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First record of Blattulidae from mid-Cretaceous Burmese amber (Insecta: Dictyoptera)
Accepted Manuscript First record of Blattulidae from mid-Cretaceous Burmese amber (Insecta: Dictyoptera) Lu Qiu, Zong-Qing Wang, Yan-Li Che PII:
S0195-6671(18)30527-5
DOI:
https://doi.org/10.1016/j.cretres.2019.03.011
Reference:
YCRES 4113
To appear in:
Cretaceous Research
Received Date: 25 December 2018 Revised Date:
27 February 2019
Accepted Date: 10 March 2019
Please cite this article as: Qiu, L., Wang, Z.-Q., Che, Y.-L., First record of Blattulidae from midCretaceous Burmese amber (Insecta: Dictyoptera), Cretaceous Research, https://doi.org/10.1016/ j.cretres.2019.03.011. 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 Blattulidae from mid-Cretaceous Burmese amber (Insecta: Dictyoptera)
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Lu Qiu, Zong-Qing Wang and Yan-Li Che*
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4 Institute of Entomology, College of Plant Protection, Southwest University, Chongqing, China
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*Corresponding author
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E-mail addresses: [email protected] (L. Qiu), [email protected] (Y.-L. Che)
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Blattulidae is an extinct cockroach family which was widely distributed around the world and
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lasted from Late Triassic to Cretaceous. Here we describe and illustrate the first blattulid genus
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found in the mid-Cretaceous Burmese amber, Huablattula gen. nov., to accommodate two new
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species, Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. Most blattulid genera were
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only described based on inadequate samples, or were based only on nymphs, isolated tegmina, or
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wings. The well-preserved adult specimens described in this work not only fill a blank in
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Blattulidae study in Burmese amber, but also advance out knowledge of this family.
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Keywords: Huablattula, new genus, new species, ovipositor, Mesozoic
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1. Introduction
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The Mesozoic cockroach family Blattulidae was established with genus Blattulites and included
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eight other genera originally belonging to the family Mesoblattinidae (Vishnyakova 1982). This
ACCEPTED MANUSCRIPT family had a history of more than 100 million-years, originating in the Late Triassic and ranging
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across the whole Jurassic and Cretaceous (Vršanský 2002; Vršanský, Vishnyakova et Rasnitsyn
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2002; Vršanský et al. 2013). Blattulidae is considered to be related to Corydiidae (=Polyphagidae)
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(Vishnyakova 1982; Vršanský 1999). Thus, this family was accepted as a member of superfamily
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Corydioidea/ Polyphagoidea (Vršanský, Vishnyakova et Rasnitsyn 2002; Vršanský 2003, 2004a,
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2004b; Vršanský et Ansorge 2007; Lee 2016); while some works also treated it as a member of the
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Blattuloidea (Anisyutkin et Gorochov 2008; Vršanský 2005a, 2008b, 2009). It was a relatively
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dominating family, being widely distributed around the world (Vršanský 1999; Wang, Liang et
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Ren 2007). However, its diversity at the generic level is poor, with about 18 genera reported to
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date (Vishnyakova 1982; Vršanský 2003, 2005a, 2005b, 2009; Martins-Neto, Mancuso et Gallego
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2005; Cifuentes-Ruiz et al. 2006; Vršanský and Ansorge 2007; Wang et al. 2007; Wang, Liang et
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Ren 2007; Anisyutkin et Gorochov 2008). Most genera have low diversity, with only a single
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species recorded (monotypic genera). The relatively diverse genera are Blattula and Elisama, the
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former containing 44 species and the latter with 20 species (Clapham et Karr 2018).
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Describing fossil cockroaches only based on nymphs (difficulty in distinguishing the exact
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families), or isolated tegmina (abundant plesiomorphies and homoplasies present) is considered to
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be less reliable (Li et Huang 2018). Such a situation is also present in the study of Blattulidae.
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Genera such as Anablatta, Argentinoblattula, Xonpepetla, Svabula, and Orbablattula were
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described based on the tegmen (Martinez-Delclos 1993; Martins-Neto, Mancuso et Gallego 2005;
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Vršanský 2005b; Cifuentes-Ruiz et al. 2006; Martins-Neto, Gallego et Zavattieri 2007). Kridla
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was described based on the hind wing (Vršanský 2005a). Globula, Nula, and Blattulites were
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described from nymphs (Vishnykova 1982; Vršanský 2008a, 2009). The credibility of these genera
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ACCEPTED MANUSCRIPT remains to be confirmed. On the other hand, genera such as Tarakanula, Habroblattula,
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Macaroblattula, and Ocelloblattula were described based on relatively well-preserved materials,
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leading to more informative studies (Vršanský 2003; Wang et al. 2007; Wang, Liang et Ren 2007;
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Anisyutkin et Gorochov 2008). Revisions of Blattulidae genera and species were also made using
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abundant materials and by re-examining the type specimens (Vršanský et Ansorge 2007; Wang et
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al. 2007; Lee 2016).
The records of Blattulidae in amber are poor, with four monotypic genera known, viz.
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Ocelloblattula in Lebanese amber (Anisyutkin et Gorochov 2008) (Ocelloblattula santanensis
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(Lee, 2016) described based on impression fossil from Brazil needs further confirmation), and
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Nula, Batola, and Globula in French ambers (Vršanský 2008a; Vršanský 2009). However, only
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Ocelloblattula was described based on an almost complete adult specimen. Batola nikolai was
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described based on an incomplete adult specimen and several nymphs and body fragments, but the
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adult specimen was designed as a paratype. Nula and Globula were totally based on nymphs,
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which are poor in classification and may produce difficulties in matching adults with nymphs in
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the future study.
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Herein, the authors report the first record of Blattulidae in Myanmar amber, with Huablattula
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gen. nov. described to include Huablattula hui sp. nov. and Huablattula jiewenae sp. nov., both of
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which are described based on well-preserved specimens. One Huablattula sp. is also presented for
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study but not described to species due to the insufficient characters present in the amber. The
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specimens presented herein all reveal some important characters to enrich our knowledge of the
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Blattulidae. These discoveries highlight the palaeodiversity of this family in the mid-Cretaceous,
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which suggests more taxa will be discovered in the near future.
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ACCEPTED MANUSCRIPT 67 2. Material and methods
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The amber pieces containing the specimens were obtained from the Hukawng Valley, Kachin
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State, in Northern Myanmar (Yin et al. 2018: fig. 1A, 26°21′33.41″N, 96°43′11.88″E). The age of
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Burmese amber is estimated as the Late Cretaceous (98.79±0.62 Ma) based on U-Pb dating of
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zircons (Shi et al. 2012). The studied specimens are deposited in the Institute of Entomology,
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Southwest University, Chongqing, China (SWU), some of which are previously in the personal
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collections of Zhengkun Hu, Guizhou, China (CZKH) and Dan Zuo, Hunan, China (CDZ). The
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amber pieces were polished using sandpaper of different grain sizes and with rare earth polishing
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powder to make them clear. All images (Figs. 1-6) were made using a Leica® M205A
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stereomicroscope plus Leica DFC 550 and Canon EOS 50D camera plus a Canon EF 100mm
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f/2.8L IS USM Macro lens combined with Helicon Focus software and modified in Adobe
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Photoshop CS6. Morphological terminology mainly follows Grandcolas (1994) and Roth (2003).
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Venation terms follow Li et al. (2018). This publication is registered in ZooBank under
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urn:lsid:zoobank.org:pub: 10AF1305-932C-41E3-9617-360E794383F2.
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3. Systematic paleontology
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Order Dictyoptera
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Family Blattulidae
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Genus Huablattula gen. nov.
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(urn:lsid:zoobank.org:act: F72A82F0-B379-4E3C-90A9-6CA195F22928)
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Type species: Huablattula hui sp. nov. here designated.
ACCEPTED MANUSCRIPT Description. Male: body small, elongate, sub-transparent, decorated with spots and stripes. Head
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broad, near triangular, not hidden by head; eyes round and large, strongly convex; interocular
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space greater than the distance between antennal sockets; the space between eyes and above
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antennal sockets distinctly prominent; three ocelli present (currently this character is only
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indicated in Huablattula jiewenae sp. nov.); antennae slender, exceeding the apex of tegmina,
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segments short, with sparse but distinct setae from antennomere 10 on, scape cylindrical, robust;
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clypeus convex but small; mouth constricted; maxillary palpomeres 3–4 slender, palpomere 5
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slightly expanded or rounded. Pronotum smooth, without pubescent, usually decorated with dark
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stripes, hind margin distinct protruded medially. Venation simple, decorated with dark stripes in
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tegmina, median of CuA with a large spot or irregular macula, which may extend to the distal
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margin of anal area. Tegmen with simple and long ScP, R branches sub-parallel, M and CuA
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usually with several branches; intercalary veins present between branches of R, M and CuA.
