Silurian and Devonian crinoids from central Victoria

114 Pages·1999·47.1 MB·English

.
Sit URIAN AND DFV0N1AN CRINOIDS PROM CENTRAL VICTORIA
PETERA-JELL
Jell.P.A. 19990630: Silurian and Devonian crinoids Fromcentral Victoria. Memoirs of'tht
Queensland Museum 431 1 1: 1-114. Brisbane. ISSN0079-8&35,
Crinoidshave longbeen known frommiddle Palaeozoicelasticsincentra] Victoriabut this\i
ihe Qi tl - nnprcb^nsivc attempt to document the fauna; 54 taxa are described; 25 are
eamcrates lea), 7 dispanels Id). 19 cladids (el) and 3 llexibles if). Nev\ taxa are the genera
Hollawayorinm cairns fca). Duncanicrinus zaJvarioht* lea), FrattkctcrtHiiS hoftnesi, F-
cnidac lea). Ptel'ihoc] ioid&e uen. run. (ca], Darraghcrinus tarn; (d), Krappacrinitt
heathcoiensis K. matkiesonensis(d), Stewbrecrinii*lerryi id), ffolmesaeritma cnidtn\H.
hlucnxis (cl) and Qimdritaxocrinus websteri (0; camerate species Ophiocrfnus miattae,
LUnwn.Kriniti's biaphhtsti. F.ttcrimis clarkae, Bmimerocrmit* ektird fill NtXocftnffl
walfUn^nsis, Hexacrfnlte& chiruM:'- ...- Qehlerti£rinu& temenni O. jeani, Atiaocrinm
Ifneatwt, Clematacrittusperforates and C ewgiItwsti disparidspecies Trichacrhna tmrleyi
;
tnid Phif/Kjrri/ins hanschi; cladid species Ciniiiicrtnus secundum, Cupiilocriilits
aUstrogr&cili&t DB}\dfacritiUi» Grfitgiutt Rtirodendrotyinm awitralis Sninfacrfnus
cometensis, S. rlchi, Ann'hoitiocrintts ehopmmri, Na&soviocrinm corcorauU On tcnocriMm
i/)//vand D. reniotusi andthe flexible MerisioirmusqwlrSmrnttx. 3 i'rwaids, Silurian,
'iiati, Victoria.
PeterAJell QueenslandMu$&um '5 - 3300.SouthBrisbane,4JO/, Australia;received
?S March 199H. t
Middle PalaeozoicSequences in the Melbourne from Heatheote (Jell, 1982)wanwd: 3. Koaptoono-
Trough of central Victoria have yielded rich crimis mitti, Codiacrittos Oendrocrintis
fossil faunas since the middle oflast century but saundvrs\ and a new genus of Dimerocnnitidae
the diverse crinoid component has been largely (-Dttnccnircnmts catvitriolits below) from near
igjiored among eehinoderm groups (Withers & [he ilunan Devonian boundary east ofMelbourne
Keble, 1934aandb;Ml, 1983}. Possibly the first (Jell m Jell & Hotloway, 1983). Numerous other
record and certainly the first illustration of records ol crinoids in these sequences (e.g.
crinoids from central Victoria was that of Williams. 1904; Gill & Caster, I960) simply
Blandowski (1855) but he figured only stem noted them among wider faunal listings without
columnalsanddidnot follow with thedescriptive naming ordescribing taxa. Other described crin-
paper promised. Bather 897) Erst described an oidsfrom VictoriaoceurinlimestonesatLilvdale
( 1
articulated cruKnd,Hapaincrinus victoria^, with (Bates. 1972), Mansfield (Jell et af. 1988) and
a linedrawing that heacknou (edgedin a footnote Toongabbie (Philip, 1961). In the rest ol
as being somewhat uncertain. Photographs of Australia, Devonian crinoids have been
Bather's (1897) specimen and its counterpart described from limestones m Queensland and
NSW
(Bather claimed it was lost) arc provided below (Jell et a!., 19KS) and Pisoertnus (referred
and showthai thecup ispoorly known. C hapman to Pantpisocrinns by Ro/hnov, 19X1) and
(Iu()3) described HelicocjrittHS piuniosus and La Liimrn,}K\ have been reported from the
flninocrjjitfs fongibraehiatus from the Silurian Silurian at Yass (Etheridge, 1904; Chapman.
ofthe Melbourne area. Both are reviewed below 1934).
with the latter re-assigned to Ntmoviotnnw
BIOSTRATIGRAPtn
i el, 191S.
