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Modern Megapode Research a Post-Frith Review

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Modern Megapode Research a Post-Frith Review Au.nralu\ittt Biftl R(tittrs Nunlb,J I Coreila,1989, 13(5): 145 154 MODERNMEGAPODE RESEARCH A POST-FRITHREVIEW DARRYL N. JONES Divi\ioD of Au\tralian Enviroomcoral Studics.Criftjlh tlnivcfsity, Nilrhan. eld .llll R..?iril .10ll4ar< h. I98t) lvlodernstudies on the megapodescommenced wth H J Frlhs studes on the Ma eelow Le4roeoce/jala during lhe 1950's S nce th s lirnemuch debatehas cenlredon the evolulon ol lhe lam y, especally on taxonomc relatonsh ps and routesof d spersionlvlore recenfly rnalor advances n the understandng ol megapodencubal on havebeen made, especla y the f!ncl oningol mounds, and lhe associaledadaptalions of eggs and ch cks Detailedslud es of sexLralnteraclions have Teveaed nLrmeToustypes of matingsysteras wthn the groLrp,eading to speculatonsas to their evoluton A/anyspeces arc currentythfeatened and manyareas ol ecoogy and behavourremaln to be sluded INTRODUCTION scattcrcdand oftcn obscurclitcraturc in two com- prchensiverevicws of incubation habitswithin thc Thc mcgapodcs (Mcgapodiidac). also known 'mound-buildcrs'. family (Frith 1956a,1959a). and also speculatccl collectivclyas thc are among on thcir cvolutionary origins. Clarr (1964a) thc most fascinating yet least studied fanrilics extcndod this inforrnationto examiDecritically ol bilds. L,vcl since thcir discovery cluring thc hypothesisof a dircct reptiliananccstly; Magcllan's l5l9 l-522expcdition (Frirh l9-59a). hc saw thc mcthods of incubation as renarkablc thc birds havc fascinatcd and perplexecltravel- sccondaryadaptations, suggcstive of cvolutionary lcrs. nilturalistsand scicntistsalikc. It has been spccialization. their usc of cnvironmcntal sourccsof heat for thc incubation of cggs that has attracted most atten- Clark (1964b) and morc rcccntly Diamond tion. Morcover, nO othcr birds provide so littlc (1983)havc provided conciscsummarics of the parcntal carc as thc mcgzrpodes:eggs and chicks researchconducted during thc I960's anci 1970's. are cffectivcly abandonedat the incubation sites. During the past dccade. howevcr, nany ma.jor Initially rcgardcd as anatomically archaic (Clark advancesin thc understandingof thc physiology. l96.la), this apparcnt reptilian trait was takcn as ecology and behavioul of nunerous nrcgapodes furthcr cvidencc of the primitive stagc of have bcen made. It is thc aim of tlris rcvicu' to evolutionary dcvclopment of the family (Frith provide a lrrief ovcn,iew of thc most pertinent of 19564;Clalk 1964b). thcse discoverics. This cannot prctcnd to be exhaustive; rathcr, the intention is Allh,'ugh therc rrc importrnr cxeeptions to updatc scientilic views of these birds, to stimulatc (Brrrrcttirnd Crrrndrl l93l: Flcrl l0-37:Coles intercst, and overall, to demonstriltc 1937), the foundations of modern megapode thc nccd, potential, and value of furthcr rcscarch werc laid by H. J. Frith. His decade or stutly. so o[ work on the Malleefowl Leipoa ocellata TAXONOMY AND MORPHOLOGY (Frith 1956b,1957, 1959b, 1962a,1962b) drew worldwide attention and providcd the basis for a Although one spccics, the Australian Brush- scientificunderstanding of the family as a whole. Tvkcy Alecturo Lothami,w.