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The cerambycids included in Baltic amber: current knowledge status with the description of new taxa (Coleoptera, Cerambycidae) PDF

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© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at The cerambycids included in Baltic amber: current knowledge status with the description of new taxa (Coleoptera, Cerambycidae) Francesco VITALI Abstract: A synopsis of all cerambycid species recorded from Baltic amber until today is provided and analysed according to the current systematic and paleontological knowledge. Only eight species result to be valid. A further new species, Encyclopidonia punctatissiman.gen.n.sp. (Cerambycidae, Lepturinae, Rhagiini), is described. The new genus Trichosieversiafor Pseudosieversia eu- ropaeaVITALI, 2004 is instituted. The analysis of the cerambycid Baltic fauna strongly implies the presence of temperate environ- mental conditions and suggests therefore to date the Baltic amber at least to the Early Oligocene. Key words: Coleoptera Cerambycidae, fossil, Baltic amber. Santrauka: Pateikiama visu˛ u–suocˇiu˛ (Coleoptera, Cerambycidae) ru–šiu˛, kurios iki šiol aprašytos pagal Baltijos gintarsa˛, apžvalga. Šie vabzdžiai analizuojami, remiantis dabartin(cid:0)mis sistematikos ir paleontologijos žiniomis. Tik aštuonios ru–šys laikomos validžio- mis. Aprašoma dar viena nauja ru–šis Encyclopidonia punctatissima n.gen.n.sp. (Cerambycidae, Lepturinae, Rhagiini). Ru–šis Pseudo- sieversia europaea VITALI, 2004 perkeliama i˛ naujai aprašoma˛ genti˛ Trichosieversia n.gen. Baltijos gintaro u–suocˇiu˛ fauna akivaizdžiai rodo, kad gintarmedžiu˛ miškas augo vidutinio klimato sa˛lygomis; jis patvirtina, jog šis gintaras turi bu–ti datuojamas bent jau anks- tyvuoju oligocenu. Raktiniai žodžiai: Coleoptera, Cerambycidae, fosilijos, Baltijos gintaras. Introduction of them just focused on Baltic amber, but a lot of work is still in preparation or remains to be done. The goal of The Cerambycidae LATREILLE, 1802 are a family of this paper is to present the actual knowledge status of Coleoptera Cerambycoidea widespread throughout all the cerambycid fauna included in Baltic amber, to the world with nearly 30,000 species. The elegance of which a further new species is presently added. forms and the attractive aspect of most of them have made cerambycids one of the most popular family among collectors, while the long antennae characteris- Material and Methods ing most species apparently render them easy to identi- Unfortunately, most of the cerambycids described fy by the non-specialist, too. In particular, this charac- from Baltic amber were conserved in European Muse- teristic has allowed to record cerambycids in amber ums that heavily suffered the dramatic consequences of since the onset of this study. Unfortunately, the fact WWII. The research done in these Museums has al- that long antennae characterise also different families lowed to discover that the holotypes of the species once and the evolution of the taxonomy and the palaeontol- preserved in Danzig (2), Leipzig (1) and Paris (1) are ogy occurred during the centuries has caused a big in- today lost (dispersed or burnt), while only the types certitude regarding the real consistence of the fossil ce- (mostly holotypes) preserved in Berlin (4) are still ex- rambycid fauna. isting (ANDRÉ, BECHLY, MÜLLER, NEUMANN, REICH, Actually, except for the paper written by Richard SZADZIEWSKI, WEITSCHAT, in litt.). Other two species ZANG(1905), all other papers regarding this topic suf- have been described on materials preserved in Ham- fer from approximate taxonomic, biological and burg (1) and London (1) after WWII, but it is actually palaeontologic knowledge about this family. Hence, for about synonyms of a well-known species whose types Denisia 26, some years the author of this paper has begun analysing are still existing. Finally, three holotypes, besides other zugleich Kataloge der oberösterreichischen fossil and sub-fossil cerambycids through several papers specimens coming from Baltic amber, are present in the Landesmuseen (VITALI2004a, b,2005, 2006a-e, 2007a-e, 2008), some author’s collection. Neue Serie 86(2009): 231–242 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Fig. 1: Habitus of probable North American origin of this genus. This Trichosieversia record is probably referable to some elongated Rhagiini, europaea(VITALI, maybe Pseudosieversia europaea. 2004); the black line represents 1 mm. Genus PachytaDEJEAN, 1821 KLEBS1910, p. 328; STATZ1938, p. 173; LINSLEY1961, p 54; ABDULLAH1967, p. 147; LARSSON1978, p. 156; SPAHR1981, p. 23; POINAR1992, p. 138. This genus was cited by KLEBS(1910) from his own material, which was identified by Edmund REITTER. This presence seems very doubtful due to the large size of their representatives, and thus probably referable to some hitherto undescribed Rhagiini closely related to the genera Evodinus LECONTE, 1850 (MULSANT’s “Pachyta“) or AcmaeopsLECONTE, 1850. Trichosieversia n.gen. The availability of further better conserved material has allowed to reconsider the taxonomic and systematic position of Pseudosieversia europaeaVITALI,2004, which is here attributed to a new genus. Genotype: Pseudosieversia europaea VITALI, 2004 (monobasic). Description: Small (4.7-5.9 mm), elongated, con- vex