Journal of Paleontology, 87(2), 2013, p. 177–182 Copyright Ó 2013, The Paleontological Society 0022-3360/13/0087-177$03.00 REMARKABLE STASIS IN A PHLOEOCHARINE ROVE BEETLE FROM THE LATE CRETACEOUS OF NEW JERSEY (COLEOPTERA, STAPHYLINIDAE) STYLIANOS CHATZIMANOLIS,1 ALFRED F. NEWTON,2 CARMEN SORIANO,3 AND MICHAEL S. ENGEL4 1 Department of Biological and Environmental Sciences, Dept. 2653, The University of Tennessee at Chattanooga, 615 McCallie Ave., Chattanooga, TN 37403, USA, ,stylianos-chatzimanolis@utc.edu.; 2Division of Insects, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA, ,anewton@fieldmuseum.org.; 3European Synchrotron Radiation Facility, BP220, 6 rue Jules Horowitz, 38043 Grenoble Cedex, France, ,carmen.soriano@gmail.com.; and 4Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive, Suite 140, University of Kansas, Lawrence, KS 66045, USA ,msengel@ku.edu. ABSTRACT—The first definitive fossil species of the rove beetle (Staphylinidae) subfamily Phloeocharinae is described and figured from a single individual preserved in Late Cretaceous (Turonian) amber from New Jersey. The species is representative of the extant genus Phloeocharis Mannerheim and is described as Phloeocharis agerata Chatzimanolis, Newton, and Engel, new species. The specimen was imaged using traditional light microscopy as well as synchrotron propagation phase contrast microtomography, permitting a detailed examination of otherwise difficult to observe features. Examination revealed remarkable homogeneity across many characters with those of extant relatives, highlighting considerable morphological stasis in the genus over the last 90 million years. INTRODUCTION T beetle subfamily Phloeocharinae is a small group of seven genera and 59 species. The monophyly of Phloeocharinae has not been tested and historically the subfamily had been a dumping ground for several primitive-looking rove beetles (Newton et al., 2000), although Herman (1972) suggested several characteristics that have served as a working definition of the group. Currently, the following genera are included: Charhyphus Sharp (with a Neotropical, Nearctic, and Palearctic distribution), Dytoscotes Smetana and Campbell (Nearctic), Ecbletus Sharp (Neotropical and Nearctic), Phloeocharis Mannerheim (Nearctic and Palearctic), Phloeognathus Steel (Australian), and Pseudophloeocharis Steel (Australian and Oceanic). Little to nothing is known of phloeocharine biology, but where data are available they are found in forest litter, frequently flightless (e.g., Assing 2003) or are in a few cases blind and endogean (e.g., Coiffait, 1957; Hernando, 2003). The fossil record of the subfamily is exceptionally poor. Scudder (1900) described a species from the Eocene–Oligocene boundary of Florissant, Colorado as ‘‘Triga coeni’’ (now placed in Charhyphus simply owing to the synonymy of Trigites Handlirsch, the generic replacement name for Triga Fauvel, with this genus). However, in his revision of Charhyphus Herman (1972) noted, ‘‘. . . the characters in the description and the photograph of the specimen offer no compelling evidence for believing the species belongs to this genus’’. Indeed, the specimen is a minute compression with little to no relief and sufficient characters for placing the individual in Phloeocharinae are simply not preserved, hence we consider Charhyphus coeni to be ‘‘Staphylinidae incertae sedis’’. Klebs (1910) and Bachofen-Echt (1949) listed individuals of Phloeocharis in midEocene Baltic amber (repeated in the catalog of Spahr, 1981), but provided no further details and the location of their private material is currently uncertain (e.g., a subset of Bachofen-Echt’s material apparently survives in Munich [Dunlop, 2006] with the remainder missing, while Klebs’s collection was thought to have perished in Königsberg during the last world war [Ley, 1951] but small portions apparently survive in Göttingen [Ritzkowski, 1990] and London [Fikáček and Engel, 2011]). Since Klebs HE ROVE (1910) indicated that his ‘‘sehr schön’’ specimen of Phloeocharis had been studied and named to genus by Edmund Reitter, a leading coleopterist who was familiar with the genus (e.g., Reitter, 1909), this identification may well be correct. Recently, Yue et al. (2010) described Megolisthaerus chinensis from the Early Cretaceous Yixian Formation in northeastern China and hypothesized that the species might be related to the subfamilies Olisthaerinae and Phloeocharinae but a new paper based on more complete material has reclassified Megolisthaerus as a staphylinine (Cai and Huang, 2013). Among amber material recovered from the Raritan Formation of New Jersey was discovered a single individual of the extant genus Phloeocharis. The individual is beautifully preserved but a series of fractures in the amber piece