A new coleopterous family Wabbelidae fam. nov. …follows: Body form oblong, head prognathous, metacoxae not meeting edges of elytral epipleura, metepimeron meeting anterior angle

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Baltic J. Coleopterol. 17(1) 2017

ISSN 1407 - 8619

INTRODUCTION

Baltic amber is comparatively young (ascompared with inclusions from Burmese,Lebanese, Spanish, and other Cretaceous amberdeposits) and includes almost modern insectfauna at the family level. The most dramaticchanges in insect fauna took place beforeCainozoic (Zherikhin 1978), the last majorextinction of insects was in the earlyCretaceous, and the rate of extinct lineages offamily rank from Eocene is low.

New insect families from Baltic amber wereestablished in different orders. At least fifteeninsect families are known: Protoxenidae inStrepsiptera (Pohl et al. 2005), Weitschatidaein Hemiptera (Koteja 2008), Ogmomyidae in

A new coleopterous family Wabbelidae fam. nov. (Coleoptera:

Cucujoidea) from Baltic amber (Cenozoic, Paleogene, Eocene)

Vitalii I. Alekseev

Alekseev V.I. A new coleopterous family Wabbelidae fam. nov. (Coleoptera: Cucujoidea)

from Baltic amber (Cenozoic, Paleogene, Eocene). Baltic J. Coleopterol. 17 (1): 29 - 41.

An unknown peculiar beetle from Tertiary Baltic amber, Wabbel cerebricavus gen. et sp.nov. designated to a new family, is described and illustrated. The new family, Wabbelidaefam. nov., is diagnosed and assigned to the superfamily Cucujoidea. The new family can beidentified by the thick moniliform antennae, modified (deeply concave) frons and vertex,longitudinally strigose pronotum with pair of basal grooves, wide gula with complete gularsutures, pentamerous non-lobed tarsi, seriate punctuation of elytra, and dorsally non-flattened body.

Key words: taxonomy, fossil resin, new family, Polyphaga, Cucujiformia

Vitalii I. Alekseev. Department of Zootechny, FGBOU VO “Kaliningrad State Technical

University”, Sovetsky av. 1. 236000 Kaliningrad, Russia. E-mail:

[email protected]

Trichoptera (Wichard 2013), Elektraphididae inHomoptera, Catastylidae and Palpigeridae inCollembola (according to Zherikhin 1970),Babidae in Ephemeroptera (Kluge et al. 2006),Yantarocentridae in Trichoptera (Ivanov &Melnitsky 2016), Hoffeinsmyiidae in Diptera(Michelsen 2009), and Paleomelittidae (Engel2001), Pelecinopteridae (Zherikhin 1970), andElectrotomidae (Rasnitsyn 1977) inHymenoptera. Three new coleopteran familieswere described based on the material fromBaltic amber: (1) Berendtimiridae, a family withthe monotypic genus Berendtimirus Winkler,1987; (2) Electrapatidae, at present consideredto be of the tribe Electrapatini Iablokoff-Khnzorian, 1962, belonging to the familySchizopodidae LeConte, 1859; (3) Circaeidae

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Alekseev V. I.

Iablokoff-Khnzorian, 1961, currently withinAderidae Csiki, 1909.

In the present paper the description of a newenigmatic taxon of beetles discovered in Balticamber is provided. The inclusion cannot beassigned to any extant family and therefore isplaced in a new genus and new family based ona combination of peculiar externalmorphological characteristics.

MATERIAL AND METHODS

The examined material for the study consistsof a single specimen (holotype) originatingfrom the private collection of Christel and HansWerner Hoffeins (Hamburg, Germany)[CCHH]. The holotype will be deposited in thecollection of Senckenberg DeutschesEntomologisches Institut in Müncheberg(Germany) [SDEI]. The amber piece wasprepared manually and embedded in a block ofpolyester resin (Hoffeins 2001). Photographswere taken with a Zeiss AxioCamICc 3 digitalcamera mounted on a Zeiss Stemi 2000stereomicroscope. Measurements were takenwith the ocular micrometer of thestereomicroscope. Reconstructions were basedon free-hand drawings made during theexamination of the original specimen. Figureswere edited using Adobe Photoshop CS8software.

