Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions.

77 Pages·2001·3 MB·English

by  Ward, P. S.

InvertebrateTaxonomy,2001, 15,589—665
Taxonomy, phylogeny and biogeography ofthe ant genus Tetraponera
(Hymenoptera Formicidae) in the Oriental and Australian regions
:
Philip S. Ward
Department ofEntomology, University ofCalifornia, Davis, CA 95616, USA. Email: [email protected].
Abstract. A revision ofthe ant genus Tetraponera in the Oriental and Australian regions reveals 33 species (18
new), belonging to four informal species-groups: allaborans-gmup (T. allaborans (Walker), T. apiculata, sp. nov.,
T. avia, sp. nov., T. bita, sp. nov., T. brevis, sp. nov, T. conica, sp. nov, T. connectens, sp. nov, T. crassiuscula
(Emery) stat. nov., T. extenuata, sp. nov, T. microcarpa Wu & Wang, and T. modesta (F. Smith)); «/gra-group {T.
aitkenii (Forel), T. atra Donisthorpe, T. attenuata F. Smith, T. binghami (Forel), T. buops, sp. nov, T. difficilis
(Emery), T. inversinodis, sp. nov., T. laeviceps (F. Smith), T. mimula, sp. nov., T. nigra (Jerdon), T. nitida (F. Smith),
T. nixa, sp.nov, T. nodosa, sp. nov, T. notabilis, sp.nov., T.polita, sp.nov, T.punctulataF. Smith, T.rotula, sp. nov,
T. tucurua, sp. nov, T. vivax, sp. nov., and T. volucris, sp. nov);pilosa-growp (T.pilosa (F. Smith)), andrufonigra-
group {T. rufonigra (Jerdon)). Keys are provided for identification ofworkers, queens and males, although the
sexual forms remain unknown in some species. Phylogenetic analysis indicates that the four species-groups
represent independent lineages, each with its nearest extant relatives in the Afrotropical region. There have been
multiple invasions of Asia from Africa, and at least four west-to-east transgressions ofWallace's line into the
Australian region. Plate tectonic eventspostulatedto havebeen importantin facilitating such dispersal includethe
collisionofIndiawithAsiaintheEoceneandthe approachoftheAustralianplatetoLaurasiainthemid-Miocene.
Introduction species-groups are defined, and aspects oftheir phylogeny
andbiogeographyare explored.
The subfamily Pseudomyrmecinae is a group of slender- Early taxonomic work on the Indo-Australian
bodied, large-eyed, arboreal ants found throughout the Old Tetraponera consists of isolated species descriptions by
andNewWorldtropics.TheOldWorldspeciesbelongtothe Donisthorpe (1948, 1949), Emery(1889, 1900, 1901), Forel
genus Tetraponera F. Smith, containing approximately 110 (1902, 1903*, 1909Z;, 1912a, 1912*, 1915Z>), Jerdon(1851),
nominal species or subspecies (Ward 1990; Bolton 1995). Karavaiev (1933), Motschoulsky (1863), Roger (1863a),
Species of Tetraponera have piqued the interests of Smith (1852, 1858, 1859, 1860, 1863, 1877), Stitz (1925),
biologists studying ant-plant interactions (Hocking 1970; Viehmeyer(1916), andWalker(1859). Keys to specieswere
Janzen 1972;Ward 1991;Kleine/a/. 1992;Kleine?a/. 1993; given in Bingham (1903), Emery (1900) andForel (1903a),
Buschinger et al. 1994; Young et al. 1997; Djieto-Lordon but these now have little utility. Several recent papers have
and Dejean 1999; Palmer et al. 2000), ant defensive dealt with species of Tetraponera from China and adjacent
chemistry(Braekmane/a/. 1987;Merline?a/. 1988;Renson regions (WuandWang 1990; DlusskyandRadchenko 1990;
etal. 1994;Devijvere;a/. 1995)andgutanatomy(Billenand Radchenko 1997; ZhouandJiang 1997). The characteristics
Buschinger2001). Yetthere has beenno modemtaxonomic ofthe genus as awhole were summarisedinWard(1990).
treatment of the genus at the species level. This paper One conclusion to emerge from the present study is that
attempts to redress this problem for the Oriental and many of the names proposed for Tetraponera species
Australianregionsbyrevisingthe Tetraponera speciesfound representjuniorsynonyms, acircumstancethatarisesinpart
in this part of the world. Some of the results must be because relevant type material was not always examined by
considered provisional because of uncertainties about previous investigators. In addition, however, it is now
species boundaries. Nevertheless, it has been possible to apparent that some species of Tetraponera can show
circumscribe most taxa with reasonable clarity and to offer remarkable variation in size and shape, both within
worker-, queen- and male-based keys to the species. Four populations and over larger geographical scales. Thus,
)CSIRO2001 10.1071/ITOlOOl 0818-0164/01/05589
590 P.S.Ward
progressonthe species-leveltaxonomyoftheseantsrequires NHMB NaturhistorischesMuseum,Basel, Switzerland
theanalysisofgeographicallyextensivepopulationsamples, NHMV NaturhistorischesMuseum,Vienna,Austria
and an appreciation of the potential for species to show NTUC Department ofEntomology, National Taiwan University,
Taipei,Taiwan
substantialphenotypic variation. OXUM Hope Entomological Collections, University Museimi,
Most collections ofIndo-Australian Tetraponera consist Oxford,UK
of isolated workers, unassociated with sexual alates, and PSWC p. S. Ward Collection, University ofCalifornia at Davis,
often lackingbiological data. Ourknowledge ofthe ecology CA,USA
and behavior of these ants would be enhanced if more RMBR Raffles Museum of Biodiversity Research, National
UniversityofSingapore, Singapore
emphasiswereplacedontheprocurementofnestseries. This RMNH Nationaal Natuurhistorische Museum, Leiden,
would also allow the accumulation of worker-associated Netherlands
queens and males, whose characteristics may prove to be ROME RoyalOntarioMuseum, Toronto,Canada
more reliable for delimiting species. Males are known for SAMC SouthAfricanMuseum,CapeTown, SouthAfrica
only about halfthe species (and in small sample sizes for STMENRKC TSrtaoaptilciaclheEscoMsuysseteummsfRuersNeaatrucrhkuCnendter,e,KaCrSlsIrRuOh,e,WiGnenremllainey,
some of these) but the available data indicate that the
Australia
genitaliaprovide gooddiagnostictraits forsome species and UASK InstituteofZoology,UkrainianAcademyofScience,Kiev,
clades. Ukraine
UCDC BohartMuseumofEntomology, University ofCalifornia
atDavis, CA,USA
Materials and methods UCRC Entomology Research Museum, University ofCalifornia
atRiverside,CA,USA
Sourcesofmaterial UMSC Institute for Tropical Biology and Conservation,
bSepleocwi.mCeondsewsefroerpeuxbalimciniensdtiatnutdi/oonrsfdoelplooswittehdosienitnhAernceotltleecttialo.ns(1l9i9s3t)e,d USNM NUUnaSitviAeornsailtiMMuaslaeyusmiaoSfaNbaathu,rKalotHaisKtionrayb,alWua,shMianlgatyosni,a DC,
whereavailable. ZMAS Zoological Institute, Russian Academy of Science, St
AMNH American Museum ofNatural History, New York, NY, Petersburg,Russia
AMSA UAuSstAralianMuseum,Sydney,Australia ZMHB MBeurlsien,umGerfimiarnNyaturkunde der Humboldt Universitat,
ANIC AustralianNational InsectCollection, CSIRO, Canberra, ZMPA Institute of Zoology, Polish Academy of Sciences,
Australia Warsaw,Poland
ASIC AndreasSchulzCollection,Leichlingen,Germany ZMUM Zoological Museum, University of Moscow, Moscow,
BMNH NaturalHistoryMuseum,London,UK Russia
CASC CaliforniaAcademyofSciences,SanFrancisco,CA,USA
CESB Centre for Ecological Sciences, Indian Institute of Additionalabbreviationsused(specimensnotexamined):
Science,Bangalore,India CFRB ChineseAcademyofForestry,Beijing,China
CMNH CarnegieMuseumofNaturalHistory,Pittsburg,PA,USA GNUC Department of Biology, Guangxi Normal University,
CPDC Jacques Delabie Collection, CEPEC/CEPLAC, Itabuna, Guilin,Guangxi, China
Bahia,Brazil
CUIC CornellUniversityInsectCollection,Ithaca,NY,USA
DZUC Department of Zoology, University of Calicut, Kerala,
Analysisofmorphology
India
FRCK ForestResearchCentre,Kuching, Sarawak,Malaysia Toquantifyinterspecificdifferencesinsizeandshapeaseriesofmeas-
HNHM HungarianNaturalHistoryMuseum,Budapest, Hungary urementswastakenonarepresentativesampleofTetraponeraworkers
HZIC HerbertZettelCollection,Vienna,Austria (n=380),queens(n=92)andmales(n=41).Foreachapparentspecies
JDMC J. D. Majer Collection, Curtin University ofTechnology, orspecies-complex specimenswerechosenfromthroughouttherange
Perth,Australia ofthetaxonand,ingeneral,nomorethanoneindividualwasmeasured
KFBG KadoorieFarmandBotanicGarden,HongKong,China fromeachnestseriesorlocal collection. Thisrestrictionwasliftedfor
KUBC Faculty of Forestry, Kasetsart University, Bangkok, rarespeciesinwhichsampleswerelimitedtooneorafewcollections.
