Table Of Contentth<n\n<tnm\ oftin- Hajwi Susieh"{5 rtWJ I WUbbh 120(T. K3-w
THE BREEDING BIOLOGY OF UTORIA SVBGLAISDVLOSA AND L CITROl'A
(ANURA: HYLJDAE), AND A RE-EVALUATION OF THEIR GEOGRAPHIC
DISTRIBUTION
by Makion Anstis* & Murray LrrrudoiiN'
.1.
Summary
Awns, M $ | iTTUyOH!^ M .1, ||o%) The breeding bwtogj v\'lJt»riu .stth,t>lwuli<h>.\u and /, itroffi) tAnur;i;
liyltdac). and u re-BYMluitfinn ol [heir geographic distribution. 7Whm. A' Sin. S
November, 14%.
'Mil- known mnge "| UWhti whxluitduiost; is cvlcnJu'J and lluu 111 /.. rmvyvj is ivvised. Population u nil
observed at the type localiiy during ihc 1960s 70s and iy$ta flit!compared.
Ihe advertisement ca;ldll. adult colourafii'ii in lilR. behaviour and enihryological development ol /
SUbxfanMo.Ht are devcriibLe.d and compared with ihoseol L. iitm/tu. The single e^'g mass ol /. \ithuhnulitii>s<
showsadaptationhidie loiicemironment. btfirujcompact and strongly adherent Flicembryos .ind larv.d slaves
o! the two species are very similar in shape and colour in life from KtUge 17 onwards, hW $t£ rradilv
distinguishable- by moulhparts. CompataMvc lotcs do lOj'Vat hctwvuuir are given.
Kiv Wwkus: Iif/nut stihtihaulttfosa, Utwitt cfflvpjtl, disiribution, population ireiidv. advcniscnieiii calls,
oviposiiion. entbryohH/y, larval behaviour.
Introduction R^6l)75 - (my f-awkes Rivei, Cbor: R7I 109-71 I 14
Back Creek (Rarwick Riven ncai Point Look<nu
xluLnt(tiourti<oKs\aNhiT>lyilte'ritl&ulo\Aunswlaiss. de1s*9c?r?ibebdutasreLnltavm)'ehdl CRd3e7ii0I7Inn-es5kn(iGiSbraolltaWralRcahnuue:);R3OR0552C9)3I50kSmaFmlowf
because of primary homnnyrny (Tyler & Ansiis. Cieek, Doiriijo, R5II7X-XO Sts \ River. Point
1983% A member of cite L it'tropn species group Lookout; R76S|g - GlouccsJer Tops: RM6K3
(Tyler & Duvics [078), L suhi>latuitihi\d was LIpper Allyn River. Fiarnivjion lops; fcUMSOS
previously Known only loan the Quecnsraiiti/NSVV Hllenborou^li River. Bul^a State Foresi, NSW.
border south 10 the New raigland ranges ol" northern
NSW (Tyler & Ansiis 1975) The type description Lttnria enount
included a description ol the larvae, but nodata were Adult v/vr/wevM csaminctl: Atislralian Museum
available on ovipositiomcrnbryoloijicnl development, R75bO_ Orbi3st:75^ AbertekkVic: 19237. 18234.
larval hehavioni or the* advertisement cull. Uttfe. IK23X Slanwcll Tops: 794^0 Sianwell Park:
The species was found 160 kin soulh ol its 245t»0-24505 2759t», l-mlc.mhridpe: 45X5X.
previous known distribution in die mid-norih coastal rhillmere LakTes; 3I6S5. 71 12. 7S^>27 1leiensburch;
ijm^cs and Burriivjton Tops region by one of US 45424. Tianjara l-'alls; 5INS. Megaloue Valley: 71 10,
iM.A.i. in 1977 lis presence there din\ ihe absence lla/elbrook: SHHL Blaekheallv. 69034. Bell.
ot / . tny>i>u, prompted a re-examination o\ the Kurfifiuns Rd.: 76625. IS km N iit l.ilhgow; S459,
distnbubon ofboth species. In addition, observations Pennant Hilts; 14495, Colo Vale: 79100. 76623,
on ovipositioiu the morphology <T embryos. )ar\ae Clljoul Range N of Colo flt>.: 4261. Bundainjon;
and adults and a comparison of the advertisemcni 7IK9S. 24 km N of Moss Vale: 15462. Goslnrd;
calls i.if L. ritn'pa and /.. sidiyjttndtdasn were ma<le /S264-26. 7S698. Kurmgai^Chase: 6<»425. Nad^ree
and are repotted here. Reserve; 794*9, (Jalslon doi-ge; 7563, Manly, NSW.