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Wings with dark venation; ScP simple, connected with RA near the base; RA with several short
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branches at distal half, the area among the short branches usually dark and thickened (forming
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pterostigma); RP simple with about 4 branches; M straight, bifurcate at distal half; CuA with 4–5
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parallel and curved branches; CuP very long and slender; Pcu with several small branches basally;
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anal area small, simply folded behind the rest part of the wing; intercalary veins present between
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RP, M and distal portion of CuA; cross veins present but sparse. Each femoral apex with a spine
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(extremely long at mid and hind femora), hind margins of femur with two rows of long but sparse
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spines; tarsal claw asymmetrical, arolia large. Cerci long, pubescent, distal segment long-spinous,
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or elongate. Subgenital plate protruded, pubescent, with a pair of segmented and elongate styli
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near the apex.
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ACCEPTED MANUSCRIPT Female: relatively larger than male, the habitus and stripes generally similar to male. Subgenital
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plate strongly bulging, apex constricted, forming a narrow valved protrusion, a pair of ovipositor
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processes extend externally, slender, apex curved and acute, base with a valved sheath.
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Etymology. Hua refers to the Chinese character 花, meaning piebald, for its body is decorated
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with multiple stripes and spots. Hua + blattula therefore means “piebald blattulid cockroach”.
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Species included. Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. described herein.
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Also with one undescribed species mentioned below.
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Remarks. Huablattula gen. nov. resembles Ocelloblattula in the Lebanese amber (Anisyutkin et
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Gorochov 2008) in its general appearance, shape of head, globous eyes, shape of pronotum and
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legs; but it differs from the latter by the ocelli and its position, which are smaller and with the
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lateral two more separated and closer to the eyes. It can be also distinguished from the latter by its
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remarkable markings on pronotum and stripes on tegmina. The apical portion of subgenital plate
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as well as the ovipositor also easily distinguish it from the latter. This new genus differs from
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Habroblattula and Macaroblattula found in the Yixian Formation (Wang et al. 2007; Wang, Liang
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et Ren 2007) by the non-branched ScP. This new genus also resembles other impression fossil taxa,
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such as Vrtula, Elisama, Tarakanula, and Svabula by the markings on its tegmen, but differs from
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Vrtula, Elisama, and Tarakanula by having wing pterostigma (Vršanský 2005a; Vršanský 2008b).
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It differs from Svabula by having a more protruded and angular apex of tegmen (Vršanský 2005b),
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and from Kridla by not having a darkened apex of the wing (Vršanský 2005a). It can be
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distinguished from Blattula by having dark macula on the tegmen (Wang, Liang et Ren 2007;
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Vršanský et Ansorge 2007). This new genus can be distinguished from the adult paratype of
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Batola nikolai by the protruded apex of the subgenital plate (Vršanský 2009).
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Huablattula hui sp. nov. (Figs. 1–4)
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(urn:lsid:zoobank.org:act: A3A7FC5C-C3EB-46FB-BC2B-412536F07045)
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complete, apices of both antennae and right tegmen lost, parts of tarsi in right mid leg and both
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hind legs lost (SWU, ex CZKH, SWUMA-0002).
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Paratypes. A well-preserved male adult in amber with tegmina and wings in resting position,
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distal half of antennae missing, the joint part of femur and tibiae in left front and mid legs, apical
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portion of right femur and the whole tibiae and tarsus, apical portion of right tarsus, the whole
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tarsus of both hind legs missing (SWU, ex CDZ, SWUMA-0003).
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A sex unknown adult in amber, with tegmina and wings in resting position, ventral side
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unobservable (SWU, ex CDZ, SWUMA-0004).
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A sex unknown adult with tegmina and wings in resting position, well-preserved in amber, but the
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hind portion of the body is missing, including the distal half of left mid tibiae and the whole tarsus,
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the whole left leg, the whole right hind tarsus, abdomen tip, and tegmen and wing tips. A
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Hymenoptera insect is also preserved in the amber (SWU, ex CZKH, SWUMA-0005).
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Sex unknown adult fragments preserved in amber, with only tegmina, mesonotum, one antenna,
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one femur left. A broken Hymenoptera insect is also preserved in the amber (SWU, ex CZKH,
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SWUMA-0006).
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A female adult in amber, head broken. Two Diptera insects and a probably Neuroptera insect are
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also preserved in the amber (SWU, SWUMA-0007).
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ACCEPTED MANUSCRIPT Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
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Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
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Diagnosis. Huablattula hui sp. nov. resembles Huablattula jiewenae sp. nov., but it can be easily
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distinguished from the latter by the markings on pronotum, the outline of pronotum, and the
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stripes of tegmina; a large triangular plaque is found along the distal margin of anal area to the
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median portion of CuA on the tegmen of Huablattula hui sp. nov., while a large spot is found
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around median of CuA on that of Huablattula jiewenae sp. nov.
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Description. Holotype SWUMA-0002. Body elongate, transparent, decorated with markings and
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stripes (Fig. 1A–B). Length 8.7 mm, including tegmina and wings 11.0 mm.
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Head (face toward left; the ventral detail is difficult to observe) nearly triangular, with dark stripes and spots, length 1.93 mm. Eyes round and large, situated at distal corners, length 0.87 mm;
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interocular space greater than the distance between antennal sockets; space between eyes and
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above antennal sockets distinctly prominent; ocelli unclear in the amber. Antennae slender, distal
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portions missing (5.40 mm and 5.81 mm for the remaining portion of left and right, respectively);
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segments short, with sparse but distinct setae from antennomere 10 on; scape cylindrical, robust,
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length 0.42 mm; pedicellus nearly conical, length 0.14 mm, the distal margin expanded, width
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0.12 mm; the rest of antennomeres cylindrical, generally becoming longer and thinner toward
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apex. Clypeus small, slightly convex. Mouth constricted; maxillary palpomeres 1–2 subequal in
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length (about 0.13 mm), 3–5 lengths 0.41 mm/0.37 mm/0.35 mm, palpomeres 1–2 short, nearly
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globous, palpomeres 3–4 slender, palpomere 5 slightly expanded.
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Pronotum (Fig. 1H) transparent, smooth, without pubescent; disc with symmetrical dark
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markings that extend in two long stripes posteriorly, basal half decorated with some light-colored
ACCEPTED MANUSCRIPT spots; margins outlined with dark line. The margins of the pronotum generally rounded, anterior
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margin round, posterior margin protruding and constricted medially, lateral-hind parts roundly
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protruding. Length 2.00 mm (measured medially), width 2.46 mm (measured at widest points).
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Tegmina (Fig. 1D–-E) length 8.19 mm (left one), width 2.12 mm. Venation decorated with
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dark thick stripes, anal veins with reticulated stripes, a large triangular plaque lies along the distal
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margin of anal area to the median portion of CuA. ScP simple, R branches sub-parallel, M
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bifurcate near base, CuA with 4 branches; intercalary veins present between branches of R, M and
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CuA (based on the opened right tegmen). Wings (Fig. 1F–G) with dark venation, apex narrow;
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ScP simple, connected with RA at 1/3 from the base; RA with 4 short branches at distal half, the
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area among the 4 branches dark and thickened (with pterostigma); RP bifurcate basally, distal
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portion with three branches; M straight, with a bifurcate branch at 2/3 from the base; CuA with 4
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parallel and curved branches; CuP slender; Pcu with three small branches basally. Intercalary veins
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present between RP, M and distal portion of CuA (based on the folded right wing).
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Legs mixed with dark maculae. Front leg: femur 1.69 mm, tibia 1.20 mm, tarsus 1.46 mm (tarsal
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segments 1–5 lengths 0.58 mm/0.28 mm/0.21 mm/0.16 mm/0.27 mm); mid leg: femur 2.42 mm,
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tibia 2.12mm, tarsus 1.62 mm (tarsal segments 1–5 lengths 0.81 mm/0.26 mm/0.18 mm/0.14
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mm/0.27 mm); hind leg: femur 2.55 mm, tibia 3.01 mm, tarsus missing. Each leg with a long
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spine at the apex of femur, and two rows of long but sparse spines at hind margins. Length of the
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apical spines from front, mid and hind femora 0.26 mm/0.70 mm/1.04 mm (extremely long for
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those of mid and hind legs). Tarsal claw asymmetrical. Arolia large (Fig. 1C).
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Meso- and metanotum dark dorsally. Cerci long, length 2.09 mm, pubescent, apical one spinous.
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Subgenital plate protruding, pubescent, slightly asymmetrical distally, median of apex with a small
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ACCEPTED MANUSCRIPT protrusion, two long styli situated laterally beside the protrusion, segmented, basal segment not
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enlarged, length 1.05 mm (Fig. 1I–J).
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Paratype SWUMA-0003. Length 7.26 mm, including tegmina and wings 8.99 mm, width 3.46
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mm. Generally similar to the holotype (Fig. 2A–B). The head can be better observed, length of
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head 1.72 mm, width of head 1.75 mm. The markings on face as in Fig. 2C figured. Due to the
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head being nearly transparent, the ocelli are still difficult to recognize, only left ocellus is distinct,
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situated next to the middle margin of the eye. Subgenital plate, styli and cerci similar to those of
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the holotype (Fig. 2D).
