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A kingfisher (Halcyonidae) from the Miocene of Riversleigh, northwestern Queensland, with comments on the evolution of kingfishers in Australo-Papua PDF

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Preview A kingfisher (Halcyonidae) from the Miocene of Riversleigh, northwestern Queensland, with comments on the evolution of kingfishers in Australo-Papua

A KINGFISHER (HALCYONIDAE) FROM THE MIOCENE OF RiVERSJ>ElGH, NORTHWESTERN QUEENSLAND. WITH COMMENTS ON THE EVOLUTION OF KINGFISHERS IN AUSTRALO-PAPUA WALTER BOLES E. Boles. WE. 1997 06 30: A kingfisher iHaloyonidaei Truni the Miocene of Rivcrslcigh, norlhwesicmQueensland,wiihcommcnlsonrhcevoluiionolkingfishcrsinAusimlo-Pupua. Memoirsofihe QueenslandMuseum41(2);229-234. Brisbane.ISSN0079-8835. A Miocene kingfisher from Riverslcigh, norlhwcstern QueensiantI, representedby a com- plete carpomelacarpus, is ihecariicsl record ofthe Hakyonidac from Australasia, h shares similariiieswithseveralmodemgenera,buiapositivegenericidentificationcannotbemade. Although it can bedlstingiiislied froin exiaiii species, this skeletalelemem is insufficient to creel a new genus. A processusdcntiformis in Tanysipwraand Mcluiam and itsabsencein Todimmphusandothergenerasuggest that the lormcrgeneraarcamongthe more pnmitixe of the Ausiralo-Papuan kingfishers. The less developed processus denliforniis in the Rivcrslcigh specimen is consistent with it being an earlier member of ihe Todiramphus lineage. Ofliving kingllshersexamined,all that retain the processusdenliformis are inhab- itantsofrainforest, \2iKifi^fisher. HakyofUdcH. Hivi-nletfi/h Miocene, evolution. Waiter E. Boles. Attsmdum Mustum, 6 C(dlege Street. Svdney N.S.W. 2000, Australki: received4NovemberJ996. The kingfishers (Alcedinidae s.l.) are subdi* bones largely follows Baumel & Wilmcr 19^>3). vicled into 3 suhfamilies. DNA-DNA hybridisa- Todiramphus and Syma are considered d(istinct tion studies (Sibley & Ahlquisi, 1990).suggested from Halcyon, following ChrisAtiMdis &. Boles that these should be recognised as families. 1994). Institutional prefixesare (Australian Cerylidae do not occur in Australasia. Al- M(useimi). ANWC (Australian National Wildlife MV QM cedinidae {/river kingfishers*) and Halcyonidae Collection), (Museum <y\ Victoria), i= Dacelonidae auct] ('tree kingfishers') arc rep- (Queensland Museum) and USNM (United resented in Ausiralo-Papua by 5 species in I States National Museum). genus and 21 species in 5-6 genera, respectively (Bcehleret al.. 1986; Fry et a!.. 1992; Christidis SYSTEMATICPALAEONTOLOGY & Boles. 1994). There are no named Tertiary forms from out- Family HALCYONIDAE side Australasia (Olson, 19S5) Mourer- Although the Halcyonidae includes some ofthe Chaurvire {\9^1) listed this family (Alcedinidae largest kingfishers in the world, si/eisnot avalid s.l.) from Eocene-Oligocene deposits at Quercy, character for family allocation o\ osieological France, and Olson ( 1985) noted thai he had ex- material. Australia's 2 Alcedinidae, Alcedo afmroimnetdhespleowceirmeEntsx:celnoeseotfoNtohristhfaAmmielyriocriagainnadtitnhge pkuisniglfliashaenrd .s4,peacziuersen.(wairnegthleecnogutnhtsry5's5msmmallaensdt medial Eocene ofGermany. All Australian Qua- 75mm. respectively), but the closely related A. ternary kingfishermaterial isreferable to modem websien of New Britain has a wing length of taxa: Alcedo cizurea, Dacelo novaeguineae, Todiramphus pyrrhopygia and To, sanctus (9e(.)gm.m,,oTvocdrilraappmipnhgusinsmia/celtehaevisim,allweirnhgalclyennngitdhs (Baird. 