render difficult many views of the specimen (Fig. 1). In order to obtain a more complete understanding of the specimen and permit a thorough characterization of the species, the amber piece was subjected to synchrotron x-ray microtomography as detailed below. Herein we provide an account of this species as well as long-term morphological stasis in Phloeocharinae and other rove beetle lineages. MATERIAL AND METHODS Light microscopy and measurements were undertaken with an Olympus SZX12 stereomicroscope. The geology, chemistry, age, and biota of New Jersey amber have been reviewed by Grimaldi et al. (2000) and Grimaldi and Nascimbene (2010). The holotype is affected by several cracks in the amber, as well as partially covered by a white froth of microscopic bubbles (Fig. 1). To resolve detail of pertinent anatomical characters the specimen was imaged at beamline ID19 in the European Synchrotron Radiation Facility (ESRF, Grenoble) using a propagation phase protocol as described in Tafforeau et al. (2006) and Soriano et al. (2010). The scan consisted of 2000 projections acquired through a 1808 rotation and 0.3 seconds of exposure time, and a beam set at 20 keV using a 17.6 m period undulator with a gap of 14.5 mm and 2 mm of aluminum as filter. The scan voxel size was 1.4 microns, and the propagation distance 25 mm. After the scan the volume was reconstructed 177 178 JOURNAL OF PALEONTOLOGY, V. 87, NO. 2, 2013 FIGURE 1—Light photomicrographs of holotype (AMNH NJ-88) of Phloeocharis agerata Chatzimanolis, Newton, and Engel, new species, as preserved. 1, dorsal view; 2, lateral view. Much of the individual is obscured by fractures from various angles of view. using a back filtered projection algorithm especially adapted for local tomography (PyHST, ESRF), and later segmented using a manual region growing approach in VGStudioMax software (ver. 2.1, Volume Graphics, Heidelberg). All of the microtomographic data including original and segmented slices, acquisition parameters, pictures and animations, and stereolitographic mesh volume will be available after publication at the ESRF paleontological online database (paleo.esrf.eu). SYSTEMATIC PALEONTOLOGY Family STAPHYLINIDAE Latreille, 1802 Subfamily PHLOEOCHARINE Erichson, 1839 Genus PHLOEOCHARIS Mannerheim, 1830 PHLOEOCHARIS AGERATA Chatzimanolis, Newton, and Engel, new species Figures 1–3 Diagnosis.—Phloeocharis agerata can be most easily distinguished from other species in the genus by having a relatively larger maxillary palpomere IV (subequal in length to and nearly half as wide at base as palpomere III) and elytra much longer than the pronotum. Recent species of Phloeocharis typically have a maxillary palpomere IV that is no more than 0.7 times as long as and a third as wide as palpomere III, and elytra that are subequal in length to or shorter than the pronotum (but elytra are about a third longer than pronotum in at least one species, P. hummleri Bernhauer). Description.—Total length (as preserved) approximately 1.7 mm; small species, body subparallel-sided, convex; integumental coloration poorly preserved, apparently dark brown to black (where evident). Head small, much narrower than pronotum, width (including compound eyes) 0.33 mm, upper interocular distance 0.25 mm; head constricted laterally immediately behind compound eyes to form broad neck; compound eyes mediumsized, with many ommatidia; epistomal suture apparently absent. Labrum slightly narrower than clypeus; mandibles short and stout; maxillae not visible except maxillary palp, maxillary palpomere I not visible, palpomere II long and slender, palpomere III subequal in length but much wider than palpomere II, palpomere IV conical, thin, nearly as long as but less than half as wide at base as palpomere III. Labium not clearly visible except labial palpi. Antennomeres I–III robust, longer than wide, gradually decreasing in width; antennomere IV small, quadrate, almost 0.5 times as long as antennomere III; antennomeres V–X subquadrate, gradually increasing in size; antennomere XI twice as long as antennomere X. Pronotum distinctly wider than head, transverse, maximum width at posterior angles, maximum width of pronotum 0.52 mm, lateral margins gently convex; posterior angle rounded but distinct, approximately orthogonal, posterior CHATZIMANOLIS ET AL.