above (Fig. 1). Head convex; forehead largely grooved; antennal tubercles widely separated, elevated; cheeks short, shorter than under eye-lobes; temples prominent; neck The following results are primarily based on the bib- distinct. Eyes relatively large, close to the basis of the liography and some checked types. Other research about mandibles, emarginate at upper side, uniformly convex the remaining types conserved in Berlin will be part of at the under one, finely faceted. Last palpomere blade- further publications. Regarding the consulted bibliogra- shaped, as long as wide at the apex. phy, several papers and general catalogues (HANDLIRSCH Antennae inserted between the eyes, reaching the 1907; STATZ 1938; LINSLEY 1961; ABDULLAH 1967; elytral apex in female; scape bowed; pedicle elongated, SPAHR 1981; HIEKE & PIETRZENIUK 1984; CARPENTER one-half longer as broad; antennomere III scarcely 1992; POINAR 1992) quote a lot of cerambycid genera longer than scape; antennomere IV scarcely shorter (sometimes even mentioned as species) from Baltic am- than scape; antennomere V five-third as long as scape; ber. Nevertheless, it is about unchecked quotations of antennomere VI-IX progressively shortened; anten- ancient approximate and often inexact records, already nomere VI scarcely shorter than previous, antennomere proved as erroneous in the past. Actually, the real VII one-fourth longer than scape; antennomere VIII amount of longhorn species included in Baltic amber is scarcely longer than scape, antennomere IX scarcely still exiguous. shorter than scape, antennomere X three-fourth as long as scape; antennomere XI two-third as long as scape (proportions according to the formula: 1.3: 0.4: 1.4: 1.2: Observation on the recorded taxa 2.1: 2.0: 1.6: 1.4: 1.2: 1.0: 0.9). Subfamily Lepturinae LATREILLE, 1802 Prothorax little convex above; sides obtuse toothed Genus StenocorusFABRICIUS, 1775 at about two-third of their length from the base, HOPE1836, p. 142; SPAHR1981,p. 24; POINAR1992, p.138. grooved by two transversal furrows, one narrower, ante- This ancient, no longer confirmed, presence was cit- rior and one wider, posterior; front and hind margins of ed by HOPE(1836) on material of DALMAN’s collection, the pronotum elevated, hind margin broadly enlarged at today partially preserved in Berlin. Actually, it seems to the outer angles; surface glabrous, extremely finely and be very doubtful due to the large size and especially the densely punctured, except for a glabrous longitudinal 232 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at 2 3 4 Figs 2-5: (2) Trichosieversia europaea(VITALI, 2004), specimen FS14B14 author’s coll., particular of the head; (3): Encyclopidonia punctatissiman.gen.n.sp., holotype, particular of the head; (4), dorsal side; (5), inclusion. Legs long, femora slightly club-shaped, tibiae linear, armed with an apical tooth at the inner apex and anoth- er shorter at the outer one; surface very finely, thick punctured, tibiae carrying also some erect setae; tarsi long, metatarsus one-half as long as metatibia; metatar- somere I very long, three-fifth as long as tibia; tarsomere II one-third as long as I; tarsomere III one-sixth as long as I, bilobed, deeply incised, onychium one-third as long as tarsomere I. Feminine genital armature with stylus apical, drop- shaped, rounded at the apex. Differential diagnosis: Trichosieversia n.gen. essen- tially differs from PseudosieversiaPIC,1902 and the relat- ed genera, with which it was originally confronted 5 (Sivana-Macropidonia-Pidonia), in the short cheeks (Fig. 2). Such archaic character makes this species more closely related to Encyclops NEWMAN, 1838,which dif- carina located on the basal half of the disc. Scutellum fers in the more elongated habitus and the advanced po- transverse, rounded posteriorly, unpunctuated. sition of the antennal insertion. Moreover, this charac- ter makes Trichosieversia a possible link both between Elytra 2.6 as long as wide at the humeri, clearly Encyclopsand the group Pseudosieversia-Sivana-Macropi- wider than prothorax, feebly enlarged posteriorly, api- donia Pidonia, as well as between Encyclops and Cor- cally suddenly convergent and separately rounded; su- todera. Actually, the tribal assignment of some genera of ture very finely grooved; surface covered with some iso- Lepturinae (Pidonia, Cortoderaand Grammoptera) is still late long semi-recumbent black setae and with a strong, uncertain, being attributed to either tribe on the basis of almost thick, regular punctuation becoming more con- different adult or larval characters. Equally arguable is fused and almost rugose to the apex. the value of the tribe Encyclopini LE CONTE, 1873, Prosternum in lateral view making a distinct angle which is either considered as a valid taxon or as syn- with the procoxae; procoxal cavities posteriorly open; onym of different tribes. However,the long elytral setae intercoxal of prosternum extremely narrow; metaster- of Trichosieversia, absent from all treated genera, are in num very finely, thick punctured and very finely pubes- all likelihood an autapomorphy that justifies the institu- cent; pygidium convergent-sides, truncate at apex, two tion of a new genus. This character also suggests that P. times longer than other visible sternites, exceeding the europaeais not the direct ancestor of the quoted genera elytral apex. but a collateral dry branch. 