SYSTEMATIC PALAEONTOLOGY

Genus Wabbel gen.nov.

Type species. Wabbel cerebricavus sp. nov.

Composition. The new genus is monotypic,represented by the type species only.

Diagnosis. Body of medium size (4.2 mm);elongate, moderately convex dorsally. Headprognathous; with small round compound eyes;without subocular antennal grooves and

postocular constriction; genae projectingforward. Frons and vertex exceptionally deepconcave. Antennae 11-segmented, moniliform,without distinct antennal club. Antennalinsertions visible from above. Pronotum finelybordered basally and laterally, without sublateralcarinae, longitudinally strigose medially, withtwo grooves basally. Prosternal process widerthan shortest diameter of a procoxal cavity.Procoxal cavities closed, without lateral slits.Mesocoxal cavities open, metacoxal cavitiesclosed. Legs moderately developed.Trochantins not visible. Procoxae stronglytransverse. Metacoxae not meeting edges ofelytral epipleura. Tibiae narrow, simple.Trochanters completely separating femora andcoxae. Tarsi pentamerous, with non-lobedsegments. Tarsomeres 1-4 equal in size,terminal tarsomere being the longest.Metaventrite without transverse suture, shorterthan prothorax and mesoventrite combined.Elytra not truncate; finely punctate-striate; withwidely flattened lateral margins and apices.Epipleuron long, without cavities. Scutellumlarge, transverse. Abdomen with five freeventrites. Femoral lines absent.

Derivatio nominis. The generic name of newtaxon is derived from the word “Wabbel” (or“Wabel”), i.e. “beetle” in the low Prussian dialectof German language.

Note: Unfortunately, morphology of genitalia,wings venation, and wing folding are notaccessible in the studied fossil specimen andcannot be seen.

Wabbel cerebricavus sp. nov.

(Figs. 1-6)

Material examined. Holotype No. 1615-4[CCHH], imago, sex unknown. Inclusion in asmall yellow amber piece, embedded in a blockof GTS polyester resin measuring 10 x 9 x 6mm. Syninclusions are absent.

Type strata: Baltic Amber. Upper to mid-Eocene.

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A new coleopterous family Wabbelidae fam. nov. (Coleoptera: Cucujoidea) from Baltic amber...

Fig. 1. Wabbel cerebricavus gen. et sp. nov. Habitus: dorsal view.

Fig. 2. Wabbel cerebricavus gen. et sp. nov. Habitus: ventro-lateral view.

Type locality: Baltic Sea coast, Yantarnysettlement [formerly Palmnicken], Sambian(Samland) peninsula, Kaliningrad region,Russia.

Diagnosis. As stated for the new genus (videsupra).

Description. Habitus. Total body length 4.2mm, maximal body width 1.6 mm. Elongate-

ovate, moderately convex dorsally andsubflattened ventrally, entirely black.

Head. Subquadrate; prognathous, not retractedinto pronotum, finely punctate, not constrictedbehind eyes. Frons and vertex exceptionallydeep concave, forming a hole in the basal medianthird of head. Clypeus dull, longitudinallytriangular with angle directed basally, sharplyseparated, slightly convex. Gular sutures well

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Alekseev V. I.

separated, entire; gula wide, smooth. Genaeprojecting forwards, knobbed in front of eyes.Temples 2.5 x longer than diameter of eye,bearing a brush of long, dense, brown hair.Compound eyes small, round, entire, withoutinterfacetal setae, not prominent, coarselyfacetted (in each eye about 50 facets). Maxillarypalps 4-segmented; first and third palpomeresshort and transverse; second palpomerelongitudinal and 3 x longer than palpomere 1 or3; terminal palpomere cylindrical, apicallyrounded, being the longest. Ratio of palpomeres1:3:1:4. Antennal insertions exposed in dorsalview; distantly in front of eyes; separated fromeach other by a distance narrower than thedistance between eyes, but wider than the firstantennomere length. Antenna with 11 freelyarticulating antennomeres, moniliform, nearlyuniform in width, thick, not flattened, extendingto elytral basis. Scape conical, pedicel andantennomeres 3-9 slightly transverse,antennomere 10 subquadrate, antennomere 11slightly longitudinal, truncate. All antennomereswith irregularly long (sometimes longer thanlength of segment), thick, black, sparce,semierect hairs.