1991 ). NoTertiary kingfishersareknown 90mm). from Australasia (Fordyce, 1991: Vickers-Rich, The carpometacarpus of the Halcyonidae can 1991). bedistinguished fromthatoftheAlcedinidaeand Described herein is a Miocene kingfisherfrom Cerylidae (Table 1) and on this basis the Rivcrslcigh. northwestern Queensland. Rivcrslcigh Ibssilisassignedlothe Halcyonidae. METHODS HaLcyonid gen. indei. Fig. IC Measurements (Stcadman, 1980) were made with vernier calipers accurate to tll)5mTn and MATERIAL QMF29719. rightcarpometacarpuswith rounded to the nearest (11mm, Temiinolooy of only mmor abrasion to some surfaces from ?niiddic 230 MEMOIRS OFTHEQUEENSLAND MUSEUM TABLE I.Characters forseparatingthecarpometacarpus ofthe Alcedinidae, Cerylidae and Halcyonidae. Character Alcedinidae Cerylidae Halcyonidae proximalborderofdorsalcarpaltrochlea moreangular morerounded moreangular developmentofos metacarpalisalulare moregracile morerobust morerobust orientationofosmetacarpalisalulare morecaudal morecaudal moreproxunal lipofprocessusextensorius rounded expanded,slightlyrugose rounded positionofprocessusintemieiacarpalis moreproximal moreproximal moredistal widthanddistalextensionofsulcus narrower,notas broad,extendingalmost broad,extendingalmost inierosseus extensivedistad tofaciesdigitalisminor tofaciesdigitalisminor planeofsynostosismetacarpalisdistalis OSmetacarpalisminor OSmetacarpalisminor anddistalendsofosmetacarpalismajor depressedbelovs plane depressedslightlybelow Hat,coplanar andOSmetacarpalisminor plane Miocenetoearly laic Miocene Last Minute Site from quent comparisons involve Todiramphus, Syma, System C (Archer et al., 1989, 1994). This site is Tanysiptera and Melidora. interpreted to represent shallow pools oreven emer- The fossil (length 15.8mm) is in the size range gent accretingsurfaces,and isdominatedbyterrestrial of Tanysiptera (sylvia 14.0-15.2mm; gaUitea vertebrates including the possums Djilgaringa 15.2-16.5mm) and Todiramphus [sanctus 14.8- gillespieae Archer el al., 1987 and Sirigocuscus reidi Fhuinery & Archer, 1987. Other avian laxa from this 15.1mm; macleayii 14.5-15.7mm; pyrrhopygia site are a range ofpasserines, including the logrunner 15.7-16.5mm; chloris 16.5-17.8mm). It is larger OrfhonyxkafdowinyeriBoles. 1993. than Syma (torotoro 13.5mm; megarhyncha \AAxw\x\)dW(l^m'iX\\cx\\\ixnMelidoramacrorrhina DESCRIPTION. Length 15.8mm. Length of (18.9 mm). It differs from Tanysiptera and spalium intcrmelacarpale 61% of length of car- Melidora and resembles Syma and Todiramphus pometacarpus. Processus dcnliformis low and by being more slender and by having spalium pointed, locatedabout midwaybetweenthedistal intcrmelacarpale longer relative to the length of edge of facies arlicularis alularis and the carpometacarpus and the dorsal rim of trochlea carpalis extending less dislally relative to the cranialmost point of facies arlicularis digitalis major. Os metacarpale majus of about equal ventral rim. The fossil differs from Todiramphus and resembles Tanysiptera, Melidora and Syma thickness forentire length, in ventral view. Spat- by having processus dentiformis present. This ium intcrmelacarpale gradually becoming wider process, however, is narrow and pointed, rather distally. Processus intermetacarpalis far proxi- than broad and tlal as in these genera (and larger mally in spalium inlermelacarpale. than in Syma), and is situated more proximally REMARKS. Among extant Australo-Papuan relative to the spalium inlermelacarpale and pro- Halcyonidae. Tanysiptera and McUdora have a cessus intermetacarpalis than in Tanysiptera and low. Hat processus dentiformis, Syma has an al- Melidora, and more distally than in Synui. From