Digitization improves dissemination, discovery, and long-term preservation in fossil collections.

Phil Barden Department of Biological Sciences New Jersey Institute of Technology Discovery

Dissemination

Improving Preservation Leptomyrmex neotropicus from ~20 million year old Dominican amber (Barden, Boudinot, & Lucky 2017) 2003 Zootaxa 1623: 47–53 (2007) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2007 · Magnolia Press ISSN 1175-5334 (online edition)

First fossil Micropholcommatidae (Araneae), imaged in Eocene Paris amber 2007 using X-Ray Computed Tomography

DAVID PENNEY1,4, MANUEL DIERICK2, VEERLE CNUDDE2, BERT MASSCHAELE2, JELLE VLASSENBROECK2, LUC VAN HOOREBEKE2 & PATRIC JACOBS3

1Earth, Atmospheric and Environmental Sciences, The University of Manchester, M13 9PL, UK. E-mail: [email protected] 2Department of Subatomic and Radiation Physics, Ghent University, Proeftuinstraat 86, 9000 Ghent, Belgium 3Department of Geology and Soil Science, Ghent University, Krijgslaan 281/S8, 9000 Ghent, Belgium 4Corresponding author

Abstract

We apply Very-High-Resolution X-Ray Computed Tomography (VHR-CT) to a minute fossil spider (~1 mm long) from Eocene amber of the Paris Basin, France. We demonstrate that the newly described and species of Micropholcom- matidae, Cenotextricella simoni, retains excellent details of the somatic and male pedipalpal morphology that allows unqualified comparison with extant species. Thus, in addition to calibrating the tree of life, such fossils can now be incor- porated into cladistic matrices and their resultant phylogenies. This is the first fossil record of the family Micropholcom- matidae, extending the known geological range of the family by 53 million years (lowermost Eocene) and that of the symphytognathoid clade by approximately 5–9 million years.

Key words: Cenozoic, new species, palaeontology, spider, Symphytognathoidea, Textricellinae

FIGURES 11–12. Photographs of Cenotextricella simoni sp. nov. (male holotype, MNHN PA 327) using traditional light microscopy. (1) dorsal view; (2) ventral view. See Fig. 5 for scale. Introduction

