PROC. ENTOMOL. SOC. WASH. 115(2), 2013, pp. 173–181

PTENIDIUM KISHENEHNICUM (COLEOPTERA: ), A NEW FOSSIL DESCRIBED FROM THE KISHENEHN OIL SHALES, WITH A CHECKLIST OF PREVIOUSLY KNOWN FOSSIL PTILIIDS

FLOYD W. SHOCKLEY AND DALE GREENWALT

(FWS) Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC–165, Washington, DC 20013–7012 (e-mail: [email protected]); (DG) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC–121, Washington, DC 20013–7012 (e-mail: [email protected])

Abstract.—Ptenidium kishenehnicum Shockley and Greenwalt,anew of feather-winged (Coleoptera: Ptiliidae), is described from 46 million year old Kishenehn oil shales in Montana, USA. This compression fossil is the first beetle species to be described from this formation. A checklist of known fossils and their ages is provided. Key Words: compression fossil, feather-winged , fossil beetles, , paleoentomology DOI: 10.4289/0013-8797.115.2.173

The feather-winged beetles (Coleoptera: The fossil record for Ptiliidae is better Ptiliidae), while worldwide in their dis- than one might expect given the numbers tribution, constitute a relatively small of described extant species and the bee- family, with approximately 635 described tles’ small physical size. Several major species (Grebennikov 2009). They are catalogs of fossil beetles (Klebs 1910, unique in that they are among the small- Spahr 1981, Carpenter 1992) have in- est known, with body lengths of cluded Ptiliidae. Ptiliids have been found 400 mm or less reported, and with fe- in amber from the Baltic, the Dominican males having a single ovary—the Republic, Lebanon, Rovno, Myanmar and can accommodate only one maturing egg Mexico (Table 1). In fact, ptiliids have at a given time (Polilov 2005). The small been reported to make up 11% of all size of the Ptiliidae may also be re- Coleoptera in the American Museum of sponsible for other unusual character- Natural History collection of Myanmar istics, such as complete lack of a heart amber (Grimaldi et al. 2002). Isolated (Grebennikov 2008) and the extremely ptiliid elytra have also been found in small wing membranes, often 20 times Holocene, Pleistocene and Pliocene de- greater in length than width, with nu- posits (Matthews 1977, Matthews and merous long setae attached to their mar- Telka 1997). Despite the relatively large ginal edges that contribute the majority of number of reports of fossil ptiliids, only the wing’s surface area (Horridge 1956, five species of Ptiliidae have been de- Grebennikov 2008). scribed(Table1).Compressionfossils 174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. References to fossils of Ptiliidae Erichson, 1845, with approximate age of formations. Ages of formations largely derived from Grimaldi and Engel (2005).

Taxon Reference(s) Fossil Type Age

Ptiliidae Poinar 1992, Poinar and Poinar 1999, Dominican amber 20–17 Ma Wu 1996 Ptiliidae Poinar 1992 Mexican amber 29.0–23.6 Ma Ptiliidae Poinar and Poinar 2008 Lebanese amber 125 Ma Ptiliidae Rasnitsyn and Ross 2000, Grimaldi 95 Ma et al. 2002, Poinar and Poinar 2008 Ptiliidae Helm 1896, Klebs 1910, Larsson 1978, Baltic amber 44 Ma Kulicka and Slipi nski 1996 sp. Matthews 1977 Disarticulated fragments 5.7 Ma Acrotrichis sp. Matthews and Telka 1997 Disarticulated fragments 9.36 Ka Acrotrichis sp. Matthews and Telka 1997 Disarticulated fragments 125 Ka Acrotrichis sp. Matthews and Telka 1997 Disarticulated fragments 5.2–1.7 Ma groehni Polilov and Perkovsky 2004 Baltic amber 44 Ma Micridium sp. Matthews 1977 Disarticulated fragments 5.7 Ma geistautsi Dybas 1961 Baltic amber 44 Ma Ptilium tertiarium Horion (in Statz and Horion) 1937 Compression 23.6–21.0 Ma Ptilium sp. Polilov and Perkovsky 2004 Baltic amber 44 Ma Ptilium sp. Polilov and Perkovsky 2004 Rovno amber 54.8–33.7 Ma oligocenica Parsons 1939 Baltic amber 44 Ma Ptinella rovnoensis Polilov and Perkovsky 2004 Rovno amber 54.8–33.7 Ma Ptenidium kishenehnicum Present paper Compression 46.2–43.5 Ma Ptenidium