The first fossil cyphophthalmid harvestman from Baltic amber

2011·7.1 MB·English

by  Jason A Dunlop

1
ArachnologischeMitteilungen40:47-54 Nuremberg,January201
The first fossil cyphophthalmid harvestman from Baltic amber
Jason A.Dunlop & Plamen G.Mitov
doi:10.5431/aramit4006
Abstractivefirstfossilcyphophthalmidharvestman(Opiliones:Cyphophthalmi)fromPalaeogene(Eocene)Baltic
amberis described.This isonlythethird fossil example ofthis basal harvestman lineage;the others beingfrom
theprobablyslightlyyoungerBitterfeldamberandthemucholder,earlyCretaceous,Myanmar(Burmese)amber.
Although incompleteand lacking mostoftheappendages,the new Balticamberfossil can be identified asafe-
male.Thesomaticcharacterspreserved,especiallyspiraclemorphologyandthecoxo-genitalregion,allowittobe
assignedwithsomeconfidencetotheextantgenusSiroLatreille,1796(Sironidae).Thisfossilisformallydescribed
hereasSirobalticussp.nov. It resembles modern North American Sirospecies morethan modern European ones,
andcan bedistinguished principallyon its relativelylargesizeandtheoutlineformofthebody.
Keywords:Cyphophthalmi,Eocene,newspecies,Opiliones,palaeontology,Siro,systematics
There are currently29validspecies offossilharvest- thesmallsizeandcrypticlifestyleofthesesuperficially
men (Arachnida: Opiliones) in the literature; see mite-like animals, which significantly reduces their
DUNLOP (2007) foranoldersummary. Since2007 a potential for fossilisation. While conceding the in-
furtherspeciesbelongingtoCyphophthalmihasbeen herentimprecisioninageestimates,thephylogenyof
describedfromearlyCretaceous(ca. 100Ma)Myan-
mar (Burmese) amber (POINAR2008). Long-legged
andrathermodern-lookingEupnoimaterial,partially
assignable to the extant family Sclerosomatidae, is
nowknownfrom the midJurassic (ca. 160-180Ma)
ofDaohugou, Inner Mongolia, China (HUANG et
al. 2009). Both records are highly significant given
the relative rarityofbothAsian fossil arachnids and
MesozoicArachnidaingeneral.DUNLOP&cMlTOV
(2009) described further specimens belonging to
EupnoiandDyspnoiintheGermanBitterfeldamber;
the age ofwhich is controversial, butwhich maybe
Oligocene (ca.24-25Ma). Someoftheserecords are
ofspeciespreviouslyrecordedfromtheolder(Eocene,
ca. 45-50 Ma) Baltic amber.Two further Bitterfeld
species were described as new, and one is effectively
indistinguishable from, and potentially conspecific
with, aliving Caucasus harvestman species.
Cyphophthalmids (mite harvestmen) are widely
recognisedasthemostbasallineageofOpiliones(e.g.
GlRIBET et al. 2002, 2010: Figs. 9-10) andyet they
show the poorest fossil record of any of the major
harvestmangroups.Thismaywellbeaconsequenceof
JasonA.DUNLOP,MuseumfürNaturkunde,LeibnizInstitute
forResearchonEvolutionandBiodiversityattheHumboldt
UniversityBerlin,Invalidenstraße43,D-10115Berlin,Germany.
E-mail:[email protected]
PlamenG.MITOV,DepartmentofZoologyandAnthropology, Figures1-2:Sirobalticussp.nov.;holotype(F2147/BB/CJW).
FacultyofBiology,UniversityofSofia,8DraganTsankovBlvd., Thefirstrecordofafossil miteharvestman(Arachnida:
1164Sofia,Bulgaria. E-mail:[email protected] Opiliones:Cyphophthalmi)from Eocene(ca.45-50Ma)
Balticamber.- 1:Dorsalview.-2:Ventralview.Scalebar
submitted:10.12.2009,accepted:26.4.2010;online:10.1.2011 equals 1.0mm.
