THE ASTEROSTOMATID ECHINOID ANTILLASTER FROM THE PARADASH GROUP (MIDDLE EOCENE) OF THE NAKHICHEVAN REGION OF AZERBAIJAN

9 Pages·2003·2.7 MB·

by  Kenneth J Mcnamara

Recordsofthe WesternAustralian Museum21: 157-165 (2002).
The asterostomatid echinoid Antillaster from the Paradash Group
(Middle Eocene) of the Nakhichevan Region ofAzerbaijan
Kenneth McNamara'andOktayH.Melikov^
J.
’DepartmentofEarthandPlanetarySciences,WesternAustralianMuseum,FrancisStreet,
Perth,WesternAustralia6000,Australia
'AzerbaijanStateOilAcademy,370102Azadlikpr.20,Baku,Azerbaijan
Abstract - The asterostomatid echinoid Antillaster has previously been
recorded only from the Caribbean region and northeastern South America.
Herewereportthe firstoccurrenceofthisgenus fromoutsidetheAmericas,
intheLesserCaucasus,whereithasbeenfoundinMiddleEocenesediments
oftheParadashGroupintheNakhichevanregionofAzerbaijan.Thematerial
is described as Antillaster bagmanovi sp. nov. It is the earliest known
occurrence of the genus. Its presence in the Paratethys, in a region far
removed from previous known occurrences, has significant
palaeobiogeographic implications. Its disappearance from the Lesser
Caucasus,butpersistence intheCaribbean, maybe linked tothecontraction
oftheTethysduringtheCenozoic.
INTRODUCTION from Eocene to Miocene sediments in Venezuela,
Eocene sediments in the Nakhichevan region of Cuba, Jamaica, Bonaire, Antigua, Puerto Rico and
Azerbaijan have long attracted the attention of Mexico (Kier 1984). Here we report the firstrecord
manyinvestigators,duetotheirgoodexposureand ofthisgenusfromoutsidetheCaribbeanregion,in
extensive foraminiferal and molluscan fauna. The Middle Eocene sediments of the Lesser Caucasus
dating of the predominantly clay-sand facies as region in Azerbaijan. This provides the earliest
beingEoceneisbasedonthisrichforaminiferaland recordofasterostomatidechinoidsintheParatethys
moUuscanfauna (Azizbekov 1961;Bagmanov 1966, region.
1980; Paffengoltc 1979). The presence ofan Eocene
uecnhkinnooiwdnfauunntailin1th9e58Nawkhheicnh,eviannardedgiitoinornemtaointehde GEOLOGICALSETTINGAND
BIOSTRATIGRAPHY
foraminifers and molluscs from sections in Juga,
Daralik, Ilandag and Bilav, M.A. Bagmanov Middle Eocene sediments are are widespread in
collected a rich fauna of echinoids from the thesouthwesternpartoftheNakhichevanregionof
Paradash Group. Unfortunately, this fauna Azerbaijan, extending to the Arax River, which
remained unstudied for many years. However, in marks the border with Iran. These sediments are
1984 Bagmanov kindly passed his echinoid separated by a pronounced unconformity from
collection over to one ofus (O.H.M). During 1985- imderlying Triassic sediments. This boundary is
87Melikov, tosupplement thiscollection,collected markedby athickbasal conglomerate. The Middle
additional material from Bagmanov's sections. Eocene sediments are divided into three groups,
These collections from the Paradash Group have which extend through six foraminiferal zones
yieldedarichechinoidfauna,comprising25species (Figure1).