most non-existent processus dentiformis as a all 4 genera, the Riversleigh specimen differs by barely raised roughened area, and Todiramphus having the caudal edge ofos metacarpale majus and Dacelo, as well as Afro-Asian Halcyon, lack siraighler and less caudally concave, making os il (Boles unpubi. data). X-!"ay photographyof metacarpale majus thicker and spalium inter- studyskins showed ActenoidesandLcicedo have metacarpale proportionally narrower relative lo a low processus dentiformis. but Halcyon the width ofthe carpometacarpus. (Pelargopsis) and Clyrocey.x do not. The x-rays, The significance ofthese differences is uncer- tain. They are individually minor, yet within the while sufficient fordetermining this process, are Halcyonidae the amount ofvariation in this bone not adequate for detailed comparisons of these is little so that these differences may assume laxa. Other than size, there are not substantive greater importance. Variation in the caqximcta- differences between the carpomclacarpi of carpus, however, is not representative ol' that of Todiramphus, Dacelo (and presumably thercmainderofthe skeletonorindeed the rest o\' Clyioceyx)\Symadiffersonlyinitslowprocessus the morphology (Boles unpubi. data). dentiformis. Because Dacelo and Clxfoceyx are The fossil cannot be assigned to Tanysiptera, considerably larger (Z). novaeguineae 32.5- Melidora, Syma or Todiramphus and can be dif- 35.6mm), they arenoiconsidered further. Subse- frereniialed from extant species ofthese genera. A MIOCENEKINGFISHER FROM RIVERSLEIGH 231 FIAG,. M1e. ClairdpoormaetmaaccarroprirhoifnAaus(lrAalNoW-PCapuCaOnRSha-l5c3y)on.idBk,inTgafinsyhseirpst,eirnavesnytlrvaila v(iAewM.O.A6ll09r2ig6h)t.sidCe,exRcievpctrsfloerigBh. Halcyonidae gen. indel. (QMF297I9). D, ToiUramphussanctus{MA0.51\%2). E,Dacelonovaeguineae (AM 0.59217). Scales =5mm. Overall it hasthegreatest resemblance tospecies mostasancientasthe mid-Cenozoicoriginofthe ofTodiramphus in size and morphology, despite Alcediniformes [sensu Feduccia( 1977)]'. By the the presence of a small processus dcntiformis. early Miocene, the present geographical config- Given therelatively limited importanceofcarpo- uration of Malesia had been reached. This, in metacarpal variation in the halcyonids and the Fry's opinion, provided 'ideal circumstances for limitedfossil material itisimprudenttorecognise the multiplication ofspecies, resulting in a fauna a new genus al this time. offorest-dwelling, non-fishing kingfishers ... At some more recent time, perhaps about the mid- DISCUSSION Pliocene, this fauna gave rise to a lineage, Hal- cyon [s.l.], adapting to more open habitats'. To place the Rivcrsleigh fossil in perspective Under this suggested sequence of events, the within halcyonid kingfisher evolution, the prim- rainforest-dwellingTanysiptera,Melidora,Acten itive members of the family must be identified. oides and Lacedo would be among the more Fry (1980a, b)employed 3 criteriaforthis. Prim- primitive genera. Todiramphus, included by Fry itive kingfishers have 1) generalised diets and (1980a) in his open habitat Halcyon, would be relatively unspecialised modes of foraging (i.e.; morederived.PresumablySynia(alsoincludedin sitting and pouncing, non-fishing); 2. stable hab- Halcyon sensu Fry [1980a]j also wasconsidered itats and not ofrecent origin (rainforest), within by Fry (1980a) to be a more derived taxon. Al- which they may bediscontinuously orrelictually though entering open country adjacent to forest, dislributed;and3.oligotypicgenera(i.e.withfew the two speciesofSyma are essentially rainforest species)withoutcloserelatives. Fry( 