Article Revision of Molopsis Schatzmayr (Coleoptera: Carabidae: Pterostichini), with descriptions of...

Accepted by J. Serrano: 21 Dec. 2011; published: 7 Feb. 2012

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Zootaxa 3185: 1–35 (2012) www.mapress.com/zootaxa/ Article

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Revision of Molopsis Schatzmayr (Coleoptera: Carabidae: Pterostichini), with descriptions of four new species

ROMAN LOHAJ1, BORISLAV GUÉORGUIEV2, GÉRARD DUBAULT3 & BERNARD LASSALLE4

1Institute of Forensic Sciences, Kuzmányho 8, SK-041 02 Košice, Slovakia. E-mail: [email protected], [email protected] Museum of Natural History, 1 blvd. Tzar Osvoboditel, 1000 Sofia, Bulgaria. E-mail: [email protected], [email protected], rue A.-Dumas, F-91600, Savigny sur Orge, France. E-mail: [email protected] rue Lefebure, F-28340, Boissy lès Perche, France. E-mail: [email protected]

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Systematic part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Molopsis Schatzmayr, 1943. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Tapinopterus (Molopsis) molopiformis (Lutshnik, 1922) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Tapinopterus (Molopsis) aenigmaticus sp.nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Tapinopterus (Molopsis) phrygius G. Müller, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Tapinopterus (Molopsis) machardi (Jeanne, 2005) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Tapinopterus (Molopsis) oyukluensis sp.nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Tapinopterus (Molopsis) molopinus (Chaudoir, 1868) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Tapinopterus (Molopsis) chaudoiri sp.nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19Tapinopterus (Molopsis) relegatus sp.nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Tapinopterus (Molopsis) wiedemanni (Chaudoir, 1850). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Tapinopterus (Pseudomolopsis) rebellis (Reiche & Saulcy, 1855) s.l. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Ovipositor morphology, seasonal activity and possible parental care in Molopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Key to the identification of Tapinopterus subg. Molopsis and Pseudomolopsis (in press) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31Checklist of the species of Tapinopterus subgenus Molopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Abstract

The species belonging to the subgenus Molopsis Schatzmayr, 1943 of genus Tapinopterus Schaum, 1858 are revised. Thestudy is based on 372 specimens and includes, for each taxon, diagnosis, description (only for the new species), references,new distributional records and illustrations. Morphological characters of the male and female genitalia are widely used todelimit the separate species, the significance of some non-gonapophyseal genital structures of the females for the system-atics of Molopsis is explicitly emphasized. For the first time, the male genitalia of T. molopiformis, T. molopinus, T. wie-demanni, and T. rebellis, including also newly described species, as well as female genitalia of Molopsis are described andillustrated. All available type material is listed and represented by habitus photographs. As a result, 9 monotypic species of Molopsis are recognized. Tapinopterus rebellis (Chaudoir, 1868) and T. rebelliskumanensis (Reitter, 1884), formerly assigned to Molopsis are recognized as outgroup-taxa. The following new speciesare described: T. (Molopsis) aenigmaticus sp.nov. (Asian Turkey, „Armenia“ imprecise locality), T. (Molopsis) chaudoirisp.nov. (Asian Turkey, Bursa Villayet, Uludağ Mt., Sakarya (Adapazari) Villayet, Gökdağ Mt.), T. (Molopsis) oyukluensissp.nov. (Asian Turkey, Konya Villayet, Fasihan Pass), and T. (Molopsis) relegatus sp.nov. (Asian Turkey, Bolu Villayet,Mengen env., Dorukhan Pass). The following nomenclatural acts are also proposed: Tapinopterus (Molopsis) molopinus(Chaudoir, 1868) = Tapinopterus (Molopsis) dipojranus brussanus Straneo, 1984, syn.nov.; Tapinopterus (Molopsis)

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LOHAJ ET AL.2 · Zootaxa 3185 © 2012 Magnolia Press

phrygius (G. Müller, 1932) = Tapinopterus (Molopsis) phrygius pisidicus G. Müller, 1932, syn.nov. = Tapinopterus (Mol-opsis) dipojranus Straneo, 1984, syn.nov. = Tapinopterus (Molopsis) dipojranus cilicius Straneo, 1984, syn.nov.; Tap-inopterus (Molopsis) machardi (Jeanne, 2005), comb.nov. of Molopsis machardi Jeanne, 2005; Tapinopterus (Molopsis)wiedemanni (Chaudoir, 1850), comb.nov. of Feronia (Molops) wiedemanni Chaudoir, 1850. Lectotypes for Feronia wie-demanni Chaudoir, 1850 and Feronia rectangula Chaudoir, 1868 are designated. In conclusion, the relations between the ovipositor morphology, seasonal activity and possible parental care in Mol-opsis are discussed. Several evident derived characters in the group are briefly considered too. Last but not least, a key tothe identification and check-list of the species are provided.

Key words: Coleoptera, Carabidae, Pterostichini, Molopsis, taxonomy, revision, new species, new synonymy, new com-bination, Palaearctic Region, Turkey

Introduction

Originally Molopsis was proposed as subgenus of Tapinopterus Schaum, 1858 by Schatzmayr (1942: 51, 1943:119) for two polytypic species: one Asiatic (T. molopinus Chaudoir) and one European (T. rebellis). Prior to thisevent (Chaudoir, 1850, 1868, 1876; Fairmaire, 1866; Kraatz, 1875; Marseul, 1880, 1882; Seidlitz, 1887, 1888;Ganglbauer, 1889; Apfelbeck, 1904; Bodemeyer, 1905; Jakobson, 1907; Breit, 1914; Lutshnik, 1922; Müller,1923, 1932, 1937; Schatzmayr, 1929; Csiki, 1930; Straneo, 1936; Mařan, 1940) and after that (Jedlička, 1963;Straneo, 1986; Kirschenhofer, 1991; Lorenz, 1998, 2005; Casale & Vigna Taglianti, 1999; Bousquet, 2003; Jeanne,2005; Guéorguiev & Lohaj, 2008), a great number of authors dealt with the taxonomy of this group. Recently, Mol-opsis is treated as subgenus of Tapinopterus including four polytypic species (Bousquet, 2003; Lorenz, 2005): T.(M.) dipojranus Straneo 1986 from Turkey, with three subspecies, T. (M.) molopinus (Chaudoir, 1868) from Tur-key, with three subspecies, T. (M.) phrygius G. Müller, 1932 from Turkey, with two subspecies, and T. (M.) rebellis(Reitter, 1884) from Greece, with two subspecies.

The species of this group belong to a quite specific and original Anatolian lineage of the carabid tribe Pterosti-chini. The adults are generally found under stones and rotting logs in moderately moist temperate forests reachingthe alpine zone. As a rule, they are wingless and are found in small numbers. The larvae of the group are unknown.

The purpose of this paper is to provide a taxonomic revision of the species from Molopsis.Aims of the study. The main objectives of this study are: (i) to revise the taxonomic structure at species level;

(ii) to revise the published and add new distributional data and (iii) to make the species recognition possible from apractical (e.i. morphological, rather than geographic) point of view. The achievement of these aims follows someconsecutive steps: collecting and analysis of published data; revision of type material; study of unpublished mate-rial; re-evaluation of hitherto used characters and looking for new ones; analysis of all data.

Material and methods

This paper is based on a study of 372 specimens of Molopsis as well as dozens of specimens belonging to othersubgenera of Tapinopterus and other genera of Pterostichini.

The morphological structures were examined using stereoscopic microscopes Carl Zeiss Jena Technival 2,Olympus SZ 60, and LEICA S8 APO. Photos were taken a LEICA S8 APO microscope equipped with a digitalcamera NIKON COOLPIX® E 4500. Dissections were made using standard techniques. To extract genitalia speci-mens were boiled in water. Male and female genitalia were dissected, cleaned and cleared in 10 % KOH. Afterstudy, they were mounted on the same labels together with specimens, on separate labels under the examined spec-imens, or in vials pinned beneath the specimens from which they were removed. If need for making drawings, gen-ital structures were replaced in glycerine and outlined using drawing tube mounted on stereoscopic microscopeCarl Zeiss Jena Amplival. Dorsal aspects of the median lobe of aedeagus were drawn using drawing tube assem-bled on Carl Zeiss Jena Technival 2.Sources of material are the collections of the following institutions and private collectors:

BMNH Natural History Museum, London, United KingdomDEI Deutsches Entomologisches Institut, Eberswalde, Germany


Zootaxa 3185 © 2012 Magnolia Press · 3REVISION OF MOLOPSIS (CARABIDAE: PTEROSTICHINI)

HNHM Hungarian Natural History Museum, Budapest, HungaryMCSNM Museo Civico di Storia Naturale di Milano, ItalyMCSNT Muzeo Zoologico di Storia Naturale, Trieste, ItalyMNHN Muséum National d´Historie Naturelle, Paris, France MNHUB Museum für Naturkunde der Humboldt-Universität zu Berlin, GermanyNMNHS National Museum of Natural History, Sofia, BulgariaNMP Národní Museum Praha, Czech RepublicNMW Naturhistorisches Museum Wien, Vienna, AustriaZISP Zoologiceskij Institut, Akademia nauk, Sankt Peterburg, RussiaZMAN Zoölogisch Museum Amsterdam, The NetherlandscAC Coll. Achille Casale, Sassari, Italy cBL Coll. Bernard Lassalle, Vanves, FrancecDW Coll. David Wrase, Berlin, GermanycDE Coll. Dominique Echaroux, Etrechy, FrancecEH Coll. Evžen Hajdaj, Ježov, Czech RepubliccGD Coll. Gérard Dubault, Savigny sur Orge, FrancecMP Coll. Maurizio Pavesi, Milan, ItalycRL Coll. Roman Lohaj, Košice, SlovakiacVS Coll. Vladimír Skoupý, Žilina, Czech RepubliccWH Coll. Walter Heinz, Schwanfeld, Germany

Other abbreviations used in the text: BL—body length measured from apex of mandibles to apex of elytra,Mt.—mountain/s; NW—northwest; S—south; SW—southwest; W—west; PL—pronotal length along midline;PW—maximum width of pronotum; p—printed label; h—handwritten label; / - used to separate different labels.

In this work, nine monotypic species of Molopsis are recognized. Tapinopterus rebellis (Chaudoir, 1868) and T.rebellis kumanensis (Reitter, 1884) will be placed to a new genus (Giachino et al., in press).

Notes about some morphological terms. In general, the terminology of morphology in the adults of pteros-tichines follows Bousquet (1999). Apart from that, we pay attention to several terms and structures that have yetnot widely used in the carabid literature.

We propose using “internal face” and “external face” for depiction of the parameres. The first term concernsthe surface of each paramere which is normally facing the median lobe of aedeagus (in life and before dissectionthe paramere and lobe are firlmy fastened). Conversely the “external face” refers to the surface of the outer face.We use “sternum VII” for the last visible, normally not retracted, abodominal sternum. In many papers this isreferred to as sternum VI, but actually it is morphologically the seventh adult sternum as sternum I in adult beetlesis atrophied (Ball & Shpeley, 1983).

The terminology of the chitinised structures of the female genitalia follows Ball & Shpeley (1983).Significance of female genitalia morphology in the systematics of Molopsis. The siginificance of some

structures of the female genitalia in the Carabidae has already been discussed (Brandmayr, 1977, 1978; Habu,1981a, 1981b; Allen & Ball, 1980; Giachino & Casale, 1983; Giachino & Sciaky, 1991). Normally, the shape ofovipositor, including both the valvifer and stylus are indistinguishable or hardly distinguishable at species level inPterostichini (Habu, 1981a, 1981b). However, after a careful study of a moderate number of samples and analysisof data, we found that, probably in result of strong regression phenomena, the ovipositor (especially basal stylus) inMolopsis has more or less constant aspect in one and same species. Besides, the shape of other structures of thefemale genitalia, namely sternum VIII and receptaculum of spermatheca exhibit little intraspecific differences andthat, in combination with other characters, is widely used in the classification of Molopsis. The occurrence of a lowintraspecific variability of the non-gonapophyseal ovipositor has been demonstrated in Pterostichini (Brandmayr,1978).

Historical remarks. The studies on the ground beetle subgenus Molopsis Schatzmayr, 1943 can be dividedinto two stages, according to the availability of material for study as well as stage of knowledge and methods ofwork.

The first mention of a representative of this group is a statement about “Steropus brevis” from “Constantinop.”(Sturm, 1843: 25), as result of incorrect assignation to Pterostichus brevis (Duftschmid, 1812). After that, Chaudoir



(1850) described Feronia (Molops) wiedemanni from Northern Anatolia. Based on material collected by M. J.Lédérer in 1865, the last species was recorded also from the Boz dagh Mt. (Fairmaire, 1866). However, Chaudoir(1868) found differencies between his F. wiedemanni and Fairmaire´s specimen from Boz dagh and announced thelatter is a separate species, Feronia (Haptoderus) rectangula. In the same work, Chaudoir described also Feronia(Haptoderus) molopina from type locality “Istanbul” (ibid.), based on the mentioned above datum of Sturm (1843).Since then the most authors working on Tapinopterus omitted Feronia wiedemanni (Csiki, 1930; Müller, 1932;Mařan, 1940; Schatzmayr, 1942; Straneo, 1986; Kirschenhofer, 1991; etc.). This taxon was recently placed as spe-cies “incertae sedis” of the genus Molops Bonelli, 1810 (Bousquet, 2003). After searching in the Chaudoir´s collec-tion in MNHP by one of us (GD), the type series of this species (together with the types of Feronia molopina andFeronia rectangula) has been discovered.

Kraatz (1875) described Pterostichus olympicus from the mountain of “Olymp”. Later, Csiki (1930) regardedmolopinus Chaudoir and molopiformis Lutshnik as different species and placed them in the genus Pterostichus,subgenus Tapinopterus, sectio Crisimus. This author considered olympicus Kraatz as a synonym of molopinusChaudoir.