233 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Trichosieversia europaea(VITALI, 2004) n.comb. tectable at the apex of the mesotibiae. Tarsi long; (Figs 1, 2) metatasomere I as long as the following two together, Pseudosieversia europaeaVITALI, 2004a, p. 1-8, figs 1-4 (Holoty- tarsomere II slightly longer than III, onychium, as long pe: coll. VITALI); VITALI2005, p. 530-531, fig. 1; VITALI2007b, as I. p. 14, fig. 1. Differential diagnosis: The prosternal shape makes Encyclopidonia n.gen. Encyclopidoniaa member of the tribe Rhagiini, while the Genotype: Encyclopidonia punctatissima n.sp. general habitus (long cheeks, antennae inserted between (monobasic). eyes, prothorax mutic and rounded at base) suggests a re- lation with the genera close to Pidonia MULSANT, 1863. Description: Small, elongated, depressed above Nonetheless, the group Pseudosieversia-Sivana-Macropi- (Figs 5-6). donia has stouter elytra and a tuberculated prothorax, Head relatively long (Fig. 3); forehead apparently while Pidonia has stouter elytra and a much less devel- squared, even; antennal tubercles widely separated, fair- oped body punctuation. On the other side, the long ely- ly elevated; cheeks well developed, scarcely but appar- tra remind of the tribe Encyclopini, which nevertheless ently visibly longer than under eye-lobes; temples short, includes genera with shorter checks, antennae inserted rounded, scarcely convergent backward, as long as the at the front margin of the eyes, and tuberculated protho- eye-lobes; neck long. Surface sculpture and mouth rax. Some Grammoptera-species (especially of the sub- pieces not visible. Eyes relatively small, widely separat- genus Neoencyclops MATSUSHITA& TAMANUKI, 1940) al- ed from the basis of the mandibles, relatively small, fee- so have a similar habitus but also shorter cheeks, a fact bly emarginate at the upper side, uniformly convex at that suggests no relationship with this fossil. Moreover, the under one, finely faceted. Encyclopidonia shows a peculiar proportion of some an- tennomeres (article IV not shorter than III), which are Antennae inserted between the eyes, reaching the absent from nearly all the previously quoted genera. This posterior fourth of the elytral length, glabrous, extreme- abnormal proportion, being only known in Macropidonia ly finely and densely punctured; scape scarcely bowed, PIC, 1901 (among the Lepturinae of the Recent), the fos- rounded at the apex; pedicle as long as broad, one- sil Paracorymbia antiquaVITALI,2005 and some Spondy- fourth as long as scape; antennomeres III-IV sub-equal, lidinae and Cerambycinae, is in all likelihood plesiomor- one-fifth longer than scape; antennomere V scarcely phic. Further peculiar characters are the close dense longer than previous; antennomere VI scarcely shorter punctuation of the pronotum (Fig. 4) and the pointed than scape; antennomere VII three-fourth as long as elytral apex (Fig. 6). In particular, the elytral apex, being scape; antennomeres VIII-IX equal, scarcely shorter unknown in both Encyclopsand Pidonia, seems to be au- than VII; antennomere X less than half as long as scape tapomorphic, suggesting that Encyclopidonia diverged (proportions according to the formula: 1.6: 0.4: 1.8: 1.8: from Encyclopswithout being an ancestor of Pidoniaor of 1.9: 1.3: 1.2: 1.0: 1.0: 0.8: >0.5). other genera of the Recent. Prothorax feebly elongated, convex above, widely Finally, the black body colour can be found among constricted at apex and at base, regularly rounded at Recent species in the melanic form of the Alleghenian sides; hind angles rounded; apex and basis finely Pidonia ruficollis (SAY, 1824), and many related genera grooved; disc without longitudinal furrow, everywhere (Encyclops, Trichosieversia, Pseudosieversia, Macropidonia, covered with a coarse dense punctuation, equal in size Idiopidonia, Cortodera, Anoplodera, Grammoptera). Prob- but more serrate than that of elytra. Scutellum small, ably, this was the original pattern of all species of this forming an equilateral triangle. Prosternum in lateral group; later, the adaptation to the life on flowers view making a distinct angle with the procoxae, procox- evolved more and more yellow patterns, analogously to al cavities posteriorly open; metepisternum 2.5 times as other species living in the same habitat. long as wide. Elytra long, 3 times as long as wide at the humeri, Encyclopidonia punctatissima n.sp. (Figs 3-6) clearly wider than prothorax, depressed above, restricted Holotype: Baltic Coast, ex coll. P. CARDWELL after humeri, then parallel-sided, apically suddenly con- A8137, author’s coll. FS39B24. The insect is partially vergent and separately acutely rounded, apex pointed; covered by turbidity and missing the apical part of the suture very finely grooved; surface covered with a coarse, last right antennomere, the last five left antennomeres, almost thick, irregular punctuation, apparently glabrous. the two last left protarsomeres, and a part of the poste- rior right leg in the knee region. Legs long, femora slightly club-shaped, tibiae linear, rectilinearly truncated at the tip, extremely finely and Y, length 9 mm, body and antennae black, legs ap- densely punctured. A very minute stout spine is de- parently reddish brown. Characters of the genus. 