Thailand Measurementsweremadeat50x usingaWildMSAmicroscopeanda
KUEC Institute of Tropical Agriculture, Kyushu University, dual-axisNikonmicrometerwiredto adigitalreadout. Measurements
Fukuoka,Japan were recorded to the nearest 0.001 mm, but arepresented here to the
KUES Seiki Yamane Collection, Kagoshima University, second decimal place. For three frequently used metrics, HW (head
Kagoshima,Japan width),HL(headlength)andEL(eyelength),anestimateofprecision
LACM Natural History Museum of Los Angeles County, Los wasobtainedbytakingfiverepeatedmeasurementsatdifferenttimein-
Angeles,CA,USA tervalson10workerspecimensofTetraponerapunctulata.Thisyielded
MCSN Museo Civico de Historia Natural 'Giacomo Doria', estimates ofmean values ofthe average deviation from the mean of
Genoa,Italy 0.003 mm,0.005mm,and0.003 mm,respectively.
MCZC Museum of Comparative Zoology, Harvard University, The linear measurements and indices employed in this study are
Cambridge,MA,USA describedbelow. The firstfourmeasurements aretakenwiththehead
MHNG Museumd'HistoireNaturelle,Geneva, Switzerland infull-face, dorsalview(headconsideredprognathous). Thisinvolves
MNHN MuseumNationald'HistoireNaturelle,Paris,France positioning the head so that its posterior margin and anterolateral
MZLU MuseumofZoology,LundUniversity,Lund, Sweden corners(abovethemandibularinsertions)areinthesameplaneofview.
Antgenus TetmponerainAsiaandAustralia 591
HW Head width: maximum width of head, including the PWI Petiolewidthindex: DPW/PL
eyes. Thefollowingquantitativeassessmentsofpilositywereused:
HL Head length: midline length ofhead proper, from the
posteriormarginoftheheadtotheanteriorextremityof CSC Cephalicsetalcount:numberofstandinghairs,i.e.those
the clypeus. When the anterolateral margins of the forminganangleof45°ormorewiththesurface(Wilson
clypeus have a greater anterior reach than the clypeal 1955), visible ontheposteriorhalfofthe dorsumofthe
midpoint, headlengthismeasuredtothemidpointofa head,as seeninlateralandposteriorviews.
linedrawnacrosstheanterolateralmargins. MSC Mesosomalsetalcount: numberofstandinghairsvisible
EL Eye length: length ofcompound eye, measured in the inprofileonthemesosomadorsum.
same view as HL. In taking this measurement it is
important to have adequate lighting to discern the Thecephalicsetalcountessentiallycoverstheareabetween,andposte-
margins oftheeye, sincethe outercircletofommatidia riorto, the middle ofthe compound eyes. Oftenthere is only a single
isusuallydarkenedandinconspicuous. pairoferectsetaehere,oneoneachsideoftheheadattheposterome-
MFC Minimum frontal carinal distance: minimum distance sialmarginoftheeye. This isreferredtoasthe supraocularpairofse-
betweenthefrontalcarinae. tae. InmostspeciesofTetmponerathestandinghairsonthedorsumof
SL Scape length: length of the first antennal segment, theheadandmesosomaareclearlydistinguishable fromthe appressed
excludingtheradicle(Fig. 1). pubescence,bytheirgreaterlengthandmoreerectappearance.Insome
LF2,LF3, Length of funicular segments 2, 3 and 4: maximum species,however,thestandinghairstendtogradeinto suberectandde-
LF4 measurable lengths of the second, third and fourth cumbent pubescence (e.g. Figs 65-68), and in such instances the ce-
funicular segments (thir4 fourth and fifth antennal phalicandmesosomalsetalcountsarenecessarilyapproximate.
segments),respectively. MandibulardentitionisanimportantcharacterbutitcanbedifficuU
FL Profemur length: length of the profemur, measured toassesswhenthemandibles aretightlyclosedagainstone anotheror
alongitslongaxisinposteriorview(Fig. 2). whentheteethareabraded. Moreover, insome Tetraponeraspeciesthe
FW Profemur width: maximum measurable width of the masticatory (chewing) and basal margins ofthe mandible meet at a
profemur,measuredfromthe sameviewasFL, atright rather oblique angle, and there may be difficulty in determining their
anglestothelineofmeasurementofFL(Fig.2). limits, and hence the number of teeth on each. The interpretation
PrWM Pronotumwidth:maximumwidthofthepronotumatthe adoptedhere is as follows: themasticatorymargin is consideredtobe
dorsolateral margins. In species in which the lateral composedofthoseteeth,beginningwiththeapicaltooth,whoseapices
marginsofthepronotumarenotwelldefine4PrWMis lie more or less in a straight line and whose basal-most tooth, the
measured at the point where the pronotal surface apicobasal tooth, lies along a straight-line extension ofthe proximal
becomesvertical. portion ofthe basal margin (Fig. 6). Thus, a line drawn between the
PDH Propodeumheight: heightofthepropodeum,measured apicobasal tooth and the origin of the basal margin more or less
in lateralview, fromthe base ofthemetapleurontothe parallels that margin. The apicobasal tooth is counted as part ofthe
maximum height of the propodeum, along a line dentition ofthe masticatory margin but not that ofthe basal margin.
MostAsianTetraponerahavefourteethonthemasticatorymargin(five
orthogonaltothelowermetapleuralmargin(Fig. 3).
MTW Metapleural width: maximum distance between the in T.rufonigra).Inthosespeciesthatareconsideredtohavethreeteeth
(allaborans-group)(Fig. 7)thereisanadjacenttooth,lyingonthebasal
metapleura,measuredindorsalview(Fig.4). marginofthemandible(asheredefined), thatmightwellrepresentan
PL Petiolelength: lengthofthepetiole,measuredinlateral
offsetfourthtoothofthemasticatorymargin.
viewfromthelateralflangesoftheanteriorpeduncleto The term mesopropodeal impression is used for the transversely
theposteriormarginofthepetiole (Fig. 3). depressed region on the mesosoma dorsum between the mesonotum
PH Pinetiloalteerhaeligvhti:ewmaaxtimriughmtheaingghltesofttohePpeLtio(lFei,g.me3a)s,urbeudt agrnodovteh'e.pIrnoTpeotdreaupmo.nerTahiwsoriksecrso,mmhoownelvyerr,eftehrerreedctaonabsetahesma'lmle,tarnaoitsaeld
excluding any protruding teeth or lobes at the welt-like structure inthe middle ofthis impression(Fig. 44),whichis
anteroventralorposteroventralextremitiesofthepetiole here interpreted as the metanotum since it is flanked by the
DPW D(eo.rgs.aFligpset5i6o,le82w).idth: maximum width of the petiole, mMeatlaatghaosryacaincdsApfirriacclaens.speIctieiss opfarTteitcrualpaornlyeraw,elblutdaelvseoloocpceudrsiinnasofmeew
measuredindorsalview. Asiantaxa.
