New ambrosia beetles (Coleoptera: Curculionidae: Platypodinae)

16 Pages·2015·4.79 MB·English

OrgDiversEvol
DOI10.1007/s13127-015-0213-y
ORIGINALARTICLE
New ambrosia beetles (Coleoptera: Curculionidae: Platypodinae)
from Miocene Mexican and Dominican ambers and their
paleobiogeographical implications
DavidPeris1&MónicaM.SolórzanoKraemer2&EnriquePeñalver3&XavierDelclòs1
Received:12February2015/Accepted:10April2015
#GesellschaftfürBiologischeSystematik2015
Abstract TwonewspeciesaredescribedfromMexicanam- resin producers. Based on the analysis of fossil and current
ber (15–20 Ma): Cenocephalus tenuis Peris and Solórzano distribution ofsuch plants, our hypothesisconsiders that the
Kraemer sp. nov. and Tesserocerus simojovelensis Peris and beetledispersionoccurredwithHymenaea,whichwaspossi-
Solórzano Kraemer sp. nov. Cenocephalus, originally de- blyitshostplant.
scribed as living in Central and South America and then as
fossilsfromEarlytoMiddleMioceneamber,isnotedasmor- Keywords Platypodinae .Cenocephalus .Mitosoma .
phologicallyindistinguishablefromMitosoma,andoriginally Hymenaea .Paleobiogeography.Madagascar
described as endemic from Madagascar. Thus, we consider
that a close taxonomic relationship exists, even if they are
notthesamegenus.Newevidenceofthespeciesalreadyde- Introduction
scribed in Platypodinae (Tesserocerini) from Mexican and
Dominican ambers (15–20 Ma) and the differences between Ambrosiabeetlesisthecommonnameforthebeetlesubfam-
those species are discussed, complementing the original de- ily Platypodinae, which contains more than 1,400 species
scriptions. The paleobiogeography of Cenocephalus and (Kuschel et al. 2000; Oberprieler et al. 2007; Kirkendall
Mitosomaisanalyzed,whichstronglysupportsthehypothesis etal.2015)thatcultivatefungiintunnelsexcavatedinwood.
ofcolonizationfromAfrotropicalMadagascartoAmericapri- AlthoughPlatypodinaewastraditionallyconsideredaninde-
ortoEarlytoMiddleMiocene (15–20Ma) via sea currents. pendent family within Curculionoidea (Wood 1986, 1993;
Hymenaea was interpreted as the Mexican and Dominican WoodandBright1992),itisnowconsideredwithinthefamily
Curculionidae (Kuschel et al. 2000; Oberprieler et al. 2007;
Alonso-ZarazagaandLyal2009;Bouchardetal.2011;Jordal
* DavidPeris
etal.2011,2014).ExtensiveanalysesbyKuscheletal.(2000)
[email protected];[email protected]
andFarrelletal.(2001)suggestedthatplatypodinesshouldbe
MónicaM.SolórzanoKraemer
better classed as a group of scolytines (Jordal and Cognato
[email protected]
2012); however, other researchers, namely Marvaldi (1997)
EnriquePeñalver
andMarvaldietal.(2002,2008),concludedtheyweremore
[email protected]
suitably described as being independent subfamilies within
XavierDelclòs
Curculionidae.
[email protected]
Jordal et al. (2011) carried out an extensive phylogenetic
1 Departamentd’Estratigrafia,PaleontologiaiGeociènciesMarines;
studyonthemostprominentwood-boringtaxa,withinwhich
andInstitutdeRecercadelaBiodiversitat(IRBio),Facultatde
they suggested that Scolytinae and Platypodinae were sister
Geologia,UniversitatdeBarcelona(UB),MartíiFranquéss/n,
08028Barcelona,Spain lineages within Curculionidae and that early diversification
2 SenckenbergForschungsinstitutundNaturmuseum, madethemasadvancedweevils.Morerecently,withtheanal-
Senckenberganlage25,60325FrankfurtamMain,Germany ysisofhigher-levelphylogeneticrelationshipsinweevils,the
3 MuseoGeominero,InstitutoGeológicoyMinerodeEspaña,Ríos polyphylyofwood-boringlineagesremainsopen.However,it
Rosas23,28003Madrid,Spain ispossiblethatthePlatypodinaemaynotbecloselyrelatedto
D.Perisetal.
Scolytinae,and sothe positionof Platypodinae remains am- their occurrence has not yet been presented (Weitschat and
biguous(Guillettetal.2014;Hulcretal.2015),andthedebate Wichard2002),andnodescriptionofthesefindingshasbeen
isongoing(Bright2014;Jordaletal.2014). madetodate(Alekseev2013;seeCognato2015).However,
Ambrosia beetles embedded in Mexican and Dominican theauthorsofthepresentstudywillbeanalyzingasampleof
Early to Middle Miocene ambers are extremely diverse (see platypodineinBalticamberinthenearfuture.
Schedl1962;Schawaller1981;BrightandPoinar1994;Davis Inthisstudy,adescriptionoftwonewplatypodinespecies
andEngel2007)andoftenincludeseveralspecimensnotyet inMexicanamberispresentedaswellasnewevidencerelat-
described.PlatypodineinvolvementinMioceneresinproduc- ing to several species already described from Mexican and
tion, together with scolytines, has been suggested by some Dominican ambers (Table 1). Based on the analysis of this
authors (Bright and Poinar 1994; Grimaldi and Engel 2005; newevidence,sometaxonomicchangesrelatedtoMitosoma
McKellar et al. 2011). The Mexican and Dominican ambers (Malagasy distribution) and Cenocephalus (Neotropical dis-
were formed by resiniferous angiosperms (Hymenaea spp.) tribution) will be required following Schedl’s work (1972).
