景德镇高专-猜灯谜大全

rnalofArachnology39:211–222
Phylogenyand classificationofPholcidae(Araneae):anupdate
:Al exanderKoenigResearchMuseumofZoology,Adenauerallee 160,53113Bonn,Germany.
E-mail:@
r100years,Eu ge`neSimon’ssystemofpholcidclassificationhasbeenus edwithonlyminor
eneticresearchoverthelastdecade hasshownthatsomefundamentalchangesarenecessaryifth e
ncladisticanalysesofmorphologicalandmolecular
dataandonqualitativecharacterassessment,thefam ilyisheredividedintofivesubfamilies:Ninetinae,Arte minae,
Modisiminae,Smeringopinae,oughthe
evi dencesupportingsomeofthenodesandassignmentsisweak, thecladogramgeneratesnumeroustestablehypotheses
andprovidesanimprovedframeworkforthemappingof‘new ’characterslikethosefromspermultrastructureand
chromosomeanalysis.
Keywords:Ninetinae,Artemina e,Modisiminae,Smeringopinae,Pholcinae
WhenSimon (1893)presentedthefirstclassificationof
Pholcid ae,hedidsointheLinnaeantradition,withsimilarity
tem(Table1)provedto
beusefulasameanstoorganize information,andforover

generawereadded,some generawereshiftedtoothergroups,
andsomeofSimon’ sgroupsweremerged,butjudgingfrom
ourcurrentview ofpholcidphylogeny,thetwomajorupdates
byPetrunk evitch(1928)andMello-Leita˜o(1946)werequite
obv iouslybasedonapoorerunderstandingofthefamilythanSimon’soriginalsystem(aviewalreadyexpressedbyBri gnoli
1972a,1972b,1980).
Thefirstphylogeneti canalysisofthefamily(inHuber2000)
indicatedthat someofSimon’sgroupswereverylikelynot
monophylet ic,butforseveralreasonsIhesitatedtopresenta
new formalsystembecause:1)someofthemajorcladesdidnotseemwellsupported(e.g.,ninetines,holocnemines);2 )thetaxon
sampleofsomemajorgroupswasinadequate( especiallytrue
forOldWorldtaxa);and3)forthcomin gprojectswerelikelyto
resultinfurtherchangestoo urunderstandingofpholcid
ecidedtouseprovisional ,informalnames
suchas‘‘ninetines’’,‘‘holocnemin es’’,‘‘NewWorldclade’’,and
‘‘pholcines’’rathert hanformalsubfamilynames(Huber2000).
Since2000,s everalfurtherphylogeneticanalyseshavebeen
publi shed,usingbothmorphologicalandmoleculardata(Astrin
´
etal.2005;Huber2001,2003a,etal.2007;Bruvo -Madaric
2003b,2003c,2005a,2005b,2007).Eventhou ghmanyofthe
samereasonsthatpreventedmefrompubli shingaformalsystem
in2000couldstillbeinvoked,it appearsjustifiedtopresentthe
systematicstatusqu oafteradecadeofphylogeneticworkonthe
jorcladesn owappearwellsupported
andlikelytobestable,andmo stgeneracanbeplacedwithsome
rentpaperisthusanattempttosummarize,update,andformalizethesubfami ly-level
r,thesystembelowisa
workinghypothes is,andongoingprojectsjustifythehopethatit
willn otlastaslongasSimon’s.
METHODS
Thephylogenet ichypothesispresentedbelowismainly
derivedfromt wosources:1)cladisticanalysesoftheentire
211
familyorofsubgroupsofPholcidae(Astrinetal.2007;Br uvo-
´
etal.2005;Huber2000,2001,2003a,2003b, 2003c,Madaric
2005a,2005b,2007;HuberinpressonPh olcusWalckenaer
1805andcloserelatives);and2)qua litativeassessmentof
particularcharacters(e.g., tarsus4comb-hairs:Huber&
Fleckenstein2008).Sinc etheaimistopresentahypothesisfor
theplacementof allnamedgenera,asupertreeorsupermatrix
approach wasnotfeasible;toomanygenerahaveneverbeen
inclu dedinanycladisticanalysis,andsomegeneraare
extr emelypoorlyknown(e.g.,somemonotypicVenezuelan
g enerarecentlydescribedbyGonza´lez-Sponga1998,2003,
2009).Asaresult,nomeaningfulbranchsupportvalue scan
d,
thesupportforeachnodeispresentedandd iscussed
separately,usingadmittedlysubjectivean dqualitativejudg-
ments,thoughinamannerthatisho pefullytransparentand
traceable.