234 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Fig. 7: Habitus of Paracorymbia antiquaVITALI, 2005; the black line represents 1 mm. genera has been demonstrated to occur (PESARINI& SAB- BADINI2004), their taxonomic position deserves to be re- Fig. 6: Habitus of Encyclopidonia punctatissima vised. n.gen.n.sp.; the black line represents 1 mm. GenusLepturaLINNAEUS,1758 GenusGrammopteraAUDINET-SERVILLE, 1835 and HOPE 1836, p. 142; BERENDT 1845, p. 46-47, 56, 58; GIEBEL “propeGrammoptera etStrangalia“ 1856, p. 132; MENGE1856; p. 21; MOTSCHULSKY1856, p. 28; KLEBS1910, p. 237-238; STATZ1938, p. 173; LINSLEY1961, p SCUDDER1885, p. 793; SCUDDER1886, p. 73; SCUDDER1891, 54; ABDULLAH 1967, p. 147; LARSSON 1978, p. 156; SPAHR p. 546; ZANG1905, p. 243; HANDLIRSCH1907, p. 787; LARS- 1981, p. 22; POINAR1992, p. 138. SON1978, p. 156; SPAHR1981, p. 23; POINAR1992, p. 138. This presence was cited by KLEBS(1910), based on To the specimen of BERENDT’s collection mentioned his own material identified by REITTER, but may possibly by HOPE(1836), BERENDT(1845) added other two ones, be referable to Encyclopidonia punctatissima. which were mentioned by GIEBEL(1856) and finally ex- Paracorymbia antiqua VITALI,2005 (Fig. 7) amined by ZANG (1905). One of them effectively be- longed to the Lepturini and was described as Strangalia VITALI2005, p. 531-533, figs 2-5(Holotype: coll. VITALI). berendtiana, while both remaining specimens were This fossil (Fig. 7), though classifiable in the genus doubtfully identified as Heteromera. Paracorymbia MIROSHNIKOV, 1998, shows some peculiar archaic characters (stout habitus, convex under margin of Adults and one larva of MENGE’s collection were the eyes, short pubescence on the pronotum) that makes mentioned by MENGE (1856), MOTSCHULSKY (1856), it a natural link with the close genus Vadonia MULSANT, SCUDDER(1885, 1886, 1891), HANDLIRSCH(1907) and 1846. However, as interbreeding between species of these LARSSON(1978) but the material is lost. Though possi- 235 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at ble, the presence of the genus Lepturain Baltic amber is closely related to the extant Nothorhina muricata(DAL- not supported by any known specimen yet. MAN, 1817) in habitus and body size. Taxonomic revi- sion and probable biology in VITALI(2006b). GenusStrangalia AUDINET-SERVILLE, 1835 KLEBS1910, p. 238; STATZ1938, p. 173; LINSLEY1961, p 54; Palaeoasemum duffyiABDULLAH, 1967 ABDULLAH1967, p. 147; LARSSON1978, p. 156; SPAHR1981, ABDULLAH 1967, p. 149-150, figs. 4-8 (Holotype: No. 1461; p. 24; POINAR1992, p. 138. Paratypes: No. 546 and No.1556, ex coll. KLEBS, Hamburg); This record, which KLEBS(1910) cited from his own ABDULLAH1975, p.397; SPAHR1981, p. 23; VITALI2006b, p. material identified by REITTER, is very probably referable 33-36. to Strangalia berendtiana since KLEBS did not mention ZANG’spaper in his catalogue. Palaeoasemum crowsoni ABDULLAH, 1967 ABDULLAH 1967, p. 149, figs. 1-3 (Holotype: No. 18796, ex Strangalia berendtianaZANG, 1905 coll. KLEBS, No. 533, London); ABDULLAH1975, p. 397; SPAHR ZANG1905, p. 243-244, fig. 3 (Holotype, coll. BERENDT, Ber- 1981, p. 23; CARPENTER1992, p. 312; VITALI2006b, p. 33-36. Both species are evident synonyms of Nothorhina lin); HANDLIRSCH1907, p. 787; LARSSON1978, p. 156; SPAHR 1981, p. 24; HIEKE& PIETRZENIUK1984, p. 305; VITALI2004a, granulicollis(VITALI, 2006b). ABDULLAHdescribed them p. 1; VITALI2005, p. 537, fig. 7. on the basis of outdated North American keys, ignoring A credible Strangalia with closer affinity to Ameri- descriptions and types of cerambycids of Baltic amber can congeners than to Asian ones. and even extant European species. The new genus and species were claimed on the basis of typical sexual char- Genus Necydalis LINNAEUS, 1758 acters of Nothorhina(pronotal shape, body proportions) HOPE1836, p. 143; BERENDT1845, p. 56; GIEBEL1852, p. 656; and well-known variable characters among cerambycids SCUDDER1885, p. 793; HANDLIRSCH1907, p.787; SPAHR1981, (body size, scutellar shape). p.23; POINAR1992, p. 138. This specimen, which HOPE (1836) recorded from GenusTetropium KIRBY, 1837 BERENDT‘s authority, was no longer mentioned as such KLEBS1910, p. 238; STATZ1938, p. 173; LINSLEY1961, p 54; since the beginning of the past century. By considering ABDULLAH1967, p. 147; LARSSON1978, p. 155; SPAHR1981, the ancient taxonomy, it should be identified as the p. 24; POINAR1992, p. 138; VITALI2006b, p. 36. “Molorchus“ of BERENDT‘s collection that ZANG (1905) This genus, which KLEBS(1910) cited from his own finally recognised as “Cantharis“. No other records are material identified by E. REITTER, remains to be verified known; however, the genus Necydalis, today Vancouver- but may possibly be valid (VITALI2006b). ian and Oriental, was in all likelihood absent from Ter- tiary Baltic forests. Genus SpondylisFABRICIUS, 1775 SCUDDER1885, p. 794, figs. 1025, 1025a; SCUDDER1886, p. 73; Subfamily Spondylidinae SCUDDER 1891, p. 583; HANDLIRSCH 1907, p. 786; LARSSON AUDINET-SERVILLE, 1832 1978, p. 155; SPAHR1981, p. 24; POINAR1992, p. 138. It is about the record of a larva; nevertheless, besides Nothorhina granulicollis ZANG, 1905 the incertitude due to the type of finding, Spondylishas Callidiumsp. I-IV BERENDT1845, 46-47; ZANG1905, p. 236. currently an Eurasian, originally Vancouverian, distri- Nothorrhinaprope muricataHELM, 1886, p. 272; HELM1896, p. bution