First record of a weevil (Coleoptera: Nemonychidae) from the Lower Cretaceous (Wealden) of southern England

Accepted Manuscript First record of a weevil (Coleoptera: Nemonychidae) from the Lower Cretaceous (Wealden) of southern England Andrei A. Legalov, Edmund A. Jarzembowski PII:

S0195-6671(17)30319-1

DOI:

10.1016/j.cretres.2017.10.006

Reference:

YCRES 3717

To appear in:

Cretaceous Research

Received Date: 11 July 2017 Revised Date:

16 September 2017

Accepted Date: 3 October 2017

Please cite this article as: Legalov, A.A., Jarzembowski, E.A., First record of a weevil (Coleoptera: Nemonychidae) from the Lower Cretaceous (Wealden) of southern England, Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.10.006. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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First record of a weevil (Coleoptera: Nemonychidae) from the Lower Cretaceous (Wealden) of southern England

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Institute of Systematics and Ecology of Animals of the Siberian Branch of the Russian Academy

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of Sciences, Frunze str., 11, Novosibirsk 630091, Russia

State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and

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b

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Andrei A. Legalov a, Edmund A. Jarzembowski b,c

Palaeontology, Chinese Academy of Sciences, 39 East Beijing, Road, Nanjing 210008, China c

Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK

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ARTICLE INFO

Article history:

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Received Accepted after revision

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Available online

Key words: Curculionoidea, Eobelinae, new species, Barremian, Early Cretaceous

ABSTRACT A new weevil, Oxycorynoides bucklowae sp. nov. is described from the lower Barremian Upper Weald Clay Formation of south Surrey, UK. The new species differs from other species of the nominative subgenus in the large body size; large, elongate eyes; and distinctly curved rostrum.

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It approaches but is also distinguished from the Berriasian-Hauterivian Oxycorynoides mongolicus Zherikhin, 1986 and O. gurvanensis Legalov, 2010 from Mongolia by the long metaventrite from the former species, and by the short rostrum from the latter.

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1. Introduction The Early Cretaceous entomofauna of the English Wealden Supergroup has been studied systematically in recent decades and 14 insect orders are now known to be present

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(Jarzembowski, 1995; Jarzembowski et al., 2016). The beetles (Order Coleoptera Linnaeus, 1758) are represented by at least nine species of Cupedidae Laporte, 1836 (Jarzembowski, 2011;

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Jarzembowski et al., 2013a, 2013b; 2015) and undescribed possible Elateroidea Leach, 1815 (Jarzembowski, 1995), Byrrhidae Latreille, 1804, Buprestidae Leach, 1815 (Jarzembowski et al., 2012), Carabidae Latreille, 1802 and Staphylinidae Latreille, 1802 (Jarzembowski et al., 2016). The weevil family Nemonychidae Bedel, 1882 was previously unknown from this deposit, but the gallery systems of bark beetles have been described (Jarzembowski, 1990).

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In general, the superfamily Curculionoidea (weevils) is well represented in the Early Cretaceous (Legalov, 2012). The first Nemonychidae appear in the fossil record on the boundary

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of the Middle and Upper Jurassic of Karatau, Kazakhstan (Gratshev and Legalov, 2014). The tribe Oxycorynoidini is represented by nine species in four genera from the Jurassic of Karatau

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to the Cretaceous (Albian) of Khetana (Legalov, 2015). The genus Oxycorynoides Arnoldi, 1977 is known from the Jurassic of Karatau to the Berriasian-Hauterivian of Mongolia. Here we describe a new species belonging to the eobeline Oxycorynoides from the lower

Barremian of the Upper Weald Clay Formation in the English Wealden.

2. Material and methods

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The unique type (holotype) is in the Natural History Museum, London (NHMUK). The comparative material is deposited in the Borissiak Paleontological Institute, Russian Academy of Sciences, Moscow. The holotype was examined dry and wetted with ethanol under a Wessex WSL1 long-arm

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binocular stereomicroscope. The long arm was needed because the specimen is near the centre of a large, oval, sideritic concretion split along the bedding plane; it is associated with numerous other insect remains (including Coleoptera), ostracods, fish scales, a gastropod and plant

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fragments including a coniferous leafy shoot. The concretions (which are sometimes phosphatic) occur in the lower part of the Upper Weald Clay exposed at Smokejack’s brickworks in the

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county of Surrey, UK (National Grid Reference TQ 116 373; Figs. 1, 2). They occur in the lower insect beds (1-6 NW: Austen et al., 2016; Fig. 3) and this one came from one of the upper courses.

A reconstruction of this beetle (Fig. 3) was painted by Biddy Jarzembowski.

terminology is used.