More recent reports of crinoids m clastic fhe sections from which these crinoids have
sequences ofcentral Victoria are restricted to been collectedspan the Ludlow and Pridoli ofthe
: I
Talent (1965:17. pi, 4. tig- 2) noted crinoid stem Silurian and the Lochkov oi' the Devonian.
ossicles as a common element throughout the However, soil cover, urban development and
Heatheote sequence and crowns at localities 41 metHmorphism around intrusions limit available
\^Hapalocrinitcsurgylensis sp. no\.. below) and sections. Virtually all the echmodcrms known
fittum where he figured the unidentifiable have been collected from quarries, road cuttings,
external mould of a crown: 2. Crotaloerimies creek beds or other man-made exposure.
MEMOIRS OF THE QUEENSLAND MUSEUM
Wenlock Ludlow Prid- Lochkov Pragian
oli
Not
Aegiriathomasi Boucotiajaneae Boucotiaaustralis
Plentiensis
1 7 3 4 5 6 7 8 9 10 11
Dendrocrinussp. X Tnchocnnusmorlevi
Dendrocrinusarmgiits X X X X Hollowcwcrinuscalvus
Xassoviocrinuscorcorani X X Ctenocrinussignaius
Hapalocrinusvictoriae X Ctenocrinuspaucidactylus
Helicocrinusplumosus Dimerocnnitesbispinosus
Nassoxio-longibrachiatus X EucrinuscUukuu
Phimocrimsamericanus X X
Clematocrinusargylensis X Clematocrimtsperforatus
Alisocrinuslineatus X o Erankocrinusholnwsi
Phimocrinushanschi X Erankocrinusenidae
Shintocrinuscometensis X o Duncanicrinuscalvanolus
Antihomocrinuschapmani X X X X
Ouadnlaxocrinuswebsteri X Nexocrinussp.
Holmesocrinusenidae X Plwodendro-australis
NexocrinuswaUanemis X Stewbrecrinustern'i
Kroppo- heathcotemis X X Codiacrinussecundus
Cupula-austrogracilis X Holmesocrinusidaensis
Hexacriniteschimsidemis X Darraghcrinustomi
Ophiocrinusnnettae X Kroppo-mathiesonensis
Eudimerocnnusgilli X Meristocrinusquatriramus
Ctenocrinusstellifer X X X Decacrinussp.
Dictenocrinusremotus X Ancyrocrinussp.
Dictenocrinusibaeypus X o Crotalocrinitespulcher
Kooptoonocrinusmitti X o Eudimerocnnuseckardti
Gerotdicrinussp. X Oehlerticrinuslemenni
Shintocrinusrichi X Oehlerticnnusjeani
Pterinocrinidaegen.nov. X Myelodactylid indet.
Dendrocrinussaundersi X
Codiacnmtsrants X
FdIeGal1t.wDiitshtriinbutthiisonpaopfecrr.inJeolilds&inHtohlelSoiwlauyri(a1n9-8D3e)voorniJaenllc(la1s9t8i2c).seTqhueenicnetseronfatcieonntarallsVciaclteorisiac;oralrlelsapteecdietsoltihseteldoacrael
BrachiopodAssemblageZoneschemeofGarratt(1983)inthetoptworows,followingGarratt&Wright(1988).
ThenumberingsequenceofcolumnsinthethirdrowreferstoaggregatesoflocalitiesaslistedintheAppendix;
detailedrelativelevelsofwidelydistributedlocalitiesisuncertainandthesemustbeseeninmanycasesasbest
approximations.OccurrenceoftaxalistedontheleftofthefigureareindicatedwithkX'andthoseontherightby
'O1.Genericnamesendinginahyphenrequireadditionofthesuffixcrimtsandareshortenedtofitasingleline.