Thorax. Pronotum transverse (in dorsal viewabout 0.69 times as long as its median length),slightly narrower than elytral base, finelybordered laterally and basally. Anterior marginstraight, posterior margin gradually arcuate;lateral margins weakly rounded. Posteriorangles rectangular, anterior angles slightlyprotruding forwards, rounded. Pronotal surfacecovered with irregular, dense and coarsepunctures medially; with rasped punctureslaterally. Pronotal pubescence long, accumbent,brown; directed forwards in anterior margin.Pronotum longitudinally strigose in apical andcentral part (nearly 20 fine keels); with twosymmetrical, deep, longitudinal, tear-shapedgrooves basally. Procoxa separated. Prosternalprocess slightly extended beyond procoxalcavities, elongate, gradually dilated apically,truncate, wider than the shortest procoxaldiameter. Notosternal suture complete.Hypomeron with dense, large, and rounded

punctures. Prosternum in front of coxal cavitieswith fine lines directed forward and accumbentlong hairs. The ratio of lengths of prothorax tomesoventrite to metaventrite to abdomen is4.5:1.0:4.5:11.5. Metaventrite withouttransverse suture, distinctly shorter than theprothorax and mesoventrite combined.Metepimeron meeting anterior angle of ventrite1. Scutellar shield strongly transverse,pentagonal with rounded angles. Elytra evenlyconvex; elongate oval in dorsal outline;concealing abdominal apex; separately roundedapically; with widely flattened lateral marginsand apices; finely punctate; shiny. Elytralpunctures small, bearing short hairs, arrangedin striae, shallower on disc, becoming confusedapically. Each elytron with 18 rows of finepunctures (formed by strial and interstrialpunctuation equal in size), 4 rows in basal thirdwith distinctly enlarged 8-10 punctures bearingerect hairs, 4 x as long as interval between strialpunctures. All elytral intervals flat. Epipleurawell developed, wide, reaching apex, withoutcavities. Metathoracic wings fully developed,partially exposed in the holotype specimen.

Abdomen. With five visible, similarlyarticulated, finely and densely punctured, finelypubescent ventrites. Relative lengths (medially)of ventrites 1� 5 equal to 3.5-1.7-1.7-1.5-3.0.Intercoxal process of abdominal ventrite 1widely triangular and acute. The first ventrite isapproximately as long as the second whenmeasured without process. Femoral lines absent.

Legs. Moderately developed. Trochantins notvisible. Pro- and metacoxal cavities closed,mesocoxal cavities open. All coxae separated:procoxae separated by a distance slightly widerthan the minimal procoxal diameter by theprosternal process; mesocoxae narrowlyseparated; meso-metaventral junction withoutmetaventral knob; metacoxae separated by adistance approximately 2 × the coxal diameterwith wide triangular process of ventrite 1.Procoxae transverse, 2 x as long as wide;mesocoxae nearly round. Metacoxae transverse,do not extend laterally to meet the elytra.

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Fig. 3. Wabbel cerebricavus gen. et sp. nov. Head and pronotum.

Fig. 4. Wabbel cerebricavus gen. et sp. nov.: A - Abdomen, thorax and head ventrally; B - Meso-,metathorax and abdomen (schematically).

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Alekseev V. I.

Trochanters of all legs large, completelyseparating femora and coxae. Femorasubflattened, simple. Tibiae slightly shorter thanfemora; with longitudinal keels; pubescent;shortly setose apically, without crenulation,denticles, or spines. Tarsal formula 5-5-5. Notarsomeres lobed. Tarsi shorter than tibiae,pubescent below. Length of apical tarsomeresubequal to combined length of tarsomeres 2-4; tarsomeres 1-4 cylindrical, almost equal insize. Tarsal claws equal in size, thin, simple,length about one third of apical tarsomere.

Derivatio nominis. The specific epithet refersto the very peculiar concave form of frons andvertex. It is formed after the Latin “cerebrum”(brain, brainpan) and “cavus, -a, -um” (empty,hollow, concave).