The second period of research is result of more intensive faunistic investigations mostly in the Anatolian area.In the early thirties of the twentieth century, three important works have been pubslihed by different Italian ento-mologists. Müller (1932) described Tapinopterus (Crisimus) phrygius, including two different subspecies. The typ-ical form was described from the Ak dağ Mt. and Sultan dağ Mt. (“Phrygic Taurus”), whereas the subspeciespisidicus was described from the Barla Dağlari Mt. and Davraz dağ Mt. Müller pointed out differential diagnosisfor T. molopinus (he treated the Kraatz´s species olympicus as identical with Chaudoir´s molopinus) and T. molopi-formis. In 1936 Straneo (1936: 156–157) differentiated four closely related species of Tapinopterus in a groupcalled by him the “gruppo rebellis” (much later, however, this famous expert of the World Pterostichini, tracing outaspects of the history of the genus, omitted his earlier arrangement, Straneo, 1986). Mařan (1940) treated T. olympi-cus and T. molopiformis as separate subspecies of T. molopinus and also presented the differencies between thenominotypical subspecies and T. molopinus olympicus. Seven years after the Straneo’s grouping Schatzmayr(1943) created subgenus Molopsis for two species. One of them is Tapinopterus molopinus (Chaudoir, 1868) desig-nated as type species. This author regarded the last taxon as polytypic and included in it the nominotypical ssp.(type locality Istanbul?), T. molopinus olympicus (Kraatz, 1875) from the Uludağ Mt. near Bursa (NW Turkey), T.molopinus molopiformis (Lutshnik, 1922) from the Boz Dağlari Mt. (= Tmolos Mt. by ancient authors) near Izmir(W Turkey), and T. molopinus phrygius G. Müller, 1932 from SW Turkey (Ak dağ, Sultan dağ, Barla Dağlari andDavraz dağ). The second species classified by Schatzmayr in Molopsis was T. rebellis (Reiche & Saulcy, 1855)from Peloponnesos, south Greece (Reiche & Saulcy, 1855). Straneo (1986) described Tapinopterus (Molopsis)dipojranus from Dipojraz dağ Mt. (SW Turkey, west of the Beyşehir Lake), which besides the nominotypical formhas included T. dipojranus brussanus from Uludağ Mt. near Bursa and T. dipojranus cilicius from Bolkar dağlariMt. (S Turkey). In this work, discrimination between T. dipojranus s. str. and T. phrygius, based upon the shape ofpronotum and median lobe of aedeagus, as well as that between the nominotypical form and T. dipojranus brus-sanus and T. dipojranus cilicius is made. Recently, in an excellent survey of the Anatolian carabid fauna, Casale &Vigna Tagliatni (1999) indicated three species of Molopsis from the Asian part of Turkey: T. molopinus (with thenominotypical form and the subspecies olympicus and molopiformis), T. phrygius (with the nominotypical formand the subspecies pisidicus) and T. dipojranus (with the nominotypical form and the subspecies brussanus and cil-icius). The same arrangement of taxa was followed by Bousquet (2003) and Lorenz (2005), but they count also inMolopsis the Greek T. rebellis (with nominotypical form and ssp. kumanensis). Recently, Jeanne (2005) describedfurther new species, Molopsis machardi Jeanne, 2005, from the area of Antalya. It is worth mentioning that, thisauthor recognized Molopsis as distinct genus and introduced some changes in the intrageneric treatment.

Systematic part

Molopsis Schatzmayr, 1943

Type species. Feronia molopina Chaudoir, 1868 by original designation.References. Tapinopterus (Molopsis): Schatzmayr, 1942: 51; Schatzmayr, 1943: 119; Straneo, 1986: 122;



Kirschenhofer, 1991: 5; Lorenz, 1998: 267; Casale & Vigna Taglianti, 1999: 381; Bousquet, 2003: 518; Lorenz,2005: 288.

Molopsis: Jeanne, 2005: 284.Original definition of Molopsis, translated. “Habitus different from the other Tapinopterus–subgenera, simi-

lar to small Molops or Styracoderus (Carabidae, Pterostichini). Basal bead of pronotum near hind angles often flat-tened; elytron without scutellar setigerous puncture and with rudimentary scutellar stria; last tarsal segment withoutsetae ventrally.” (Schatzmayr, 1943).

Redescription. Medium-sized pterostichine (BL 11.2–16.2 mm), piceous brown to black, smooth, shinythough some females matt. Habitus very similar to Molops s. str., mainly representatives of the “piceus” speciesgroup, which also represented in NW Turkey by a single polytypic species.

Head relatively large and robust, smooth, with prominent eyes. Frons with two supraorbital setigerous punc-tures. Clypeus with two long setae, labrum anteriorly with six marginal setae. Mentum with pair of setigerouspunctures; tooth prominent and bifide at apex. Submentum with two pairs of setae, outer setae much shorter thaninner ones. Glossal sclerite rectangular with two setigerous punctures on anterior margin; paraglossae apparentlylonger than glossal sclerite. Antennae relative short, reaching anterior angles of elytra, Antennomere I twice as longas II and much thicker then remaining, antennomeres IV–XI covered with dense pubescence.

Pronotum almost as wide as long, or slightly wider than long, with rounded lateral sides. Anterior anglesslightly prominent, rounded at apex. Posterior angles distinct, acute (T. molopinus, T. wiedemanni, T. phrygius) torectangular (T. molopiformis), without setigerous punctures. Base of pronotum on each side with single modestlydeep, smooth (sometimes with fine transversal wrinkles), linear posterolateral impression, spreading over basalthird. Median longitudinal impression distinct, not reaching basal or anterior margins. Lateral groove evenly nar-row along sides, with single setigerous punctures in anterior third.

Elytra oval or more or less cylindrical, with distinct, finely punctate striae, with rudimentary scutellar stria,

scutellar setigerous punctures missing. Third elytral interval with two setigerous punctures (normal 2nd and 3rd dor-sal elytral pores in carabids) situated in apical half of elytra; exceptionally with one or three punctures. Umbilicateseries consisting of 13–17 pores in all but one species (10 pores in T. relegatus sp. nov., see below); one setigerouspunctures at the end of 7th elytral stria.

Mesepisternum without punctation. Last abdominal segment (sternum VII) with two pores (T. machardi, somespecimens of T. phrygius) or four pores (other species).

Legs relatively short, robust, tarsomeres 1–4 glabrous above, pubescent ventrally. Onychium without setaeunderneath (except T. machardi - with one or two pairs of setae). Two setigerous punctures on middle coxa, onlyone on middle trochanter. Three setigerous punctures on hind coxa, including medial one situated below meetingpoint of coxae; single seta on hind trochanter. Profemur with two setigerous punctures on anterior ventral margin;mesofemur with three medio-ventral seta on anterior margin and 3 (rarely 4) on posterior ventral margin; metafe-mur with two setae on posterior ventral margin. Mesotibial ctenidium clear but poorly differentiated (as in Molopsspp. and Myas). Males with first three protarsomeres dilated, ventrally with densely arranged adhesive setae.

Male genitalia: Aedeagus relatively large, median lobe rectangulary arcuate and apex variable (ventral view),short and subtriangular (dorsal view) in T. molopiformis to very long, narrow, and hooked at extremity in T. wiede-manni, apical orifice deflected to left lateral position. Left paramere with transverse apophysis; right paramereelongate apically.

Female genitalia: Tergum VIII with convex or subtriangular distal margin and ± short proximal “legs”. Ster-num VIII consisting, in most species, of two chitinised plates connected by a membrane which Connected by mem-brane which is disappeared in T. wiedemanni. Syntergum IX+X long, narrow, bearing a pair of small, articulatedovipositors. Ovipositor consisting of large valvifer and one-segmented stylus (in the most species) or two-seg-mented stylus (in T. molopiformis). Spermatheca with seminal canal and receptaculum discrete; receptaculumshorter than seminal canal; spermathecal canal long or short, inserted at junction of receptaculum and seminalcanal.

Larvae. Unknown.Distribution. The presence of T. (M.) molopinus (cf. Chaudoir, 1868) in Europe is doubtful. In Asia, Molopsis

inhabit the western half of the Anatolian Peninsula but the eastern limits of the range remain poorly documented atpresent. The species occur in forested regions, elevated alpine meadows, near wet places, streams, thawing snow,under stones, leaves, and pieces of wood.



Tapinopterus (Molopsis) molopiformis (Lutshnik, 1922)(Figs. 3–4, 26–27, 46, 56, 81, 96, 111, 125, 139, 145, 151, 155–156, 163, 167)

Type locality. Anatolie, Bosch-Dagh.Taxonomic decisions. Feronia (Haptoderus) rectangula Chaudoir, 1868: 246.

Platysma (Crisimus) molopiforme Lutshnik, 1922: 77, replacement name for Feronia rectangula Chaudoir, 1868:246, nec Fairmaire, 1866: 252.

ReferencesMolops wiedemanni Chaudoir: Fairmaire, 1866: 252.Feronia (Haptoderus) rectangula: Chaudoir, 1868 : 246Feronia wiedemanni Kraatz, 1875: 419Feronia (Steropus) rectangula: Marseul, 1882: 48.Haptoderus rectangulus: Ganglbauer, 1889: 52.Feronia rectangula: Apfelbeck, 1904: 244.Platysma (Tapinopterus) rectangulum: Jakobson, 1907: 352.Platysma (Crisimus) molopiforme: Lutshnik, 1922: 77.Crisimus (Tapinopterus) rectangulus: Schatzmayr, 1929: 330.Pterostichus (Tapinopterus) molopiformis: Csiki, 1930: 707.Tapinopterus (Crisimus) molopiformis: G. Müller, 1932: 217.Platysma (Tapinopterus) molopiformis: Straneo, 1936: 156.Tapinopterus molopiformis: G. Müller, 1937: 132.Tapinopterus molopinus molopiformis: Mařan, 1940: 57.Tapinopterus (Molopsis) molopinus molopiformis: Schatzmayr, 1943: 119; Straneo, 1986: 122; Kirschenhofer,

1991: 6; Casale & Vigna Taglianti, 1999: 381; Bousquet, 2003: 518.Pterostichus molopinus: Jedlička, 1963: 19.Tapinopterus (Molopsis) molopiformis: Lorenz, 1998: 267.Molopsis molopiformis molopiformis: Jeanne, 2005: 385.Tapinopterus (Molopsis) molopinus molopiformis: Lorenz, 2005: 288.

Material examined. 29 specimens.Type material. Lectotype ♀ of Feronia rectangula here designated, relatively well preserved, “rectangula

Chaud. Anatolie. Lederer.” (h label pinned on the bottom of box)/ LECTOTYPE Feronia (Haptoderus) rectangulaChaudoir G.Dubault désign. 2003 (red p)/ Tapinopterus (Molopsis) molopiformis G.Dubault détermin. (p) (Fig. 3)(MNHN, “Collection Chaudoir”, box no 216 “Argutor”).

Other material examined. Boz daglari: 2 ♂♂ 2 ♀♀, “Caucas” / “Coll. Türk 188” (NMW); 1♀, “BoszdaghColl. Javet” (MNHN); 1 ♀, “Bosdagh” (MNHN); 4 ♂♂ 5 ♀♀, “Tmolos-Gbg., Lydien, West-Kleinasien, Weirather,Innsbruck” (MCSNM, MCSNT, MNHN; ZMAN); 1 ♂, “Asia Minor Bos-Dagh.” / “Coll. Türk 188” / “Pt. rectan-gulus Chd. Coll. Reitter” (HNHM); 1 ♂, “Bos dagh As. min.” / “rectangulus” / “Coll. Apfelbeck” / “Tapinopterusrectangulus Chd. det. Ing. Jedlička” (HNHM); 1 ♂, Caucase? (h) (ZISP); 1 ♀, Klein Asien, Bosz-Dagh, Lederer(h) (DEI); 1 ♂ 1 ♀, TR (Izmir) Boz-dağ, ob. Bozdagköy, 1500–1700 m, 2.–3.5.1996, Heinz lgt. (cWH); 2 ♂♂,TURKEY, vil. Izmir, Boz dağlari, Boz dag köy env., 1200–1500 m, 29.5.2003, R. Lohaj & I.Smatana lgt. (cRL)(Fig. 4); 3 ♂♂ 2 ♀♀, NW Turkey, prov. Izmir, Boz dag, 1520 m, NE Ödemish, 6.5.07 lgt. E.Hajdaj (cEH, cRL).

Diagnosis. BL 11.2–13.8 mm.Male genitalia. Median lobe of aedeagus short, with subtriangular apex, apical orifice deflected to left lateral

position (Figs. 26–27); basal bulb short, with concave basal orifice, apical part longer, copulatory sclerite straight,obliquely situated in medial part, apex slighlty bent downwards (Figs. 46, 56). Right paramere relatively short,slightly curved medially, with thicker basal part and thinner apical one (Figs. 81, 96). Left paramere with long, thintransverse apophysis and oblique paramedial apophysis (Figs. 111, 125).

Female genitalia. Tergum VIII with convex distal margin and short proximal “legs” (Fig. 139). Sternum VIIIconsisting of two chitinised plates connected by membrane, with well-developed “legs” (Fig. 145). SyntergumIX+X long, narrow, bearing pair of small, articulated ovipositors (Fig. 151). Ovipositor consisting of a large valvi-fer and smaller two-segmented stylus (Figs. 155–156); valvifer with both internal and external margins each withpointed tip and well-developed process across; basal stylomere relatively larger than apical one, latter with single



minute (probably ensiform) seta. Spermatheca with seminal canal and receptaculum discrete; seminal canal long;receptaculum slighty shorter than seminal canal, coiled apically (Fig. 163); spermathecal gland prolonged, sper-mathecal canal long, inserted at junction of receptaculum and seminal canal (Fig. 167).

Taxonomic notes. This taxon was first recorded as Molops wiedemanni Chaudoir from Boz dag by Fairmaire(1866: 252). Chaudoir (1866) noted that Faimaire's specimens were different from his specimens of F. wiedemanniand described the taxon as Feronia rectangula. The name Feronia rectangula was preoccupied by Feronia(Orthomus) rectangula Fairmaire, 1859, currently a synonym of Orthomus (Orthomus) barbarus barbarus(Dejean, 1828) (see Bousquet, 2003: 477). Lutshnik (1922: 77–78) first noted this homonymy and proposed a newname, Platysma molopiforme, for Feronia rectangula Chaudoir, 1868.

Müller (1937: 132) studied a series of T. molopiformis from the type locality and compared it with specimensof his T. phrygius s.str. and T. phrygius pisidicus and stated that the former is a distinct species.

Distribution. Boz dağlari (Tmolos Mountains by the ancient authors) near Izmir (“Lydia”), W Turkey.

Tapinopterus (Molopsis) aenigmaticus sp.nov.(Figs. 5, 28–29, 47, 57, 82, 97, 112, 126)

Type locality. Turkey ?Material examined. One specimen.Type material. Holotype ♂, with left hind leg and three right metatarsomeres missing, Armenie (h) / Fer.