48 J.A.Dunlop&P.G.Mitov
GlRIBETetal.(2010)predictedapossiblediversifica- sequential series ofca. 10 images at different focal
tionage(asopposedtoorigins)forCyphophthalmiof planesthroughthespecimenusingaLeicastereomi-
Ma
ca.345 (i.e. earlyCarboniferous).This postdates croscope with the LeicaApplication Suite software.
the oldestknown harvestman (probablyan eupnoid) Stacks ofimages were assembled into a single final
which is earlyDevonian (ca. 410 Ma) in age. pictureusingAutoMontage.Itwasdrawnandstudied
The extant cyphophthalmid fauna currently underastereomicroscopewithacameralucidaattach-
comprises 168 species (MURIENNE et al.2010).The ment (Fig. 5-6).The fossil was compared to extant
first fossil example ofa cyphophthalmid, Siroplaty- cyphophthalmids in the collections ofthe Museum
pedipusDunlop5cGiribet,2003,wasdescribedfrom fürNaturkunde Berlinandthepersonalcollectionof
Bitterfeld amber and assigned to the extant genus P.G. Mitov, as well as to the literature; particularly
Siro Latreille, 1796 (Sironidae). A second species, Juberthie (1970), Giribet 5cBoyer (2002) and
Palaeosiro burmanicum Poinar, 2008 was described KARAMAN (2009), from whom the morphological
fromCretaceousMyanmaramber(seeabove).Itwas terminology used is largely adopted. Terms for the
assigned to a new (extinct) genus, also in the family description ofthe coxo-genital region are based on
Sironidae,basedonthecombinationofitssmallsize, Hoffman (1963) andvan der Hammen (1985).
thepresenceoftype2ozophores,roundspiracles,and Furtherdataonthedistributionofmoderngenera
a large sternal gland on the first sternite (POINAR wasretrievedfromthe catalogue ofGlRIBET (2000),
2008). In terms ofbiogeography (e.g. BOYER et al. nowupdatedonline andincludingnumerous images
2007, Fig. 1) its assignmentto Sironidae is probably ofextanttaxaunder <http://giribet.oeb.harvard.edu/
incorrect (Giribet pers. comm.), given that only the Cyphophthalmi/>.An exactprovenance forthe new
family Stylocellidae occurs in this region ofSE Asia fossil specimenis notrecorded,butmuchofthe cur-
today. Based on the putative presence ofa cuticular rently available amber derives from the Kaliningrad
lensandmicrovilliintheozophoresPoinarsuggested regiononthe BalticcoastofRussia.Anexactagefor
that these structures may also have functioned as amberis difficultto determine objectively,butBaltic
light-sensitiveorgans,inadditiontotheirfunctionin amber is traditionally dated at Paleogene (Eocene),
releasing defensive secretions. A more prosaic inter- or about 45-50 Ma.
pretationwould be that that the translucent areas at Forcomparativepurposes,ascanningelectronmi-
theendsoftheozophoresmerelyreflectfluidreleased crograph (SEM) photographofthegenitalregionof
duringthe entrapmentprocess. arecentcyphophthalmidharvestmanisalsoincluded
Here,we report on only the third fossil example here (Fig. 4): 1 female Cyphophthalmus duricorius
NW
ofacyphophthalmid (Figs. 1-3,5-6).The specimen Joseph, 1868: Slovenia, 4 km from Postojna,
isincompletewhichmakesdetailedcomparisonswith PivkaJama camp site, under stones in the old pine
otherextantandfossiltaxa (cf. KARAMAN2009 and forest (N 45°48’18.28" E 14°12T6.01", 550-579 m
references therein) difficult. However some genus- a.s.l.), 16.IX.1989, leg. and det. P. G. MITOV. The
specific characters can indeed be recovered and we SEM study was made at 10-20 kV with a Philips
can provisionally assign it to a new species of Siro 515 machine. The specimen was sputter-coated
;
representingthefirstrecordofthegenus from Baltic with a 300-400 Ä gold layer. Further SEMs ofthe
amber. American species Siro exilis Hoffman, 1963 were
kindly provided by Günther Raspotnik (Figs. 7-8):
MaterialandMethods 1 female Siro exilis Hoffman, 1963: West Virginia,
The new fossil described here is from the Jörg Randolph Co., Monongahela National Forest, nr.