LHT Metatibialength: lengthofthehindtibia,excludingthe ThetermsforintegumentsculpturefollowHarris(1977). Sculpture
proximomedial part ofthe articulation that is received isbestobservedwithsoftlighting,whichcanbeachievedbyplacingan
into the distal end of the metafemur (Fig. 5). This opaquefilter(e.g. Mylarplastic)betweenthe specimenandthe source
measurement is taken with the extensor surface ofthe ofillumination.
tibia in full view, so that the line ofview corresponds
withtheplaneoftibialflexion. Inferringspeciesboundaries
CI Cephalicindex: HW/HL
REL Relativeeyelength: EL/HL Torhgeanivsimesw,saudcohptaesdTeitnrapthoinserap,apsepreciiesstahraetmoinstsuesxeufaullllyycroenpcreoidvuecdinags
REL2 Relativeeyelength,usingHW: EL/HW groups ofpopulations linkedtogetherbyrecentorongoing gene flow
FCI Frontalcarinal index: MFC/HW and having inherited characteristics that cause them to be
SI Scapeindex: SL/HW reproductively isolated from other populations (Mayr 1942). This
512 Scapeindex,usingHL: SL/HL biological species concept motivates the search for discrete
513 Scapeindex,usingEL: SL/EL morphological gaps that indicate the occurrence of reproductive
FI Profemurindex: FW/FL boundaries. The resulting inferences about species limits are working
PDI Propodealindex:PDH/MTW hypotheses, subject to further testing with additional morphological
PLI Petiolelengthindex: PH/PL characters, genetic markers, or direct behavioural observations. In
592 P.S.Ward
PDH
4 7
MTW
LHT
mm mm
1.0 0.5
Figs 1-7. Workermorphology, Tetraponerapunctulata (1-6) and T. allaborans (7), illustrating various measurements (SL, FL, FW, PDH, PL,
MTW
PH, andLHT:seetext)andmandibulardentition, 1,scape,dorsalview;2,profemur,posteriorview;3,metapleuron,propodeumandpetiole,
lateral view; 4, mesopleuron, metapleuron andpropodeum, dorsal view; 5, metatibia, lateral (external) view; 6, 7, mandible, dorsal view, with
masticatorymarginbracketed. Scalebar= 1.0mmforFigs 1-5 and0.5mmforFigs6-7.
many animal taxa the male genitalia exhibit species-specific Pseudomyrmecinae (Ward 1991) was reanalysed, restricting it to
characteristics(Eberhard1986).ThisappearstobetrueofTetraponera, species in the genus Tetraponera and one outgroup taxon,
andwhere male specimens were available I used features ofthe male Pseudomyrmex gracilis. The result was a dataset of 88 characters,
genitaliatorefine decisions about species limitsbasedonworkerand drawn from the set of 125 characters listed in Ward (1991), but
queenmorphology. excluding 37 characters that became invariant forthis more restricted
Not all observed morphological discontinuities indi—cate species setoftaxa. This exemplardata setcontained 16representative species
boundaries.Theymaybeartifactsofinsufficientsampling especially of Tetraponera, seven from the Indo-Australian region and the
where th—e only available specimens are from geographically distant remainder from the Afrotropical region. The Indo-Australian species
locations or they may reflect the existence ofdiscrete intraspecific included examples from all four species-groups recognised in the
polymorphism. Bearing in mind that Tetraponera has not been presentstudy.
intensivelysampledinmostareas,Ihavetakenaconservativeapproach To examine in more detail the history of one species-group in
towardsspeciesdelimitation. Someofthepolytypicspeciesrecognised Southeast Asia and Australia, a new set ofmorphological characters
in this study, such as T allaborans and T. punctulata, encompass a was developedforthe nigra-gxowp. This datasetcovered 16 ofthe 20
considerabl—e range of phenotypes, her—e interpreted as geographical known species (fourtaxabeingtoopoorly sampled). Alsoincludedin
variants or where they are sympatric as intrapopulation variation, the datasetwerethree otherAsian Tetraponera (one fromeach ofthe
perhapsecotypic innature. Carefulgenetic andecological studieswill other three species-groups), and two Afrotropical species of
beneededtodemonstratewhetherthevariationin suchpolytypic taxa Tetraponera(T.clypeata, Tnatalensis).TwospeciesofPseudomyrmex
represents different species, intraspecific polymorphism, or an (P.gracilis,P.termitarius)wereusedasoutgroups. Totestthestatusof
intermediate situation involving assortative mating and partial (but the «/gra-group, it was not constrained to be monophyletic. The
incomplete) reproductive isolation. Of course, for closely related morphologicalcharactersandtheiralternatestatesareasfollows.
populations that are strictly allopatric and partly differentiated there
maybenonon-arbitraryresolutionofspeciesstatus. (1) Worker,basalmarginofmandible(0)subequalto,orshorterthan,
masticatory margin (Fig. 6); (1) longerthan masticatory margin
(Fig. 7)
Phylogeneticanalysis
(2) Worker,numberofteethonmasticatorymarginofmandible(0)3;
ForthepurposesofassessingwhethertheIndo-AustralianTetraponera (1)4;(2)5;(3)>5
faunacomprisesamonophyleticgrouporapolyphyleticassemblage,a (3) Worker,proximaltoothonbasalmarginofmandible(0)present;
morphological data set developed for the entire subfamily (1)absent
1
593
Antgenus TetraponerainAsiaandAustralia
(4) WoAer,withheadin full-faceviewandmandibles closed, dorsal (26) Worker, queen, mesosternum (0) mostly smooth and shiny; (1)
abductorswellingofmandible(0)visible; (1)notvisible,covered denselypubescent
bytheanterolateralmarginofclypeus (27) Worker, standingpilosityonmesosoma(0) sparse, MSC0-8; (1)
(5) Worker,labrum,proximalhalf(0)lackingteethorprotuberances; common,MSC>8
(1) with a single median tubercle; (2) with median tubercle, (28) Worker, queen, posteroventral margin ofpetiole (0) unmodified:
flankedbyalateralpair;(3)withlateralpairoftuberclesonly forming ventral part of structure against which helcium
(6) Worker, median portion ofclypeus, ventral surface adjacent to articulates; (1) modified as a ventral, protruding hood, which is
junction with labrum (0) lacking transverse carina; (1) with separatedfromthesurfaceagainstwhichthehelciumarticulates
transversecarina (29) Worker, queen, mid-posterior portion of petiolar sternite, as
(7) Workerhead(0)relativelybroad,CI >0.78; (1)moreelongate,CI viewed in profile (0) flat to weakly convex; (1) strongly convex
<0.78 (Figs94-96)
(8) Worker, compound eye (0) small to medium-sized, REL2 0.32- (30) Worker, queen, petiole (0) lacking posteroventral teeth; (1) with
0.53; (I)relativelylarge,REL2 >0.54,andfallinginuppercloud posteroventralteeth(Figs79-84)
ofpointsinbivariateplotofELv. HW(Fig. 110) (31) Worker, petiole, in lateral view (0) relativelyrobust, PLI usually
(9) Worker,eyelengthrelativetoLHT(0)small,EL/LHT<0.42; (1) >0.52(butvaryingwithbodysize: seeFigs 119-120);(1)slender
(10) mWoerdkieurm,-leayrege(,0)EeLlo/nLgHatTe,0.>412.-50x.8l8o;ngseeretahlasnowFiigd.e;11(1)oval,<1.5x aponidntasttiennubaitvea,riPaLteIpulsoutaslloyfP<0H.5v2.,PaLndanfdalPliLnIgvi.nHloWwe(rFicglsou1d19o-f
longerthanwide 120)
(11) Worker,scapelengthinrelationtoHW(0)short,SIusually<0.60; (32) Worker,petiolelengthinrelationtoheadlength(0)morethanhalf
(1) longer, SI >0.60, and falling in upper cloud of points in HL (PL/HL 0.54-0.92); (1) shorter, about one-half HL or less