(Melikov in prep.). Current work by Melikov
suggests that these species can be assigned to 14 DaralikGroup
genera in 11 families, and 5 orders. Itis interesting
that this species richness is in strong contrast to a Lowersubgroup
fauna of equivalent age occurring in sediments in Along the south-west limb of the Nakhichevan
the neighbouring territory of Armenia, a mere 40- synclinalbasin, particularlyatDaralik(ontheright
50kmaway.Hereonlysixspeciesareknown. bank ofthe Alinjachai River), thisbasal unit ofthe
One of the more surprising echinoid finds from Middle Eocene sequence encompasses the
thesesedimentswerelargespecimensofAntillaster. Nummulites laevigatus zone and is represented
The asterostomatid echinoid Antillaster has mainly by conglomerates containing a series of
previouslybeen describedonlyfromtheCaribbean isolatedpacketsandlensesofLimestones,clay-shale
region and northern South America, specifically and sandstones. It is characterised by its reddish-
158 K.J.McNamara,O.H.Melikov
Upper
Globerigina corpulenta
Eocene zone
>
n
c c
0 CO Truncorotaloides rohri d.
OT
'c zone S
r“ o Paradash o
0 c. Group c:
CO Acarinina rotundomarginata e
o CD zone CC3O)
•Q
o
Globeriginatheka subconglobata CO
-S
LJJ c zone
C=
0 CO
Nummulites atacicus Bilav
-1—> zone Group
0)
ID Morozovella aragonensis
zone
Daralik
2
Group
Nummulites laevigatus
zone
Figure1 Biostratigraphic chart showing the foraminiferal zonation, stratigraphic groups and range of Antillaster
bagmanoviintheNakhichevanregionofAzerbaijanRepublic.
brown colouring. The thickness of the subgroup postiloculinoides Chalilov. Acarinina bulbrooki occurs
m
variesbetween5-10 to120-150m.Thissubgroup in the Lower Lutetian, P9 and PIO. N. uranensis
contains abundant Nximmulites of Middle Eocene occurs in PIO. Echinoids occurring in this upper
age, notably N. laevigatus (Brug.), N. laevigatas var. subgroup are Schizaster (Schizaster) rindensis
lantaensis Lerisch (B) and the echinoids Porosoma Poretzkaia,S.(S.)sp.nov.,MaretiahojfmaniGoldfuss
lamberti Checcia-Rispoli, Triplacidia veronensis andM. sp.nov.
Bittner, Echinolampas daralagesensis Poretzkaja, £.
blaviensisCotteau, Echinanthuscf.issyaviensis(Klein) BilavGroup
and Cyclastersubquadratus (Desor). The presence of This is represented by tuffaceous conglomerate,
N. laevigatus indicates a Lower Lutetian age, tuff, tuffite of andesite to andesite-dacite
foraminiferalzonePIO. composition and lenses of sandy shale. The total
thickness of this group is 875 m. Bagmanov (1966)
Uppersubgroup identified Nummulites laevigatus (Brug) within this
This encompasses the Morozovella aragonensis group. Highly developed olistolites of very
zone. Up the section this predominantly different-sizes is a characteristic feature of this
conglomeratesubgroupisintermixedwithlayersof group.
shelly-sandstone, argillites and rarely limestones,
containing abundant Nummulites. Within this ParadashGroup
sguypbsgurmoiuspedinshtahleeJaulnfda arleeguiroolni,teos,cc1u5r0lmenstehsicok.f biPoasltareaotnitgorloagpihciaclally thziosngersoupfirsodmividtehde inltoowfeosutr
These sharply decrease in thickness along both Globigerinatheka subconglobata zone up through the
limbs of the Bilav fold. The most common Acarininarotundomarginatazone,theTruncorotaloides
foraminifers in the zone are Nummulites laevigatus rohri zone to the Globigerina corpulenta zone. The
(Brug.) and N. uranensis Heim. Other foraminifers lowerthreeareincludedwithintheMiddleEocene,
include Acarinina bulbrooki Bolli and Globigerina theGlobigerinacorpulentazonebeingUpperEocene.