Wunderlich collection, specimen number F2147/ Bowden; Otter Creek Wilderness trailhead near
BB/CJW.This materialwillprobablybe transferred Alpena Gap, deep litterofmixed riparianforest (red
eventually to the Senckenberg Museum, Frankfurt/ maple, yellowbirch, eastern hemlock,white spruce)
Main, Germany,orone ofits alliedinstitutions.The withdenseunderstoryofrhododendron,930ma.s.l.,
specimenliesinasubrectangularblockofclear,yellow 38° 56.505’ N, 79° 40.084’W, 13.VI.2006, leg. and
amber; dimensions ca. 25 x 15 mm. Itwas extracted det. RoyA. Norton.
from a larger amber piece (number F2159) in order The specimenwas air-dried,mountedon an alu-
to reveal theventral surface ofthe harvestman more minium stub and sputteredwith gold (AGAR sput-
clearly. tercoater,Gröpl,Tulin,Austria).TheSEMstudywas
The fossil was photographed (Figs. 1—2) by taking made at20kVwithaPhilipsXL30 ESEM (Philips/
SiroinBalticamber 49
FEI,Vienna,Austria) athighvacuum mode. 0.11 and 0.12 respectively. Posterior end ofopistho-
OrderOpilionesSundevall, 1833 soma bluntly rounded to slightly angular in dorsal
SuborderCyphophthalmiSimon,1879 view.
FamilySironidae Simon, 1879 Ventralprosomalcomplexincomplete.Chelicerae,
Genus SiroLatreille, 1796 pedipalps, coxae oflegs 1-2 and allwalkinglegs be-
Sirobalticussp.nov. yondthetrochanterabsent.Conceivably,themissing
coxae(I—II,orperhapsIIonly)werefree,i.e.notfused
Material: Holotype $ and onlyknown specimen,JÖRG tocoxaeIII—IV;whichcouldexplainwhythebreakage
Wunderlichcollection,F2147/BB/CJW.FromBalticam- point in the fossil lies between the second and third
ber,exactlocalitynotrecorded;Paleogene,Eocene. coxae.Third coxae triangular, comingto apointim-
Diagnosis: Relativelylarge (length 2.34 mm) fossil mediatelyabove the thoracic complex,but not quite
Siro species, specifically without the projecting rear reaching the midline. Third trochanter somewhat
endtypicalofmodernEuropeanforms,andwithbody
proportions differing from those in the - probably
morecloselyrelated-extantNorthAmericanspecies
(see Remarks for details).
Derivation ofname: From Baltic amber and the
Baltic region; the source ofthis material.
Description: Partially complete female specimen
preservedinbothdorsal (Figs. 1,5) andventralview
(Figs.2,6).Allmeasurementsinmm.Bodyoval;pale
brown in colour within the amber, but with darker
patches acrossthebody;totallength2.34;maximum
prosomalwidthbehindtheozophores1.33;maximum
opisthosomal width 1.34. Lengthrwidth ratio 1.75.
Distancebetweenfrontofscutum (i.e.anteriormar-
gin ofprosoma) to an imaginaryline connectingthe
anterior (front) bases ofozophores 0.23; totalwidth
across (andincluding) ozophores 1.09.
Entire body with pustulate ornament of small,
roundedtoovaltubercles,generallylargerinanterior
body regions and smaller posteriorly and on the leg
trochanters.Eyesabsent.Ozophoresconicaltoslightly
pointedandangular,dorsolaterallyprominentonthe
scutuminthetype2positionsensuJUBERTHIE(1970,
Fig.2). Exactposition ofthe ozopore itself-i.e. the
opening ofthe repugnatorial gland - difficult to re-
solve,butmaybeterminal.Lengthofozophores0.13;
widthatbase0.22. Slightbulgetobodyimmediately
behind the ozophores. Frontal ridge ofscutum (i.e.