andtheirscolytineandplatypodinefaunaprobablyusedthese The morphological similitude, at generic level, of
plant species as hosts, which would have damaged the trees Cenocephalus with living Mitosoma is discussed.
and induced the resin production (Poinar and Poinar 1999; Additionally, we will analyze a hypothesis that attempts to
CognatoandGrimaldi2009). explain the current distribution of Mitosoma and
Hymenaea (Fabaceae (=Leguminosae): Detarieae) is a Cenocephalus. This hypothesis takes into account the fossil
resin-producing angiosperm tree. It currently comprises 14 record and ecology of living representatives, as well as the
species, which are generally well distributed in the tropical analysisofdatarelatedto the Mioceneresin-producing trees
and subtropical forests of Central America, the West Indies, intheNeotropics.
andmostofSouthAmerica.Hymenaeaverrucosaisaunique
species that is found in East Africa and Madagascar and is
consideredthe mostprimitive species ofthe genus (Leeand Materialsandmethods
Langenheim1975;Langenheim2003;Fougère-Danezanetal.
2010).Nevertheless, a completephylogeneticand molecular Samples with bioinclusions described in this paper are from
studyofthewholespecieshasnotbeencarriedoutuntilnow. Dominican and Mexican ambers. They are housed at three
The species Hymenaea mexicana and H. allendis have differentinstitutions:BEPGM-RD^Dominicanambercollec-
been found in Mexican amber (Poinar and Brown 2002; tion of the Laboratori de Paleontologia, Department of
Calvillo-Canadelletal.2010)whileH.proterahasbeenfound Estratigrafia, Paleontologia i Geociències Marines,
inDominicanamber(Poinar1991).Thesefindingsmayhave Universitat de Barcelona (Barcelona, Spain); BMCAM^
beenmadepossiblethroughtheanalysisofflowersandleaves Museu de la Ciència Ambre, CosmoCaixa Barcelona
preservedasbioinclusions.Thesefossilspeciesarethoughtto (Barcelona, Spain); and BMx^ Staatliches Museum für
betheproducersofbothMexicanandDominicanfossilresins, Naturkunde, Schloss Rosenstein (SMNS) (Stuttgart,
respectively. Furthermore, a morphological study of Germany).
H.proteraandH.mexicanaindicatedacloserelationshipwith The Dominican amber pieces are fromLa Toca mine, lo-
the Afrotropical species H. verrucosa (Poinar and Brown cated at Cordillera Septentrional in northern Dominican
2002).Giventhelackofamolecularphylogeneticstudythat Republic (Fig. 1a). The Dominican amber was reviewed by
includesallHymenaeaspeciesandtheabsenceofHymenaea several authors, and following discussions regarding the age
spp.inthemacro-remainsandpollenfossilrecordsofAfrica andoriginofthespecimen,itwasconcludedthatitdatedback
and South America before the Middle Miocene, there are to the late Early to early Middle Miocene (15–20 Ma)
varyingwaystointerprettheevolutionaryhistoryofthistree (Grimaldi 1994; Iturralde-Vinent and MacPhee 1996;
genus(seebelow). Iturralde-Vinent 2001; Grimaldi and Engel 2005; Penney
DavisandEngel(2007)andKirejtshuketal.(2014)carried 2010). The Mexican amber pieces are from Simojovel de
out studies in which they summarized all the fossil species AllendeinthestateofChiapas,Mexico(Fig.1b).Thisamber
known in Platypodinae. They reported a total of 14 species is considered to be of Early to Middle Miocene in age (15–
in one fossil and three extant genera, plus a highly dubious 20Ma)andwasthereforecorrelatedwiththeDominicanam-
assignment to an extant species. The first fossil containing berdeposits(SolórzanoKraemer2007,2010).
ambrosiabeetlewasdescribedfromMioceneMexicanamber Dominican pieceshavethe following accession numbers:
asCenocephalusbySchedl(1962),whiletheearliestevidence EPGM-RD-0106, EPGM-RD-0107, EPGM-RD-0108,
wasreportedfromCretaceousMyanmaramberandthenfrom EPGM-RD-0110, EPGM-RD-0111, MCAM-0034, MCAM-
Eocene Baltic amber (Larsson 1978; Poinar 1992; Cognato 0530,MCAM-0590,andMCAM-0623.Theaccessionnum-
comm. pers.). Platypodinae were occasionally listed among bersoftheMexicanpiecesareasfollows:Mx-213,Mx-214,
the beetles fromBaltic amber;however, reliableevidenceof Mx-238, Mx-275, Mx-347, Mx-354, and Mx-379. All
Paleobiogeographicalimplicationsofnewambrosiabeetles
ectionoftheNaturkunde, Depository 3 3 1,2 1,2 1 2 swNaaemsrcepilmeaslbsewoneeereamncbdueStdiadlvneeddrspitneoilnias2ht0erd0a,0n)sa.pnadrethnet EepPoGxMy-RreDsinpie(sceees
ollür Specimens were examined under a Leica MS5 stereomi-
cf
erm croscope and an Olympus BX41 compound microscope.
piecesstudiedfromthe(1)Dominicanambarcelona,Spain),and(3)StaatlichesMuseu Newsamples Mx-214,Mx-275 Mx-213,Mx-347 EPGM-RD-0106,EPGM-RD-0107,EPGM-RD-0108,MCAM-0530EPGM-RD-0106,MCAM-0034,MCAM-0623EPGM-RD-0110 MCAM-0590 wheretheoriginalspeciesweredescribed GeslwPmaTuorraheecaerfbonWartniwdleatneteosatergatmsa2ariehalunecemstopshtruLteaphgheemdceodeenhiEtdmcttoedtlhaodegeafueomIsrrsMitatclkzihoeplrneeoei1nhpgsytw0tshtLsti0etswhfoeh0r1enioeLe.Ac0srmaDaeesluaiaoorcMtrtnaWnahefawdtkosSwioseorni5Ctonna-eamdZgrornsueesrtadis(eoelrwi1ClrlmnayieD9meogzeo9iramracemea3argrt)bwoemLiao,cirsefnsarcbaXio.edeocunsae6aptdcZce.uDothP.hsLApFiePseynelCph,lagdteaoluim4caxttsi2eo(oeeic2d0gnsnaa0r.gsocmaw0uampte9rmihhteri)hceas--.