Three adults iitcd as /. vitrofiu b> lyler & Ansiis
Materials and Method* (1975) irom Barrinylon Tops localities- l)epl/ool_,
UtliV. Melbourne (MC/I)| 1792/64 - t'].»pei Allyu
Liuftfa \ifh's>i(tti(lnl('\<t and MV/O 1690^1/63 - Wombat Creek, were re-
\didt specimens examined: Australian Museum examined because of appareni overlap in ranjjc with
(AM) RI7577. 35525, 4293i-33. 50I6.V 510&&-7, Ihc Barrirtrton Tops localities lor A_ vuhithmdidtna.
51104, 517354V. Point Lookout; R3445N - 14km These specimens have since been registered by the
f-.asi ol [dim: K36724 - Oafesv Creek near Poor, National Museum ttf Victoria (NMVt as D32666
(tipper Allyn Riyei) and 03260T65 (Wombat
KWhtevrm Ril BetcrWRi Hv.yhls NSW 2oH2. CreekI- Similarly, two specimens <NMV D670'J-10).
l)r|iiiMiiuiii ul /...iL^y.t'hH'i-,Hv ,.t WUiMi.iH. I'iied by Copland (1957) i\$ I. tftfQpn from nr.n
P.HkvillrVK- KJSJ ( irafton. iMMtheaslern NSW. wore examined,
'
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BREEDING BIOLOGY OF L SUBGLANDULOSA & L CITROPA 87
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Observations on three captive breeding pahs ol Illustrations were made using a drawing tube
i-;iili species, collected by MA-, art* summarised in attached US the microscope. Preserved and living
lahlcs 1 and ?, In each case-, acaUin*i male was first larvae ol /„ \\th^landuh>su fmtn sites I u were
collected al night, then a gravid female was found examined for comparison with those Irom Ihe type
durin;: d oylight the next day. in (lie same vicinity aa bhmIh\ and measurement* are given m Table 6.
the male The pans were each placed i^ a large ("ceding and swimming behaviour of several larvae
ipfl&icd ptolU b&g ejOnftU&iflg siieain water, fi flat of boih speuesw.r-obsefved mcaptivity uud in Mien
ruck and aquatic vegetation. The bag was covered natural Mfc environmenl.
Willi opaque material lor the duration ofamplcxus.
Adventsnwtu nd(\
(Jtttriti \iih\;ktntlitlo\u 'Ilie calls off. \tih^luruhiit>Mi were recorded at a
SlftgCS |*23 (Gosncr, (9601 were studied ffOW lapcspeedof 4 7fruiiMf.c"'. usiu^ aSons Tt-ttSfRO
three separate £gg masses one from (lie type locality portable cassette rccordei with a Lher M5I6
(locality 10. Table I i and die others from \nc new microphone and a Grampian paia6olu lerlivtoi.
localities 3 dlld 4b Hereafter, numhered localities Calls of /.. L'hr&p& were recorded with a Nagra 4 2
will refer lo "fable I (unless otherwise stated*. open reel lape recorderal <t tapespeed ot |4cm m\
plirthci '-.aiupte9 froilleeg masses foundill (liestream and a Beyer M-XN eardioid dynamic microphone.
al locality # were maintained until stage 25 to for L, suh^lunduh'SOy the tape cassette was
confirm idcniitv. b\ I> A. White ot the National replayed on a Nuknnuchi Dragon kipe deck, and lot
Patks&Wildlife Service. NSW(NP.VWS). I'mbryos /., i ///r//w. the open-reel tape was replayed on either
and larvae Were held in dishes (40 cm dram.) attevOVB 77oruSo.n'!(. S I0O l;ipe nxoidei. the
i'oni.iiMimj Mui.iiM Wiik-r. toekv sediment^ and calls ^rffi analysed Oil a Kay r:lcmelnes Digital
aquatic vegetation, and maintained al 14 21 C Soiiii-Graph. Model DSP-5^00, Addiltonal analyses
(locality 9*. and 15 -24 C (localities 3 and 4b). ol waveforms were made by way ol a Sound-Blaster
Phc egg nwws TOW locality 4b laid on 7 aii.1'/Htl K16 caid (Creative Technology! installed in an IRM
< lahle Ji. was submerged wilhm a metal lea strainer -compatible desktop computer, and using the
in thetool, flowing w.iter*.)'the stream I'm the initial Wiivv Studio (CreativeTechnology and Spectra Plus
>
two days ol development, but bolh Ihe egg masses Professional. Release 3.0 (Pioneer Hill) software.