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Paratype SWUMA-0004. Length including tegmina 9.76 mm, width including tegmina 4.03 mm
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(Fig. 2E).
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Paratype SWUMA-0005. The antenna is complete in this specimen, the total length of the
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antenna is 9.19 mm, which is distinctly longer than the body length of this species. Right ocellus
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can be recognized in this specimen (Fig. 2F–G).
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Paratype SWUMA-0006. Length of left tegmen 8.20 mm, width 2.45 mm. Since the holotype
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specimen slightly overlaps in the amber (probably due to the struggling of the insect), the venation
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of the holotype is not as clearly observed. From this specimen, the intercalary veins can be better
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recognized (Figs. 3A–F).
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Paratype SWUMA-0007. The only known female (Fig. 4A–B). Body length from antetior
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margin of pronotum to tegmina tip 12.9 mm, length from antetior margin of pronotum to the apex
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of ovipositor valves 14.7 mm. Pronotum length 2.68, width 2.89 mm. The marking on pronotum is
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similar to male, but shows some reduction (Fig. 4C). Subgenital plate (Fig. 4D) large, strongly
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bulging and protruded, with a narrow and valved posterior lobe. External ovipositor present (Fig.
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ACCEPTED MANUSCRIPT 4E) consisting of two processes and a sheath: a pair of very elongate, flat and pubescent processes
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extend from the posterior lobe, apical portions of both processes curved and with needle-like apex;
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a valved sheath extends from the posterior lobe dorsally, and covers the base of the two long
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processes.
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Etymology. Named after Mr. Zheng-Kun Hu (Guizhou, China) who kindly donated his amber
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collection to us for study.
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227 Huablattula jiewenae sp. nov. (Fig. 5)
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(urn:lsid:zoobank.org:act: CC3A1364-8C8F-4A69-8581-992109A1D6B4)
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Type material. Holotype. A well preserved male adult in amber with tegmina and wings in resting
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position, apices of antennae missing, median of left antenna missing, mouth parts broken, the
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jointed portions of femur and tibia are missing for front legs, apex of tarsi are missing for mid legs,
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left tibia broken, tarsus missing, right tibia with broken tibia. Two Diptera insects and a small
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Coleoptera insect are also preserved in the amber (SWU, ex CDZ, SWUMA-0008).
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Paratype. A sex unknown adult preserved in amber with tegmina and wings in resting position,
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mostly complete, but ventral side unobservable. A Hymenoptera insect, a Staphylinidae beetle and
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a Eucnemidae beetle are also preserved in the amber (SWU, ex CZKH, SWUMA-0009).
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Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
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Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
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Diagnosis. See the diagnosis of Huablattula hui sp. nov.
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Description. Holotype SWUMA-0008. Body elongate, transparent, decorated with stripes and
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ACCEPTED MANUSCRIPT spots (Fig. 5A–B). Length 7.2 mm, including tegmina and wings 9.3 mm, width 4.3 mm (the
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tegmina in the amber are slightly opened, thus this measured body width may be wider than
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actual).
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Head nearly triangular, vertex with three longitudinal stripes, length 1.38 mm (the head is upward
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in the amber, thus this measurement is shorter than usual), width 1.80 mm. Eyes round and large,
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situated at distal corners; interocular space greater than the distance between antennal sockets
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(0.76 mm versus 0.37 mm); the space between eyes and above antennal sockets prominent, a dark
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band transversely across the prominent portion; three ocelli present, situated on the transverse
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band (two near the eyes, one in the center, intervals between the three ocelli about 0.3–0.4 mm).
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Antennae slender, median part of left one missing, apex of right one missing, measurement for the
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remaining parts is difficult; segments short, with sparse but distinct setae from antennomere 10 on;
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scape cylindrical, length 0.26 mm; pedicellus nearly cylindrical, length 0.17 mm; the rest of
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antennomeres cylindrical, generally becoming longer toward apex. Clypeus broken. Mouth parts
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constricted; maxillary palpomeres broken, palpomeres 4–5 length 0.29 mm/0.27 mm, palpomere 5
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roundly enlarged (Fig. 5D–E).
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Pronotum smooth, without pubescent, lateral margins transparent; nearly pentagonal, anterior
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margin nearly straight, lateral portions and hind margin protruding; four longitudinal bands cross
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throughout from the anterior margin to the posterior margin, the lateral two thicker than the inner
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two; margins of pronotum outlined with a dark line very narrowly. Length 2.03 mm (measured
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medially), width 2.20 mm (measured from widest point) (Fig. 5D).
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Tegmina length 7.29 mm, width 2.23 mm. Venation decorated with dark stripes, median of CuA
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with a large irregular spot. ScP simple, R branches sub-parallel, distal two with multiple branches,
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ACCEPTED MANUSCRIPT M bifurcate medially, CuA with 4 branches; intercalary veins present between branches of R, M
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and CuA (Fig. 5H). Wings unobservable, apex narrow.
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Legs mixed with some small spots. Front leg: the jointed portion of femur and tibia missing for
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both, tarsus 1.17 mm (tarsal segments 1–5 lengths 0.47 mm/0.22 mm/0.17 mm/0.13mm/0.24mm);
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mid leg: femur 1.79 mm, tibia 1.68 mm, tarsus with apical portion missing for both (tarsal
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segments 1–3 lengths 0.63 mm/0.27 mm/0.17 mm); hind leg: femur 2.20 mm, tibia 3.06 mm, left
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tarsus with apical portion missing, right tarsus all missing (tarsal segments 1–2 lengths 0.82
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mm/0.41 mm). Each femur with two rows of long but sparse spines at hind margins, mid and hind
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femora with extremely long spines at apex. Tarsal claw asymmetrical. Arolia large.
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Cerci long, length 1.79 mm, pubescent, apical two elongate. Subgenital plate protruded, pubescent,
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apex unobservable, styli segmented, length 1.06 mm, basal segment not enlarged (Fig. 5F).
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Paratype SWUMA-0009. Length including tegmina and wings 9.60 mm. Generally similar to the
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holotype (Fig. 5C), antenna exceeding the tip of tegmina. Tarsal claw asymmetrical (Fig. 5G).
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Etymology. Named after Ms. Jie-Wen Zhao (Hunan, China). Her mother Ms. Dan Zuo kindly
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provided her ambers for this study and hopes this honor inspires her daughter’s interest in natural
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history.
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Huablattula sp. (Fig. 6)
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Material examined. Male adult fragments preserved in amber, only wings, metanotum, tergum 1
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and subgenital plate left (SWU, SWUMA-0010).
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Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
ACCEPTED MANUSCRIPT Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
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Description. Wing (Fig. 6A–D) rounded, length 6.82 mm, apical portion not narrowed, relatively
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wider than that of H. hui sp. nov.; ScP simple, connected with RA at 1/3 from the base; RA with
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3–4 short branches at distal half, the area among the branches dark and thickened (with
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pterostigma); RP with 4–5 parallel branches; M simple, bifurcate at distal half; CuA with 5
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parallel and curved branches; CuP very long; Pcu with two small branches basally, distal one
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bifurcate; intercalary veins present between RP, M and CuA; anal area simply folded once,
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overlapped behind the remaining part of the wing. Subgenital plate similar to that of H. hui sp.
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nov., with a small protrusion at apex, two basal lobes of the subgenital plate present (this character
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is the same as in extant cockroaches); stylus segmented, length 1.28 mm, pubescent, basal
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segment slightly enlarged (Fig. 6E).
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Remarks. This specimen has a wider wing, the apex is rounded, the basal segment of stylus is
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slightly enlarged. Thus we definitely consider it to be a different species from H. hui sp. nov. and
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H. jiewenae sp. nov., but since no complete specimen is available, we temporarily treat it as an
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undescribed species.
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4. Discussion
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Although the ocelli of H. hui sp. nov. cannot be clearly observed, Huablattula hui sp. nov. and
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Huablattula jiewenae sp. nov. are considered congeners by their general appearance,
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triangular-shaped head, globous and large eyes, shape of pronotum, the long spines on legs,
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asymmetrical tarsal claws, protruded subgenital plate, segmented and elongate styli, their similar
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venation, the pronotum with markings, and the tegmina decorated with dark stripes in both
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species. The external ovipositor in this family has been reported from some impression fossils
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(Vishnykova 1982; Wang et al. 2007). Before this research, the holotype of Ocelloblattula
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ponomarenkoi was the only one where the ovipositor was observed in amber (Anisyutkin et
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Gorochov 2008). This newly described female of Huablattula hui gen. nov. et sp. nov. is the
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second amber record of the ovipositor in this family. Compared to O. ponomarenkoi, H. hui sp.
