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Head Structures of Priacma Serrata Leconte (Coleptera, Archostemata

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Head Structures of Priacma Serrata Leconte (Coleptera, Archostemata JOURNALOFMORPHOLOGY252:298–314(2002) HeadStructuresofPriacmaserrataLeconte(Coleptera, Archostemata)InferredFromX-rayTomography ThomasHo¨rnschemeyer,1*RolfG.Beutel,2 andFreekPasop3 1Institutfu¨rZoologie&Anthropologie,AbteilungMorphologie&Systematik, D-37073Go¨ttingen,Germany 2Institutfu¨rspezielleZoologieundEvolutionsbiologie,FSUJena,Jena,Germany 3ThermoScientificB.V.,Breda,TheNetherlands ABSTRACTInternalandexternalfeaturesofthehead shapedprotuberancesonthedorsalsideoftheheadis ofPriacmaserratawerestudiedwithX-raymicrotomog- consideredanautapomorphyofCupedidae.Thegalea raphyandwithhistologicalmethods.Thecomparisonof withanarrowstalkandaroundandpubescentdistal bothtechniquesshowsthatX-raytomographyisaprom- galeomereisanotherautapomorphyofthisfamily.Ithas isingnewtechniquefortheinvestigationofinsectanat- probablyevolvedasanadaptationtopollen-feeding.The omy.ThestillsomewhatcoarseresolutionoftheX-ray shapeofthemandibleofCupedidaeisplesiomorphiccom- dataiscompensatedforbyadvantageslikethenonde- paredtowhatisfoundinadultsofOmmatidae.Thever- structiveandartifact-freedataacquisition.TheheadofP. ticalarrangementofapicalteethisanautapomorphyof serrataandotheradultsofArchostemataischaracterized thelatterfamily.Thelateralinsertionoftheantennain bymanyderivedfeatures.MuscularfeaturesofPriacma, PriacmaisagroundplanfeatureofCupedidae.Thedorsal especiallymusclesofthelabiumandpharynx,differ shiftisasynapomorphyofallothercupedidgenera.A stronglyfromwhatisfoundinothergroupsofColeoptera. cladisticanalysisofcharactersoftheheadandadditional Severalcharacterstatesareconsideredasautapomor- dataresultedinthefollowingbranchingpattern:((Crow- phiesofArchostemata:scale-likesurfacestructures,con- soniellaϩ(OmmaϩTetraphalerus))ϩ(Micromalthusϩ strictedneck,stronglyreducedtentorium,andtheplate- (Priacmaϩ(Paracupesϩ(CupesϩTenomergaϩProlixo- like,enlargedprementum.Thescalesprovideaprotecting cupesϩRhipsideigmaϩDistocupesϩ(AdinolepisϩAs- surfacepatternandmayhaveevolvedwithamoreex- cioplaga)))))).J.Morphol.252:298–314,2002. posedlifestyle.Theenlargedprementumformsalid, ©2002Wiley-Liss,Inc. whichclosesthemouthandcoverstheligulawhenitis pulledbackbycontractionoftheunusuallystrong KEYWORDS:micro-CT;head;skeletomuscularsystem; submento-prementalmuscle.Thepresenceoffourcone- X-raytomography;Coleoptera;Archostemata;phylogeny PriacmaserratabelongstoCupedidae,oneoffouror KnowledgeofArchostemataisverylimited,de- fiveextantfamiliesoftheArchostemata.Theearliest spitetheobviousphylogeneticimportanceofthe fossilspeciesofCupedidaeareapproximately250mil- group.Availabledataoninternalmorphologyand lionyearsold(Permian:Sakmarian;Labandeira, biologyarerestrictedtoveryfewrepresentatives.As 1994).Cupedidaeistheoldestextantbeetlefamily farasadultmorphologyisconcerned,Priacmaser- accordingtothepresentlyavailablefossilrecordand rataisthebestinvestigatedspeciesbecausethe ArchostematatheoldestsuborderofColeoptera(Law- malescaneasilybecollectedinlargenumbers(Ed- rence,1999).TheextantArchostemataretainanum- wards,1951;Atkins,1957).Besidesthethoracic berofprimitivecharacters,themostobviousofwhich morphology(Baehr,1975),themalegenitaliawere isthereticulationoftheelytra.Thisfeaturewasal- examinedbyEdwards(1953a,b)andtheflightis readypresentintheoldestknownbeetlesfrom280- alsowelldocumented(Atkins,1958).Nevertheless, million-year-oldsedimentsofthelowerPermian (Ponomarenko,1969;Beutel,1997).Otherprobably ancestralcharactersincludethelargelyexposedpro- pleuraandprothoracictrochantin,therelativelylarge Contractgrantsponsor:DAAD;Contractgrantnumber:D/96/ mesothoraxwithatransversesutureontheventrite, 05967.* andthepresenceofseveralthoracicmusclesthatare absentfromallotherbeetles(Baehr,1975;Beuteland *Correspondenceto:ThomasHo¨rnschemeyer,Institutfu¨rZoologie Haas,2000).Nevertheless,therearealsoderivedchar- &Anthropologie,Abt.Morphologie&Systematik,BerlinerStr.28, acters,especiallyinthemorphologyofthelarvae(Beu- D-37073Go¨ttingen,Germany.E-mail:[email protected] tel,1997)andinthemalegenitalarmatures(Ed- PublishedonlinexxMonth2002in wards,1953a,b;Neboiss,1984),thatallowusto WileyInterScience(www.interscience.wiley.com) identifytheArchostemataasamonophyletictaxon. DOI:10.1002/jmor.1107 ©2002WILEY-LISS,INC. HEAD STRUCTURES OF PRIACMA SERRATA 299 even for this species, considerable information is tion of slices during the cutting process or the ne- wanting. Females of P. serrata are very rare and cessity to align the slices correctly. This makes it their genitalia are yet undescribed. Only the first- relatively easy to reconstruct the X-ray data into a instar larva is known (Ross and Pothecary, 1970). three-dimensional model of the structures of inter- From the publications of Edwards (1951, 