—STASIS IN ROVE BEETLES 179 FIGURE 2—PPClCT of holotype (AMNH NJ-88) of Phloeocharis agerata Chatzimanolis, Newton, and Engel, new species, imaged at ESRF. Voxel size 1.4 microns. margin relatively straight in dorsal view; medial length of pronotum 0.28 mm. Elytra about as wide as pronotum, without epipleural fold laterally, at suture clearly longer than pronotum, combined width of elytra 0.53 mm, length of elytron 0.48 mm, posterior margin weakly convex, almost straight medially; elytra not impressed near mesoscutellum; medial length of mesoscutellum 0.09 mm. Legs relatively short; procoxa without mesal groove; protibia without spines, meso- and metatibia with spines apically; tarsomeres short except distitarsi (distitarsus¼apicalmost tarsomere) elongate, slightly (pro- and mesodistitarsomeres) to much (metadistitarsomere) longer than combined length of respective basitarsi (basitarsus¼basalmost tarsomere) and mediotarsi (mediotarsus¼medial tarsomeres, or those between the basitarsus and distitarsus), tarsal formula 5-5-5, pro- and mesobasi- and mediotarsomeres distinctly widened; pretarsal claws arcuately curved. Abdomen nearly as wide as combined width of elytra, maximum width of abdomen 0.50 mm; individual abdominal segments largely transverse except segment VII enlarged and more quadrate (Fig. 2), segments III–VII each with a single pair of slender paratergites laterally, terga with scattered punctures over otherwise finely imbricate integument; similar sculpturing on sterna; terga and sterna with scattered, caudallydirected, appressed to suberect, simple setae mostly along posterior quarters to thirds of length, setae appearing rather stiff. Distal margin of sternum VII with a slight V-shaped emargination. Tip of aedeagus visible. Holotype.—Male, AMNH NJ-88 (Fig. 1); Late Cretaceous (Turonian), New Jersey, E. Brunswick, Sunrise Landing site, G.R. Case coll.; deposited in the Division of Invertebrate Zoology (Entomology), American Museum of Natural History, New York, New York. The specimen is relatively complete, with small bubbles of air appressed against the body and emanating from beneath the apices of the elytra, at the abdominal apex and extending dorsoanteriorly over the apicalmost segments, between the abdomen and the right hind leg, and from the mouth posteriorly along the venter of the head to the right procoxa (Fig. 2). Etymology.—The specific epithet is taken from the Greek word 180 JOURNAL OF PALEONTOLOGY, V. 87, NO. 2, 2013 FIGURE 3—Synchrotron radiation computerized tomographic detail of head of Phloeocharis agerata Chatzimanolis, Newton, and Engel, new species (AMNH NJ-88). ageratos, meaning, ‘‘ageless’’, as a reference to the remarkable bradytely of the species. DISCUSSION The species described here keys with some difficulty (not all of the pertinent characters are visible, even with the PPClCT reconstruction due to preservational problems) to the subfamily Phloeocharinae based on the staphylinid keys provided by Newton et al. (2000). Herman (1972) and Smetana (1983) suggested several diagnostic characters for the placement of genera in Phloeocharinae, these being: procoxa without mesal groove, antennal insertions more or less concealed from above, abdominal terga IV and V with a pair of cuticular combs, and a distinctive hypopharynx. Of these characters, only the first two are clearly visible and present in P. agerata, while the hypopharynx is not visible and the reconstruction from the synchrotron tomography is not sufficient clear for the morphology of terga IV and V. However, the species described here has the other diagnostic characteristics mentioned by Newton et al. (2000) in the description of the subfamily such as procoxal fissures open, tarsal formula of 5-5-5, abdomen with six visible sterna, and a single paratergite per segment. The combination of those preserved features all support placement in Phloeocharinae rather than any other lineage of staphylinids. Using the key to Nearctic genera of Phloeocharinae in Newton et al. (2000), which includes all five northern hemisphere genera of Phloeocharinae, the species described here keys readily to Phloeocharis, and differs from the other four genera in many characteristics as evident in that key. However, Phloeocharis was considered closely related by Steel (1950, 1953) to the two remaining phloeocharine genera that are not included in this key, namely Phloeognathus and Pseudophloeocharis from the Australian and Oceanic regions, and Newton (1985) likewise treated these three genera together as a ‘‘probably monophyletic’’ Phloeocharis generic group. This group (including a subsequently discovered undescribed genus CHATZIMANOLIS ET AL.