AsterostomatidechinoidAntillasterfromAzerbaijan 159
Globigerinathekasubconglobatazone Globigerinacorpulentazone
This overlies the tuffaceous conglomerate of the Sediments within this zone consist of repeated
BilavaGroupandisrepresentedinitslowerpartby beds of nummulitic sandstones, clays and
repeated beds of argillites, sandstones and clay/ conglomerates, 50 m thick. Foraminifers present
shales, having a total thickness of 126 m. These include Nummulites striatus (Brug.), N. brongniarti
rocksareoverlainby70mofrepeatedbedsofdark D'Archiac and Haime, N. paradashensis Mamedov,
red clay/shale, argillites and subordinate N.fabiani Prever, N. rectusCurrey. N. brongniarti is
interbedded sandstones. Over these lie a 100 m Bartonian, and N. fabiani uppermost Bartonian to
thick sequence of greyish clay. This lowest LowerPriabonian.
subgroup is capped by a sequence of repeated The Paradash Group would therefore seem to
m m
sandstone and clay/shale 55 thick, with a 4 range from near the Lower-Middle Lutian
thick conglomerate at its base. The beds of this boundary to the Upper Bartonian or earliest
subgroupextendalongthesouthernlimbofIlandag Pribonian. The sequence at Daralik probably
Mountain. occupies the entire Middle Eocene, possibly
Clay/shale of all packets contain a foraminiferal extendingintothelowestpartoftheUpperEocene.
assemblage containing Acarinina bulbrooki (Bolli),
whichrangesfromP9-P10,A. punctocarinataFleish,
Morozovella spinulosa Cushman, which ranges from SYSTEMATICPALAEONTOLOGY
P10-P14, Globigerina bozueri Bolli and Nummulites
laevigatus Brug., which occurs in PIO, and an OrderSpatangoidaClaus,1876
echinoid fauna of Schizaster (Schizaster) rindensis
Poretzkaia, Linthia (Linthia) soudanensis Bather, FamilyAsterostomatidaePictet,1857
Guallieria (Gualtieria) damesi Koch, Eupatagus GenusAntillasterLambert,1909
(Eupatagus) formosus Loriol and Maretia hoffmani
Goldfuss. G. subconglobata occurs in Pll. Thus this Typespecies
sequence is probably near the Lower-Middle Asterostoma cubensis Cotteau, 1871; by original
Lutetianboundary. designationofLambert,1909,p. 103.
Acarininarotundomarginatazone Diagnosis
This is represented by repeated beds of clay, Test large, low to high; apical system ethmolytic
sandstone and argillites having a total thickness of with 3 or 4 genital pores; petals long, wide, open,
391 m. Foraminifers present include Subbotina flush with test, or very slightly depressed; anterior
frontosa Subbotina (P11-P14), Acarinina vievensis ambulacrum with small pores, in slight groove or
Morozova, Morozox^ella lehneri Cushman (P12-P13), flush with test; some plates occluded at ends of
Hantkeninaalabamensis Cushman (P12), Nummulites petals; peristome large; periproct large,
acutus(Sowerby)andN.striatus(Brug.).Thespecies inframarginal;tuberclessmall,nofascioles;plastron
of echinoids recovered from this zone are: mesamphisternous with large labrum, narrow
Antillaster sp. nov., Gualtieria (Gualtieria) damesi sternal plates, large episternal plates; in some
Koch, Triplacidia veronensis Bittner, Schizaster speciesfirstplateininterambulacrum1 followedby
(Schizaster) sp. nov., Schizaster (Paraster) rimosus L. singleplate(afterKier1984:131).
Agassiz,ConoclypusconoideusLeske,andC. leymeriei
Cotteau. The planktic forams indicate a Remarks
foraminiferalzone ofP12, whichisUpper Lutetian SpeciesofAntillasterareparticularlycharacterised
toearlyLowerBartonian. by their very large test size, typically reaching in
mm
excessof100 testlength,butsometimestoover
Tnmcorotaloidesrohrizone 150 mm. Antillasterlacks any evidence of fascioles,
Thisconsists ofred-brownish (43m) and greyish hence its assignment to the Asterostomatidae.
(25m)claywithinterbedsofsandstonescontaining Antillaster bears a superficial resemblance to the
nummulites. This zone is characterised by the brissid Pharaonaster, which is known from the
presence of numerous Truncorotaloides rohri Eocene ofNorthAfrica (RomanandStrougo 1994).