Thesubfamil ynamespresentedbelowaremostlytaken
fromSimon’s( 1893)system,eventhoughSimonrecognized
onlytwosu bfamilies:NinetidinaeandPholcinae(Table1).
Howe ver,thelattersubfamilywasdividedintoseven
subgr oups,andfourofthesefamily-groupnamesarehere
,th ePholcinaeasdelimited
hereinismuchnarrowerinsco pethanSimon’sPholcinae,but
eistrueof
netinae arefullycongruentin
bothsystems,butthisissimply duetothefactthatSimon’s
(1893)Ninetidinaewasmon otypic.
CLASSIFICATION
1850
Themonophylyo fPholcidaehasneverbeenseriously
questioned,anda goodnumberofcharactersareboth
synapomorphicandu sefulforeasydiagnosis(Huber2000).
Themalepedipa lpisusuallyheavilymodified,includingeven
thepro ximalsegments(Figs.3–7).Aprocessofthepalpal
tar sus(the‘procursus’)occursinalmostallspecies(pin
Figs.3–7).Themalecheliceraearealsousuallymodified
(Fig.2),andthesemodificationsarefunctionallyre latedto
modificationsofthefemaleexternalgenital ia(Huber1994,

212
Table1.—Euge`neSimo n’s(1893)schemeofclassificationforthe
Pholcidae .*BlechroscelishasbeensynonymizedwithPriscula(Hube r
2000);HedypsilushasbeensynonymizedwithModisim us(Huber
1996b).
HighertaxaIncludedgenera
Pholcidae
NinetidinaeNinetisSimon1890
Pholc inae
ArtemeaeArtemaWalckenaer1837
PholceaePh ysocyclusSimon1893,Pholcus
Walckenaer1805,Holoc nemusSimon
1875,SpermophoraHentz1841,
Metago niaSimon1893
LeptopholceaeLeptopholcusSimon1893 ,Micromerys
Bradley1877
SmeringopodeaeSmerin gopusSimon1890,UthinaSimon
1893,CrossoprizaSimo n1893
PrisculeaePrisculaSimon1893
Blechrosce leaeMecolaesthusSimon1893,Psilochorus
Simon1893 ,CoryssocnemisSimon1893,
BlechroscelisSimon1893 *,Litoporus
Simon1893,SystenitaSimon1893
Mod isimeaeModisimusSimon1893,HedypsilusSimon
1893*
1995,1997a,1998,2002).Thelatterareoftensculptu redand
moreorlesssclerotisedexternally,resultin ginanepigynum
thathasevolvedindependentlyofthee ntelegyneepigynum
(andthatisfunctionallyrelated tothemalecheliceraerather
thantothemalepedipalp asinEntelegynae).Otherputative
synapomorphiesar ethehighclypeus(Figs.8,9),the
arrangementofeyes (twotriadsseparatefromtheanterior
medianeyes–th elattermaybeabsent;Figs.8–11),three
trichobothr iaonthelegtibiae,tarsalpseudosegments,comb-
hai rsonthefourthtarsi(Figs.12,13),andeggsacscarriedby
femaleswiththeirchelicerae.
SubfamilyNineti naeSimon1890
Typegenus.—NinetisSimon1890(bymono typy).
Ninetinae:Simon1890:93.
Ninetidinae:S imon1893:oughthisisthe
grammaticallycorrectspel ling,theelidedoriginalformis
inprevailinguseand ismaintainedinaccordancewith
Article29.3.1.1oft heInternationalCodeofZoological
Nomenclature(In ternationalCommissiononZoological
Nomenclature1 999).
ThemonophylyofNinetinaeissupportedbytwopu tative
morphologicalsynapomorphies(Huber2000):a narrow
openingofthecapsulatetarsalorgan(compare Figs.17and
18)andarelativelydistalpositionofthe retrolateral
trichobothriumonthelegtibiae(beyon d45%oftibialength).