sp. Klebs 1910, Handlirsch 1925, Baltic amber 44 Ma Bachofen-Echt 1949, Polilov and Perkovsky 2004 of Ptiliidae are even more rare, with only the Kishenehn Formation, these speci- a single such fossil described (Statz and mens were never formally described. We Horion 1937). The small number of fossil herein describe the ptiliid fossil Ptenidium species is not only a result of their ex- kishenehnicum Shockley and Greenwalt, tremely small size, but also the result of new species, the first species of any a reliance on anatomical details rarely Coleoptera to be described from the preserved in fossilized specimens but re- Kishenehn oil shales and the first fossil quired for ptiliid species identification. ptiliid to be described from the New The oil shales of the Kishenehn For- World. mation in northwestern Montana have re- cently been shown to contain exquisitely MATERIALS AND METHODS preserved insects with a bias for the pres- The compression fossil described herein ervation of very small insects (Greenwalt is housed in the National Museum of et al. 2011). For example, six new species of Natural History (NMNH). It was col- Mymaridae (Hymenoptera: Chalcidoidea), lected in 2010 at the Constenius Spring the first ever to be described from com- site along the Middle Fork of the Flathead pression fossils, were recently described River in northwestern Montana under the from the Kishenehn Formation (Huber and auspices of USFS Permit HUN281. It was Greenwalt 2011). collected from the middle sequence of Although Constenius et al. (1989) the Coal Creek member of the Kishenehn recorded two different beetle families, Formation, which has been estimated to be Scarabaeidae and Chrysomelidae, from 46.2 +/– 0.4 Ma by 40Ar/39Ar analysis and VOLUME 115, NUMBER 2 175

43.5 +/– 4.9 Ma by fission-track analysis (Constenius et al. 1989, Constenius 1996). An interesting feature of fossils re- covered from this formation is that their details are best seen when the shale is wetted. The compression fossil was there- fore immersed in 95% ethanol for exami- nation and photography. The dorsal habitus image was captured using a Canon EOS 7D attached to a Visionary Digital Imaging System (Visionary Digital™,Palmyra, VA). Images were then montaged and edited using Adobe PhotoshopÒ. The following measurements were re- corded as part of the description. Total length (TL) was measured from the an- terior margin of the head capsule to the apex of the abdomen, and total width (TW) was measured at the widest point across the elytra. Head length (HL) was measured at the midline from the anterior to posterior margin of the head capsule, and head width (HW) was measured at the widest point across the head capsule. Pronotal length (PL) was the length mea- sured at the midline from the anterior to posterior margin, and pronotal width (PW) Fig. 1. Ptenidium kishenehnicum Shockley and was measured at the widest point across Greenwalt, new species. Scale bar = 0.1 mm. the pronotum. Elytron length (EL) was the length measured from the anterior margin from amber. Its relative small size (0.65 to the apex along the suture, and elytron mm) distinguishes it from Microptilium width (EW) was measured at the widest gestautsi Dybas, Ptinella oligocoenica point across one elytron. Abdomen length Parsons, and Ptinella rovnoensis Polilov (AL) was measured at the midline from and Perkovsky, which are all 0.80–0.91 the anterior margin of abdominal ventrite mm in length. The large mesoscutellum 1 to the apex of abdominal ventrite 6, and and the shape and sculpturing of the pro- abdomen width (AW) was measured at notum readily separates P. kishenehnicum the widest point across the abdomen. from Micridium groehni Polilov and Perkovsky and Ptilium tertiarium Horion, RESULTS the only other species described from a Ptenidium kishenehnicum Shockley compression fossil. and Greenwalt, new species Description.—Overall body shape elon- gate oval, body widest across middle of (Figs. 1–2) elytra. TL = 0.65 mm, TW = 0.35 mm. Diagnosis.—This new species differs Body coloration reddish-brown, head dark markedly from the other five described brown, antennae lighter. Clypeus is dark species of fossil ptiliids, known mostly gray in appearance. 176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