from localities 3 and 4b were maintained at higher Both systems yielded consistent results tor analysis
temperatures of up to 24"C away Irom the stream ofthe same signals.
from the third day alter deposition. hmhryoiOL Ihe dominant (= peak) frei|uencu-\. wetv
development was observed under a Wild \!J calculated as those ol greatest amplitude in a power
stereoscopic microscope, spectrum or an averaged spectral display. Numbers
of pulses were determined by inspection ol
Liioriit ettnuiu waveforms. Pulse mlcs were calculated from the
Stages I 25 were studied from (wo egg masses interval between Ihe peak of (he first pulse and the
Iron Darkes forest and one from Ourimbah peak ol the last pulse ina pulse tram and the number
i
(localities I? & 16, Table 3). Samples ofeggs found of pulses reduced by ona tie, n-1 pulses) Because
fullered overthe substrate in Ihe stream were raised of the difficulty in determining ihe beginnings and
to stage 25 io confirm identity. Adults in breeding ends (i.e.. zero amplitudes! of pulses and pulse
condiliun were placed in an inflated plastic hag trains, the peak - pc;tk interval was taken as the
covered with opaque material during amplcxus and duration. Where appropriate, pulse trains arc termed
lUcic^uUinc embryos maintained al lb -23 C\ 'notes'. II two distinctly different types oftemporal
unit are present in acall, then Ihe signal is described
Liirvar a* diphasic isensit Uttlejohn Sf. Harrison 19851,
tadpoles were measured (lo 0.1 mm) with vernier
callipers and an ocular micrometer attached lo the Results
microscope They were anaesthetised in Chlorbutol
solution before preservation in 37/ lotntuhn. The Di>lrdvitton undhahitui
slaving system is that of Gosncr (IVoOl. LUoriu snlygjitndnlosn
Abbreviations for larval measurements shown in The new loeahues{ I-9)recorded inTable I extend
Table 0. follow Anslis (1976): I'L - total length, BL the known southern range of this species about I.H0
= body length. BD - maximum body depth, TD = km All localities are permanent streams/rivers oi
maximum tail depth. TM = tad musculature depth basalt ormetamorphic rock country associated with
(measured in line with TD). IO = inierorbital span. rainforest, montane or wei selerophyll forest (except
IN = intcrnarial span. LN - the distance between eye for Ma iSc Mb) anil are al S10 m or hicher. The
go M AKSTISA M I I IT!I PJOIIN
Observations by John de Bavay and Paul Webber
confirm thai (herehas been littleevidenceofthis frog
overrecent years at the type locality, suggesting lhat
the species may be undergoing a decline there. The
National Parks and Wildlile Biodiversity Study has
records ol live nudes of Ihis species calling at three
sites on 2.i\.l995 in the Styx River Stale Poresl in
(he region ofthe l\pc loealiiy:
Holy Creek -"lat./long. 30 _W 39" K, 152 14'
I j
4VS. (altitude 1(Kid ml
2> h'elv Creek - 31]' 35' 26" b" 152 13' IK" vS,
(890 m)
U
3) Watile I Camping Area - 30" }5'IS"P, \?1
l2'3X"S(X70m).
Observations on 20,\ii.|994 at localities 12. 13b
and 14 (all northern localities), indicated the
presence olL. \nb^iaiuin(nsa tadpoles
Litoviti i iiVDpNaMV
Specimens D33&66 fl.'ppor Allyn Riven and
FH2664-65 (Wombat Cicek) weie examined ami. on
O — L.citropa die basis (4* the indisiincl tympanum, promineni
*=tiMibgtaiKfulosa sLupHriailh\)>}iuntp)a<lnuiict>Kldo-ldNaMndVh1e)a6d70w9i-dt1h0,cwietredeblyouCnodpltcainbde
1 1957) as (. < ifrofHi from near Oration in the north-
easlcoasl ofNSW. form (he basisofihe slalemenl by
Heaiwnle ti uL (1995) (hat L vflrdfHi "extends from
100 200 norlhcasicrn New Souih Wales (o southeastern
Victoria". Upon examination, these specimens were
I irl.uIe.nAu ruevfibsigolnaomlh'(ihUewdisptrroibvuitdieodn <>b)'yItTtyorluetr<'&7//tAyrWttsttruhd huuid l»' have (he body proportion Ctf /
i l<W5k incHnhny ;t numberofne% Inc;i1ilux lur/. fcufa*. suhi>Uui(/ulifsat but because both were collected m
iatiflufo.sa. 1X65 and in a po<»rstate ofpreservation, it is diffieuh
in come lo a dellniie conclusion as to their identity.