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Photographs were taken with an Olympus TG-4 digital camera. Standard beetle

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3. Systematic palaeontology

Class: Insecta Linnaeus, 1758

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Order: Coleoptera Linnaeus, 1758 Family: Nemonychidae Bedel, 1882 Subfamily: Eobelinae Arnoldi, 1977 Tribe: Oxycorynoidini Arnoldi, 1977 Genus: Oxycorynoides Arnoldi, 1977 Subgenus: Oxycorynoides s. str. Oxycorynoides bucklowae sp. nov. urn:lsid:zoobank.org:act:63BDA93F-A154-4062-B18D-F4C47A39C755

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Figs. 4-6.

Holotype, NHMUK II. 3102 [field no. S. 3073] a, b, part and counterpart, coll. E. and B.

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Jarzembowski.

Locality and strata. Smokejacks, nr Ockley, Surrey, England ( N 51 deg. 7 mins 53 secs, W 0 deg. 25 mins 10 secs; Fig. 1); Upper Weald Clay below British Geological Survey Bed 5c, lower

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Barremian (Liao et al., 2016: figs 1, 2; Fig. 2).

palaeontology, NHMUK.

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Etymology. The epithet of this species is after Jenny Parry née Bucklow, volunteer in

Description of holotype. Body length (without rostrum) 5.0 mm, length of rostrum 1.1 mm. Body dark-brown, moderately sclerotized, elongate, and moderately convex dorsally.

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Head capsule 0.9 times as short as wide, not constricted behind eyes. Mandibles comparatively long. Rostrum elongate, weakly curved, weakly dilated at base, about 10.0 times

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as long as wide at apex, 4.4 times in middle, 6.2 times at base; about 0.9 times as long as pronotum. Antennae long and thin, not geniculate, inserted at middle of rostrum. Antennomeres

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probably subconical. Forehead weakly convex. Eyes large, oval, 1.3 times as long as wide. Temples short, 0.3 times as long as longitudinal diameter of eyes. Pronotum with flattened disc. Lateral carinae of pronotum distinct. Elytra 2.6 times as long as pronotum, punctuate, striae narrow and distinct; punctures in

elytral striae medium size; intervals 2.5-5.0 as wide as striae; elytral apices gently curved. Precoxal portion of prosternum long, about 1.2 times as long as procoxal cavity. Procoxal cavities located at base of prosternum. Metaventrite long, 3.2 times as long as length of metacoxal cavity. Abdomen convex; ventrites 1 - 3 subequal in length; ventrite 1 about 0.7 times

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as long as metacoxal cavity; ventrite 4 about 1.1 times as long as ventrite 3; ventrite 5 subequal in length to ventrite 4. Femora slightly thickened.

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Comparison. The new species differs from other species of the nominative subgenus in the large body size, large elongated eyes, and distinctly curved rostrum. It is closest to the BerriasianHauterivian species O. mongolicus Zherikhin, 1986 and O. gurvanensis Legalov, 2010 because

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the elytra have deep, punctate striae and it has a thick rostrum; it is distinguished by the rostrum

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being shorter than in O. gurvanensis and metaventrite longer than in O. mongolicus.

Discussion

This beetle adpression does not belong to the subfamily Brenthorrhininae Arnoldi, 1977 because the procoxae are located closer to the basal margin of the prothorax. The pronotum with a distinct lateral carina shows that it belongs to a group of extinct subfamilies including

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Eobelinae Arnoldi, 1977, Cretonemonychinae Gratshev et Legalov, 2009 and Paleocartinae Legalov, 2003. The antennae inserted submedially suggest placement in the subfamily

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Eobelinae. Four tribes (Eobelini, Probelini Legalov, 2009, Oxycorynoidini and Karataucarini Legalov, 2009) are recognised in this subfamily. This species does not belong to Eobelini and

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Probelini because it has the rostrum slightly shorter than the pronotum and a small body size. The tribe Karataucarini with one genus Ampliceps Arnoldi, 1977 is known only from the Jurassic of Karatau and is characterized by the tarsomeres being dilated and wider than the tibiae, but these are not visible on the Wealden beetle. Nevertheless, Ampliceps is distinguished from the latter by the long rostrum, round eyes and thicker femora. The Wealden species is most like members of the Oxycorynoidini, representatives of which are found not only in the Jurassic of Karatau, but also in the Early Cretaceous of South America and Asia. The rostrum dilated at base in the Wealden form excludes it from the genera Microprobelus Liu, Ren et Shih, 2006 from the

