A new species of Litoria (Anura: Hylidae) from New Guinea and a redefinition of Litoria leucova (Tyler, 1968)

7 Pages·1994·2.4 MB·English

A NEW SPECIES OFUTORIA ANURA: HYLIDAE") FROM NEW GUINEA ANDA
(
REDEFINITIONOFLITORIA LEUCOVA (TYLER, 1968)
G.R.JOHNSTON ANDSJ.RICHARDS
Johnston,G.R.&Richards,S.J. 1994 J201;AnewspecksofLituria(Amita. Hylidae)ttoiii
NewGuineaandaredefinitionofLitoriaieiicova(Tyler, 1968) MemoirsoftheQueensland
Museum37(1): 273-279. Brisbane ISSN 0079-8835.
AtpresentLitoriateucova is known from theholotypecollectedat Busilminnearthenorth
flowing May River and seven specimens from the south flowing Ok Tcdi in the Stai
MountainsofNew Guinea. We show that the specimens from the north andsouthfacesof
thecentralmountainsrepresenttwodistinctspecies.Thispaperdescribesthe southerntaxon
as Litoriamajikthisesp. nov, andredefinesL teucovaconfiningittothe northerntaxon.
Hylidae, Litoriateucova, Litoriamajikthise, newspecies. NewGuinea.
GregJohnston, SchoolofBiologicalSciences, Flinders UniversityofSouthAustralia, GPO
Hox 2/00, Adelaide, South Australia 5(MX), Australia; Stephen Richards, Department of
Zoology,James Cook University, Townsville, Queensland48]/, Australia; 16June 1994.
Litoria teucova (Tyler, 1968) was described mencetal., 1984)wasdonetotest fot-significant
froma single adult female collected from Busil- multivariatedifferencesinrawmeasurementsbe-
minneartheMay Riveronthenorthern slopesof tween the three taxa.
the Star Mountains ofPapua New Guinea. Men- Calls were recorded in the field with a SONY
kes (1993) redescribed the species based on Professional Walkman easette recorder and an
sevenspecimensfromthesouthflowingOkTcdi, Electret Condenser microphone ECM Z200.
without reference to the holotype, and placed it Calls were analysed using ULTRASOUND
in theLitorianigropunctata species group. vh10(Jordan, 1988).
During field workin the Westernand Sandaun
Provinces of Papua New Guinea we collected
Comparisons
specimens which, after comparison with type
mbayteMreinalz,iemsad(e19i9t3c)leiasrptohlayttLy.pitce.ucTohveasapsedceifmiennesd basLiistoorfiaftlseiuncgolveafweamasloeriwgiitnha:llIy,dSiVaLgn=os3e0d,o4nmtmh;e
Menzies(1993)referredtoL teucovaare notthis 2, an immaculate blue dorsum and stippled blue
species but represent an undescribed species. In on thethighs, 3,an eyetonarisdistance less than
this paper we report further specimens of L intcrnarial span; 4, no vomerine teeth; 5, an im-
teucova and describe and illustrate the new maculateunpigrncnledventralsurface;6,noflash
markings in the groin oron posteriorsurfaces of
species.
thighs; and 7, unpigmented ova (Tyler, 1
M \TERJALS AND METHODS This specimen was collected on the northern
slopes ofthe Star Mountains.