1980a)con- inhabitants, particularly in New Guinea (Coates, cludedthat kingfishers(all familiesasrecognised 1985). here) arose in Malesia, the area between Indo- In addition to sharing the primitive characters china and the Coral Sea. proposed by Fry (1980a, b), species of Tan- Prominent among the primitive forms were ysipteraandMelidora (as well asActenoidesand halcyonid kingfishers, many o\ which live in Lacedo) have a low but well-defined processus Malesian rainforests. Fry (1980a) speculatedthat dentiformis on the carpometacarpus. This is ab- 'the Daceloninae [= Halcyonidae] have a history sent in the otherhalcyonid genera examined, the of evolution in eastern equatorial rainforests al- Alccdinidae andCerylidae, and someothercora- 2i2 MEMOIRSOFTHEOUEENSUNDMVSEUM ciitbrmfamiliesleg.Momotidac.Coraciidac).If Bell (1981J alsoconsidered that MflUhntwas any of ihese fyrtiilies is used for outpA^up com- closest to Tanysiptera. He iK>ted thai the call parisons, the suggested polarity oi Ihc processus notes ofthese two species were similar and the dcnliforrnis h thaiitspresenceisdcnved. A sim- distress notes identical. They have simihirskele- ilarcomparison usingolhcrcoraciiform families tal proportions, particularly in therehttivc length (Tod due. Pho^niculidue, Upupidac, ofIhe legs wlienconiparcd toTodiramphus,Hid- i Bucerotidae). in which the structure is present. cyan or Pacelo {Boles, unpuhl. data), These two ^ivt*N the opposite conclusion: presence of the genera thus have more similarities than might lie pmcessus dcnliforrnis is primitive, its absence immediaiely obvious, They also share habitat derived. Within the Mcropidae, the structure is prelercnces. Although Mvlidora martonhiua prcseni in some species {Merops ormtms) arni and Tanysipteni species will enter miingrovex, ahseni in others {M. optaster). The significance teak pl.iotations and drieradjacent country, they ofIhisvariationisunknown, asarcthe tunclumal arc primarily occupants of rainforests. Lacedo aspects ol the processus denliformis. Thus itie puJchvllaund the6.speciesi)iAnefundt's inhabit polarityofthischaracter'spresencei.snotknown. rainforest, preferably in a primary, undisturbed (This, oi course, assumes thai the processus slate deniilormis is homologous across ihe order. Species oi'Todiramphus are unifonn in pUnri- Whetherthis isso, and what relationship it hasto age. Most have a variation ofthe basic pallcni of tlie siiniliir process lound in the majority of the greenorblueuppcrpartsandwhileorlightorange Passeriformes, is unknown.) Within the underparts and collar. Several subgroups can be halcyonidkin^dshcrsthe presenceoftheproces- discerned, but Ihey still show only small diver- susdenlifontiisexhibilsastrongcorrelation with gences from thisgeneral form. These species arc Fry's(I980a,hj primitivecriteria. almost all sit andpounce feeders, Habiiai prefer- Suporficiallythereseemstobelidleinconmion ences aniong species arc more varied, ranging exieriially between Mciidora and lanystpwra Iromrainforesttottpcncountry. Williin Ausiralo- beyondthehiisickingfishersimilarhics. Hachhas Papua. hi»wever, fewoccurinrainlorestandthere specialised generic characters: a hooked bill in isadecided biastowards open loreM. woodland. Mvlidom (Hooked-hilled Kingfisher) and elon- mangroves, clearings and open country. Most gated, spalulaic central iccirices in Tan\sipiera rainforest inhabiting forms are found on islands (paradise kingfishers). Metiiioni macronhimi is ofthe southwest Pacific. a ralher drably coloured species, Otherihan blue The processus denliformis in the Riversleigh scalloping on the