Discussion

The Micropholcommatidae are currently pSomeoorly deli mneiteontd, butolo cang beis cotsnsi cdreriedti ctoi zecons paist ofla teontwo subfolami-ogy in their belief that preservation of fossils is inadequate in terms of lies: the Micropholcommatinae and Textricethllein aeimpor (Platnickta &nt For tastxoer 1986nomic). Based c onhar soacmatteric mso rtphoheylog yreveal (see Grimaldi et al. 2000). For example, Wanless (1984) con- and the morphology of the male pedipalp, Cenotextricella appears to belong in the Textricellinae. This is the first fossil record of the Micropholcommatisidade,er exedtendi ang s mthe aklnownl nu gmeolbogierca lo rfan Bge alof ttheic famamilyb byer 5sp3 iders in his review of the salticid spider subfamily Spartaeinae, but million years (to the lowermost Eocene). Theempha known sigeolzedogica thl agee ofta txheonomi superfamcily iSymphnferiytoorgnatityhoi odefa many amber specimens as originally preserved. The novel applica- can also be extended by approximately 5–9 million years, from the previously oldest recorded specimens of Anapidae in Baltic amber (Wunderlich 2004)ti.on All etoxtant fos micrsophilsolc ommatiof exid tasxtai n(33g specimieas ginin eighgt gteechnera;nologies, designed for non-palaeontological purposes, is yielding Platnick 2007) are tiny (<1.5 mm), so the discovery of the new fossil species is a remarkable find and repre- sents the only northern hemisphere record ofre them faamrkilya. ble results (Grimaldi et al. 2000; Polcyn et al. 2002; Alonso et al. 2004; Siveter et al. 2004; Ascaso et The taxonomic status of the Textricelallin.ae 200 has r4;ecei Tveda faf oconreasideruabl ete degral. ee2 0of0 a6ttent), ioanl tihn orecuentgh in some cases, these methods destroy the specimen (Siveter et al. decades. Initially erected by Hickman (1945), the family Textricellidae (along with the Micropholcomma- tidae) was synonymized with the Symphyt2004ognathid).ae Vbye Fory-rstHer igh(1959)-Res. Howeolveruti, mononophyl X-Ray of tyhe ComputSym- ed Tomography (VHR-CT) (Grimaldi et al. 2000; Polcyn et al. phytognathidae was questioned by several2002 worker;s, Alonwho conssoide etred altha.t 2most00 cha4; raHcteersn udseedr itoc kjuxstif ety it al. 2006; Dierick et al. 2007) provides a non-destructive, minimum were simplifications, reductions or losses that could have been the consequence of miniaturization (Schütt FIGURES 1–10. VHR-CT scans of Cenotextricella simoni sp. nov. (male holotype, MNHN PA 327). (1) dorsal view; 2003). Micropholcommatidae and Textriceprllideparae werateio thusn mremoveethodd fr ofom rsy imanonymygin witgh mSymphytinuteog- morphol(2) venogitral vicalew; det(3) anaitelrisor, viincew;l (u4d) poinstgerior int vierew;n (al5) lmateroalrph viewo; lo(6)gy late,ra anl sedcti ogenened vieraw; tes(7) right pedipalp dor- nathidae (Forster & Platnick 1977) and placed in the superfamily Palpimanoidea (Forster & Platnick 1984). sal view; (8) right pedipalp posterior view; (9) left pedipalp retrolateral view; (10) right pedipalp prolateral view. Abbre- Shortly thereafter Micropholcommatidae an3-d dimenTextricellsidaeional were reconsconidersedt rsynuconymoustions thawith tea cchan ot herbe sectiviatoneions: AdL anE, adnt eviriorewed lateral e yefr;o AmME m, antueltrioiplr med iaanngl eye;es c, co(esndusctor;enti cyal, clyymbiu permim; ds, dttoirsnalg abdominal scutum; (Platnick & Forster 1986) (the former namedi takgiitnga lpr ‘ecedidenssce)ec tonion the’ b asiofs tthahet the sp geenuscim Teeutnon wielliat hiBrin- thee, a emmbbolues;r )f.e , Hfemeurre; mwt/t,e m aeppltatarsyus /tarVHsusR jo-inCt; Tpa ,t poa tellaa s; piPLE,de pro sterbioioir nclatelruals eyioe;n PME, in ~po5ster3ior median eye; st, gnoli could not be assigned to either taxon. Several workers did not accept the assignment to Palpimanoidea sternum; th, tibial hook; ti, tibia; vs, ventral abdominal scutum. (e.g. Schütt 2003), and a more recent cladismiltic anlialyon-sis yeasuggesr-otedld that Eo somcene palep imParanoidis taamberxa belong (Nelin the et al. 1999; Nel 2004) and show that the specimen is preserved with superfamily Araneoidea (Griswold et al. 2005). The Micropholcommatidae were recently synonymized with 50 · Zootaxa 1623 © 2007 Magnolia Press PENNEY ET AL. Anapidae (Schütt 2003), but this conclusionli wasfe -lnoikt acce eptfiedd elby iPltyat,nic rkeve (2007)al ingdue to suthe flifmiticiedent numbe der oftail of palpal morphology for phylogenetic comparison with extant genera included in the analysis and is not fotallowedxa, herthuse for dethe msamoenst reasonra.t ing that such fossils are not as taxonomically sub-standard as previously thought.