The iwo species have not been found in syiupauy ai
southernmost locfiljl) .il Wbicll the $pCCh$& has been any siteexamined, and this fact, in coihhiiiation with
lound is locality c>. Kal Brook, Mounl Royal Slate the examination o\ museum material, indicates Mui
Porvst. NSW. Hie National Parks & Wildlife North- the drainaec o( the Hunter River appears to be a
east Forests Biodiversity Study ( 1091 I'ftH) riscprds natural geographic barriei separating (hem iFie.l)..
/.. siibgUineiuinsa at a number of sites between ihe Larvae were observed by MA. on Pi.1976 and
Harrington Tops region and Ihe northernmost forests 25.i.1996 at Boardinghonsc Dam in the Watugan
«»i nsvv, including DoyJcs Rjvftr state Forest, mi Slate Poresl, south ol the Hunter iivei NSW (33 00'
Boss Suite Poresl, Nowciidoe, Wcrnkinihc National 01"H. 151 24'l5\S)and by R.Wells lurlhcruoiih
Park Gibraltar Range National Park, Sty \ Stale in the Pokolbin Stale Forest, near Cessnock.. in
Pnrcst Spirnho State Forest and Boonoo State January 1993. Ibis fc ihe noiihcrnmosi known
Forest. Phis indicates the species has a fairly locality lorthis species.
continuousdistributionalongdierangecountry, from Lttorid stihgUimlulosa appears to replace /.. citro/ut
locality 9 in the south lo near Slanthorpe. jusl north in the Bamuglon lops region north of Newcastle
.,( the Old/NSW border (151 40'30"h, 2S'40'20" (lig- 1)_ /.. tirrttfhi occupies a wider variety of
SMl-ip. 1). huhitau than L, .sttbxhitulalosa, including permanent
On a daytime visit to localities 10 M-lTu I I on streams in basalt country associated with wet
19 \h.I994. no tadpolesofIbis or olhcrspecies wcic sclerophyll or rnontane foresl, to similar streams in
located. This was at a tune when numerous tadpoles sandstone country. Although found at an altitude of
of L. MfhghHNfftln.stt, L~ h{)f>moh>m;fnsis and 1066 m at Abertcldy, Vic and Rlnekhemfv NSW. /,
MtMipliyc.s hitlhus would he expected !o be present iinttpd alsobas been lound m lowercoastal areas lo
(based on annual studies in (be 1960s and '70s). 50 m (locality 16, Table 3>.
BKI-LDINC UIOLOliV Ol t'lUUANW'iOW A I., t </hy>r\ '•I
Atltili vchmr in life distinctly polyphonic chorus Thecallsoi frogsat the
Iitnriti sah^hintlulesa souihcrn localilies could nt>( be diHercnliulcd from
Specimens from northern localities werti those of males ,u the lype localilv.
predominantly grefiji. whcicus those from mid-multi Malesobserved calling al nightwereoften petched
i <>,isi;tl localities ( I 9) ranged I'rom uniform golden on broad leaves of trees and shrubs approximately
biown with scaltered darker imtilling o\c\' the 0,> 1 ,5 m above streams, on terns at the edge of the
dorsum, lit specimens vvilh some small areas ol stream, or on vcgclulion lurlhcr from the waters
given often alone- the canthus rosiralis or underthe edge. They were frequently found calling in small
eye. Two males liom hiealiiy I each had ;i broad groups, iwo or moiv metres apart- On 22.\AW-\ at
dorsal paich ol i-jven over ihe head or dorso lateral Inealiiy 5K. 40 males were catling at riigtil in groups
ledums Two specimens. AMR7ro M), fuan •)f up to six alone; a 50 m >nvtcli of the stream iS.
tiloiiecslerTops NSW. and another observed b> II. Ciow pers, eomm.». Al locali'y 7b on 7,\i.I^'M. lour
limes (NKVAVSi al Fal Brook (locality S?K were males were ealline 25 m apart (K, Thumm |vr>.
umlotm bright gisctl, apart rfbm the ehurucU'ii-..u comm r
ijolden dor.so-laleral snipes. An analysis ol the advertisement call is piostded
Some vuldemhrown specimens dcvclopc. large* below and comparison made with thai <d / fftm^i
hntihl green palehes over llie dorsum al nielii iS. lw<> adiitliiuud call sequencfs. alliibutable 10 /..