ThespecimensMenzies (1993 referredtojs t
)
Eighi morphometric measurement (in mm) leucovowereall maleandquitedifferentfromthe
were made of specimens of Litoria iris, L holotype: they were larger (SVL = 31.0-
leucovu and the new species held in the South 35.4mm). mottled on the dorsum, had brigir
AustralianMuseum(SAM)and the University of flash markings in the groin and on the hidden
Papua New Guinea (UPNG). These measure- surfacesofthethighs, largepurplepatchesonthe
mentswere: snout-ventlength(SVL),headwidth ventrolateral surfaces, and possessed vomerine
(HW). head length (HL), eye diameter(ED),eye teeth. All of these specimens were from the
to narisinterval (EN), inlernarial span (IN), tibia southern slopesofthe Star Mountains
length (TL)and footlength(FL) followingTyler Since the holotype was female and all of
11968). Menzies' (1993)specimensweremale,thesedif-
Descriptive statistics and a number of mor- ferences may have reflected sexual dimorphism
phometric ratios were calculated from these in one species. However, this is unlikely foi a
measurements. All measurements and propor- numberofreasons.Firstly,maleAnuraareusual-
tions given are for adult males unless otherwise ly smaller than females. The existence of a
staled. A discriminant function analysis (Rey- specie* wittl ;i leveisa] ofthis pattern would he
274 MEMOIRSOFTHEQUEENSLANDMUSEUM
L.'leucova' were correctly identified. All
misidentifiedsouthernfrogswereassignedtothe
group representing L iris by the analysis. A
L
posteriori placement of the type specimen of
leucova onto the discriminant functions
demonstratedclearlythatitfallswithinthenorth-
erncluster, butwelloutside oftheclusters repre-
seniing L iris and the southern taxon. On the
basisofthismorphologicalanalysisanddifferen-
ces in mating call we describe the southern L
'lei4cova' as a new species, and redefine L.
leucova (sensu stricto) to include only northern
specimens.
Litoria majikthise sp. nov.
(Figs 1-4)
-3-1
-y • a MaterialExamined
Holotype: SAMR44093, 6kra SSW Tabubil.
rtMANT FUNCTiON I) Western Province. Papua New Guinea, 5'18'S,
141a12*E. G.R. Johnston, S.J. Richards, 10,viU993,
FIG. 1. Plot ofindividual Litoria iris(circles), Liiaria male.
leucova(triangles)andLitoriamajikthise(squares)on Paratypes: UPNG6734, Megalsimbip, Ok Menga,
thefirsttwodiscriminantfunctionaxesbasedonseven Western Province, Papua New Guinea, 5J18*S,
measurements. Hollow pentagons=group eentroids. 141°20'E, J.C. Pemetta, viU981; UPNG7305-7309,
Hollow star=holotypeofLitorialeucova, Lower Kam valley, Western Province, Papua New
Guinea, 5'12'S. 14ri4'E, J. Menzies, i.1985;
very unusual, especially in the absence ofmor- UPNG8501-S508, 6km SSW Tabubil, Western
phological or behavioural indications chat male Province, Papua New Guinea, 5°18'S, 14l°12*E, S.J.
combatisimportantinthisspecies(Shine, 1979). Richards, G.R. Johnston, 28.xi,199l; SAMR44094,
Secondly, we have collected a female of the 1km SSW Tabubil. Western Province, Papua New
1
"southern' taxon and found it to be larger than Guinea. 562TS 141'12'E, G.R. Johnston, S.J.
males (SVL=39 2mm) but entirely consistent Richards,22.vii 1993;SAMR44095-44101,6kmSSW
with them in all other respects. Thirdly, several Tabubil. Western Province, Papua New Guinea,
males collected near the type locality of L 5°18'S 141'12'E, G.R. Johnston, S.J. Richards, 9-
leucovacorrespondinallrespectstotheholotype PIrO.ovviiin.c1e9,93P;aUpuPaNGN8e6w02G-u0i3ne.a,125k°m22S'STa1bu4briil7.'WEe,stGe.Rr.n
of that species, but differ from males of the
Johnston, S.J. Richards, l-20.vu.1993.