ListofalreadydescribedspeciesofCenocephalusfromMexicanorDominicanambersthathavebeenfoundandanalyzedintheoridePaleontologiadelaUniversitatdeBarcelona(Barcelona,Spain),(2)MuseudelaCiènciaAmbre,CosmoCaixaBarcelona(BRosenstein(Stuttgart,Germany) SpeciesOriginaldescription –nC.succinicaptusSchedl,19621962C.succinicaptusSchedl:Figures14andText-Fig.1–nC.hurdiSchedl,19621962C.hurdiSchedl:Figures58andText-Fig.1––canC.rhinoceroide(Schwaller,1981)1981MitosomarhinoceroideSchwaller:FiguresAbb.13and1420 ––canC.exquisitusBrightandPoinar,19941994C.exquisitusBrightandPoinar:Figures56and1819 –canC.quasiexquisitusDavisandEngel,20072007C.quasiexquisitusDavisandEngel:Figures15––canC.spinatusBrightandPoinar,19941994C.spinatusBrightandPoinar:Figures910and2223 ^B^aterialsandMethodstofindinformationrelatedtothegeologyoftheseambersandReferencestofindthecompletereferences SONGAGlnsaDPaC(‘itasCtssesyrlpireeleeicyesooldswnSSFSTCFErHAPDTtpsentencnhmlemtehuynuuauiaretsteoiholdegrgedyiamlresssodepbpban:baomleoclaugp:maopneeoefryroheCrdttiiernrgahCer,yyly’attrn:ceoayoiseypmyfoif.ocdodpp)et:neCaTplltlr:ispe2l.ehoalpaoecenaim-eieeieorhrCeclatTset:(:hah:gsoocnys:ty:ri:ti)CeMMaasiuyhCcplamyal:pfPpFenyeruaMelrper:tlpPeuseaouaereclxeex:nposa:yo,cmeSrrllhsrdudl--xdhCmsaycateCca2,3opla,te-ahtaubbpicouee1o5yts3leLroelallyMhoyurMonoresn3un4pci7nnsiicsvoaturn,e,nisospwto9yicueytgfeendmidoittnasdc,iploxsClPnxhhiraoecayipilStaeieeaioieeihMgosPnahiecehpnEncdtldonnararlegaoioullsahociLmleoscioooearpnlswnyy;solnannaiiohpaccmui,muascdeScl)odhipitaamivsua1i.trlrneadmeehollshctiyueisbm7Iiieninoeoa,suou,ttafitsoi5rltyyyiedntlo1nmcsc1iboaebLelhid8eroekes8.naae8Snloey,maawrp,dpean6nn,u8r,agomrtl.rr1r5iddeaCpe16ilpdoaetfss8wóIlbthc9oeryt,icddteee0rleoim1ietit1szleectic2rrostatse4shsppuasra8hapein,sunelfioos4tld.feee1rshbedosse0oocsCioS8.eeiilomiAnmKtty0cMhmmnbooeLla2rarsruriealavNyy,ypaASetsnceiatuiitSpaahimwnpmybiincrrensidtyeelicev(,ocihetSwaardehtti1jairhhlmorteastor8uiyseemp,alv,b6ryte.lsedtssfta5ilnghhrnaaaldcroaogtmmrofnoadhenororivltdnteeeeeesfr-,.,
Table1LaboratSchloss Amber Mexica Mexica Domini Domini Domini Domini BSeeM evxetxenwdiitnhgaipnatoiraofshaocrutt,eascpuitneifporromcepssr,oftraucseioonfsdienctloivtihtyeivnetreyrsctroiane-
D.Perisetal.
Fig. 1 Miocene amber mines; a La Toca, Cordillera Septentrional, northern Dominican Republic; b Simojovel de Allende, Chiapas, Mexico;
arrowsindicatepitheads
3,protibiaearmedwithtwotransverserugaeandouterfaceof byshortspiniformprotrusions,apairofacutespiniformprotru-
metatibiaearmedbyfourtransverserugae. sions into the face of the declivity, interstriae 3; apical margin
Description: Female. Body length 3.64 without mandibles, extendingintoashort,acuteprocess.
maximum width 0.59 at the elytral apex, 6.2 times longer than
wide.Bodycompletelycollapsed.Pubescencepresentdorsallyin
Table2 Taxaandauthornamedalongthetext(withtheexceptionof
thehead(longhairs)andelytra(shorthairs),pronotumglabrous.
thealreadynamedtaxainTable1)
Headdistinctlylongerthanwide,approximatelytwotimes
longerthaneyediameter,slightlyconstrictedbehindtheeyes, Plant
marked dorsally. Eyes lateral, small, round, easily visible in Hymenaea Linnaeus 1753
dorsalview,veryfinelyfaceted.Fronsaswideashead,strong- H.allendis Calvillo-Canadell, 2010
Cevallos-FerrizandRico-Arce
ly concave transversely, surface with long, straight setae,
H.courbaril Linnaeus 1753
which are curved onperiphery. Limit between the frons and
H.mexicana PoinarandBrown 2002
therostrumunidentifiable.Antennaeinsertionbasal,antennae
H.protera Poinar 1991
length very short, not extending to cephalic insertion. Scape
H.verrucosa (Gaertner)Oliver 1791
straight, only slightly curved at the insertion with the head,
extending until the fore margin of the eyes; funicle four-
Animal
segmented; very little pedicel inserted at the scape apex, as
Acanthotomicus Blandford 1894
wide as long; antennomere 3 conical, narrower at base and
A.hymenaeae Eggers 1933
wider at apex, as long as the apical diameter; antennomeres
Batrachorhina Chevrolat 1842
4 and 5 stronglytransverse; club compressed, almostround.
