2016 | 69/2 | 177–185 | 10 Figs. | 2 Tabs. | www.geologia-croatica Journal of the Croatian Geological Survey and the Croatian Geological Society
Palaeobiogeography of the Late Carboniferous brachiopoda from Velebit Mt. (Croatia) Mirko Japundžić1 and Jasenka Sremac2
1 Gruška 16, 10 000 Zagreb, Hrvatska; ([email protected]) 2 University of Zagreb, Department of Geology, Division of Geology and Paleontology, Horvatovac 102a, 10 000 Zagreb, Croatia; (corresponding author: [email protected]) doi: 10.4154/gc.2016.23
Abstract Article history: An abundant and diverse Late Carboniferous brachiopod fauna from Velebit Mt. (Croatia) com- Manuscript received September 30, 2015 prises 63 brachiopod taxa dominated by Productida and Spiriferida. The Spiriferinida, Athyridida, Revised manuscript accepted June 21, 2016 Orthotethida and Rhynchonellata are less common, while the Orthida, Dictyonellida and Tere- Available online June 29, 2016 bratulida occur in very small numbers. Brachiopods are mostly preserved as casts and moulds in shales, limestones and sandstones. Associated fusulinid foraminifera and calcareous algae
Keywords: Brachiopoda, palaeobiogeography, indicate a Kasimovian to Gzhelian age for the brachiopod–bearing deposits. The global biogeo- palaeoecology, Late Carboniferous, Velebit Mt., graphic distribution of brachiopod taxa indicates the probable seaways and brachiopod migra- Croatia. tion routes, along the Euramerican shelves.
1. INTRODUCTION Brachiopods are common marine macrofossils in the Late Car- to 6 km wide, representing the core of an anticline, with a NW– boniferous sedimentary rocks of Velebit Mt. They have been col- SE strike (Fig. 1). They exhibit a variety of ancient environments lected since the beginning of the 19th century and stored in the varying from shoreline forests and swamps, through coastal and Croatian Natural History Museum. shallow shelf biomes (SREMAC, 2012; CLEAL et al., 2015). Car- Upper Carboniferous (Pennsylvanian) sedimentary rocks boniferous shelves were densely populated with foraminifera from Velebit Mt. crop out in an elongate belt, 40 km long and up (mostly fusulinids), calcareous algae, brachiopods, crinoids, bry-
Figure 1. Simplified geological map of the research area, with the location of brachiopod observations (black squares?) (after CLEAL et al., 2015). Geologia Croatica 2015 Database; www.fossilworks.org and/or revised ( (underMuseum History 96 numbers).inventory 2014. The brachiopod collection is stored in the Croatian Natural SALOPEK by (1942, collected were 1948) study and this by from the present brachiopods authors between Carboniferous 2012- MATERIALS2. AND METHODS (SREMAC, horizons Gzhelian 2012). fossils, including brachiopods from increases the Kasimovian to (KOCHANSKY, 1955; SREMAC, 2012). The diversity of marine lowish shales and siltstones of Late Kasimovian to Gzhelian age se to quences, be estimated of Kasimovian age, Carboniferous with overlying yel Late the of horizons lower the in common ing to SALOPEK, 1942, 1948). sandstones“ „Fusulinid are more 1942, 1948) and sandstones“,greywackes („fusulinid SALOPEK, 1942, 1948), limestones ( and siltstones ( shales yellowish-brown types: lithological three in discovered rine macrofossils, including the brachiopods (SIMIĆ, 1935), were Mt. were recognized quite early by SALOPEK (1942, 1948). Ma SREMAC, 2012; et CLEAL al., 2015). (KOCHANSKY-DEVIDÉ, 1955,megaflora land 1970; and sils (late Kasimovian age or earliest Gzhelian) stratigraphic was The determined from microfosregion. this of Carboniferous Late SOKAČ et al., 1974, 1976a, b, c; VELIĆ et al., 2009). (IVANOVIĆ1:3001973,al., Croatia 000 of et 1976; Map gical gical maps of Yugoslavia on the 1:100000 scale and Basic Geolo geological of Basic in setting the iswider area described Geolo his students between 1935 and 1938 (SALOPEK, 1942, lected 1948). during mapping campaigns undertaken by The M. Salopek and CHANSKY-DEVIDÉ, 1955,KO 1948; 1970;1942, SREMAC, 2005, 2012).