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nov. has a different subgenital plate, which is valved posteriorly (while rounded posteriorly in O.
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ponomarenkoi). The apical portions of the ovipositor processes are also different (needle-like and
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roundly curved in H. hui sp. nov., while acuminate and extended caudad to the apex of posterior
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lobe of genital plate in O. ponomarenkoi). The two ovipositor processes are much more extended
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in H. hui sp. nov. than in O. ponomarenkoi. A valved sheath is present in H. hui sp. nov., but seems
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not to be indicated in O. ponomarenkoi.
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From the delicate body, large eyes, long antennae, well-developed wings and the gorgeous
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stripes on the pronotum and tegmina, Huablattula gen. nov. is believed to be an active insect in
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daytime. The slender legs (with long spines), large arolia and weak and asymmetrical tarsal claws
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indicate that this genus inhabited the leaves of trees.
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5. Conclusion
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This paper presents the first record of the extinct cockroach family Blattulidae in Upper
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Cretaceous Burmese amber, with description of Huablattula gen. nov. to accommodate two new
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species, Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. Most of the studied material is
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well-preserved, enriching our knowledge of Blattulidae.
ACCEPTED MANUSCRIPT 331 Acknowledgments
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We thank Mr. Zheng-Kun Hu (Guizhou, China) and Ms. Dan Zuo (Hunan, China) for providing
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these precious amber materials. We also give our thanks to John Richard Schrock (Department of
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Biological Sciences, Emporia State University) for revising the manuscript before submission.
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Anonymous reviewers are acknowledged for their valuable comments and suggestions to the
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manuscript. This work has been supported by the National Natural Sciences Foundation of China
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(Nos. 31772506), Program of Ministry of Science and Technology of the People’s Republic of
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China (2015FY210300), and Natural Science Foundation Project of Chongqing (No.
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cstc2016jcyjA0487).
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341 References
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Blattulidae from Lebanese amber (Dictyoptera, Blattina). Paleontologicheskii Zhurnal 1, 44–
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Clapham, M., Karr, J., 2018. Taxonomic occurrences of Blattula and Elisama recorded in the
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Formation, Difunta Group in northeastern Mexico. Geologica Carpathica 57, 347–354. Grandcolas, P., 1994. Phylogenetic systematics of the subfamily Polyphaginae, with the assignment of Cryptocercus Scudder, 1862 to this taxon (Blattaria, Blaberoidea,
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Polyphagidae). Systematic Entomology 19, 145–158. Lee, S.-W., 2016. Taxonomic diversity of cockroach assemblages (Blattaria, Insecta) of the Aptian Crato Formation (Cretaceous, NE Brazil). Geologica Carpathica 67, 433–450. Li, X.R., Huang, D., 2018. A new Cretaceous cockroach with heterogeneous tarsi preserved in
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terminology. Zoomorphology 2018, 1–15. https://doi.org/10.1007/s00435-018-0419-6
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Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q., Li, X., 2012.
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Age constraint on Burmese amber based on U-Pb dating of zircons. Cretaceous Research 37,
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155–163.
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Vishnyakova, V.N., 1982. Jurassic cockroaches of the new family Blattulidae from Siberia. Paleontologicheskii Zhurnal 1982, 69–79. Vršanský, P., 1999. Two new species of Blattaria (Insecta) from the Lower Cretaceous of Asia, with comments on the origin and phylogenetic position of the families Polyphagidae and
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Blattulidae. Entomological Problems 30, 85–91.
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Vršanský, P., 2002. Origin and the early evolution of mantises. AMBA Projekty 6, 1–16.
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Vršanský, P., 2003. Unique assemblage of Dictyoptera (Insecta - Blattaria, Mantodea, Isoptera)
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Vršanský, P., 2004. Cretaceous Gondwanian cockroaches (Insecta: Blattaria). Entomological Problems 34, 49–54.
Vršanský, P., 2004. Transitional Jurassic/Cretaceous cockroach assemblage (Insecta, Blattaria)
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from the Shar-Teg in Mongolia. Geologica Carpathica 55, 457–468. Vršanský, P., 2005a. A fossil insect in a drilling core sample - cockroach Kridla stastia gen. et sp. nov. (Blattulidae) from the Cretaceous of the Verkhne-Bureinskaya Depression in eastern
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Russia. Entomological Problems 35, 115–116.
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Vršanský, P., 2008a. A complete larva of a Mesozoic (Early Cenomanian) cockroach (Insecta:
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Blattaria: Blattulidae) from the Sisteron amber (Alpes de Haute Provence, SE France).
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Geologica Carpathica 59, 269–272.
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Vršanský, P., 2008b. New blattarians and a review of dictyopteran assemblages from the Lower
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Cretaceous of Mongolia. Acta Palaeontologica Polonica 53, 129–136. Vršanský, P., 2009. Albian cockroaches (Insecta, Blattida) from French amber of Archingeay. Geodiversitas 31, 73–98. Vršanský, P., Ansorge, J., 2007. Lower Jurassic cockroaches (Insecta: Blattaria) from Germany and England. African Invertebrates 48, 103–126.
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Cockroaches probably cleaned up after dinosaurs. PLoS ONE 8, e80560, 80561–80511.
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Vršanský, P., Vishnyakova, V.N. & Rasnitsyn, A.P., 2002. Order Blattida Latreille, 1810. The
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cockroaches. In: Rasnitsyn, A.P. & Quicke, D.L.J. (eds.). History of Insects. Kluwer
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Academic Publisher, Dodrecht, 263–271.
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Wang, T., Ren, D., Liang, J.-H., Shih, C., 2007. New Mesozoic cockroaches (Blattaria: Blattulidae)
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from Jehol Biota of western Liaoning in China. Annales Zoologici (Warsaw) 57, 483–495.
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Wang, T.-T., Liang, J.-H., Ren, D., 2007. Variability of Habroblattula drepanoides gen. et. sp. nov.
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(Insecta: Blattaria: Blattulidae) from the Yixian Formation in Liaoning, China. Zootaxa 1443,
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17–27.
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Yin, Z., Cai, C., Huang, D., 2018. Last major gap in scydmaenine evolution filled (Coleoptera:
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Staphylinidae), Cretaceous Research 84, 62–68. doi: 10.1016/j.cretres.2017.10.026.
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Fig. 1. Huablattula hui sp. nov., habitus and detailed characters of holotype (SWUMA-0002),
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male. A. Habitus, dorsal view. B. Habitus, ventral view. C. Tarsal claws, rear view. D–E. Right
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tegmen (intercalary veins are omitted in fig. E). F–G. Right wing. H. Pronotum (outlined). I.
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Abdominal tip, ventral view. J. Detail of right stylus (arrows indicate the segmented parts).
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Abbreviations: arolium (ar), cercus (ce), stylus (st), subgenital plate (sg), tarsal claw (tc),
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tarsomere 5 (t5). Scale bars: A–B = 2 mm; C = 0.1 mm; D–G = 1 mm; H–I = 0.5 mm; J not to
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scale.
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(SWUMA-0003~0005). A–B. Habitus and detailed characters of SWUMA-0003, male; A. Dorsal
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view; B. Ventral view; C. Head; D. Abdominal tip. E. Habitus of SWUMA-0004, sex unknown,
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dorsal view. F–G. Habitus of SWUMA-0005, sex unknown; F. Dorsal view; G. Ventral view.
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Abbreviations: cercus (ce), eye (ey), ocellus (oc), stylus (st), subgenital plate (sg). Scale bars: A–
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B, E–G = 2 mm; C–D = 0.5 mm.
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Fig. 3. Huablattula hui sp. nov., detail characters of paratype (SWUMA-0006). A. General
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condition of the amber; B. Basal portion of the antennal fragment; C. Right tegmen; D. Leg
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fragment (show tibiae and the spines); E–F. Left tegmen. Scale bars: A = 2 mm; B, D = 0.5 mm; C,
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E–F = 1 mm.
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Fig. 4. Huablattula hui sp. nov., habitus and detail characters of paratype (SWUMA-0007), female.
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A. Habitus, dorsal view; B. Habitus, ventral view; C. Pronotum; D. Abdominal tip; E. Apical
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portion of subgenital plate and ovipositor. Abbreviations: cercus (ce), subgenital plate (sg),
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ovipositor (ov), bubble (bu), sheath (sh), ovipositor process (pr), posterior lobe (pl). Scale bars:
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A–B, D = 2 mm; C = 1 mm; E = 0.5 mm.
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Holotype (SWUMA-0008), male; A. Habitus, dorsal view; B. Habitus, ventral view; C. Paratype
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(SWUMA-0009), sex unknown, dorsal view; D. Head and pronotum, holotype, dorsal view; E.
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Head (arrows indicate the ocelli), holotype, ventral view; F. Abdominal tip, holotype, ventral view;
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G. tarsal claw, paratype, dorsal view; H. Right tegmen, holotype. Abbreviations: arolium (ar),
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cercus (ce), stylus (st), subgenital plate (sg), tarsal claw (tc), tarsomere 5 (t5), tibiae (ti). Scale bars:
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A–C, H = 2 mm; D–F = 0.5 mm; G = 0.1 mm.
