1 fossil species of Cerambycidae, Disteniidae, Vesperidae and Oxypeltidae, including one species of Chrysomelinae as an outgroup. Only unam- reaching the base of ventrite V, metatibia longer than pro- and biguous character changes are shown. The black circles indicate nonhomoplasious mesotibia, 1.2 times as long as the metafemur, metatarsus shorter changes; the white circles indicate changes in homoplasious characters. Numbers than metafemur. Tarsomere II cleft to 1/2, III deeply cleft to 5/7 above branches are character numbers; numbers below branches are character (Fig. 1e). states. Etymology. This species is named after Mr. Zhihao Qi, the amber specimen owner. Dorcasominae. Unfortunately, the trees did not provide strong ev- 3.2. Cladistic analysis idence to differentiate the cerambycoid complex or elucidate the position of Qitianniu clearly. For example, the four genera of The maximum parsimony analysis of the 51 morphological Spondylidinae are very widely separated, and the eight subfamilies characters (Supplementary Method S2) yielded three most parsi- of Cerambycidae are not in a monophyletic clade. Similar problems ¼ ¼ ¼ monious trees (tree length 118 steps, CI 0.66, RI 0.81) and 3 have also appeared in previous studies based on morphological or nodes were collapsed in a strict consensus tree. Morphological molecular data (Napp, 1994; Hunt et al., 2007) in which the fi characters were optimized with parsimony on the rst of the four phylogenetic relationships among the lineages of the cerambycoid most parsimonious trees (Fig. 3), showing only unambiguous complex were not well resolved. Some cerambycoid relationships changes. Black circles indicate nonhomoplasious changes, and have been proposed by previous authors (Linsley, 1961 on the basis white circles indicate changes in homoplasious characters. of various structures; Villiers, 1978; Danilevsky, 1979; Nakamura,   Numbers above branches denote character numbers, and those 1981; Napp, 1994; Svacha et al., 1997; Svacha & Lawrence, 2014); below branches denote character states. however, it is beyond the scope of this study to perform a very Strict consensus (Fig. 4) of 3 MPT from the parsimony analysis of broad taxa sampling and character selection to determine the 22 extant species and 1 fossil species from Cerambycidae, Dis- phylogenetic relationships among subfamilies in the cerambycoid teniidae, Vesperidae, Oxypelidae and Chrysomelidae. Bootstrap complex. In this paper, we do not try to analyze the synapomorphic support values from 1000 replications are presented above support for any clades. branches only for those nodes for which support was calculated.

4. Discussion 4.2. Systematic position of Qitianniu

4.1. Phylogeny of cerambycoid complex On the basis of the characters that were able to observe clearly in the fossil specimen, we can conclude that Qitianniu does not Napp (1994) is the only phylogenetic study designed for the belong to Disteniidae, because it lacks the long recumbent setae in cerambycoid complex (Haddad & Mckenna, 2016). Initially, we large sockets in shallow longitudinal grooves on the posterior incorporated part of the data from Napp (1994) with our data from antennal face, and it does not belong to Oxypeltidae or Anoplo- amber to determine the systematic position of the amber. The most dermatinae, as indicated by the densely setose elytra. Of the parsimonious trees produced in our analyses are quite similar to Cerambycidae s. str., it does not belong to Parandrinae, because of Cladograms 2 and 13 of Napp (1994). The additional taxon Apa- the long filiform antennae and broadly expanded tarsomeres, nor tophysis was not included in Lepturinae (Necydalinae included) or is it likely to belong to Necydalinae, because of the pronotum with Cerambycinae clades, thus supporting the subfamily position of lateral margin and complete elytra. Although the long and dense M.-Y. Lin, M. Bai / Cretaceous Research 75 (2017) 173e178 177

homology is dubious) and the moderately thick body (although this characteristic is not reliable).