southern taxon in morphology and structure of
the mating call. Diagnosis
Thus, two separate taxa, from the southern and AmemberoftheLitorianigropuncnituspc
northern slopes ofthe Star Mountains, appearto group (Menzies 1972, 1993). Distinguishable
be included in Menzies* (1993) concept of L from all other members of this group by the
leucova. This hypothesis was tested using a dis- following combination of characters: 1, green
criminant function analysis of seven measure- dorsallyinlife:2.srnHit-ventlength30.5-3.S4niiii
ments using L iris, the southern and northern L. inadultmales;3, thighsandgroincrimsoninlife;
'leucova' asapriorigroupings(Fig. 1).L Mswas 4, purple speckling on lateral surfaces of belly
included in this analysis because phenetically it tending to form patches which do not separate
is the species of LUoria most similar to the distinct large, white lateral patches from venter:
southern taxon. 5, distinct white postocular spot; 6, call with
There were two significant discriminant func- dominantfrequencyof2.5kHz,apulserepetition
tions, the first of which accounted for 81.4% of rateof0. 16-0.24pulscs/ms.
the variance. All measurements contributed sig-
nificantly to decrease Wilk's Lambda. The Description
analysis resulted in correct identification of Bodysizesmall(SVL=30.5-35.4mminmales,
93.8%ofspecimensoverall. AllZ,. insandnorth- 39.2mm in a female). Head slightly broader than
ern L, 'leucova', and 85.7% of southern long oras broad as long (HL/HW = 0.81-1.03).
NEW HYLIDFROG 275
TABLE I. MeasurementsofadultmaleUtoriairis> L. faces of hindlimbs, concealed parts of thighs,
itucovaandL.majikthise. Valuesarcmeans(standard groin and in some specimens much of the belly
deviations).Themeasurementsaredefinedinthetext. pale pink Lateral surfaces of throat and body
P=probabilities associated with oneway ANOVAs with blue-grey stippling, sometimes developing
comparing thetaxa. intolargedarkpatchesinthe groinandonthroat.
' Some stippling may also occur on the posterior
Character mt.ajikthise L iris L teucava P surface of thighs. A distinct, broad white pos-
(N=I4) (N=I3) (N^5) tocular bar between eye and foielimb, rum
SVL 32.7(1.659) 31.4(3.111) 26.1(0.391) 0.000 belowthetympanum.
HW 9.8(0.395) 9.5(0.647) 8.8(0.363) 0.006 Inlifedorsalsurfacesimmaculatedarkgreenor
HL 8.9(0.622) 8.1 (0.806) 7.6(0.466) 0.003 pale emerald green with darker green mottling.
ED 4.0(0.412) 3.6(0.456) 3.0(0.164) 0.000 Ventral surfacesgenerally white;ventralsurfaces
of hindlimbs and belly red. Lateral surfaces of
EN 3.0(0.283) 2.6(0.166) 2.3(0.123) 0.000 throatandbody speckledwithpurple,sometimes
tN 3.frilJ 1K1> 3.5(0.420) 2.7(0.205) 0.000 developing into large dark purple patches in the
Tl. 18.4(0.547) 18.2(1.719) 14.2(0.383) 0.000 groin and on throat. Stippling also present on
posteriorsurface ofthighs in some specimens A
Head length about one quarter of body length distinct broad, pearl-white postocular bar. be-
(HL/SVL = 0.24-0.29). Snout prominent, round tweeneyeandforearm,belowtympana. Irisgold
when viewed from above and slightly round to en yellow.
truncate in profile. Nostrils lateral. Eye-to-naris Measuretnents of holotype (in mm). SVL =
distance less ih;m or equal to inlernarial span 34.0;HL =9.0;HW= 10.8;ED= 43:EN =3.I:
iEN/IN =0.72-1.03). Nostrilsmuchclosertoend IN =3,9;TL= 18.7; FL= 231,
ofsnoutthaneye Canthusrostralisrounded,con-
caveand slightlyto well defined. Eyes large and MatingCall
protuberant, eye diameter generally greater than Two types ofCall are uttered (Fig. 4B.D). One
0d.i9s7t-a1n.c5e5)f.roTmymepyeantuomnosctorvielrenodstwriilth(sEkDi/n,ENon=e hia$russhho'rchii{p2'6-co8n0siims,tminegano=f46(-3,1S6Dpu=lsIe8s.0(6m\eNa=n7)=.