Chaetastus Numberg 1953
Mandibles large, with one large tooth at the apex and one
Cladoctonus Strohmeyer 1911
toothattheapicalthirdofthebitingedge,smoothandconvex
C.ruber BrightandPoinar 1994
onupperface.Submentumseparatedoneachsidefrommar-
Coccotrypes Eichhoff 1878
ginoforalfossabydeepcleft.
Coecephalophonus (Schedl) 1965
Pronotum2.1timeslongerthanwide,wideratapicalfifth,
Cryptocarenus Eggers 1937
narrower towards the base; pronotal disk smooth. Posterior
Electroborus Cognato 2013
margin of prothorax strongly procurved in pleural area.
Hypothenemus Westwood 1836
Scutellumsmall,slender,apicallypointed.Procoxaecontigu-
Mitosomaantiquus BrightandPoinar 1994
ous,precoxalpieceonprosternumobtuselypointed.
Mitosomabiconicus BrightandPoinar 1994
Elytrasubparallel,becomingwiderapically,baseofelytraas
Nesanopliumpuberulum (FleutiauxandSallé) 1889
wide as base of pronotum. Elytra 2.9 times longer than wide
Platypicerus (Nunberg) 1953
(measuredfromthebasetothedeclinationandthewider,apical
Scolytogenes Eichhoff 1878
portion); 1.5 times longer than pronotum. Elytral disk shallow
Tesserocerusprimus BrightandPoinar 1994
punctured, deeply striate, total number of striae and interstriae
T.retusus Guérin-Méneville 1838
notdiscernible.Basesofinterstriae3and5transverselyrugose;
Wendilgarda Keyserling 1886
interstriae 3 carinate along the basal third, interstriae 5
subcarinate close to the base. Elytral declivity beginning near
Differences between plants and animals are presented here, ordered
apicalfourth, convex,lateralmarginevenlyarcuate and armed alphabetically
Paleobiogeographicalimplicationsofnewambrosiabeetles
Fig.2 C.tenuissp.nov.from
MioceneamberfromSimojovel
deAllende,Chiapas,Mexico;
aholotypeMx-379,female,
illustrationandcameralucida
drawingofthedorsalhabitus;
bholotype,detailedcameralucida
drawingoftheelytraldeclivity;
callotypeMx-354,male,detailed
illustrationoftheheadandthe
antennae;dparatypeMx-213,
detailedillustrationoftheelytral
declivity
Mesosternum with mesepisternum convex, unarmed. Theuniquecharacterthatdifferentiatesthisgenusfromthe
Metasternum long, with distinct femoral impression; Magalasy Mitosoma is the current distribution, which is a
metacoxaecontiguous.Legslong;femorawide,compressed; relatively weak piece of evidence when describing fossils
protibiaewithtwotransverserugaeandalongweaklyhooked (seeBDiscussion^).C.tenuissp.nov.hasseveralsimilarities
innermucro;outerfaceofmetatibiaearmedbyfourtransverse withtheAfricangenusChaetastus,buttheydifferinthean-
rugae.Tarsislender,longerthantibia;tarsomere1aslongas tennaetypeandelytraldisk(seealsoBDiscussion^abouttax-
thefollowingtarsomerescombined;tarsomeres2and3equal onomic remarks). These characters were highlighted by
in length; tarsomere 4 the shortest, 0.5 times the length of Browne(1971),whodescribedthemasessentialindifferen-
tarsomere 3, tarsomere 5 as long as the previous three com- tiating Chaetastus from Mitosoma and Cenocephalus.
bined,slender,withlong,slenderclaws. However,severalothercharactersinC.tenuissp.nov.warrant
Male.Ofsimilarsize,proportions,andgeneralsculpture.It amorecorrectassignmentofthenewspeciesinCenocephalus
differs from the female in frons feeble depressed, glabrous, (although see also BDiscussion^ about taxonomic remarks);
and elytral declivity with spiniform protusions on the these include dimensions and proportions of the body parts,
interstriae3morestronglydeveloped. which lack an apparent transverse band of fine pores along
Comments:ThespecimensfitwithinCenocephalusbythe basalportionofthepronotaldisk,numberoftransverserugae
posterior margin of prothorax strongly procurved in pleural atthepro-and metatibiaeouter surface,positionand sizeof
area,submentumseparatedoneachsidefrommarginoforal spines along the declivity face and margins, and American
fossa by deep cleft, procoxae contiguous, scutellum small, distribution.
slender, pointed, antennal funicle four-segmented (counting C.tenuissp.nov.isnotasstoutasmembersofthisgenus
the pedicel), eye with circular profile in lateral view, tend to be, and the pronotum is slender; however, the set of
mesepisternum convex, unarmed, precoxal piece on charactersareconcordantwithCenocephalus.Thefossilspe-
prosternum obtusely pointed, second segment on antennal cies Cenocephalus exquisitus, Cenocephalus antiquus, and
funicle longer than wide, outer face of metatibiae armed by Cenocephalus spinatus also have a slender pronotum.
fourtransverserugae,elytraldeclivityconvex,baseofdecliv- Although C. tenuis sp. nov. has a similar spine distribution
ity armed by spiniform protrusions, and Central American alongthedeclivitytothefossilC.biconicus,thenewspecies
distribution(Browne1971;Schedl1972;Wood1993). is more slender, with the declivity margin armed with short
D.Perisetal.