(Tapinopt.) rectangula Chd. (pale orange h)* / Zool.Inst St. Petersburg (yellow p) / HOLOTYPE Tapinopterus (sg.Molopsis) aenigmaticus n. sp. Lohaj, Guéorguiev, Dubault & Lassalle, 2004 (red p)/ (Fig.5) (ZISP).

*Note: One male of Molopsis molopiformis in ZISP bears label “Caucase?” written with the same handwritingand on same pale orange paper as the first two labels of the new species. We assume these labels were later addi-tions.

Description. BL 12.1 mm, maximum width 4.4 mm. Head, pronotum and elytra black, shiny, antennae, legsand abdomen piceous brown. Head large, robust, almost as long as wide, shiny, smooth, with very fine microsculp-ture. Antennae relatively short, reaching hind angles of pronotum, with dense, decumbent pubescence on antenno-meres 4–11.

Pronotum wider than long, ratio PL/PW 0.67, with rounded lateral sides, shiny, smooth, with very fine trans-versal wringles along the midline. Anterior edge strongly sinuate, with prominent, rounded, apical angles. Posteriorangles small, almost rectangular. Base of pronotum with single deep linear posterolateral impression on each sideof midline, reaching basal third, smooth, with very fine transversal wrinkles. Midline distinct, not reaching basal oranterior margins.

Elytra parallel-sided, slightly flattened, with distinct, finely punctate striae. Third elytral interval with two set-

igerous pores situated in apical half of elytra, 2nd in middle and 3rd in apical fourth. Umbilicate series consists of 14setigerous punctures on each side, in the middle widely interrupted, with distribution: left side 5 + 1 + 8, right side6 + 8.

Male genitalia. Median lobe of aedeagus relatively short, straight, with short and fine apex in dorsal view anda small twisted tooth in ventral position (Figs. 28–29); apical orifice deflected to left laterad position, ventral mar-gin forming slight acute angle at medial part, lobe with distinct lateral process in left lateral view (Figs. 47, 57).Right paramere strongly curved apically, with thicker basal part and thin apical part truncate towards tip (Fig. 82,97). Left paramere with subangular plate, long, thin transverse apophysis, slightly bent inwards, and oblique para-medial apophysis (Figs. 112, 126).

Female genitalia. Unknown.



PLATE 1. Habitus of Tapinopterus species (Figs. 1–12). Fig. 1. Tapinopterus (Pseudomolopsis) rebellis (Greece, Taygetos);Fig. 2. T. (Pseudomolopsis) rebellis kumanensis (holotype); Fig. 3. T. (Molopsis) molopiformis (lectotype of Feronia rectan-gula); Fig. 4. T. (Molopsis) molopiformis (Turkey, Boz dağ); Fig. 5. T. (Molopsis) aenigmaticus sp. nov. (holotype); Fig. 6. T.(Molopsis) phrygius (syntype); Fig. 7. T. (Molopsis) phrygius (Turkey, Davraz dağ); Fig. 8. T. (Molopsis) phrygius (Turkey,Kumalar dağlari); Fig. 9. T. (Molopsis) phrygius (holotype of Tapinopterus dipojranus); Fig. 10. T. (Molopsis) phrygius (para-type of Tapinopterus dipojranus cilicius); Fig. 11. T. (Molopsis) phrygius (Turkey, Beyşehir); Fig. 12. T. (Molopsis) phrygius(Turkey, Sultan dağlari).



Etymology. From the Latin adjective aenigmaticus, meaning “enigmatic”. The name reflects the uncertaintyabout the type locality of this species.

Diagnosis. This species is easily recognized from all other species of the group by the shape of the median lobeof aedeagus and right paramere; median lobe dorsally with short, fine apex, with distinct lateral process at left lat-eral position which forms a small tooth ventrally; ventral margin of forming slight acute angle at medial part (Figs.28–29, 47, 57). Right paramere strongly curved apically, with thin apical part not pointed towards tip (Fig. 82, 97).

Distribution. The type locality of this new species remains uncertain. If the locality label “Armenie” is correct(see also Note above), the new species originated either from Armenia (former republic of USSR) or from “Arme-nian Taurus” of ancient authors. This region, also called “Small Armenia” is located in Kurdistan, south of LakeVan (see Casale & Vigna Taglianti, 1999: 401, note 21). Correct type locality needs to be confirmed by the exami-nation of additional material.

Tapinopterus (Molopsis) phrygius G. Müller, 1932(Figs. 6–12, 30–34, 48–52, 58–62, 83–84, 98–99, 113–114, 127–128, 146, 157)

Type locality. Kleinasien, Ak-Dagh im Phrygischen Taurus. Taxonomic decisions.

Tapinopterus (Crisimus) phrygius G. Müller, 1932: 216 (typ. loc. “Ak dagh”)Tapinopterus (Crisimus) phrygius pisidicus G. Müller, 1932: 217 (typ. loc. “Barla dağlari”) , synonymum novum.Tapinopterus (Molopsis) dipojranus Straneo, 1986: 124 (typ. loc. “Dipojraz”), synonymum novum.Tapinopterus (Molopsis) dipojranus cilicius Straneo, 1986: 126 (typ. loc. “Bolkar daglari”), synonymum novum.

References.Tapinopterus molopinus: Bodemeyer, 1905: 36.Pterostichus (Tapinopterus) molopinus (= olympicus): Bodemeyer, 1905: 103.Tapinopterus (Crisimus) phrygius: G. Müller, 1932: 216.Tapinopterus (Crisimus) phrygius pisidicus: G. Müller, 1932: 217.Tapinopterus phrygius: G. Müller, 1937: 132; Mařan, 1940: 57.Tapinopterus phrygius pisidicus: G. Müller, 1937: 132; Mařan, 1940: 57.Platysma (Tapinopterus) phrygius: Straneo, 1936: 156.Platysma (Tapinopterus) phrygius var. pisidicus: Straneo, 1936: 156.Tapinopterus (Molopsis) phrygius: Schatzmayr, 1943: 119; Straneo, 1986: 122; Kirschenhofer, 1991: 6; Casale &

Vigna Taglianti, 1999: 381.Tapinopterus (Molopsis) phrygius pindicus (!): Schatzmayr, 1943: 119.Tapinopterus (Molopsis) dipojranus: Straneo, 1986: 124; Kirschenhofer, 1991: 6.Tapinopterus (Molopsis) dipojranus cilicius: Straneo, 1986: 126; Kirschenhofer, 1991: 6; Lorenz, 1998: 267; Bous-

quet, 2003: 518.Tapinopterus (Molopsis) phrygius pisidicus: Kirschenhofer, 1991: 6; Lorenz, 1998: 267; Bousquet, 2003: 518.Tapinopterus (Molopsis) dipojranus dipojranus: Lorenz, 1998: 267; Bousquet, 2003: 518.Tapinopterus (Molopsis) phrygius phrygius: Lorenz, 1998: 267; Bousquet, 2003: 518.Molopsis molopiformis dipojranus: Jeanne, 2005: 385.Molopsis molopiformis cilicius: Jeanne, 2005: 385.Molopsis molopiformis phrygius: Jeanne, 2005: 385.Molopsis molopiformis pisidicus: Jeanne, 2005: 385.Tapinopterus (Molopsis) dipojranus dipojranus: Lorenz, 2005: 288.Tapinopterus (Molopsis) dipojranus cilicius: Lorenz, 2005: 288.Tapinopterus (Molopsis) phrygius phrygius: Lorenz, 2005: 288.Tapinopterus (Molopsis) phrygius pisidicus: Lorenz, 2005: 288.

Material examined. 72 specimens.Type material. One syntype (♂) of Tapinopterus (Crisimus) phrygius, Ak dagh bei Tschiwril Asm. (h)/ phry-

gius Müll. (h)/ Paratypus (p) (red label)/ 322 (h) (red label)/ (Fig.6) (MCSNM).Two syntypes (♂♂) of Tapinopterus (Crisimus) phrygius pisidicus, “Klein Asien Pisidischer Taurus Barla



Gruppe N. O. von Isparta” (green p/h) / “vend. Weirather Tapinopterus (Crisimus) phrygius pisidicus J. Müll.” (p/h) / “Tapinopterus phrygius pisidicus J. Mü.” (pink h) (NMP).

Holotype ♂ of Tapinopterus (Molopsis) dipojranus, Dipojras Gbg. Pisid. Taurus, Klein-Asien, Weirather, Inns-bruck. (p)/ Holotypus (p) Tapinopt. dipojranus (h) (red label)/ Holotypus Tapinopterus dipojranus (h),det.S.L.Straneo 1985 (p)/ TAPINOPTERUS (sg. MOLOPSIS) phrygius Müller 1932 R.LOHAJ det.2003 (p)/ (Fig.9) (MCSNM).

Four paratypes (♂♂) of Tapinopterus (Molopsis) dipojranus ssp. cilicius, TURCHIA—vil. Mersin, Namrun(Çamliyayla) 5.VI.1983, G.Sama leg. (p)/, Bolkar Dağ, altipiano m 2400- (h)/PARATYPUS (p) Tap. dipojr.s.cil-icius (h) (red label)/ Paratypus T.dipojranus s.cilicius (h), det.S.L.Straneo 1985 (p)/ TAPINOPTERUS (sg. MOL-OPSIS) phrygius Műller 1932 R.LOHAJ det. 2003 (p)/ (Fig. 10) (2 ♂♂ MCSNM, 1 ♂ cBL, 1 ♂ cWH).

Other material examined. Barla daglari: 1 ♂, Barla-Gbg. Pisid. Taurus (MCSNM); 1 ♀, Gelindschik Ana-Gbg. nö Isparta, Pisidia, Asm. (MCSNM); 1 ♂, Turcia, 8.5.1991, Barla dagi, 200–2500 m, M. Mikát lgt. (cVS); 1♀, “Barla-Gbg. Pisid. Taurus” (MNHN).

Kumalar daglari: 2 ♂♂ 2 ♀♀, Turkey occ., 21.vi.1999, Kumalar Daglari mts., alp., Basören S, 1900–2200 m(50 km S-of Afyon) S.Benedikt leg. (Fig. 8) (cVS, cRL).

Sultan daglari: 1 ♀, Turkey, 23.–26.5.96, Sultan Daglari, Cankurtaran, 1800 m, Smatana lgt. (cRL) (Fig.12); 1♂ 2 ♀♀, “Asia minor Sultan Dagh v.Bodemeyer” (MNHN, ZMAN); 1 ♀, Asia minor, Sultan Dagh, v. Bodemeyer,O.Leonhard (DEI); 1 ♀, Asia minor, Sultan Dagh, v. Bodemeyer, ex-coll. Dr. Melichar (BMNH); 1 ♀, Anatolien,Ak-Chehir, 1900, Korb. (DEI); 1 ♂ 1 ♀, Anatolia centr, Sultan Dagh b. Çay, 1900–2200 m, 18.7.1965, Heinz lgt.(cWH); 4 ♂♂ 2 ♀♀, Asia minor, Sultan Dagh, v. Bodemeyer (HNHM, NMP, ZISP).

Davraz dag: 9 ♂♂ 2 ♀♀, TURKEY W, 1900 m, 22.5.2001, Isparta, Davras dag, lgt. M.Snížek (Fig. 7) (cVS,cDW, cRL); 1 ♂, TR. Vil. Isparta, 24.5.2001, 10 km W of Yukarigökdre, Kasnak, J. Mertlik lgt. (cVS); 9 ♂♂ 2 ♀♀,Davraz, Isparta Turquie, VI.96, G. Dubault, B. Lassalle (sGD, cBL); 1 ♀, Davraz Dağ, w. Anatolien, 6.5.72 (cWH);4 ♂♂ 2 ♀♀, “Turquie (Isparta) Mt. Davras 2.000/2.200 H. Coiffait. 20.V.54” (MNHN).

Dedegöl dağlari: 1 ♂, Dedegöl-dağ, Nordsei, alpine zone, s üdl. Egirdir, 28.VII.1971, 1800–2300, Anatoliamer, Heinz lgt. (cWH); 1 ♂ 1 ♀, TR vill. Isparta, Yenisar Bademli env., Dipoyraz daği, Josef Mertlik lgt, 1.6.1995(cVS).

Bolkar daglari: 1 ♀, Dumbelek, Taurus Cil, Anatolien (MNHUB); 2 ♂♂ 1 ♀, Namrun, Anat.m., 10.5.–3.6.63,leg. F. Schubert (MNHUB, NMP); 3 ♂♂ 2 ♀♀, Turkei, Mersin, Findik pinari, Arslanköi, 2.5.01, leg. Megger(cDW, cRL).

Konya Villayet: 1♂1♀, “Turkey, 90 IV 18. Konya. Beysehir leg. Podlussány A.” (Fig.11) (HNHM).! unspecified localities: 1 ♂, “76” (green h)/ “T. phrygius pisidicus” (yellow h) (MCSNT); 1 ♂, “89” (green h)

/ “Tapinopt. phrygius m det. J. Müller” (h/p) (MCSNT); 1 ♀, “89” (green h); 1 ♂, “H 406” (h) (MCSNT).Diagnosis. BL 11.8–15.9 mm. Male genitalia. Median lobe of aedeagus short to medium-sized, apical lamella long, thin, more or less regu-

larly curved to left (Figs. 32–34) or rarely slightly sinuous before left-orientated tip (Figs. 30–31); apical orificedeflected to left lateral position; basal bulb relatively long, with basal orifice concave (Figs. 48, 50–52), apical partlonger than basal one (Figs. 50–52, 60–62), rarely almost equal in size (Figs. 48, 58), curved copulatory sclerite insubapical position (Figs. 49, 51, 59, 61), apex becoming thinner towards tip, hardly deflexed (Figs. 48, 58) orstraight (Figs. 50–52, 60–62). Aedeagus of syntype of T. phrygius pisidicus from Barla-dağlari (Fig. 31) showsslight deformation very probably caused by aging of this old specimen. Examination of another male from thislocality confirms that it is identical with other specimens of T. phrygius s.l. Right paramere relatively long, more orless curved medially, basal part thicker than apical one, with thickened marginal part in internal view, apical partpointed towards apex (Figs. 83–84, 98–99). Left paramere with shortened transverse apophysis, and long, obliqueparamedial apophysis (Figs. 113–114, 127–128).

Female genitalia. Sternum VIII very long, consisting of two chitinised plates characteristic in shape, connectedby membrane, proximal part with reduced, round “legs” (Fig. 146). Ovipositor with equal in size valvifer and sty-lus, as longer axis of former ± perpendicular to longer axis of latter (Figs. 157); valvifer with long, cylindrical prox-imal part, plate-like distal one, and small process across at distal part; stylus consisting of one, basal stylomere ofsubcylindrical shape (apical stylomere disappeared).

