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New Finds of the Fossil Ant Genus Prionomyrmex Mayr (Hymenoptera, Formicidae, Myrmeciinae) in Late Eocene European Amber A

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New Finds of the Fossil Ant Genus Prionomyrmex Mayr (Hymenoptera, Formicidae, Myrmeciinae) in Late Eocene European Amber A ISSN 0031-0301, Paleontological Journal, 2020, Vol. 54, No. 6, pp. 617–626. © Pleiades Publishing, Ltd., 2020. Russian Text © The Author(s), 2020, published in Paleontologicheskii Zhurnal, 2020, No. 6, pp. 60–67. New Finds of the Fossil Ant Genus Prionomyrmex Mayr (Hymenoptera, Formicidae, Myrmeciinae) in Late Eocene European Amber A. G. Radchenkoa, * and E. E. Perkovskya, b, ** aSchmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiev, Ukraine bBorissiak Paleontological Institute, Russian Academy of Sciences, Moscow, Russia *e-mail: [email protected] **e-mail: [email protected] Received November 26, 2019; revised January 24, 2020; accepted February 3, 2020 Abstract—A new species of the fossil genus Prionomyrmex, P. gusakovi sp. nov., represented by three workers from Baltic amber (Late Eocene, Priabonian, 33.9–37.2 Mya), is described. Prionomyrmex longiceps Mayr was first identified in the coeval Rovno amber from Ukraine. The differences between the species described and other known species of the genus are presented. The main problems of evolution and systematics of the extinct representatives of the subfamily Myrmeciinae are discussed. Keywords: ants, Formicidae, Prionomyrmex gusakovi sp. nov., paleontology, Baltic, Bitterfeld, and Rovno ambers, systematics, evolution DOI: 10.1134/S0031030120060088 INTRODUCTION the tribe Prionomyrmecini to the subfamily Myrmeci- The extinct ant genus Prionomyrmex was estab- inae, but Baroni Urbani (2000) raised the tribe Prion- lished by Mayr (1868), based on P. longiceps Mayr, omyrmecini to a subfamily level and synonymized the 1868, represented by a single worker from Baltic generic name Nothomyrmecia Clark, 1934 with Prion- amber. Later, Wheeler (1915) studied nine Prionomyr- omyrmex, so that he regarded the name of the subfam- mex workers from the Baltic amber (but not the holo- ily Nothomyrmeciinae as a junior synonym of Priono- type!), reported additional important diagnostic fea- myrmecinae. Dlussky and Perfilieva (2003) as well as tures of this genus, and first described the male (a single Ward and Brady (2003) independently transferred Pri- specimen without the abdomen), which he identified as onomyrmex back to Myrmeciinae and resurrected the P. longiceps. Unfortunately, all the material mentioned name Nothomyrmecia from synonymy. However, Bar- above was destroyed during World War II. oni Urbani (2005) returned to his own earlier opinion on the subfamily status of the Prionomyrmecinae and The genus was considered monotypic until the end th the synonymy of Prionomyrmex and Nothomyrmecia. of the 20 century, but then Baroni Urbani (2000) Finally, Archibald et al. (2006), and later, Dlussky described P. janzeni, a new species from Baltic amber, (2012), restored the “status quo”, so that Prionomyr- which is very close to P. longiceps and differing from mex and Nothomyrmecia are currently regarded as sep- the latter mainly by the absence of erect setae on the arate genera of Myrmeciinae; and we fully support this scape. Finally, Dlussky (2012) recently described a opinion. new species, P. wappleri, represented by an imprint from the Upper Oligocene Rott fossil locality in Ger- To date, thirteen workers and one male of P. longi- many, and a large (approximately 20 mm long) Myr- ceps have been reported: the holotype, 10 specimens meciinae worker, which has not yet been described, described by Wheeler, and photographs of two specimens was recorded from the Early Eocene Oise amber are available on the AntWeb website (https://www.ant- (France) (Aria et al., 2011; La Polla et al., 2013). web.org/images.do?subfamily=myrmeciinae&genus= Prionomyrmex was first assigned to the subfamily prionomyrmex&rank=genus&project=allantwebants); Ponerinae (Mayr, 1868). Wheeler (1915) also regarded importantly, they all were found in Baltic amber. In this genus as a representative of Ponerinae, but addition, the holotype and the paratype of P. janzeni, assigned it to a new tribe, Prionomyrmicini (sic!), even and the holotype of P. wappleri are also available. though Emery (1877) earlier placed this species into We investigated five specimens (workers) in five the “group” Myrmeciidae. Brown (1954) transferred pieces of amber. We identified two specimens from 617 618 RADCHENKO, PERKOVSKY Rovno and Bitterfeld amber as P. longiceps (see the frontal carinae; FLW, distance between the exter- below), and assigned three workers from Baltic amber nal edges of the frontal lobes; SL, scape length from its to the species P. gusakovi sp. nov., described below. apex to the articulation with condylar bulb; EL, eye The specimen from Rovno amber is the first record of length (maximal diameter); MDL, mandible length; a representative of the subfamily Myrmeciinae in ML, thorax (mesosoma) length in side view measured Ukraine; thus, the overall number of Rovno from the anterior margin of the neck shield to the