\s initially classificdas Frith also provided a detailed coverage of the a vulture (Gould 1865),there has becn little doubt 145 14ti D. N- Joresr Modernmegapode research Corela 13(5) thirl rnegapod!'surc closclv allicclto othcr specics posscssirrga largc lrclr cl likc casqucon thc back ()rdcr (ialliforrres. within the Within this i)rdcr of thc skull. Stark (l98ii) hasshown rhis ro be oi th-c'nlcgap()(les i:re thc ntost clistinctivc fantily pncLrnatizcdbonc. not horn as oltcn rcportcd. (Olson l9l.i0).arrcl. uith thc Ncw Worltl Familr.i. Thc structlll c uDdallachtrcnt of tirccasquc t() thc thc Cnrciclue(chrrchalacas ctc.). hirvebccn classicl skill is analogucoust{) tlltl fi)unclin sornc rvoocl ils thc n)os1plinritil c. pcckcrs und sullgestsir sinrilarshock-ubsorbing Afpurent trlliniticsbctu,ccn thc rncgapoclesancl mcchanisn.'l'hisnltv bc rclatcclto thcir habil of cnLciilshavc l)ccn greatlv strcngthcncclby the hammcring hartl shcllcd nuts obscrvcil tluring reccDltaxononlv of Siblcl. Ahlquist anclMonroc fccding(R. I)ckkcr. grcls.corrrlr.). (l9Sli). In u rc\1rlutionarvstcp thesc autlrors havc 'I'lrc cxtanl Mcgapodiiclac consists oI six scpalatcd lhcse two groLtps liorr thc olhcr gcncril.currcntlv contprising l9 spccics(Tablc I). (iirllilormcs into ir scplnltc C)rcler (thc Tltt trttttll (t'rtt:rr' lhr(( ntllttrt\li\ rir: ( racilirrmcs).Thc '"clacitvol this cllrssilication !rn! A l cc l u rt, Lc ip o tt ttttt-)M ut r ot c lut kt rt. A l cLl tt t u is will ncccssililtctcsting lfotrr othcr sourccsol' lt crrnsidcrctlto bc closclv rclatcd to Ta[a{ulla c\.rdc n cc. 'liid Acpvpodius, a group kriown as thc trrusli-tulkcvs.'lrt All nrcgirpodcsshow grcat strucluralsirnilrritv r.rilr hrrrirtg ;, 111111lrqek rrrtrl lrrr , u lrir'lrnt;rr .u)(lirL rli.lil(tl\ |I I r ' I I , , I r I I \ IL. ti (. in ,rriglsl 11 1,,1 I'riglrtl)e,,i,'ure,l ((,,irl(. lrrNi LleLlrl..r.irrrn l96.lrr). Thc M,rlco Motntct'phulon nmlco sltttu,s and Zimnrernrln lt)E6). Alettrrru arrd thc two lhc ltrcatcstskclctal divcrgcncc within lhc farnily. Acplltodius sl.rccicsalso posscssirrllitlable ncck TABLE 1 l hf Mcllrpodcs: rrmcs. dislribution\ llrrd \lllu\. (icncrLtl distr ibutionr MrllccliNl AuslrrhLr.inland l Au\lrirliaIlIlrush lurkey I: Ar\lrtliir J Rcd billcrl tsf ush-turkc_v J l]Lrck billed Bfush-ttrfkey S \cw (i ui|rca l Brown-billcd Llrushlurkc\ N Nc$ (;uinel l Wrtllcd []rush-trrrkcv trpland Nc!v (i uincu l Ilruiin s I} Lrshlurkcy I Nlrlco : Nicol)u, Scrubfo$ l Nicobrr Isllnd I PhrlippincScrublo$l 1)hilippinc\to Suh$csi I Sulr Scruhl()$l ttarrggriLrrrd Su|, l ()rafigc-l(n)lcd Sct uhli)\r I Kangcun IsLrnd1(, N J Dusky Scfubio\\| N Moluccir\ Ncr (;uinca Scrul)l()wl N Now (;uincr. .l ollshorcislrnds Mclanesirn Scrubft)\vl Krrkar to Sol(,In()n l Isllnds Vanualu Scrublbrvl llrlvncsi:rnScr ubl()$,1 Mrr ir na and l'rlul Island\ Mrcron0sranScr ublowl Nivirlo ou Island. Tongl N{(tuccrn Scruhfowl MolLrccanlsla d\ LAllcr I)ckker l9iilJir:Whitc ar)d Brucc 1988. rl:or rlclailed disrr;burions scc Beehlcr ./ ./1 1986r BLrkers./ d/. 