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Fig. 6. Huablattula sp. (SWUMA-0010), male. A–B. General condition of the amber; C–D.
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Wings; E. Subgenital plate. Abbreviations: basal lobe (bl), protrusion (pr), stylus (st), subgenital
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plate (sg). Scale bars: A = 2 mm; B–E = 1 mm.
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DOI:
https://doi.org/10.1016/j.cretres.2019.03.011
Reference:
YCRES 4113
To appear in:
Cretaceous Research
Received Date: 25 December 2018 Revised Date:
27 February 2019
Accepted Date: 10 March 2019
Please cite this article as: Qiu, L., Wang, Z.-Q., Che, Y.-L., First record of Blattulidae from midCretaceous Burmese amber (Insecta: Dictyoptera), Cretaceous Research, https://doi.org/10.1016/ j.cretres.2019.03.011. 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 Blattulidae from mid-Cretaceous Burmese amber (Insecta: Dictyoptera)
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Lu Qiu, Zong-Qing Wang and Yan-Li Che*
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4 Institute of Entomology, College of Plant Protection, Southwest University, Chongqing, China
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*Corresponding author
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E-mail addresses: [email protected] (L. Qiu), [email protected] (Y.-L. Che)
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Blattulidae is an extinct cockroach family which was widely distributed around the world and
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lasted from Late Triassic to Cretaceous. Here we describe and illustrate the first blattulid genus
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found in the mid-Cretaceous Burmese amber, Huablattula gen. nov., to accommodate two new
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species, Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. Most blattulid genera were
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only described based on inadequate samples, or were based only on nymphs, isolated tegmina, or
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wings. The well-preserved adult specimens described in this work not only fill a blank in
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Blattulidae study in Burmese amber, but also advance out knowledge of this family.
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Keywords: Huablattula, new genus, new species, ovipositor, Mesozoic
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1. Introduction
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The Mesozoic cockroach family Blattulidae was established with genus Blattulites and included
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eight other genera originally belonging to the family Mesoblattinidae (Vishnyakova 1982). This
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across the whole Jurassic and Cretaceous (Vršanský 2002; Vršanský, Vishnyakova et Rasnitsyn
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2002; Vršanský et al. 2013). Blattulidae is considered to be related to Corydiidae (=Polyphagidae)
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(Vishnyakova 1982; Vršanský 1999). Thus, this family was accepted as a member of superfamily
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Corydioidea/ Polyphagoidea (Vršanský, Vishnyakova et Rasnitsyn 2002; Vršanský 2003, 2004a,
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2004b; Vršanský et Ansorge 2007; Lee 2016); while some works also treated it as a member of the
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Blattuloidea (Anisyutkin et Gorochov 2008; Vršanský 2005a, 2008b, 2009). It was a relatively
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dominating family, being widely distributed around the world (Vršanský 1999; Wang, Liang et
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Ren 2007). However, its diversity at the generic level is poor, with about 18 genera reported to
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date (Vishnyakova 1982; Vršanský 2003, 2005a, 2005b, 2009; Martins-Neto, Mancuso et Gallego
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2005; Cifuentes-Ruiz et al. 2006; Vršanský and Ansorge 2007; Wang et al. 2007; Wang, Liang et
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Ren 2007; Anisyutkin et Gorochov 2008). Most genera have low diversity, with only a single
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species recorded (monotypic genera). The relatively diverse genera are Blattula and Elisama, the
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former containing 44 species and the latter with 20 species (Clapham et Karr 2018).
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families), or isolated tegmina (abundant plesiomorphies and homoplasies present) is considered to
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be less reliable (Li et Huang 2018). Such a situation is also present in the study of Blattulidae.
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Genera such as Anablatta, Argentinoblattula, Xonpepetla, Svabula, and Orbablattula were
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described based on the tegmen (Martinez-Delclos 1993; Martins-Neto, Mancuso et Gallego 2005;
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Vršanský 2005b; Cifuentes-Ruiz et al. 2006; Martins-Neto, Gallego et Zavattieri 2007). Kridla
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was described based on the hind wing (Vršanský 2005a). Globula, Nula, and Blattulites were
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described from nymphs (Vishnykova 1982; Vršanský 2008a, 2009). The credibility of these genera
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Macaroblattula, and Ocelloblattula were described based on relatively well-preserved materials,
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leading to more informative studies (Vršanský 2003; Wang et al. 2007; Wang, Liang et Ren 2007;
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Anisyutkin et Gorochov 2008). Revisions of Blattulidae genera and species were also made using
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abundant materials and by re-examining the type specimens (Vršanský et Ansorge 2007; Wang et
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al. 2007; Lee 2016).
The records of Blattulidae in amber are poor, with four monotypic genera known, viz.
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Ocelloblattula in Lebanese amber (Anisyutkin et Gorochov 2008) (Ocelloblattula santanensis
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(Lee, 2016) described based on impression fossil from Brazil needs further confirmation), and
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Nula, Batola, and Globula in French ambers (Vršanský 2008a; Vršanský 2009). However, only
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Ocelloblattula was described based on an almost complete adult specimen. Batola nikolai was
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described based on an incomplete adult specimen and several nymphs and body fragments, but the
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adult specimen was designed as a paratype. Nula and Globula were totally based on nymphs,
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which are poor in classification and may produce difficulties in matching adults with nymphs in
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the future study.
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Herein, the authors report the first record of Blattulidae in Myanmar amber, with Huablattula
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gen. nov. described to include Huablattula hui sp. nov. and Huablattula jiewenae sp. nov., both of
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which are described based on well-preserved specimens. One Huablattula sp. is also presented for
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study but not described to species due to the insufficient characters present in the amber. The
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specimens presented herein all reveal some important characters to enrich our knowledge of the
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Blattulidae. These discoveries highlight the palaeodiversity of this family in the mid-Cretaceous,
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which suggests more taxa will be discovered in the near future.
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The amber pieces containing the specimens were obtained from the Hukawng Valley, Kachin
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State, in Northern Myanmar (Yin et al. 2018: fig. 1A, 26°21′33.41″N, 96°43′11.88″E). The age of
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Burmese amber is estimated as the Late Cretaceous (98.79±0.62 Ma) based on U-Pb dating of
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zircons (Shi et al. 2012). The studied specimens are deposited in the Institute of Entomology,
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Southwest University, Chongqing, China (SWU), some of which are previously in the personal
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collections of Zhengkun Hu, Guizhou, China (CZKH) and Dan Zuo, Hunan, China (CDZ). The
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amber pieces were polished using sandpaper of different grain sizes and with rare earth polishing
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powder to make them clear. All images (Figs. 1-6) were made using a Leica® M205A
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stereomicroscope plus Leica DFC 550 and Canon EOS 50D camera plus a Canon EF 100mm
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f/2.8L IS USM Macro lens combined with Helicon Focus software and modified in Adobe
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Photoshop CS6. Morphological terminology mainly follows Grandcolas (1994) and Roth (2003).
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Venation terms follow Li et al. (2018). This publication is registered in ZooBank under
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urn:lsid:zoobank.org:pub: 10AF1305-932C-41E3-9617-360E794383F2.
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3. Systematic paleontology
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Order Dictyoptera
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Family Blattulidae
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Genus Huablattula gen. nov.
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(urn:lsid:zoobank.org:act: F72A82F0-B379-4E3C-90A9-6CA195F22928)
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Type species: Huablattula hui sp. nov. here designated.
ACCEPTED MANUSCRIPT Description. Male: body small, elongate, sub-transparent, decorated with spots and stripes. Head
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broad, near triangular, not hidden by head; eyes round and large, strongly convex; interocular
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space greater than the distance between antennal sockets; the space between eyes and above
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antennal sockets distinctly prominent; three ocelli present (currently this character is only
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indicated in Huablattula jiewenae sp. nov.); antennae slender, exceeding the apex of tegmina,
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segments short, with sparse but distinct setae from antennomere 10 on, scape cylindrical, robust;
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clypeus convex but small; mouth constricted; maxillary palpomeres 3–4 slender, palpomere 5
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slightly expanded or rounded. Pronotum smooth, without pubescent, usually decorated with dark
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stripes, hind margin distinct protruded medially. Venation simple, decorated with dark stripes in
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tegmina, median of CuA with a large spot or irregular macula, which may extend to the distal
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margin of anal area. Tegmen with simple and long ScP, R branches sub-parallel, M and CuA
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usually with several branches; intercalary veins present between branches of R, M and CuA.