Taxonomic notes. In the course of studying the types of T. dipojranus s.str. and T. dipojranus cilicius, oneparatype of phrygius as well as a numerous specimens from the type localities of phrygius s.str., phrygius pisidicus



(Sultan dağlari, Kumalar dağlari, Barla daği, Davraz dağ, Dipojraz dağlari), and additional material from Bolkardağlari (type locality of dipojranus cilicius), we concluded that members of all above mentioned species and sub-species are conspecific with the typical phrygius since no significant morphological differencies were found. T.(Molopsis) phrygius is, just like the other species of this subgenus, a very variable species. Specimens from variouspopulations vary in size (significant size discrepancies were discovered even in the populations of different yearsfrom Davraz dag; specimens from 2001 were obviously smaller and more slender than those collected in 1996),and in shape of pronotum and median lobe of aedeagus. Understanding this, it was not possible to establish subspe-cies that would have more constant features. The population from Bolkar daglari (described as diporjanus ssp. cil-icius), which lives the farthest from the phrygius type-locality, is the most different from those of typical phrygius,but in fact there are no distinct features to fullfil the criteria for establishing a valid subspecies.

Distribution. Taurus mountain ranges about lakes Beyşehir, Eğirdir and Burdur: Ak dağ, Sultan dağlari,Kumalar dağlari, (“Phrygic Taurus”), Barla daği, Davraz dağ, Dipojraz dağlari (“Pisidic Taurus”), SW Turkey, Bol-kar dağlari (Bulghar Dagh by the ancient authors, “Cilician Taurus”), S Turkey.

Tapinopterus (Molopsis) machardi (Jeanne, 2005)(Figs. 13, 35, 53, 63, 85, 100, 115, 129)

Type locality. Turquie, Antalya, Col Karaovabeli, 1560 m.Taxonomic decisions. Molopsis machardi Jeanne, 2005: 383.

Tapinopterus (Molopsis) machardi (Jeanne, 2005), combinatio novum of Molopsis machardi Jeanne, 2005.References. Molopsis machardi Jeanne, 2005: 383, 385.Material examined. Four specimens.Type material. One paratype ♂ of Tapinopterus (Molopsis) machardi, Col. KARAOVABELI 1400 Antalya

Turquie, 03.05.96 B.Lassalle (p)//PARATYPE (red p) / Molopsis machardi m. Jeanne det. 2002 (p) (Fig.13) (cBL).Other material examined. 3 ♂♂, Col. KARAOVABELI 1400 Antalya Turquie, 03.05.96 G. Dubault &

D.Echaroux (cGD, cDE).Diagnosis. BL 14.5–16.2 mm. Male genitalia. Median lobe of aedeagus long in dorsal view, apical lamella long, straight, and thin, apical ori-

fice deflected to left lateral position (Figs. 35); basal bulb relatively long, with concave basal orifice, apical partlonger than basal one, curved copulatory sclerite in subapical position, apex becoming thinner towards tip, hardlybent downwards (Figs. 53, 63). Right paramere long, regularly curved medially, thicker basally than apically, withwell-differentiated, curved subbasal part, apical part shortly acute towards apex (Figs. 85, 100). Left paramere withshort transverse apophysis and long oblique paramedial apophysis (Figs. 115, 129).

Female genitalia. Unknown.This species differs from all other species of Molopsis by the presence of very fine setae underneath the ony-

chium and by the specific shape of the median lobe of aedeagus and right paramere (Figs. 35, 53, 63, 85, 100).Taxonomic notes. The single paratype studied possesses different numbers of setae underneath the onychium,

respectivelly: 1+1 (left anterior leg), 1+1 (right anterior leg), 1+1 (left middle leg), 1+0 (right midlle leg), 2+1 (leftposterior leg), 1+0 (right posterior leg). The setae vary in size and thickness, and generally are shorter and finerthan typical setae in the most species of Tapinopterus s.l. We consider that the presence / absence of ventral setaeon onychium may have different phyletic value to specify monophyletic lines in Pterostichini. This process reflectsmorphological regressions appeared in different, unrelated clades of the tribe as a result of an adaptation to differ-ent environments. The two states of this character occur either in species from different subgenera of the samegenus (Ancholeus Dejean, 1828 and Poecilus Bonelli, 1810 of Poecilus s.l.) or among species from one and samesubgenus—Phonias Gozis, 1886 (of the genus Pterostichus Bonelli, 1810; cfr. Hůrka, 1996), Molopsis Schatzmayr(this paper).

Distribution. South Taurus, Karaovabeli pass between Kaş and Elmali, W of Antalya (“Lycian Taurus”).



PLATE 2. Habitus of Tapinopterus species (Figs. 13–24). Fig. 13. Tapinopterus (Molopsis) machardi (paratype); Fig. 14. T.(Molopsis) oyukluensis sp. nov. (holotype); Fig. 15. T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 16. T. (Mol-opsis) molopinus (holotype of Tapinopterus dipojranus brussanus); Fig. 17. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 18.T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Fig. 19. T. (Molopsis) chaudoiri sp. nov. (paratype, Oylat); Fig. 20. T.(Molopsis) chaudoiri sp. nov. (paratype, Gök dağlari); Fig. 21. T. (Molopsis) wiedemanni (lectotype); Fig. 22. T. (Molopsis)wiedemanni (Turkey, Abant); Fig. 23. T. (Molopsis) wiedemanni (Turkey, Sundiren dağlari); Fig. 24. T. (Molopsis) relegatus sp.nov. (holotype).



PLATE 3. Median lobe of aedeagus (Figs. 25–36), dorsal view (Figs. 25–28, 30–36) and ventral view (Fig. 29) of Tapinopterusspecies. Fig. 25. Tapinopterus (Pseudomolopsis) rebellis (Greece, Kalamata); Fig. 26. T. (Molopsis) molopiformis (Tmolos-Gbg., Lydien, West-Kleinasien, Weirather); Fig. 27. T. (Molopsis) molopiformis (Coll Türk 1888); Figs. 28–29. T. (Molopsis)aenigmaticus sp. nov. (holotype); Fig. 30. T. (Molopsis) phrygius (syntype); Fig. 31. T. (Molopsis) phrygius (syntype of T.phrygius pisidicus); Fig. 32. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 33. T. (Molopsis) phrygius (Turkey,Beyşehir); Fig. 34. T. (Molopsis) phrygius (paratype of Tapinopterus dipojranus cilicius); Fig. 35. T. (Molopsis) machardi(paratype); Fig. 36. T. (Molopsis) oyukluensis sp. nov. (holotype). Scale bar: 1 mm.

Tapinopterus (Molopsis) oyukluensis sp.nov.(Figs. 14, 36, 54, 64, 86, 101, 116, 130)

Type locality. Faşikan pass, 1600 m, Konya, Turkey.Material examined. One specimen.Type material. Holotype ♂, Faşikan pass, 1600 m, Konya Turquie, VII.86 B.Lassalle (p)/ HOLOTYPE Tap-

inopterus (sg. Molopsis) oyukluensis n. sp. Lohaj, Guéorguiev, Dubault & Lassalle, 2007 (red p)/ (Fig. 14). Theholotype is deposited in cBL.

Description. BL 13 mm, maximum width 4.1 mm. Head, pronotum and elytra dark brown, shiny, antennae,legs and abdomen piceous brown. Head large, robust, almost as long as wide, shiny, smooth, with very finemicrosculpture. Antennae relatively short, reaching hind angles of pronotum, with dense, decumbent pubescencefrom antennomere 4.

Pronotum weakly transverse, wider than long, ratio PL/PW 0.73, with rounded lateral sides, shiny, smooth,with very fine transversal wringles along the midline. Anterior edge strongly sinuated, with prominent, at apexrounded angles. Posterior angles small, almost rectangular. Base of pronotum with single deep linear posterolateralimpression on each side of midline, reaching basal third, smooth, disc without transversal wrinkles. Midline dis-tinct, not reaching basal or anterior margins.

Elytra oval, slightly flattened, with maximum width in posterior third, with distinct, impunctate striae. Third

elytral interval with two setigerous pores situated in apical half of elytra, 2nd behind the middle and 3rd behind apicalthird, situated in second elytral striae. One irregular setigerous pore is also present in the middle of first left elytral



stria. Umbilicate series consists of 17, respective 16 setigerous punctures, in the first fourth interrupted, with distri-bution: left side 6 + 1 + 10, right side 6 + 10.

Male genitalia. Median lobe of aedeagus in dorsal view relatively short, apical lamella wide, directly curved toleft (Figs. 36); apical orifice deflected to left lateral position, ventral margin forming right angle at medial part,apex regularly deflexed (Figs. 54, 64). Right paramere with basal part thicker than apical part, in middle constrictedand curved, apex pointed towards tip (Fig. 86, 101). Left paramere subelongate, transverse apophysis reduced,paramedial apophysis, oblique, well-developed (Figs. 116, 130).

Female genitalia. Unknown.Etymology. Topotypic, referring to the type locality, the mountain range Oyuklu daği, northern part of Taurus

Mountains, ca 100 km south of Konya.Diagnosis. T. oyukluensis sp. nov. is closely related to T. phrygius, a species widely distributed in various

mountain ranges of the Taurus (see also Check list). This species differs from all other species of Molopsis by theshape of the median lobe of aedeagus and left paramere; median lobe in dorsal view (Fig. 36) with apical lamellawide, directly curved to left (this state occurs also in T. phrygius, but here the lamella is thinner and either regularlycurved to left (see Figs. 32–34) or slightly sinuous before apex (see Figs. 30–31); left paramere subelongate, withreduced transverse apophysis (Figs. 116, 130).

Distribution. Type specimen was collected in Faşikan geçidi (pass) between the villages of Taşkent and Sariv-eliler, ca 30 km southwest of ridge of mountain range Oyuklu daği (Central Turkey, villayet Konya).

Tapinopterus (Molopsis) molopinus (Chaudoir, 1868)(Figs. 15–17, 37–38, 65–67, 70–72, 87–88, 102–103, 117–118, 131–132, 140, 147, 152, 158, 164, 168, 170)

Type locality. Constantinople.Taxonomic decisions. Feronia (Haptoderus) molopina Chaudoir, 1868: 245.

Pterostichus olympicus Kraatz, 1875: 418 (typ. loc.: “Olymp”) (synonymy by Apfelbeck, 1904: 244, 405).Tapinopterus (Molopsis) dipojranus brussanus Straneo, 1986: 126 (typ. loc.: “Brussa”), synonymum novum.

References. Feronia (Haptoderus) molopina: Chaudoir, 1868: 245; Ganglbauer, 1889: 52. Pterostichus olympicus: Kraatz, 1875: 418; Ganglbauer, 1889: 52.Feronia molopina: Kraatz, 1875: 419.Feronia (Cophosus) olympica: Marseul, 1880: 306.Feronia (Steropus) olympica: Marseul, 1882: 47.Feronia (Steropus) molopina: Marseul, 1882: 48.Pterostichus (Pterostichus) molopinus: Seidlitz, 1887: 40; Seidlitz, 1888: 44.Pterostichus (Pterostichus) olympicus: Seidlitz, 1887: 40; Seidlitz, 1888: 44.Pterostichus (Haptoderus) molopinus: Heyden et al., 1891: 35.Tapinopterus (Crisimus) olympicus: Heyden et al., 1891: 38.Haptoderus molopinus: Heyden et al. 1891: 397.Pterostichus (Tapinopterus) molopinus: Apfelbeck, 1904: 244.Pterostichus (Tapinopterus) molopinus: Heyden et al., 1906: 82.Platysma (Tapinopterus) molopinum: Jakobson, 1907: 352.Pterostichus (Tapinopterus) molopinus: Breit, 1914: 55; Csiki, 1930: 707.Tapinopterus molopinus: G. Müller, 1923: 111.Crisimus (Tapinopterus) molopinus: Schatzmayr, 1929: 330.Tapinopterus (Crisimus) molopinus: G. Müller, 1932: 216–217.Platysma (Tapinopterus) molopinus: Straneo, 1936: 156.Tapinopterus molopinus molopinus: Mařan, 1940: 57.Tapinopterus molopinus olympicus: Mařan, 1940: 57.Tapinopterus molopinus: Schatzmayr, 1942: 45.Tapinopterus (Molopsis) molopinus: Schatzmayr, 1943: 119; Straneo, 1986: 122; Kirschenhofer, 1991: 5; Lorenz,

1998: 267; Casale & Vigna Taglianti, 1999: 381.Tapinopterus (Molopsis) molopinus olympicus: Schatzmayr, 1943: 119; Kirschenhofer, 1991: 5; Bousquet, 2003:

518.



Tapinopterus (Molopsis) dipojranus brussanus: Straneo, 1986: 126; Kirschenhofer, 1991: 6; Lorenz, 1998: 267;Bousquet, 2003: 518.

Tapinopterus (Molopsis) molopinus molopinus: Bousquet, 2003: 518.Molopsis molopinus molopinus: Jeanne, 2005: 384.Molopsis molopinus olympicus: Jeanne, 2005: 384.Tapinopterus (Molopsis) dipojranus brussanus: Lorenz, 2005: 288.Tapinopterus (Molopsis) molopinus molopinus: Lorenz, 2005: 288.Tapinopterus (Molopsis) molopinus olympicus: Lorenz, 2005: 288.

PLATE 4. Median lobe of aedeagus (Figs. 37–44), dorsal view (Figs. 37–39, 41–44) and ventral view (Fig. 40) of Tapinopterusspecies. Fig. 37. T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 38. T. (Molopsis) molopinus (holotype of Tap-inopterus dipojranus brussanus); Fig. 39. T. (Molopsis) chaudoiri sp. nov. (paratype, Oylat); Fig. 40–41. T. (Molopsis) wiede-manni (lectotype of Feronia wiedemanni); Fig. 42. T. (Molopsis) wiedemanni (Turkey, Abant Lake); Fig. 43. T. (Molopsis)wiedemanni (Turkey, Aladağlari); Fig. 44. T. (Molopsis) wiedemanni (Turkey, Sundiren dağlari). Scale bars: 1 mm (Figs. 37–39); 2 mm (Figs. 40–44).

