Mesozoic relative of the common synanthropic German cockroach (Blattodea)

Mesozoic relative of the common synanthropicGerman cockroach (Blattodea)

Peter Vrsansky*

Geological Institute, Slovak Academy of Sciences, D�bravska c. 9, P.O. BOX 106, 840 05 Bratislava, Slovakia andPaleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow, Russia

Introduction

Throughout the over 350 Myr history of the order,cockroach taxa were mostly rather uniform, providingthe decomposition function in diverse ecosystems.Their diversity has usually been moderate, rarely low.The Early Cretaceous assemblage from the Shin-Khu-dukh in Mongolia with a single dominant species(Vrsansky 2008a); and rather high diversity assem-blages from Mongolia and Kazakhstan (Middle Jurassicof Bakhar Bed 275/1: 16 species, 319 specimens; EarlyCretaceous of Bon Tsagaan: 20 species, 646 specimens;Late Jurassic of Karatau: 3 different assemblages with72 species, 2,000 specimens (Vrsansky 2003a)) are un-ique.

Deviating forms have been restricted to a few raretaxa, like the aquatic (Takahashi 1926) Opisthoplatiaorientalis (Burmeister, 1838), two semiaquatic species(Shelford 1909); the single bioluminescent genus Luci-hormetica (Zompro & Fritzsche 1999); the exclusiveblattelid pollinator of Uvaria elmeri (Nagamitsu & In-oue 1997); the translucent troglobitic Spelaeoblattamyugei (Vidlicka et al. 2003); the single Late Jurassic

jumping Skok svaba (Vrsansky 2007); a single earwig-like species from the Middle Jurassic of China repre-senting a separate family (Vrsansky et al. submitted);the metallic Oxyhaloa buprestoides (Saussure 1862);the monogeneric semisocial Cryptocercidae Brues &Melander, 1932 (Steinmiller 2001; Nalepa 2003;Grandcolas et al. 2005); a single recorded eusocialcockroach from the Albian French amber (P. Vrsansky,submitted); a single ovipositor-bearing living cockroach(Vidlicka & Vrsansky, in preparation); but also therather common carnivorous (Vishniakova 1973) LateJurassic-Late Cretaceous Raphidiomimidae Vishniako-va, 1973, the beetle-like Late Jurassic-Late CretaceousUmenocoleidae Chen et Tian, 1972 (Vrsansky 2003b),and descendants of cockroaches: the predatory mantises(Vrsansky 2002) and eusocial termites (Vrsansky et al.2002).

The wide historical distribution of cockroaches, thatwere dominant in diverse Paleozoic as well as Meso-zoic localities, mostly in warm, tropical and subtropicalzones (Schneider 1978a, 1978b, 1980a, 1983b, 1983;Vrsansky et al. 2002) is in contrast to their prevailingconservativeness.

Dtsch. Entomol. Z. 55 (2) 2008, 215–221 / DOI 10.1002/mmnd.200800022

museum fur naturkundeder Humboldt-Universita t zu Berlin

# 2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Received 7 May 2008Accepted 17 June 2008Published 5 November 2008

Key Words

Fossil insectsMesozoic amberBlattida ¼ Blattaria ¼ BlattodeaLiving genusBlattella germanica

Abstract

Cockroaches, with an evolutionary history going back 350 Myr and with over100,000 fossil specimens collected so far, form one of the most consistent fossil re-cords in terrestrial arthropods. In addition to their descendants, the eusocial termitesand predatory mantises, their variability is presented by such diverse forms as biolumi-nescent, somatically translucent, beetle-like, predatory, aquatic, semi-social, eusocialand viviparous species. In spite of their conservativeness at higher taxonomic levels,the evolutionary tempo of cockroach species is comparatively high. The modern fa-milies of cockroaches only appear as early as the Cretaceous, and the oldest taxonclosely resembling a living genus described here from the Early Cenomanian (ca.96 Ma) French amber greatly increases, by 46 Myr, their expected antiquity. ?Blattella

lengleti sp. n. – a close relative of a common synanthropic German cockroach, indi-cates that this genus and/or its very close relative shared environments with dinosaurs,almost 100 Myr before it occupied human households.