thirdtoonehalfofeyediameter. Vomerineteeth 8.9, SD = 3.29) It a pulse rule of0.20-0.24 pul-
on two slight elevations between the choanae. ses/ms(mean=0.22,SD=0.013).Thisshortcall
Tongue broad andoval in shape. beginsat 2.2-3.2kHzand finishesat 1.4-4.3kH7„
hungers short with moderate lateral fringes; bui has a dominant frequency around 2.5kH7
finger lengths 3>4>2>l. Terminal discs large. Thesecond longer(29Q-360m.s, mean =330, SD
Webbingtosubarticulartubercleatbaseofpenul- =36.1, N = 3)call is a harsh *raagh\ incre*iMng
timate phalanx of fourth finger. Hindlimbs in volume, consisting of 49-63 pulses (mean =
moderately long (TL/SVL = 0.53-0-60); toe 57.3, SD= 737)at a rate of0.16-0.19 pulses/ms
lengths 4>3>5>2>1. Webbing between toes to mean=0.174,SD=0.010).Thislongcallbegins
I
base ofdisc on all toes, except fourth and fifth at 1.8-2.2krteand ends at l.8-5.0kHz, but has a
Webbing on fourth toe to base of penultimate dominantfrequencyaround2.5kHz.Anumberof
phalanx. Webbing on fifth loe reaches midway individualfrogswerebeardtoutterbothshortand
alongpenultimatephalanx,continuingtodiscvia longcalls(including theholotype). Twentycalls
a narrow fringe. A small, oval inner metatarsal uttered by fourindividualsgave a meancallrate
tuberclepresent; noouter metatarsal tubercle. of one per 26s. All calls were recorded at
Skin of dorsal surfaces minutely tubercular temperaturesof21.0to22.2C.
Throatandchestslightly granular. Fewtubercles
onposteriorsurfaceofforearmandouteredgeof Etymology
each foot. Few conspicuous tubercles on From Majikthise a farcical comedycharacter
'
ventromedial surfaces of thighs, below anus. (Adams. 1979). referring to the vividly coloured
Posteriorsurfaceofthighsgenerally smooth, but thighs and groin.
interrupted by few large tubercles. Supratym-
panic fold inconspicuous. DiSTRJBLTIONAND HABITAT
In preservative dorsal surfaces immaculate Known only from altitudes between 550 and
blue (one specimen) or densely mottled with 650m in theheadwatersofthe soMtl draining Ok
darker blue (all other specimens). Ventral sur- Tedi in the StarMountains. Specimens werecol-
faces generally cream to white, but ventral sur- lectedatnightonfoliageattheedgeofcultivated
;
276 MEMOIRSOFTHEQUEENSLANDMUSEUM
MaterialExamined
Holotype: SAMR6461, Busilmin, Sandaun
Province, Papua New Guinea, 4*55'S 141'00'E, B.
Craig, 1.v.1965,female.
OtherMaterial:SAMR44091-44092,UPNG8604-
06, Stolle Mountain, Sandaun Province, Papua New
Guinea, 4#48*S 141'39'E, G.R. Johnston, S.J.
Richards,4.vii.1993.
Diagnosis
Distinguishable from all other Papua New
GuineanLitoriabythefollowingcombinationof
characters: 1,Greendorsallyinlife;2,Snout-vent
length30.5-35.4mminadultmales;3,thighsand
groin translucent pink with yellow spots in life;
4,nopurplespecklingonlateralsurfacesofbelly
5, no white postocular spot; 6, calls with
dominant frequency of 4.0-5.0kHz, a pulse
repetitionrateof0.11 pulses/ms.
Description
Bodysizesmall (SVL=30.5-35.4mminmales,
30.4mm in a female). Head slightly longer than
broad (HL/HW=0.81-0.91). Head length about
one third of body length (HL/SVL=0.26-0.31).
