Latvijas Entomologs 2012, 51: 27-39. 27

Macratriinae (Coleoptera: ) of the Baltic amber

Dmitry Telnov

Stopiņu novads, Dārza iela 10, LV-2130, Dzidriņas, Latvia; e-mail: anthicus@ gmail.com

Telnov D. 2012. Macratriinae (Coleoptera: Anthicidae) of THE Baltic amber. – Latvijas Entomologs 51: 27-39.

Abstract: A new species of Macratriinae (Coleoptera: Anthicidae) from Baltic amber is de- scribed and illustrated – Macratria alleni sp. nov. The genus Protomacratria Abdullah, 1964 is placed as a junior synonym of Macratria Newman, 1838. A key to fossil species of Mac- ratriinae is presented for the first time. Additional information on known species of Baltic amber Macratriinae is also provided.

Key words: Macratria, Protomacratria, Macratriinae, Anthicidae, Coleoptera, Eocene, Baltic amber, .

Introduction specimens of Macratria from Baltic amber for identification which stimu- The Macratriinae is one of eight lated this critical review of hitherto subfamilies of the Anthicidae sensu described Baltic amber Macratriinae. stricto (Chandler 2010). There are 5 genera and two tribes recognized within this subfamily – Macratriini Material and methods LeConte, 1862 and †Camelomorphi- ni Kirejtshuk, Azar et Telnov, 2008 Specimens were studied using (Telnov 2011). a Leica S6D stereomicroscope. Two genera and three species Specimen photographs were taken of Macratriinae have been previously using a Canon EOS 450D SLR known from Baltic amber (Abdullah camera attached to the microscope, 1964, 1965, Kirejtshuk 2002). De- and CombineZP software was used scriptions of these three species have for image stacking. All taxa are listed never been revised since their origi- alphabetically, since a phylogenetic nal descriptions. I received additional arrangement is impossible. All label 28 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

data is reproduced exactly, with no most diverse in the subfamily, which corrections or additions; labels (if includes 295 recent and fossil species more than one for the same specimen) (Telnov 2011). No fossil representa- are separated by slashes ( / ). If not tives are hitherto known from the otherwise stated, all labels are printed. other genera of Macratriinae, namely My comments are placed in square Salimuzzamania Abdullah, 1968 brackets [ ]. (Central America) and Thambospas- ta Werner, 1974 (southern U.S.A.). The following abbreviations are used: According to recent studies (Telnov 2009), the Madagascan genus Telesi- BMNH – British Museum (Natural History), London, U.K.; nus Fairmaire, 1903 is a member of GPIH – Geologisch-Paläontologisches Eurygeniinae and not Macratriinae, Institut und Museum Universität where it was erroneously placed by Hamburg, Germany; Chandler (2010). ISZP – Institute of Systematic and evolution In this publication, one new syn- of (Instytut systematyki i ewolucji zwierząt), Polish Academy of onymy and two new combinations are sciences, Krakow, Poland; made and one species new to science LDM – Latvian Natural History Museum is described and illustrated. An origi- (Latvijas dabas muzejs), Rīga, Latvia. nal identification key to all fossil spe- cies of Macratriinae (Anthicidae) is also provided for the first time. Systematic part Protomacratria Abdullah, 1964 Among the five macratriine gen- era, two are known from amber inclu- Abdullah (1964: 332) placed his sions. †Protomacratria Abdullah, new genus into ‘Pedilinae Macratri- 1964 (Macratriini) was established ini’, now Macratriini of Macratriinae. for two Baltic amber species, but Already in the original description it †Camelomorpha Kirejtshuk, Azar was stated, that (citation from Abdul- et Telnov, 2008 (†Camelomorphini) lah 1964: 332) ‘in general form and (coauthors of this taxon are three and shape of maxillary palpi the two spec- not two as specified by Bouchard et imens described below remind me al. 2011) is a monotypic genus de- of Macratria Newman but they lack scribed from Lebanese amber. A pubescence on the tibial spurs and fourth amber fossil macratriine spe- have unusual punctures on the ver- cies is known from the cosmopolitan tex. When the revision of the world genus Macratria Newman, 1838, the species of Macratria on which I am Latvijas Entomologs 2012, 51: 27-39. 29