spiniform protrusions and face of declivity convex; timeslongerthanwide,almostflat,slightlyvisibleindorsalview,
C. biconicus is more robust, with the declivity margin un- subtriangular in outline, anterior margin entire; posterior border
armedanditsfaceobliquelytruncated. delimitedbysomelongandstoutsetae,assimilarlyoccuralong
mandible insertions. Vertex coarsely punctured, punctures two
Genus:TesserocerusSaunders,1837 timeslongerthanwide.Fronsslightlyconcavetransversely,sur-
Typespecies:PlatypusinsignisSaunders,1837 face with relatively long, fine, erected setae. Antennal insertion
TesserocerussimojovelensisPerisandSolórzanoKraemer basal,closertomandiblesthantotheeyes;antennaeshortextend-
sp.nov. ingtocephalicinsertion;scapelarge,slender,extendinguntilthe
Figure3 fore margin of the eyes, completely covered by long and stout
Etymology:SpecificnamesimojovelensisafterSimojovel setae,beinglongertowardapex;funiclefour-segmented;pedicel
deAllende(Mexico),localitywherenewspecieswasfound. inserted at the pedicel apex, rounded in shape; antennomere 3
Holotype: Mx-238, Miocene amber from Simojovel de conical,narroweratbaseandwiderapically,aslongastheapical
Allende, Chiapas, Mexico; housed at the SMNS (Stuttgart, diameter; antennomeres 4 and 5 strongly transverse; club com-
Germany). The holotype is a complete specimen, the entire pressed,almostround.Mandiblesarelarge,withonelargetooth
habitusvisibleandwellpreserved.Itisafemalebyitsfrons at the apex and one tooth at the apical third of the biting edge,
flattenedwithsurfacecoveredbylongsetaeandroundedbase smoothandconvexonupperface.Submentumseparatedoneach
ofthedeclivity,somethingexplanatebutwithoutprotrusions. sidefrommarginoforalfossabydeepcleft.
Diagnosis:Female.Body5.5timeslongerthanwide;head Pronotum1.8timeslongerthanwide,narrowertowardsthe
distinctlylongerthanwide, about two times longer thaneye base;pronotaldiskofroughappearance,completelycovered
length; scape, mandible insertion, and prosternum strongly by rounded punctures; pronotal lateral margins thickened,
pubescent, with long and stout hairs; elytral declivity with with a set of long and stout setae along the margins.
three spiniform protrusions on anterior border; protrusions Scutellumverysmall,apicallypointed.Prosternumlong,cov-
becoming sequentially shorter from the first to the third; ered by short and stout setae, precoxal piece on prosternum
metasternumlong,impressednearmetacoxaforreceptionof acutelypointed.Procoxaecontiguous.
femur, anterior margin of impression armed by one pointed Elytra subparallel, becoming wider apically, base of elytra
spine. slightly wider than base of pronotum. Elytra 2.9 times longer
Description:Bodylength5.35,maximumwidth0.97atthe thanwide;1.7timeslongerthanpronotum.Elytraldiskstrong-
elytral apex, 5.5 times longer than wide. Pubescence present ly punctured, deeply striate; interstriae 3 carinate along the
along the entire body, very conspicuous on the head, frons, basal third, interstriae 5 remarked with small tubercles close
scape,mandibles,andprosternum;elytraldiskalmostglabrous. tothebase.Elytraldeclivitybeginningnearapicalfourth,steep,
Head distinctly longer than wide, about two times longer with three spiniform protrusions on anterior margin of elytral
thaneyelength.Eyeslateral,elongate,big,approximately1.5 declivityandarisingfromelytralstriae3,5,and7;protrusions
becomesequentiallyshorterfromthefirsttothethird,although
the first and the second are much bigger than the third.
Declivityfacecoveredbyshortandfinehairsandapicalmar-
ginsextendingintoashort,roundedlobeineachelytron.
Metasternumlong,impressednearmetacoxaforreception
offemur,anteriormarginofimpressionarmedbyonepointed
spine; metacoxae contiguous. Legs short, robust; femora
wide, compressed; profemora wider than the rest; protibiae
armed withtwo transverserugae on the posterior face and a
long weakly hooked inner mucro; becoming wider from
protibiaetometatibiae;outerfaceofmetatibiaearmedbythree
transverserugae.Tarsistout,longerthantibiaeinallthelegs;
tarsomere 1 as long as tarsomeres 4 and 5 combined,
protarsomeres 2 and 3 equal in length, protarsomere 4 the
shortest, 0.5 times the length of tarsomere 3, tarsomere 5
slightlylongerthanthepreviousthreecombined,slender,with
long,slenderclaws.
Comments: The specimen fits withinTesserocerusby the
posterior margin of prothorax strongly procurved in pleural
Fig.3 T.simojovelensissp.nov.fromMioceneamberfromSimojovelde
area,submentumseparatedoneachsidefrommarginoforal
Allende, Chiapas, Mexico; holotype Mx-238, female, illustration and
cameralucidadrawingofthedorsalhabitus fossa by deep cleft, procoxae contiguous, scutellum small,
Paleobiogeographicalimplicationsofnewambrosiabeetles
slender, pointed, antennal funicle four-segmented (counting
thepedicel),eyeelongate,1.5timeslongerthanwide,anterior
marginentire,pronotum1.8timeslongerthanwide,precoxal
piece on prosternum acutely pointed, and pedicel attached
nearitsapex(Wood1993).AllTesserocerusspp.arebetween
3.0and4.5timeslongerthanwide,includingtheuniquefossil
species (Wood 1993; Bright and Poinar 1994), while
T. simojovelensis sp. nov. is 5.5 times longer than wide.
Except for this morphological feature, we have decided to
keepitinTesserocerusbecause allthecharactersinthe new
species are consistentwiththose for thisgenus inthe key to
worldplatypodinegenerafromWood(1993).