bivariateplotofSLv. HW(Fig. 112) (PL/HL usually _0.52, and falling in lower region ofpoints in
(12) Worker, scape length, inrelationto EL (0) short, S13 0.83-0.98; bivariateplotofPLv. HL; seeFig. 121)
(1)medium,SB 1.02-1.54;(2)long,SI3 >1.55; seealsoFig. 113 (33) Worker,queen,pubescenceonabdominaltergiteIV(0)consisting
(13) Worker, head (0) with three distinct ocelli; (1) with two distinct ofarelativelydensematoffine, overlappinghairs; (1)relatively
ocelli, the third (median) ocellus weak or absent; (2) lacking sparse,hairsscattere4 separatedbytheirlengthsormore
distinctocelli (34) Workersize(0)smaU-medium(HW<0.95);(1)large(HW>0.95)
(14) Worker,queen,head(0)denselypunctate,lackingextensiveshiny (35) Male, head (0) notably broader than long (CI >1.06); (1)
interspacesbetweenthepunctures;(1)lessdenselypunctate,with approximatelyaslongasbroad,orlonger(CI<1.06)
conspicuousshinyinterspacesbetweenthepunctures (36) Male, antenna (0) moderately long, its length subequal to, or
(15) Worker, queen,puncturesondorsumofheadbetweencompound greater than, mesosoma length; (LF2 + LF3 + LF4)/HL 0.80-
eyes (0) coarser, mostly 0.010-0.020 mm in diameter and 1.40;(1)veryshort,lengthofantennalessthanmesosomalength;
s0e.p0a1r5atmemdbiynadbioaumtettheeriranddiaumseutaelrlsyosrelpeasrsa;t(e1d)bfyinesre,vemroasltdliya0m.e0t0e5r-s (37) M(aLlFe2,+scLaFp3el+eLnFg4th)/reHlLataivpeprtooxhiemaadtewliydt0h.5(00)smaller,SI<0.35;(1)
(16) Worker and queen, standing pilosity on posterior half of head larger, SI~0.36
dorsum (0) sparse: 0-8 hairs, usually arranged in pairs; (1) (38) Male, scape lengthrelative to head length (0) larger, S12 ~ 0.30;
common: 10ormorehairsscatteredacrossupperheadsurface (1)smaller, SI~0.20-0.27
(17) Worker, queen, pronotal humeri as seen in dorsal view (0) (39) Male, length ofsecond funicular segment (0) notably less than
subangulate(Figs59-60); (1)rounded(Figs30-36) combinedlengthofscapeandfourthfunicularsegment(LF2/(SL
(18) Worker,profemur(0) slender, FIusually<0.40; (1)broader, FI> +LF4)0.40-0.72); (1)aslongas,oralmostaslongas,combined
0.42, and/orfallinginleftupperregions inbivariateplotsofFW length ofscape and fourth funicular segment (LF2/(SL + LF4)
V. FLandFIv. HW(Figs 114-115) 0.82-1.00)
(19) Worker,profemurlength,relativetoHL(0)short,FL/HLusually (40) Male,mesoscutum,asseenindorsalview(0)narrowinggradually
<0.65; (1) longer, FL/HL usually >0.65, and falling in upper towards anterior end; (1) broadened posteriorly, anterior part
regioninbivariateplotofFLv. HL(Fig. 116) suddenlyandstronglyconstricted
(20) Worker, mesopropodeal impression (0) short and transverse, (41) Male, mesoscutum (0) denselypunctate-reticulate, thepunctures
widthofimpression(measuredasdistancebetweenthemetanotal subcontiguous and the integument subopaque; (1) less heavily
spiracles) four or more times its length; (1) longer and sculptured, with scatteredpunctures separated by one to several
subquadrate, width of impression (distance between metanotal diameters, andinterspaces sublucid
spiracles)notmorethanaboutthreetimes itslength (42) Male, hypopygium(sterniteIX) (0) semicircularto subtriangular
(21) Worker, mesopropodeal impression (0) lacking pit-like inshape(Figs 126, 128-133); (1) subrectangular(Fig. 127)
depression;(1)withapit-likedepressionposteriorly,precededby (43) Male, hypopygium(sternite IX), anterolateral extremity,position
an open, transverse strip of integument; (2) with a pit-like inrelationtomedianapodeme(0)posterior; (1)anterior
depression,occupyingmoreorlesstheentireimpression (44) Male, hypopygium (sternite IX), anterolateral arms (0) simple
(22) Worker,dorsalfaceofpropodeum(0) flattenedorweaklyconvex (Figs 126-128); (1) subtended by a thin, weakly sclerotised,
in profile (Figs 94-99); (1) moderately to strongly convex in lamellateextension(Figs 129-133)
profile,decliningposteriorly(Figs91-93) (45) Male,hypopygium(sterniteIX),posteromedial sectionthatbears
(23) Worker, metatibia (0) relatively long, LHT/HL 0.80-0.97; (1) mostsetae(0)moreorlessflat; (1)depresseddorsad
short,LHT/HL0.58-0.79; seealsoFigs 117-118 (46) Male, hypopygium (sternite IX), posterior margin, as seen in
(24) Worker, queen, metabasitarsal sulcus (0) absent; (1) present, ventral view (0) convex (Figs 128-129, 133); (1) straight or
simple; (2)present,subtendedbyaraisedcarina weakly (broadly) concave (Figs 126, 127); (2) strongly and
(25) Worker, queen, anteriorquarterofpronotum (0) lacking a dense narrowlyconcave(Figs 130-132)
patch of punctation, that contrasts with sparser punctures (47) Male,hypopygium(sterniteIX),posteromedialextremity,asseen
elsewhereonthepronotumandondorsumofhead; (1)withsuch inventralview(0)lackingsmall,thin,ligulateprotrusion;(1)with
sculpture suchaprotrusion, extendedposteriorly(Fig. 129)
594 P.S.Ward
(48) Male,hypopygium(sterniteIX),posteromedialmargin,asseenin forming an angularjunction atmidpoint (Fig. 140), and slightly
posteriorview(0)relativelythin(Fig. 129);(1)notablythickened weakenedhere
(Figs 130-133) (70) Male, aedeagus, oblique carina on posterodorsal extremity of
(49) Male,hypopygium(sterniteIX),posteromedialmargin,asseenin external face (0) absent (Fig. 139); (1) present (Figs 137-138,
posterior view, thin lamellate extension (0) absent; (1) present, 140-141)(hereaftertermed 'upperobliquecarina')
depresseddorsad;(2)present,depressedposterodorsad (71) Male,aedeagus,upperobliquecarina,ifpresent(0)situatedmuch
(50) Male,hypopygium(sterniteIX),posteromedialmargin,asseenin closertoposterodorsalmarginthanto lowerobliquecarina(Figs
posterior view, thin lamellate extension (0) absent; (1) present, 140-141); (1) situated about midway between lower oblique
broadlytransverse,notprotrudingstrongly(Fig. 131);(2)present, carinaandposterodorsalmargin(Fig. 137)
narrower,obviouslyprotruding(Fig. 130, 132) (72) Male,queen,typicalnumberofforewingcubitalcells(0)2; (1) 1
((5521)) MMmgiaradallldeeiu,,naelp,plaayrartaafmamreeorsrmheea,,mripidliilymnienrnreoerpurn(pFodrixeogid.xmiaa1mnl6ag7lld)eo;rd(s(oFa1ri)lsgasdmlia1vrm6ega8rir-gnig1ni7(g8n0))s(un0do)dtednsiluvydedrfegrniolnymg uusnneooqrruIddeneenrtrceheeded.lagisbaTtovhevedeesaribdemoaimvtleaaa.irnsAeritenssgucelhptcaashr,raaarbctautectetrawesnriastl5,hywss4iel9sri,egihn5tt0lwryheaairnteceddhdu5acal4esldcworhereardsreeoarlceuttdtre,ieroasnit.newdeFtrohaeres
directed lateroventrally and passing below inner distal dorsal
margin; (1)ofsuchaform bsoeatrhcthhees8fo8r-cthhaermacotsetrp(a1r7s-itmaoxnoin)ouasntdre7e2s-cwhearreacctaerrri(e2d3o-uttaxuosni)ngdaPtAaUsPet*s
(53) Male, paramere, distal end (0) lacking digitiform processes; (1) (version4.0b4a)(Swofford2000),with 100randomadditionreplicates
withapairofdigitiformlobes(Fig. 154) and TBR branch swapping. Bootstrap analyses employed the same
(54) Male,paramere,distalend,asseeninlateralview(0)pointed;(1) heuristic search strategies, with 1000 bootstrap replicates. MacCIade
t(rFuingscat1e56a-n1d6r6o)unded(Figs 153, 155)(2)truncateandsubquadrate (version3) (MaddisonandMaddison 1992)wasusedtotracetraits of
interestonthetrees.