Cow pt rs, coium.k The inner surfaces ol the hind stfh<;iiitutf>h>\iiy 3TC in the Btoaeouslic Library ol Ihe
limb and gTQiil area were uanslucenl yellow, as Depal'tutenl of Zoology VniNersKy of Melbourne,
touud in adulls irom the type locality. both recorded bv M. J. Liltlcjolui and bis associates.
The first, from tiuy Kiwke> Creek hbor NSW iMi
24' 20" L. I.S2 20' 4h" Si \\<i\s a-emded on
l.iioriu titrofht ha.-v a uniform golden brown dorsal 2JS.\,1^64al awel bulbairtemperatureot :<S C.and
W
colouration Iwiib green along ihe canthus rosiralis Ihe second, from Flat Rock Creek H km t^'Poiiil
and sides of the bodyi. similar to most specimens of Lookout NSW (close to ihe f&W silt), on IL\.1V63S
/. sith^/atulnhtstt from localiliC-% 1-9. The principal ai a wet bulb ail lemperaiure of IVC They ate
difference belween the specie*, is the colour of the similar in all pertinent respect> to (he ljII ilcseriheo!
inrici surfacesolThehind hmb and eroin, winch m /.. here.
i iit.'fii is brick red.
Lt!<.-riii ci/ropa
I ttftfHH >t<'tntt\ Males al Darkes horest (locality 15. /'able \\ were
l.tlcnn mkglttitthifoiii obsei^ed dunn» s[iriuy and summercatlm;j lo>t(j low
Culling heems in spring and vvns observed on branches neside Ihe stream,on rocks near theotj&eol
20a.I91>4 at liualiiv X. when water icmperaiuies al the ^.itei. iv on exposeil rock shelf in midstream
IligllC were very low. e.:: . n C and the div bulb an clove to shallow, slowly flowing water \> \\n\\ I
temperature al locality 5b (1900 h) was 9,8 C i.A. MibyJiittJiilt^tt nulcs called while two or more
While. S, Cow |vis_ LOinm.t, Olhei obseiVatioUs by melres aiarr and activil\ mcicised on oVercasi
M \. al ihe type locality during annual three-week evenings during or aflcr rain. Pry bulb air
periods (Dec/Jan. I9fi(i-74). and al all other temperatures on several nighls when males were
localities listed in lable i. mdieale ihut calling calling in Septembei LXxernbet. 1972-1VS0. wete
persists ilirouidiout December/January ina variety of 14 22 'C No diurnal calling vsa% obsetved.
ur.iihcrconditions, with increased i.etivily during,or
aflet. heht rum. livening dry bulb air temperatures Aihcrfiu'iwtil *v(h
taken during pcrn'dsol spring/summeraclrvily al the l.ctena \nbitlutuhilosit
localiOlies in Table I wete |V"C-I9.5~C (mean Tlte .idvertisement call of /.. wfa$IiWfItfif}.st{ was
15 7 At the lower temperatures (>V-I4"C). recorded by .1. Courtney al Diehard Creek, Clen
calling wa.s less intense and by aural comparison Innes (locality Hai. on 20.\»3. The dry-bulb au
onlv, notes were al a slower repetition rate. icmpcrature was I V C\ The lollowine, data were
S(M»iadic dunoal calling was common during the obtained from the fourth call ill Ihe sec)uence (I:\ii.
bteedmeseason bul males were most active at me,it. ^A). The call hasa duration ol 9.375 s and consist,-,
Diurnal calling look place from concealed positions of 13 pairs (doublets! of pulse trains (notes), svilh
such us under rocks or from within vegetation, either each of iho.se in the first five pairs all beina of
near the siieam. ot at time* up to abotil i'om meues relatively low amplitude ibi", -mi. In the subsequent
away from the water. A single male or a small seven pans t)l noies. ihe second note is of much
number oi individual-., called from as early as 0742 greaiet ampliiude than the firsi. Thus, all but one o\'
h (e.g.. locality 7b). Nocturnal calling was initiated (he firs! notes (which is of equal amplitude) are
by one forjg, in-imal!*, followed by others m ;i Softer ft 'I' 'he ani|4iliide of second notes bejf\£
M
l)2 ANSTIS & M. J. LITTLEJOHN
Fig. 2. Live egg mass pfLitoria sub#lanchitt/.\u attached to a leaf from submerged overhanging foliage inTuckers Creek,
BurringtonTops ( locality X). Scalebar- 10mm.