The new genus exhibits a unique combinationof characteristics warranting its recognition asa new family. Further investigations by X-raymicro-computed tomography (micro-CT) couldhelp to study internal features in this specimenand may add data on the systematic placementof the taxon. If additional conspecific orcongeneric specimens will be found in Balticamber material in the future, this can be usefulfor understanding the position of the taxon inthe system of Coleoptera, too. At the moment,it is proposed to establish the new familyWabbelidae fam. nov. based upon the monotypicgenus Wabbel gen. nov. and to place this familyin the system as follows:

Order Coleoptera Linnaeus, 1758Suborder Polyphaga Emery, 1886Series Cucujiformia Lameere, 1938Superfamily Cucujoidea Latreille, 1802Family Wabbelidae fam. nov.

Type genus. Wabbel gen. nov.

Composition. The new family is proposed forone generic group only, consisting of the onenew species Wabbel cerebricavus sp. nov. fromBaltic amber.

Description. The new family includes one genusand one species as described above in detail.The list of the most important (Leschen et al.2005) characteristics can be summarized asfollows: Body form oblong, head prognathous,metacoxae not meeting edges of elytralepipleura, metepimeron meeting anterior angleof ventrite 1, metaventrite shorter thanprothorax and mesoventrite combined, tarsi notlobed, tarsal formula 5-5-5, metaventritewithout transverse suture, basal ventrite free,antennal insertions not concealed, protibiaesimple (without crenulation, denticles, orspines), procoxal cavities closed, mesocoxalcavities open, metacoxal cavities closed,procoxae strongly transverse, head withoutsubocular antennal grooves, elytra not truncateand concealing abdominal apex, ventrite 1without femoral lines, pronotum withoutsublateral carinae, elytra without ridges,antennae 11-segmented with antennal clubindistinct, epipleuron without deep cavity,procoxal cavities without narrow lateral slits,prosternal process wider than the shortestdiameter of a procoxal cavity, head withoutpostocular constriction, genae projectingforward, elytral punctuation seriate, mesocoxalcavities narrowly separated.

Diagnosis. First ventrite undivided bymetacoxae and prothorax without notopleuralsutures. These findings suggest placement ofWabbelidae fam. nov. in the suborder ofPolyphaga, excluding a relationship of the newtaxon to Rhysodidae (fairly similarly thickmoniliform antennae, 5-segmented tarsi, smalleyes, dorsal surface of head with grooves andlobes). Not strongly projecting procoxaeexclude a possible membership ofStaphylinoidea with similarly formed tarsi andantennae. The appendages’ morphology(antennae, legs) and some other habitualcharacteristics (e.g. elytral punctuation) doresemble those of the families JacobsoniidaeHeller, 1926 and Ptinidae Latreille, 1802 fromthe series Bostrichiformia.

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Figure 5. Wabbel cerebricavus gen. et sp. nov. Habitus dorsally, reconstruction.

Jacobsoniidae can be recognized by a minute,narrowly elongate body, an elongate prothorax,lack of a visible scutellum, and a markedlyelongate metaventrite (at least 2.5 x as long asmesoventrite), combined with a relatively shortabdomen and extremely slender metanepisterna,which are potential synapomorphies of thefamily (Löbl & Burckhardt 1988; Lawrence&Leschen 2010; Bi et al. 2015; Cai et al. 2016).Additionally, all known modern Jacobsoniidaehave antennal insertions concealed from above

and a pronotal disc without paired basalimpressions (Lawrence & Leschen 2010). Thecharacteristics of Wabbel gen. nov. representingWabbelidae fam. nov., i.e. comparatively shortmetaventrite, distinct transverse scutellum,metepisterna not slender, abdomen notshortened, paired pronotal impressions, anddorsally exposed antennal insertions, excludethe beetle from the systematical vicinity ofapparently similar Jacobsoniidae.