(SALOPEK, macrofossils other and ozoans preserved shell and inner cast with visible medianFigure septum 2. – area C; c) cast with visible external ornamentation, surrounded with fusulinid casts – area C. 178 All together 110 brachiopod specimens were determined determined were specimens 110brachiopod together All Velebitof Carboniferous the in units lithological Different the for typical is deposition siliciclastic-carbonate Mixed col been has deposits these from assemblagefossil rich A ) and asserted to 63 taxa. ) and asserted Modes of preservation of Late Carboniferous brachiopods in the Velebit Mt.: a) inner cast (with gaps in places of dissolved median septa) – area C; b) partly cau SELDEN , equivalent to to the Beds according Auernig , 2007; and references therein; August EMIG cv, et al., 2013; Paleobiology according to SALOPEK, cf, accord ------ ltl b poutd ( productids by pletely chiopods (Fig. 1, Tab 1., 2. Figs. 1, 3–7) bra different with C) and B (A, clusters three into grouped be (SELDEN, 2007). Localities Palaeontology with carboniferous brachiopods can Invertebrate on Treatise the to according vised 1884; SCHELLWIEN, 1892, 1900; MOORE, 1979). Taxa are re 1816; (e.g. VONSCHLOTHEIM, papers WAAGEN, published 1.). Initial brachiopod taxonomy studies were based upon several Spiriferida, Rhynchonellida, DictyonellidaOrthida, Spiriferinida, and Terebratulida (Table Athyridida, Orthothetida, ductida, level. (Fig. 2), and it is not always possible to determine them to species of Velebit Mt. (SIMIĆ, 1935; SALOPEK, 1942, 1948). sedimentary core the ofslopes on anticline continental the forming the rocks Carboniferous in discovered were Brachiopods TAXA 3. BRACHIOPOD (2015) reconstruction. Blakey’supon based is localities brachiopod Carboniferous the horizons, while shales and scarce limestones prevail in the the in prevail limestones numerous youngerBrachiopod fossils particularly horizons. are scarce and shales while horizons, wackes („ sandstones“) are Fusulinid more common in the older pling (SALOPEK,areas 1942, 1948; SREMAC, 2012), but grey lids (Tabs. 1., 2.; Figs. 3 c, 6). dictyonel and orthids rhynchonellids, spiriferids, spiriferinids, most a hundred specimens of productids, orthotethids, athyridids, al assemblages, with variable(C)brachiopod highly comprises of part belt the Carboniferous Velebitin Mt. and the Lika regio ids and rhychonellids occur (Tabs. 1., 2.; Figs. 3 b, 5). wards The (B),Eastern brachiopod diversity increases and, the first orthoteth this area are stored in the Natural History Museum. Moving east from specimens brachiopod 6 4).Altogether, (Tab. a, 1.;Fig.3 granulifer The The Carboniferous palaeogeographic map chosen to present All the aforementioned facies types occur in all three sam three all in occur aforementionedtypes faciesthe All com almost (A)localitiesdominated are westernmost The taxa belong The determined to nine taxonomic groups: Pro Brachiopods are in most cases preserved as casts and moulds , Paramesolobulus latesinuata Paramesolobulus iordcu lineatus Linoproductus , with small small with , Geologia Croatica 69/2 Croatica Geologia , , Neochonetes Neochonetes (Scale bar 1cm) Dielasma n ------
Geologia Croatica 179 cf
cv FACIES
cau
C (7-11) C
E) ® B (5-6) B
LOCALITIES (W LOCALITIES A (1-4) A
WAAGEN ) PRATTEN ) CHAO ) and and ) ) ) ) ) in in ) ) ) ) ) ) ) ) CHAO ) ) SCHLOTHEIM ) ) WAAGEN ) ) ) SCHELLWIEN STUCKENBERG ) WAAGEN WAAGEN GRAB SCHELLWIEN ) ) MARTIN ( BEYRICH SOWERBY GRABAU ’ORBIGNY KROTOV ( OWEN COY PHILLIPS NIKITINO