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Wings with dark venation; ScP simple, connected with RA near the base; RA with several short
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branches at distal half, the area among the short branches usually dark and thickened (forming
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pterostigma); RP simple with about 4 branches; M straight, bifurcate at distal half; CuA with 4–5
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parallel and curved branches; CuP very long and slender; Pcu with several small branches basally;
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anal area small, simply folded behind the rest part of the wing; intercalary veins present between
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RP, M and distal portion of CuA; cross veins present but sparse. Each femoral apex with a spine
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(extremely long at mid and hind femora), hind margins of femur with two rows of long but sparse
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spines; tarsal claw asymmetrical, arolia large. Cerci long, pubescent, distal segment long-spinous,
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or elongate. Subgenital plate protruded, pubescent, with a pair of segmented and elongate styli
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near the apex.
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ACCEPTED MANUSCRIPT Female: relatively larger than male, the habitus and stripes generally similar to male. Subgenital
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plate strongly bulging, apex constricted, forming a narrow valved protrusion, a pair of ovipositor
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processes extend externally, slender, apex curved and acute, base with a valved sheath.
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Etymology. Hua refers to the Chinese character 花, meaning piebald, for its body is decorated
115
with multiple stripes and spots. Hua + blattula therefore means “piebald blattulid cockroach”.
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Species included. Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. described herein.
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Also with one undescribed species mentioned below.
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Remarks. Huablattula gen. nov. resembles Ocelloblattula in the Lebanese amber (Anisyutkin et
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Gorochov 2008) in its general appearance, shape of head, globous eyes, shape of pronotum and
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legs; but it differs from the latter by the ocelli and its position, which are smaller and with the
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lateral two more separated and closer to the eyes. It can be also distinguished from the latter by its
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remarkable markings on pronotum and stripes on tegmina. The apical portion of subgenital plate
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as well as the ovipositor also easily distinguish it from the latter. This new genus differs from
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Habroblattula and Macaroblattula found in the Yixian Formation (Wang et al. 2007; Wang, Liang
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et Ren 2007) by the non-branched ScP. This new genus also resembles other impression fossil taxa,
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such as Vrtula, Elisama, Tarakanula, and Svabula by the markings on its tegmen, but differs from
127
Vrtula, Elisama, and Tarakanula by having wing pterostigma (Vršanský 2005a; Vršanský 2008b).
128
It differs from Svabula by having a more protruded and angular apex of tegmen (Vršanský 2005b),
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and from Kridla by not having a darkened apex of the wing (Vršanský 2005a). It can be
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distinguished from Blattula by having dark macula on the tegmen (Wang, Liang et Ren 2007;
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Vršanský et Ansorge 2007). This new genus can be distinguished from the adult paratype of
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Batola nikolai by the protruded apex of the subgenital plate (Vršanský 2009).
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ACCEPTED MANUSCRIPT 133 134
Huablattula hui sp. nov. (Figs. 1–4)
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(urn:lsid:zoobank.org:act: A3A7FC5C-C3EB-46FB-BC2B-412536F07045)
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136 Type material. Holotype. A well-preserved male adult in amber with opened right tegmen almost
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complete, apices of both antennae and right tegmen lost, parts of tarsi in right mid leg and both
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hind legs lost (SWU, ex CZKH, SWUMA-0002).
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Paratypes. A well-preserved male adult in amber with tegmina and wings in resting position,
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distal half of antennae missing, the joint part of femur and tibiae in left front and mid legs, apical
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portion of right femur and the whole tibiae and tarsus, apical portion of right tarsus, the whole
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tarsus of both hind legs missing (SWU, ex CDZ, SWUMA-0003).
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A sex unknown adult in amber, with tegmina and wings in resting position, ventral side
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unobservable (SWU, ex CDZ, SWUMA-0004).
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A sex unknown adult with tegmina and wings in resting position, well-preserved in amber, but the
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hind portion of the body is missing, including the distal half of left mid tibiae and the whole tarsus,
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the whole left leg, the whole right hind tarsus, abdomen tip, and tegmen and wing tips. A
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Hymenoptera insect is also preserved in the amber (SWU, ex CZKH, SWUMA-0005).
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Sex unknown adult fragments preserved in amber, with only tegmina, mesonotum, one antenna,
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one femur left. A broken Hymenoptera insect is also preserved in the amber (SWU, ex CZKH,
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SWUMA-0006).
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A female adult in amber, head broken. Two Diptera insects and a probably Neuroptera insect are
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also preserved in the amber (SWU, SWUMA-0007).
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ACCEPTED MANUSCRIPT Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
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Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
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Diagnosis. Huablattula hui sp. nov. resembles Huablattula jiewenae sp. nov., but it can be easily
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distinguished from the latter by the markings on pronotum, the outline of pronotum, and the
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stripes of tegmina; a large triangular plaque is found along the distal margin of anal area to the
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median portion of CuA on the tegmen of Huablattula hui sp. nov., while a large spot is found
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around median of CuA on that of Huablattula jiewenae sp. nov.
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Description. Holotype SWUMA-0002. Body elongate, transparent, decorated with markings and
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stripes (Fig. 1A–B). Length 8.7 mm, including tegmina and wings 11.0 mm.
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Head (face toward left; the ventral detail is difficult to observe) nearly triangular, with dark stripes and spots, length 1.93 mm. Eyes round and large, situated at distal corners, length 0.87 mm;
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interocular space greater than the distance between antennal sockets; space between eyes and
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above antennal sockets distinctly prominent; ocelli unclear in the amber. Antennae slender, distal
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portions missing (5.40 mm and 5.81 mm for the remaining portion of left and right, respectively);
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segments short, with sparse but distinct setae from antennomere 10 on; scape cylindrical, robust,
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length 0.42 mm; pedicellus nearly conical, length 0.14 mm, the distal margin expanded, width
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0.12 mm; the rest of antennomeres cylindrical, generally becoming longer and thinner toward
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apex. Clypeus small, slightly convex. Mouth constricted; maxillary palpomeres 1–2 subequal in
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length (about 0.13 mm), 3–5 lengths 0.41 mm/0.37 mm/0.35 mm, palpomeres 1–2 short, nearly
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globous, palpomeres 3–4 slender, palpomere 5 slightly expanded.
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Pronotum (Fig. 1H) transparent, smooth, without pubescent; disc with symmetrical dark
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markings that extend in two long stripes posteriorly, basal half decorated with some light-colored
ACCEPTED MANUSCRIPT spots; margins outlined with dark line. The margins of the pronotum generally rounded, anterior
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margin round, posterior margin protruding and constricted medially, lateral-hind parts roundly
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protruding. Length 2.00 mm (measured medially), width 2.46 mm (measured at widest points).
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Tegmina (Fig. 1D–-E) length 8.19 mm (left one), width 2.12 mm. Venation decorated with
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dark thick stripes, anal veins with reticulated stripes, a large triangular plaque lies along the distal
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margin of anal area to the median portion of CuA. ScP simple, R branches sub-parallel, M
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bifurcate near base, CuA with 4 branches; intercalary veins present between branches of R, M and
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CuA (based on the opened right tegmen). Wings (Fig. 1F–G) with dark venation, apex narrow;
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ScP simple, connected with RA at 1/3 from the base; RA with 4 short branches at distal half, the
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area among the 4 branches dark and thickened (with pterostigma); RP bifurcate basally, distal
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portion with three branches; M straight, with a bifurcate branch at 2/3 from the base; CuA with 4
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parallel and curved branches; CuP slender; Pcu with three small branches basally. Intercalary veins
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present between RP, M and distal portion of CuA (based on the folded right wing).
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Legs mixed with dark maculae. Front leg: femur 1.69 mm, tibia 1.20 mm, tarsus 1.46 mm (tarsal
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segments 1–5 lengths 0.58 mm/0.28 mm/0.21 mm/0.16 mm/0.27 mm); mid leg: femur 2.42 mm,
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tibia 2.12mm, tarsus 1.62 mm (tarsal segments 1–5 lengths 0.81 mm/0.26 mm/0.18 mm/0.14
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mm/0.27 mm); hind leg: femur 2.55 mm, tibia 3.01 mm, tarsus missing. Each leg with a long
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spine at the apex of femur, and two rows of long but sparse spines at hind margins. Length of the
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apical spines from front, mid and hind femora 0.26 mm/0.70 mm/1.04 mm (extremely long for
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those of mid and hind legs). Tarsal claw asymmetrical. Arolia large (Fig. 1C).
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Meso- and metanotum dark dorsally. Cerci long, length 2.09 mm, pubescent, apical one spinous.
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Subgenital plate protruding, pubescent, slightly asymmetrical distally, median of apex with a small
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ACCEPTED MANUSCRIPT protrusion, two long styli situated laterally beside the protrusion, segmented, basal segment not
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enlarged, length 1.05 mm (Fig. 1I–J).
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Paratype SWUMA-0003. Length 7.26 mm, including tegmina and wings 8.99 mm, width 3.46
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mm. Generally similar to the holotype (Fig. 2A–B). The head can be better observed, length of
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head 1.72 mm, width of head 1.75 mm. The markings on face as in Fig. 2C figured. Due to the
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head being nearly transparent, the ocelli are still difficult to recognize, only left ocellus is distinct,
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situated next to the middle margin of the eye. Subgenital plate, styli and cerci similar to those of
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the holotype (Fig. 2D).