198.1:Coa(c\ 1985r Dckkcr l9tt); Riflcy 1961)rWhirc irnd Bfucc l(186. '(irnsc|r,ation slirtus: l: Scriously end ngcrcd; 2: lhrcltencd. conscr\.rtron nruirsur,rs rcquircdl l: Sccurc hut unconlnon or \omc popLrlittioo\ lhrcltcncd: .1: Widcsprcad and al)undrnt (scc Ci)a1cs1985: Mcgap(xle Specialisl(jK)up 1988) December,19Bg D. N. Joresr l,4odernmegapode research sacs or wattlcs (.Toncs1987; Kloska and Nicolai of thc Phasianidlrc(phcasants and quails). the l9ti8). lilrgcst lanlily in thc Galliforrncs. cxtcrlds west- ward throughout this rcgion with virtually no Thc gcnus Mcgupotlius, or scrublowl, arc thc ovcrlap with ncgapoiles (Olson 1980). Olson str:,llc{lmegirpr)rlts. Geographie \irriati,,n is con- (19U0)attributcd this tlistributionto sonrelornr sitierablc but most are chicken-sizedbilds with of compctitiveexclusion. noting that despitethe short tails and a short pointed nuchal crest nregapodcs superior abilities to dispcrsc ovcr (Beehlerct a1.1986). watcr (Riplcy 1960).phasianids appcar to havc Rclationshipsamong the extant megapodesarc displaced mcgapodcs whcrcver they mcct, thus far from ccrtain. Clark (196:ta)proposed a two preventinS thcir spread northwurds. This picture branch systcm: one sidc diverging to the brush- has been complicrtcd somewhat by rclativcly turkcys with Lelpoa branching off bcforc Alec rccent translocittionsof Junglcfowl (ia11u.rspp. /&r.r; thc other wrth Mu<rocephalonas an early and scrubfowl onto marv Indo Pacilic Islands branch. with thc scrubfowl bcing the most rcccnt (Riplcy 1960). spccics.This provisionalphylogeny is likely to be Thc almost pcrfcctly contplerncntrry distri- challcngcdbv a numbcr of currcnt studiesinvcs- butions of thc nrcgapodcs phasianicls tigi]ting comparittive anatomy and crnbryokrgy, and the is striking. Dckker ( nl,rtPh,rrenctic(l(\r lopment. lhc mi\'ri,\tru!lur(. 19139).howcvcr. doubtcd whcthcr r)l lcillh(rs- rrnJ egg chrrrlctr.risticr ccological cornpctition oflers suflicicnl explanation. Rathcr. hc ar€lucd tltat thc occur- ORIGINS AND BIOGEOGRAPHY rcncc of certain carnivorcs was a kcv lactor not Onc of thc implicationso[ thc rcvisedrelation- ,)nl)ol lhc distrilruti,rrr,'lthe nrc!rrn,'.1t., I'ut rrlr,' ships bctween thc mcgapodesancl other fanilies in the typcs of incubation (ntound vcr.susburrow) (Siblcy ct rrl. l9lill) is thirt of reviving support for devclopcdin the easterndistributcd spccics. a (londwanic origin for thc family. This vicw, Currcntly mcgapocicsarc vcrv widclv clistri rdvanccd mainly by Cracrati (1973), buted thloughout the south cast Asiitn and hypothcsizccla trans-Antarctic clispcrsalhistory Mclancsian islantl chain (Fig. t). from thc ol irnccstral mcgapodes into Australia. Thcsc Nicobar Islandsto Niuafi)'ou.ncar' 1'onga. and birds. anti a suggcstedproto-cracid group, wcrc throughout an area 22 dcgrcesnorth and soutlt of clcrivcd fronr ancestral galliforms inhabiting thc cquator (Blakcls, Davies anci Rcillv lt)t1.1: Gondwanulanilin thc Crctaceous(Cracraft 1973). Rirke lt)x()1.Within rhi. \ir{l irrer.rhr' s1r11iq1 This idca was cliticized by Olson (1980),who havc splcad onto nrany cxtrcnlcly remote and favourccla North Amcricalr ccntrc for the cracicls, small islands as wcll as occurring on nost of thc arguingthat this group would not havc becn in largcr land nrasscs.C)lson (l9lt0) and Dckkcr Soutlr Amcrica at a timc appropriate for dispersal (1989) provide thc most accurate maps of acrossAntarctica.
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