T. simojovelensis sp. nov. is clearly different from
Tesserocerus primus, which is currently a unique fossil spe-
cies ofthis genus,described from Dominican amber (Bright
andPoinar1994).Thenewspeciesislarger,withtworugaeon
the posterior face of the protibiae and with three spiniform
protrusions on anterior margin of elytral declivity, while
T. primus is smaller, with one ruga on the posterior face of
protibiae,andfourspiniformprotrusionsontheelytraldecliv-
ity, also with different proportions between them.
T. simojovelensis sp. nov. is similar to Tesserocerus retusus
innumberandpositionofspiniformprotrusions,butthefirst
pairofprotrusionsinthenewspeciesislongerthanthesecond
pair,whileinT.retususallthespinesareofsimilarlength.
SpeciesalreadyknownfromMexicanandDominican
ambers
Fig. 4 Diverse illustrations of two species of Cenocephalus from
Miocene amber from Simojovel de Allende, Chiapas, Mexico;
Several species already described from Mexican and
a C.succinicaptus, Mx-275,dorsal illustration ofthe elytral declivity;
Dominicanamberswereobservedintheamberpiecesstudied. b detailed camera lucida drawing of the elytral declivity; c C. hurdi,
Thedifficultyindifferentiatingbetweensomeofthesespecies Mx-347,lateralhabitus
usingthepublisheddatajustifiesthefollowingreview,which
includes comments on certain aspects of the species (see illustratesmoreclearlythemaledeclivityofC.succinicaptus.
Table1fordetails). BrightandPoinar(1994)notedthattheholotypeofthisspe-
cies is a completely distorted and broken specimen and that
MexicanspeciesofPlatypodinae theallotype(male)isalsodistortedexceptfortheelytralapex;
the new specimens found herein will be very helpful in the
CenocephalussuccinicaptusSchedl,1962 identificationofthespecies.Weknowthatbothspecimensare
InFig.4a–b,theoverallhabitusofthenewambrosiabee- malesduetothewell-developedspiniformprotrusionsalong
tles Mx-214 and Mx-275 is indicative of C. succinicaptus the elytral declivity margins and interstriae 3, the feebly de-
based on its original description and the figures 1–4 in pressed frons, and the lack of notable pubescence (Schedl
Schedl (1962). However, the original description was also 1962).ThedeclivitymargininMx-275waspartiallypolished,
accompanied by the Text-Fig. 1, which contains an error in andtheillustrationisbasedonlyinpartononeelytron.Mx-
thefigurelegend:themaledeclivityillustrationspresentedby 275 was found in syninclusion with a possible
Schedlcorresponding toC.succinicaptus(left) andC.hurdi Ceratopogonidae (Diptera) specimen. Mx-214 was found in
(right)werereferencedonthecontraryinthelegend,accord- syninclusionwithotherprobablePlatypodinaespecimens,but
ingtotheoriginaldescription.Takingintoaccountthismod- its preservation is not sufficient to enable us to identify this
ification, the original illustration for the male declivity in specimeninanymoredetail.
C.succinicaptus(Schedl1962)isconsistentwithFig.4bhere-
in,whichillustratesthemaledeclivityofC.succinicaptus. C.hurdiSchedl,1962
TheoriginallinedrawinginSchedl(1962)islackingdetail In Fig. 4c, there are two specimens consistent with the
and differs in some ways from the picture. Figure 4b herein description and illustration of the declivity that Schedl
D.Perisetal.
(1962)describedinC.hurdi.Mx-213andMx-347areprob- thepresenceofmycangia(KnízekandBeaver2004).Females
ablymalesduetotheirflatfrons(Schedl1962).Mx-213was of C. exquisitus can be differentiated from the male by their
foundinsyninclusionwithC.tenuissp.nov.andonepossible longer and stouter bodies, frons broadly flattened, covered
Anthribidae specimen (Coleoptera: Curculionoidea).
Unfortunately, the new material cannot be compared with
theholotypebecauseitislost;BrightandPoinar(1994)were
unable to locate the specimen described by Schedl (1962)
during their search in the Museum of Paleontology of the
University of California and the Naturhistorisches Museum
(Wien). Furthermore, the authors of the present study were
unable to locate the holotype of C. hurdi in the
NaturhistorischesMuseumcollection.Noparatypesweredes-
ignatedintheoriginaldescription.
DominicanspeciesofPlatypodinae
C.exquisitusBrightandPoinar,1994
ShowninFig.5a–c,therearetwospecimens(onemaleand
one female, both in EPGM-RD-0106) which are consistent
with the general description of C. exquisitus (Bright and
Poinar 1994). The male has a particular set of spiniform pro-
trusionsalongelytraldeclivitythatisdescribedbyBrightand
Poinar(1994)andlaterbyDavisandEngel(2007).Theorig-
inal illustration for the male declivity in Bright and Poinar
(1994)hasamistake:Thelengthofthespiniformprotrusions
on the anterior margin of the elytral declivity in the dorsal
aspect does not reflect the similar proportion as the same
spiniformprotrusionsinthelateralaspect(Figures5and6in
BrightandPoinar1994).This,combinedwiththeconclusions
ofDavisandEngel(2007),leadsustobelievethatthesecond
pair of spiniform protrusions on the anterior margin of the
elytraldeclivityshouldbeshorterthanthefirstpair.Indeed,this
characteristic is typically used to differentiate between
C.exquisitusandC.quasiexquisitus.Assuch,thepresentstudy
offersnewillustrationsofmaleelytraldeclivity(Fig.5a–b).