and was in all likelihood absent from Europe un- 229; HANDLIRSCH1907, p. 786; SPAHR1981, p. 23. Nothorrhinasp. KLEBS1910, p. 238; STATZ1938, p. 173; AB- til the Pleistocene (VITALI,in prep.).This larva, if still DULLAH1967, p. 147; HIEKE& PIETRZENIUK1984, p. 305; POI- existing, might be referred to one species belonging to NAR1992, p. 138; WEITSCHAT& WICHARD2002, p. 166, fig. the Asemini (LARSSON1978), eventually, to Nothorhina 63g. granulicollis. Nothorrhina granulicollisZANG, 1905, p. 236-240, fig. 2 (Holo- type and 4 paratypes, three of which still exist, coll. BERENDT, Spondylis crassicornis GIEBEL, 1856 Berlin); HANDLIRSCH 1907, p. 787; LINSLEY 1961, p 54; AB- GIEBEL1856, p. 127 (Holotype, Leipzig, lost); SCUDDER1885, DULLAH1967, p. 150; LARSSON1978, p. 155; SPAHR1981, p. p. 793; HANDLIRSCH 1907, p. 785; LARSSON 1978, p. 155; 23; HIEKE& PIETRZENIUK1984, p. 305. SPAHR1981, p. 24; VITALI2006b, p. 36-37. Palaeoasemum crowsoniABDULLAH, 1967, p. 149, figs. 1-3; AB- The morphological characters provided in the orig- DULLAH1975, p. 397; SPAHR1981, p. 23; CARPENTER1992, p. inal description do not surely allow to identify this fos- 312. sil as Spondylis but maybe as the oldest synonym of Palaeoasemum duffyi ABDULLAH, 1967, p. 149-150, figs. 4-8; ABDULLAH1975, p. 397; SPAHR1981, p. 23. Nothorhina granulicollis (LARSSON 1978; VITALI 2006b). Nothorhina granulicollisVITALI,2006b, p. 30-41, figs. 1-11. Nonetheless, this statement might open some practical The most common cerambycid in Baltic amber, problems, due also to the fact that the type is lost today. 236 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at This species should be deemed as incertae sedis,or bet- Genus GraciliaAUDINET-SERVILLE, 1834 ter as nomen oblitum. KLEBS1910, p. 237; STATZ1938, p. 173; ABDULLAH1967, p. 147; ABDULLAH 1967, p. 147; LARSSON 1978, p. 156; SPAHR Subfamily Cerambycinae LATREILLE, 1802 1981, p. 22; POINAR1992, p. 138. This presence, which KLEBSrecorded from his own GenusCerambyx LINNAEUS, 1758 material identified by REITTER, is possible but not veri- BERENDT1830, p. 30; HOPE1836, p. 141; MENGE1856; p. 23; fied yet. SCUDDER1885, p. 794; SCUDDER1886, p. 73; SCUDDER1891, p. 488; ZANG 1905, p. 232-233; HANDLIRSCH 1907, p. 786; Genus MolorchusFABRICIUS, 1792 SPAHR1981, p. 22; POINAR1992, p. 138. BERENDT1845, p. 56; GIEBEL1856, p. 128; SCUDDER1885, p. This specimen of BERENDT‘s collection, which HOPE 793; ZANG1905, p. 233; SPAHR1981, p. 23; POINAR1992, p. (1836) had cited together with material of STRONG’s 138. collection (British Museum), was examined by ZANG The specimen recorded by BERENDTand mentioned (1905), who recognised a tiger beetle. This fossil was by other authors was finally examined by ZANG,result- later studied by HORN (1906), who recognised the ing to be a Cantharis-species with damaged elytra. American Tetracha carolina (LINNAEUS, 1766). More re- cently, RÖSCHMANN (1999) deemed such species very Obrium propetestaceumand Obriumsp. closely related to, but not conspecific with, the Caroli- BURMEISTER1832, p. 635; GIEBEL1856, p. 129; SCUDDER1885, na tiger beetle. p. 793; HANDLIRSCH1907, p. 786; KLEBS1910, p. 238; STATZ MENGE(1856) also recorded a larva (cited by SCUD- 1938, p. 173; LINSLEY 1961, p 54; ABDULLAH 1967, p. 147; DER and HANDLIRSCH), whose identification is erro- LARSSON1978, p. 156; POINAR1992, p. 138. neous in all evidence (no Cerambyx-species lives in The former species was recorded by BURMEISTER conifers). This record is maybe referable to some larvae (1832), while the latter one was recorded by KLEBS (1910) from three specimens of his own collection. This of Cerambycinae Callidiini (VITALI, in prep.). latter record was later mentioned by all following au- thors but probably it is the same, as yet undescribed Genus CallidiumFABRICIUS, 1775 species. The presence of the genus Obrium, though pos- HOPE1836, p. 141; BERENDT1845, p. 46-47;GIEBEL1856, p. sible, has not been confirmed yet. 128; SCUDDER1885, p. 793; ZANG1905, p. 236; KLEBS1910, p. 237; STATZ1938, p. 173; LINSLEY1961, p 54; ABDULLAH1967, p. 147; LARSSON 1978, p. 155; SPAHR 1981, p. 21; HIEKE & Genus ClytusLaicharting, 1784 PIETRZENIUK1984, p. 305; POINAR1992, p. 138. HOPE1836, p. 142; MOTSCHULSKY1856, p. 28; SPAHR1981, p. The six Callidium-specimens that BERENDT (1845) 22. recorded and GIEBEL(1856) mentioned, were examined This genus was recorded from specimens of DAL- by ZANG (1905), who described five of them as MAN‘s (HOPE1836) and MENGE‘s (MOTSCHULSKY1856) Nothorhina granulicollisand recognised the last one as a collection. This material, whose destiny is uncertain, representative of Cantharidae or Lampyridae. This last might be referred to Clytus pici. specimen should be the same “Callidium“ which HIEKE & PIETRZENIUK(1984) recorded from BERENDT‘s collec- Clytus piciPITON, 1940 tion, since ZANGdidnot mention other specimens. Clytus (Xylotrechus) piciPITON, 1940, p. 63-64. Clytus piciSPAHR, 1981, p. 22. HOPE‘s (DALMAN‘s collection) and KLEBS’ records The holotype is lost today, so that only speculations might be the same species or even the same specimen. about the description are possible. KLEBS‘ specimen was identified by REITTERand is, in all likelihood, related to some still undescribed species of PITON(1940) described this specieson the basis of a Callidiini, possibly Semanotus MULSANT, 1839.LARSSON single female, 