now engaged is complete, the posi- tion by Abdullah (1964: 333)) and of tion of the two specimens from am- P. tripunctata, I was unable to observe ber can be considered’; ‘It is doubtful any differences separating the above- if the large punctures or pits on the mentioned species from typical Mac- vertex of Protomacratria are dorsal ratria. Moreover, characters given by ocelli’ (Abdullah 1964: 334). In his Abdullah - ‘lack of pubescence on the “Natural classification of the family tibial spurs’ cannot be confirmed (all Anthicidae...” Abdullah (1969: 353) tibial spurs of both species are clearly erroneously stated that tibial spurs densely pubescent by short and strong are absent in Protomacratria, what setae), but ‘unusual punctures on the was obviously wrong. All other char- vertex’ are no more than three vague acters of Protomacratria given in the circular impressions on the head (pos- original description (reduced tem- sibly caused by damage) and because pora, distinct clypeolabral and indis- of this are only visible in one speci- tinct frontoclypeal suture, obsolete men (P. appendiculata) and ‘the two frontal ridge, 4-segmented maxillary posterior punctures are not visible [in palpi, lateral and convex eyes, width P. tripunctata] but I think they will be of the head nearly equal to that of found to be present’ (Abdullah 1964: pronotal base, filiform 11-segmented 334). In any case, this character alone antennae, antennomeres 9-10 (or 11) (presence of impressions on the head) slightly serrate, terminal antenno- could not be enough to keep this ge- mere slightly longer than penultimate nus separately from Macratria. So a one, longitudinal pronotum, non-con- new synonymy is proposed: tiguous metacoxae, short tibial spurs, tarsal claws simple or appendiculate, Macratria Newman, 1838 presence of epipleural fold (carina) = Protomacratria Abdullah, 1964 on elytra, striato-punctate dorsal sur- syn. nov. face of elytra and 5 visible ventrites) are in fact not distinctive and do Macratria alleni sp. nov. (Figs 6-7) not separate Abdullah’s taxon from Holotype ?♀ LDM: No. G 404/3 / Macratria (most of these characters HOLOTYPE [label red] / MACRATRIA are discussed by Telnov (2011) in a alleni sp. nov. det. D.Telnov, 2011. monograph of Indo-Australian Mac- ratria). inclusion in a slice After studying the type specimens of amber with some circular dim of Protomacratria appendiculata microcracks around upperside of the (type of the genus, original designa- body, upperside is invisible; Baltic amber collected in Lithuania or 30 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

Latvia, Upper Eocene. maxillary palpomere elongate cultriform. Pronotum [strongly Derivatio nominis: damaged] with anterior rim, Patronymic. This species is devoted dorsally densely but shortly suberect to Mr. Albert Dean Allen (U.S.A.), pubescent. Elytra [strongly damaged] the first owner of the holotype. with distinct longitudinal sulcus Measurements [due to the along the lateral margin. Punctures position of the inclusion which is not distinct, partly forming more or less available in dorsal view, and strong regular rows in basal half of elytra. deformation (specimen is flattened Pubescence subdecumbent, short and dorsally) it is not possible to measure dense. Dorsally with separate longer the width of any of body parts]: Total and erect setae. Legs with distinctly body length ~3.0 mm. Head ~0.5 mm clavate femora. Tibial spurs long, long, pronotum ~0.7 mm long, elytra minutely setose. Three basal pro- and 1.8 mm long. mesotarsomeres strongly widened Colouration: Forebody brown and dorso-ventrally flattened. Basal to reddish, elytra darker brown to metatarsomere longer than combined black. Antennae and palpi paler length of three remaining tarsomeres. reddish, 2-3 terminal antennomeres All tarsal claws appendiculate. Hind darkened. Legs reddish except wings well developed. Last visible darkened tibiae. Trochanters reddish. ventrites without apical appendages. Abdomen reddish to reddish brown. Differential diagnosis: Description: Head [strongly Among the known fossil species, M. damaged] elongate, constricted alleni sp. nov. is remarkable due to posteriad to eyes. Base narrower the reddish colouration of the ventral than median part. Eyes large. Dorsal surface and legs (meso- and metatibi- pubescence with some separate long ae look darker than femora and tarsi), and erect setae. Groups of long setae head constricted posteriad, long tibial also on posteriolateral margins of spurs and strongly widened three bas- the head. Antennae reaching elytral al tarsomere of pro- and mesotarsi. humeri. Antennomeres 3-4 and 5-6 slender, almost equal in size and Macratria appendiculata (Abdul- proportions. Antennomeres 8-9 lah, 1964) comb. nov. (Figs 1-4) slightly shortened and widened Abdullah (1964: 333, plate 1: figs 1-3), as distally. Terminal antennomere Protomacratria appendiculata; Abdullah (1969: 353), as Protomacratria appendicu- elongate conical, almost twice so lata; Larsson (1978: 146), as Protomacratria long as penultimate one. Terminal appendiculata; Spahr (1981: 10), as Proto- Latvijas Entomologs 2012, 51: 27-39. 31