Thecompactocularareawiththe anteriormedianeyesina
veryhighposition(sometime sevenabovetheanteriorlateral
eyes;Fig.9)egs
(maletibia1onlyupto2.53carapacewidth)aretypicalfor
ninetines,butthisispossiblytheplesiomorphicpho lcid
lardatahavesofarsupportedthe
monophylyo fninetines,butthesampleofbothtaxa(three
species )andgenesisstillinadequate(Astrinetal.2007).
Ni netinesmayindeedbesistertoallotherpholcids(asimpli ed
THEJOURNALOFARACHNOLOGY
inSimon’s1893syst em),butphylogeneticsupportforthe
latterclade(cl ade1ainFig.1)isweakandambiguous:awide
sternuman dlonglegsmaybemorphologicalsynapomorphies;
mole cularsupportisalsoweak(Bruvo-Madaric
´
etal. 2005;
Astrinetal.2007).
Withinninetines,theg enerainclade2asharealongspine-
likeprocessonthe malepalpalbulb(Huber2000:figs.321,
335);thegene rainclade2bsharereducedepiandrousspigots
andane xposedtarsalorgan(acharactercombination
reminis centofModisiminae:Figs.19,22;Huber2000;Huber
&E lHennawy2007);thegenerainclade2csharealarge
dor salflapontheprocursus(Huber2000:figs.374,383);the< br>generainclade2dsharealongprocursuswithcharacter istic
proximalcourse(Huber2000:figs.353,360);th egenerain
clade2eshareasimplified(orevenreduced )procursus(Huber
2000:figs.437,454,466).Thesign ificanceoffrontalhumpson
themalesternumisdubiou s;theyoccurinvariousNinetinae
butalsoinModisimi nae(Huber2000).

numberssuggestrelativelyhig hgenus-leveldiversitybutalack
netinesaretinyspi dersthatlive
closetotheground,e
largelyrestr ictedtosemiaridregions(Huber&Brescovit
2003),highestdiversityoccursintheNewWorld(Fig.24),with only
NinetisandNitawidespreadinAfricaandtheMidd leEast.
TheMalaysianMystesBristowe1938isadubiou smonotypic
kaalaHuber
2000fromNewCaledoniais theonlyrepresentativeofthe
genusoutsideChile.Composition:AucanaHuber2000;ChisosaHuber2000;
EneteaHuber2000;GalapaHuber2000;GertschiolaBrigno li
1981;GuaranitaHuber2000;IbotyporangaMello-Le ita˜o1944;
KambiwaHuber2000;MystesBristowe1938; NerudiaHuber
2000;NinetisSimon1890;NitaHuber&El Hennawy2007;
PapiamentaHuber2000;Pholcophora
Banks1896;Serrato-
chorusWunderlich1988;Toltec aHuber2000.
SubfamilyArteminaeSimon1893
Type genus.—ArtemaWalckenaer1837(bymonotypy).
Arteme ae:Simon1893:463.
Theonlyknownmorphologicalsyna pomorphyofArteminae
isapairofdistinctivestructu resonthemaleprocursus:adorsal
apophysisandavent ralpocket(Fig.6).InPhysocyclus
globosus
(Tac zanowski1874),thefunctionofthesestructures
hasb eenstudiedinsomedetail:duringcopulation,the
apo physisofoneprocursusislodgedinthepocketoftheother< br>procursus,resultinginasymmetricinsertionofthepr ocursi
(Huber&Eberhard1997).Moleculardataalsote ndtogroup
thethreearteminegenerasequencedsofar( Bruvo-Madaric
´
et
al.2005;Astrinetal.200 7).Arteminaemaybesisterto
Modisiminae(clade1bin Fig.1),butthisrelationshipwasonly
supportedbyac ombinedanalysisofmolecularandmorpho-
logicaldat ainBruvo-Madaric
´
etal.(2005).Thereductiono f
epiandrousspigotsandmoleculardatasuggestthatt hetwo
generainclade3aaremorecloselyrelatedtoeac hotherthan
botharetoArtema(Huber2001;Bruvo-Mada ric
´
etal.2005).
Arteminaecurrentlyinclu des64speciesin5genera,withthe
largemajorityofsp eciesinthegeneraPhysocyclusSimon1893

HUB ER—PHOLCIDPHYLOGENY
213
Figure1.—Proposedcla dogramforthefamilyPholcidae,ednodesarediscussedint hetext.