—STASIS IN ROVE BEETLES and species from Chile) can be characterized now by sharing the following characteristics: head much narrower than pronotum and with a small lateral (but not dorsal) constriction immediately behind eyes, forming a very broad neck; epistomal suture absent; elytron without an epipleural fold; at least pro- and mesotarsi with basal tarsomeres expanded and bearing tenent setae in both sexes; abdomen with a single pair of paratergites on most segments; abdominal tergite III only with a subbasal carina; and abdominal sternite III without carina-delimited coxal cavities for the metacoxae. Phloecharis agerata clearly belongs in this group as far as these characteristics can be seen (last two characters not clearly visible), and agrees uniquely with Phloeocharis in the small size of maxillary palpomere IV relative to III (in all southern hemisphere genera palpomere IV is at least as wide as, and about twice as long as, palpomere III). Additionally, P. agerata and other Phloeocharis lack the slender falcate mandibles of Phloeognathus. Together, all of these extinct and extant members of the Phloeocharis group form a probably monophyletic core of the subfamily Phloeocharinae, although monophyly of this core remains to be rigorously tested. The inclusion of the remaining northern hemisphere genera (especially the morphologically isolated Ecbletus and Vicelva) in this subfamily remains more doubtful. As mentioned above, Phloeocharis is Holarctic in distribution and includes 44 described species, most of which occur in the Western Mediterranean region (Herman, 2001; Hernando 2003; Assing, 2003, 2004, 2006a, 2006b; Feldmann, 2004). Based on our examination of 10 of these 44 species, and descriptions of others, P. agerata evidently differs from the modern species only in having a maxillary palpomere IV that is slightly larger, and elytra that are slightly longer, than the range seen in modern species. In both cases these are likely to be plesiomorphic conditions within Phloeocharis, but we can find no justification for separating P. agerata generically from the modern species. The beetle family Staphylinidae is one of the most hyperdiverse of all animal lineages, with over 58,500 described living species (Newton, unpublished data). The family is also one of the more long-lived, with recognizable individuals extending back well into the Triassic and many of the extant subfamilial lineages extending into Jurassic or earliest Cretaceous (Chatzimanolis et al., 2012). Indeed, current paleontological evidence tends to suggest that significant diversification took place within the family early in its history, with many of the principal higher monophyla having originated during the Jurassic, none of which seem to represent radiations which subsequently failed to survive, such as is the case for non-avian dinosaur clades relative to birds. In addition to the great longevity and persistence of rove beetle lineages, a growing number of Mesozoic fossils are being discovered which are representative of modern genera, or so strikingly similar as to be scarcely diagnosable relative to their living counterparts (e.g., Clarke and Chatzimanolis, 2009). The present species of Phloeocharis certainly represents yet another dramatic example of bradytely (‘‘arrested evolution’’; Simpson, 1944) among beetles. Clarke and Chatzimanolis (2009) described a case of bradytely in the rove beetle genus Octavius Fauvel (Staphylinidae, Euasthetinae) and they hypothesized that the constant presence of mesic habitats over geological time is responsible for the stasis in this group. In other words, the constant presence of mesic habitats, along with the relative small size of these beetles has protected them from extinction and strong selection. In another example of bradytely, Cognato and Grimaldi (2008) ascribed the stasis observed over 100 million years in the scolytine beetle Microborus Blandford to the constraints of their subcortical habitats. This same kind of habitat likely explains the bradytely 181 observed in Zorotypus Silvestri (Zoraptera), along with the general antiquity of many polyneopterous lineages (Engel and Grimaldi, 2002). Many rove beetle lineages, particularly basal lineages of individual subfamilies, have a similar mode of life, with much speciation centering around microhabitat differentiation or feeding biologies, and this kind of conservatism is likely a contributing factor to the long-term stability of clades. Naturally, rigorous cladistic tests as well as a comprehensive revision of the fossil record of Staphylinoidea are needed but even what limited data is available attests to a group of considerable ecological and evolutionary success, and significant long-term morphological stasis as evidenced by a relatively stable groundplan morphology at least 220 million years old and genera in multiple subfamilies dating back at least 90–100 million years. ACKNOWLEDGMENTS Partial support was provided by U.S. National Science Foundation grants DEB-0741475 (to S.C. and M.S.E.) and DEB-0542909 (to M.S.E.). We are grateful to D. A. Grimaldi, and the late J. S. 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Megolisthaerus chinensis get. et sp. n. (Coleoptera: Staphylinidae incertae sedis): An enigmatic rove beetle lineage from the Early Cretaceous. Insect Systematics and Evolution 41: 317–327. ACCEPTED 2 NOVEMBER 2012