macratria appendiculata; Kirejtshuk (2002: Upper Eocene; ISZP 1 specimen: no pagination), as Protomacratria appen- MP/1348 Coleoptera: Anthicidae: diculata. Protomacratria appendiculata 1 det: D. Kubisz, 1999 waga: 3.4 g Dar: Holotype (BMNH): In. 18788 Jacek Serafin [single specimen], [Baltic amber, Upper Eocene]. Baltic amber [collecting locality Additional material 1?♀ unknown], Upper Eocene. LDM: No. G 404/4, beetle inclusion Notes: Abdullah gave the wrong in a slice of amber, with air bubble ID number of type specimen in his held on setae all around the specimen; original description of this species. Baltic amber collected in Lithuania or The holotype specimen is numbered Latvia, Upper Eocene. 3893 and not ‘no. 1065’ as mentioned Characters additional to the by Abdullah (1965). original description: Eyes very large and strongly prominent, roughly Macratria tripunctata (Abdullah, facetted. Tempora short. Head 1964) comb. nov. (Figs 4-5) base with short but broad median Abdullah (1964: 333, plate 1: fig. 4), as Pro- longitudinal impression. Tibial spurs tomacratria tripunctata; Abdullah (1969: minutely setose. Three basal pro- and 353), as Protomacratria tripunctata; Larsson mesotarsomeres strongly widened (1978: 146), as Protomacratria tripunctata; Spahr (1981: 10), as Protomacratria tripunc- and dorso-ventrally flattened. Basal tata; Kirejtshuk (2002: no pagination), as metatarsomere longer than combined Protomacratria tripunctata. wide of three remaining tarsomeres. Holotype (BMNH): In. 17723 Macratria succinia Abdullah, 1965 [Baltic amber, Upper Eocene]. (Figs 8-11) Characters additional to the Abdullah (1965: 38, plate 1: figs 1-2); Kire- original description: Eyes very jtshuk (2002), erroneously mentioned de- large and strongly prominent. scription year of this species as 1964; Kubisz (2000: 227), incorrectly identified as Proto- macratria appendiculata. Macratria sp. Klebs (1910: 237); Bachofen-Echt (1949: Holotype (GPIH): Typ.-Kat. 112); Abdullah (1964: 331); Larsson (1978: 3893 [Baltic amber, Upper Eocene]. 146); Spahr (1981: 10). Additional material, LDM 2 There are no indet. Macratria specimens: No. G 404/5 & G 404/6 in Klebs’s collection’s, part which is [two beetle inclusions about 3 mm stored at BMNH (C. Mellish pers. apart in a large slice of amber], Baltic comm.). I was not able to locate these amber collected in Lithuania or Latvia, 32 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

5 specimens at any museum. At least a 4 Legs red, underside reddish to reddish part of Kleb’s collection is considered brown. Head distinctly constricted posterior to be lost or even destroyed in to eyes ...... Macratria alleni sp. nov. Königsberg during World War II. – Legs dark to yellowish brown but never red. Underside dark brown to black. Head is not or only shortly constricted posterior to Identification key to fossil eyes ...... 5 Macratriinae (Anthicidae) 5 Head base narrowly rounded, notched medially. Third maxillary palpomere I have included all known fossil widened distally but distinctly longer than species of Macratriinae in this key, broad. Tempora short but clearly visible. and not only species described from Antennomeres 6-8 widened distally but the Baltic amber. elongate ...... Macratria appendiculata (Abdullah, 1964) 1 Pronotum strongly narrowed in posterior – Head base broadly rounded, notched part, anterior half being distinctly broader than medially. Third maxillary palpomere short posterior one. Three terminal antennomeres and strongly triangular, not longer than broad. very elongate, almost as long as combined Eyes very strongly prominent and large, length of 8 preceding antennomeres. almost covering all tempora. Antennomeres Lebanese amber (Lower Cretaceous) ...... 6-8 shortened and widened, looks ovoid ...... Camelomorpha longicervix ...... Macratria succinia Abdullah, 1965 Kirejtshuk, Azar et Telnov, 2008 – Pronotum only slightly narrowed posteriad, anterior half indistinctly broader than Acknowledgements posterior one. Three terminal antennomeres much shorter ...... 2 I am deeply indebted to Aneta 2 Large species, total body length (excluding Garbula and Daniel Kubisz (both exposed abdominal ventrites) ~8.0 mm. ISZP), Dr. Ulrich Kotthoff (GPIH), Pronotum elongate and narrow. Florissant Claire Mellish (Department of formation of Colorado (Eocene) ...... Palaeontology, BMNH) for providing ...... Macratria gigantea Wickham, 1910 material for study and friendly – Species of distinctly smaller body size. assistance in preparing photographs of Pronotum not very narrow. Species from previously described species. Albert Baltic amber (Upper Eocene) ...... 3 Dean Allen (Boise, Idaho, U.S.A.) is 3 Tarsal claws of all tarsi appendiculate .... 4 thanked for his altruism for providing – Tarsal claws of at least one leg pair simple . highly interesting Macratriinae .... Macratria tripunctata (Abdullah, 1964) specimens from Baltic amber. I Latvijas Entomologs 2012, 51: 27-39. 33