214
THEJOURNALOFARACHNOLOGYFigures2–7.—Cheliceraeandpedipalps,illustratingi mportantpholcidcharacters(p:procursus;ta:trochante rapophysis).elicerae
withfrontalandlateralapoph yses(la)(Pholcusquinquenotatus).lpwithdorsalattach mentofbulb(b)totarsus(Buitingakadogo).4.
Pedipa lpwithdistinctsclerite(arrow)betweentarsusandbulb( b)(Pholcusquinquenotatus).lpwithcoxaapophysis(ca)a ndpointed
andupwardprojectingfemoralapophysis(f a)(Tupigeapenedo).lpwithprocursusprovidedwithdorsa lapophysis(ap)andventral
pocket(po)(Trichocyclu sarawari).lpwithshiftedtibia-tarsusjoints(arrows;c omparewithFig.4)(Zatavuazanahary).FromHuber
200 1,2003a,2003b,inpress,Huber&Rheims2011.
teminae arerelatively
largespiderswithlong,stronglegsan dhighglobose
keNinetinae,theyoftenoccurinrather
dryregions,sometimesevenindeserts,liketheAustr alian
logyofthepantropicalPhysocyclus
Simon1 893hasbeen
studiedinsomedetail(Eberhard1992a;Eb erhardetal.1993;
Huber1996a;Huber&Eberhard1997; Perettietal.2006).

HUBER—PHOLCIDPHYLOGEN Y
215
Figures8–23.—Scanningelectronmicrograp hsillustratingimportantpholcidcharacters.8,ata,fro ntalviews,showingeye
arrangementandhighclypeus( 8,Pholcusbourgini;9,Ninetissubtilissima).areaandth oracicpit(arrow)inCrossoprizacylindrogaster.
ar eaofTupigeateresopolis,showingreductionofanteriorm edianeyes.12,-hairsonthefourthtarsi,complex(12)and
simple(13)types(12,Belisanaketambe;13,Pholcusc hattoni).14,orlateralspinnerets,withcomplete(14)an dreduced(15)setof
spigots(14,Pholcuslamperti;15 ,Calapnitasaluang).alhairsinhighdensityonthefemuro fModisimuspalvet.17–tarsal
organs,‘normal’capsu latetype(17),ninetinetypewithsmallopening(18),ande xposedtype(19)(17,Belisanahormigai;18,Ninetissubti lissima;
19,Tainoniaserripes).edtipofmalepalpal trochanterapophysis(Leptopholcusdebakkeri).21,nopo res,withandwithout
epiandrousspigots(21,Leptoph olcusgracilis;22,Tupigeaangelim).umwithknob-likest ructure(arrow)(Pholcuskwamgumi).From
Huber2005a ,2009,inpress,Huber&Rheims2011,Huberetal.2010,Hube r&vanHarten2001.

216
THEJOURNALOFARAC HNOLOGY
Figure24.—KnowndistributionofNinetinae.
PhysocyclusistheonlyNewWorldgenusinArteminae;t he
othergeneraoccurinAsiaandAustralia(Fig.25).W iththe
exceptionoftwopantropical,synanthropicsp ecies(P.
globosus,ArtemaatlantaWalckenaer1837)A rteminaeseems
tobeabsentfromSouthAmericaandAfri ca.
Composition:ArtemaWalckenaer1837;Holocnemin us
Berland1942;PhysocyclusSimon1893;TibetiaZhan g,Zhu
&Song2006;TrichocyclusSimon1908.
Subfa milyModisiminaeSimon1893
Typegenus.—ModisimusSi mon1893(bysubsequent
designation—herein).
Mo disimeae:Simon1893:484.
‘‘NewWorldclade’’:Huber 2000:36.
ThemonophylyofModisiminaesensustricto( clade4bin
Fig.1)iswellsupportedbythepresenceofa retrolateral
apophysisonthemalepalpalcoxa(Fig.5 )thatstabilizesthe
palpinitsrotatedpositionatth eonsetofcopulation(Huber
1998,2000)andbytheredu ctionofepiandrousspigots
(compareFigs.21and22). Moleculardataalsosupportthe
´
ylyofthiscoreg roupofgenera(Bruvo-Madaric
2005;Astrinetal.2007 ).TheAustralianWugigarraHuber2001
shareswithtax ainclade4bthereductionofpiriformgland
spigotson theanteriorlateralspinnerets(asshowninFig.15for
arepresentativeofPholcinae;Huber2001),butitsphylo genetic
sofWugigarrasharewithArteminae
thedi stinctiveprocursusmodificationsmentionedabove(cf.< br>Fig.6);moleculardataarenotyetavailableforWugiga rra.