anteriormarginofcarapace) slightlyrecurved.Sulcus
beginningimmediatelybehindthe ozophores curves Figures3-4:Detailsofthethoraciccomplexregion.-3:Siro
down towards the midline and defines a posteriorly balticussp.nov.;holotype(F2147/BB/CJW).Therounded
deeply recurved anterior area of the body; length spiracle(sp)withdenticlesinsidethelumenisaconvin-
0.84 on the midline.Areabehind itincorporates the cingcharacterofthegenusSiroLatreille,1796.Coxae
bulge inthebodylaterallyandis alsorecurvedatthe anruomubnedretdh.eTchoexaalrpeoarebse)twheasenancooxbatlusleo,becsa.II9I0a°,ndanIgVle(i.e.
midline, length here 0.12. This region followed by (outlined byblackbars:arrowed);anotherSirocharacter.
eight,quiteclearlydefined,tergitesallwithessentially Scalebarequals200pm.-4:Sameregionina moderncy-
straightposteriormargins.Gapsbetweentergiteslack phophthalmid,CyphophthalmusduricoriusJoseph,1868,SEM.
tuberculation.Anteriorfourtergiteslonger,lengthsc. aSrcealaebbeatrwseeeqnuaclox1a0l0lpobme.sInIIICaypnhdopIhVthhaalsmuas,vebryycaocnutrtaesta,ngtlhee
0.2;posteriorthree are notablyshorter,lengths 0.13, (outlined byblackbars:arrowed).
50 J.A.Dunlop&P.G.Mitov
Figures5-6:Interpretativecameralucidadrawingsofthespecimen shown in Figs.1-2.5:Dorsalview.-6:Ventralview.Abbrevi-
ations:ca=expectedsiteofcoronaanalis,cp=carapace,cx=coxa,oz=ozophore,sp=spiracle,st=sternites2+3 (subse-
quentsternitesnumberedsuccessively),tc=thoraciccomplex,tg1 =tergite 1 (subsequenttergitesnumberedsuccessively),
tr=trochanter.Scalebarequals 1.0mm.
roundedandcup-shaped,butdetailsequivocal.Fourth linesulcus.Eithersideofthis,i.e.ontheanterolateral
coxaemuchlargerthanthe thirdandmore quadratic margins of the gonostome, are areas surrounding
in shape. Distinct suture line originates near the an- the coxalpores (= orifices ofthe coxalglands). Pores
terolateralcornersofthethoraciccomplexandcurves themselves clearlyvisible as small,butdistinctholes,
towards the posterior margin ofthe coxae,bisecting diameter 0.0125, lying between coxal lobes III and
it about a third ofthe way along its length towards IV.
the distal margin. Fourth trochanter emerges from Sternites two and three apparently fused into a
the posterior part ofthe fourth coxae and is associ- single, large subtriangular plate, projecting anteri-
ated with a small, but distinct indent into the coxal orly between the leg 4 coxae with a procurved and
margin.Trochanter 4 tubular, longer than wide and bluntly rounded anterior margin pointing towards
with a slightlyoval outline,wideningdistally,length the thoracic complex. Tuberculation here heavier,
0.28. withlargerandmoreovaltubercles.Spiraclespresent
Thoraciccomplexwiththe outlineofaninverted as prominent,roundto oval-shaped elements (maxi-
mum
subtriangularstructure, dominated centrallybylarge diameter0.16)inalateralpositionimmediately
genital opening (or gonostome) ofthe female type behindthefourthcoxae;ringoftinydenticlespresent
(Fig. 3). Gonostome anteriorly with semicircular within the lumen, expressing - at least on the right
outline;width0.17,length0.16;nogenitalstructures side-adistinctinvaginationtowardsthecentreofthe
visiblewithinthisopening.Thoraciccomplexflanked spiracle.Noobvioussternalpores,orotherstructures,
laterallybysutured-offpartofthe fourth coxae,pos- located on the midline between the spiracles.
teriorlybythesecondsterniteandanteriorly(inpart) Five sternal elements (presumably sternites 4-8)
by the third coxae. More anterior elements missing. preserved behind the large, spiracle-bearing sclerite.