Material examined. 97 specimens.Type material. Holotype ♂ of Feronia molopina, designated by monotypy, well preserved, “molopinus

Chaud. Constantinople Sturm.” (h label pinned on the bottom of box) / HOLOTYPE Feronia (Haptoderus) mol-opina Chaudoir 1868 G.Dubault désign. 2003 (red p)/ Tapinopterus (Molopsis) molopinus Chaudoir G.Dubaultdétermin. (p) (Fig.15) (MNHN, “Collection Chaudoir”, box no 216 “Argutor”).



Holotype ♂ of Tapinopterus (Molopsis) dipojranus brussanus, “Brussa Anatolien G.Frey IV.32.” (p)/ “Tap-inopterus molopinus Chd.” (h) / “Holotypus T. dipojran. s. brussan. (red p/h) / “Tapinopterus dipojranus s. brus-sanus (h), det. S. L. Straneo 1985 Holotypus” (p/h)/ TAPINOPTERUS (sg. MOLOPSIS) molopinus (Chaudoir,1868) R.LOHAJ det.2003 (p) (Fig.16) (MCSNM).

Other material examined. Uludag: 1 ♂, Turkei 1848 (NMP); 1♂, Asia minor (HNHM); 1 ♂, BALKANFRIV. (HNHM); 3 ♂♂, Olympe (ZISP); 2 ♂♂, Bithynischen Olymp, leg. Merkl, coll. Leonhard (DEI); 1 ♀,Olymp, coll. O.Leonhard (DEI); 1 ♂, Olympos Bithyn. Asia min. Dr. Jureček 1931 (NMP); 4 ♂♂, BRUSSA, 1870,leg. Pável (HNHM); 5 ♂♂ ♀♀, “Uludag Bursa 1.500 West. Turkey J. y S. Klapperich.” (MNHN); 6 ♂♂ ♀♀, “UluDagh Brousse 1400-VI66” (MNHN); 6 ♂♂ ♀♀, “Ulu Dagh Turquie 1700 m VI 68” (MNHN); 2 ♂♂, Merkl.92,Türkei (NMNHS); 1 ♂, Turcia Merkl 98. (MCSNT); 1 ♂, “rebellis Turcia” (MNHN); 1 ♂, “Museum Paris 1906Coll Léon Fairmaire” / “Pter. olympicus Turquie” (MNHN); 1 ♂, “olympicus Kr. Turkei Merkl.” (MNHN); 1 ♀,Türkei, Ulu dagh 2000 m, leg. Kenyery, VI.1963 (NMP); 3 ♂♂ 1 ♀, Anatolia occ, Uludagh bei Bursa (gipfelzone)1500 m, 28.V.1966, Blumental leg. (NMP); 8 ♂♂ 4 ♀♀, Turchia, Bursa Uludag, 29.5.1974, lg. Sama, Coll.Gudenzi (cMP); 3 ♂♂, Anatol. Bursa Uludag 25.5.82 Drioli (MCSNT); 1 ♂ 1 ♀, Turkei, Uludag, 13.IV.1983,H.+L.Freude (cMP); 8 ♂♂ 7 ♀♀, Uludag 1500, Bursa Turquie, VII 87 G.Dubault & D.Echaroux lgt. (cGD, cDE);1 ♀, Uludağ 1500 m, (Bursa) 15.V.89, leg. R.Pittino (cWH); 2 ♀♀, Uludag 1500 Bursa VI 90” / “Lassale” (cBL);7 ♂♂ 3 ♀♀, TR (Bursa) Uludağ b. Soğurpinar, 1200–1400 m, 26.IV.1992 Heinz leg. (cRL); 4 ♂♂, Tr.occ., Bursaenv. Uludag, 27.5.1994, Skoupý leg. (cVS); 1 ♂, “NW Turkey—Bursa Uludag pineto—abietum, 1500 m 3.10.97,D. Čatlos lgt. (cRL) (Fig. 17); 1 ♂, Tr.occ, Bursa Uludag 1650 m, 15.5.2000, Skoupý leg. (cVS); 1 ♂, Tr.occ, BursaUludag 2100 m, 15.5.2000, Skoupý leg. (cVS); 1 ♀, Turkey, Cumalikizik, Bursa, Ulu dağ, 24.5.2004, 5–900 m,T.Tichý (cEH).

Diagnosis. BL 10.8–15 mm.Male genitalia. Median lobe of aedeagus long in dorsal view, with long, thin, almost straight apical lamella,

apical orifice deflected to left lateral position (Figs. 37–38); basal bulb relatively long, with concave basal orifice,apical part longer, copulatory sclerite curved, situated in medial part, apex slighlty bent downwards (Figs. 65–67,70–72). Right paramere relatively long, more or less strongly curved and constricted medially, with thicker basaland thinner apical parts (Figs. 87–88, 102–103). Left paramere with short, more or less reduced transverse apophy-sis and fairly long, oblique paramedial apophysis (Figs. 117–118, 131–132).

Female genitalia. Tergum VIII with convex distal margin and short, thin proximal “legs” (Fig. 140). SternumVIII consisting of two chitinised parts connected by membrane, without “legs” on proximal margin (Fig. 147).Syntergum IX+X long, narrow, bearing pair of small, articulated, distant each other ovipositors (Fig. 152). Ovipos-itor with large valvifer and smaller stylus, as longer axis of former ± perpendicular to longer axis of latter (Figs.158); valvifer with short cylindrical part, more acute internal margin, less acute external margin, and with devel-oped, slightly curved externally, process at distal part; stylus consist of one, basal, elongate stylomere (apical sty-lomere disappeared), having incised outer margin. Spermatheca with seminal canal and receptaculum discrete;seminal canal long; receptaculum shorter than seminal canal, slightly curved apically (Fig. 164).

This species differs from its closest relatives, T. chaudoiri sp. nov. and T. relegatus sp. nov. in the structure ofthe male and female genitalia. T. molopinus has a smaller aedeagus, with relatively shorter and thinner apicallamella, about 1/3 entire length of lobe (in dorsal view). T. chaudoiri sp. nov. has a larger aedeagus, with apicallamella about 2/5 that of median lobe (compare Figs. 37–39). In lateral view, the aedeagus of T. molopinus issmaller, with narrower medial part and shortened apical lamella (Figs. 65–74). The transverse apophysis on leftparamere in T. molopinus is more or less reduced while in T. chaudoiri possesses a longer, well-developed trans-verse apophysis (Figs. 117–120, 131–134). The females of this species are easily distinguished from the females ofT. chaudoiri in the absence of incision at the apex of elytra and apex of sternum VII (Figs. 168–171). Besides, ster-num VII has finer posterior border.

For the distintions between T. molopinus and T. relegatus, see paragraph “Diagnosis” under the latter species.Taxonomic notes. The first mention of this taxon has been made by Sturm (1843: 25), but the species was

described by Chaudoir in 1868 as Feronia (Haptoderus) molopina based on single male specimen. The last authorwrote “C’est le Steropus brevis du catalogue de Sturm, indiqué comme venant de Constantinople, et que j’ai achetéà la vente de sa collection.”. In view of the relativeness of the geographical designation in the old-days, it should benoted that the type locality of T. molopinus is doubtful. We are unaware of any subsequent record or material ofMolopsis in collections from Constantinople (= Istanbul). Regarding the present bionomy of the species, we sup-



pose that the type material was collected in the area of the Uludag Mt.; the same has already been supposed(Apflelbeck, 1904: 244, note 1).

PLATE 5. Median lobe of aedeagus (Figs. 45–54) in left lateral view of Tapinopterus species. Fig. 45. Tapinopterus (Pseudo-molopsis) rebellis (Greece, Kalamata); Fig. 46. T. (Molopsis) molopiformis (Tmolos-Gbg., Lydien, West-Kleinasien, Wei-rather); Fig. 47. T. (Molopsis) aenigmaticus sp. nov. (holotype); Fig. 48. T. (Molopsis) phrygius (syntype); Fig. 49. T.(Molopsis) phrygius (syntype of T. phrygius pisidicus); Fig. 50. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 51. T.(Molopsis) phrygius (Turkey, Beyşehir); Fig. 52. T. (Molopsis) phrygius (paratype of Tapinopterus dipojranus cilicius); Fig. 53.T. (Molopsis) machardi (paratype); Fig. 54. T. (Molopsis) oyukluensis sp. nov. (holotype). Scale bar: 1 mm.

The holotype of Feronia molopina was found in MNHN. Meantime, two of us (BG, RL) tried to find the holo-type of Pterostichus olympicus in DEI and MNHUB. According to Dr. Lothar Zerche (personal communication),no type of this species exists in DEI. The search of BG in NMHUB led nowhere too. The material of Tapinopterusfrom DEI, very probably including also the type of Pterostichus olympicus, was many years ago loaned by Dr.Jozef Mařan (Praha, Czech Republic) and maybe have never been returned (Dr. L. Zerche, personal communica-tion). According to the information provided by Dr. Josef Jelinek, the former curator of Coleoptera in NMP, theTapinopterus loan was in possession of Dr. J. Mařan at the time of his death. The type material was, however, notlocated after his demise.

Kraatz (1875) did not indicate clearly the exact massif from which the type series of Pterostichus olympicuscomes, e.g. the Olymp Mt. in Central Greece or the Uludag Mt. (Bythinic Olymp near Bursa by the ancientauthors) in Northwestern Anatolia. The title of the Kraatz paper has points that the author dealt with species fromEurope (“Drei neue europäische Pterostichus Bon.“) and author noted: „Mein Ex. stammt aus der Schaum´schenSammlung und wurde wahrscheinlich von Dr. Krüper aufgefunden“. Dr. Theobald Johaness Krüper (1829 –1921) aGerman ornithologist and entomologist and long-time curator and director of University Museum in Athens. He



collected in Greece and also often in Turkey. Authors (RL, BG) could not find the type sample in the studied col-lections which may have successfully associated with the specimen/s from the type series. For this reason, until thetype material is located and examined, we prefer to treat this taxon as synonym of T. molopinus.

Pterostichus olympicus Kraatz was first synonymized with Pterostichus molopinus Chaudoir by Apfelbeck(1904: 244, 405). Judging from the context of the Apfelbeck’s work we suppose that this author examined typematerial of this taxon. The most of the subsequent authors (see the list above) have not accepted this opinion. OnlyCsiki (1930: 707) and Müller (1932: 217) agreed with the Apfelbeck’s decision. Mařan (1940: 57) first removed T.olympicus from the synonymy and considered it as subspecies of T. molopinus. His view was followed by all subse-quent authors (Schatzmayr, 1943: 119; Kirschenhofer, 1991: 5; Bousquet, 2003: 518; Jeanne, 2005: 384).

Pterostichus (Crisimus) kyparissis Jedlička, 1963 was described by Jedlička (1963: 19) by single female spec-imen from the southwestern part of the Peloponnesos. The author compared the new species with T. molopinus andT. rebellis, and stated that it possesses setigerous pore in scutellar stria. Despite still not clear relationships of T.kyparissis, we exclude it as belonging to Molopsis based on the last mentioned character.

Distribution. This species inhabits the massif of Uludağ near Bursa, NW Anatolia.

PLATE 6. Median lobe of aedeagus Figs. 55–64) in right lateral view of Tapinopterus species. Fig. 55. Tapinopterus (Pseudo-molopsis) rebellis (Greece, Kalamata); Fig. 56. T. (Molopsis) molopiformis (Tmolos-Gbg., Lydien, West-Kleinasien, Wei-rather); Fig. 57. T. (Molopsis) aenigmaticus sp. nov. (holotype); Fig. 58. T. (Molopsis) phrygius (syntype); Fig. 59. T.(Molopsis) phrygius (syntype of T. phrygius pisidicus); Fig. 60. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 61. T.(Molopsis) phrygius (Turkey, Beyşehir); Fig. 62. T. (Molopsis) phrygius (paratype of Tapinopterus dipojranus cilicius); Fig. 63.T. (Molopsis) machardi (paratype); Fig. 64. T. (Molopsis) oyukluensis sp. nov. (holotype). Scale bar: 1 mm.



PLATE 7. Median lobe of aedeagus (Figs. 65–79), left lateral view (Figs. 65–69, 75) and right lateral view (70–74, 76–79) ofTapinopterus species. Figs. 65, 70. T. (Molopsis) molopinus (holotype of Feronia molopina); Figs. 66, 71. T. (Molopsis) mol-opinus (“olympicus”, HNHM); Figs. 67, 72. T. (Molopsis) molopinus (holotype of Tapinopterus dipojranus brussanus); Figs.68, 73. T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Figs. 69, 74. T. (Molopsis) chaudoiri sp. nov. (paratype, Oylat);Figs. 75–76. T. (Molopsis) wiedemanni (lectotype of Feronia wiedemanni); Fig. 77. T. (Molopsis) wiedemanni (Turkey, AbantLake); Fig. 78. T. (Molopsis) wiedemanni (Turkey, Aladağlari); Fig. 79. T. (Molopsis) wiedemanni (Turkey, Sundiren dağlari).Scale bars: 1 mm (Figs. 65–74); 0.5 mm (Figs. 75–79).

Tapinopterus (Molopsis) chaudoiri sp.nov.(Figs. 18–20, 39, 68–69, 73–74, 89–90, 104–105, 119–120, 133–134, 141, 148, 159, 169, 171)

Type locality. Forest between Domanic and Tahtaköprü, 1000–1500 m, NW Turkey.Material examined. 65 specimens.Type material. Holotype ♂, TR Wld zw. Domanic u. Tahtaköprü, IV 1973, 1000–1500 m, Heinz leg. (p)/

HOLOTYPE Tapinopterus (Molopsis) chaudoiri n. sp., Lohaj, Guéorguiev, Dubault & Lassalle des. 2007 (red p)/(cWH).

Paratypes: 24 ♂♂ 12 ♀♀, TR Wld zw. Domanic u. Tahtaköprü, IV 1973, 1000–1500 m, Heinz leg. (Fig. 18)(p)/ PARATYPE Tapinopterus (Molopsis) chaudoiri n. sp., Lohaj, Guéorguiev, Dubault & Lassalle des. 2007 (red



p)/, (MCSNM, NMNHS, cBL, cGD, cRL, cWH), 9 ♂♂ 5 ♀♀, Turkey, Domanic—Bursa, 03.05.98, G.Dubault &D. Echaroux lgt. (p)/ PARATYPE Tapinopterus (Molopsis) chaudoiri n.sp., Lohaj, Guéorguiev, Dubault & Lassalledes. 2007 (red p)/ (cDE, cGD), 3 ♂♂, Oylat 1200 Bursa Turquie, 07.05.98 G. Dubault (Fig. 19) (p)/ PARATYPETapinopterus (Molopsis) chaudoiri n.sp., Lohaj, Guéorguiev, Dubault & Lassalle des. 2007 (red p) (cBL, cDE,cGD), 2 ♂♂ 8 ♀♀, Anatolia occ. Heinz leg. (p)/ Gök-dağ: Keltepe, ca. 1100–1400 m, 14.IV.1982 (h)/ PARATYPETapinopterus (Molopsis) chaudoiri n.sp., Lohaj, Guéorguiev, Dubault & Lassalle des. 2007 (red p) (Fig. 20)(MCSNM, NMNHS, cRL, cWH), 1 ♂, Cangaz, Anat 1962 (h)/ Tapinopterus olympicus Kr. (h) det. Ing. Jedlička(p)/ PARATYPE Tapinopterus (Molopsis) chaudoiri n.sp., Lohaj, Guéorguiev, Dubault & Lassalle des. 2007 (redp) (NMP).