Bronnimann and Bermudez (P11-P14), Globigerina However, Antillaster can be distinguished by its
incretacea Chalilov, G. praebulloides Blay., much larger test size and absence of fascioles;
Nummulites brongniartiD'Archiac and Haime (P12- peripetalous and subanal fascioles are present in
P14), N. perforatus (Mont.)(P12-P14), N. striatus Pharaonaster. Another superficially similar form is
(Breys.). Echinoids occurring in this zone are Hypsopatagus, which is known from the Eocene to
Conoclypus conoideus Leske and Antillaster sp. nov. Oligocene of parts of Europe, Asia and North
The foraminifers indicate a possible Upper America. However, unlike Antillaster it has a
Bartonianage. peripetalous fasciole; is smaller; and has closed,
ratherthanopen,petals.Onaccountofitslargesize.
160 K.J.McNamara,O.H.Melikov
absence of fascicles, distally open petals that are Paradash Group (Middle Eocene - Upper Lutetian
almost flush with the surface of the test, the form toearlyLowerBartonian).
described herein from Azerbaijan is considered to
beaspeciesoiAntillaster. Etymology
Named in honour of palaeontologist M. A.
Bagmanov.
Antillasterbagmanovisp.nov.
Figures2-7
Description
mm
Diagnosis Test very large, reaching up to 154 TL;
Species of Antillaster with very slightly sunken cupola-shaped, withapex anteriorofcentre;height
petals;anteriorlydivergentpetalswithverynarrow about 50%TL; test longer than wide, ranging
interporiferous zone; peristome sunken; between 90-947oTL. Aboral surface declines more
ambulacrumIIIdeeplysunkenorally. steeply anteriorly; ambitus broadly rounded.
Shallow anterior notch present. Apical system
anteriorly eccentric, 41-46%TL from anterior
Material
WAM ambitus; ethmolytic, with four genital pores,
Holotype 99.428, from Daralik,
madreporite extending posteriorly the same
Nakhichevan region of Azerbaijan Republic;
paratypes WAM 99.429 and 99.432 from the same distance again as the distancebetween the anterior
horizonandlocality;otherspecimensWAM99.427, andposteriorpairsofgenitalpores.
99.430, 99.431 and 99.433 from the same horizon AmbulacrumIIIveryslightlydepressed aborally;
and locality. All specimens collected by O. H. pore pairs very small. Anterior petals slightly
Melikov from the Acarinina rotimdaniargmata zone. flexuous, curving anteriorly distaUy; very slightly
WAM
Figure2 Antillasterbagmanovi sp. nov. 99.428; holotype, aboral view; from Paradash Group (Middle Eocene),
Daralik,NakhichevanregionofAzerbaijanRepublic;xl.
AsterostomatidechinoidAntillasterfromAzerbaijan 161
elongate, not conjugate; interporiferous zone
narrow, being less than width of pore pairs in
smallest specimen (TL 89.5 mm) toslightly greater
in largest specimen. Posterior petals also very
slightly sunken; straight; slightly longer than
anterior,being46-52%TL; withup to51 pore pairs
in each row; nature of pore pairs and width of
interporiferous zone same as in anterior petals;
slightlywider than anteriorpetals,being8-10%TL.
Terminal ambulacrum plates of petals sometimes
occluded.
Aboral tubercles relatively sparsely distributed;
mm
range in diameter from 0.8 to 1.6 in largest
specimen. Set in a dense field ofmiliary tubercles.
Fasciolesabsent.
Oral surface with moderately convex
interambulacralareasandsunkenambulacra.These
are very weakly sunken ambitally, increasing in
depth to relatively deeply sunken peristome.
Ambulacrum III deeply sunken throughout.
Peristome width 11%TL; moderately crescentic;
posterior of peristome situated 30-35%TL from
anterior ambitus. Labrum strongly arcuate
anteriorly. Plastron length 24-34%TL, being
relatively longer in smaller specimens; width 17-
21%TL; gently convex. Epistemal plates abouthalf
the length of sternal plates, but epistemal and
preanal plates combined are longer than sternal
plates. Periproct inframarginal, transversely oval,
width 12-13%TL. Adoral tuberculation slightly
more dense than on aboral surface; primary
tubercles of more even diameter; up to 1.5 mm;
slightly smaller on plastron than on lateral
interambulacra; set in field of dense miliary
tubercles.