15 mm long, blackish grey, having a (1978) also recorded a larva of Callidiumfrom the Muse- cephalic carina, a silver short pubescence but no visible um of Copenhagen. Actually, these records should not pattern. Among the Clytini currently living in Eurasia be related to Callidiumsince this genus has a Vancouver- only the genus Xylotrechus CHEVROLAT,1860 is charac- ian-Manchurian distribution (LINSLEY1961), which sug- terised by a frontal carina; hence, no misidentification gests an arrival in Eurasia since the Pleistocene and its seems to be possible. Moreover, though no larva of the consequent absence in Europe during the Tertiary (VI- European species feeds on conifers (BENSE,1995), some TALI, in prep.). Other authors quoted Callidium-speci- North American species – X. annosus (SAY, 1826), X. mens from KLEBS’or undetermined collections. sagittatus(GERMAR, 1821) and X. undulatus(SAY,1824) – have this biology (CRAIGHEAD,1923). Therefore, the generic attribution seems to be correct and this species 237 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at might be transferred to Xylotrechus, according to au- (1845) added other four specimens of his own collec- thor’s intention. tion, which were mentioned by GIEBEL (1856) and fi- Nonetheless, PITON also noticed a vague resem- nally examined by ZANG(1905). Some of them are lost, blance with Xylotrechus cinereus LAP., actually Pseu- others belong to strange Lamiinae, which ZANG pre- ferred not to describe, and a last one was described as dosphegesthes cinerea(LAPORTEDECASTELNAU& GORY, “Dorcaschema“succineum. 1836), because the frontal carina looked feeble. In real- ity, such species has no frontal carina. Other specimens belonging to MENGE‘s collection The disappearance of the type does not allow to were mentioned by subsequent authors (MOTSCHULSKY solve the question; moreover, the attribution to Xy- 1856; HANDLIRSCH 1907; LARSSON 1978) but are also lotrechus will open some taxonomic problems since X. lost today. SCUDDER (1885) recorded a larva that was pici is a still existing species. While waiting for further later mentioned many times (SCUDDER 1886, 1891; investigations, it should certainly be better to maintain HANDLIRSCH 1907; LARSSON 1978; SPAHR 1981); the name Clytus pici. nonetheless, at least this record is surely taxonomically incorrect since Saperdadoes not feed on conifers (LARS- GenusAnaglyptus MULSANT, 1839 SON1978). KLEBS1910, p. 237; STATZ1938, p. 173; LINSLEY1961, p 54; Actually, the presence of this genus in Baltic amber ABDULLAH1967, p. 147; LARSSON1978, p. 156. remains to be verified. This genus was recorded by KLEBS from one speci- men of his own collection identified by E. REITTER. “Dorcaschema“succineum ZANG, 1905 Though no Recent Anaglyptus-species bores conifers Saperdasp. III BERENDT1845, p. 47. (LARSSON 1978), the presence of mixed forests during ZANG1905, p. 240-243, fig. 6 (Holotype: coll. Berendt, Ber- Baltic amber formation renders this occurrence possible. lin); HANDLIRSCH1907, p. 790; LINSLEY1961, p 53; LARSSON Nonetheless, all European species have a Euro- 1978, p. 156; SPAHR 1981, p. 22; KLAUSNITZER & SANDER Siberian or Euro-Caucasian distribution, while the 1981, p. 57, fig. 23; HIEKE& PIETRZENIUK1984, p. 305; POINAR genus is mainly Oriental. Hence, Anaglyptus should 1992, p. 138. have colonised Europe from Asia only after the draining The characters provided in the original description of the Turgai Sea at the end of Eocene. This presence is do not agree with any Recent genus of Dorcaschemati- therefore very interesting regarding the amber dating ni (BREUNING 1949); hence, this species might belong and deserves to be verified. Finally, this record might al- to a new genus, possibly close to the North American so be referable to Xylotrechus pici. Dorcaschema HALDEMANN,1847. Subfamily Lamiinae LATREILLE, 1825 Parmenops longicornis SCHAUFUSS, 1891 Aenictosoma doenitzi SCHAUFUSS,1891 SCHAUFUSS1891, p. 60-62 (Holotype: coll. HELMNo. 40, Dan- zig, lost);HELM1897, p. 89; HANDLIRSCH1907, p. 788; KORS- SCHAUFUSS1891, p. 58-60 (Holotype: coll. HELMNo. 87, Dan- CHEFSKY1939, p. 12, pl. 1, fig. 3a-b; SPAHR1981, p. 23; CAR- zig, lost); HELM1897, p. 89; HANDLIRSCH1907, p. 788; KORS- CHEFSKY1939, p. 12; SPAHR1981, p. 21; CARPENTER1992, p. PENTER1992, p. 312; POINAR1992, p. 138. The original description implied a representative of 312;POINAR1992, p. 138; VITALI2006a, p. 99-101. the tribe Parmenini, but the original drawing, which the The holotype belonged to HELM‘s collection and is lost today. The species was also mentioned several times author had not published and KORSCHEFSKY(1939) pro- by many authors without comments, but the characters vided only nearly 50 years later, suggests that this provided in the original description (pentamerous tarsi, species was actually winged. The missing of the type and elbowed antennae, pointed palpi) clearly imply that it the current puzzling systematics of Lamiinae do not al- was actually a representative of the Scydmaenidae low identifying even the tribe; nonetheless, this fossil Mastiginae Clidicini (VITALI2006a). seems somehow related to the tribe Apomecynini THOMSON, 1860. GenusSaperda FABRICIUS, 1775 HOPE1836, p. 137, 141; BERENDT1845, p. 47, 56; GIEBEL1856, Pogonocherus jaekeli (ZANG, 1905) p. 132; MENGE1856; p. 21; MOTSCHULSKY1856, p. 28; SCUD- Lamiasp. I BERENDT1845, p. 56. DER 1885, p. 793, figs 1023, 1023a; SCUDDER 1886, p. 73; Pogonochaerus jaekeliZANG,1905, p. 233-236, fig. 5 (Holotype: SCUDDER1891, p. 577; ZANG1905, p. 240-243; HANDLIRSCH coll. BERENDT, Berlin); HANDLIRSCH 1907, p. 789; LINSLEY 1907, p. 790; LARSSON1978, p. 156; SPAHR1981, p. 24; POI- 1961, p. 53; LARSSON 1978, p. 156; HIEKE & PIETRZENIUK NAR1992, p. 138. 