New Data on the Family Blaberidae (Dictyoptera) from Southeast Asia
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ZOOSYSTEMATICA ROSSICA, 24(1): 14–30 25 JUNE 2015 New data on the family Blaberidae (Dictyoptera) from Southeast Asia: new species, morphological diversity and phylogeny on the base of ribosomal DNA sequences Новые данные о семействе Blaberidae (Dictyoptera) из Юго-Восточной Азии: новые виды, морфологическое разнообразие и филогения на основании последовательностей рибосомальной ДНК V.A. MAVROPULO, L.N. ANISYUTKIN*, M.V. ZAGOSKIN, A.S. ZAGOSKINA, S.V. LUKYANTSEV & D.V. MUKHA В.А. МАВРОПУЛО, Л.Н. АНИСЮТКИН, М.В. ЗАГОСКИН, А.С. ЗАГОСКИНА, С.В. ЛУКЬЯНЦЕВ, Д.В. МУХА V.A. Mavropulo, Vavilov Institute of General Genetics, Russian Academy of Sciences, 3 Gubkin Str., Moscow, 119991, Russia L.N. Anisyutkin, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia. E-mails: [email protected], [email protected]. Corresponding author M.V. Zagoskin, Vavilov Institute of General Genetics, Russian Academy of Sciences, 3 Gubkin Str., Moscow 119991, Russia A.S. Zagoskina, Vavilov Institute of General Genetics, Russian Academy of Sciences, 3 Gubkin Str., Moscow 119991, Russia S.V. Lukyantsev, Tomsk State University, 36 Lenina Str, Tomsk 634059, Russia D.V. Mukha, Vavilov Institute of General Genetics, Russian Academy of Sciences, 3 Gubkin Str., Moscow 119991, Russia. E-mail: [email protected] New cockroaches of the family Blaberidae are described from Southern Sumatra: two new spe- cies of the genus Cyrtonotula Uvarov, 1939, C. secunda sp. nov. and C. tertia sp. nov. (Epilam- prinae); and one new species of the genus Paranauphoeta J.W.H. Rehn, 1951, P. pullata sp. nov. (Paranauphoetinae). Detailed morphological descriptions of the new species are given. Struc- tures of the male genitalia of the genus Cyrtonotula are described for the first time. Hypothesis on the relationships of these new taxa as well as Morphna maculata Brunner von Wattenwyl, 1865, Rhabdoblatta sp. and Pseudophoraspis sp. based on 28S ribosomal DNA sequences is discussed. Из Южной Суматры описываются новые тараканы семейства Blaberidae: два новых вида рода Cyrtonotula Uvarov, 1939, C. secunda sp. nov. и C. tertia sp. nov. (Epilamprinae), и один новый вид из рода Paranauphoeta J.W.H. Rehn, 1951, P. pullata sp. nov. (Paranauphoetinae). Для новых видов дается детальное морфологическое описание. Строение гениталий сам- цов впервые описывается у представителей рода Cyrtonotula. Обсуждается выдвинутая на основании изучения 28S рибосомальной ДНК гипотеза о родственных отношениях новых таксонов, а также Morphna maculata Brunner von Wattenwyl, 1865, Rhabdoblatta sp. и Pseudophoraspis sp. Key words: cockroaches, taxonomy, morphology, phylogeny, 28S ribosomal DNA, Southeast Asia, Dictyoptera, Blaberidae, Epilamprinae, Paranauphoetinae, Cyrtonotula, Paranauphoeta, new species Ключевые слова: тараканы, таксономия, морфология, филогения, 28S рибосомальная ДНК, Юго-Восточная Азия, Dictyoptera, Blaberidae, Epilamprinae, Paranauphoetinae, Cyr- tonotula, Paranauphoeta, новые виды © 2015 Zoological Institute, Russian Academy of Scienсes V.A. MAVROPULO ET AL. NEW DATA ON BLABERIDAE FROM SOUTHEAST ASIA 15 INTRODUCTION The ratio of “distance between eyes” to “length of eye” was measured as the in- In the present paper, new representatives terocular distance on the vertex (i.o. in Fig. of the family Blaberidae Brunner von Wat- 1A) to the dorsoventral length of eye (d.e. tenwyl, 1865 are described from Southeast in Fig. 1A). Types of the anterior margin of Asia. There are two new species of the genus fore femur armament are given according to Cyrtonotula Uvarov, 1939, belonging to the Bey-Bienko (1950) and Roth (2003). The subfamily Epilamprinae Brunner von Wat- terminology of the male genital sclerites fol- tenwyl, 1865, and one new species of Pa- lows Klass (1997) but with some modifica- ranauphoeta Brunner von Wattenwyl, 1865, tions (Anisyutkin, 2014; Anisyutkin et al., belonging to the subfamily Paranauphoeti- 2013). The terminology used by Grandcolas nae J.W.H. Rehn, 1951. Ribosomal DNA se- (1996) for genital structures is given in pa- quence data of the described taxa and males rentheses following the author’s designa- of the genus Cyrtonotula are considered for tions. The terms introduced by the author the first time. A provisional hypothesis re- are given in quotation marks. garding the relationships in the studied taxa The material studied (including type is proposed on the base of the consensus specimens) is deposited at the Zoological cladogram that was inferred from the 28S Institute of the Russian Academy of Sci- ribosomal DNA fragment sequence data. ences, St Petersburg, Russia. The subfamily Epilamprinae is one of the largest and most diverse subfamilies Abbreviation