Structure within the Melbourne Trough consists the shelly faunas have been placed in sequence
of tight folding with long approximately N-S (Garratt, 1983). The crinoids described herein
axes, also limiting measurable sections. Fossils comemainlyfromthefewhorizonswherefossils
occur in profusion in a few horizons; they are areprolificandpreservationissuperb;verylarge
relatively rare in many other horizons; but most numbers ofarticulated echinoderms occurthere.
of the sedimentary sequence is unfossiliferous. For example, NMVPL252 provides the greatest
Brachiopods arethemostcommon group among diversity ofcrinoids and of echinoderms in the
theshellyfaunaandhavebeenusedtoestablisha MelbourneTrough.Thislocalitywasoriginallya
preliminary biozonation (Fig. 1) against which small road metal quarry but most available
CRINOIDS FROM CENTRAL VICTORIA
material came from a new excavation of the SYSTEMATIC PALAEONTOLOGY
qbEuynaitrdhreyHaoiulntmhteohreswsaiuntmdhmteShtreessvueosftEac1ik9na8er0dd-ht1e9la8pn3dofuonFcdrceaarsntikaoaknneadln VicMtaotreiraia(lNdMeVscPr)ibaendditshheoluoscealditiinetshaereMuresgeisutmereoid'
in the same Museum (NMVPL). Specimens are
help ofseveral others. Those localities that yield
preserved in decalcified elastics so are found in
only one or 2 species mostly represent localities
the form of internal and external moulds often
where fossilsarerareandtheiragedeterminedby
stratigraphic position. The crinoid localities partially infilled with iron oxides making the
mould lessthan faithfull.Morphological detail of
(Appendix) are placed in approximate relative
the moulds improves withdiminishinggrain size
stratigraphic sequence (Fig. using these best
available data. The brachi1o)pod zonation of ofthe matrix. All illustrations are of latex casts
taken from these moulds and whitened with a
Garratt is related to the international stage scale sublimate ofammonium chloride. Terminology
twhirthougghratphteolsiteeqsuetnhcaet (oGcacrurarttat&aWrfiegwht,ho1r9i8z8o)n.s follows Moore&Teichert(1978). Measurements
are given as: length, parallel to the central axis;
The crinoids described herein are not signif- width, transverse to.butnevercuttingormeeting
icantly useful biostratigraphically as they occur the central axis; and depth, normal to, and may
relatively rarely in terms of both localities and join the central axis.
horizons.Neverthelesstherearecleardifferences
between the Ludlow and Lochkov faunas for Class CRJNOIDEA Miller, 1821
CAMERATA
example and the continued study of this fossil Subclass
groupwithotherelementsoftheshellyfaunawill Wachsmuth & Springer, 1885
continue to refine what is at present a very Order DIPLOBATHRIDA
preliminary brachiopod zonation. Moore & Laudon, 1943
Suborder EUDIPLOBATHRINA Ubaghs, 1953
PALAEOECOLOGY
Superfamily RHODOCRINITOIDEA
Palaeoecological settings in the Melbourne Roemer, 1855
Trough were discussed in a number of papers Family OPSIOCRINIDAE Kier, 1952
(Garratt, 1983; Cas, 1983; Vandenberg & Frest & Strimple (1981) and Ausich (1986a)
Wilkinson, 1982; Vandenberg, 1988, 1992). Cas recognised the close similarity of Opsiocrintts
(1983, fig. 17) provided a schematic diagram of Kier. 1952 and Opkiocrinus Salter, 1856 and
regional palaeogeography for SE Australia assigned them both to this family; they are
showingthe MelbourneTrough as a longnarrow consideredsynonymous by Jell & Theron, 1999.
embayment ofthe ocean with ample neighbour- Ausich (1986a) also erected 2 new genera from
ing relief and continued subsidence to provide the Llandovery of Ohio in this family and
thethick pileofclastic sediments. Localities rich discussed the group in detail. Inclusion of the
in articulated echinoderms occur in distal Australian Holtowaycrinus nov. is discussed
turbidites (PL252) or extensive thin sand units under that heading below. This addition
(within a finer grained sequence) (PL300, 229, necessitates widening the family concept to
1924). Both these settings indicate catastrophic includea form with radials in lateralcontact; this
events causing the death and fast burial ofthese difference has been sufficient to separate taxa at
animals (Jell, 1983) but the cause or causes of superfamily level mthepast(Ubaghs, 1978b).To
such events remain unclear. take an even more heretical view 1 question
PALAEOBIOGEOGRAPHY whether Stelidiocrinus Angelin, 1878 from the
UpperSilurianofGotland shouldnotbe included
These crinoid faunas provide few useful data in this family. It has been classified in the
for palaeobiogeographic reconstruction as they Monobalhrida because it lacks the second circlet
contain taxa otherwise restricted to North ofplates below the radials but it (particularly S.