NORWOOD ( TSCHERNYSCHEW ( ( WAAGEN
( WAAGEN SCHELLWIEN MORTON (d
SCHELLWIEN ( SCHLOTHEIM MARCOU WAAGEN ) grandis grandis KEYSERLING MARCOU MARCOU SCHELLWIEN TOULA MARTIN SCHELLWIEN SCHELLWIEN ( sp. KUTORGA cf. cf. sp. SCHELLWIEN sp. SCHELLWIEN ( sp. sp. longispinus longispinus sp. sp. sp. sp. cf. sp. KUTORGA sp. List of Carboniferous brachiopod taxa grouped through localities in the Velebit Mt. and facies types. and facies types. Mt. Velebit the localities in through taxa grouped brachiopod List of Carboniferous sp. sp. sp. sp. sp. sp. sp. TEREBRATULIDA Dielasma Rhipidomella pecosi ( pecosi Rhipidomella Enteletes DICTYONELLIDA paotechovensis Isogramma Hustedia mormoni ( mormoni Hustedia confinensis aff. Rhynchonella ( alpinum Stenoscisma ( timorensis Uncinunellina ORTHIDA Reticularia Spirifer Squamularia RHYNCHONELLIDA Neospirifer cameratus ( cameratus Neospirifer ( fasciger Neospirifer ( zitteli Plicatocyrtia ( lineata Reticularia Elinoria carnica ( carnica Elinoria ( lyra Eliva ( semiplana Martinia Martinia Ambocoelia Ambocoelia rostrata Bajkuria ( fritschi Choristites Choristites Spiriferellina cristata ( cristata Spiriferellina Spiriferina (Mc laminosa Spiriferina SPIRIFERIDA trautscholdi Alphachoristites Retzia ( mormoni Hustedia SPIRIFERINIDA Waagen ornata Callispirina Streptorhynchus semiplanus ( semiplanus Streptorhynchus Streptorhynchus RHYNCHONELLATA ATHYRIDIDA Derbya (Derbyia) Derbya (Derbyia) Derbya Meekella pelargonatus Streptorhynchus Transennatia gratiosa ( gratiosa Transennatia punctatiformis Vediproductus Waagenoconcha ORTHOTHETIDA altestriata (Derbyia) Derbya Productus mölleri Productus transversalis Productus Productus Productus Neochonetes variolata variolata Neochonetes strophomenoides (Sommeriella) Neochonetes latesinuata Paramesolobus sp. Plicatifera Linoprodutus pusilla Marginifera aagardi Megousia granulifer Neochonetes Echinoconchus elegans Echinoconchus punctatus Echinoconchus incisus Incisius lineatus Linoproductus Chonetes mamontovi mamontovi Chonetes papilionacea Chonetes Chonetes curvirostris Comuquia subpunctata Cubacula STROPHOMENATA PRODUCTIDA echidniformis Breileenia gruenewaldti Chaoiella CLASS/ORDER/GENUS/SPECIES Table 1. Table revised). research 1942, 1948 and own SALOPEK, (SIMIĆ, 1935; MirkoJapundžić JasenkaSremac:and Palaeobiogeography CarboniferousLate the of brachiopoda Mt.(Croatia) fromVelebit Geologia Croatica Figure 4. limestones) (Tab. 1.). siltstones; (shalesand units lithological three the of two in and or gratiosa Transennatia limestones), and greywackes siltstones, and shales – units lineatus ductus as such environments, ofmarine variety a to adapting taxa (Tab. 1). The exceptions are several tolerant capabletaxa, of rarelythe same comprise very areas to note three these tant that evidently towards increases the East (Tab. 2., Fig. 8). It is impor ferent fossil assemblages (Tabs. 1., 2., Fig. 3). Palaeobiodiversitydif with areas three into grouped clearly are finds Brachiopod of brachiopod genera 4.1. Palaeoecology and the local distribution DISCUSSION 4. to SALOPEK, 1942, 1948) (Tab. 1.). sandstones“, („fusulinid greywackes the in served ob were taxa brachiopod few 1942,1948).a SALOPEK, Only fossil assemblage is preserved in the limestones ( to SALOPEK,Beds according Auernig 1942, 1948). A different ( siltstones and shales the in diverse and (area C) comprise an abundant and variable brachiopod fauna, with the predomination of Productida. comprises large fenestellid casts and rather small brachiopods; b) Central area (B) is characterizedFigure3. by the domination of flattened orthotethids; c) Eastern localities 180 Mould of a Differentiation faciesof and brachiopod assemblages the threein examined areas with Carboniferous deposits in Velebit Mt.: a) Westernmost area (A) (present in all three areas and in all lithological all in and areas three all in (present Linoproductus , present in the central and eastern areas, areas, eastern and central the in present , brachial valve (a) and bioeroded cast of cau , equivalent to the the to equivalent , cv, cf, according to