(55) Male,paramere,distalend,asseeninlateralview(0)notelongate
subrectangularandextendedposteroventrally(Figs 156-161);(1)
soshaped(Figs 162-166)
Geography
(56) Male, paramere, distal end in posterolateral view (0) lacking a
deep, obliquely transverse, dorsal impression; (1) with such an The expression 'Indo-Australian region' is used to refer to the area
impression formed by the combination of the Oriental and Australian
(57) Male,paramere,distalendindorsalview(0)notnotablynarrowed biogeographic regions. The boundary between these two regions
(Figs 167-170); (1)narrowingtowardstip(Figs 171-178) involves a transition zone in the area between Wallace's line and
(58) Male, paramere, inner distal margin, as seen in dorsal view (0) Lydekker'sline, i.e.betweentheeasternmarginofthe Sundashelfand
lackingbeak-likemesialprotrusion(Figs 171-176);(I)withsuch thewesternlimitoftheSahulshelf(Whitmore 1981;HallandHolloway
aprotrusion(Figs 168-170, 177-178) 1998). As used here the term 'Wallace's line' refers to the original
(59) Male,paramere, distalend, inner(mesial)surface, saucer-shaped versionoftheline(Wallace 1863),runningeastofBali,Borneoandthe
concavity (0) not visible in dorsal view; (1) partially visible in Philippines.
dorsalview(Figs 168-173); (2) fiillyvisibleindorsalview(Figs Inthe lists of'Materialexamined' therecords foreach speciesare
174-178) arranged alphabetically by country and by primary adminisfrative
(60) Male,paramere,mesiodistalconcavity(0)notextendingtodorsal divisionwithineachcountry. Administrative divisions are omittedfor
margin ofdistal exttemity ofparamere; (1) extending to dorsal Brunei, Bhutan, the Seychelles and Singapore. Most collection data
marginofdistalextremityofparamere have been abbreviated to locality and collector, with the source
(61) Male, paramere, mesiodistal concavity (0) not visible or only collections indicated at the end of each list. A more complete
slightlyvisible inposteriorview(e.g. Figs 145, 152); (1) largely enumerationofthecollectiondataisavailableas 'AccessoryMaterial'
exposedinposteriorview(Figs 142-144, 147) on the Invertebrate Taxonomy webpage (http://www.publish.csiro.au/
(62) Male, paramere, mesiodistal concavity, posterior margin (0) journals/it/).Insomeinstancesthelocalitynamesonthespecimenlabel
mostlythick,non-franslucent; (1)thin, semi-translucent have been emended forclarity, withthe original orthographygiven in
(63) Male, paramere, mesiodistal concavity, posterior margin (0) quotes (e.g. Bengkulu [as 'Benculen']). The abbreviation 'c.u.'
sfraight orbroadly concave, as seen in posterior view (e.g. Figs signifiesthatthecollectorcouldnotbedetermined.
145, 152); (1)narrowly concave, as seeninposteriorview(Figs Coordinates(latimdeandlongitude)weredeterminedforalmostall
142-144, 148-149) collection sites, except those with imprecise locality data (such as
(64) Male, volsella, distal end (0) free from paramere; (1) fused to 'Borneo', 'Sumatra' and 'N. Guinea'). The shareware program
innerfaceofparamere Versamap (Version 2.05) was used to plot species distributions. The
(65) Male, aedeagus, lateral apodeme (0) shorter than anterior following references provided useful information about old locality
apodeme (Figs 135-136); (1)as longas, orlongerthan, anterior names not found in modem gazetteers or atlases: Andersson (1913),
apodeme(Figs 134, 137-141) Beccari (1921), Broersma (1916), Chapman and Chapman (1947),
(66) Male, aedeagus, single posteroventral tooth or spine (0) absent Dohrn(1898),Fea(1896),Fruhstorfer(1903),Gressitt(1936a, 19366),
(Figs 134-136);(1)present(Figs 137-141) Karny (1922), Mantero (1903), McClure (1929, 1934), Mjoberg
(67) Male, aedeagus, posteroventral extremity, small lamellate (1930),Modigliani(1892), Soderberg(1919)andWallace(1890).
protrusion anteriorto posteroventral tooth (0) absent (Figs OS-
MO);(1)present(Figs 137, 141)
(68) Male, aedeagus, external face, arched carina originating Synopsis ofthe genus TetraponeraF. Smith 1852
aabnsteenrto;ven(t1r)alplryesaenndte(nFdiignsg1a3t7o-r14n1e)ar(hpeorsetaefrtoevrentterarlmetdoot'hlow(e0r) Tetmponem is one of three genera in the ant subfamily
obliquecarina') Pseudomyrmecinae. For a foil listing ofgeneric synonymy
(69) Male, aedeagus, lower oblique carina, if present (0) broadly seeWard(1990)orBolton(1995). Theworkersofthisgenus
arched, well developed throughout (Figs 137-139, 141); (1) havethe following characteristics:
..
Antgenus TetraponerainAsiaandAustralia 595
1 Mandible short, usually with 3^ (rarely 5-6) teeth on of the diversity is apportioned between two groups
the masticatorymarginand0-2 smallteethonthebasal (allaborans-growp, nigra-group), with the two remaining
margin;basalmarginlackingproximaltooth(Figs6-7). groups each containing a single, taxonomically isolated
2. Palpformula6,4(reducedto4,3 inoneAfricanspecies). species.
3. Posteromedial margin of clypeus not prolonged
backwardsbetweenthe frontal carinae. allaborans-group
4. Anterodorsal surface of median portion of clypeus allaborans(Walker, 1859: 375)
continuous, non-truncate (in confrast to =minuta(Jerdon, 1851; 112)syn. nov.(nomenoblitum)
Pseudomyrmex), its anterior margin with a line of =rufipes(Jerdon, 1851: 112)syn.nov.(nomenoblitum)
clypeal setae, and varying from straight and entire to =compressa(Roger, 1863a: 179)(synonymybyF.Smith, 1877:69)
protruding anddentate (e.g. Figs 8-18, 61-64). =femoralis (Motschoulsky, 1863: 21) (synonymy by Dalla Torre,
5. Mcoevdeirainnglomboestofofanttheenbnaaslalsccleornidteyleexpoafntdheedanltateenrnaallwyhaennd = ce1y8l9o3n:ic5a3)(Motschoulsky, 1863: 22) (synonymy by Dalla Torre,
1893:53)
head is observedin frill-face view. =subtilis(Emery, 1889: 500)(synonymybyDallaTorre, 1893: 53)
6. Antenna 12-segmented. =longinoda (Forel, \9Q9b: 394) syn.nov.
7. Scape short, lessthanthree-quarters ofheadlength(SI2 =allaboranssumatrensis(Emery, 1900: 676)syn. nov.
0.38-0.68 inOriental/Australian species). apiculata, sp.nov.
8. Frontal carinae relatively well separated, the minimum avia, sp.nov.
distance between them greater than basal scape width bita, sp. nov.
brevis, sp. nov.
(FCI 0.08-0.20 inOriental/Australian species).
conica, sp.nov.
9. Compoimd eye relatively large (REL 0.25-0.53 in connectens, sp.nov.
Oriental/Australian species), width ofeyetwo-thirds or crassiuscula(Emery, 1900: 677)stat.nov.
more oflength. extenuata, sp. nov.
10. Ocellipresent (usually 3) orabsent. microcarpaWu&Wang, 1990: 515
11 Pronotum andmesonotum not frised, freely articulating modesta(F. Smith, 1860: 106)
withone another. =fulva(Viehmeyer, 1916: 117)syn.nov.
=pisarskiiRadchenko, 1997:480syn.nov.
12. Mesopropodeal impression usually well marked,
sometimescontainingaplate-like sclerite, apparentlyof
nigra-gronp
metanotal origin.
13. Propodeal spiracle circular to elongate, located well aitkenii(Forel 1902: 245)
atraDonisthorpe, 1949:493
forwardonupperthirdofpropodeum. attenuataE Smith, 1877: 71
14. Metapleural glandwell developed, the opening directed = attenuata tenuissima (Emery, 1900: 675) (synonymy by Forel,
ventrolaterally or posterolateral^, and preceded by an 19126:54)
impression along the lower margin ofthe metapleuron, =birmana(Forel, 1902: 245)syn. nov.
whose dorsal margin is marked by an oblique =thagatensis(Forel, 1902:249)syn.nov.
longitudinal carina. binghami(Forel, 1902: 243)
15. Metabasitarsal sulcus nearly always present, reduced in =binghamilindgreeni(Forel, 1902: 245)syn.nov.
some species (absentinthree African species). buops,sp.nov.
dijficilis(Emery, 1900: 677)
16. Meso- and metatibiae each with a pair of the apical =nitens(Stitz, 1925: 117)syn.nov.
spurs, the posterior spur well developed and pectinate, =dilatata(Karavaiev, 1933: 267)syn.nov.
theanteriorspursmaller, sometimesveryreduced. =stipitum(Forel, \9\2b: 54)syn.nov.