America orto Europe and othertaxa common to laevis Angelin, 1878 as figured by Ubaghs
both those areas. Few other parts of the world (1978b. fig. 307, le)) resembles the
havewellenoughdescribedcrinoidfaunasofthis Opsiocrinidae very much in cup shape, median
age for meaningful comparison. At the generic column ofanal plates in CD interray, 10 biserial
level there are slightly more representatives of arms, few slightly depressed interradial plates
the European fauna as is the case with brachio- and presence of an intersecundibrach. Its first
pods (Boucot et al., 1969) but among camerates primibrach is shorter and wider than in most
there are more American affinities. Opsiocrinidae but is comparable with that ofO.
—
MEMOIRS OF THE QUEENSLAND MUSEUM
Ophiocrinus nnettae sp. nov.
Eifelian Ophiocrinusmariae
(Fig- 3)
Ophiocrinusslangen - 'Ophiocrinus-*srp.-—cf,O.mariae ETYMOLOGY. Ananagramfrom Annette.
Pragian MATERIAL. NMVP149344 fromNMVPLI990.
\
Lochkov Holkrw/aycrinw DIAGNOSIS Intcrrays narrow; primanal in
Ophiocrinusnnettae posterior interray contacting C and D radials and
V-
1-1iki*-I: 1stprimibrachs,supporting3 analplatesdistally;
t* central columnoflargerhexagonal plates in anal
SteHdiacrirms sac. Arms 10, with triangular axillary 2nd
Wenlock h primibrach, distinctly biserial above distal to
Llandovery Rliachicrinus SUfanocrbaa secundibrach. Stem circular, with small but
distinct marginal projections on nodals and
Ordovician Gaurocrinus-"TUceancestor
internodals.
FIG. 2. Sketch of phylogeny of the Opsioerinidae DESCRIPTION. Crown >40mm long, >4 times
followingAusicb (1986a) and discussion herein. as long as wide at cup, with arms spreading
gently (tips not preserved). Cup low conical,
nnettaedescribedbelow. Throughout the Opsio- approximately5mmlong,widerthanlong;plates
erinidaetheinfrabasalsareconcealedbythestem smooth, convex so sutural margins depressed.
attachment and Opsiocrinus itselfwas originally Infrabasals presumably concealed by stem.
describedasmonocyclic; I suggesttherefore,that Basals longer than radials, hexagonal, longer
a careful examination ofthe Swedish material of than wide. Radials pentagonal, wider than long,
Stelidiocrinus should be made to determine this separated from each adjoining radial by basals
feature. Stelidiocrinus would join Holloway- and 1stinterprimibrach,withdistinctlymargined
crinusasthesecondOpsioerinidaewithradialsin outer ledge across distal margin matched by
&
lateral contact. Witzke Strimple (1981) sug- similarledgeon 1stprimibrach,similarmatching
gested that evolution from rhodocrinitid to ledgesonsucceedinginterplatesuturesalong the
dimerocrinitid stage (i.e. withdrawl of the first arm gradually diminishing in distinctness and
interbrachial to allow lateral contact of radials) size; 1st primibrach rectangular, as wide as
occurred several times. They postulated 3 radial, short; 2nd primibrach axillary, triangular,
separate occasions from different Ptychocrinus as wideas 1st primibrach. Arms 10, subquadrate
stocks so my suggestion of2 further occasions insection,withflattenedouterface;eachraywith
above is in line with their thinking. 2 main rami, uniserial as faras 6th secundibrach,
becoming biserial abuptly (3rd in one arm
Currentknowledgeofthis family suggeststhat
observed) distally, with long pinnule of 5 or 6
only small sections ofthe lineages involved are
kStnroiwmpnle((Fi1g9.81)2)thaantdagIreaagtrdeeealwmiotrheWbiatszickedat&a pfiinxendulianrscuopn;eianctherfprreiembirbarcahcihasl;smoanlllyaprnidmfiebwr;ac1hsst
largest, contacting basal, 2 radials and 1st
about Silurian forms in particular will be
primibrachs, supporting 2 long narrow plates
necessary to refine our understanding.