Snout not prominent, round when viewed from
above and truncate in profile. Nostrils lateral.
Eye-to-naris distance less than internarial span
(EN/IN=0.78-0.96). Nostrils closer to end of
snout than to eye, Canthus rostralis straight and
slightly defined. Eyes moderate in size, not
prominent.Eyediameterfractionallygreaterthan
distancefromeyeto nostril(ED/EN=1.25-1.43).
Tympanum visible, one half of eye diameter.
Vomerine teeth absent. Tongue oval in shape
withstrongly indentedposteriorborder.
Fingers short withbroad lateral fringes; finger
lengths3>4>2>1.Terminaldiscslarge.Webbing
to subarticular tubercle at base of penultimate
phalanx of fourth finger. Hindlimbs long and
slender (TL/SVL=0.53-0.55); toe lengths
4>5>3>2>1. Webbing between toes to base of
disconalltoes,exceptfourth.Webbingonfourth
reacheshalfwayuppenultimatephalanxandcon-
FIG.2.A,B,Litoriamajikthise;C,Litorialeucova.A,C, nectedtodiscbybroadlateralfringe. Small,oval
lateralviews;B,ventralview. inner metatarsal tubercle present; no outer
sago swamps, among grasses in uncultivated metatarsaltubercle.
swampsandonvegetationalongaroadsideditch, Skin of dorsal surfaces smooth, except for
allwithin 15kmofthetownshipofTabubil.Perch weakly tubercularuppereyelids.Throatsmooth.
heights varied between 1.0m and2.5m. Chest and anterior portion of abdomen, and
ventral surfaces of thighs granular. Large
Litoria leucova(Tyler, 1968) prominenttubercleson ventromedial surfacesof
(Figs 1-4) thighs, below anus. Supratympanic fold
prominent, extending from posterior corner of
HylaleucovaTyler, 1968: 119. eye, covering upperrimoftympanic annulus.
1
NEW HYLID FROG 277
cleared swampandfromvegetationoverhanging
a swift-flowing stream at 1600m altitude.
COMPARISON OFUTORIA LEVCOVA
mTWLMAJIKTHfSE
Uloriuleucova issmallerand therefore differs
in most absolute body measurements from L
majikihise (Table 1). The correlations between
the measurementsandthediscriminantfunctions
indicate that, apart from its smaller size, L
leucovahasalongerhead, moreprominentsnout
and smaller eyes than L majikthtse. Vomerine
teeth ai>; absent in L leucova but present in L
majikihise.
FIG. 3. A, Ltioria majikihise; B, /* iris. Ventral view The two species also differ in colour (Fig. 2).
showing thatthe purple (black)ofL majikthtsedoes In life the presence ofcrimson red in the groin
not separate distinct white lateral patches from the andonthethighsinLmajikthtseclearlydifferen-
remainderofthe venlerasit docs in Z- iris. tiates it fromL leucova, in which the thighsarc
In preservative dorsal surfaces immaculate translucentpinkwithyellow spots.However, this
dark blue. Ventral surfaces white. Hidden sur- difference is lessclearinpreservedspecimensas
faces of the thighs stippled with dark blue; this the crimson colour of L majikihise fades. The
stippling isabsentfrom numeroussmall circular presence ofpurple stippling on the ventrolateral
areassothatgroundcolourappears ntheformof surfacesofthebodyclearlydistinguishL majik
i
circularspots. Subanal tuberclescreamishwhite. thisefromL leucova,inwhichtheventralsurface
In lifedorsal surfacespaleemeraldgreen with is white. All but one specimen of L majikihise
occasional paler green spots on some in- was strongly mottled dorsally, whereas L
dividuals.. Ventral surfaces generally white leucova is generally immaculate pale greendor-
Thighs and anterior lateral surfaces translucent sally with the occasional paler spot on some
pink with yellow spots. Insgolden yellow individuals. TheonlyL majikihise tolackdorsal
mottling wasdark greenon rhedorsum A pearly
Mating Call white postocularspotis present in L. majikthtse.