according according Linopro Productus transversalis - - - - to to the productids. thotethids and rhyncnonellids (Tabs. 1., 2; Figs. 5 b, c) in addition size small probably food sources. restricted indicate soft with bottom, low depositional rates. Their low diversity and the on(Tabs. spines by1.,b). anchoredlived Figs.a, 2., They 4 larger the of exception the with taxa, productid 1948; KOCHANSKY-DEVIDÉ 1955, 1970; SREMAC, 2012). 7 a), bivalves (Fig. 7 b) and gastropods (SIMIĆ, 1935; with SALOPEK, calcareous algae, brachiopods (Figs. 6 a-h), bryozoans (Fig. with a predomination of and crinoid fusulinids ossicles, together land of vicinity the (SREMAC, 2012;indicates CLEAL etmegaflora al., 2015). terrestric Marine fossils are preserved diverse, Slovenia (SCHELLWIEN, 1892). Velebitonly in neighbouring deposits in Mt. Permian the in and 1993). most probable deep subtidal depositional area (KALASHNIKOV, areas. to open subtidal, basinal from ronments, discovered in this area could have lived in different envi marine The western area (A, Fig. 1) is characterized by several small The The eastern area (C, Fig. 1) is generally rich in fossils. A well The presence of Figs. (B, area central 1, The 3 b, 5 some a-d), or comprises Rhynchonella Rhynchonella confinensis TSCHERNYSCHEW (b) (Scale bar 1cm) Linoproductus Linoproductus Neochonetes Neochonetes variolata from locality cluster A. (Scale bar 1cm) seems seems to be endemic, occurring and and Transennatia (Fig. 4 a) indicates a Geologia Croatica 69/2 Croatica Geologia Linoproductus (Figs. 5 a, d) - -
Geologia Croatica - 181 Choris sp.) (g) and the spiriferid ( spiriferid the and (g) sp.) Spiriferina (f), spiriferinid ( (f),spiriferinid Marginifera Marginifera (d) and and (d) (d) with visible spine bases from the locality B. (Scale bar 1cm) bar (Scale B. locality the from bases spine visible with (d) Linoproductus Linoproductus (b), (b), Linoproductus Dictyoclostus (a, c, e), e), c, (a, , (b,c) and segment of of segment and (b,c) , Productus Derbya (a), (a), Productid genera (a-f): genera Productid Transennatia Transennatia ) (h) brachiopoda from locality group C. (Scale bar 1cm) bar (Scale C. group locality from brachiopoda (h) ) MirkoJapundžić JasenkaSremac:and Palaeobiogeography CarboniferousLate the of brachiopoda Mt.(Croatia) fromVelebit Figure 6. 6. Figure tites Figure 5. 5. Figure Geologia Croatica Figure 8. Table 2. biota. marine diversification of significant the enabled and ents beaches to the morefrom ronments, favourable niches.subtidal of envi variety a indicate moulds(Figs. features 1-6). these All tests numerous and highly diverse, are also mostly present as orcasts fusulinid of 2012).(SREMAC, influence fresh-water although Brachiopods, dissolution probable a indicate 1955) complete KOCHANSKY, 1948; (SALOPEK, or Partial ment. in some sulinids sandstone layers indicate a the environ coastal to the Scolithos ichnofacies.be attributed can layers, and the within butalso surfaces, bedding on present Figure 7. 