Ontogeneticvariation
While the partially weathered and usually
somewhat crushed nature of the specimens
precludes any detailed ontogenetic assessment of
thespecies, somedifferences areapparentbetween
the smallest specimen (WAM 99.429, test length 89
mm), compared with the other, larger, individuals
(125 to 154 mm). The plastron in the smaller
individual is distinctly narrower, with a length to
width ratioof 1:1.77, compared with 1:1.27- 1:1.40
(n=5)inthelargerones.Furthermore,thepetalsare
slightymoresunken in largerindividuals, as is the
WAM anterior ambulacrum on the oral surface. The
Figure3 Antillaster bagmanovi sp. nov. 99.429; interporiferous zone increases slightly in relative
paratype. A, aboral view, B, adoral view;
from Paradash Group (Middle Eocene), width during ontogeny, being slightly narrower
Daralik, Nakhichevan region of Azerbaijan than the width of the pore pairs in the smallest
Republic;xl. specimen,becomingwider thanporepairwidth in
thelargestspecimens.
sunken; width 8-9%TL; diverging anteriorly at Remarks
about 140°; long, 43-48%TL, extending to the
Kier(1984)consideredthatthereweretwospecies
ambitus; distally open; with up to46 pore pairs in groups of Antillaster present in the Caribbean
each row; innerpores circular, outer pores slightly region. One hecharacterisedby its possession of a
162 KJ.McNamara,O.H.Melikov
WAM
Figure4 Antillasterbagtmnovi sp. nov. 99.432; paratype, adoralview; from Paradash Group (Middle Eocene),
Daralik,NakhichevanregionofAzerbaijanRepublic;xl.
large,veryhigh,steep-sidedtest,withaflatventral EoceneofJamaicaandA.albeariKier, 1984fromthe
surface and very wide petals. Species within this Middle to Late Eocene of Cuba. The apical system
grouprangein age from theOligoceneto Miocene. is less anteriorly situated in A. bagmanovi than in
Kier characterised the second group (which these two species. Moreover, it has sunken petals
comprises species ranging in age from Eocene to that are more anteriorly divergent and narrower
Miocene) by the possession of relatively lower, interporiferous zone. Although sharing slightly
more elongate test, more rounded ventral surface sunken petals, A. bagmanovi can be distinguished
and narrower petals. In its possession of narrow from A. albeari by possessing more anteriorly
petals andrelativelylowertestwithmorerounded divergentpetals, verymuchnarroweranteriorand
ventralsurface,A. bagmanovi resembles thesecond, posterior petals and interporiferous zone, more
probablymoreprimitive,group. vaultedtestanddeeperanteriorambulacrumonthe
Antillasterbagmanovicanbereadilydistinguished oralsurface.
from all other described species of Antillaster by LikeA.elegansJackson, 1922 from theMioceneof
virtue ofitsslightly sunken petals thathave a very PuertoRico,A. bagmanovihasanapical system ina
narrow interporiferouszone, and orally possessiiag similar relative position, a little anterior of central,
a relatively deeply sunken ambulacrum 111. The and slightly anteriorly divergent petals. However,
only other known Eocene species ofAntillaster are tlie Azerbaijan species has slightly sunken petals,
A.amoldiClarkin Arnold andClark, 1927fromthe narrower interporiferous zone, deeper anterior
AsterostomatidechinoidAntillasterfromAzerbaijan 163
A
WAM
Figure5 Antillaster bagmanovi sp. nov. 99.432; paratype. A, partial adoral plating, B, lateral profile; from
ParadashGroup(MiddleEocene),Daralik,NalAichevanregionofAzerbaijanRepublic;xO.9.
164 K.J.McNamara,O.H.Melikov
contracted rapidly to the west in the Oligocene.