* E-mail: [email protected]

Among living cockroaches, the most widely distribu-ted is the common synanthropic German cockroach Blat-tella germanica. Its genome length is comparable to thehuman one (Bier & M�ller 1969), with peculiar homeo-tic mutant 10 Pw, which is a reciprocal translocation ofchromosomes 9 and 10 (in heterozygotes; homozygotesare lethal), characterized by pronotal extensions resem-bling a third pair of wings (Tanaka & Ito 1997) – sug-gesting that a shift from 3-pairs of wings in the most pri-mitive extinct insects to 2-pairs in living insects mighthave been caused by a single chromosomal translocation.

The German cockroach is extremely resistant to en-vironmental stress and irradiation (Koval 1983) and ithas also been used for experiments in orbit. Insecticidesspent against Blattella germanica reach a billion dollarsannually.

The genus Blattella comprises 51 rather uniform de-scribed species (Brenner et al. 1993; Roth 1995, 2003).Its cosmopolitan distribution, contrasting with a conser-vative morphological standard, might be explained by along history without major evolutionary changes:hitherto the oldest representatives were known from theEocene (50 Myr) Baltic amber (Poinar & Poinar 1992)and also from Eocene sediments of the Green RiverFormation of Colorado, U.S.A. (P. Vrsansky, L’. Vi-dlicka & C. Labandeira, in preparation), characterizedby subtropical fauna and flora, where the genus was al-ready highly diversified.

Cockroaches are rather frequent fossils in Mesozoicamber though there are only a very few described spe-cimens (Tab. 1). Only two adults and two immatures

are described from the Lebanese and New Jersey am-bers (Vrsansky 2003b, 2004; Anisyutkin & Gorochov2008) and two immatures from the Burmese (Grimaldi& Ross 2004) and Sisteron ambers (Vrsansky 2008b).

Systematic paleontologyBlattaria Latreille, 1810

Blattellidae Karny, 1908

Blattella Caudell, 1903

Type species. Blattella germanica Linnaeus, 1767: 688; Caudell 1903:234; Melville 1982: 243.

Diagnosis (Roth 1985). Tegmina and wings usually fullydeveloped, but sometimes they may be reduced; . . . firstabdominal tergite of male always unspecialized. Abdom-inal tergite 7 (T7), or tergites 7 and 8 specialized (tergalglands). Female abdominal tergites unspecialized. Malesubgenital plate weakly or distinctly asymmetrical. Stylesvariable. In female, subgenital plate usually convex, hindmargin truncate, or slightly rounded, rarely excised medi-ally (1 species). Front femur with anteroventral marginarmed with spines which are smaller in size distad, pos-teroventral margin with a few elongate spines; antero-and posteroventral margins of mid and hind femurs withsome spines; tarsal joints elongate, covered with smallsetae, the first 3 with a very small pulvillus distad, fourthjoint with distal half bearing a small pulvillus; tarsalclaws symmetrical, usually simple, rarely minutely ser-rated on their inner margins.

Vrsansky, P.: Mesozoic amber cockroach (Blattodea)216

Table 1. Distribution of cockroaches in Mesozoic ambers.

Amber Country Age Ma n Described Adults Families Reference

Canadian Canada Campanian � 77 ? 0 0 – Blattulidae – Carpenter et al. (1937)

Tajmir Russia Santonian � 85 50 þ 0 0 – Blattulidae

– Mesoblattinidae

– in preparation

– Zherikhin & Sukatsheva

(1973)

Burmese Myanmar Cenomanian or

Turonian

� 90 � 50 1 ? – Blattulidae

– ?

– Grimaldi & Ross (2004)

New Jersey U.S.A. Turonian 92 5 2 1 – Blattulidae

– Mesoblattinidae

– Umenocoleidae

– in preparation

– Vrsansky 2003b

Sisteron France Cenomanian 96 1 1 0 – Blattulidae – Vrsansky (2008b)

Buzinie France Cenomanian 96 1 1 0 – Blattellidae – this publication

Archingeay France Albian 98 17 17 2 – Blattulidae

– Caloblattinidae

– Mesoblattinidae

– Blattellidae

– new family

– Vrsansky (in press)

Alava Spain Aptian � 120 50 þ 0 ? – Blattulidae

– ?

– Delclos et al. (2007)

Lebanese Lebanon Barremian or

Aptian

� 125 15 15 4 – Blattulidae

– Mesoblattinidae

– Caloblattinidae

– Umenocoleidae

– Vrsansky (2003b, 2004,

in prep.)