crown, Ihcplumageisacombi- kingfisher is smaller than thai in primitive mod- nation of bixjwns and white. The underside is em torms. This could indicate thai it has been white, while the hack, rump. tall, and wings arc undergoing reductionsincethe splitofitslineage dark brown with paler broAO Vv-alloping. This from that of TanysipK^rn-Mfluiofn. In this re- plumageisquite unlikethatofthepariidise-king- spectitisconsistentwiihwhatwouldbe[>Tcdicted lishcrs'/anysiptera,adultsofwhicharcstrikingly foraprimitivespeciesofTodiramphtis.Thebone patternedinunmarkedbluesandblacks,andusu- exhibils a hirgely lodirarnphtts chacacict while allyeitherwhiteorbuff/rose. Thejuvenile plum- retaining this more primilive halcyonid lealure. ;ige of Tonysiptera, however, is brown with The ideniilicalicvn ofthe Riversleigh fossil asa scalloping,andFry( l^SOb)was 'impressedbyits halcyonidiscompatiblewithFry's(1980a)inter- {MvHdora''s\plumageresemblancetothedistinc- pretation, as IS considering ihe piesence of the tivejuvenile of Tanysiptera i^ahfea\That these processusdenliformisas primitive. According to two genera mighi be closely related was sug- Fry's scenario, un early Miocene kingfisher gested by Fry (1980b). who thought it 'possible shouldbeapi'imitiveform. Hiscriteria, however, that Melidont and Tunysipteru are of immediate arenotusefulinthissilualion.The foragingmeth- common descent and the former is a specialised odsofthefossilcannothe determined, norcan its derivative that has retained, in the adult, the an- systematic isolation be ascertained. The cesiral juvcnde plumage'. The presence ol sim- Riversleigh habitat is considered (Archer cl al, ilar plumages is also evident in female Lacedo 1992) to have been rainlorest. but thisciinnot be and Nume species ol Actenoidcs. niMably A used as a characior for making a taxonnmii. de- pHmeps and A. Undsayiofall ajtcs.This resem- terrninaiion, blance beiwecni Actenoides and Ikmysipura and Although this fossil permiis idenlilicadon as a l»eiwe«n LmTdv and MvUdont w;js commented halcyonid kingfisher, it is noi clear whether ihis onhyFiry(1980h). species belongsloanexistinggenus orshould be A MIOCENE KINGFISHERFROM RIVERSJL£1GH «3 allocatedma new one. Thepresenceoi a feature BAUMEL, J,J. & L.M. WITMER. 1993. Osteologia. found in living primitive genera and ihe Publications of ihe. Nuilal! Ornithological Club mhkiaonvryete'ibpserheiemnri'tsriatvi;enK.l\fb*roicmisrirt.ensTccuhgiigsnesw\^thaciothnasiisisciiio.cnntsloiodw.ietrwhe;idnthleou BEEH2DL3AE:R.4,51-91BS3.62M.,.B.irPdRsAoTfTN.ewTGKuin&ea.Z(IPfMiaMeEelRoiMtAUNm.- versity Press: Princeton). sicqucnceofevolutionarycventssuggesltidbyFry BELL, H.L. 1981. Jnfurmation on New Guinea king- fishers. Aleedinidae. IIms 123; 51-61. ACKNOWLEDGFMFNTS BOLES. W.H. 1993. A iogrunncr Onhoffvx from iho Miocene o( Rtvcrsicigh. nonhwesiem Qucen*^ land. Emu 93.44-49. For access to specimens I ihcink Wayne CHRISTIDIS. L. & BOLES, W.E. 1994. laxunoiny Longmore(QueenslandMuseum).Jerry vanTcts andsi>ecicsofbirdsofAustraliaanditslerriltiriev and RichardSchiulde (Australian National Wild- RAOU Monograph 2 (Royal Australasian OnU- hlc Culleclion). LesChrislidis and Rory O'Brien tJioIogists Union: Melbourne). (Museum ofVieiotia). itnd Slorrs Olson (United COATES, B.J. 1985. Thebirds ofPapua NewGuiiitM. vol (Dove: Brisbane). sSftueanuldmessp1N0raotsviuiopdnpeaodlrtaMtuvhsiesenuurmees)e.ianrTcwhhh.eiTcAhhuestlorRaiwlvoicarrnsklMeaiung-dh FLANptNlfiiEdailRia.Yangn.edTridTFsr,U(&hMoaArsRsunCrpuHisaElRidai,:cMkPM.hmui1l,9a8ntg7wconS)tirnidefewiao)altif.s"r*aio.siimlT material was collected via an ARC Pnj^Tainme the Miocene of noithwcsiern Quccnslajid. Pp. Gtant 10 M. Archer; support Irotii the University 527-536. In Archer. M. (cd) Possums and op<j.s of New South Wales; a grant tiom the rvp;iri- sums; .Snidics inevolution- (SurreyBratly:Chip- ment of Atls, Sport, the r-lnvironmenl. Tounsm ping Norton). GaonddlhTeelrrpi;toarigersunttofMromAtrhceheNra.tiSo.naHlaEnsdtataendPrHo.