used in figures of Blaberidae. It is widely distributed in (see text for further details): regions with wet tropical climate and is a.s., “additional spines”, i.e. spines bor- known since at least the Paleocene epoch dering euplantulae from inside and outside; š (Vr ansky et al., 2013). The subfamily Pa- ap.scl., “apical sclerite” of sclerite L2D in ranauphoetinae includes a single genus of the male genitalia; uncertain systematical position, Paranau- b.hla., basal membranous part of sclerite phoeta. It undoubtedly belongs to Blaberi- L3; dae, but its relationships within the fam- b.L2D, basal part of sclerite L2D in the ily are unclear (for details see: Anisyutkin, male genitalia; 2003a). There are no paleontological data b.L3, basal subsclerite of sclerite L3 in on Paranauphoeta yet. the male genitalia; c.b.m.a., “chaetae-bearing membranous MATERIAL AND METHODS area” located between right phallomere and sclerite L2D; The material studied was collected in c.b.m.l., “chaetae-bearing membranous tropical forests at light or during night lobe” located above basal part of sclerite work with the use of a flashlight on the L2D; leaves of trees and bushes. The material was c.p.R1T, caudal part of the sclerite R1T preserved in 70% ethanol. in the male genitalia; The male genitalia were processed with cr.p.R1T, cranial part of the sclerite R1T alkali by means of a standard procedure in the male genitalia; (Anisyutkin et al., 2013) for 12–24 hours d.e., dorsoventral length of the eye; for maceration of the soft tissues. The il- f.s., “folded structure” of the sclerite L3 lustrations were sketched by means of a in the male genitalia; drawing tube on a Leica MZ 16 binocular hge., groove of the sclerite L3 in the male microscope; further drawing and examina- genitalia (= hge); tion were made with an MBS–10 binocular hla, sclerotized apical part of the sclerite microscope. L3 in the male genitalia (sensu Klass, 1997); © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(1): 14–30 16 V.A. MAVROPULO ET AL. NEW DATA ON BLABERIDAE FROM SOUTHEAST ASIA hl., hollow on the sclerite R2 in the male QIAquick Gel Extraction Kits (QIAGEN, genitalia; Hilden, Germany). Cleaned products were i.o., interocular distance on the vertex; cloned into pGEM-T Easy vectors (Pro- L2D, L3, L4U, sclerites in the male geni- mega), and transformed into Escherichia coli talia; JM109 competent cells (Promega) follow- Par., paraproct; ing the manufacturer’s protocol. Amplified pr.s., preputial spines located at the “api- DNA fragments and several clones were se- cal sclerite” of sclerite L2D of the male geni- quenced for each specimen. talia; Automated sequences were generated pv., sclerite at the base of the cerci (= pv- on an ABI PRISM® 310 Genetic Analyzer sclerite); according to Sanger et al. (1977) with a Big- r.plm., right phallomere of the male geni- Dye Termination kit (Applied Biosystems). talia; Opposite strands were confirmed for all R1T, R2, R3, R4, R5, sclerites in the male templates. ABI trace files were edited and genitalia. contigs were assembled using the program ChromasPro v. 1.7.6 (http://technelysium. DNA analysis com.au/?page_id=27). DNA extraction, PCR amplification Nucleotide alignment and phylogenetic and sequencing analysis Total genomic nucleic acids were ex- Sequences from this study for Paranau- tracted from ethanol-fixed insects using phoeta pullata sp. nov., Cyrtonotula tertia a standard DNA extraction buffer (Tris– sp. nov., C. secunda sp. nov., Pseudophoras- HCl, proteinase K, SDS) according to the pis sp. and Rhabdoblatta sp. are under the protocol described in Mukha et al. (1995). accession numbers KJ194457, KJ184844, DAMS-18 (gtccctgccgtttgtacaca) and KJ194458, KJ194459 and KJ194460, re- DAMS-28 (ctactagatggttcgattagtc) prim- spectively. Furthermore for the reconstruc- ers were used for DNA amplification and tion of the phylogenetic tree, the following sequencing. The priming sites for DAMS- rDNA sequences were used: Archiman- 18 and DAMS-28 are highly conserved in drita tesselata J.A.G. Rehn, 1903 (acc. # eukaryotes; these primers are “universal” GU384924), Blaberus atropos (Stoll, 1813) and may be used for the amplification of the (acc. # AF321252), Morphna maculata (acc. corresponding ribosomal DNA fragments # KM659175), Symploce pallens (Stephens, across a broad taxonomic range of organ- 1835) (acc. # KF922887), Parathemnop- isms (Mukha & Sidorenko, 1995, 1996; teryx couloniana (Saussure, 1863) (acc. # Mukha et al., 2000; Mukha et al., 2002). KF922888), Parcoblatta lata (Brunner von PCR amplification of the analyzed rDNA Wattenwyl, 1865) (acc. # AF321247), Blat- fragments was carried out using Master Mix tella vaga Hebard, 1935 (acc. # AF321246), (2X) (Fermentas, Vilnius, Lithuania) ac- Blattella lituricollis (Walker, 1868)