17. Terga and sterna ofabdominal segments II (petiole), III inversinodis, sp.nov.
(postpetiole) and IVnotlaterallyfused. laeviceps(E Smith, 1859: 145)
=humerosa(Emery, 1900: 674)syn.nov.
18. Postpetiole distinctlydeveloped.
=dentifera(Karavaiev, 1933: 266)syn.nov.
19. Pupanaked. =platynota(Karavaiev, 1933: 269)syn.nov.
20. Larva with trophothylax (food pocket) on ventral mimula,sp.nov.
surface ofthorax. mgra(Jerdon, 1851: 112)
For practical purposes workers of Tetraponera can be =atrataE Smith, 1852: 44(synonymybyDallaTorre, 1893: 54)
distinguishedfromthoseofallotherantsbythecombination ==pneitgiroalaktraaFm.aS(mFiotrhe,l,1817971:2a7:01s0y5n).snyonv..nov.
of well-developed postpetiole, short mandibles, large oval =nigrainsularis(Emery, 1901: 113)syn.nov.
eyes, andaflexible promesonotal suture. =nigrafergusoni(Forel, 1902: 248)syn.nov.
n/ftWa(F Smith, 1860: 106)
Synonymic list ofTetraponera species from the Oriental =carbonaria(F. Smith, 1863: 20)syn.nov.
andAustralian regions =hrevicornis(Emery, 1900: 675)syn.nov.
=difficilislongiceps(Forel, 1902: 247)syn.nov.
In this study, 33 species ofIndo-Australian Tetraponera are =siggi(Forel, 1902:246)syn. nov.
recognised(18new),belongingtofourspecies-groups.Most =sigginebulosa(Forel, 19036: 404)syn.nov.
596 P.S.Ward
=siggisetifera(Viehmeyer, 1916: 119)syn. nov. lengths or more; relatively small species, HW 0.49-0.93
=bidentata(Karavaiev, 1933:264)syn.nov. (allaborans-gTOup) 4
=bidentataangusticeps(Karavaiev, 1933: 266) syn.nov. Mandiblemorerobust,withfourteethonthemasticatorymargin,
=mq^M!Donisthorpe, 1948: 591 syn. nov. and0-1 denticles onthebasalmargin (Fig. 6); basalmargin of
=shankouensisZhou&Jiang, 1997: 72 syn.nov. mandible subequal to, or shorter than, masticatory margin;
nixa,sp.nov. posteroventralmarginofthepetiolecloselyassociatedwiththe
nodosa, sp.nov. helciumventer(e.g.Figs68,94),althoughitmaybeflankedby
notabilis,sp.nov. ventrolateral flanges (Figs 79-84); most of the mesosternum
polita,sp. nov. devoid of pubescence; abdominal tergite IV usually densely
HW
punctulataF. Smith, 1877: 72 pubescent;sizevariable, 0.63-1.48(«!gra-group) 4
=punctulatakimberleyensis(Forel, \9\5b: 37)syn.nov. Small, black species (HW 0.58-0.61, LHT 0.50-0.52), with
rotula,sp. nov. disproportionately small eyes (REL 0.32-0.34) (Fig. 13), short
tucurua,sp. nov. scapes (SI2 0.42-0.45) and broad profemur (FT 0.47-0.48);
vivax, sp.nov. pronotal dorsum rounding into sides, lateral margins poorly
volucris,sp. nov. developed (Fig. 24); mesopropodeal impression lacking a
distinct metanotal plate but may be bisected by a weak
p/Zoifl-group transverse ridge that interrupts the longitudinally rugulate
pilosa(F, Smith, 1858: 160) sculpture(Thailand) T.connectens,sp.nov.
=nicobarensis(Forel, 19036: 402)syn.nov. Size, color, scapesandprofemurvariablebutifsmall(HW<0.65
and LHT <0.55) and black, then eyes larger (REL 0.35-0.41);
rufonigra-groxxip either lateral pronotal margins better developed or
mesopropodeal impression with a distinct, flattened metanotal
rufonigra(Jerdon, 1851: 111)
=rufonigrayeensis(Forel, 1902: 248)syn. nov. plate 5
=rufonigratestaceonigra(Forel, 19036: 402)syn.nov. Mesopropodeal impression with irregular longitudinal rugulae,
=rufonigraceylonensis(Forel, 19096: 394)syn.nov. interruptedbya small, raisedtransversewelt(metanotalplate),
whichisboundedlaterallybythemetanotalspiraclesandwhich
lacks rugulate sculpture (Fig. 44); pronotum lacking distinct
lateral margins, the dorsum rounding gently into the sides, as
Tetraponera ofthe Oriental andAustralian regions: key seen inposteriorview(Figs20-23); profemurshortandbroad,
to species, based on theworker caste FI0.44—0.53;darkbrowntoblackspecies 6
Mesopropodeal impression with irregular longitudinal rugulae,
This key excludes T. vivax and T. volucris, two species sometimes crossed at the midpoint by a broken transverse
knownonly fromthe queencaste. rugule, but lacking a raised metanotal plate; pronotum with
1. Head with three distinct ocelli; in dorsal view pronotal humeri more orless distinctlateral margins, whichvary from sharp to
appearingsubangulate(Figs59-60);headdenselypunctate,and blunt-edged; in posterior view pronotal dorsum meeting the
lackingextensiveshinyinterspacesbetweenthepunctures;large sides at a sharply rounded angle (Figs 19, 25-29); profemur
species,HW 1.14-2.07 2 usuallymoreslender(Fl0.36-0.48);colorvariable 9
- Head almost always lacking ocelli, very rarely with two orthree Median clypeal lobe subtriangular, protruding, and pointed
faintocelli(inafewlargeworkersofZnigraandT.punctulata); (Fig. 9);petiolenarrowindorsalview(DPW/MTW0.61-0.64)
pronotalhumerivaryingfromnarrowlytobroadlyrounded(e.g. (Borneo) T.apiculata,sp.nov.
Figs 30-36), but not subangulate; head usually less densely Median portion of clypeus broadly convex, not prominently
punctate and with conspicuous shiny interspaces between the protruding andpointed (Figs 10-12); petiole broader in dorsal
punctures (always the case in species with HW >1.10); size view(DPW/MTW0.65-0.73) 7
variable(HW0.49-1.48) 3 Largerspecies,withbroadhead(HW0.82,CI0.92)andlargeeyes
2. Larger species (HW 1.62-2.07), with smaller eyes (REL2 0.35- (REL 0.42); anterior clypeal margin edentate and non-
0.37) (Fig. 55); usually bicolored, the dark head and gaster protruding (Fig. 10); petiole slender (PLl 0.51) (Fig. 45);
contrasting with the orange-brown mesosoma (the latter mesopleuron extensively longitudinally carinulate (West
infuscated in some populations); standing pilosity common on Malaysia) T.avia,sp.nov.
the mesosoma dorsum, including the propodeum, MSC 20-66 Smaller, withmore elongatehead(HW 0.63-0.75, CI 0.73-0.84)
(Fig.56)(PakistantosouthernChina,southtoSumatraandJava; and smaller eyes (REL 0.35-0.39); anteriormargin ofclypeus
introducedintotheSeychelles) T.rufonigra(Jerdon) withamodestlyprotrudingandcrenulatemedianlobe(Figs 11-
Smaller species (HW 1.14-1.51), with larger eyes (REL2 0.49- 12); petiole more robust (PLI 0.58-0.67) (Figs 47^8);
0.56)(Fig. 57); bodyunicolorous darkbrown; standingpilosity mesopleuronpredominantlysmoothandshining 8
sparse on the mesosoma dorsum, absent from the propodeum, Smallerspecieswithmoreelongatehead(HW0.63-0.65,CI0.73-
MSC 3-6 (Fig. 58) (Myanmar to Vietnam, south to Palawan, 0.75)(Fig. 11); eyelargerinrelationtoscapelength(SI3 1.19-
Borneo,SumatraandJava) T.pilosa(F.Smith) 1.21);petiolerelativelyslender(PLI0.58-0.62)(Borneo,?West
3. Mandibleslender,withthreeteethonthemasticatorymargin, and Malaysia) T.bita,sp.nov.
1-2 denticles on the basal margin (Fig. 7); basal margin of Largerspecieswithbroaderhead (HW 0.73-0.75, CI 0.82-0.84)
mandiblemuch longerthanmasticatorymargin; posteroventral (Fig. 12);eyesmallerinrelationtoscapelength(SI3 1.25-1.30);
margin ofpetiole in the form of a thin, ventrally protruding petiole shorter and more robust (PLI 0.63-0.67) (West
hood, which is distinctly separated from the helcium venter Malaysia) T.brevis,sp.nov.