Thoraciccomplexdivided anteriorlybyashortmid- Sternites 4—7 decrease successively in length: i.e.
S0i.roinBalticamber 51
20, 0.19, 0.16 and 0.10.
Sternite8formstheanterior
border ofa fairly large oval
opening, width 0.32, which
in life would probably have
1.
contained the corona analis.
Posteriormarginofopistho-
soma somewhat blunt and
rectangular in ventral view;
e. not smoothly rounded,
but details of any specific
sclerites here difficult to
resolve. In general, tergites
slightlywider than sternites
and marginal overlap from
the overlyingtergites canbe
seen ventrally at the lateral
edges ofthe body.
In cyphophthalmids the
form ofthe gonostome dif-
fers between the sexes and
this specimen clearly has Figures7-8:ComparativeSEM imagesofafemaleoftheRecentspeciesSiroexilisHoffman,
the female type. Since the 1963fromtheeastern USA.TheseeasternSirospeciesarethoughttobeclosertothe
gonostomeispreservedopen eEquuraolpsea1n.0fmamun.aNtohtaentthheoosveeroafltlhseimwielsartietryntoUSthAe(cffo.ssSihleSairro1s9p8e0c)i.e7s:iOnvteerrvmisewo.fStchaeleabbsarence
(Fig. 3), it must be an adult ofa projecting rearend (arrowed);afeatureusuallyseen inthe(modern)European
and is thus clearly a mor- forms.-8:Detailofspiraclesandthoraciccomplexregion.Scalebarequals200pm.
tality rather than a moult.
Intuitively, the newfossilwith its distribution in the fossil.An alternative character (cf. KARAMAN 2009:
EoceneofnorthernEuropeislikelytobeamemberof p. 262) which would have been useful is the shape
Sironidae (see e.g. GlRIBET(2000: Fig.2) orBOYER and structure oftheprosomalcomplex (i.e. the coxal
et al. (2007: Fig. 1) for distribution maps) because lobes oflegs II, immediately in front ofthe genital
none of the other extant cyphophthalmid families opening), but unfortunately this feature is equivocal
occurinthisregiontoday.Thefossilisunquestionably in the fossil.
moderninappearanceandaffinitieswithanumberof AcharacterdiscussedbyKARAMAN(2009)which
extantgeneraneedtobeconsidered:i.e.SiroLatreille, ispreservedisthe form ofthe spiracles andthis does
1796, Cyphophthalmus Joseph, 1868 (re-established offersusefuldataabouttheanimal’s affinities.In Cy-
byBOYERetal.2005 foraBalkanradiation; see also phophthalmusthespiraclesaresemicircular,eachwith
KARAMAN 2009 and MURIENNE et al. 2010) and aconicalcuticularprojectiononitsposteriormargin.
thefourIberiangeneramostrecentlyinvestigatedby This is notseen in the fossil,whichbycontrast (Fig.