Althoughthespeciesisonlydescribedbasedonthemale
form, this study located a possible female specimen in
syninclusion with the male. Due to its proximity with the
male, it has been tentatively interpreted as a female of
C.exquisitus. Sexualdimorphismisa verycommoncharac-
teristic in nearly all Platypodinae, mainly in structures that
include the frons, elytral declivity, abdominal ventrites, and
„
Fig. 5 Diverse illustrations of four species of Cenocephalus from
Miocene amber from La Toca, Cordillera Septentrional, Dominican
Republic; a C. exquisitus, MCAM-0623, illustration of the dorsal
habitus; b EPGM-RD-0106, male, detailed camera lucida drawing of
the elytral declivity in lateral view; c female, detailed camera lucida
drawing of the elytral declivity in dorsal view; d C. rhinoceroide,
EPGM-RD-0106, illustration of the habitus of three females and one
male; e male from the samepiece, illustrationof the dorsalhabitus; f
C. quasiexquisitus, EPGM-RD-0110, male, illustration of the dorsal
habitus; g C. spinatus, MCAM-0590, male, illustration of the dorsal
habitus
Paleobiogeographicalimplicationsofnewambrosiabeetles
withlongerecthairs,andelytraldeclinationconvex,withtwo entire male. MCAM-0530 contains one female of
poorly developed spiniform protrusions in each elytron C. rhinoceroide in syninclusion with one Curculionoidea
(Fig.5c). (Coleoptera), one Diptera, one Psocoptera, one
InadditiontothetwospecimensofC.exquisitusinEPGM- Hymenoptera, one Trichoptera, one Arachnida, and one
RD-0106, there were initially seven specimens of Hymenaeaflower.
C. rhinoceroide, one Scolytinae (Cladoctonus ruber), and This species was defined originally within the genus
oneAcari.Followingpreparationoftheamber,onlytwospec- MitosomabySchwaller(1981).Ithasalreadybeendescribed
imensofC.exquisitusandfivespecimensofC.rhinoceroide and carefully illustrated, so it is unnecessary to give a new
remained.Thefactthatbothspecieswerefoundinthesame illustration herein. The existing description notes the differ-
pieceofamberconfirmstheircoexistenceinthepast. ences between males and females, which is also seen in the
The male specimens of C. exquisitus resemble males of new pieces of Dominican amber. Although the frons of the
C. rhinoceroide, C. quasiexquisitus (both from Dominican females in EPGM-RD-0106 and EPGM-RD-0107 are cov-
amber), and C. succinicaptus (from Mexican amber). ered by hairs in a more compact fashion than those of both
However, C. exquisitus differs from C. rhinoceroide by the the female in MCAM-0530 and the original illustration
longerspiniformprotrusionsonthedeclivitymarginandde- (Schawaller 1981), all other characters are consistent with
clivityfaceandbythesecondpairofspiniformprotrusionson the original female description. Thus, this character was not
the anterior margin of the elytral declivity, which is shorter considered sufficient to divide them into two species, as it
thanthefirstpair.Incontrast,inC.rhinoceroide,thespiniform couldbeapreservationaleffect.
protrusions are shorter and the two first pairs of spiniform C.rhinoceroideisverysimilartoC.exquisitus,but,given
protrusionsontheanteriormarginoftheelytraldeclivityare the features commented on in the previous species and the
equallylong. comparisonofbothspeciesinEPGM-RD-0106(Fig.5aand
The male specimens of C. exquisitus differ from 5e,elytraldeclinationinbothspecies),itispossibletodiffer-
C.quasiexquisitusfirstlybythesecondpairofspiniformpro- entiatethem.
trusionsontheanteriormarginoftheelytraldeclivity,whichis
shorterthanthefirstpair,andsecondlybytheprotrusionsat C.quasiexquisitusDavisandEngel,2007
thebaseofthedeclivity,whichareshorterandsituatedfurther Shown in Fig. 5f, the species was initially described by
apart. C. quasiexquisitus differs from the second pair of DavisandEngel(2007)fromanisolatedfossilspecimenfrom
spiniform protrusions on the anterior margin of the elytral Dominican amber, who noted its resemblance with
declivity as they are slightly longer than the first pair, and C.exquisitus.However,somecommentsintheirpaperallow
theprotrusionsatthebaseofthedeclivityareverylongand forthepossibleseparationofthetwospecies(seecomments
aresituatedclosertogether.MalesofC.exquisitusdifferfrom ofC.exquisitushereinandinDavisandEngel2007).EPGM-
C.succinicaptusbythemuchlongerspineonthelowerouter RD-0110 contains one entire male, which can be clearly
apicalmarginofthedeclivityandbythedifferentconfigura- alignedwiththedescriptionofC.quasiexquisitus,andapos-
tionofthespiniformprotrusionsonthelateralmarginofthe teriorhalfofanotherspecimen.Theremainsofthelatterare
elytral apex (Schedl 1962; Schawaller 1981; Bright and alsoconsistentwiththeelytraldeclivityofC.quasiexquisitus;
Poinar1994;DavisandEngel2007). however, it is impossible to make a definitive taxonomical
As with the amber piece described above, MCAM-0623 determination. In the same piece, there are syninclusions of
also contains two males and one female of C. exquisitus as three Acari: two specimens associated with its metasternum
syninclusions. By contrast, in MCAM-0034, there are 23 and one free specimen, which is very close to the
specimensofC.exquisitus,whicharemainlymales,although metasternum.
thereare possiblysomefemalesvisible.Inthis lastcase, the
piece also contains seven specimens of the scolytine C.spinatusBrightandPoinar,1994
Cladoctonus ruber, which is very commonly seen as Shown in Fig. 5g, one isolated male (MCAM-0590) is
syninclusionwithplatypodines(BrightandPoinar1994). presentinacompletelytransparentpieceofamber.Thisspe-
ciesisclearlydifferenttothespeciespreviouslydiscusseddue
C.rhinoceroide(Schwaller,1981)WoodandBright1992 toitselytraldeclivityinmales,whichisarmedbyapeculiar
Shown in Fig. 5d–e, there are four new pieces of amber setoflargespiniformprotrusions.