expanded apically (possibly an artifact of compression), convex along anterior margin. Mouthparts directed ventrally and obscured dorsally by head capsule. Antennae 11-segmented with loose 3-segmented club. Scape and pedicel subequal in length, enlarged and barrel- shaped; antennomere 3 smaller; anten- nomeres 4–6 elongate and narrow, each subequal in length to pedicel but only 0.5X as wide; antennomeres 7–8 signif- icantly shorter and progressively wider than preceding segments, bead-like in ap- pearance; antennomeres 9–11 forming a loose 3-segmented club, as long as pre- ceding 4 segments combined; antennomeres 9–10 as long as wide; terminal anten- nomere more elongate, 1.5X as long as wide. Lengths of antennomeres (mm): 25.5, 21.3, 8.4, 21.3, 23.4, 14.9, 14.9, 19, 25.5, 31.9. No anterior whorls of setae visible on any of the segments. Thorax: Pronotum strongly convex and transverse, much wider than long; PL = 0.12 mm, PW = 0.27 mm. Lateral margins slightly arcuate, widest near middle; pro- Fig. 2. Ptenidium kishenehnicum Shockley and notum nearly as wide as the elytra. Ante- Greenwalt, new species. Labeled structures: bsw = rior margin nearly straight (but appearing basal strut of hind wing; cly = clypeus; e = eye; concave due to distortion), anterior angles = = = = el elytron; fov fovea; N1 pronotum; scl narrowly rounded, indistinct; posterior scutellum; sclg = scutellar groove; st2 = meso- sternum; st3 = metasternum; t = tergite; v = ventrite. margin nearly straight (but appearing Scale bar = 0.1 mm. convex due to distortion), posterior angles obtuse to subrectangular. Pronotum pos- teriorly bearing 4 faint punctiform struc- Head: Head broadly oval, 1.4X as tures, possibly foveae (difficult to discern broad as long, HL = 0.14 mm, HW = as they lie on top of anterolateral notches 0.20 mm. Partially obscured from above of mesosterna). Mesonotum with scutel- by pronotum such that the anterior an- lum large, acutely triangular posteriorly. gles approach the posterior margin of the Metanotum with dorsomedial portion of eyes. Antennal insertions fully exposed metascutum, scutellar groove and alary (difficult to see on the left), positioned ridges visible through the elytra, extend- medially on vertex near anteromedial ing to a point just beyond 1/3 length of margin of eye. Eyes prominent, coarsely elytra. Elytra elongate (EL = 0.41 mm, faceted. Fronto-clypeal suture conspic- EW = 0.18 mm) and complete, widest just uous due to discoloration of clypeus, anterior to midlength, narrowing api- evenly arcuate between the antennal in- cally, exposing part of tergite VI and py- sertions. Clypeus large and laterally lobed, gidium (likely an artifact of compression). VOLUME 115, NUMBER 2 177

Metathoracic wings visible under the appear convex rather than straight, as it elytra, membrane narrow, basal stalk with would have been in life. Although dor- a single strut, trichia faint but visible soventral compression introduced some posteromedially. artifacts into the fossil, it also made Legs: Legs entirely missing or buried observation of more ventral structures in the next layer down from the re- possible as the entire specimen was mainder of the body, making examina- compressed into the same focal plane. tion impossible. For example, the right antenna which Abdomen: Abdomen strongly com- bends medially at the 4th antennomere is pressed, appearing wider (due to flatten- obscured by the head and pronotum but ing of the abdominal pleura) and longer is still visible due to this phenomenon. (due to stretching of the intersegmental Similarly, some abdominal features, the membranes); AL = 0.25 mm, AW = struts