- FORDbYraCiEe.reRc.oEr.d 1o9f9N1e. wA Zneeawlalnodo.kPapt.thIeIf9ois-sIi3l1(v>e.acI-n jjratnme Grams Scheme to M. Archer and A. Vickcrs-Rich. P.V.. Monaghun. J.M.. Baird. R.F, Bartholomai; andgrants m iiid lo the Riversleigh &A.nsRltrcahla,siTa.H(.Pi(ocndse)erVre)ncesbiginai.eSmdpiatkrieMoenltokl^ougmyeo)l Research Project from Wang Australia Ply Ltd, FEDUCCIA. A. 1977. A model for the evolution of ICI Australia andthe Auslralian Geographic So- pcreliingbirds. SystematicZoology26: 19-31. ciety. FORSHAWJ.M. 1987. Kingfishersand related birds. vol. 2. Alcedinidae: Hakyon to Tfinyxipk^m, REFERENCES (Lansdowne: Sydnev). FRY. C.U- 1980a. The originofAfroimpiealkingllsh- ARCHKR, M.. GODTMULP, M, HAND, SJ. Si crs. Ibis 122:57-72. MEOIRIAN. D. I98S). Fossil manitnals of 1980b.TheevolutionarybiologyOfkinglishers<Al- Hiversleigh. nunhweMcni Quccnslarul: prclitni- cedinidae). Living Bird !8* 1 13-160 ntiryoverview ofbiosiraiigraphy.conclalionand FRYX-H.FRY,K <fe HARRIS, A, 1992. Kingllshcrj, cnvironniemal change. Au.sLralian ZotKjgisi 25: l>cc-e3iersi.utd njlleis, tCrooin Hehn; London). 29-65. MOURER-CHALIRVIRE. M 1982 Les ojscau.N ARCHER. M., HAND, S.J & GODTHCLP, II 1991. fn^silcN des Phosphorites du Quercy (Eocene Rivcrsleigh. (Reed;Sydney). Supi^ricur a OligocL^nc Supericur): implications 1994 Riversleigh. 2ndedition.(Reed: Sydney). paleohiog<5ogrnphiques. Geobios, mem. sp6c 6; 1995. Terliarv environmental ;md hiotic change in 413-426. Australia Pp.77-90. lnVrba.E.S..Denion,G.H.. OLSON, S.L. 1985. The fossil record of birds. I»p. Panndge,T.C-&Burkle.L.H.(cds)Palcoclimaic 79-238. hi Fanicr,D.S., Kmg.J.R.ikf^arkes,AK.wC. andcvolulion, wiiti tmphnsisim humanorigins. (eds) Avian biology, vol 8. (New York: (Yale University Press: New Haven). dcniic EYcss). ARCMHR. M., TEDFORD. R H it RICH. T.H. 1987. SIBLEY, C.G & AHLQUIST.J.E. 1990. Phylogcny Tliir Pilkpildrid.ic. n new family and four new and chissirication of birds: A Study In molecular species of ?peiaurid possums (Marsupialia: evolution, (YalellniverMly Pres.';: New Haveni. Phalungotida) from ihc Au'^tralian Miocene. Pp- STEADMAN. D W. 1980- A review ot the oslcoUigy 607-627, In Aivher, M. (edt Possums and opos- and paleontology of turkeys (Aves; sums: Studies mevolution.(Surrey Bcatiy. Chip- Melcagridinae).Coniribut]onstoSciencefromthe ping Norton) Nauiral History Museum ofLxts AngelesCiiunty BAIRD, R.h*. 1991. Avian fossils from Ihc Quaicniarv 330: 131^207. ofAusu-alia. Pp. 809-870. In Vickers-Ridi. P.V'. JEDFORD. R.H. 1967. Fos.sil mammalsfromtheCarl Mtmaghaii. .I.M.. Baird. R.F. & Rich, T.H, (cds) Creek limesione, northwestern Queensland. Bul- Vertebrate palaconiologyofAustralasia.(Pioneer letinotWxBureauofMineralResourcesGeology Design Studio: Melhoume). andOcophy-sicN92: 2l7-23ft. 234 MEMOIRS OFTHEQUEENSLAND MUSEUM VICKERS-RICH, P. 1991. The Mesozoic andTertiary VICKERS-RICH, P.V., MONAGHAN,J.M., BAIRD, history ofbirds on the Australian plate. Pp. 721- R.F. & RICH, T.H. (eds) (1991). Vertebrate pal- 808. In Vickers-Rich, P.V., Monaghan, J.M., aeontology of Australasia. (Pioneer Design Stu- Baird, R.F. & Rich, T.H. (eds). Vertebrate pal- dio: Melbourne), aeontology of Australasia. (Pioneer Design Stu- dio: Melbourne).

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