Description. BL 11.3–15 mm (Holotype 12.6 mm), maximum width up to 5.5 mm. Head, pronotum and elytrablack, shiny, some females matt, antennae, legs and abdomen in some specimens paler, piceous brown. Head large,robust, almost as long as wide, shiny, smooth, with very fine microsculpture. Antennae relatively short, reachinghind angles of pronotum.

Pronotum weakly transverse, slightly wider than long, ratio PL/PW 0.65–0.76 (HT 0.67), sides rounded, shiny,smooth, with very fine transversal wrinkles along the midline. Anterior margin concave, anterior angles prominent,rounded at apex.. Posterior angles strongly prominent, sharp, acute-angled. Base of pronotum with single deep lin-ear posterolateral impression on each side of midline, reaching basal third, smooth. Midline distinct, not reachingbasal or anterior margins.

Elytra elongate oval, with maximum width in apical third, slightly convex, with distinct, finely punctate striae.Apex of elytra with deep and symmetric incision, which is wide in females, and very narrow, but visible in males(Fig. 169). Third elytral interval with two setigerous punctures situated at apical half of elytra, 2nd behind middleand 3rd in apical fifth-sixth of elytral length. Umbilicate series consists of 14–17 setigerous punctures at each side,in the middle widely interrupted.

Distal margin of last visible abdominal sternum (sternum VII) with thick posterior border, surface smooth, notpunctate, with two pairs of SP; apex of sternum VII in females with very deep circular incision at tip (Fig. 171).

Male genitalia. Median lobe of aedeagus in dorsal view long, with long, wide, almost straight apical lamella,apical orifice deflected to left lateral position (Figs. 39); basal bulb short, with basal orifice slightly concave, apicalpart longer, copulatory sclerite curved, situated in medial part, apex more or less bent downwards (Figs. 68–69, 73–74). Right paramere relatively long, more or less strongly curved and constricted medially, with thicker basal partand thinner apical one (Figs. 89–90, 104–105). Left paramere with long transverse apophysis and fairly long,oblique paramedial apophysis (Figs. 119–120, 133–134).

Female genitalia. Tergum VIII with regularly convex distal margin and very short proximal “legs” (Fig. 141).Sternum VIII consists of two round, chitinised parts connected with membrane, without “legs” on proximal margin(Fig. 148). Ovipositor with valvifer and stylus, as longer axis of former more or less perpendicular to longer axis oflatter (Figs. 159); valvifer with short cylindrical part, internal margin strongly one-humped, external margin regu-larly convex, and with clear process across at distal part; stylus consist of one, basal, elongate stylomere (apicalstylomere disappeared), with slightly incised internal margin.

Etymology. Patronymic, dedicated to baron Maximilien Stanislavovitch de Chaudoir (1816–1881), famousRussian entomologist and outstanding specialist on the carabid beetles.

Diagnosis. This species is distinct from its closest relatives, T. molopinus and T. relegatus sp. nov. in the struc-ture of the male and female genitalia. T. chaudoiri sp. nov. has a larger aedeagus, with longer and wider apicallamella, with length about 2/5 of entire length of lobe (in dorsal view); T. molopinus has smaller aedeagus, with rel-atively shorter and thinner apical lamella only 1/3 of length of median lobe (compare Figs. 37–39). In lateral view,the aedeagus of the new species is larger, with wider medial part and prolonged apical lamella (Figs. 65–74). Thetransverse apophysis on left paramere in T. chaudoiri sp. nov. well-developed while in T. molopinus it is more orless reduced (Figs. 117–120, 131–134).

T. chaudoiri sp. nov. is easily distinguished from the closely distributed T. molopinus from the Uludağ Mt. bythe presence of deep, symmetric incision on the apex of elytra (in both sexes) and by the presence of deep circularincision on the apex of sternum VII in females, which is missing in T. molopinus (Figs. 168–171). Besides, the ster-num VII in T. chaudoiri sp. nov. has thicker posterior border.

T. chaudoiri sp. nov. differs also morphologically from T. relegatus sp. nov. in the different shape of tergumVIII (Figs. 1410142), sternum VIII (Figs. 148–149), and valvifer and stylus of ovipositor (Figs. 159–160).



PLATE 8. Right paramere (internal face) (Figs. 80–94) of Tapinopterus species. Fig. 80. Tapinopterus (Pseudomolopsis) rebel-lis (Greece, Kumani); Fig. 81. T. (Molopsis) molopiformis (Coll Türk 1888); Fig. 82. T. (Molopsis) aenigmaticus sp. nov. (holo-type); Fig. 83. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 84. T. (Molopsis) phrygius (paratype of Tapinopterusdipojranus cilicius); Fig. 85. T. (Molopsis) machardi (paratype); Fig. 86. T. (Molopsis) oyukluensis sp. nov. (holotype); Fig. 87.T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 88. T. (Molopsis) molopinus (holotype of Tapinopterusdipojranus brussanus); Fig. 89. T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Fig. 90. T. (Molopsis) chaudoiri sp. nov.(paratype, Oylat); Fig. 91. T. (Molopsis) wiedemanni (lectotype of Feronia wiedemanni); Fig. 92. T. (Molopsis) wiedemanni(Turkey, Abant Lake); Fig. 93. T. (Molopsis) wiedemanni (Turkey, Aladağlari); Fig. 94. T. (Molopsis) wiedemanni (Turkey,Sundiren dağlari). Scale bar: 0.5 mm.



PLATE 9. Right paramere (external face) (Figs. 95–109) of Tapinopterus species. Fig. 95. Tapinopterus (Pseudomolopsis)rebellis (Greece, Kumani); Fig. 96. T. (Molopsis) molopiformis (Coll Türk 1888); Fig. 97. T. (Molopsis) aenigmaticus sp. nov.(holotype); Fig. 98. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 99. T. (Molopsis) phrygius (paratype of Tap-inopterus dipojranus cilicius); Fig. 100. T. (Molopsis) machardi (paratype); Fig. 101. T. (Molopsis) oyukluensis sp. nov. (holo-type); Fig. 102. T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 103. T. (Molopsis) molopinus (holotype ofTapinopterus dipojranus brussanus); Fig. 104. T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Fig. 105. T. (Molopsis)chaudoiri sp. nov. (paratype, Oylat); Fig. 106. T. (Molopsis) wiedemanni (lectotype of Feronia wiedemanni); Fig. 107. T. (Mol-opsis) wiedemanni (Turkey, Abant Lake); Fig. 108. T. (Molopsis) wiedemanni (Turkey, Aladağlari); Fig. 109. T. (Molopsis)wiedemanni (Turkey, Sundiren dağlari). Scale bar: 0.5 mm.



PLATE 10. Left paramere (internal face) (Figs. 110–123) of Tapinopterus species. Fig. 110. Tapinopterus (Pseudomolopsis)rebellis (Greece, Kumani); Fig. 111. T. (Molopsis) molopiformis (Coll Türk 1888); Fig. 112. T. (Molopsis) aenigmaticus sp.nov. (holotype); Fig. 113. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 114. T. (Molopsis) phrygius (paratype of Tap-inopterus dipojranus cilicius); Fig. 115. T. (Molopsis) machardi (paratype); Fig. 116. T. (Molopsis) oyukluensis sp. nov. (holo-type); Fig. 117. T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 118. T. (Molopsis) molopinus (holotype ofTapinopterus dipojranus brussanus); Fig. 119. T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Fig. 120. T. (Molopsis)chaudoiri sp. nov. (paratype, Oylat); Fig. 121. T. (Molopsis) wiedemanni (lectotype of Feronia wiedemanni); Fig. 122. T. (Mol-opsis) wiedemanni (Turkey, Abant Lake); Fig. 123. T. (Molopsis) wiedemanni (Turkey, Sundiren dağlari). Scale bar: 0.5 mm.

The several unique morphological differences between the populations from the massifs of Gök Daglari andUludağ as well as the lack of any records from the intermediate area between them suppose presence of two differ-ent and apparently isolated (allopatric) species as the different aedeagus manifests the isolation.

Distribution. This species inhabits southern and eastern slopes of Uludağ mountain range (pass betweenDomanic and Tahtaköprü, surrounding of Oylat) and mountain of Gök Daglari, situated south of the Sapanca Lake,(villayet Sakarya), NW Anatolia. Authors tried to designate the position of the locality “Cangaz”, by which is onemale paratype specimen labelled, but without success.



PLATE 11. Left paramere (external face) (Figs. 124–137) of Tapinopterus species. Fig. 124. Tapinopterus (Pseudomolopsis)rebellis (Greece, Kumani); Fig. 125. T. (Molopsis) molopiformis (Coll Türk 1888); Fig. 126. T. (Molopsis) aenigmaticus sp.nov. (holotype); Fig. 127. T. (Molopsis) phrygius (holotype of T. dipojranus); Fig. 128. T. (Molopsis) phrygius (paratype ofTapinopterus dipojranus cilicius); Fig. 129. T. (Molopsis) machardi (paratype); Fig. 130. T. (Molopsis) oyukluensis sp. nov.(holotype); Fig. 131. T. (Molopsis) molopinus (holotype of Feronia molopina); Fig. 132. T. (Molopsis) molopinus (holotype ofTapinopterus dipojranus brussanus); Fig. 133. T. (Molopsis) chaudoiri sp. nov. (paratype, Domanic); Fig. 134. T. (Molopsis)chaudoiri sp. nov. (paratype, Oylat); Fig. 135. T. (Molopsis) wiedemanni (lectotype of Feronia wiedemanni); Fig. 136. T. (Mol-opsis) wiedemanni (Turkey, Abant Lake); Fig. 137. T. (Molopsis) wiedemanni (Turkey, Sundiren dağlari). Scale bar: 0.5 mm.

Tapinopterus (Molopsis) relegatus sp.nov.(Figs. 24, 142, 149, 160, 165)

Type locality. Dorukhan Pass near Zonguldak, NE Turkey.Material examined. One specimen.Type material. Holotype ♀, relatively well preserved, only apical borders of elytra bitten, Turkey, col

Dorukhan, Zonguldak, VII.1990 (p)/ Holotype Tapinopterus (Molopsis) relegatus sp. n. Lohaj, Guéorguiev,Dubault & Lassalle, 2007” (red p) (Fig. 24) (cBL).

Description. BL 12.2 mm; maximum width 4 mm. Body smooth and glabrous (excluding antennae), head andpronotum without microsculpture, elytra with distinct fine isodiametric microsculpture. Head, pronotum and elytrablack, shiny, antennae, palpi, legs and body ventrally piceous brown.

Head large, robust, subconvex, 1.33 times longer than wide; disc smooth. Mandibles stout and protruding.Antennae relatively short, not reaching hind angles of pronotum, with dense, decumbent pubescence from antenno-



mere 4, antennomere 2 nearly two times shorter than scapus. Length of eyes almost equal to length of scapus; tem-porae not protruding, shorter than length of eye. Frontal furrows short, very smooth. Mentum with short bifid tooth.Pronotum 1.48 times wider as long and 1.53 times wider than head, subcordate and subconvex, smooth. Lateralsides more convergent posteriorly than anteriorly; ratio anterior margin / basal margin 1.09. Anterior margin mod-erately sinuate, with rounded and slightly prominent angles. Posterior margin almost straight; angles sharplypointed, prominent outside. Basal surface with single impunctate basal fovea on each side of midline, fovea withsingle linear posterolateral impression slightly divergent anteriorly and not reaching basal margin. Midline distinctand deeper in basal third, not reaching anterior, and hardly reaching basal margin.

Elytra suboval, coalesced along suture, 1.32 times longer than wide and 1.41 times wider than pronotum; discconvex. Shoulders obtusely rounded. Scutellar stria absent; basal border not reaching scutellum; striae fine, barelypunctate. Intervals slightly convex; interval 3 of left elytron with single setigerous puncture at apical third, sameone of left elytron without puncture. Umbilicate series of each elytron consist of 10 setigerous punctures (leftelytron 5+0+5, right elytron 5+0+5), in the middle largely interrupted. Prosternum, mesosternum, metasternum andabdominal sterna smooth, glabrous, impunctate; anterior margin of metaepisterna rather longer than internal mar-gin; abdominal sterna II–VI with two setigerous punctures, sternum VII with four marginal setigerous punctures.

Legs moderately long, femora stout; protibia with two clip setae; mesotibial ctenidium well differentiated; hindcoxa with medial setigerous puncture below meeting point of coxae; both median and hind trochanter with setiger-ous punctures; tarsomere 5 without ventral setae.

Female genitalia. Tergum VIII with convex, slighlty angled at top, distal margin and very short proximal“legs” (Fig. 142). Sternum VIII consists of two oblong chitinised parts connected with membrane, without “legs”on proximal margin (Fig. 149). Ovipositor very small, with relativelly bigger valvifer and smaller stylus, as longeraxis of former more or less perpendicular to longer axis of latter (Figs. 160); valvifer having two-humped internalmargin, smoother external margin, and well-developed process across; stylus consisting of one basal degeneratestylomere (apical stylomere disappeared). Spermatheca with seminal canal and receptaculum discrete, seminalcanal longer, receptaculum shorter than seminal canal, strongly curved apically (Fig. 165).

Male genitalia. Unknown.Etymology. The specific epithet derives from the Latin adjective relegatus, meaning “exile” or “ejected” (in

English), reflecting its isolation from the ranges its known to be inhabited by its congeners.Diagnosis. This species is distinct from its congeners by: (i) the reduction of setigerous punctures in interval 3

of elytra, i.e presence of single puncture on left elytron, and none on rigth elytron; (ii) umbilicate series on eachelytron consisting of 10 setigerous punctures; (iii) peculiarities in the structure of the female genital armature (seeabove paragraph “Female genitalia”).