containing specimens of this species; however, EPGM-RD-
0106andEPGM-RD-0107constitutedthesamepieceofam-
ber prior to preparation. EPGM-RD-0106 contains only one Unnamedsample
entiremale,threeentirefemales,andoneveryspoiledfemale
lackingtheposteriorhalfofthebody.EPGM-RD-0107con- EPGM-RD-0111: The fossil is the posterior half of a
tainsonemaleandonefemale.EPGM-RD-0108containsone Platypodinae. By the elytral declivity, it is possible to infer
D.Perisetal.
itsresemblancetomalesofC.quasiexquisitus,butitspreser- separate them morphologically by certain features on the
vationprecludesanydefinitedetermination. elytral declivity treated as Busual^ and by the difference in
the geographic distribution of the living species, i.e.,
Mitosoma is endemic to Madagascar and Cenocephalus to
Discussion CentralandSouthAmerica.
C.succinicaptus,afossilfromMexicanamber,hasavery
TaxonomicremarksonCenocephalusandMitosoma obliqueelytraldeclivity(Schedl1962).Inseveralfossils de-
scribedasCenocephalusfromDominicanamber,thebaseof
Aftertheauthor’sobservationofextantplatypodinesfromthe themaledeclivityhaslargespiniformprotrusions(Brightand
MNHN(Paris,France)andNHMW(Wien,Austria),themor- Poinar1994).ThelasttwocharactersweretreatedasBusual^
phological confusion between the genera Mitosoma and for Mitosoma by Wood (1993). Based onthe observed vari-
Cenocephalusincurrententomologicalcollectionswasnoted. ability, the unique argument to separate both genera is
Chapuis (1865) described both genera. However, after biogeographic.
some new descriptions of living and fossil species, Schedl Although it is not the objective of this study to review
(1972) decided to synonymize Cenocephalus (Neotropical), the current genera, it is necessary to consider some close
Chaetastus (Afrotropical: Africa), Platypicerus, and relationships between Cenocephalus and Mitosoma. The
Coecephalophonus (Afrotropical: Madagascar) with use of current distribution as a definitive character to sep-
Mitosoma (Afrotropical: Madagascar). The reasons for this arate two genera is not suitable and even less so when
are based on previous research by Browne (1971), who considering fossil forms. Thus, consistent with the
reviewedthegenusChaetastusandnotedtheclearlydefined approach by Browne (1971) and Schedl (1972), it is nec-
group comprised Mitosoma, Cenocephalus, and Chaetastus, essary to critically assess the relationship between these
and a possible similarity to Coecephalophonus. The author genera, particularly the group comprising Cenocephalus,
wasonlyabletodistinguishChaetastusfromtheothersimilar Mitosoma, and Chaetastus (clade with a possible category
generabythesecondsegmentoftheantennalfunicleasitwas of subtribe Mitosomatina sensu Jordal comm. pers.). In
longerthanitwaswide(thirdantennomere),ageneralsculp- the present work, the synonymy between Mitosoma and
tureoftheelytradrawninhiswork,anditsAfricandistribu- Cenocephalus is not formally established. This is firstly
tion(Browne1971).Schedl(1972)didnotconsiderthechar- because it is not the focus of the study and secondly
actersdescribedbyBrowne(1971)sufficienttomaintainthe because this would require an analysis of their phylogeny
diversity and synonymized all the genera cited previously and morphological features (including fossil and living
withMitosoma. species). However, it is considered that they are closely
This proposal was maintained for 30 years and related through a common ancestor and are morphologi-
Cenocephalus rhinoceroide was the first fossil platypodine cally indistinguishable from each other.
described from Dominican amber in Mitosoma (Schawaller
1981).ThesituationchangedwhenWoodandBright(1992) Hymenaeadispersal
consideredMitosoma, Cenocephalus, and Chaetastusasdif-
ferent genera in their catalogue with no explanations about Hymenaeacurrentlyincludes14species.H.verrucosa,from
theirgenericnames.Later,Wood(1993)specifiedtherestora- Madagascar and Eastern Africa, is considered the most an-
tionofallthreenamestofullgenericstatusandproposedakey cient representative of the genus. The other 13 species have
to differentiate the diverse genera in Platypodinae Neotropical distribution (Lee and Langenheim 1975;
(Platypodidae sensu Wood 1993). In that key, Chaetastus Langenheim 2003; Fougère-Danezan et al. 2010). There are
wasclearlyseparatefromtheothertwogenerabythefollow- varioushypothesesthatattempttoexplainthehistoricdistri-
ing:posterioronethirdofpronotumwithatransversebandof bution of the genus, which are proposed and discussed as
mycetangiaporesinsteadofmycetangiaporesnotdiscernible; follows:
anterior face of metatibiae with only one transverse ruga in-
stead of anterior face of metatibiae armed by three or more 1. Poinar (Poinar 1991) and Poinar and Brown (2002) ar-
transverserugae;AfricaninsteadofMalagasyorCentraland guedthattherecentdistributionofHymenaeaspp.isthe
SouthAmericandistribution;andsomeadditionallessvisible resultofavicarianceprocess.
characters.Wood(1993)didnotconsideroneofthecharacters
usedbyBrowne(1971) important indifferentiatingbetween BasedonthepresenceoffossilcaesalpinioidpolleninLate
Chaetastusanditstworelatedgenera.Thischaracteristicwas CretaceousofSouth America (Muller 1981),theysuggested
thethirdsegmentoftheantenna,whichwaslongerthanitwas that Hymenaea existed in Gondwana during the Cretaceous
wide.Furthermore,althoughWood(op.cit.)decidedtosepa- periodbeforetheriftingofSouthAmericaandAfrica(atabout
rate Mitosoma and Cenocephalus again, he could only 100 Ma). However, vicariance in Hymenaea is difficult to