distally. Primanal hexagonalthough contacting8
plates, almost as large as basals but not quite as
Ophiocrinus Salter. 1856 long, supporting 3 anals distally; anal plates
numerous, decreasing in size distally, with
average size larger than that of interbrachials,
TYPESPECIES. OphiocrinusstangeriSalter, 1856 from distinct central column ofdecreasing hexagonal
theLowerDevonian Bokkcveld Series, SouthAfrica. plates. Stem circular in section, heteromorphic
with slightly different sized (length and
REMARKS. Th&is genus is discussed in detail diameter) columnals alternating, noditaxis Nl,
elsewhere (Jell Theron, 1999). Its occurrence with epifacet on each columnal having an outer
in the Lower Devonian of Victoria in close circlet of small but distinct tubercles or
proximity geographically and temporally with
pseudocirri.
Hollowaycrinus gen. nov. makes itquite feasible
thatone gave rise to the otheras remarked under REMARKS. This Victorian species differs from
that genus below. the South African genotype (Jell & Theron,
CRINOIDS FROM CENTRAL VICTORIA
ETYMOLOGYForDavid1lollowaywhogreatlyassisted
Willi fieldwork,curationmid scientific advice.
DIAGNOSIS. Infrabasals 5, concealed by stem
attachment. Radials large, heptagonal. incontact
witheachotherlaterally except across CD inter-
ray; 1st primibrach hexagonal; 2nd primibrach
axillary. Primanal in radial circlet, supporting 3
platesdistally.with mediancolumnoflargeranal
plates involving 4 and probably more plates.
Anns 10-20, with each ray having 2 main amis
becoming biserial distal to the 6th or 8th
secundibrach. with some arms branching irreg-
ularly once or twice distal to primaxil even
though other rami in the same individual are
unbranched.
REMARKS. This genus is placed in the Opsio-
crinidae basedonthe similaritiesincup shape. 5
concealed infrabasals. median column of anal
CD
plates in interray. primanal supporting 3
plates, large 1st intciprimibrach supporting 2
plates followed by large number of very small
irregularplates. 10-20 arms, uniserial tobiserial.
Conventional classification following Ubaghs
(1978) would place this species in Dimero-
crinites because of the radials being in lateral
contact. However, the arms being uniserial
proximally then becoming cuneate and finally
biserial. the intcrradial plates being mam small
onesaboveafew largeonesandthe radialsbeing
barely inlateralcontactallpointtoanoriginfrom
theOpsiocrinidacanditisclassifiedaccordingly.
Hollowavcrinuscalvus sp. nov.
FIG. 3. Opkiocrinus nnettae sp. nov., incomplete (Fig. 4)
holotvpe crown in C ray view, NMVP149344 from
NMVPL1990,x3. ETYMOLOGYLatincalvus,bald;referringtothelackof
median ray ridgesandornament
1999) in having fewer interprimibrachs. no
intersecundibrachs.distinct ledges oneach plate MAI'LRIAL.HOLOTYPL: NMVP100158. PARATYPES:
oneachinterplatesuturefromradialsuptoabout NMVP100150, 100153, 107099-107104, all from
NMVPL229.
3rd secundibrach. its arms becoming biserial
muchearlier(secundibrach 3 in some anus) and DIAGNOSIS. As forgenus.
in the stem with tubercles on internodals. From
theNorthAmericanO. mariaeKier, 1952andO. DESCRIPTION. Crown subcylindricaf averag-
henderi (Kesling. 1968). if those are separate ing 30mm long, at least 4 times as long as wide.
speciesas suggestedby Frcst & Strimple (1981) Cup low to medium conical. 2-6mm long, just
contrary to Kesling & Chilman (1975). the wider than long, of unornamented plates.