1953a,b), est. Further development in the achievable spatial Atkins (1957, 1958), Roth and Pothecary (1970), and resolution now makes it possible to use X-ray tomog- from the personal experience of the authors one can raphy for the study of insect morphology, as is dem- summarize the knowledge of the biology of P. serrata onstrated in the following. as follows: The species has an activity period of approximately 4 weeks each year. This period usu- ally starts in spring or early summer with calm, MATERIALS AND METHODS sunny weather and temperatures above 15°C. Dur- Materials: Fixed Specimens ing these 4 weeks Priacma males are attracted to Priacma serrata LeConte, 1861, from Montana, solutions of sodium hypochlorite and sodium chlor- USA, collected by one of the authors (T.H.). ate and can be collected in large numbers. It is unknown whether these 4 weeks are the pairing season of the species and adult beetles are around Dried Specimens From Museum Collections for a longer time of the year or whether adult beetles Adinolepis youanga (Neboiss, 1960), from Australia, are only present during this period. There seems to Australian National Insect Collection, Austra- be an affinity of Priacma to conifers, because wher- lia. ever Priacma was collected there were conifers (of- Distocupes varians (Lea, 1902), from Australia, Aus- ten Douglas fir) in the vicinity. A direct proof of such tralian National Insect Collection, Australia. an association is still missing. There is no informa- Omma stanleyi (Newman, 1839), from Australia, tion available about the food sources of larvae and Australian National Insect Collection, Austra- adults. lia. Head morphology is also largely unknown, espe- Priacma serrata (LeConte, 1861), from USA, Mu- cially internal features. It was demonstrated in ear- seum of Comparative Zoology, Cambridge, MA, lier studies that head structures are not only of USA; Los Angeles County Natural History Mu- great importance in terms of coordination of the seum, USA; Field Museum of Natural History, central nervous system, sensory perception, and Chicago, IL, USA. food processing, but also provide much phylogenetic Prolixocupes latreillei (Solier, 1894), from Chile, information due to their complexity (e.g., Beutel, Staatliches Museum fu¨ r Tierkunde, Dresden, 1997). Therefore, the main goal of this contribution Germany. is to provide anatomical data on the head of Pri- Prolixocupes lobiceps (LeConte, 1874), from Los An- acma, to compare the findings with characters of geles County Natural History Museum, USA. other archostematan representatives, to offer func- Rhipsideigma cretacaeotincta (Kolbe, 1897), from tional interpretations, and to give a preliminary Tanzania, Africa, Field Museum of Natural His- phylogenetic interpretation. tory, Chicago, IL, USA. To investigate head structures of Priacma serrata, Rhipsideigma lugubris (Fairmaire, 1895), from an innovative technology—X-ray tomography—was Madagascar, Field Museum of Natural History, applied for the first time in insect morphology and Chicago, IL, USA. compared with other techniques. X-ray tomography, Rhipsideigma raffrayi (Faimaire, 1884), from Mada- besides its application for medical purposes, has so gascar, Zoologisches Forschungsinstitut und far mainly been used to investigate bone and tooth Museum Alexander Koenig, Bonn, Germany. structures in vertebrates (Ru¨ egsegger et al., 1996; Tetraphalerus bruchi (Heller, 1913), from Argentina, Bjørndal et al., 1999; Barou et al., 1999). The rea- holotype, Staatliches Museum fu¨r Tierkunde, sons for these restrictions are that mineralized tis- Dresden, Germany. sues give distinctly better X-ray images than soft tissues and that the spatial resolution of the avail- Some of the dry specimens were dissected (Omma able X-ray equipment did not allow the investigation stanleyi, Distocupes varians). Others were used to of smaller organisms. Due to the usage of phase compare the external morphology of head capsule contrast and improvements in the spatial resolution and its appendages. this method could also be used for studies of soft tissues (Beckmann et al., 1999) and very small Methods structures like the inner ear of the mouse (Van Spaendonck et al., 2000). Important advantages of All specimens of Priacma serrata were collected X-ray tomography over other methods are the pos- directly into alcoholic Bouin’s solution (ϭDuboscq- sibility of a nondestructive investigation of internal Brasil) (Romeis, 1989: 97). After 2 days they were structures and the avoidance of preparation arti- transferred to 80% ethanol. The specimens for sec- facts. There are no such problems as the deforma- tioning were softened in Diaphanol (Romeis, 1968: 300 T. HO¨ RNSCHEMEYER ET AL. source and the CCD detector. As the source produces a cone beam of X-rays, magnification is achieved by moving the object towards the X-ray source (Fig. 1). After
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