1984, p. 305. This genus was firstly recorded by HOPE (1836) Pogonocherus jaekeliSPAHR1981, p. 24; POINAR1992, p. 138; about one specimen of BERENDT‘s collection. BERENDT VITALI2007c, p. 15-16. 238 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at According to ZANG,this fossil was so similar to the One of them was described as Pogonochaerus jaekeli, while extant Pogonocherus ovatus (GOEZE, 1777) in body size the other three belonged to two very characteristic un- and aspect that it might be considered as its direct an- known species, which ZANGpreferred not to describe. cestor. P. ovatusis actually a polyphagous species fairly rare Discussion in southern Europe but widespread in northern regions The cerambycid fauna of Baltic amber lists today to Scandinavia, interesting facts in order to understand only 9 valid species: Trichosieversia europaea (VITALI, the climate related to the Baltic amber. 2004); Encyclopidonia punctatissima n.sp.; Paracorymbia antiquaVITALI,2005; Strangalia berendtianaZANG,1905; Dorcadionoides subaeneus MOTSCHULSKY, 1857 Nothorhina granulicollis ZANG, 1905; Clytus pici PITON, MOTSCHULSKY 1857, p. 27 (Holotype: coll. MENGE, lost); 1940; Parmenops longicornis SCHAUFUSS, 1891; Dor- SCUDDER 1885, p. 793; HANDLIRSCH 1907, p. 789; LARSSON caschema succineumZANG,1905 and Pogonocherus jaeke- 1978, p. 156; SPAHR1981, p; CARPENTER1992, p. 312; POINAR 1992, p. 138; VITALI2007c, p. 15-16. li(ZANG,1905). Other species have been recognised as not belonging to Cerambycoidea (Aenictosoma doenitzi The characters provided in the original description (very minute size, long antennae) may suggest that this SCHAUFUSS, 1891), as synonyms of already described fossil was the oldest synonym of Pogonocherus jaekeli, de- ones (Palaeoasemum crowsoni ABDULLAH, 1967 and P. scribed nearly 50 years later (VITALI 2007c). Nonethe- duffyiABDULLAH,1967), or should be deemed as nomi- less, the missing of the type and the very approximate na oblita (Spondylis crassicornisGIEBEL,1856 and Dorca- description suggest to consider this species as a nomen dionoides subaeneus MOTSCHULSKY, 1857). These last species unfortunately belong to the most ancient de- oblitum. scriptions of cerambycids in amber, but their rough diag- GenusAcanthocinus GUÉRINDEMÉNEVILLE, 1826 noses and the missing of the types make them absolute- ly unrecognisable. HOPE1836, p. 142; SPAHR1981, p. 21; POINAR1992, p. 138. This very ancient record of three specimens of DAL- The first analysis of the Baltic cerambycid fauna MAN‘s collection was no longer mentioned by subse- (HOPE 1836) underlined the presence of a “South quent authors. However, the presence of Acanthocinus American relationship“ and of “considerably warm“ cli- in Baltic amber is very suspect since this genus, which mates. Nonetheless, the genera mentioned in order to has the same distribution asCallidium, was in all likeli- sustain the former (Saperda, Gyrinus) and the latter hood absent from Europe during the Tertiary. In fact, claim (Cerambyx, Clytus, Callidium, Acanthocinus, the only credible fossil Acanthocinus-species of Europe is Lamia, Leptura) reveal the complete groundlessness of that described by SCHMIDT (1967) from Late Pliocene both hypotheses. shales of Willershausen am Harz (Germany). Further analyses (ZANG1905; KLEBS1910; LARSSON 1978) rightly noticed the great abundance of Spondy- GenuspropeDorcadion DALMAN, 1817 Saperda sp. I BERENDT 1845, p. 47, 56; GIEBEL 1856, p. 132. lidinae, relating it to habitats rich in conifers. LARSSON also considered Nothorhina as a Mediterranean genus ZANG1905, p. 240; SPAHR1981, p. 22. ZANG(1905) recorded a species scarcely resembling and this mistake was recorded by subsequent authors this genus when examining a relatively large specimen (HIEKE& PIETRZENIUK1984; POINAR1992; WEITSCHAT of BERENDT‘s collection previously identified as “Saper- & WICHARD2002), contributing to sustain the idea of a da“ (BERENDT, 1845). Actually, Dorcadion has never relative warm climate. Actually, the extant Nothorhina been recorded from Baltic amber. have only a relict distribution in the Mediterranean, be- ing primarily widespread in Siberia and in the Hi- GenusLamia FABRICIUS, 1775 malayan region, where they bore mountain pines (Pinus HOPE 1836, p. 142; BERENDT 1845, p. 56; SCUDDER 1885, p. sylvestris L., P. mugo uncinata RAMOND, P. nigra laricio 793; ZANG1905, p. 233; SPAHR1981, p. 22; POINAR1992, p. POIRET, P. roxburghi SARG.). On the other side, the 138. claim that Lepturini are indicators of conifers (HIEKE& This genus was first recorded by HOPE (1836) from PIETRZENIUK 1984; WEITSCHAT & WICHARD 2002) is material of DALMAN‘s collection and later by BERENDT still not verified since the species described until today (1845) from material of his own collection. The presence are not evidently related to such plants (VITALI2005). of Lamiain Baltic amber is obviously erroneous and this In fact, though the hosts of Trichosieversiaand Encyclopi- genus should be understood as Lamiinae sensu lato in donia are unknown, all genera morphologically related both cases. Four of BERENDT‘s specimens were examined to them have larvae living on oaks and other broadleaf by ZANG (1905) and effectively proved to be Lamiinae. trees. 