PrisculaSimon1893shareswithWugigarraandMod isiminae
sensustrictoanexposedtarsalorgan(Fig.1 9)andalarge
distancebetweentheanteriorlateralan dposteriormedianeyes
(.0.553diameterofposterior medianeyes).Moleculardata
haveeithersuggestedaw eakaffinityofPrisculatoModisiminae
sensustricto oranunresolvedbasalposition(Astrinetal.2007).
R elationshipswithinModisiminaehaveprovendifficultto
nera(clade4c;the‘‘Modisimusgroup’’sensu
Hub er1998)shareadistinctivelyshapedapophysisonthe
Figure25.—anthropicPhysocyclusglobosusandArtemaatl antahavepantropicaldistributionsand
eminuspirit arsisBerland1942iswidelydistributedinthePacific,th
AmericanPhysocyclusviridisMello-Leita˜o1940isc onsideredmisplaced.

HUBER—PHOLCIDPHYLOGE NY
217
Figure26.—anthropicModisimusculicinus (pantropical)andPsilochorussimoni(introducedto
EuropeandNewZealand)areexcluded.
malepalpalfemu r(pointedandupwardprojecting)(Fig.5;
Huber1998, 2000),butmoleculardatahavesofarnot
supportedthe monophylyofthistaxon(Huber&Astrin
2009).Withint hisclade,somegenera(clade4d)shareshort
vertical hairsinhighdensityonthemalelegtibiaeandor
femor a(Fig.16;Huber2000).Suchhairsalsooccurinthetwo
generainclade4h,agroupthatisotherwiseonlysupported by
superficialsimilarity(asaretaxainclade4e).Cl ade4fincludes
mostlyVenezuelangenerathatsharean apophysisonthemale
palpalfemurthatisdirectedtow ardsproximal(orawayfrom
thefemuratarightangle;H uber2000:figs.989,1016,1066).
Themorphologicala ndecologicaldiversitywithinthisgroup
isremarkab leandexplainstheconsiderablenumberofgenera
desc ribedbySimon(1893)afterhisexpeditiontoVenezuelain< br>1887–1888(Levi1964).Recently,thesituationinthis cladehas
beencomplicatedfurtherbythedescription ofnumerous
monotypicgenerabyGonza´lez-Sponga(19 98,2003,2009).
Finally,acloserelationshipbetwee nthegenerainclade4gis
suggestedbymoleculardata( Astrinetal.2007).

isoneofthetwolargestsubfa miliesofPholcidae,withseveral
veryspecies-richg enera(e.g.,AnopsicusChamberlin&Ivie
1938,Psiloc horusSimon1893,Modisimus,Mesabolivar
Gonza´lez- Sponga1998)andmanyundescribedspecies.
Modisimin aearethetypicalpholcidsofthehumidNeotrop-
ics,w heretheyoccupyawidevarietyofmicrohabitatsfrom
l ogicalvarietyisparalleled
byawiderangeofbodyfor ms,fromtinyleaf-andlitter-
dwellingformstosomeo fthelargestpholcidswithlegspans
ofover15cm(Hube r&Astrin2009).Basicbiologicaldata
andsomedetail saboutsexualbehaviorhavebeenstudiedina
numberof species(Bricen˜o1985;Eberhard1992b;Eberhard
&Br icen˜o1983,1985;Fu¨rst&Blandenier1993;Huber1994,1996b,1997b,1998,2005c).Withtheexceptionofthe
dubiousWugigarra(seeabove)andtwosynanthropicspeci es
(Modisimusculicinus[Simon1893];Psilochorussi moni[Ber-
land1911]),Modisiminaeisrestrictedtot heNewWorld
(Fig.26).