Two types ofcall were uttered (Fig.4A,C). One but absent in L Jeuroxa
isashort(15-20ms,N=23),harsh 'chip' consisting ThecallsofL leucovahave ahigherdominant
of a single pulse. This short call begins at 3.5- frequency (4-5kHz) than those of L majikihise
60kHz for 10-15ms, then broadens to 2.0-6.0kHz. (2-3kHz). Both species utter distinct short and
for the last 5ms. The dominant frequency was 4- longcalls. TheshortcallsofL leucovaconsistof
5kHz.TTiesecondlonger(90ms,N=lKailisaharsh asingle pulseand are 15-20mslong, whereas the
raagh\ consisting of 10 pulses at a rate of 0.1 shortcallsofLmajikihiseconsistsof6-16pulses
pulses/ms. This call began with three pulses be- and are 26-80rrts long. A single long call of L
tween 4.0 and 5.5kHz, followed by nine pulses leucova consisted of 10 pulses and was 90ms
between 2.0 and 5.5kHz.The dominant frequency long, whereas the long call ofL majikihise w<js
was 4.0-5.0kHz for the first nine pulses and then longer (290-360ms) and contained more pulses
shifted to 2.0-3.0kHz for the last pulse. Two in- (49-63). L. leucova calls at a much faster rate (1
dividuals were heard to utter both short and long call/2.6s) than does L. nuijikthise(1 call/26s), so
calls in long, series. Twenty four calls by a single that it is difficult to know whether the series of
frog gaveamean call rateofoneper2.6s forshort short and long calls uttered by L leucova is a
Callsandonecallper60sforlongcalls.Callswere series of separate mating calls or a single call
recordedata temperatureof 17.6C. consistingofthese subunits.
Utorialeucovaoccursatmuchhigheraltitudes
Distributionand Habitat (1600m) than docs L. majikihise (550-650m).
Knownonlyfromtwolocalitiesonthenorthern They may also differ in habitat preferences; L
slopesofthecentral mountains. Specimens were magikthisewasfoundinswampsoralongasmall,
collected while they called from piles of socks slow-flowing creek, whereas L. lateova was
overhanging streams running into a recently found calling from streams flowing through ••>
1 ,
27H MEMOIRSOFTHE QUEENSLAND MUSEUM
ia]A B clearly distinguishLable from theharsh 'chip' ora
harsh 'raagh* of majikthise (Menzies, 1993;
pers. obs.).
* COMPARISONOFLITORIA LEUCOVA
WITHOTHERSPECIES OFLITORIA
Tyler & Davies (1978) placed L leucova in a
0-1 B monotypicspeciesgroup,definedbybeingsmall
o
uniformlygreen, montane frogs, withshort, half-
wehbed fingers, fully webbed toes and unpig-
mentcdcggs.Tyler(1968)notedthatthegranules
on the ventral surface of the holotype of L
leucovatend toformtransverseridges.Examina-
tion of the type and live specimens suggest that
this characteristic is an artefact of preservation
u H and not a structural feature ofthe frogs themsel-
0.1 0.5 ves.
TIME(seconds! The only other species of similar size to L
leucova which lay unpigmented eggs include
FILG.ma4j.iSktohniasge,rahmolsootyfpcea.llUsp.pAe,rC:,sLhiotrotrciaalll.eLucoowvea;rBl.oDn,g aslolmmeemmbeemrbserosfothfetLhebL.ecnkiigrgorpouunpct(asetnasgurToyulpearn&d
call. Note thai the scales on the abscissae differ be- Davies, 1978).SpeciesintheL.beckigroupdiffer
tween panel Dandallotherpanels. ftotn L leucova in having long unwebbed (vs
short 1/3 webbed) fingers and a ventral surface
clearedareathathaduntilrecently beenaswamp variegated with dark pigments (vs white). L
and along a well vegetated, swiftly flowing leucova is similar to members of the L becki
stream. group in that it has been found along a swiftly
flowing mountain stream, but some L. leucova
COMPARISON OFUTORIA MAJ1KTHISE werecollected from small streamswhich flowed
WITH OTHER SPECIESOFLITORIA through what had recently been a swamp. Thus
the true habitat preferences of L leucova are
Litoria majikthise has purple ventolateral difficult toassess.
patches in common with L iris and L ollauru Of the species belonging to the L.