The new species possess receptaculum strongly curved apically that differs quite clearly from the receptaculumin T. molopinus, which is only slightly curved apically. In the structure of the female genital components, i.e. ter-gum VIII, sternum VIII, stylus of ovipositor, and spermatheca (Figs. 140–142, 147–149, 158–160, 164–165), T.relegatus sp. nov. looks like T. molopinus and T. chaudoiri sp. nov. Most probably, three aforementioned species ofMolopsis form a monophyletic complex.

It worth noting that the species inhabits an area situated very closely to the range of T. wiedemanni, but beyondthe ranges of T. molopinus and T. chaudoiri sp. nov. On the other hand, T. relegatus sp. nov. and T. wiedemanniseem to be not closely related phyletically. There are clear distinctions between the last two species, as well asbetween T. wiedemanni and the pair T. molopinus - T. chaudoiri sp. nov., in the structure of the female genitalia(see Figs. 140–143, 147–150, 158–162, 164–166). From a zoogeographic perspective, we may postulate the fol-lowing two hypotheses. In the past the lineage “T. molopinus - T. chaudoiri sp. nov. - T. relegatus sp. nov.” had awider range in NW Anatolia and occurred east of the Sakarya River. After that, probably as a result of climaticchanges, the spreading of this group was reduced as the central and most of the eastern parts of former range wereoccupied by T. wiedemanni. Today, two taxa of the complex live in the western part of the former range, and a thirdtaxon survived in a restricted area in the east. Or, alternatively, a single migration of the same lineage, started fromthe area of west of the Sakarya River, reached the region of Dorukhan Pass. The migration took place along thenorth parts of the Bolu Range. These assumptions should be tested by phylogenetic and molecular analyses.

Distribution. The type specimen was found in the Dorukhan geçidi (pass), between Mengen and Devrek. Thepoint lies ca 20 km N of Mengen (NW Turkey, villayet Zonguldak).



PLATE 12. Terga VIII (dorsal view) (Figs. 138–143) of Tapinopterus species. Fig. 138. Tapinopterus (Pseudomolopsis) rebel-lis (Greece, Taygetos, Anogia); Fig. 139. T. (Molopsis) molopiformis (Tmolos-Gbg., Lydien, West-Kleinasien, Weirather); Fig.140. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 141. T. (Molopsis) chaudoiri sp. nov. (paratype, Gök-dağ); Fig. 142. T.(Molopsis) relegatus sp. nov. (holotype); Fig. 143. T. (Molopsis) wiedemanni (Turkey, Bolu, Kartalkaya). Scale bar: 0.5 mm.

Tapinopterus (Molopsis) wiedemanni (Chaudoir, 1850)(Figs. 21–23, 40–44, 75–79, 91–94, 106–109, 121–123, 135–137, 143, 150, 153, 161–162, 166)

Type locality. Nord de l´Anatolie.Taxonomic decision. Tapinopterus (Molopsis) wiedemanni (Chaudoir, 1850), combinatio novum of Feronia

(Molops) wiedemanni Chaudoir, 1850: 145 (nec Fairmaire, 1866: 252).References

Feronia (Molops) wiedemanni Chaudoir: Chaudoir, 1850: 145; Marseul, 1882: 51.Feronia wiedemanni Chaudoir: Kraatz, 1875: 419.Feronia (Steropus) wiedemanni Chaudoir: Marseul, 1880: 313.Molops wiedemanni Chaudoir: Ganglbauer, 1889: 52; Bousquet, 2003: 475.Platysma (Molops) wiedemanni Chaudoir: Jakobson, 1907: 355.

Material examined. 103 specimens.Type material. Two syntypes (♂ ♀) of Feronia wiedemanni (lectotype male and paralectotype female by pres-

ent designation), well preserved, “Wiedemanni Chaud. Anatolie C. Falderm.” (h label pinned on the bottom ofbox)/ LECTOTYPE and PARALECTOTYPE Feronia (Molops) wiedemanni Chaudoir D.Dubault désign. 2003(red p)/ Tapinopterus (Molopsis) wiedemanni G.Dubault détermin. (p) (Fig. 21) (MNHN, “Collection Chaudoir”,box no 216 “Argutor”).

Other material examined. Abant: 29 ♂♂ ♀♀, “Aband 1.300 m. Turkey J. y S. Klapperich.” (MNHN); 1 ♂,TR.bor.occ. ABANT lake, 5.6.1992, V.Skoupý lgt. (cVS); 1 ♂ 1 ♀, TURKEY occ. Abant Gölü env. (35 km SW ofBOLU), 1993-06-21, Kuboň lgt. (cGD); 2 ♂♂, Türkie bor., Abant, 9.–10.5.1994, A.Trmal lgt. (cRL); 1 ♂ 3 ♀♀,TURKEY (Bolu), Elmacik dagi, Golu lake, 10.V.1994, Major (cDW); 3 ♂♂ 1 ♀, N TURKEY, Abant env., 5.1995,F.Moravec lgt. (cRL) (Fig. 22); 1 ♂, TR-prov. Bolu, Abant gölü, 1200 m, 4.6.1996, Z.Malinka lgt. (cVS); 1 ♂ 1 ♀,TURKEY, Abant Golu, 30 km W Bolu, 4.5.1997, leg J.Bašta (cDW); 1 ♂, TURKEY, Abant Golu, W Bolu,8.6.1998, lgt. E.& P. Hajdaj (cDW); 1 ♀, TR vill. Bolu, Abant Gölü, 23.–24.5.2000, Josef Mertlik lgt. (cVS).

Aladaglari: 15 ♂♂ 8 ♀♀, Anatolia bor., Aladağ ca 20 km südl. Bolu, 1400–1600 m, 27.IV.1981, Heinz leg.(MCSNM, cWH, cBL, cRL).

Köroğlu dağlari: 1 ♀, Anatolia bor, Tasliyayla, ca 21 km ne Seben (Bolu), 1500–1600 m, 27.IV.1984, W.Heinz lgt. (cWH); 13 ♂♂ 9 ♀♀, Kartalkaya, 1800 Bolu Turquie, 19.05.94, G. Dubault (cGD, cRL, cBL).

Sündiren dağlari: 7 ♂♂ 1 ♀, Anatolia centr., Wald n. Mihaliççik, 1500–1600m, Heinz leg. (cBL, cGD) (Fig.23).

Diagnosis. BL 11.3–15.8 mm.Male genitalia. Median lobe of aedeagus in dorsal view long, with long, thin, and slightly curved to the left



apical lamella, which formed more or less distinct transverse disc in lateral position (Figs. 40–44, 75–79); apicalorifice deflected to left lateral position; basal bulb shorter than apical, with basal orifice concave, apical part long,copulatory sclerite curved, with widened medial part, situated in paramedial position, apex of lobe bent down-wards, but reflexed before end. Right paramere relatively short, strongly curved and slightly constricted medially,with thinner apical and thicker basal part (Figs. 91–94, 106–109). Left paramere with long transverse apophysisand long, oblique paramedial apophysis (Figs. 121–123, 135–137).

Female genitalia. Tergum VIII relatively long, with convex, subpyramidal distal margin, straight margin andshort “legs” in proximal position (Fig. 143). Sternum VIII unified, in proximal position with two closed “holes”and without “legs” (Fig. 150). Syntergum IX+X long, narrow, bearing pair of small, articulated, distant each otherovipositors (Fig. 153). Ovipositor with valvifer and stylus (Figs. 161–162); valvifer with twin-peaked internal mar-gin, convex external margin, and process across at distal part; stylus consist of one, basal, subelongate stylomere(apical stylomere disappeared), having strongly incised internal margin. Spermatheca with seminal canal andreceptaculum discrete; receptaculum shorter than seminal canal, slightly curved apically (Fig. 166); spermathecalgland elongate, spermathecal canal short, inserted at junction of receptaculum and seminal canal.

Taxonomic notes. This species was described from three males and one female as Feronia (Molops) wiede-manni from the north of Anatolia (Chaudoir, 1850: 145–146). Later, Gemminger & Harold (1868: 333) placed itunder Molops, but Chaudoir (1876: 348) specified that it did not belongs to that genus. Kraatz (1875: 419) is first toindicate that F. molopina and F. wiedemanni were close relatives. Marseul (1882) put F. wiedemanni Chaudoiramong the Molops species. For the lack of basolateral setigerous punctures and presence of single basal impressionof pronotum, Ganglbauer (1889: 52) supposed that this taxon belonged to Tapinopterus. It seems that since thework of Jakobson (1907) this taxon disappeared from the literature. It appears again only recently, when F. wiede-manni Chaudoir, 1850 was considered either as a representative of Molops (Mlynář, 1977) or was placed amongthe species “incertae sedis” of the same genus (Bousquet, 2003: 475).

In fact, no real explanation about the systematic position of this species has been taken after the above note ofChaudoir (1876). After prolonged search for the Chaudoir´s types in MNHN by one of us (GD), two syntypes ofFeronia wiedemanni were eventually found. Their study revealed that both belong to a valid species of Tap-inopterus, and not to Molops.

Distribution. Mountain ranges around Bolu: Abant, surrounding of the Abant Lake, Aladağlari, Köroğludağlari, Sündiren dağlari, NW Turkey.

Tapinopterus (Pseudomolopsis) rebellis (Reiche & Saulcy, 1855) s.l.(Figs. 1–2, 25, 45, 55, 80, 95, 110, 124, 138, 144, 154)

Material examined. 24 specimens.Type material. Holotype ♂ of Pterostichus rebellis v . Kumanensis, Morea Cumani Brenske (p)/ rebellis, v.

Kumanensis m.(h)/coll. Reitter (p)/Holotypus (p) 1884 Pterostichus rebellis v. Kumanensis Reitter (h) (red label)/(Fig. 2) (HNHM).

Other material examined. Peloponnesos: 3 ♂♂ 1 ♀, GREECE (Peloponissos) (Arkadia) Taygetos Geb., Lan-gadapaβ 1000 m, bei Abzweigung Kokkinilakka (Gesiebe an Platanen in schmalem Bachtal), 28.IV.1999 IngoWolf (Fig. 1) (cDW); 3 ♂♂ 2 ♀, Mt. TAYGETE 1300, Péloponnése, Gréce V.94, G.Dubault, B.Lassalle (cGD,cBL); 1 ♂, Mt. TAYGETE 1300, Péloponnése, Gréce 17.05.93, G.Dubault (cGD); 1 ♂, “Rte KALAMATA-Spartekm 28 1000, Péloponnése, Gréce, 20.10.99, G.Dubault” (cGD); 1 ♀, GRECE Péloponnése, Taygéte Forét, Au des-sus d´Anogia, 7.V.2001, G. Ledoux (cGD); 1 ♂, TAYGETE, Péloponnése Gréce, X.00 G.Dubault (cGD); 4 ♂♂ 1♀, KOSMAS, Péloponnése Gréce, X.00 G.Dubault, B.Lassalle (cGD, cBL); 1 ♀, Kalamata GR, Oxihori, Taygétes,1300m, 26/IV/2000, Péloponnése, Gréce, B.Lassalle (cBL); 1 ♂, Morea Cumani Brenske (HNHM); 2 ♂♂, Grece(ZISP); 2 ♂♂, N37°47´EO21°44´, Griechenland Pelop Umg. Koumani 630 m Lompe 1.4.1997 (cDW).

Taxonomic notes. In the course of this work, we came to the conclusion that Tapinopterus rebellis s.l. is notclosely related to the known species of Molopsis, which form a distinct monophyletic complex. Diagnostic featuresthat distinguish these groups of Tapinopterus are: 1/ general shape of the pronotum; 2/ length of the antennae; 3/puncturation of the abdominal sterna; 4/ setation of the hind coxa; 5/ structure of the aedeagus; 6/ structure of theovipositor. Our results were confirmed by the subsequent study of Italian authors and species of Tapinopterusrebellis s.l. will be placed to the new genus Pseudomolopsis (Giachino et al, in press).



Distribution. Greece: Peloponnesos.

PLATE 13. Sterna VIII (ventral view) (Figs. 144–150) of Tapinopterus species. Fig. 144. Tapinopterus (Pseudomolopsis)rebellis (Greece, Taygetos, Anogia); Fig. 145. T. (Molopsis) molopiformis (Turkey, Boz dağ); Fig. 146. T. (Molopsis) phrygius(Turkey, Davraz dağ); Fig. 147. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 148. T. (Molopsis) chaudoiri sp. nov. (paratypeGök dağ); Fig. 149. T. (Molopsis) relegatus sp. nov. (holotype); Fig. 150. T. (Molopsis) wiedemanni (Turkey, Bolu, Kartal-kaya). Scale bar: 0.5 mm.

Ovipositor morphology, seasonal activity and possible parental care in Molopsis

As it was demonstrated above, the regression of the ovipositor occurs in all the species of this subgenus (Figs. 151–153, 155–162). As well, the reduction of sternum VIII (Figs. 145–150) occur in the most species having reducedovipositors except T. wiedemanni. Only T. molopiformis, the species which in our view stays closest to the hypo-thetical ancestor of the group, possesses both apical stylomere with seta/e (Figs. 155–156) and sternum VIII withwell-developed “legs” (Fig. 145). Surprisingly, the sternum VIII in T. wiedemanni is not so reduced as expected,compared with those in the other species with degenerate ovipositors (excl. T. molopiformis), but actually its struc-ture seems rather modified for pterostichines. The regression of the ovipositor consists of reduction of both the sty-lus and valvifer. This process mostly affected the apical stylus, which either present having setae highly reduced (asin T. molopiformis), or it is totally absent (as in all other species). The valvifer has strongly reduced posterior lobesin all species (Figs. 151–153, 155–162). The strong reduction in the styli-valvifera complex in Pterostichini impliesan inactive ovipositor (see also Brandmayr & Brandmayr, 1978).

The data about the phenology of the species (Table 1) suppose a spring-early summer activity peak, a late sum-mer-early autumn break, and a questionable late autumn activity. An imaginal aestivation has been noticed for spe-cies of Molops, as a result of change in the reproductive strategy (Brandmayr & Zetto Brandmayr, 1978). Besides,a correlation between the ovipositor reduction and presence of parental behaviour was documented in Pterostichini(Brandmayr, 1978; Brandmayr & Zetto Brandmayr, 1978; Brandmayr & Zetto Brandmayr, 1979). In accordancewith Thiele (1977), the brood care Pterostichini are predominantly montane species and the change in the reproduc-tive strategy represents adaptation to cooler climates.