Composition:AnopsicusCh amberlin&Ivie1938;Aymaria
Huber2000;BlancoaHube r2000;CanaimaHuber2000;
CarapoiaGonza´lez-Spong a1998;CarbonariaGonza´lez-
Sponga2009;ChibcheaH uber2000;CiboneyaPe´rez2001;
CoryssocnemisSimon 1893;IxchelaHuber2000;Litoporus
Simon1893;Maimi reGonza´lez-Sponga2009;Mecolaesthus
Simon1893;M esabolivarGonza´lez- Sponga1998;Modisimus
Simon1893;NasutaGonza´lez- Sponga2009;OtavaloaHuber
¨
zdikmen&Demir2000 ;PisaboaHuber2000;PlatnickniaO
2009;PomboaHuber 2000;Priscula
Simon1893;Psilochorus
Simon189 3;QueliceriaGonza´lez- Sponga2003;Sanluisi
Gonza´lez- Sponga2003;StenosfemuraiaGonza´lez-Sponga
1998; SystenitaSimon1893;Tainonia
Huber2000;Teuia
Huber2000;TonoroGonza´lez-Sponga2009;TupigeaHuber< br>2000;VenezuelaKoc¸ak&Kemal2008;WaunanaHuber2000 ;
WugigarraHuber2001.
SubfamilySmeringopinae Simon1893
Typegenus.—SmeringopusSimon1890(bysub sequent
designation—herein).
SmeringopodeaeS imon1893:474.
‘‘Holocnemus-group’’:Timm1976:70.
ThemonophylyofSmeringopinaeisweaklysupportedby
thepresenceofalargethoracicpitonthecarapace(Fi g.10;
ratherthananarrowfurroworanevenlydomedcar apaceas
inFigs.8,9,11).Moleculardatahavepartlys upportedthe
´
etal.2005),monophylyofthetaxas equenced(Bruvo-Madaric
andpartlytheyhavesuggest edratherobscurerelationships,
castingdoubtonthe sequencesorontheusefulnessofthe
molecularmarker smorethanontherelationshipssuggested
bymorpholo gy(e.g.,thepositionofHolocnemusamong
Ninetinaei nAstrinetal.2007).Moleculardatahave
consistentl ytendedtosuggestacloserelationshipbetween
Smeri ngopinaeandPholcinae(clade1cinFig.1),andsucha
r elationshipisalsosupportedbythedistributionofcomb-
hairsonthefourthtarsus(overtheentiretarsusleng thrather
thanrestrictedtothetarsustip:Huber&Fle ckenstein2008).

218
Figure27.—an-
thropicCrossoprizalyoni,SmeringopuspallidusandHolo cnemus
plucheihaveworldwidedistributionsandaree xcluded.
WithinSmeringopinae,somegenera(clade5a )shareweb
ornamentsmadeofnumeroussmallsilkballs (notall
individualswithinaspeciesmakethesesilkb alls,butthe
exactconditionsunderwhichwebornamen tsarebuiltare
controversial:Hajer&R
ˇ
eha ´kova´2003;Japyassu´&Macagnan
2004;.),andwithin thisgroupthe
generainclade5bsharespottedlegs.With56speciesineightgenera,Smeringopinaeisarathe r
smallsubfamily,andSmeringopusiscurrentlytheon lygenus
netinaeand
Arteminae,Smeringopinaear eoftenfoundinratherarid
peciesarewidespreadsyna nthropes(Smer-
ingopuspallidus[Blackwall1858],C rossoprizalyoni[Blackwall
1867],Holocnemuspluch ei[Scopoli1763]),andmostofwhat
weknowaboutthebi ologyofSmeringopinaerelatestooneof
thesethreesp ecies(Jackson1992a,1992b;Jacksonetal.1992,
1993 ;Huber1995;Jakob1991,1994,2004;Jakob&Dingle
199 0;Blanchongetal.1995;Kaster&Jakob1997;Sedey&
Ja kob1998;Johnson&Jakob1999;Jakobetal.2000;
Stric kmanetal1997;Calbacho-Rosaetal.2010;-forother
s peciesseeSenglet2001;Hajer&R
ˇ
eha´kova´2003 ;Huber
2009).Theoriginaldistributionofthesubfam ilyisAfrica,the
Mediterranean,andtheMiddleEast( Fig.27).
Composition:CenemusSaaristo2001;Cerato pholcus
Spassky1934;CrossoprizaSimon1893;Holocn emusSimon
1875;HoplopholcusKulczyn
´ski
1 908;SmeringopinaKraus
1957;SmeringopusSimon1890 ;StygopholcusKratochvil1932.
1850
Typegenus. —PholcusWalckenaer1805(bymonotypy).
Pholcidae:1 850:31(InternationalCodeof
ZoologicalNomenclatu re,Article36:PrincipleofCoordi-
nation;Internat ionalCommissiononZoologicalNomen-
clature1999).