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Paratype SWUMA-0004. Length including tegmina 9.76 mm, width including tegmina 4.03 mm
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(Fig. 2E).
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Paratype SWUMA-0005. The antenna is complete in this specimen, the total length of the
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antenna is 9.19 mm, which is distinctly longer than the body length of this species. Right ocellus
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can be recognized in this specimen (Fig. 2F–G).
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Paratype SWUMA-0006. Length of left tegmen 8.20 mm, width 2.45 mm. Since the holotype
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specimen slightly overlaps in the amber (probably due to the struggling of the insect), the venation
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of the holotype is not as clearly observed. From this specimen, the intercalary veins can be better
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recognized (Figs. 3A–F).
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Paratype SWUMA-0007. The only known female (Fig. 4A–B). Body length from antetior
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margin of pronotum to tegmina tip 12.9 mm, length from antetior margin of pronotum to the apex
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of ovipositor valves 14.7 mm. Pronotum length 2.68, width 2.89 mm. The marking on pronotum is
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similar to male, but shows some reduction (Fig. 4C). Subgenital plate (Fig. 4D) large, strongly
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bulging and protruded, with a narrow and valved posterior lobe. External ovipositor present (Fig.
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ACCEPTED MANUSCRIPT 4E) consisting of two processes and a sheath: a pair of very elongate, flat and pubescent processes
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extend from the posterior lobe, apical portions of both processes curved and with needle-like apex;
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a valved sheath extends from the posterior lobe dorsally, and covers the base of the two long
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processes.
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Etymology. Named after Mr. Zheng-Kun Hu (Guizhou, China) who kindly donated his amber
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collection to us for study.
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227 Huablattula jiewenae sp. nov. (Fig. 5)
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(urn:lsid:zoobank.org:act: CC3A1364-8C8F-4A69-8581-992109A1D6B4)
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Type material. Holotype. A well preserved male adult in amber with tegmina and wings in resting
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position, apices of antennae missing, median of left antenna missing, mouth parts broken, the
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jointed portions of femur and tibia are missing for front legs, apex of tarsi are missing for mid legs,
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left tibia broken, tarsus missing, right tibia with broken tibia. Two Diptera insects and a small
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Coleoptera insect are also preserved in the amber (SWU, ex CDZ, SWUMA-0008).
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Paratype. A sex unknown adult preserved in amber with tegmina and wings in resting position,
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mostly complete, but ventral side unobservable. A Hymenoptera insect, a Staphylinidae beetle and
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a Eucnemidae beetle are also preserved in the amber (SWU, ex CZKH, SWUMA-0009).
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Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
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Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
241
Diagnosis. See the diagnosis of Huablattula hui sp. nov.
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Description. Holotype SWUMA-0008. Body elongate, transparent, decorated with stripes and
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ACCEPTED MANUSCRIPT spots (Fig. 5A–B). Length 7.2 mm, including tegmina and wings 9.3 mm, width 4.3 mm (the
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tegmina in the amber are slightly opened, thus this measured body width may be wider than
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actual).
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Head nearly triangular, vertex with three longitudinal stripes, length 1.38 mm (the head is upward
247
in the amber, thus this measurement is shorter than usual), width 1.80 mm. Eyes round and large,
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situated at distal corners; interocular space greater than the distance between antennal sockets
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(0.76 mm versus 0.37 mm); the space between eyes and above antennal sockets prominent, a dark
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band transversely across the prominent portion; three ocelli present, situated on the transverse
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band (two near the eyes, one in the center, intervals between the three ocelli about 0.3–0.4 mm).
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Antennae slender, median part of left one missing, apex of right one missing, measurement for the
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remaining parts is difficult; segments short, with sparse but distinct setae from antennomere 10 on;
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scape cylindrical, length 0.26 mm; pedicellus nearly cylindrical, length 0.17 mm; the rest of
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antennomeres cylindrical, generally becoming longer toward apex. Clypeus broken. Mouth parts
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constricted; maxillary palpomeres broken, palpomeres 4–5 length 0.29 mm/0.27 mm, palpomere 5
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roundly enlarged (Fig. 5D–E).
258
Pronotum smooth, without pubescent, lateral margins transparent; nearly pentagonal, anterior
259
margin nearly straight, lateral portions and hind margin protruding; four longitudinal bands cross
260
throughout from the anterior margin to the posterior margin, the lateral two thicker than the inner
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two; margins of pronotum outlined with a dark line very narrowly. Length 2.03 mm (measured
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medially), width 2.20 mm (measured from widest point) (Fig. 5D).
263
Tegmina length 7.29 mm, width 2.23 mm. Venation decorated with dark stripes, median of CuA
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with a large irregular spot. ScP simple, R branches sub-parallel, distal two with multiple branches,
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ACCEPTED MANUSCRIPT M bifurcate medially, CuA with 4 branches; intercalary veins present between branches of R, M
266
and CuA (Fig. 5H). Wings unobservable, apex narrow.
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Legs mixed with some small spots. Front leg: the jointed portion of femur and tibia missing for
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both, tarsus 1.17 mm (tarsal segments 1–5 lengths 0.47 mm/0.22 mm/0.17 mm/0.13mm/0.24mm);
269
mid leg: femur 1.79 mm, tibia 1.68 mm, tarsus with apical portion missing for both (tarsal
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segments 1–3 lengths 0.63 mm/0.27 mm/0.17 mm); hind leg: femur 2.20 mm, tibia 3.06 mm, left
271
tarsus with apical portion missing, right tarsus all missing (tarsal segments 1–2 lengths 0.82
272
mm/0.41 mm). Each femur with two rows of long but sparse spines at hind margins, mid and hind
273
femora with extremely long spines at apex. Tarsal claw asymmetrical. Arolia large.
274
Cerci long, length 1.79 mm, pubescent, apical two elongate. Subgenital plate protruded, pubescent,
275
apex unobservable, styli segmented, length 1.06 mm, basal segment not enlarged (Fig. 5F).
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Paratype SWUMA-0009. Length including tegmina and wings 9.60 mm. Generally similar to the
277
holotype (Fig. 5C), antenna exceeding the tip of tegmina. Tarsal claw asymmetrical (Fig. 5G).
278
Etymology. Named after Ms. Jie-Wen Zhao (Hunan, China). Her mother Ms. Dan Zuo kindly
279
provided her ambers for this study and hopes this honor inspires her daughter’s interest in natural
280
history.
282 283
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Huablattula sp. (Fig. 6)
284
Material examined. Male adult fragments preserved in amber, only wings, metanotum, tergum 1
285
and subgenital plate left (SWU, SWUMA-0010).
286
Locality and horizon. Hukawng Valley, Tanai Township, Myitkyina District, Kachin State,
ACCEPTED MANUSCRIPT Myanmar; the lowermost Cenomanian (98.79 ± 0.62 Ma), mid-Cretaceous (Shi et al. 2012).
288
Description. Wing (Fig. 6A–D) rounded, length 6.82 mm, apical portion not narrowed, relatively
289
wider than that of H. hui sp. nov.; ScP simple, connected with RA at 1/3 from the base; RA with
290
3–4 short branches at distal half, the area among the branches dark and thickened (with
291
pterostigma); RP with 4–5 parallel branches; M simple, bifurcate at distal half; CuA with 5
292
parallel and curved branches; CuP very long; Pcu with two small branches basally, distal one
293
bifurcate; intercalary veins present between RP, M and CuA; anal area simply folded once,
294
overlapped behind the remaining part of the wing. Subgenital plate similar to that of H. hui sp.
295
nov., with a small protrusion at apex, two basal lobes of the subgenital plate present (this character
296
is the same as in extant cockroaches); stylus segmented, length 1.28 mm, pubescent, basal
297
segment slightly enlarged (Fig. 6E).
298
Remarks. This specimen has a wider wing, the apex is rounded, the basal segment of stylus is
299
slightly enlarged. Thus we definitely consider it to be a different species from H. hui sp. nov. and
300
H. jiewenae sp. nov., but since no complete specimen is available, we temporarily treat it as an
301
undescribed species.
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4. Discussion
304
Although the ocelli of H. hui sp. nov. cannot be clearly observed, Huablattula hui sp. nov. and
305
Huablattula jiewenae sp. nov. are considered congeners by their general appearance,
306
triangular-shaped head, globous and large eyes, shape of pronotum, the long spines on legs,
307
asymmetrical tarsal claws, protruded subgenital plate, segmented and elongate styli, their similar
308
venation, the pronotum with markings, and the tegmina decorated with dark stripes in both
ACCEPTED MANUSCRIPT 309
species. The external ovipositor in this family has been reported from some impression fossils
311
(Vishnykova 1982; Wang et al. 2007). Before this research, the holotype of Ocelloblattula
312
ponomarenkoi was the only one where the ovipositor was observed in amber (Anisyutkin et
313
Gorochov 2008). This newly described female of Huablattula hui gen. nov. et sp. nov. is the
314
second amber record of the ovipositor in this family. Compared to O. ponomarenkoi, H. hui sp.