– Anisyutkin & Gorokhov

(2008)

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? Blattella lengleti sp. n.

Type material. Holotype specimen MNHN-BUZ-2.3, a nearly com-plete third instar (immature) female lacking head and apical part ofthe pronotum, preserved in Upper Cretaceous, Lower Cenomanianamber (from the lithological subunit B2 sensu N�raudeau et al. 1997)from La Buzinie in Champniers near Angoul�me, department ofCharente, SW France (Perrichot et al. 2007a). The piece of amber israther opaque, obscured by detritus, allowing the complete dorsalview but ventral view of the fore tarsi only. Deposited in the ambercollection of the Earth History Department, National Museum of Nat-ural History (MNHN), Paris, France.

Differential diagnosis. The strongly vaulted and elon-gated apically subgenital plate is diagnostic and identi-cal with the living B. beybienkoi, B. bisignata andB. nipponica, all of which belong to the Blattella ger-manica group (sensu Roth 1985). The form of subgeni-tal plate is specific to the species-groups of Blattella,and this particular type excludes affiliation of this fos-sil with the related genus Symploce, characterized by aplainer (not as distinctly vaulted) subgenital plate.

The species may be defined as a chronospecies, butit also differs from B. beybienkoi, B. bisignata andB. nipponica by having dark base and apex of cerci,from B. beybienkoi and B. bisignata by lacking medianposterior extension of pronotum. B. beybienkoi hasmore reduced pronotum coloration, B. bisignata haslongitudinal bands usually converging posteriorly (Roth1985), B. nipponica has pronotal stria more approxi-mated.

The present species additionally differs from all theknown Blattella species in its coloration of legs withdark midtibia and coloured basal and terminal part ofthe hind tibia, while the lateral margins are pale.

Description. Fragment length 3.2 mm. Antenna long,with 3 rows of sensilla – basal visible segment with 1 set,apical visible segments with 3 sets of 3 sensilla (20 seg-ments are preserved). Labial palp comparatively short,with the 4th segment (0.45 mm) narrow near basis, den-sely covered by sensilla; terminal segment (0.69 mm)sparsely covered by sensilla, with terminal protrudedelongation (Fig. 1). Pro-, meso- and metanotum withtwo longitudinal dark stripes. Pronotum incompletelypreserved, apparently transverse (1.1 þ / 1.9 mm), lat-eral margin covered by long setae. Lateral margin ofmesonotum with 4 long setae, posterior margin with 9or 10 setae (including the posterior lateral ones); meta-notum with 3 lateral and 10 posterior setae (9 are visi-ble on the holotype, while one additional is expectedon the basis of the free space).

Lobes on the meso- and metanotum absent.Fore tarsi (1 :0.46 : 0.34 :0.31 :0.81) with extremely

long claws (longer than tarsomere 4) and with long sen-silla chaetica. The remaining parts of the fore legs aresubmerged in the partially opaque amber and are thusunavailable for optical study.

Mid femur (1.5 mm) with anterior ridge colouredand covered by two rows of short sensilla. Posteriormargin with 9 strong and long setae. 8 additional setae

are present in central ridge. Mid tibia (0.9 mm) co-loured, with transversal ridges, strongly carinated. Tarsicoloured.

Hind femur (1.6 mm) with carination restricted to thelarge terminal spur and one posterior spur, other sensil-lae are fine. Hind tibia (2.0 mm) with basal and term-inal parts coloured, heavily carinated, but without shortsensilla. Hind tarsi (0.84 :0.26 : 0.13 :0.10 :0.13 mm)with very fine and short sensilla chaetica.

Abdomen dark, with central pale stripe and addi-tional pale macula on each side; 9 terga with posteriormargin bearing long setae (2, 9–10, 9–10, 9–10, 9–10,0, 0, 0, 4) (Some of the setae are invisible or weredestroyed during life and/or burial. Nevertheless, theyare distributed regularly throughout the surface andthere is a free space available for them.).

Cercus with 10 segments (1st reduced to a great ex-tent), with large lateral sensillae (terminal in the lastsegment) and small chaetica mainly in the bottom half.

Remarks. On the basis of the general habitus, and charac-teristic leg morphology the species can be attributedwithin the Blattellidae (including the subfamily Pseudo-phylodromiinae sensu Roth (2003), characterized by themirror, right position of the phallomere, unrotated oothe-ca and some insignificant differences – in contrast toGrandcolas (1996), who regards it as a separate family).