and membrane of the hind wings 0.30 mm. Tergites and ventrites visible and the scutellar groove, are all visible through elytra, 7 visible tergites, 6 visible directly through the elytra and abdominal ventrites (posteriormost ventrites difficult tergites. Unfortunately, this “transparency” to discern). Pygidium distinct, posterior also makes it difficult to discern the two margin smooth, acute apically, apex bear- lateral pronotal foveae (already quite ing a small tuft of setae medially, no faint) because they lie directly above serrations or teeth apparent. Genitalic the anterolateral notches of the meso- structures not preserved. sternum, which are heavily sclerotized Material examined.—Holotype (Sex un- and plainly visible in the specimen. One known), labeled “Ptenidium kishenehnicum of the most conspicuous diagnostic fea- Shockley and Greenwalt. Holotype USNM tures of the family is the whorls of # 545816”. Deposited in NMNH. A setae present on the antennae. However, second specimen from an adjacent lo- poor preservation of setae and surface cality, Constenius Park, collected in 2012 sculpturing is a known artifact of com- was also examined, labeled “Ptenidium pression fossils so their absence in this kishenehnicum Shockley and Greenwalt. specimen is not particularly surprising. USNM # 553512”. Deposited in NMNH. Likewise, soft genitalic structures are Etymology.—The specific epithet is a often not preserved and are absent in Latinized adjective based on the geo- this fossil as well, so in the absence of logical formation where the fossil was secondary sexual characteristics it is discovered. impossible to determine the sex of the Comments.—The Kishenehn ptiliid specimen. specimen appears slightly distorted, most The other specimen found at an ad- likely due to dorsoventral compression jacent locality in 2012 is not as well immediately postmortem or during fossil- preserved as the holotype. Therefore, ization. Without a doubt the elytra and the we are reluctant to declare it a para- abdomen have been distorted from their type, despite it presenting a nearly antemortem position by this compression, identical habitus to the holotype. The thus making the abdomen much wider relatively poor condition of this sec- than it would have appeared naturally. ondary fossil makes it impossible to Similarly, the pronotum appears slightly definitively place as P. kishenehnicum, rotated with the anterior margin appear- but it is not unreasonable to assume that ing higher than the posterior margin, it is the same species, based on what making the anterior margin artificially features are visible. 178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

DISCUSSION for generic assignment, are unavailable. Carpenter (1992) first suggested a However, the elytra slightly shortened, Tertiary origin for the family, most likely pygidium hindmargin without a conspic- during the late , as they were well uous apical tooth, possible presence of known from Baltic amber. Ponomarenko pronotal fovea along hind margin, pro- (1995) also concluded that Ptiliidae orig- notum not constricted basally, hind an- inated in the late Eocene or early Oli- gles of pronotum not acute, and prothorax gocene, later revising his estimate for without median longitudinal depression the origin of the family to the mid to late suggest its appropriate assignment to (Ponomarenko 2002), an es- Ptenidium (Hall 2000, 2005). Prior to the present study, only five timate congruous with the discovery of fossil ptiliids had been described at the ancient ptiliids in Lebanese and Myanmar species level. The first to be described amber, dated to 125 and 95 Ma, respec- was a compression fossil (Statz and tively (Rasnitsyn and Ross 2000, Grimaldi Horion 1937). This particular fossil, Ptilium et al. 2002, Poinar and Poinar 2008). The tertiarium, consisted of both a part and presence of large numbers of ptiliids in 95 counterpart, which allowed for exami- Ma Myanmar