4.4. The bilobed tarsomeres III & II

Among the three fossils from the Early Cretaceous, tarsomeres II and III are both bilobed (Wang et al., 2014; Yu et al., 2015; this paper); in fact, tarsomere I is also bilobed (Wang et al., 2014). Although in most of extant species, only tarsomere III is bilobed, a bilobed tarsomere II occur in some groups (such as Philus and Spiniphilus of Philinae, Prosopocera and Sternotomis of , Oplatocera and Plectogaster of Cerambycinae, Macrodontia, Eur- ypoda and Aegosoma of Prioninae, Peithona prionoides and Caraphia of Lepturinae, etc.), and a slightly bilobed tarsomere I also occurs in several groups, especially on the fore tarsi (Anacolus sanguineus of Prioninae, Rhagium of Lepturinae, Massicus, Purpuricenus and Pyr- estes of Cerambycinae; Mesechthistatus of Lamiinae). The pseudo- tetramerous padded tarsus might be adapted for walking on  smooth surfaces such as leaves and smooth bark (Svacha & Lawrence, 2014).

4.5. The lateral pronotal margin

The polarity, degree of homoplasy and the phylogenetic signif- icance of the lateral pronotal margin in chrysomeloids is prob- lematic (Reid, 1995). Its reduced and incomplete state in some Prioninae (e.g., many Aegosomatini) and most Philinae indicates Fig. 4. A strict consensus of the three most parsimonious trees from the parsimony that the long and complete lateral margin distant from the procoxal analysis of 22 extant species and one fossil species from Cerambycidae, Disteniidae, Vesperidae, Oxypeltidae, with one species of Chrysomelidae as an outgroup. Bootstrap sockets (as present in anoplodermatines and many prionines) may   support values from 1000 replications are shown above the branches. Bremer support be derived (Svacha & Lawrence, 2014). The lateral pronotal margin values (in 10 steps) are shown below the branches. (or carina) in this fossil is weak and difficult to observe and it is not be sufficient to place it in Prioninae or Philinae, or exclude it from Cerambycinae or Spondylidinae. erect flagellar pubescence can occur in some prionines, we believe Qitianniu does not belong to Prioninae, because the prionine 4.6. The sinuate foretibiae ancestor was probably a larger beetle; extant prionines are rarely below 10 mm in length, and the small forms (some Neotropical Lamiinae protibiae usually have an oblique pubescent groove Anacolini) are strongly derived. Qitianniu probably does not (antennal cleaner) on the inner face and mesotibia (seldom also belong to Lepturinae, because of its short face and lack of a metatibia) sometimes with a similar groove on the outer face; distinctly constricted neck; nor is it likely to belong to Lamiinae, protibiae with similar structure are rare in Cerambycinae (e.g., because of its prognathous head and pronotum with lateral  some Methiini) (Svacha & Lawrence, 2014). In this fossil, the margin. It probably does not belong to Vesperini, which have bare apically broadened/modified fore tibia could neither place it in elytra and whose heads are quite different. Compared with Philini, Lamiinae nor exclude it from Cerambycinae. the small size and long and hairy antennae also are not consistent with any extant group. We cannot exclude it from Cerambycinae, Dorcasominae and Spondylidinae, but it does not clearly resemble 4.7. The compound eyes any extant group. Thus, the systematic position of Qitianniu is still questionable The coarsely facetted eyes and the dull color are suggestive of a and we provisionally place it as Cerambycidae incertae sedis. nocturnal species.

4.3. Sexual dimorphism 5. Conclusions

For Cerambycidae, sexual dimorphism ranges from virtually The first cerambycid beetle found in Cretaceous Myanmar none to extreme, and males may be distinctly smaller or distinctly amber, Qitianniu zhihaoi gen et sp. nov., is described and illustrated larger than females. Generally, however, sexual dimorphism is at on the basis of one well-preserved specimen. These findings pro- most moderate in Cerambycinae, Dorcasominae and Spondylidi- vide solid evidence for the origin of Cerambycidae. The presence of nae. For this amber specimen, the following characters indicate it to some very important characters, previously unobserved in Meso- be male: long antennae with 5 antennomeres surpassing the end of zoic fossils, are unambiguously confirmed for the first time. These the body (if it were female, then the male might have had antennae include the highly setose antennae and the hairy elytra. In addition, more than triple the length of the body); the long mandibles there is sufficient information to establish a new genus rather than (Fig. 2d), and the somewhat bent abdomen (Fig. 1b). Nonetheless, place this specimen in an extant genus. However, the systematic some characters might indicate it to be female: a structure at the position of Qitianniu is still questionable; it is provisionally classi- abdominal end (Fig. 1f, 1g) that might be a stylus (although its fied as Cerambycidae incertae sedis. 178 M.-Y. Lin, M. Bai / Cretaceous Research 75 (2017) 173e178

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