MURIENNE8cGlRIBET(2009).Thesetaxaaremor- 3) seems to share acharacterseen,sofar,onlyin Siro
phologically rather conservative and appear similar and which consists of a more rounded spiracle (of
in overall habitus, while a number oftaxonomically circular type, sensu GlRIBET &,BOYER 2002), with
important details are missing from the new fossil denticles inside the lumen (cf. BlVORT &,GlRIBET
which hinders its unequivocalgeneric assignment. 2004: Fig. 36c, KARAMAN 2009: Figs. 2A-B). For
KARAMAN (2009: table 1) established a series of thisreasonweareconfidentinourgenericassignment
characters useful in separating Siro from Cyphoph- ofthis fossilto Siro.In supportofthis hypothesiswe
thalmus.Someofthese,suchasthenumberofpaired, alsonotethattheformandwidthoftheareabetween
movablefingers associatedwiththespermatopositor, coxallobes III and IVis verydifferentbetween Siro
the number ofanal glands, and the shape ofthe ad- and Cyphophthalmusspecies. Specifically, the endites
enostyle ofthe tarsal gland apophyses, onlyoccur in (= coxapophysis) ofcoxae III and IVform either an
males and are thus unhelpful in placing this female area (aroundthe coxalpores)with averyacute angle
52 J.A.Dunlop&P.G.Mitov
Discussion
(in Cyphophthalmus’.Fig.4)oraright/obtuseangle(in
Siro:Fig.3);thusinthiscontexttheamberfossilmore Asnotedabove,GlRIBETetal.(2010)inferredabasal
closely resembles female Siro species. For compara- divergencetimeformoderncyphophthalmidlineages
tive figures ofSiro see e.g. RAFALSKI (1958: Fig. 7) perhaps as far back as the Carboniferous (ca. 345
for Sirocarpaticus Rafalski, 1956,JUBERTHIE (1967: Ma);atwhichtimeEuropeandNorthAmericawere
Fig. 7) for SirorubensLatreille, 1804, andNOVAKSc part ofthe single palaeocontinent Laurasia. Indeed,
GlRIBET (2006: Figs. 5, 7, 12) for SirocrassusNovak SHEAR (1980, p. 4) commented on the strong simi-
ScGiribet,2006.Forvarious Cyphophthalmusspecies larities between some ofthe (eastern) N. American
wereferalsotoElSENBEISdcWlCHARD(1987:Plate cyphophthalmids and the European fauna: “... the
27, d) (sub Siro duricorius), MlTOV (1994, Fig. 23) original divergence took place between the western
(sub Sirobeschkovi) or KARAMAN (2008,2009). speciesandS[iro]exilis(Figs.7-8)plustheEuropean
Although incomplete,we feel able to assign this forms.The movement ofNorthAmerica awayfrom
new fossil to its own species. Ofparticular interest Europe and Africa, which resulted in the opening
is the posterior end ofthe body. Females ofmodern ofthe present Atlantic Ocean, may account for the
EuropeanSirospeciestypicallyshowaprojectingrear separation of S. exilis from its European relatives.”
end(seee.g.figuresinJUBERTHIE 1970),whereasthe ThehypothesisthattheNorthAmericanSirospecies
Baltic amber example has a more smoothlyrounded arenotmonophyleticwasagainsupportedinarecent
backendwhichthusresembles,theNorthAmerican studybyGlRIBETScSHEAR(2010),who furtherre-
Siro species (e.g. NEWELL 1943, 1947, HOFFMAN covered European affinities for some American taxa
1963, SHEAR 1980; see also Fig. 7). Conceivablyour undersomeparameters ofanalysis andreiteratedthe
fossil was part ofthis (formally Laurasian?) lineage idea that Siro maybe averyancientgenus.
(see Discussion). At 2.34 mm in body length the What this implies is, first, that there is no fun-
new fossil sironid is somewhat larger than typical damental objection to recovering an American-like
American Siro species. Bodylengths offemales vary SirospeciesfromBalticamber.Thislineagemaywell
mm
from 1.10 (in Siro sonoma Shear, 1980) to 2.08 have been originally distributed more widely across
mm(inSiroexilisHoffman,1963-thelargestmodern the Palaearctic, and was present in north-central
American sironid); see also NEWELL (1943, 1947). Europe during the presumably warmer conditions
ThelargestEuropeansironidistheepigeanSirocrassus associated with the Baltic amber forest. A possible
NovakScGiribet,2006inwhichfemalesreach2.40- parallel example ofthis would be the eupnoid har-
2.61 mm. vestman genus Caddo Banks, 1892 which was also
Direct comparisonwith the previouslydescribed presentinEuropeduringtheearlytomid-Paleogene
Europeanamberspecies,Siroplatypedibus,is difficult (DUNLOP ScMitov 2009, and references therein),
giventhatthefossils arepreservedincompletelydif- but is absent from the modern European fauna and
ferentorientations.Thecoxo-genitalandanalregion yet still found todayin North America. Second, the
ofthe younger Bitterfeld fossil cannot be resolved. opening ofthe Atlanticbegan during in theTriassic
The originally proposed diagnosis was based on its (ca.200-250Ma),andthis,inturn,offersaminimum
flattened legs; a characterwhich cannotbe tested in divergence time, based on geological evidence, for a
the Baltic form. In any case this has recently been commonancestorofthesesimilar-lookingAmerican
challengedasapossibleartefactbyKARAMAN(2009). and European Siro species; the latter also including
Henotedthatlimbflatteningcanoccurwhilemaking Sirobalticus.