amber suggests that past nation of both the ventral and dorsal abundance and diversity may be compa- surfaces. Assignment of this specimen to rable or even greater than that found to- the Ptilium was apparently based day. The ptiliids of the Eocene likely lived on the presence of two “tubercles” on the in much warmer and wetter environments posterior margin of the pygidium and than those that exist in the Nearctic today, the pronotum with a median groove and and the known Kishenehn fossils thus far two smaller adjacent transverse grooves. represent a subtropical/temperate fauna Polilov and Perkovsky (2004) dismissed (Constenius et al. 1989). this assignment and suggested that, since Ptenidium kishenehnicum Shockley Ptilium tertiarium dates from the Early and Greenwalt is a remarkably preserved Miocene (23.6–21.0 Ma) and could not specimen that can be readily identified as be distinguished from modern species, it belonging within the subfamily Ptiliinae, is probably an extant species. However, tribe Ptiliini. Members of this tribe are the lifespan of an insect species is thought generally recognized by the following to be 3 to 10 million years (Grimaldi and combination of characters: 11-segmented Engel 2005) and an age of only 20+ antennae, elytra complete or only slightly million years is not a scientifically valid shortened (last abdominal tergite only basis for assuming that P. tertiarium is an exposed), eyes normal, procoxal cavities extant species. open or coxae moderately separated by a Ptinella oligocoenica and Microptilium narrow prosternal process, and posterior geistautsi were both described from Baltic margin of the pygidium variable in form amber (Parsons 1939, Dybas 1961). The (but generally not armed apically) (Hall description of P. oligocoenica did not 2000). Unfortunately, as with many fossil include a basis for the assignment to the descriptions, its placement in the genus genus Ptinella and the assignment of Ptenidium cannot be considered defini- M. geistautsi appears to be based on a tive given that the ventral aspect of the comparison of drawings of the specimen insect is not visible and characters such as to an accompanying description of the the relative placement of the coxae and extant species M. pulchellum. The de- mesosternal processes, characters integral scription of M. geistautsi consists only of VOLUME 115, NUMBER 2 179 a detailed description of the antennae, the ptiliids remain undescribed or unassigned fossil’s color and presence of dorsal setae. generic or species names, evaluating those The two species described by Polilov and records becomes impossible without some Perkovsky (2004) were both identified explicit method for identifying specific as female. Ptinella rovnoensis retained a specimens and determining if those re- small spherical spermatheca, a remark- cords are distinct or duplicate. able example of preservation given that Although P. kishenehnicum is the first this fossil insect is only 910 mmlong. fossil of its family to be described from Ptinella rovnoensis lacks hind wings and the New World, specimens have been is thought to be a vestigial morph, a form recorded from both Dominican and known to exist within the genus Ptinella Mexican amber. Thus, we hope that its (Dybas 1978). Polilov and Perkovsky description will renew interest in de- (2004) also reported two poorly pre- scribing those specimens as well, with- served specimens identified to the genus out the constraints adopted by Polilov Ptenidium and five additional specimens and Perkovsky (2004). Increased input of Ptilium to which they did not assign into the comparative database of fossil species names as they were unable to ex- Ptiliidae is much needed and desirable amine the male genitalia and thus could for a greater understanding of the evolu- not distinguish them from recent species. tion of this unique and interesting family We would counter that inability to of miniscule beetles. distinguish new