FIRST FOSSIL MICROPHOLCOMMATIDAE Zootaxa 1623 © 2007 Magnolia Press · 51 Accepted by M. Rix: 26 Sept. 2007; published: 26 Oct. 2007 47 Haidomyrmex scimitarus in ~99 million year old (Barden & Grimaldi 2012) Dlussky 1996 Haidomyrmex cerberus Dlussky 1996

Perrichot et al.Haidomyrmodes 2008 mammuthus Perrichot et al. 2008

Antweb & Vincent Perrichot Haidomyrmex scimitarus in ~99 million year old Burmese amber (Barden & Grimaldi 2012) Haidomyrmex scimitarus in ~99 million year old Burmese amber (Barden & Grimaldi 2012) Haidomyrmex scimitarus in ~99 million year old Burmese amber (Barden & Grimaldi 2012) Linguamyrmex vladi in ~99 million year old Burmese amber (Barden et al. 2017) Courtesy Ed Stanley

What kind of data does CT scanning generate?

Brighter areas result from sample absorption meaning less photons Darker areas result from sample transmittance meaning more photons Linguamyrmex vladi in ~99 million year old Burmese amber (Barden, Herhold, & Grimaldi 2017) Linguamyrmex vladi in ~99 million year old Burmese amber (Barden, Herhold, & Grimaldi 2017) Haidomyrmex scimitarus from~99 million year old Burmese amber (Barden, Towbin, & Heiss in prep) Haidomyrmex scimitarus from~99 million year old Burmese amber (Barden, Towbin, & Heiss in prep) Haidomyrmex scimitarus from~99 million year old Burmese amber (Barden, Towbin, & Heiss in prep) Haidomyrmex scimitarus from~99 million year old Burmese amber (Barden, Towbin, & Alaric in prep) Haidomyrmex scimitarus from~99 million year old Burmese amber (Barden, Towbin, & Alaric in prep) Zigrasimecia tonsora from~99 million year old Burmese amber (Barden & Grimaldi 2013) Zigrasimecia tonsora from~99 million year old Burmese amber (Barden & Grimaldi unpublished) Zigrasimecia tonsora from~99 million year old Burmese amber (Barden & Grimaldi unpublished) Leptomyrmex neotropicus from ~20 million year old Dominican amber (Barden, Boudinot, & Lucky 2017) Leptomyrmex neotropicus from ~20 million year old Dominican amber (Barden, Boudinot, & Lucky 2017) Leptomyrmex neotropicus from ~20 million year old Dominican amber (Barden, Boudinot, & Lucky In Press) (Barden & Grimaldi 2016) (Barden & Grimaldi 2016)

Models: NJIT industrial design students Jackson Fordham, Oliver Budd, Victor Nzegwu Models: NJIT industrial design students Jackson Fordham, Oliver Budd, Victor Nzegwu

[email protected] Many thanks to BardenLab.org @haidomyrmex

Andrea Lucky / University of Florida Jessica Ware / Rutgers University – Newark Bo Wang / Chinese Academy of Sciences John LaPolla / Towson University Brendon Boudinot / University of California: Davis Mark Riccio / Cornell University & General Electric Christine Sosiak / New Jersey Institute of Technology Manpreet Kohli / Rutgers University – Newark David Grimaldi / American Museum of Natural History Martina Decker / New Jersey Institute of Technology Henry Towbin / Columbia University Michael Engel / University of Kansas Hukam Singh / Birbal Sahni Institute of Palaeobotany Susan Perkins / American Museum of Natural History James Carpenter / American Museum of Natural History Vincent Perrichot / Université de Rennes

Imaging, Animation Martina Decker / New Jersey Institute of Technology Jackson Fordham, Oliver Budd, Victor Nzegwu / New Jersey Institute of Technology Alaric Heiss / American Museum of Natural History Hollister Herhold / American Museum of Natural History James Thostenson / Duke University Morgan Hill / American Museum of Natural History Steve Thurston / American Museum of Natural History How are X-rays used in imaging?

X-ray Source

Detector or Film Worker in ~52 million year old Cambay amber (Barden & Grimaldi unpublished) Worker in ~52 million year old Cambay amber (Barden & Grimaldi unpublished) Worker in ~52 million year old Cambay amber (Barden & Grimaldi unpublished)