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In addition, Wabbel gen. nov. is similar to somemembers of the subfamily Ptininae by thewidely separated metacoxae without coxalplates, by the open mesocoxae, by the hiddentrochantins, by the long femora, by the non-clubbed antennae, by the legs with pentameroustarsi and the terminal tarsomere being thelongest, by the separation of procoxae byprosternal process (present in numerous generaof spider-beetles), and by the triangular clypealform. The reasons to exclude the Ptinidae frompossible groups to which the new taxon can beassigned are the following: Wabbel gen. nov. hasa large head with long temples and gula, whereasPtininae have a head moderately deflexeddownwards and constricted in pronotum(Mynhardt & Philips 2013); in Ptininae theantennal insertions are closely together betweeneyes (Belles 1978; Philips 2000), whereas theantennal insertions in Wabbel gen. nov. aremoderately wide, separated from each other andfrom eyes; Wabbel gen. nov. has prothoraciclateral margins, whereas in ptinids suchstructure is absent (Mynhardt & Philips 2013);in Ptininae the first 3 ventrites are connate whileventrite 4 is often reduced (Philips 2000),whereas the new taxon has basal ventrites withcomplete sutures and nearly equal in length; incontrast to the newly described taxon, ptinidshave open procoxal cavities (Philips 2000).To the author’s opinion, the attempt to assignthe new genus to Jacobsoniidae or Ptininae(Ptinidae) would destroy the modern conceptsof these taxa. The new genus has a highlyspecialized head, whwereas specialization inptinids (and, possibly, in the whole bostrichoidlineage) is connected with different pronotalmodifications. The trend towards integration ofthe head and prothorax in a single unit inPtininae and the apparent absence of suchintegration in Wabbel gen. nov. are an importantobstacle to evolutionary relation. The antennalform in Ptininae and Jacobsoniidae distinctlyvaries between the genera (moniliform inSarothrias, distinctly clubbed in Derolathrus

and Saphophagus) and species of one genus(e.g., Polyplocotes Westwood 1869 hasantennae with compressed, widened, or reducedsegments in different species). A similar

antennal form may be observed in severalunrelated coleopteran myrmecophiles while thenumber and form of antennal segments may varywithin one genus (Lawrence & Reichardt 1969;Bell & Philips 2009). Thus, such characters arenot of paramount importance. The moreimportant dissimilarities between Wabbelidaefam. nov., Ptinidae, and Jacobsoniidae are listedabove.

The not concealed head without a medianocellus, not excavated metacoxae withoutvertical posterior face, absent metacoxal plates,not approximated antennal insertion, nottruncate elytra, tarsal formula 5-5-5, simpleclaws, well-developed intercoxal process offirst abdominal ventrite, and long antennae allowfor tentatively placing Wabbel gen. nov. amongCucujoidea.

This superfamily includes 25 recent families andremains difficult to characterize owing to thephenotypic heterogeneity exhibited in thisgroup. There are no characteristics that uniteall the families of Cucujoidea as the superfamilyis characterized by the following combinationof features only (Robertson et al. 2015): Adultswith procoxal cavities internally open (mostly),tarsal formula 5-5-5 in female and 5-5-5 or 5-5-4 in male animals (rarely 4-4-4). However, ithas not been possible to definitely assign thetaxon to any known family of the superfamily.

The new family Wabbelidae fam. nov. iscompared with all families of the superfamilyin consecutive order as the new taxon obviouslydiffers from:Passandridae by the open mesocoxal cavitiesand by the not parallel-sided body form;Helotidae by the metaventrite withouttransverse complete suture and by the transverseprocoxae;Nitidulidae, Kateretidae, and Monotomidae bythe absence of an antennal club and by notexposing the pygidium;Laemophloeidae by the pronotum withoutsublateral carinae and by elytra withoutlongitudinal ridges;

Alekseev V. I.