‘‘Pholcus-group’’:Huber1995:298.
THEJOURNAL OFARACHNOLOGY
ThemonophylyofPholcinaeiswellsupp ortedby
eliceraeusually
areprovidedwithapair ofproximallateralapophyses
(Fig.2).Inaddition,t hetarsalcombhairsarereducedtoa
singleventralrow (Huber&Fleckenstein2008:figs.19,30).
Moleculard ataareavailableforonlyalimitednumberof
genera,b utsofartheresultsarelargelycongruentwiththose
f rommorphology(Bruvo-Madaric
´
etal.2005;Astr inetal.
2007).AllPholcinaeexceptZatavuaHuber200 3andNyikoa
Huber2007(clade6binFig.1)shareanapop hysisonthe
malepalpaltrochanter(Figs.3,4)thatin teractswiththe
proximallateralcheliceralapophys isduringcopulation
(Huber1995,2003a).Zatavuaand Nyikoa(clade6a)share
apeculiarshiftofthemalepal paltibia-tarsusjoints(the
retrolateraljointismo vedtowardsdorsal,theprolateral
jointtowardventr al,sothatbothjointsarevisiblein
prolateralview: Fig.7)(Huber2007).Membersofalarge
groupofgenera includingSpermophoraHentz1841and
MetagoniaSimon 1893(clade6c)shareadorsal(ratherthan
prolateral )attachmentofthebulbtothetarsus(Fig.3).
Withint hisgroup,fivegeneraandAfrican‘Spermophora’
(cla de6d)sharethereductionofpiriformglandspigotson
theanteriorlateralspinnerets(compareFigs.14and15;< br>Huber2003a,b,2005b).Anothermajorgroup(clade6e)< br>sharesastrongandlargescleriteconnectingthegenit albulb
tothetarsus(Fig.4;Huber2003c).Withinthis group,
Pholcusanditsclosestrelatives(clade6f)ar echaracte-
rizedbysimplifiedtarsus4comb- hairs(Fig.13;Huber
&Fleckenstein2008)andaknob-l ikestructureonthe
epigynum(Fig.23).Relationship swithinthisgroupare
difficulttoresolve(Huberinp ress),butLeptopholcusSimon
1893andMicromerysBra dley1877(clade6h)areverylikely
closerelatives(i nagreementwithSimon1893,butcontrary
toTimm1976) .Theyshareaderivedmalecheliceral
armature(later alapophysesverydistallyandnofrontal
modificatio n)andaserratedtipofthepalpaltrochanter
apophysi s(Fig.20;Huberinpress).
Withcurrently548species in24genera,Pholcinaeisthemost
species-richsubfa milywithinthePholcidae,includingthe
largestgenu s(Pholcus,currently174species)andseveralother
s pecies-richgenera(e.g.,MetagoniaandBelisanaThorell 1898).
Inasense,PholcinaeistheOldWorldcounterpa rtto
Modisiminae,withhighestdiversityinthehumid tropicsand
subtropicsandalargevarietyofbodyform sreflecting
bodyofliterature
dealswiththebio logyofthecosmopolitanPholcusphalan-
gioides(Fue sslin1775)(e.g.,Kirchner1986;Jackson&
Brassingt on1987;Kirchner&Opderbeck1990;Uhl1993,
1996,199 8;Uhletal.1995,2004;Scha¨fer&Uhl2002;Schaefer
& Uhl2003;Japyassu´&Macagnan2004;Scha¨feretal.2008),
butlittleisknownaboutthebiologyofotherspecies( Deeleman-
Reinhold1986;Huber1997a,2002;Senglet2 001,2008;Huber
&Wunderlich2006;Huber&Schu¨tte20 09;Chen&Li2005a,
2005b;Chenetal.2008;Xiaoetal.2 009,2010).Mostgeneraare
restrictedtotheOldWorld ,withthenotableexceptionofthe
NewWorldendemicMe tagoniaandafewrelictspecies
currentlyassignedto PholcusandLeptopholcus(Huber2000,
inpress;Huber etal.2005)(Fig.28).Somesynanthropicspecies
have attainedworldwidedistributionsorextendedtheirrange s
toanothercontinent(Pholcusphalangioides,Sperm ophora

HUBER—PHOLCIDPHYLOGENY
219
Figure28.—anthropicPholcusphalangioidesandSpermoph orasenoculata(bothcosmopolitan),
Micropholcusfa uroti(pantropical)andPholcusmanueli(introducedtoNo rthAmerica)areexcluded.
senoculata[Duge`s1836], Micropholcusfauroti[Simon1887],
PholcusmanueliG ertsch1937).