315
nov. has a different subgenital plate, which is valved posteriorly (while rounded posteriorly in O.
316
ponomarenkoi). The apical portions of the ovipositor processes are also different (needle-like and
317
roundly curved in H. hui sp. nov., while acuminate and extended caudad to the apex of posterior
318
lobe of genital plate in O. ponomarenkoi). The two ovipositor processes are much more extended
319
in H. hui sp. nov. than in O. ponomarenkoi. A valved sheath is present in H. hui sp. nov., but seems
320
not to be indicated in O. ponomarenkoi.
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From the delicate body, large eyes, long antennae, well-developed wings and the gorgeous
322
stripes on the pronotum and tegmina, Huablattula gen. nov. is believed to be an active insect in
323
daytime. The slender legs (with long spines), large arolia and weak and asymmetrical tarsal claws
324
indicate that this genus inhabited the leaves of trees.
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5. Conclusion
327
This paper presents the first record of the extinct cockroach family Blattulidae in Upper
328
Cretaceous Burmese amber, with description of Huablattula gen. nov. to accommodate two new
329
species, Huablattula hui sp. nov. and Huablattula jiewenae sp. nov. Most of the studied material is
330
well-preserved, enriching our knowledge of Blattulidae.
ACCEPTED MANUSCRIPT 331 Acknowledgments
333
We thank Mr. Zheng-Kun Hu (Guizhou, China) and Ms. Dan Zuo (Hunan, China) for providing
334
these precious amber materials. We also give our thanks to John Richard Schrock (Department of
335
Biological Sciences, Emporia State University) for revising the manuscript before submission.
336
Anonymous reviewers are acknowledged for their valuable comments and suggestions to the
337
manuscript. This work has been supported by the National Natural Sciences Foundation of China
338
(Nos. 31772506), Program of Ministry of Science and Technology of the People’s Republic of
339
China (2015FY210300), and Natural Science Foundation Project of Chongqing (No.
340
cstc2016jcyjA0487).
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341 References
343
Anisyutkin, L.N., Gorochov, A.V., 2008. A new genus and species of the cockroach family
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Blattulidae from Lebanese amber (Dictyoptera, Blattina). Paleontologicheskii Zhurnal 1, 44–
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47.
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Clapham, M., Karr, J., 2018. Taxonomic occurrences of Blattula and Elisama recorded in the
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Polyphagidae). Systematic Entomology 19, 145–158. Lee, S.-W., 2016. Taxonomic diversity of cockroach assemblages (Blattaria, Insecta) of the Aptian Crato Formation (Cretaceous, NE Brazil). Geologica Carpathica 67, 433–450. Li, X.R., Huang, D., 2018. A new Cretaceous cockroach with heterogeneous tarsi preserved in
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from the Shar-Teg in Mongolia. Geologica Carpathica 55, 457–468. Vršanský, P., 2005a. A fossil insect in a drilling core sample - cockroach Kridla stastia gen. et sp. nov. (Blattulidae) from the Cretaceous of the Verkhne-Bureinskaya Depression in eastern
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Russia. Entomological Problems 35, 115–116.
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Vršanský, P., 2005b. Lower Cretaceous cockroaches and mantids (Insecta: Blattaria, Mantodea) from the Sharin-Gol in Mongolia. Entomological Problems 35, 163–167.
Vršanský, P., 2008a. A complete larva of a Mesozoic (Early Cenomanian) cockroach (Insecta:
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Blattaria: Blattulidae) from the Sisteron amber (Alpes de Haute Provence, SE France).
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Geologica Carpathica 59, 269–272.
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Vršanský, P., 2008b. New blattarians and a review of dictyopteran assemblages from the Lower
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Cretaceous of Mongolia. Acta Palaeontologica Polonica 53, 129–136. Vršanský, P., 2009. Albian cockroaches (Insecta, Blattida) from French amber of Archingeay. Geodiversitas 31, 73–98. Vršanský, P., Ansorge, J., 2007. Lower Jurassic cockroaches (Insecta: Blattaria) from Germany and England. African Invertebrates 48, 103–126.
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Vršanský, P., van de Kamp, T., Azar, D., Prokin, A., Vidlicka, L.U., Vagovic, P., 2013.
Cockroaches probably cleaned up after dinosaurs. PLoS ONE 8, e80560, 80561–80511.
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Vršanský, P., Vishnyakova, V.N. & Rasnitsyn, A.P., 2002. Order Blattida Latreille, 1810. The
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cockroaches. In: Rasnitsyn, A.P. & Quicke, D.L.J. (eds.). History of Insects. Kluwer
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Academic Publisher, Dodrecht, 263–271.
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Wang, T., Ren, D., Liang, J.-H., Shih, C., 2007. New Mesozoic cockroaches (Blattaria: Blattulidae)
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from Jehol Biota of western Liaoning in China. Annales Zoologici (Warsaw) 57, 483–495.
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Wang, T.-T., Liang, J.-H., Ren, D., 2007. Variability of Habroblattula drepanoides gen. et. sp. nov.
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(Insecta: Blattaria: Blattulidae) from the Yixian Formation in Liaoning, China. Zootaxa 1443,
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17–27.
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Yin, Z., Cai, C., Huang, D., 2018. Last major gap in scydmaenine evolution filled (Coleoptera:
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Staphylinidae), Cretaceous Research 84, 62–68. doi: 10.1016/j.cretres.2017.10.026.
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Fig. 1. Huablattula hui sp. nov., habitus and detailed characters of holotype (SWUMA-0002),
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male. A. Habitus, dorsal view. B. Habitus, ventral view. C. Tarsal claws, rear view. D–E. Right
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tegmen (intercalary veins are omitted in fig. E). F–G. Right wing. H. Pronotum (outlined). I.
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Abdominal tip, ventral view. J. Detail of right stylus (arrows indicate the segmented parts).
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Abbreviations: arolium (ar), cercus (ce), stylus (st), subgenital plate (sg), tarsal claw (tc),
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tarsomere 5 (t5). Scale bars: A–B = 2 mm; C = 0.1 mm; D–G = 1 mm; H–I = 0.5 mm; J not to
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scale.
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(SWUMA-0003~0005). A–B. Habitus and detailed characters of SWUMA-0003, male; A. Dorsal
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view; B. Ventral view; C. Head; D. Abdominal tip. E. Habitus of SWUMA-0004, sex unknown,
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dorsal view. F–G. Habitus of SWUMA-0005, sex unknown; F. Dorsal view; G. Ventral view.
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Abbreviations: cercus (ce), eye (ey), ocellus (oc), stylus (st), subgenital plate (sg). Scale bars: A–
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B, E–G = 2 mm; C–D = 0.5 mm.
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Fig. 3. Huablattula hui sp. nov., detail characters of paratype (SWUMA-0006). A. General
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condition of the amber; B. Basal portion of the antennal fragment; C. Right tegmen; D. Leg
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fragment (show tibiae and the spines); E–F. Left tegmen. Scale bars: A = 2 mm; B, D = 0.5 mm; C,
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E–F = 1 mm.
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Fig. 4. Huablattula hui sp. nov., habitus and detail characters of paratype (SWUMA-0007), female.
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A. Habitus, dorsal view; B. Habitus, ventral view; C. Pronotum; D. Abdominal tip; E. Apical
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portion of subgenital plate and ovipositor. Abbreviations: cercus (ce), subgenital plate (sg),
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ovipositor (ov), bubble (bu), sheath (sh), ovipositor process (pr), posterior lobe (pl). Scale bars:
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A–B, D = 2 mm; C = 1 mm; E = 0.5 mm.
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Holotype (SWUMA-0008), male; A. Habitus, dorsal view; B. Habitus, ventral view; C. Paratype
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(SWUMA-0009), sex unknown, dorsal view; D. Head and pronotum, holotype, dorsal view; E.
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Head (arrows indicate the ocelli), holotype, ventral view; F. Abdominal tip, holotype, ventral view;
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G. tarsal claw, paratype, dorsal view; H. Right tegmen, holotype. Abbreviations: arolium (ar),
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cercus (ce), stylus (st), subgenital plate (sg), tarsal claw (tc), tarsomere 5 (t5), tibiae (ti). Scale bars:
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A–C, H = 2 mm; D–F = 0.5 mm; G = 0.1 mm.
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Fig. 6. Huablattula sp. (SWUMA-0010), male. A–B. General condition of the amber; C–D.
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Wings; E. Subgenital plate. Abbreviations: basal lobe (bl), protrusion (pr), stylus (st), subgenital
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plate (sg). Scale bars: A = 2 mm; B–E = 1 mm.
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