182 BRACHIOPOD GROUP Terebratulida Dictyonellida Orthida Rhynchonellida Spiriferida Spiriferinida Athyridida Orthothetida Productida Well sorted conglomerates and the aligned orientation of fu siltstones, and sandstones the in common are fossilsTrace Erosion of the uplifted Variscan Mountains providednutri VariscanMountains Erosion uplifted of the Total number of brachiopod taxa in three different Carboniferous areas in Velebit Mt. Total brachiopod diversity and area specific diversity for the three Carboniferous areas in Velebit Mt. Cast of a fenestellid bryozoan (a) and a scallop TAXA 63 16 26 1 1 2 4 4 2 7 Acantopecten A (1–4) 6 1 0 0 0 0 0 0 0 5 LOCALITY GROUP A,B, C sp.(b) from locality group C. (Scale bar 1cm) - - - - B (5–6) 7 0 0 0 1 0 0 0 3 3 shelves and possible sea-ways and climate gradients, recently recently gradients, climate and sea-ways possible and shelves continental of position the to related closely was distribution geographic Their oceans. and seas all in present were ganisms or these and 2006) al., (SHU-ZHONGet diversity brachiopod 4.2. Palaeobiogeographic studies 2007; MERINO-TOMÉ et al., 2009). PRINS, (WINKLER Spain from weredescribed trends Similar ZAROVA et al., 2009; SREMAC, 2012; et CLEAL al., 2015) area was a ofpart NE Gondwana, situated near the equator (VO study the Carboniferous, Late the 2014). ANGIOLINI, During bystudied several authors (ANGIOLINI et al., 2007; & BERRA The Late Carboniferous was a time of a slight increase in in increase slight a of time a was Carboniferous Late The C (7–11) 56 16 24 0 1 2 3 4 2 4 cau 54 15 23 1 1 2 2 3 2 5 FACIES 16 cv 0 0 0 0 4 1 0 2 9 Geologia Croatica 69/2 Croatica Geologia cf 2 0 0 0 1 0 0 0 0 1 - - Geologia Croatica ------183 ap ttp://fossil Megousia Megousia h was also wide also was is very peculiar Chonetes Chonetes Echinoconchus , observed in Velebit Mt. (records from: PED from: (records Mt. Velebit in observed , ). They lived all along the continental continental the along all lived They ). Chonetes , , Derbya , , Neospirifer 2016 and references therein references and con 2016 southern the of shelves the on also but Euramerica, long- of rather a shelves had they that possible is It (Fig.9). islands and tinents genus The stage. larval swimming lived spread, present all around Euramerica (Fig. 9). However, geographic the range of the genus Pan of shelves the along migrations possible indicating 10) (Fig. The thalassa, genus rather than Palaeotethys. those of was common on the northern of Euramerica shelves and its ) were widely widely were ) ossilworks.org, ossilworks.org, have been recorded in almost been recorded have Neospirifer, Derbya Neospirifer, ). Palaeobiogeography based upon Blakey’s (2016) reconstruction. Grey colour: continents; white colour: oceans/seas; PS: Palae PS: oceans/seas; colour: white continents; colour: Grey reconstruction. (2016) Blakey’s upon based Palaeobiogeography ). Neospirifer Neospirifer Distribution of the Late Carboniferous cosmopolitan brachiopod genera ( genera brachiopod cosmopolitan Carboniferous Late the of Distribution and references therein references and The SHNIKOV, 1998; SOBOLEV et al., 1998; WENDT et al., WENDT et al., 1998; et al., SOBOLEV 1998; SHNIKOV, Tolerant brachiopod taxa ( brachiopod Tolerant In order to identify possible migration routes, some of the 2001; SCHNEIDER, 2003; GONG et al., 2007; http://f 2007; al., et KALA GONG 2003; SCHNEIDER, 2001; spread. Disocveries of of Disocveries spread. a thousand Carboniferous and Permian1995; collectionsKORA, all over the 1991; SUTHERLAND, 1954; (PEDERSON, world common Carboniferous brachiopod genera from Velebit Mt. were were Mt. Velebit from genera brachiopod Carboniferous common further for studies. selected palaeobiogeographic MirkoJapundžić JasenkaSremac:and Palaeobiogeography CarboniferousLate the of brachiopoda Mt.