Whether Antillaster originated in the west or the
eastern parts of the orginal range is hard to
establish. Evidence from other echinoids indicates
that up until the Early Oligocene there was a
relatively free exchange of taxa across the Atlantic
Ocean,mainlyinawesterlydirection. Forexample,
Clypeaster appeared earlier in the Tethyan region
than in the Caribbean (Ali 1983). Likewise
Echinolampas first appeared in the Mediterranean
region during the Late Paleocene, but did not
appear in the Caribbean region until the Middle
Eocene (Roman 1977). SimilarlyRhyncholampas first
appeared in the Caribbean at this time, although it
is found in the Early Eocene in France (Roman
1977). Comparable east to west migrations are
shown by Eupatagus (Roman 1970) and Maretia
(Roman 1977). Perhaps of more interest than the
likelihood ofAntillasterforming part of this east to
west migration of faunal elements in the Middle
Eocene, is the fact that it disappeared from the
easternpartofitsrangesosoonafter.Thismaybea
function of its mode of life. Kier (1984) suggests
WAM that,likelivingasterostomatids,Antillasterprobably
Figure6 Antillaster bagmanovi sp. nov. 99.429;
paratype, partial adoral plating; from lived on the top of the substrate, lacking features
Paradash Group (Middle Eocene), Daralik, associated with a burrowing mode of life, and in
Nakhichevan region of Azerbaijan Republic; relatively deep water, perhaps up to 800 m deep,
xl. likelivingmembersofthefamily.Itsdisappearance
fromtheLesserCaucasusmayhavebeenassociated
with a shallowing of the sea as Tethys contracted
ambulacrum orally and narrowerlabrum. Another through the Cenozoic and a separation of the deep
species to possess anarrow interporiferous zone is basinsoftheEasternParatethysfromdeeperbasins
A. fernandezi (Sanchez Roig, 1952) from the tothewest.
Oligocene-Miocene of Cuba (Kier 1984). However,
A. bagmanovihasnarrowerpetalsthatareanteriorly
more divergent, a less vaulted test and four, not
three,gonopores.
Antillaster bagmanovi can also be distinguished
from A. vaughani (Jackson, 1922) from the
Oligocene-Miocene of Antigua, Mexico and Cuba
(Kier 1984) by its narrower interporiferous zone,
sunken petals, deeper anterior ambulacrum orally,
narrower peristome and narrower plastron. A.
sancheziLambertin Lambertand Thiery, 1924from
the Early - Middle Miocene of Cuba (Kier 1984)
has, like A. bagmanovi, a very narrow
interporiferouszoneandslightlysunkenpetals,but
the Azerbaijan species has a more central apical
system, more anteriorly divergent petals that are
not so parallel-sided, and a deeper anterior
ambulacrumorally.
BIOGEOGRAPHICIMPLICATIONS
The highly disjunct Eocene populations of WAM
Antillaster, with nine Eocene to Miocene species in Figure7 Antillaster bagmanovi sp. nov. 99.431;
theCaribbeanandoneMiddleEocenerecordinthe pPaarraatdyapseh,Graopiucpal(MsidydslteemEocpelnaet)i,ng;Darfalriokm,
LesserCaucasus,suggestsanoriginallywidespread Nakhichevan region of Azerbaijan Republic;
distributionasfareastastheEasternParatheys that xl5.
AsterostomatidechinoidAntillasterfromAzerbaijan 165
ACKNOWLEDGEMENTS
Jackson, R.T. (1922). Fossil Echini of the West Indies.
We thank Dr KevinKelly, Director ofAzerbaijan ContributionstotheGeologyandPaleontologyoftheWest
Relief International, for his assistance; Vasif Indies306: 1-103.
Melikov for helping with computer services; Dr Kier, P.M. (1984). Fossil spatangoid echinoids of Cuba.
AndrewSmithforhisconstructivecommentsonthe SmithsonianContributionstoPaleobiology55:1-336.
manuscript; Dr Stefan Rivets for assistance with Lambert,J. (1909). Descriptiondes Echinidesfossilesdes
foraminiferal biostratigraphy; Kris Brimmell for terrains mioceniques de la Sardaigne. Memoires de la
photography and Danielle West for help with the SocietePaleontologiqueSuisse35:74-142.
linedrawings. Lambert,J. and Thiery, P. (1924). Essaie deNomenclature
Raisonnee des Echinides. Chaumont, Librairie L.
Ferriere.
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