pos- hymenopteran species, including P. longiceps and terior margin of the metasternal lobes; MH, thorax another nine chrysidoid and encyrtid species found in (mesosoma) height in side view, measured from the Rovno amber (Martynova et al., 2019; Colombo et al., uppermost point of the promesonotum perpendicu- 2020; Perkovsky et al., 2020; Simutnik and Perkovsky, larly to the lowermost point of the mesopleuron; 2020), reached 128. Only 64 hymenopteran species PRW, pronotum width from the top; PL, petiole from Rovno (50%) are also known from Baltic amber length; PH, petiole height in side view; PW, petiole (Perkovsky, 2018). width from the top; PPL, postpetiole length; PPH, postpetiole height in side view, from the uppermost to the lowermost point, measured perpendicular to the MATERIALS AND METHODS tergo-sternal suture; PPW, postpetiole width at the One P. longiceps specimen (worker) from Rovno top; L-IV, length of abdominal segment IV; H-IV, amber is deposited in the private collection of height of abdominal segment IV; GL, gaster length; N.R. Khomich’s (Rovno, Ukraine), no. F-125. HTL, hind tibia length; body length was calculated as Beside this, photographs of a specimen (worker with- the total length of the head, thorax (mesosoma), peti- out a head) from Bitterfeld amber (Germany) identi- ole, and gaster. fied as P. longiceps by Dlussky were available to use; Index abbreviations are not given here to facilitate the photographs were taken by Dlussky in 2003. We the perception of information, but rather the size also studied three workers of P. gusakovi sp. nov. (the ratios of various structures are presented: for example, holotype and two paratypes) from Yantarnyi, Kalinin- HL/HW, rather than HI. grad Region, Russia. Holotype and paratype No. 964/1320 of the species described are deposited at Paleontological Institute, Russian Academy of Sci- SYSTEMATIC PALEONTOLOGY ences (PIN RAS) in Moscow, and the paratype is Subfamily Myrmeciinae Emery, 1877 deposited at the Geologisch-Paläontologisches Insti- tut des Universität Hamburg (GPIH; currently named Genus Prionomyrmex Mayr, 1868 the Centrum of Natural History, CeNak) (GPIH Type species. P. longiceps Mayr, 1868, by no. 4993; no. 6762 of C. Gröhn’s collection). monotypy; Baltic amber, Late Eocene. D i a g n o s i s. Workers large, polymorphic with STATE OF THE INCLUSIONS regard to size, body length 9.4–14.8 mm. Body slen- der, elongated, with long legs. Antennae with 12 seg- The holotype of P. gusakovi is visible in the right ments, scape long, somewhat protruding beyond the lateral position; it is a whole specimen that lacks occipital margin of the head, funiculus filiform, with- antennae; a Lasius sp. worker is found in the same out a club. Mandibles long, narrow triangle-shaped, piece of amber. The paratype PIN no. 964/1320 is vis- masticatory margin with a series of small denticles and ible from the ventral side; it is a whole specimen coarse short bristles. Eyes large, located approxi- largely obscured by fissures in the amber; the anten- mately in the middle of the lateral sides of the head or nae, maxillary and labial palps, tibial spurs, tarsal slightly posterior to them. Maxillary palps of six seg- claws, and decumbent pubescence on the ventral sur- ments, which is in agreement with the report by face of the head, coxae, and upper part of the petiole Wheeler (1915), labial palps with four segments. It is are well visible; a Dolichoderus sp. worker is found in necessary to note that Baroni Urbani (2000) reported the same piece of amber. The paratype GPIH no. 4993 a palp formula of 5, 2 for P. janzeni but did not rule out is a whole specimen with most diagnostically signifi- the possibility of the basal segments being poorly visi- cant details visible. ble in the specimen studied. The petiole without the A Leica Z16 APO stereomicroscope connected to a anterior peduncle and with a well-developed node Leica DFC 450 camera was used to take the photo- with a narrowly rounded dorsum. The constriction graphs, which were further processed in LAS Core between the abdominal segments III and IV is sharp software. and deep. Middle and hind tibiae with two spurs (sim- It was impossible to measure all structural details in ple and pectinate). Tarsal claws with an additional all specimens, and therefore we measured as many of preapical tooth (see also: Mayr, 1868; Wheeler, 1915; them as possible with a 0.01 mm precision: HL, head Baroni Urbani, 2000; Dlussky, 2012). length from the anterior edge of the clypeus to the The reader is referred to the study by Wheeler middle of the occipital margin; HW, maximal head (1915, p. 27) for a characterization of the only male of width behind the eyes; FW, minimal distance between this genus described so far. PALEONTOLOGICAL JOURNAL Vol. 54 No. 6 2020 NEW FINDS OF THE FOSSIL ANT GENUS PRIONOMYRMEX MAYR 619 Species composition. Two species, P. lon- segment ІІІ) is much more robust than the petiole, giceps and P. janzeni, from Baltic amber; the former convex, expands gradually backwards, bell-shaped, also found in Bitterfeld and Rovno amber, as reported distinctly lower and shorter that the first gastral seg- in the present study; P.
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