(Fig.40) when the postpetiole is in its normal horizontal Small species (HW 0.60-0.64), with short scapes (SI 0.52-0.57,
position;mesosternumdenselypubescent;abdominaltergiteIV S12 0.41-0.44); median clypeal lobe bidentate (Fig. 18);
sparsely pubescent, the appressed hairs separated by their pronotum relatively narrow (PrWM/MTW 1.14-1.21), with
Antgenus Tetraponera inAsiaandAustralia 597
sharp, subparallelmargins,asseenindorsalview(Fig. 36),and see discussionin text; smaller species (HW 0.51-0.64) (north-
appearing rather flattened in posteriorview(Fig. 29); profemur eastIndiatoChina,southtoNewGuinea)..J.modesta(F.Smith)
broad(FI0.42-0.48)(China,Vietnam) 14. Larger species (HW 0.95-1.48), with long legs (LHT/HL 0.80-
T.microcarpaWu&Wang 0.97); standingpilositycommon,MSC 6-71 (usually>10) and
- Scapeslonger(SI0.57-0.68,SI20.45-0.57);medianclypeallobe CSC 10^0, the cephalichairs scatteredoverthedorsal surface
usuallywiththreeorfourteeth(Figs 8, 14-16),orlackingteeth oftheheadandoftengradingintoshorter,suberectpubescence;
altogether, rarely with a pair of well developed teeth; size mesopropodeal impression flanked laterally by raised
variable but iffalling within the above range then usually the prominences(containingthemetanotalspiracles)butotherwise
pronotalmarginsaresoft-edgedandconvexindorsalview(Figs more orless open, notboundedby lateralridges that enclose a
p3r0-o3n5o)tuamnmdortheecopnrvoefxemiunrpoisstemroiorreviselewnd(eFri;gsd1o9r,sa2l5-s2u8r)fa.c.e. 1o0f - Smpailtl-elirkespdeecpireess,soionna(vaesrhaaglel(owHpWit0p.r6e3s-e1n.t4i4n);T.ibfiHngWha>m0i.)92.,..t.h1e5n
10. Larger species (HW 0.62-0.93, usually >0.70); body standing pilosity less common (MSC 0-22, CSC 0-4) and the
predominantly black, although petiole, postpetiole and limb sparse cephalic hairs arranged in pairs on the dorsum ofthe
appendages may be lighter in color; propodeum typically low head,distinctfromthemuchshorter,appressedpubescence;legs
and broad, such that PDI 0.91-1.09 (Figs 38, 40); in one rare generally shorter (LHT/HL 0.58-0.86, rarely >0.80);
aberrantmorphwithHW >0.79 the propodeum is inflated and mesopropodealimpressionpartlyorentirelyflankedlaterallyby
prominentlyraised(Figs37,43);pronotalmarginvaryingfrom raisedridgesthatencloseapit-likedepression 18
sharp- to soft-edged, and maximum width of the pronomm 15. Head elongate (CI 0.70-0.77) (Figs 61-62) and petiole very
generallyoccurringbelowthemargin(Fig. 19)(widespreadand slender(PLI0.34-0.43)(Figs65-66) 16
highly variable species, distributed from India to southern - Headbroader(CI0.76-0.94,usually>0.80)(Figs63-64);petiole
China,southtoNewGuineaandnorthernAustralia) shapevariablebutifCI<0.80(afewindividualsofT.nigra)then
T.allaborans(Walker) petiolemorerobust(PLI>0.50) 17
Smallerspecies (HW 0.49-0.79), colorvariablebutoftenwith at 16. Smaller species (HW 0.96-0.97), with relatively large and
leastthepostpetiole—andsometimesmostofthebody—yellow conspicuous eyes (REL 0.40-0.42) (Fig. 62); profemur short
ororange-brown; ifHW>0.65 thenbodymostlydarkbrownto andbroad(FI0.45-0.47)(Borneo) T.buops,sp.nov
black but propodeum notably tall (lateral view) and slender Largerspecies(HW 1.06-1.27),withrelativelysmallereyes(REL
(posteriorview),suchthatPDI 1.12-1.24(Figs39,50);pronotal 0.25-0.30)(Fig.61);profemurslender(FI0.37-0.40)(Indiaand
marginusually relatively soft-edged andoccurring atthepoint Nepal,easttosouthernChina, southtoWestMalaysia)
ofmaximumwidthofthepronotum(Figs25-28) 11 T.binghami(Forel)
11. Small species (HW 0.64) with large eyes (REL 0.42) (Fig. 14); 17. Petiole long and slender, PLI 0.38-0.47, PL/HL 0.74-0.92
propodeum conical in profile, the prominent apex located far (Fig. 67); mesosoma, petiole and postpetiole, when viewed in
forward, sothatthe shortinclineddorsalfaceofthepropodeum profile, with scattered standing pilosity accompanied by, and
rounds into a much longer, sloping declivitous face (Fig. 52); oftengradinginto,adensematofshortersuberecthairs,present
petioleshortandbroad(PWI0.54,PL/LHT0.72),subtriangular onalldorsalsurfaces(Fig.67);(north-eastIndiatoChina,south
in profile and without a well differentiated anterior peduncle toSumatra,Java,BorneoandPalawan).. ..T.attenuataF.Smith
(Fig.52);castaneousbrown(Borneo) T.conica,sp.nov. - Petiole shorterandhigher,PLI0.52-0.64,PL/HL0.57-0.72 (Fig.
Eyes smaller(REL0.34-0.41); propodeumnotconical inprofile, 68);mesosoma,petioleandpostpetiole,whenviewedinprofile,
the dorsal face convex and rounding gradually into a steep with standing pilosity and underlying suberect pubescence
declivitousfaceofapproximatelythesamelength (Figs50, 53- variably developed (and variably distinguishable), but at least
54); petiole longer and narrower (PWI 0.38-0.54, PL/LHT thepromesonotumandthe anteriorpeduncle ofpetiole lacking
0.83-0.98),adifferentiatedanteriorpeduncleandposteriornode a dense mat of short suberect hairs (Fig. 68); (Pakistan to
evident in profile (Figs 50, 53-54); size and color variable Thailand,southtoBorneoandJava) T.nigra(Jerdon)
(modesta-comp\sx) 12 18. Petiole with a pair of acute, posteroventral teeth, formed from
12. Smallspecies(HW0.53-0.61),witharelativelyshort,highpetiole ventrolateral extensions ofthe petiolar sternite (Figs 79-84);
(PLI 0.60-0.68, PWI 0.46-0.54, PL/SL 1.09-1.19) (Fig. 53); pronotum with dense punctate sculpture on its anterior quarter
body and legs dark to medium brown (pronotum, petiole and which contrasts with the shiny (and less densely sculptured)
postpetiolemaybelighterincolor); standingpilositytendingto posterior half of head and with the more sparsely punctate
berathercommon,with 8-10 longsetaeoftenvisibleinprofile posteriorregionsofthepronotum;scapesshorterthaneyelength
onthepromesonotum(butsparseorabradedinsomespecimens) (SI30.83-0.98) 19
(Thailand,WestMalaysia,Borneo, Sumatra) Petiole lacking apairofposteroventral teeth(Figs 77-78, 91-99,
T.crassiuscula(Emery) 102-103); pronotal sculpture variable but punctures more
- Petiolemoreslender(PLI0.45-0.59,PWI0.38-0.48,PL/SL 1.20- evenly distribute4 not concentrated solely on the anterior
1.41)(Figs50, 54); standingpilosityrelativelysparse, 1-2pairs quarter(althoughtheymaybe sparsemedially) andusuallynot
oflong erect setae visible inprofile on the pronotum, none on occurring in a density that contrasts strongly with that ofthe
themesonotum;size(HW0.51-0.79)andcolorvariable 13 posterior halfofthe head; scapes longer than eye length (SI3
13. Head,mesosoma,andmostofgasterblackordarkbrownishblack, 1.02-1.55) 22
the other body parts variable in color, postpetiole and tibiae 19. Headelongate (CI0.73-0.77)(Fig. 76);petioleveryslender(PLI
often a contrasting lighter yellow or orange-brown; larger 0.43-0.49)(Fig. 84)(Thailand,WestMalaysia)
species, on average (HW 0.54-0.79, usually greaterthan 0.60) T.notabilis,sp.nov.