The coloration of the femoral apex, base and apex oftibia, terminal tarsal and terminal cercal segments(Figs 2–3), and the form of palp (Fig. 1) and subgenitalplate (Fig. 5, subapical terga are destroyed), are, to-gether with small tarsal pulvilli, diagnostic for thegenus Blattella (Roth 1985), (Fig. 6), which belongs tothe subfamily Blattellinae.

All the preserved characters indicate categorizationof the present larva within the B. germanica group (seediagnosis), and any diagnostic character which wouldindicate a different generic position are absent (pre-sence of meso- and metanotal lobes vary within thirdinstar of living Blattella, but is conservatively presentin the fourth instar). Moreover, the concept of thegenus is rather wide, with significant variability withinforms, subspecies and even species (Roth 1985). Never-theless, the presence of a closely related indigenousgenus differing in the morphology of adult stage cannotbe excluded. It is the sole reason for the questioned ca-tegorization within Blattella.

The size of the fossil corresponds to the third instarof Blattella germanica (usually 6–7 instars) and also tothe third instar of the most ancient representative of theBlattellidae, Piniblattella vitimica (Vishniakova, 1964)from the Lower Cretaceous of Baissa in Siberia(Vrsansky 1997). The second instar might be excludedalso on the basis of the position of the head which wasat least partially concealed below the pronotum withapparent paranotalia, and the fourth instar excluded be-cause of the absence of wing pads.

The significant morphological differences and thetime gap of 96 myr supports erection of a new species,even when based on the female nymph alone.

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Unless homoplastic, the coloured cerci are plesio-morphic for the genus, as they are preserved both inthe present species, and in indirectly related Blattellafrom Thailand (Fig. 6). The presence of a specific formof subgenital plate in B. lengleti and in few speciesfrom the B. germanica group suggests that this charac-ter is plesiomorphic. The absence of the pronotal med-ian posterior extension also appears plesiomorphic, andits absence in B. nipponica indicates that this species isthe most primitive in the species group. The reducedpronotum coloration of B. beybienkoi, longitudinal

bands usually converging posteriorly in B. bisignata,and pronotal stria more approximated in B. nipponica,all appear apomorphic because of the regular striationboth in ancient Piniblattella and derived Symploce.Dark midtibia and coloured basal and terminal parts ofthe hind tibia, with lateral margins pale are autapo-morphic for B. lengleti (dark areas on hind left tibia,anteriorly to the terminal part, and also antero-apicallyon the femur are absent in the right leg, and most prob-ably represent changes addicted with the preservationstate).


Figures 1–6. ?Blattella lengleti sp. n. from the Mesozoic amber of France. Holotype MNHN-BUZ-2.3 (Earth History Department,Paris). A third instar (immature) female. La Buzinie, Champniers, Charente, SW France. Early Cenomanian amber. 1. Maxillarypalp (terminal segment 0.69 mm long); 2–3. General view (fragment length without cerci 3.2 mm); 4. Fore tarsus (fragment lengthca. 0.9 mm); 5. Cerci (left cercus length 0.8 mm); 6. Living immature of Blattella sp. Canopy forest, Thailand (Zoological Insti-tute, Bratislava) – note the coloration of hind femora, tarsi and cerci, identical to that of B. lengleti (specimen length without legsand cerci 6.5 mm). (This figure is available in colour online at: museum-dez.wiley-vch.de)


The occurrence of all these three related living spe-cies in Asia is geographically obscure: B. beybienkoi isknown from East Asia, B. bisignata from South EastAsia, and B. nipponica from Japan. According to thepresence of apparent plesiomorphies in B. nipponica, itcan be presumed that the occurrence of this species,limited to Japan, is relictuous.

Generally, the relations presented above are based onthe morphology of adult specimens, but the most sig-nificant character, the shape of the subgenital plate isnot expected to change in the course of ontogeny. Thusthe present species can be categorized on the basis ofnymphs.

Derivation of name. lengleti is after Thierry Lenglet,the collector of the specimen.