(Menzies. 1993), and which distinguishes these nigropunctata group, L leucova differs from L.
three species from all other species of Litoria. havma Menzies, 1993, L macro Menzies, 1993
The purple patches ofL majikthise are smaller andLpronimia Menzies, 1993 inthatmaleslack
than those in L. tris, and do not separate distinct a rostral spike (Menzies, 1993). L. vocivincens
large, white lateral patches from the venter as in Menzies, 1972 occurs only on the southern
L iris (Fig. 3). L. majikthise has crimson in the lowlandsofNewGuinea andthehiddensurfaces
groinandonthethighs,whereasLirishasyellow ofthethighsareblack(vspink)(Menzies, 1972).
toorangeandL oliauro hasorange(Tyler, 1962, L oliauro and L. iris have large purple ventral
Menzies, 1993). patches, which L. leucova lacks (Tyler, 1962;
In the Ok Tedi area L iris occur within 20 Menzies, 1993). Dorsally L nigropunctata
kilometres of L majikthise (Hyndman & Men- (Meyer. 1874) is usually brown orgreen-brown
zies, 1990), but L iris is a montane species and (vs pale green) and the hidden surfaces of the
occurs at much higher altitudes (1300-2300m; thighs of arc black (vs pink) (Menzies, 1972).
Tyler,1968) than does L majikthise, Litoria chloronata (Boulenger, 1911) is the
Comparison ofthe call of L. majikthise with species which mostcloselyresemblesL. leucova
thm of /. iris (based on Menzies, 1972, 1993) in the L. nigropunctatagroups, but it's call has a
show nodifferences in the fundamental frequen- lower dominant frequency (2.6kHz) than L.
cy, but thepulserateis lowerinL. iris(0.09-0.1 leucova and the concealed surfaces ofthe thighs
pulses/ms) than in L. majikthise (0,16-0.24 pul- andgroinarebrightorange(Menzies, 1993). The
ses/ms). Menzies (1975) described the call ofL only species in the L. nigropunctata group that
iris as a slow buzzing or spluttering which is occurat altitudes higher than 1000m is L.iris,
NHW HYLIDFROG 279
Litorialeucovadiffers frommostcongeners in lyse calls Our work in New Guinea could not
lacking vomerine teeth. Litoria chloronata and have been done without the help of Roselyn
someL irissharethelackofvomerine teeth with Busasa (Institute ofPapua NewGuineaStudies)
L leLucova (Tyler, 1962, 1968)as dom&embersof DrNavu Kwapena (Department ofEnvironment
the bicolor species group (Tyler Davies, andConservation,PNG),andDrsJamesMenzies
1978). Litoria leucova differs from all members and Ian Burrows (University of Papua New
ofthe L bicolorgroup in being of more robust Guinea). Dr Margaret Davies and Mike Tyler
habitus, and having unpigmented eggs. commentedon the manuscript.