(Croatia) fromVelebit works.org/ works.org/ arrow. the by indicated area Study Euroamerica. EA: Gondwana; G: Panthalassa; PA: otethys; ERSON, 1954; SUTHERLAND, 1991; KORA, 1995; KALASHNIKOV, 1998; SOBOLEV et al., 1998; WENDT et al., 2001; SCHNEIDER, 2003; GONG et al., 2007; 2007; al., et GONG 2003; 9. Figure SCHNEIDER, 2001; al., et WENDT 1998; al., et SOBOLEV 1998; KALASHNIKOV, 1995; KORA, 1991; SUTHERLAND, 1954; ERSON, Geologia Croatica groups, with a predominance of awith and spiriferids. groups, productids predominance brachiopod nine to belonging determined, were taxa 63 gether to All preserved. well not cases most in but diverse. and rich very is VelebitMt. from fauna brachiopod Carboniferous The 5. CONCLUSIONS continental shelves of Euramerica, near the equator (Fig. 10). the genus (Fig.10). interesting is Alternatively,equator the near pearance PA: Panthalassa; G: Gondwana; EA: Euroamerica. Study area marked by arrow. based upon Ron Blakey’s reconstruction ( PEDERSON, 1954; SANDO et al., 1975; SCHNEIDER, 2003; GONG et al., 2007; HEIM et al., 2009; Figure 10. 184 Distribution of the Late Carboniferous specialist brachiopod genera ( Transenatia is rather endemic, appearing on the eastern h ttp://cpgeosystems.com/300Marect.jpg, July 2015). Grey colour: continents; white colour: oceans/seas; PS: Palaeotethys; - Echinoconchus the associated fusulinid fauna. the fusulinid associated rockcropping belt.Carboniferous in observed was coastal diversity fine-grained clastic deposits atthe eastern part of brachiopod the out highest The ognized. ent areas with diverse brachiopod associations can be clearly rec differ clastic deposits.Three the in fine-grained highest abundance with graywackes, and limestones siltstones, and shales in A Kasimovian to Gzhelian age was proposed on the basis of Brachiopods were observed in the form of moulds and casts h , ttp://fossilworks.org/ Megousia , Transennatia and references therein , observed in Velebit Mt. (observations from: Geologia Croatica 69/2 Croatica Geologia ). Palaeobiogeography - - - Geologia Croatica ------
185 Basic Geologic Map of SFRY SFRY of Map Geologic Basic 848A:1–83 Basic Basic Geologic Map of SFRY Basic Geologic Map of SFRY 1:100000, 1:100000, SFRY of Map Geologic Basic Basic Geologic Map of SFRY 1:100000, 1:100000, SFRY of Map Geologic Basic − in Croatian].− Geološki zavod, Zagreb, Sa Zagreb, zavod, Geološki Croatian].− in − − in Croatian].− Geološki zavod, Zagreb, Sa Zagreb, zavod, Geološki Croatian].− in − April 2016 April Clastic and carbonate deposits (Carboniferous, Permian – C, Permian (Carboniferous, deposits carbonate and Clastic Treatise Treatise on Invertebrate Paleontology, Part H, Brachiopoda last checked checked last , , − in Croatian].− Geološki zavod, Zagreb, Savezni geološki zavod, zavod, geološki Savezni Zagreb, zavod, Geološki − in Croatian].− zavod, geološki Savezni Zagreb, zavod, Geološki − in Croatian].− – in Croatian].− Rad JAZU, 274, 75 p. 75 274, JAZU, Rad Croatian].− in – – in Croatian].– In: VELIĆ, I. & VLAHOVIĆ, I. 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STEPHENSON, G., MUTTONI, M., GAETANI, l., ANGIOLINI, History Museum) for the macrophotography of theHistory the brachiopodsmacrophotography of for Museum) Zagreb) of University Science, of (Faculty KOŠČAL Robert and preparation for graphics,of and to both theirreviewers for con ACKNOWLEDGEMENT Natural (Croatian JAPUNDŽIĆ Sanja to grateful are authors The sphere records are less common and represented by tolerant gene tolerant along by the continentalrepresented shelves and of Euroamerica. common Southern less hemi are records sphere cosmopolitan the e.g. MirkoJapundžić JasenkaSremac:and Palaeobiogeography CarboniferousLate the of brachiopoda Mt.(Croatia) fromVelebit