Discussion

Cockroaches are peculiar with regard to their conserva-tive morphological standard, with general habitus hav-ing remained virtually unchanged since their first ap-pearance in the Mississippian some 350 Myr ago. SomePaleozoic genera persisted even into the Early Cretac-eous, which is for over 250 Myr, and genera occurringin geological strata 100 Myr apart are not exceptional(Vrsansky 2002, 2008a, b). Nevertheless, there aresome reservations to their conservativeness, namely therapid evolutionary tempo of species, and absence of liv-ing genera in the rich Mesozoic record counting30,000 specimens. The rich Turonian cockroach assem-blages of the New Jersey amber and Orapa sedimentsfrom Botswana are entirely Mesozoic in their taxo-nomic composition; and the same is true for the term-inal Mesozoic (Campanian) of Coahuila in Mexico (Ci-fuentez-Ruiz et al. 2006).

The absence of living genera in the late Mesozoicand Paleocene faunas indicates that most of them ra-diated after the Cretaceous/Tertiary boundary, and be-came abundant only during the Eocene.

This study provides direct evidence for the occur-rence of a taxon, indistinguishable from a living genusat the larval stage, deep in the Late Cretaceous. How-ever, the potential occurrence of other living genera inthe Mesozoic must be taken with reservation – Blattel-la is a rather primitive genus, closely related to thestem Blattellidae. This oldest still living family is re-corded since the basalmost Cretaceous of Baissa inTransbaikalian Siberia, represented by the extinct genusPiniblattella Vrsansky, 1997, related to the living Sym-ploce Hebard, 1916 (Vrsansky 1997), which is a closerelative of Blattella (Roth 1985). On the other hand,parental care, namely the caring for ootheca, must beconsidered a progressive pattern, partially responsiblefor the success of the German cockroach and its rela-tives. Blattella could have originated in forest areas andradiated later after entering more open environments.

An alternative possibility is that care for the oothecamight have evolved after the origin of the genus.

In contrast to cockroach families and genera, cock-roach species have evolved rapidly, and a single spe-cies, Archimesoblatta altera (Vrsansky 1997), is re-corded in strata of geographically adjacent localitiesextending in time more than 200 ka (at localities Baissaand Bon Tsagaan, distant in time for 5–15 Myr(Vrsansky 1997)).

Ecologically, the dark and distinct coloration shown byB. lengleti is characteristic for species living in humidand dense forest habitats, such as those present in Blat-tella sp. originating from the canopy forest in Thailand(Fig. 6), in contrast to Blattella living in more open evn-vironments (see Roth 1985), supporting a (coastal) forestenvironment of the source area, which was under strongmarine influences as indicated by diverse marine organ-isms such as crustaceans, ciliates and diatoms embeddedin amber (Perrichot 2005; Perrichot et al. 2007a, 2007b,2007c; Girard et al. in press). When compared with mostMesozoic cockroach families, the reduction of carinationis apparent, and is also expressed in the present species.It suggests that in modern cockroach families, the activeescaping strategy was prevailing over the passive mor-phological protection.

To summarize, the present study provides direct evi-dence for the presence of a taxon closely resembling aliving genus in the Mesozoic, most likely expandingthe range of any living cockroach genus to 96 Myr (of46 Myr). This result is particularly important as all theliving genera, in contrast to the Paleozoic and Meso-zoic ones, were, considered to be rather short-living.

Even more unexpectedly, this exception is repre-sented by a close relative of the common synanthropicGerman cockroach. It can be presumed that the earlyorigin of caring for the ootheca, as the most progressivetrait of Blattella, was responsible for its great success.

Acknowledgements

I thank Vincent Perrichot and Didier N�raudeau (Univ. Rennes 1) forextensive help during the study. I also thank Alain Couillard and Gi-nette Lambert who discovered the amber deposit and together withRomain Vullo, Bernard Gomez (Univ. Rennes 1) and Thierry Lengletparticipated in the field trip. I thank J�rg Ansorge (EAMU Greifs-wald) and Andrey Gorochov (ZIN St. Petersbourg) for revising themanuscript, L’ubom�r Vidlicka (ZIN Bratislava) for consultations,Martin Styan for linguistic revision and L’ubka Puskelov� (GlU SAVBratislava) for technical help. Supported by VEGA 6002, MVTS; Lit-erary Fund. This article is a contribution to the CNRS ANR program“AMBRACE” (project no. BLAN 07-1-184190).

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Documents

Mesozoic relative of the common synanthropic German cockroach (Blattodea)