Victorian species is distinguished by the same Infrabasals5.concealedby stem.Basals5.longer
featuresasseparatethegenotypeaswellasa lack than radials. hexagonal except for heptagonal
of ornament on the interprimibrachs and very posterior one. longer than wide. Radials hept-
short 1st primibrach. agonal. wider than long, with longer sides
contacting basals and 1st primibrach. with short
Hollowaycrinusgen. now sides contactingadjoining radials orprimanal in
CD interray: 1st primibrach hexagonal, not as
TYPF SPECIES. Hollowaytvmuscalvussp. nov. wide as radial and little more than halfas long;
MEMOIRS OF THE QUEENSLAND MUSEUM
D
m
ZM-k
1
1 E
H
mM
CRINOIDS FROM CENTRAL VICTORIA
2nd primibrach axillary, pentagonal, as wide as Strimple, 1981 amongothers). Since most that is
1stprimibrach; each raywith 2 main arms, some known about this family comes from the
without further branching, others with 1 or 2 Northern Hemisphere it is outside my aims to
furtherbranchings irregularly, uniserial as faras review its phylogeny. However, in adding the
6th or 8th secundibrach, becoming biseriai several Australianformsthatfollow havehadto
1
distally, with long pinnule of 5-6 pinnulars on make some inferences about Northern Hemi-
each free brachial; arms free distal to 2nd sphere taxa. The simplest way to recognise
secundibrach; interprimibrachs numerous, with members ofthe family until recently hadbeen to
large heptagonal proximal one supporting 3 identify that a crinoid was dieyclic and that its
plates in a row distally, with size decreasing radials were all in contact except for interrupion
rapidly distally, with distal medial plates by the primanal in the CD interray. Witzke &
depressed. Primanal hexagonal, almost as large Strimple (1981) questioned the universality of
as basals, supporting 3 anals; anal plates num- this single feature and 1 agree with their
erous, decreasing distally, with average size discussion ofevolution between the Rhodocrini-
larger than interprimibrachs. Stem circular in loidea and Dimerocrinitoidea. In assigning
section, with slightly different sized (length and Hollowaycrinus to the Opsiocrinidae and
diameter) nodals and internodals alternating suggestingthatSielicliocrinusmayalsobelongto
proximallybutdistallywithnoditaxisN212,may that family I follow their line ofthought. 1 also
be long (no complete stem known but an follow Witzke & Strimple (1981) in the generic
incomplete one in excess of9cm known), distal divisionsofthe familybasedonnumbersofarms
terminationunknown. Allplatessmooth,without (Fig. 5) although there is good reason to believe
ornament. that numbers of arms may have changed on
several parallel lineages within the family.
REMARKS. This species is distinguished from
cOopnhtiaocctrianunsd bbyymovsatriraabdliealasrjumstbmraaknichnigngla.terIatls weMonvoiwnghaalovnegWaivtaizlkaeb&leSagmepnleer'isc(n19a8m1e)sidefaosr
separation from dimerocrinitids is discussed dimerocrinitoids with uniserial armsthatdivided
above. The amis are highly variable in their once {Duncanicrinus gen. nov.)„ twice (Nexa-
branching pattern and also m the arrangement of crinus Eckcrt, 1984) or more than twice in each
brachials. Most observable rays divide once and ray (Ptychocrinus) and for biseriai represent-
have only 2 arms whereas others may have one ativeswith 10(Dimerocrinites\ 20 (Eucrinus) or
undivided arm from the first branchingwhilethe more than 20 arms (Eudimerocrinites
second may divide twicemore. Most subsequent (—Ambicocrinus), Griphocrinus). Whether latest
divisions appearto be at fairly high and variable OrdovicianPtychocrinusfunbrialus, Llandovery
angles to growth direction; there is a question of P. longibrachialis and Early Devonian
whether these could represent a regrowth Duncanicrinus constitute a lineage or not will
response to breakage or predation to which no only be answered by gaining further knowledge
answer is readily available. Brachials begin to ofSiluriantaxaandthusdetailed phytogeniesbut
becomecuneatejustdistal to thecup and in most at present this is a reasonable hypothesis worth
cases become biseriai cuneate by about the 8th further testing (Fig. 5). These generic groupings
secundibrach; a few remain cuneate farther accommodate Brower's (1973) idea that the 3
distally and none become rectilinear biseriai. latest Ordovician species of Ptychocrinus
represented 3 separate lineages and should be
Superfamily DIMEROCRINITOIDEA separated generically and Witzke & Strimple's
ZitteL 1879 (1981) ideas on the existence of3 lineages in the
Family DIMEROCRINIT1DAE Zittel, 1879 Dimerocrinitidaebased primarily on numbers of
amis. Therearenodoubtnumerousexceptionsto
This family is a regular component ofcrinoid these general groupings and the most obvious is
faunas from Ordovician to Devonian and has the high probability that the transition from
been discussedby manyauthors (Ausich, 1986a; uniserial to biseriai arms occurred more than
Brower, 1973; Breimer, 1962; Frest & Strimple, once in each lineage. However, it is necessary to
1981; Macintosh, 1981, 1987; Witzke & continueassimilatingnewdataintoan hypothesis
FTC. 4. HoUowarcrinuscalvus gen. et sp. nov., all crowns with orwithout stem attached, fromNMVPL229. A,
NMVP10710U *2.5. B, holotypc. NMVP1O0158, x4. C. NMVPI00T50, *5. D, NMVP100153, *4 E,
NMVP107I02, x3.5.F,NMYP107l()4, *3.5. G, NMVP107I03, *3. H,NMVPI07099, x5.