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Protocucujidae by the simple non-lobedtarsomeres;Smicripidae by the 5-segmented tarsi;Silvanidae by the absence of lobed tarsomeresand by the transverse fore coxae;Sphindidae by the absence of an antennal cluband by a tubercle or cavity on the mandible;Cyclaxyridae by the elongate, not stronglyconvex body and by the absence of a deep cavityon the epipleuron;Hobartiidae by the non-clubbed antennae;Tasmosalpingidae, Lamingtoniidae, andPhloeostichidae by the closed procoxal cavities;Cucujidae by the markedly convex dorsalsurface;

Cavognathidae by the absence of narrow lateralslits on the procoxal cavities, the frons withoutdeep pits, and the closed procoxal cavities;Myraboliidae by the absence of slits on theprocoxal cavities, the body form not flattenedand parallel-sided, and the non-3-segmentedclub;Agapythidae by the prosternal process widerthan the shortest diameter of a procoxal cavityand by the head without a postocularconstriction;Priasilphidae by the narrowly separatedmesocoxal cavities, the abdomen with freeventrites, and the seriate punctuation of theelytra;

Fig. 6. Wabbel cerebricavus gen.et sp.nov. Forebodyventro-laterally: (1) compound eye [small]; (2) gula[wide]; (3) gular sutures [complete]; (4) notosternal suture[complete]; (5) pronotal border [present]; (6) procoxae[transverse, i.e. longer than wide]; (7) prosternal process[wide]; (8) hypomeron; (9) protrochanter; (10) elytralepipleuron

Cryptophagidae by the openmesocoxal cavities, the transverseprocoxae, the moniliform antennae,and the seriate elytral punctuation;Phalacridae by the elongate, notstrongly convex body, the moniliformantennae, and the simple (non-toothed) claws;Erotylidae by the non-clubbedantennae;Boganiidae by the mandibles withoutsetose cavities on the dorsal side andby the all-simple tarsomeres (non-lobed ventrally);Cybocephalidae by the not roundedbody, the non-lobed tarsomeres, andthe 5-segmented tarsi.

According to Bouchard et al. (2011),the superfamily Cucujoidea Latreille,1802 includes two families that arealso extinct: ParandrexidaeKirejtshuk, 1994 and SinisilvanidaeHong, 2002. The position ofSinisilvanidae within Silvanidae isconsidered unclear (according to itsoriginal description probably withinthe subfamily Silvaninae sensustricto) by Kirejtshuk (2011). TheMesozoic family Parandrexidae (asdiagnosed by Soriano et al. 2006)differs from newly describedWabbelidae fam. nov. by thesubfiliform antennae (moniliform in


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Wabbelidae), the long mandibles, the longpalpomeres, the narrowly separated and flatcoxae, and the 4-segmented tarsi.

Wabbelidae fam. nov. can readily bedistinguished from other Cucujoidea by thefollowing unique combination ofcharacteristics: (1) thick moniliform antennaewithout distinct 3-segmented club, (2) modified(deeply concave) frons and vertex, (3)longitudinally strigose pronotum with pair ofbasal grooves, (4) wide gula with complete andwell-separated gular sutures, (5) pentamerousnon-lobed tarsi with first tarsomere equal to thesecond, (6) seriate punctuation of elytra, and(7) dorsally non-flattened body.

Taken separately, these characteristics are notunique and occur in several taxa of modernbeetles. However, the complex of thecharacteristics listed above is unique for thesuperfamily. It allows assuming that Wabbel

gen. nov. is a representative of a formerlyunknown taxon, whose rank is equal to otherfamilies of Cucujoidea.

In the most recent key to cucujoid families(Leschen et al. 2005), Wabbel cerebricavus

(and correspondingly Wabbelidae fam. nov.)runs to couplet 31, Priasilphidae/Agapythidae,having occiput without stridulatory files.Insertion of a couplet “31a” at this point wouldseparate Wabbelidae:

31a. Antennae moniliform, frons and vertexdeeply concave…...………………...................……Wabbelidae-Antennae clubbed, head without deep groovesor concavity.........……………..…………… 31