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Barremian/Aptian of northeast China and Cratomacer Zherikhin et Gratshev, 2004 from the Aptian-Albian of Brazil. We therefore place this species in the genus Oxycorynoides Arnoldi, 1977 because of the structure of the rostrum, forehead being weakly convex, and punctures on the elytral striae of medium size. The new species belongs to the nominative subgenus because

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the rostrum base is weakly dilated and the elytral apices are gently curved (in lateral view). Weevils from the Barremian are also represented by two genera and two species of the tribe Oropsini Legalov, Azar, Kirejtshuk, 2017 (Nemonychidae) and the oldest bark-beetle

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Cylindrobrotus Kirejtshuk, Azar, Beaver, Mandelshtam et Nel, 2009 in Lebanese amber (Kirejtshuk et al., 2009; Legalov et al., 2017) plus ten species in the families Nemonychidae and

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Ithyceridae Schoenherr, 1823 from the Yixian Formation in China (Legalov, 2015). In addition, from the Hauterivian–Barremian of Las Hoyas in Spain has been described Distenorrhinus xavieri Zherikhin et Gratshev, 2003 (Brenthorrhininae: Distenorrhinini Arnoldi, 1977) and from the Barremian–Aptian of the Zhixin Basin in China Longodorsum generale Zhang, 1997 (Nemonychidae: Eobelidae) (Zherikhin and Gratshev, 2003; Zhang, 1997). The Berriasian-

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Hauterivian fauna is slightly richer than the Barremian one being represented by 16 species in two families (Legalov, 2015). Nineteen species in the families Nemonychidae, Ithyceriodae and

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Curculionidae are found in the Aptian (Legalov, 2015). There is thus a slight decrease in diversity of weevils (14 species) in the Barremian fossil record compared to the Berriasian-

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Hauterivian fauna (16 species) as well as late Aptian one (19 species). Present-day nemonychids are sometimes called pine flower weevils although Mesozoic

eobelines are usually associated with bennettitalean fossil plants (Gratshev and Legalov, 2014). The unique specimen of O. bucklowae sp. nov. is associated with a conifer (see above) and bennettitaleans are unknown from the type locality, although four pinopsidan (cheirolepidiacean) form genera are recorded (Ross and Cook, 1995): it is therefore possible that host transfer was already under way. Interestingly, a circular, weevil-like boring 1.3 mm in diameter has been found recently in an unnamed gymnospermous seed from the Wealden of the Isle of Wight

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(Barremian Wessex Formation at Barnes High on the southwest coast: Dr S. Sweetman, pers. comm.).

Conclusions

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This new find of a species in the genus Oxycorynoides shows that Eobelinae were widely distributed in the Early Cretaceous of Eurasia. The new species is closer to the BerriasianHauterivian species from Mongolia than the Late Jurassic species of Kazakhstan. This is the

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latest find of the genus Oxycorynoides. It is suggested that this species was probably associated

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with an extinct pinopsidan.

Acknowledgements

The studies of the first author were partially supported by the Federal Fundamental Scientific Research Programme for 2013–2020 (grant number VI.51.1.5) and the Russian Foundation for Basic Research (grant number 15-04-02971a). This is a Leverhulme Emeritus

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Fellowship contribution for the second author. The authors thank A.G. Ponomarenko, I.D. Sukacheva and A.P. Rasnitsyn (Moscow) for

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the opportunity to study comparative material, P. Austen (Seaford) for photographic help, and

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Dr. G.O. Poinar, Jr. (USA) and Dr. A.G. Kirejtshuk (Russia) for reviewing the manuscript.

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ACCEPTED MANUSCRIPT CAPTIONS TO FIGURES Fig. 1. Regional stratigraphy of the Weald Clay Group in SE England with localities in S-central

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Surrey from which fossil beetles have been described.

Fig. 2. Location map of quarries in the Weald Clay Group of S-central Surrey in SE England from which fossil beetles have been described: S, Smokejacks brickworks referred to herein (N

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51° 7’ 53” / W 0° 25’ 10”).

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Fig. 3. Photograph of new NE face of Smokejacks brickworks quarry (pit): the higher figure is standing on top of the upper insect bed (brown coloured), the figure below is in the lower insect beds (grey coloured) at about the level of the weevil find. Scale: higher figure is 1.9 m tall.

Fig. 4. Holotype of Oxycorynoides bucklowae sp. nov. A. part of adult beetle, lateral view. B.

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counterpart of adult beetle, lateral view. Scale bar 1.0 mm.

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Fig. 5. Holotype of Oxycorynoides bucklowae sp. nov. A. Interpretation of part of adult beetle,

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lateral view. B. Interpretation of counterpart of adult beetle, lateral view. Scale bar 1.0 mm.

Fig. 6. Reconstruction of Oxycorynoides bucklowae sp. nov.

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