Composition:AetanaHuber2005;Anansu sHuber2007;
BelisanaThorell1898;BuitingaHuber20 03;CalapnitaSimon
1892;KhorataHuber2005;Leptoph olcusSimon1893;Metago-
niaSimon1893;MicromerysB radley1877;Micropholcus
Deeleman-Reinhold&Prins en1987;NyikoaHuber2007;
OssinissaDimitrov&Riber a2005;PanjangeDeeleman-
Reinhold&Deeleman1983;P aramicromerysMillot1946;
ParaspermophoraWunderl ich2004;PehrforsskaliaDeeleman-
Reinhold&vanHar ten2001;PholcusWalckenaer1805;
QuamtanaHuber200 3;SavarnaHuber2005;Spermophora
Hentz1841;Spermo phoridesWunderlich1992;UthinaSimon
1893;Wanniya laHuber&Benjamin2005;ZatavuaHuber2003.
DISCUSSI ONANDOUTLOOK
Thenumerouslargepolytomiesinthecla dograminFig.1
showthatmanynodeswithinthePholcid aeremainunresolved.
Inaddition,somenodesareweak lysupportedandareinneed
evelofsubfamilies,gener iccomposition
maychangeinseveralcases:Ninetinae maynotcontainclade2b
(i.e.,Aucana,Nita,Chisosa) ;ArteminaemaycontainWugigarra
andbeparaphyletic withrespecttoModisiminae;and
Smeringopinaemaybe paraphyleticwithrespecttoPholcinae.
Similarprob lemsoccuratthelevelofgenera,andsome
terminaltax aaspresentedinFig.1maynotevenbemonophy-
especia llytrueofthegeneraBelisana(Huber2005),
Quamtana Huber2003(Huber2003c),MesabolivarandMeco-
laest husSimon1893(Huber2000),Spermophora(Huber2005b),LeptopholcusandPholcus(Huberinpress).
Atthele velofspecies,manyhundredsoftaxaremain
undescrib ed,andthespeciesnumbersabovemaynoteven
mple,are centrevision
ofthegenusBelisanahaselevatedspeci esnumbersfromtwo
to64(Huber2005a).Othergenerath atareknowntocontaina
largenumberofundescribedsp eciesareMetagonia
,Mesabo-
livar,Modisimus,S meringopina,andPholcus.
Manyhypothesescanbederi vedfromthecladogramin
Fig.1,andsomecanbeeasilyt estedasmaterialbecomes
mple,thecladogrampredict sseveral
charactersforthegenusTonoroGonza´lez-S ponga2009,none
ofwhichismentionedintheoriginald escription:1)exposed
tarsalorgan;2)complexcomb- hairsrestrictedtoapatchnear
thetipoftarsus4;3)r educedpiriformglandspigotsonthe
anteriorlateral spinnerets;4)reducedepiandrousspigots;5)
presen ceofamalepalpalcoxaapophysis;and6)aventral
male palpalfemoralapophysisthatisnotdirecteddistally.Ataphylogeneticlevel,alargescaleanalysisofmolecu lar
datawillmostlikelybenecessarytofurtherresol vepholcid
relationshipsandtoadvancepholcidsyste maticssignificantly.
Eventhoughsomeresultsfromr ecentmolecularworkappear
dubious,mostrelationsh ipssuggestedareeithercongruent
withmorphologica ldataorsuggestreasonablealternatives.
Theunexpl oredpotentialoftraditionalmorphologicalchar-
ac tersseemsverylimitedinthePholcidae,butsperm
ult rastructureandchromosomemorphologyareamongthe
n on-molecularcharactersmostpromisingtocontributetoo ur
understandingofpholcidphylogeny(Michalik&Uhl 2005;
Michaliketal.2005;Michalik&Huber2006;Oliv eiraetal.
2007;Ramalhoetal.2008;.).
ACKNOWLE DGMENTS
IthankMatjazˇKuntnerandMichaelRixforman yhelpful
commentsandsuggestionsforimprovementof aprevious
versionofthemanuscript.
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