DISCUSSION

The new taxon may represent a highly modifiedform with special adaptation associated withinquilinous habits. Such myrmecophiles arealways a problem to taxonomists. Many beetlefamilies originally proposed to comprise such

inquilines no longer qualify (Lawrence &Reichardt 1969). Antennal, leg, and body shapemodifications as well as hairs are known fordifferent groups of myrmecophilous beetles(Parker 2016). So, surprisingly, the pronotalform (lobes and grooves) of the recent SouthernAfrican myrmecophilous ptinine genusDiplocotidus Péringuey, rediscribed by Bell &Philips (2008), is fairly similar to the uniqueform of the head of our new taxon. Externallysimilar and, possibly, functionally equivalentforms of the forebody could arise onlyconvergently in these beetles. Elongation of theclypeus, protection of mouthparts, and reductionof body hairs or scales, so that the surfaceappears shiny, probably represent adaptationsassociated with myrmecophilous habits(Lawrence & Reichardt 1966; Philips &Mynhardt 2011) as similar trends are found inPaussidae, clavigerine Pselaphidae, severalrepresentatives of Dacoderinae (Salpingidae),different and non-related groups of Ptininae(Diplocotes, Diplocotidus, Gnostus, Fabrasia

etc.), and other inquilines. Some othersuperficially similar beetle groups (Rhysodidae,Jacobsoniidae) are mentioned in the diagnosisof the family. The way of life of the fossil taxonremains unknown as the factors explaining suchstriking head morphology are left to speculation.

The accurate dating of Baltic Tertiary amber isa debatable question: Either Lower Oligocene[Rupelian, 33.9 - 28.1 Ma], Upper Eocene[Priabonian, 38.0 - 33.9 Ma], or Lower Eocene[Ypresian-Lutetian, 56.0 - 41.3 Ma] is estimatedas the time of amber forest existence bydifferent authors (Ritzkowski 1997; Jaùoszyñski& Perkovsky 2016; Vitali & Daamgard 2016).The support of distinct branches of scienceshould be used for final conclusions. So, Vitali& Damgaard (2016) supposed Early Oligoceneas the age of Baltic amber origin, because ofthe predominance of temperate taxa ofcerambycid beetles in the Baltic amber fauna.According to the opinion of the authors, theEocene of Northern Europe was warmer thanthe Oligocene, and biota of temperate Asianareas could only immigrate after the drying of

Alekseev V. I.

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the Turgai strait, which had taken place betweenEocene and Oligocene. However, geologicaldata contradict this hypothesis. According toBaraboshkin et al. (2007), continental Asia hada northern land connection with Fennosarmatiain Bartonian and Priabonian. Therefore, thefaunal migrations may have taken place muchearlier than in Oligocene. Moreover, the widthof the narrow Turgai gate of Turgai strait spanabout 250 km in Ypresian (Baraboshkin et al.2007). Such distance over water was surely notan insurmountable obstacle for taxa possiblytransported by floating wood during millions ofyears. A similar direct migration route acrossnarrow sea straits from Fennosarmatia toNorthern America via Britain and Greenlandduring Eocene (and earlier) can be easilyassumed, too. The absence of climatic and widegeographical barriers between all land parts ofthe Northern Hemisphere in Eocene should haveresulted in the mixed and more or lesshomogeneous insect fauna of the whole area.The effectiveness of isolation of southerncontinents (considering north/south directed seacurrents and atmospheric circulations) forinsect migrations could be questioned, too. Withregard to this question, the paper of Graham(2006) is of interest since it deals with possiblecauses for the similarity of modern-daybiodiversity between the Caribbean and Africa.Arguments stating that the age of Baltic amberas a fossilized Oligocene resin based on faunalpeculiarities only and neglected the geologicaldata concerning the sedimentation of amber andpalaeo-geographic data from the Eocene, canthus be rejected.

ACKNOWLEDGEMENTS

The author gratefully acknowledges Dr. Pavel I.Alekseev (Saint Petersburg, Russia) for takingthe photographs and Mrs. Christel Hoffeins andMr. Hans Werner Hoffeins (Hamburg,Germany), who enabled the study of the amberwith the unique inclusion under their care. Theauthor is greatly thankful to Dr. Adam Úlipiñski(CSIRO, Canberra, Australia) and Mrs. Christel

Hoffeins for consultations during the work andcomments on an earlier version of this paper.The study was supported by the RussianFoundation for Basic Research, project No. 14-04-00262.

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Received: 07.11.2016

Accepted: 30.12.2016

Published: 02.08.2017


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A new coleopterous family Wabbelidae fam. nov. …follows: Body form oblong, head prognathous, metacoxae not meeting edges of elytral epipleura, metepimeron meeting anterior angle