.
MEMOIRS OF THE QUEENSLAND MUSEUM
A B
Biserial;>20Arms Uniserial;>20Anns Biserial;20Arms Uniserial;20Arms
Eifelian E. whitfieldi
A. arborescens 1
r
Emsian 11
T
1
i
i
Pragian
I
1
Eu.eckardti E. clarkae Nexocrinussp.
V'
Lochkov
V Eu.gilli
~1
ii
Pridoli
1 i
1
N. wallanensis
Ludlow i 1 i
1 EI UCRINUS —N1EXOCRINUS
I 1 _|
1
l E.laevis, nodibasis,
Eu. multibrachiatus N. occidentalis
\ lockportensis
Wenlock
1
\ 1
E. icosidactylus
v\ V 1
\
E.pentlandicus 1
\
~^"^^
Llandovery \ . N.A1damsensis
P. medinensis
\ ' A^. delicatulus
\
Ordovician *P. splendens N.parvus
c
Biserial; 10Arms Uniserial; 10Arms
Eifelian
Emsian
Pragian
Lochkov D. bispinosus D. calvariolus
1
Pridoli DIMEROCRINJTES DU1NCANICRINUS
Ludlow D. longimanus
l
i
D.planus,milligani,roemeri,
Wenlock Hlliformis,brachialus
FIG. 5. A, sketch of phylogeny of the
D. decadactylus, inornatus Dimerocrinitidae with more than 20 arms
includingPtychocrinus,Eudimerocrinus and
D. sculptus, Ambicocrinus. B, phylogeny of the
Llandovery hopkintonensis, elegans I D. longibrachialis Dimerocrinitidae with 20 arms including
Nexocrinus and Eucrinus. C, phylogeny of
the Dimerocrinitidae with 10 arms including
Ordovician nD.fimbriatus
Duncanicrinus andDimerocrinites
CRINOIDS FROM CENTRAL VICTORIA
FIG 6, Dimemcrinitesbispinosussp. now from
NMVPL252. A,B, D ray view and
enlargement ofposteriorinterray ofholotype
NMVP108581a,x2.5 and *5,respectively. C.
Aray viewofcounterpart(B rav onleft. E on
right). NMVP1()S5SU\ -2.5' D splayed
NMVP s
crownin basal view. 10S615. <2.5.
MEMOIRS OF THE QUEENSLAND MUSEUM
10
until all the details are worked out. To that end 1 primibrach axillary, heptagonal; each arm
offer a possible phylogeny (Fig. 5) for known uniserial with cuneate brachials as far as 6th or
speciesofDimerocrinitidaeforfuturetestingand 8th secundibrach, biserial distally, with pair of
modification where necessary. Just how many strong solid spines on outer surface of rami
lineagesand which features remainedconstantto placed about every 5 brachials; pinnules on each
be able to recognise separate lineages remain free brachial (i.e. distal to 2nd secundibrach) on
largelyunanswered forthe dimerocrinitids today each side of ramus, long, of 8-12 pinnulars.
as when Witzke & Strimple (1981) raised many Interprimibrachs numerous, with heptagonal
questions on the origins oftheir groupings. proximal one supporting 2 plates of about the
same size, with size decreasing rapidly distally.