(Malay Peninsula south and east to Lombok, Sulawesi andthe - Headbroader (CI 0.78-0.90) (Figs 71-75); petiole much shorter
Philippines) T.extenuata,sp.nov. (PLI0.60-0.79)(Figs79-83) 20
- Body color predominantly yellow- or orange-brown, the gaster 20. Larger species (HW 0.83-0.95), with dense pubescence on
sometimes partially or wholly dark brown; problematic postpetiole andabdominaltergite IV, which obscuresthe sheen
specimenswithmore darkenedheadandmesosomaalsooccur. oftheintegument; frontalcarinaemorewidelyseparated(MFC
. . .
598 P.S.Ward
0.12-0.15,FCI0.15-0.16) (Fig. 75)(Borneo, Thailand) and broad (PLI 0.58-0.61, PWI 0.55-0.57) (Fig. 93) (New
T.nodosa,sp.nov. Guinea) T.mimula,sp.nov.
- Smallerspecies(HW0.63-0.83),pubescencegenerallysparseron - Frontal carinae less widely separated (FCI 0.12-0.16) and eyes
postpetiole(hairs separatedbyabouttheirlengths)andvarying larger (REL 0.40-0.45, REL2 0.47-0.53) (Fig. 86), such that
from sparse to moderately dense on abdominal tergite IV, not MFC/EL 0.25-0.30; pronotum slightly to sttongly expanded
obscuring the sheen of the integument; frontal carinae less laterally (PrWM/MTW 1.15-1.37); petiole shape variable but
widelyseparated(MFC0.07-0.10,FCI0.10-0.14)(Figs71-74) generally longer andmore slender (PLI 0.45-0.59, PWI 0.40-
21 0.49) (Fig. 92) (New Guinea and adjacent islands; northern
21 Shortstandingpilosity(0.03-0.05mminlength)commononmost Australia) T.laeviceps(F.Smith)
bodysurfaces,includingsidesofhead(Fig.74),dorsumofhead 27. Small species (HW 0.73-0.81); petiole with short anterior
(CSC 15-25), andmesosoma(MSC 30-56) (Fig. 81)(northern peduncle and large globose node, with steep anterior and
Australia) T.nixa,sp.nov. posteriorfaces(Fig.95)(FW/PH0.51-0.62);puncturesonhead
- Standingpilositymuch less common on head (CSC 0-6), absent and pronotum mostly very fine, about 0.005 mm in diameter
orsparse on the sides when head is observed in full-face view (NewGuinea,northernAustralia) T.rotula,sp.nov.
(Figs 71-73); standing hairs generally sparse on mesosoma - Larger species (HW 0.80-1.44), with less globosepetiolarnode,
(MSC usually <10), but occasionally quite common; ifMSC the anterior and posterior faces more gently sloping (Figs 94,
>20 then some hairs relatively long (0.10-0.20 mm in length) 97-99) (FW/PH 0.60-0.88); punctures onhead andmesosoma
(IndiatosouthernChina, southtonorthernAusfralia) mostlylarger,approximately0.010-0.015mmindiameter(New
T.nitida(FSmith) Guinea,Ausfralia) T.punctulataF.Smith
22. Mesopropodealimpressionflankedmoreorlessentirelybylateral 28. Standing pilosity common on head and mesosoma (CSC 18-28,
ridges,sothatthepit-shapeddepressionextendstotheposterior MSC 26-54), including mesonotum andpropodeum (Figs 77-
margin ofthe mesonotum; species foundeast ofWallace's line 78) 29
(Australia,NewGuinea,andadjacentislands) 23 - Standingpilositysparseonheadandmesosoma(CSC 2^, MSC
- Mesopropodeal impressionwithflankingridgesmuchreducedor 1-5),absentfrommesonotumandpropodeum(Figs 102-103)
lackinganteriorly,sothatthepit-shapeddepressionisseparated 30
from the posterior margin of the mesonotum by an open, 29. Eyes large (REL 0.41-0.44) (Fig. 69); lateral pronotal margin
transverse strip of integument, with longitudinally rugulate sharp-edged; appressed hairs present in moderate density on
sculpture; species found west of Wallace's line (India to the abdominal tergite IV, in addition to scattered standing hairs
Philippines,Borneo,SumatraandJava) 28 (India,WestMalaysia) T.aitkenii(Forel)
23. Petiole shortandverybroad(PL/HW0.57-0.59; PWI0.79-0.88) - Eyes smaller (REL 0.33-0.36) (Fig. 70); lateral pronotal margin
(Fig. 96); postpetiole about 1.4x broader than long; frontal notwell developed; abdominal tergite IV with abtindant, short
carinaewidelyseparated(FCI0.19-0.20)(Fig. 90)(Australia) standing pilosity but appressed hairs very sparse and
T.tucurua,sp.nov. inconspicuous(Borneo) T.polita,sp.nov.
- Petiolelongerandlessbroad(PL/HW0.62-0.96;PWI0.33-0.70) 30. Eyes larger (REL2 0.51-0.56) (Fig. 101); profemur shorter (FL/
(Figs 91-95); postpetiole approximately as long as, or longer HL 0.53-0.62, EL/FL 0.74-0.82); petiolarnodeusually with a
than, broad; frontal carinae less widely separated (FCI 0.12- short, steep anterior face and much longer, more shallowly
0.19)(Figs85-89) 24 inclinedposteriorface(Fig. 103)(Borneo,Palawan)
24. Posteriorhalfofpetiolar sternite flatorweaklyconvex inprofile T.inversinodis,sp.nov.
(Figs 91-93); petiole relatively slender (PLI 0.37-0.61, PWI - Eyes smaller(REL2 0.44-0.48) (Fig. 100);profemurlonger(PL/
0.33-0.57); eyes larger (REL 0.36-0.45); punctures on head HL0.60-0.67,EL/FL0.60-0.66);anteriorfaceofpetiolarnode
between compound eyes relatively coarse, mostly 0.010-0.020 usuallynotmuch steeperthanposteriorface (Fig. 102) (Malay
mm in diameter; propodeum somewhat elevated (PDI 1.10- Peninsula south and east to Sumatra, Java, Borneo and the
1.34), its dorsal face usually convex in profile, inclined Philippines) T.difficilis(Emery)
downward posteriorly, and grading insensibly into declivitous
face(Figs91-92) 25
- Posterior half of petiolar sternite with prominent ventral
protrusion(Figs94-95);petioleusuallymorerobust(PLI0.57- Tetraponera ofthe Oriental and Australian regions:
0.80,PWI0.50-0.70)andeyestendingtobesmaller(REL0.30- provisional keyto species, based on the queen caste
0.41);puncturesonheadbetweencompoundeyesfiner,mostly
0.005-0.015mmindiameter;propodeumlowerinprofile(PDI Thiskeyshouldbeusedwithcaution. Thequeencasteisnot
1.00-1.19), its dorsal usually flatter and more strongly known for all species, and the limits of variation remain
differentiatedfromthedeclivitousface(Figs94-95,97-99) . . . uncertain in species for which only a few specimens are
27 known. Whenarangeofmetricmeasurementsiscitedinthe
25. Head densely punctate, opaque; larger species (HW 0.94-1.04, HW
LSIH2T0.05.38-40-.15.50)0)(,NweiwthGluoinngeal)egsandscapes(LT.HaTt/raHLDo0n.i7s9t-h0o.r8p6;e kney=, t5h)i.s iTshfeolslaomwepdlebysitzhee sisamtphleessiazmee(ef.go.r all s1u.b8s8e-q2u.e29n;t
- Head less densely sculptured, the punctures separated by about measurements givenwithinthe same lugofacouplet. There
theirdiameters andthe interspaces shiny; smallerspecies (HW are four species inthe «/gra-group (out of20) forwhichthe
0.75-0.94;LHT0.63-0.78),withshorterlegsandscapes(LHT/ queencaste isunknown {T. buops, T. mimula, T. nodosa and
HL0.65-0.74;SI20.43-0.50) 26 T. polita). I have inserted them in the key where I would
26. Frontal carinae widely separated (FCI 0.17-0.19) and eyes
relatively small (REL 0.36-0.39, REL2 0.41-0.45) (Fig. 87), expectthequeenstocomeout,extrapolatingfromdistinctive
such that MFC/EL 0.38-0.45; pronotum slender, as seen in features ofworker morphology. Such inferences are always
dorsal view (PrWM/MTW 1.04-1.16); petiole relatively short accompaniedby anexplicit statementthatthe queencaste is