preparations ofextant material in various mounting Toreiterate,NorthAmericayields Siro(cf.GIRI-
media,andthattheamber-formingresinasasimilarly BET ScSHEAR2010),while both Siro and Cyphoph-
concentrated viscose medium could have induced thalmusoccur in Europe. In recent studies Cyphoph-
comparableeffects.OurnewfossilandS.platypedibus thalmuswas treated as a specificallyBalkanradiation
are,atca.2 mmlong,similarinoverallsize.While it (BOYER et al. 2005, KARAMAN 2009); although we
maybepossible to drawsome comparisonsbasedon should note that Siro has also been reported at least
the profile ofthe body sculpture in these respective as far south as Slovenia (NOVAK ScGlRIBET 2006).
fossils,wecurrentlyhavelittledatatoargueeitherfor Recentwork byMURIENNE et al. (2010) inferred a
oragainsttheconspecificityoftheseextincttaxafrom late Cretaceous radiation for Cyphophthalmus ofca.
the different amberfaunas.
SiroinBalticamber 53
94Ma and attempted to tie the explosive evolution DunlopJ.A. &c P.G. MlTOV (2009): Fossil harvestmen
ofits numerous endemic species into thewidergeo- (Arachnida: Opiliones) from Bitterfeld amber. -
logical development ofthe Balkan Peninsula. Slow ZooKeys 16: 347-375-doi: 10.3897/zookeys.16.224
ratesofevolutionforcyphopthalmidsingeneralwere ElSENBEISG.&W.WlCHARD(1987):Atlasonthebiology
postulated by SHEAR (1980) - see also GlRIBET & ofsoilarthropods.Springer-Verlag,Berlin,Heidelberg,
SHEAR (2010) - and the modern appearance ofthis NewYork,London,Paris,Tokyo.437pp.
amberfossil(cf.Figs.1-2and7-8)comparedtoextant GlRIBET G. (2000): Catalogue ofthe Cyphophthalmi of
theworld (Arachnida: Opiliones).-RevistaIbericade
forms tends to support this supposition: at least for
Arachnologia2:49-76
Siro.Unfortunately,westilllackPalaeozoiccyphoph- GlRIBET G. & S.L. Boyer (2002): A cladistic ana-
thalmids which should date from the earlyphase of
lysis of the cyphophthalmid genera (Opiliones,
theirevolution aspredictedbymolecular analyses. Cyphophthalmi). - Journal of Arachnology 30:
110-128 - doi: 10.1636/0161-8202(2002)030[0110:
Acknowledgments ACAOTC]2.0.CO;2
We thankJörgWunderlich (Hirschberg) for making this Giribet G., G.D. Edgecombe, W.C. Wheeler &
material available for study and preparing the specimen C. BABBITT (2002): Phylogeny and systematic posi-
from alargerblockofresin,aswellasIvo Karaman (Novi tion ofOpiliones: a combined analysis ofchelicerate
Sad)andGonzaloGiribet(Harvard)forvaluablecomments relationships using morphological and molecular data.
and to Gonzalo for providing pre-publication copies of -Cladistics18:5-70-doi: 10.1111/j.1096-0031.2002.
relevantpapers.GünterRaspotnik(Graz)kindlyprovided tbOO140.x
aSvEaiMlabilmeabgyesDor.fSRioroyexNiolirstmoante(riSaUl,NwYh-iEcShFi,nStyurrancwusaes,mNaedwe GlRIBETG.&W.Shear(2010):ThegenusSiroLatreille,
1796(Opiliones,Cyphophthalmi,Sironidae),inNorth
York). Finally, the reviewers are also thanked for helpful
Americawithaphylogeneticanalysisbasedonmolecular
remarks. dataandthedescriptionoffournewspecies.-Bulletinof
theMuseumofComparativeZoology160: 1-33-doi:
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