Dimerocrinites Phillips in Murchison, 1839 First intersecundibrach hexagonal, subsequent
TYPE SPECIES. Dimerocrinitesdecadaciylus Phillips in plates not clear but presumably part oftegmen.
Murchison, 1839 from the Wcnlock of England; by Posterior interray with central anitaxis of hex-
subsequentdesignation ofRoomer, 1855. agonal plates; hexagonal primanal supporting 3
REMARKS. 1 apply the concept of Witzke & daencarlesasiinngnedxisttalrloy,w;wiatnhalavpelraatgeessnizuemesrmaolulse,r
Strimple (1981) to Dimerocrinites (Dimero- than that ofinterbrachials. Stem unknown.
crinites) which is to restrict the genus to those
formsofthebroadergenericgroupwith 1 arms. pRlEacMeAdRiKnSD.imWeriotchri1nibtiesseroiaflWaintuzskethi&ssSpterciimesplies
Dimerocrinites bispinosus sp. nov. (1981) and is distinguished within that taxon by
(Fig. 6) thespinesontheouterarmfaces,long hexagonal
ETYMOLOGY. Latin/»/-,twoandspinosus,spine, fordie intersecundibrachs and the heptagonal axillary
pairsofspineson theoutersidesofarms. 2nd primibrach.
MATERIAL. HOLOTYPE: NMVP108581. PARATYPE: Eucrinus Angelin, 1878
NMVP108615 from NMVPL252.
TYPESPECIES. EucrinuslaevisAngelin, 1878 fromthe
DIAGNOSIS. Ten arms, with prominent reg- Wenlock of Gotland; by subsequent designation of
ularly spaced pairs ofspines every fifth brachial Wachsmuth & Springer, 1881,
on outer face, with hexagonal intersecundibraeh, REMARKS. This taxon is used in the sense of
withlowbutdistinctanitaxisofhexagonalplates. Witzke & Strimple (1981) to include species of
DESCRIPTION. Crown subcylindrical, up to Dimerocrinites with 20 arms; there seems little
45mm long, up to 4 times as long as wide. Cup point in maintaining subgeneric distinction since
subcorneal, about cm long, longer than wide, we have no clear ideaofphylogeny amongthese
with convex tegmeIn; cup plates thin, smooth. many species and when lineages are established
Infrabasals concealed by stem. Basals pent- it seems likely that both Dimerocrinites and
agonal, except for hexagonal posteriorone, with Eucrinus will prove to be polyphyletic. The
narrowest proximal margin forming part of Victorian species is assigned on the basis that all
circular rim to basal concavity, about as long as rays countablehave4 anus except foronewhich
wide, with marked proximal medial projection has5andisconsideredaberrant;thusatotalof20
(suggesting apentagonal stem withthe angles of armswouldbeexpectedifbranchingisuniform.
the stem in the centres of basals beneath this Eucrinus clarkae sp. nov.
projection), with Y-shaped median ray ridge
(Fig. 7)
dividing about midlength and extending onto
adjacentradials. Radialsheptagonal, muchwider ETYMOLOGYForPennyClarkwhohelpedgreatlywith
than long, with inverted Y-shapedray ridge, with photography,curationand fieldwork.
2aumniisfseocorufmntdhicibrsorYsascrhussnenciftniigxoenodnttihonraodcujugaphc.enaAtvarbimalssaabllse1.0,Fairrosmft MTYAPTEESR:INAMLV.P1HO08L6O43T,YP10E8:645NaMilVfPro1m0N8M64V2P.L2P5A2R.A-
length; 1st primibrach hexagonal, not as wide as DIAGNOSIS. Basals, radials and radianal with
radial and little more than half as long; 2nd distinctive coarse ornament of short fat radial
FIG.7.Eucrinusclarkaesp. nov.allfromNMVPL252.A,C,lateralviewofcrownwithstemandenlargementof
distal cupandproximal arms ofNMVP108645a, x2.5 and *4, respectively. B, lateral view ofNMVP108643,
x2. D, lateral view ofNMVP108645K *2.5. E, CD interrayviewofholotype NMVP108642, x4.