am also grateful to Dr. Stephen G. group names in Coleoptera Compton (Leeds University, Leeds, (Insecta). – ZooKeys 88: 972 pp. United Kingdom) for reading and Chandler D.S. 2010. 11.26. Anthicidae commenting on the manuscript. Latreille, 1819: 729-741. In: Leschen R.A.B., Beutel R.G. & Lawrence J.F. (eds) Coleoptera, References . Volume 2: Morphology and systematics (Elateroidea, Abdullah M. 1964. New heteromerous Bostrichiformia, Cucujiformia beetles (Coleoptera) from the partim). Arthropoda Insecta. Baltic amber of eastern Prussia Handbook of zoology. Berlin & and gum copal of Zanzibar. New York, De Gruyter: 786 pp. – Transactions of the Royal Kirejtshuk A.G. 2002. Catalogue of Entomological Society of fossil Coleoptera [permanent London 116, No. 13: 329-346, website] – http://www.zin. pls. I-II. ru/Animalia/Coleoptera/eng/ Abdullah M. 1965. New Anthicidae paleosys.htm [last accessed: and Pyrochroidae (Coleoptera) 15.08.2011]. from the Baltic Amber Klebs R. 1910. Über Bernstein (Oligocene). – The Entomologist Einschlüße im allgemeinen 98: 38-42. und die Coleopteren meiner Abdullah M. 1969. The natural Bersteinsammlung. – classification of the family Schriften der Physikalisch- Anthicidae with some ecological Ökonomischen Gesellschaft zu and ethological observations. Königsberg 51, No. 3: 217-242. – Deutsche entomologische Kubisz D. 2000. Fossil beetles Zeitschrift 16, No. 4/5: 323-366. (Coleoptera) from Baltic amber Bachofen-Echt A. 1949. Der in the collection of the Museum Bernstein und seine Einschlüße. of Natural History of ISEA Springer Verlag, Wien: 1-204 + in Krakow. – Polskie pismo 1 pl. + 188 figs. entomologiczne 69, No. 2: 225- Bouchard P., Bousquet Y., Davies 230. A.E., Alonso-Zarazaga M.A., Larsson S.G. 1978. Baltic Amber Lawrence J.F., Lyal C.H.C., - a palaeobiological study. Newton A.F., Reid C.A.M., Entomonograph 1: 192 pp. Schmitt M., Ślipiński S.A., Spahr U. 1981. Bibliographie der Smith A.B.T. 2011. Family- Berstein- und Kopal-Käfer 34 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

(Coleoptera). – Stuttgarter In: Telnov D. (ed.) Biodiversity, Beiträge zur Naturkunde, Serie biogeography and nature B (Geologie und Paläontologie) conservation in Wallacea 80: 1-107. and New Guinea: Rīga, The Telnov D. 2009. Review of the genus Entomological Society of Telesinus Fairmaire, 1903, with Latvia: 434 pp. + 92 pls. a general discussion on African Eurygeniinae (Coleoptera: Anthicidae). – Mitteilungen Received: September 25, 2011. des Internationalen Entomologischen Vereins e.V. 34, No. 3/4: 93-113. Telnov D. 2011. Taxonomische Revision der Gattung Macratria Newman, 1838 (Coleoptera: Anthicidae: Macratriinae) aus Wallacea, Neuguinea und den Salomonen: 97-286, pls. 17-37. Latvijas Entomologs 2012, 51: 27-39. 35

Figures 1-4. Macratria appendiculata (Abdullah, 1964), holotype BMNH (photo: P. Hurst, © BMNH). 1 - habitus (dorsal), 2 - forebody (dorsal), 3 - habitus (lateral). 36 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

Figures 4-5. Macratria tripunctata (Abdullah, 1964), holotype BMNH (photo: P. Hurst, © BMNH). 4 - elytra and legs (lateral), 5 - habitus (lateral). Latvijas Entomologs 2012, 51: 27-39. 37

Figures 6-7. Macratria alleni sp. nov., holotype LDM. 6 - habitus (lateral), 7 - legs. 38 Macratriinae (Coleoptera: Anthicidae) of the Baltic amber

Figures 8-9. Macratria succinia Abdullah, 1965, holotype GPIH (photo: U. Kotthoff, © GPIH). 8 - habitus (dorso-lateral), 9 - elytra (dorsal). Latvijas Entomologs 2012, 51: 27-39. 39

Figures 10-11. Macratria succinia Abdullah, 1965, holotype GPIH (photo: U. Kotthoff, © GPIH). 10 - head & mesosternum (lateral), 11 - forebody (dorsal).