239 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Actually, this first screening reveals that the Baltic firm that the Baltic habitats were analogous to those we fauna included genera typically related to temperate, or find in central Europe today, as studies on the Tricho - even cold, middle-European mixed forests (Paracorym- ptera (WICHARD 1988) and Isoptera (WEIDNER 1995) bia,Nothorhina,Pogonocherus, possibly alsoTrichosiever- also pointed out. sia andEncyclopidonia), and other ones apparently relat- Since the hypothesis that Baltic amber included ed to taxa widespread in the tropics today (Dorcaschema, species of different altitudes (HEER 1865) has been Parmenops). proved erroneous (WEITSCHAT1997), the most logic ex- Nonetheless, considering this latter group as a trop- planation is that Baltic amber must be referred to ages ical element might be erroneous. Regarding Dor- much more recent than it is currently hypothesised. By caschema succineum, this fossil has uncertain relation- observing the temperature curve based on oxygen iso- ships with American species. Though Dorcaschematini tope measurement (BUCHARDT 1978), the Baltic bio- are today primarily tropical, the least specialised taxa cenosis should be dated at least to the Early Oligocene, (Dorcaschema HALDEMANN,1847 and HetoemisHALDE- as the first dating already indicated (NOETLING 1883, MANN,1847), to which this fossil seems mostly related, 1888). This younger dating of Baltic amber can also bet- live in temperate-cold habitats of North America, in- ter explain the fact that many fossils still have a great cluding Canada (BREUNING 1949). However, the pres- resemblance with extant species. ence of mostly tropical genera does not necessarily im- ply the existence of tropical habitats. In fact, the cur- Zusammenfassung rent genus Anoplophora HOPE, 1939, though primarily tropical, includes species that optimally live in habitats Ein Verzeichnis aller bis heute in Baltischem Bern- with cold winters as in Japan or in the surroundings of stein registrierten Bockkäfertaxa wird entsprechend der Milan, Italy (LINGAFELTER& HOEBEKE2002). heutigen systematischen und paläontologischen Kennt- nisse zusammengefasst und analysiert. Nur acht fossile Therefore, such apparently “tropical“ species were Bockkäferarten erweisen sich als gültig. Eine weitere probably the more northern boreal representatives of neue Art, Encyclopidonia punctatissima n.gen.n.sp. (Ce- taxa widespread in sub-tropical zones; possibly, they had rambycidae, Lepturinae, Rhagiini), wird beschrieben. their counterparts in taxa living in the temperate cli- Zudem wird eine neue Gattung, Trichosieversian.gen. für mates of the more southern zones of the Austral Hemi- Pseudosieversia europaea VITALI, 2004 eingeführt. Die sphere. VITALI(2008) identified in the Prioninae Xyleo- Analyse der baltischen Bockkäferfauna deutet sehr auf conites proavus HAUPT, 1950 from Geiseltal (Sachsen- das Vorherrschen von gemäßigten Umweltbedingungen Anhalt, Germany, Middle Eocene) the boreal counter- hin. Es wird daher vorgeschlagen, den Baltischen Bern- part of some genera of Macrotomini today widespread stein, aufgrund der zu der Zeit vorherrschenden klimati- only in Madagascar and South Africa. Although schen Bedingungen, zumindest dem Frühen Oligozän Geiseltal is located more southerly than the main Baltic zuzuordnen. amber occurrences, analogue observations have been made by other authors (WHEELER1914; HENNING1964, 1965, 1966; BARONIURBANI2000), especially regarding Acknowledgements the Baltic ant Prionomyrmex janzeni BARONI URBANI, The author thanks for the collaboration Dr. André 2000 and the dipteran Archiphora robusta (MEUNIER, NEL, Muséum National d’Histoire Naturelle, Paris 1905), Psosphyracephala succini(LOEW,1873) and Para- (France), Dr. Günter BECHLY, Staatliches Museum für corsomyza crassirostris(LOEW,1850). Naturkunde Stuttgart (Germany), Prof. Dr. Arnold Due to the well-known geographic reasons, unlike MÜLLER, Geologisch-Paläontologische Sammlung, Uni- their Austral counterparts, the Baltic boreal taxa were versität Leipzig (Germany), Dr. Christian NEUMANN, not able to survive the ice ages of the early Pleistocene Museum für Naturkunde, Humboldt-Universität, Berlin and became extinct, leaving the fauna that we can ob- (Germany), Dr. Mike REICH, Geowissenschaftliches serve today. This fact explains why such “tropical“ ele- Zentrum der Universität Göttingen (Germany), Prof. ments are difficult to ascribe to current taxa or have Dr. hab. Ryszard SZADZIEWSKI, Museum of Amber Inclu- been described as belonging to extinct genera, while the sions, University of Gdańsk (Poland), and Dr. Wolfgang temperate elements are fairly similar to extant taxa. WEITSCHAT, Geologisch-Paläontologisches Institut, Universität Hamburg (Germany).The Denisia-Team is Consequently, the impression of the Baltic ceramby- thanked for their efforts. cids is that we face a fauna richer and more diversified than today but, however temperate, similar to that cur- rently surviving in Korea or in Japan. This seems to con- 240

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