fossil ptiliids from ex- tant species should not, in and of itself, ACKNOWLEDGMENTS preclude their designation as new spe- We thank Karie Darrow (Dept. of cies. Although there are large numbers Entomology, National Museum of Nat- of fossil Ptiliidae reported from several ural History, Smithsonian Institution) for amber sites around the world ranging taking the high resolution habitus im- in age from 20 Ma to 120 Ma, only six ages. We also thank the two anonymous specimens of Ptiliidae have been officially reviewers whose comments and sugges- designated as new species. Dismissal of tions significantly improved the manu- Ptilium tertiarium as an extant species and script. This is contribution number 279 of the requirement by Polilov and Perkovsky the Evolution of Terrestrial Ecosystems (2004) of comparing genitalia to recent consortium at the National Museum of species are based on unrealistic criteria Natural History in Washington, D.C. that inhibit efforts to describe the many ptiliid fossils that exist. We do not mean Literature Cited to propose that, when feasible, such comparisons should not be made, but Bachofen-Echt, A. 1949. Der Bernstein und seine Einschlu¨sse. Springer-Verlag, Wien. the literature is rife with descriptions of 204 pp. fossil species that have never been com- Carpenter, F. M. 1992. Treatise on Invertebrate pared to extant species or are anatomically Paleontology. Part R, Arthropoda 4, Volume indistinguishable from extant species. 4. Superclass Hexapoda. Geological Society Similarly, specimens assigned the status of America, Boulder, CO. 655 pp. of “incertae sedis” or “species indeter- Constenius, K. N., M. R. Dawson, H. G. Pierce, R. C. Walter, and M. V. H. Wilson. 1989. Re- minate”, the only alternative to formal connaissance paleontologic study of the designation as new species, are rarely Kishenehn Formation, northwestern Montana fully described in the scientific litera- and southeastern British Columbia. Montana ture. Unfortunately, since so many fossil Geological Society 1989 Field Conference, 180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Montana Centennial. Geological Resources of Hall, W. E. 2005. Ptiliidae, pp. 251–261. In Montana 1: 189–203. Beutel, R. G. and R. A. B. Leschen, eds. Constenius, K. 1996. Late Paleogene extensional Handbook of Zoology, Volume 4. Arthropoda: collapse of the Cordilleran foreland fold and Insects: Part 38. Coleoptera, Beetles. Volume thrust belt. Geological Society of America 1: Morphology and Systematics. De Gruyter, Bulletin 108(1): 20–39. doi:10.1130/0016- Berlin/New York. 567 pp. 7606(1996)108<0020:LPECOT>2.3.CO;2 Handlirsch, A. 1925. Pala¨ontologie, pp. 117–306. Dybas, H. S. 1961. A new fossil feather-wing In C. W. M. Schro¨der, ed. Handbuch der beetle from Baltic amber (Coleoptera: Entomologie, Vol. 3. Gustav Fischer, Jena, Ptiliidae). Fieldiana. Zoology (Jena, Germany) Germany. 1201 pp. 44(1): 1–9. Helm, O. 1896. Beitra¨ge zur Kenntniss der Insecten Dybas, H. S. 1978. Polymorphism in featherw- des Bernsteins. Schriften der Naturforschenden ing beetles, with a revision of the genus Gesellschaft in Danzig N.F. 8: 220–231. Ptinelloides (Coleoptera: Ptiliidae). Annals Horridge, G. A. 1956. The flight of very small of the Entomological Society of America 71: insects. Nature 178: 1334–1335. doi:10.1038/ 695–714. 1781334a0 Erichson, W. F. 1845. Vol. 3. Naturgeschichte der Huber, J. T. and D. Greenwalt. 2011. Compres- Insecten Deutschlands. Erste Abtheilung, Cole- sion fossil Mymaridae (Hymenoptera) from optera. Verlag der Nicolaischen Buchhandlung, Kishenehn oil shales, with description of two Berlin. 320 pp. new genera and review of Tertiary amber Grebennikov, V. V. 2008. How small you can go: genera. ZooKeys 130: 473–494. doi:10.3897/ Factors limiting body miniaturization in winged zookeys.130.1717 insects with a review of the pantropical genus Klebs, R. 1910. U¨ ber Bernsteineinschlu¨sse im Discheramocephalus and description of six allgemeinen und die Coleopteren meiner new species of the smallest beetles (: Bernsteinsammlung. Schriften der Physikalisch- Coleoptera: Ptiliidae). European Journal of O¨ konomischen Gesellschaft zu Ko¨nigsberg Entomology 105: 313–328. 