Litorialeucovasharesfeaturesincommonwith
members of both the L becki and the L LITERATURECITED
&
nigropunctataspecies groups ofTyler Davies
(1978). Unfortunately, neither of these groups
canbedefinedbyasinglesynapomorphy andare ADAMS, A. 1979 "The hitch-hiker's guide to the
, galaxy' (Pan Ltd: London.)
almostcertainly not monophyletic. Indeed, there HUTCHINSO• N. M.N & MAXSON, L.R. 1987
is little correspondence between hypothesised Phylogcnetie relationshipsamong Au^ualian wee
rmeolratpihoonslhoigpsicaamlon(gTyspleecries&ofDLaivtioerisa,ba1s9e7d8o)n. mfruongoslo(gAincuarla:aHpyplriodaaceh:. PAeulsotdrraylaidainnacJ)o:urannalimof
k(aHruytocthyipniscon(K&ingM,ax19s8on1),an19d8i7m).muWnoelohgaivcealtednattaa- HYNDZMooAlNog.yD3.5:C6&1-M74E.NZIES,J.I. 1990.Rainforests
tively referred L majikthise to the L of the Ok Tcdi headwaters. New Guinea: an
nigropunctata group because of its obvious ecological analysis. Journal ofBiogeography 17;
similaritytoL. iris,aspecieswhichhaslongbeen 241-273.
included in this group (Menzies, 1972, 1993). JORDAN, B. 1988. 'Ultrasound vl.10'. (Queensland
TheaffinitiesofL. leucovaare lessclearandwill University: St Lucia).
only be resolved by a thorough pbylogfinetic KING, M. 1981. A cytotaxonomic analysis of
analysis of this speciose genus based upon all A1u6s9t-r1a7l5i.anIhnylBiadnkfsr,ogsC.of&theMagretinnu,s ALi.tAoria(.edPsp).,
available data.
'Proceedings of the Melbourne HerpetolopL-a.
Symposium May 19-21, 1980'. (Zoological
Comparative Material Examined
BoardofVictoriaand Royal MelbourneZoologi-
litoria ins: UPNG4434, 4478-79. Moiyakabip, Hin- calGardens Parkville. Melbourne).
dBeankobnearbgipR;anUgPeN;G2UIP0N1G,23M1ar8g,ariUu.PrN.GU7P3N1G32-17I37I-K MENZIES, J. 197:2. Papuan tree frogs of the Litoria
2158, 2241-2281, Nipa; UPNG3115-3135, Tan; nigropunctatagroup. Herpetologica28: 291-300.
UUPPNNGG33814540-,31T7a8,mbKuolm;o;UPUNPGN8G2386974-,905,52HU,baMiegnudbki 19a9n3.dSityssatlelmieastiincsNoefwLiGtuoirnieaairainsd(Aannuortae:oHnylsiedxauea:l)
Lodge; UPNG6992-3, 7168, 7655, Ml Gilewc; dimorphism in the group. Australian Journai oi
Zoology 14:225-255.
UUPPNNGG75082132--7I032.4,5A8n9g8a-5l9i0m3p,caHmapl;alUiPnNjGa7v1i4a8-N7i1p6a7;, REYMENT.R.A., BLACKJTH, RE &CAMPBELL,
Porgcra; SAM524I, Okapa; SAM5423, 5874. N.A. 1984. 'Multivariate morphometries*. 2nd
Telefomin; SAM5726, Dumun. Edn, (Academic Press * London),
SHLNE, R. 1979. Sexual selection and sexual dimor-
ACKNOWLEDGEMENTS phism in the Amphibia. Copeia 1979: 297-306.
TYLER. M.J. 1962. A new hylid frogfrom theCc
Highlands ofNew Guinea. Recordsofthe South
Thiswork wassupportedinpartbygrantsfrom Australian Museum 14: 253-257.
the PeterRankinTrustFundforHerpetology and
1968. Papuan hylid frogs of the genus Hyla,
James Cook University. Ok Tedi Mining Ltd Zoologixche Verhandelingen96:1-203.
provided accomodation, transport and logistic TYLER, M.J. & DAVTES, M. 1978. Species-groups
support in the field. Phil and Sue Gregory, and within the Australopapuan hylid frog genus
KcytFischerprovidedadditional assistance. Ken Litoria Tschudi. Australian Journal of Zoology
Sandersonallowed us to use his software to ana- Supplementary Scries63:1-47.