TABLE 1. Data for seasonal activity among the species of Molopsis, based on the studied material.

PLATE 14. Syntergum IX+X with articulated ovipositor and bursa copulatorix (Figs. 151–153), ventral view of Tapinopterusspecies. Fig. 151. T. (Molopsis) molopiformis (Turkey, Boz dağ); Fig. 152. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 153.T. (Molopsis) wiedemanni (Turkey, Aladağlari). Scale bar: 0.5 mm.

PLATE 15. Ovipositor (a,left; b, right) (Figs. 154–162) of Tapinopterus species. Fig. 154. T. (Pseudomolopsis) rebellis(Greece, Taygetos, Anogia); Fig. 155. T. (Molopsis) molopiformis (Tmolos-Gbg., Lydien, West-Kleinasien, Weirather); Fig.156. T. (Molopsis) molopiformis (Turkey, Boz dağ); Fig. 157. T. (Molopsis) phrygius (Turkeya, Davraz dağ); Fig. 158. T. (Mol-opsis) molopinus (Turkey, Uludağ); Fig. 159. T. (Molopsis) chaudoiri sp. nov. (paratype, Gök dağ); Fig. 160. T. (Molopsis) rel-egatus sp. nov. (holotype); Fig. 161. T. (Molopsis) wiedemanni (Turkey, Bolu, Kartalkaya); Fig. 162. T. (Molopsis) wiedemanni(Turkey, Aladağlari). Scale bar: 0.2 mm.

Species Specimens examined (seasonal data) April May June July October

T. molopiformis 29 (9) 9

T. phrygius 72 (52) 2 29 18 3

T. machardi 4 (4) 4

T. oyukluensis sp.nov. 1 (1) 1

T. molopinus 97 (71) 13 27 15 15 1

T. chaudoiri sp.nov. 65 (64) 47 17

T. relegatus sp.nov. 1 (1) 1

T. wiedemanni 103 (64) 28 32 4



PLATE 16. Spermathecal complex (Figs. 163–166) and spermathecal gland (Fig. 167) of Tapinopterus species. Fig. 163. T.(Molopsis) molopiformis (Turkey, Boz dağ); Fig. 164. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 165. T. (Molopsis) releg-atus sp. nov. (holotype); Fig. 166. T. (Molopsis) wiedemanni (Turkey, Bolu, Kartalkaya); Fig. 167: T. (Molopsis) molopiformis(Turkey, Boz dağ). Scale bars: 0.5 mm. Abbreviations: ov—oviduct; rc-receptaculum (spermatheca); sg—spermathecal gland;smc—seminal canal (spermatheca); spc—spermathecal canal; v—vagina.

Based on both the morphology of the female genitalia and the data for phenology, we assume a highly evolvedbrood care after oviposition (“Brutpflege” type) in Molopsis just like that existing in the genera Abax Bonelli, 1810and Molops. Brandmayr & Zetto Brandmayr (1978) noted that the regression of the ovipositor in Molops is “pri-mary and less recent”. Apparently, a similar decline took place independently in the genus Tapinopterus, moreespecially in the subgenus Molopsis. In fact, it is the first time when a such regression is documented for a Palae-arctic lineage of Pterostichini, out of the “Pterostichini Molopina” (sensu Casale & Ribera, 2008). This view con-tradicts the opinion of Brandmayr & Zetto Brandmayr (1978: 62) that the postnatal brood care “is an evolutionarytarget reached only by recent Jeannel’s Molopina”. In fact this state has arisen independently in Pterostichini (seealso Horne, 1990).

Conclusions

Presently, the genus Tapinopterus Schaum, 1858 consists of seven subgenera and 53 species (Bousquet, 2003;Guéorguiev & Lobo 2006; Guéorguiev, 2007). Guéorguiev & Lohaj (2008: 128–129) concluded that probably thegenus together with Speluncarius Reitter, 1886 s.l. represents a lineage. At present, the complex “Tapinopterus -



Speluncarius” unites 12 subgenera with at least 78 described species, as these figures are now provisional andexcludes undescribed taxa (Giachino et al., in press; present authors, unpublished data).

There are several evident derived characters in Molopsis in regard to their supposed ancestor and in terms of themost other lineages of Tapinopterus. Adults of all species have elytra without scutellar setigerous puncture and withrudimentary scutellar stria, last tarsomere on ventral side without seta (excl. T. machardi) and strong reduction ofsternum VIII and ovipositor. The first three characters occurs in several other species of the genus and most probablythey represent homoplasies (independently evolved derived character-states). Only the reduction of the ovipositor inMolopsis seems to be sure synapomorphy that may prove the group represents a monophyletic unit. The last state isshared by all species of the subgenus, and presently is unknown in other lineages of Tapinopterus s.l.

PLATE 17. Apex of female elytra (dorsal view) (Figs. 168–169); female abdominal sternum VII (ventral view) (Figs. 170–171) of Tapinopterus species. Fig. 168. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 169. T. (Molopsis) chaudoiri sp. nov.(paratype, Gök-dağ); Fig. 170. T. (Molopsis) molopinus (Turkey, Uludağ); Fig. 171. T. (Molopsis) chaudoiri sp. nov. (paratype,Gök dağ). Scale bars: 1 mm (Figs. 158–159); 0.5 mm (Figs. 160–161).

Key to the identification of species of Molopsis and Pseudomolopsis (in press)

1 Species smaller in size, 7.5–12 mm. Penultimate antennomeres exceed base of pronotum. Elytral striae deep, intervals convex.Hind coxa without medial setigerous puncture below meeting point of coxae. Abdominal segments coarsely punctate. Rightparamere with fine basal part (Figs. 80, 95). Tergum VIII with distal margin almost rectilinear and “legs” as long as half of ter-gum’s length (Fig. 138). Ovipositor well-developed: longer axis of valvifer more or less parallel to stylomeres (Fig. 154); api-cal stylomere well developed with two dorsolateral and one dorsal ensiform setae on tergal position, and with sensorial foveabearing two minute nematiform setae on sternal position; basal stylomere one time and half longer than apical one. Balkan spe-cies (South Greece: Peloponnesos) . . . . . . . . . . . . . . . . . T. (Pseudomolopsis) Giachino, Picciau, Vailati & Casale (in press)

- Species larger in size, 10.8–16.2 mm. Penultimate antennomeres not reach base of pronotum. Elytral striae superficial, inter-vals flat. Hind coxa with medial setigerous punctures below meeting point of coxae. Abdominal segments finely punctate.Right paramere with broadened basal part (Figs. 81–94, 96–109). Tergum VIII with distal margin convex and “legs” shorterthan half of tergum’s length (Figs. 139–143). Ovipositor highly reduced as left and right parts set widely apart each other (Figs.151–153); longer axis of valvifer more or less perpendicular to stylomeres (Fig. 155–162); apical stylomere reduced (if presentit is minute with only single hardly visible ensiform seta, but without nematiform setae, Figs. 155–156) or completely absent(Figs. 157–162); if present apical stylomere many times shorter than basal one. Anatolian species. . . . . . . . . . . . . . . . . . . . . . 2

2 Elytra without or with single setigerous punctures in interval 3 as well as with umbilicate series of 10 pores [formula: 5 + 5].Distal margin of valvifer of ovipositor forming two distant protuberances divided by hollow (Fig. 160). 12.2 mm. NW Anato-lia, Zonguldag . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. (Molopsis) relegatus sp.nov.

- Elytra with two to three (rarely with one) setigerous punctures in interval 3 as well as with umbilicate series of 13–17 pores[formula: 6 (7) + 7–8 (9–10)]. Distal margin of valvifer of ovipositor forming mostly single protuberance (Figs. 155–159, 161–



162) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 Onychium ventrally with 1–2 pairs of setae. Sternum VII in female with 2 setigerous punctures. Relatively larger and more

robust species, 14.5–16.2 mm. S Anatolia, Taurus, Karaovabeli pass . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) machardi - Onychium without setae. Sternum VII in female usually with 4 setigerous punctures (exceptionally with two - in few speci-

mens of T. phrygius from Bolkar daglari, or with 5–6 setigerous punctures—in few specimens of T. chaudoiri sp.nov.). Rela-tively smaller species, 10.8–15.9 mm. W, NW Anatolia, Taurus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4 Species with median lobe of aedeagus having apex in dorsal view short and subtriangular (Figs. 26–27). Ovipositor with dis-tinct apical stylomere bearing single very small ensiform seta (Figs. 151, 155–156). 11.2–13.8 mm W Anatolia, Boz dag.... . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) molopiformis

- Species with median lobe of aedeagus having apex in dorsal view more or less long and thin (Figs. 28–44). Ovipositor withoutapical stylomere (Figs. 152–153, 157–162). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5 Species with median lobe of aedeagus forming an acute angle (in medial part) and with distinct lateral process in left lateralview (Figs. 29, 47, 57). Right paramere with apical part not pointed towards tip (Figs. 82, 97). 12.1 mm Turkey ? . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) aenigmaticus sp.nov.

- Species with median lobe of aedeagus forming obtuse or right angle (in medial part) and without lateral process in left lateralview (Figs. 48–54, 58–64, 65–79). Right paramere with apical part more or less pointed towards tip (Figs. 83–94, 98–109). . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6 Species with apical lamella of median lobe of aedeagus wide, abruptly turned to left (Fig. 36) and left paramere subelongate,with reduced transverse apophysis (Figs. 116, 130). 13 mm. Nothern Taurus, Oyuklu daği . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) oyukluensis sp.nov.

- Species with thinner apical lamella of median lobe of aedeagus, which is either more or less regularly curved to left (Figs. 30–34, 40–44) or straight (Figs. 37–39) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7 Species with apical lamella of median lobe of aedeagus more or less regularly curved to left (Figs. 30–34, 40–44). . . . . . . . 8- Species with apical lamella of median lobe of aedeagus straight (Figs. 37–39). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Species with apical lamella of median lobe of aedeagus short, not exceeding 1/3 of entire length of lobe (Figs. 30–34). 11.8–

15.9 mm. Taurus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) phrygius- Species with apical lamella of median lobe of aedeagus long, exceeding 1/3 of entire length of lobe (Figs. 40–44). 11.3–15.8

mm. NW Turkey, surroundings of Bolu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) wiedemanni9 Elytrae of both sexes with deep, symmetrical incision at apex (Fig. 169); sternum VII in female with deep circular incision at

tip (Fig. 171). 11.3–15 mm. NW Anatolia, Uludağ, Gök Daglari. . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) chaudoiri sp.nov.- Elytrae of both sexes without incision at apex (Fig. 168); sternum VII in females without incision at tip (Fig. 170). 10.8–15

mm. NW Anatolia, Uludağ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (Molopsis) molopinus

Checklist of the species of Tapinopterus subgenus Molopsis:

Tapinopterus (Molopsis) molopiformis (Lutshnik, 1922)W Turkey, Boz dağlari

Tapinopterus (Molopsis) aenigmaticus sp. nov.Turkey (?)

Tapinopterus (Molopsis) phrygius G. Müller, 1932= Tapinopterus (Molopsis) phrygius pisidicus Müller, 1932= Tapinopterus (Molopsis) dipojranus Straneo, 1986= Tapinopterus (Molopsis) dipojranus cilicius Straneo, 1986

SW Turkey, Taurus Mts., mountain ranges about lakes Beyşehir, Eğirdir and Burdur: Ak dağ, Sultan dağlari,Kumalar dağlari, (“Phrygic Taurus”), Barla daği, Davraz dağ, Dipojraz dağlari (“Pisidic Taurus”), S Turkey, TaurusMts., Bolkar dağlari (Bulghar Dagh by the ancient authors, “Cilician Taurus”)

Tapinopterus (Molopsis) machardi (Jeanne, 2005)S Turkey, Taurus Mts., Karaovabeli geçidi (pass) (“Lycian Taurus”)

Tapinopterus (Molopsis) oyukluensis sp. nov.Central Turkey, Konya, Taurus Mts., Oyuklu dağlari, Fasihan geçidi (pass)



Tapinopterus (Molopsis) molopinus (Chaudoir, 1868)= Pterostichus olympicus Kraatz, 1875= Tapinopterus (Molopsis) dipojranus brussanus Straneo, 1986NW Turkey, Bursa, Uludağ montain range, Istanbul ?

Tapinopterus (Molopsis) chaudoiri sp. nov.NW Turkey, Uludağ montain range, pass between Domaniç and Tahtaköprü, surrounding of Oylat and Gök

Dağlari Mts. near Sapanca

Tapinopterus (Molopsis) relegatus sp. nov.NW Turkey, Dorukhan geçidi (pass), between Mengen and Devrek

Tapinopterus (Molopsis) wiedemanni (Chaudoir, 1850)NW Turkey, mountain ranges about Bolu: Abant, surrounding of Abant lake, Aladağlari, Köroğlu dağlari, Sün-

diren dağlari

Acknowledgements

We thank our colleagues and friends for enabling us to study material from institutional and or private collections,namely Maxwell Barclay (BMNH), Ben Brugge (ZMAN), Achille Casale (Sassari, Italy), Andrea Colla (MCSNT),Thierry Deuve (MNHN), Dominique Echaroux (Etrechy, France), Arnaud Faille (MNHN), Conrad Gillet(BMNH), Evžen Hajdaj (Ježov, Czech Republic), Jiří Hájek (NMP), Walter Heinz (Schwanfeld, Germany), BerndtJaeger (MNHUB), Boris Kataev (ZISP), Maurizio Pavesi (MCSNM), Fabrizio Rigato (MCSNM), István Rozner(Budapest, Hungary), Heinrich Schönmann (NMW), Vladimír Skoupý (Žilina, Czech Republic), Győző Szél(HNHM), Manfred Uhlig (MNHUB), David Wrase (Berlin, Germany) and Lothar Zerche (DEI). Thanks are alsodue to Milan Štrba (Bratislava, Slovakia), Gejza Dunay (Kráľovce, Slovakia) and Katarína Jurková (Košice, Slova-kia) for habitus photos. Special thanks are dedicated to Jon Cooter (Oxford, UK) for the language revision of thetext. Careful reviews of a previous draft of the final manuscript were made by professor Achille Casale (Sassari,Italy) and two anonymous referees.

Part of the material in this study was examined during the work of BG in the course of the EU-funded Inte-grated Infrastructure Initiative "Synthesys" under FP6 (applications AT-TAF-758; DE-TAF-725; HU-TAF-817;FR-TAF-2938; NL-TAF-4177; GB-TAF-4263).

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