51(3): 217–242. Grebennikov, V. V. 2009. Discheramocephalini, Kulicka, R. and S. A. Slipi nski. 1996. A review of a new pantropical tribe of featherwing bee- the Coleoptera inclusions in the Baltic amber. tles (Coleoptera: Ptiliidae): description of Prace Muzeum Ziemi 44: 5–11. new taxa and phylogenetic analysis. System- Larsson, S. G. 1978. Vol. 1. Baltic Amber – a atic Entomology 34: 113–136. doi:10.1111/ palaeobiological study. Scandinavian Science j.1365-3113.2008.00444.x Press Ltd., Klampenborg, Denmark. 192 pp. Greenwalt, D. E., F. Marsh, and C. C. Labandeira. Matthews, J. V., Jr. 1977. Tertiary Coleoptera 2011. Preliminary Characterization of the En- fossils from the North American Arctic. Co- tomofauna of the Middle Eocene Kishenehn leopterists Bulletin 31(4): 297–308. Basin. Proceedings of the GSA Joint Rocky Matthews, J. V., Jr. and A. Telka. 1997. Insect Mountain/Cordilleran Sections meeting 43(4): fossils from the Yukon, pp. 911–962. In H. V. 13. Danks and J. A. Downes, eds. Insects of the Grimaldi,D.A.,M.S.Engel,andP.C.Nascimbene. Yukon. Biological Survey of Canada. Ter- 2002. Fossiliferous Cretaceous amber from restrial , Ottawa. 1034 pp. Myanmar (Burma): Its rediscovery, biotic diver- Parsons, C. T. 1939. A ptiliid beetle from Baltic sity, and paleontological significance. American amber in the Museum of Comparative Zoology. Museum Novitates 3361: 1–72. doi:10.1206/ Psyche 46: 62–64. doi:10.1155/1939/86451 0003-0082(2002)361<0001:FCAFMB>2.0. Poinar, G. O., Jr. 1992. Life in Amber. Stanford CO;2 University Press, Stanford, CA. 350 pp. Grimaldi, D. and M. S. Engel. 2005. Evolution of Poinar, G. O., Jr. and R. Poinar. 1999. The Amber the Insects. Cambridge University Press, New Forest: A Reconstruction of a Vanished World. York. 755 pp. Princeton University Press, Princeton. 239 pp. Hall, W. E. 2000. Ptiliidae Erichson, 1845, pp. Poinar, G. O., Jr. and R. Poinar. 2008. What 233–246. In Arnett, Jr., R. H., and M. C. Bugged the Dinosaurs? Insects, Disease, and Thomas, eds. American Beetles. Vol. 1. Arch- Death in the Cretaceous. Princeton Univer- ostemata, , , : sity Press, Princeton. 264 pp. . CRC Press, Boca Raton, FL. Polilov, A. A. 2005. Anatomy of the feather- 443 pp. winged beetles Acrotrichis montandoni and VOLUME 115, NUMBER 2 181

Ptilium myrmecophilum (Coleoptera, Ptiliidae). Rasnitsyn, A. P. and A. J. Ross. 2000. A pre- Entomological Review 85: 467–475. liminary list of families present in Polilov, A. A. and E. E. Perkovsky. 2004. New the Burmes amber collection at The Natural species of late Eocene feather-winged beetles History Museum, London. Bulletin of the (Coleoptera, Ptiliidae) from the Rovno and Natural History Museum. London (Geology) Baltic amber. Paleontological Journal 38(6): 56: 21–24. 664–668. Statz, G. and A. Horion. 1937. Ein fossiler Ponomarenko, A. G. 1995. The geological history Ptiliidenfund aus den mitteloligocanen Abla- of beetles, pp. 155–171. In J. Pakaluk and gerungen von Rott am Siebengebirge. Ento- S. A. Slipi nski, eds. Biology, Phylogeny, and mologische Blatter 38: 8–10. Classification of Coleoptera: Papers Celebrating Spahr, U. 1981. Systematischer Katalog der Bern- the 80th Birthday of Roy A. Crowson. Muzeum stein- und Kopal-Ka¨fer (Coleoptera). Stuttgarter i Instytut Zoologii PAN, Warszawa. 1092 pp. Beitrage zur Naturkunde. Serie B, Geologie und Ponomarenko, A. G. 2002. Superorder Scar- Palaontologie 80: 1–107. abaeidea Laicharting, 1781. Order Coleoptera Wu, R. J. C. 1996. Secrets of a Lost World: Do- Linne, 1758. The beetles, pp. 164–176. In A. P. minican Amber and its Inclusions. R. J. C. Rasnitsyn and D. L. J. Quicke, eds. History of Wu, Santo Domingo, Dominican Republic. insects. Kluwer, Dordrecht. 524 pp. 222 pp.