(Surgeonfish) from the Miocene Deposits of the Medvednica Mt

2016 | 69/2 | – | 4 Figs. | www.geologia-croatica Journal of the Croatian Geological Survey and the Croatian Geological Society

First record of Acanthuridae (surgeonfish) from the Miocene deposits of the Medvednica Mt. Kristina Tripalo1, Sanja Japundžić2, Jasenka Sremac3 and Marija Bošnjak2

1 Šestinski vrh 43a, Zagreb, Croatia; ([email protected]) 2 Croatian Natural History Museum, Department of Geology and Paleontology, Demetrova 1, Zagreb, Croatia; ([email protected], [email protected]) 3 Faculty of Science, University of Zagreb, Department of Geology, Division of Geology and Paleontology, Horvatovac 102a, Zagreb, Croatia; ([email protected]) doi: 10.4154/gc.2016.15

Article history: Abstract Manuscript received September 02, 2015 Middle Miocene deposits at the Dubravica locality in SW Medvednica Mt. (N Croatia) contain Revised manuscript accepted March 03, 2016 various marine fossils, including one tooth of the surgeonfish (Acanthuridae). This is the first Available online May 16, 2016 such record in Croatia and the second in the wider region, besides the Vienna Basin. The appearance of surgeonfishes in the Paratethys coincides with the Miocene thermal maximum. Their Keywords: Central Paratethys, Medvednica Mt., distribution was probably controlled by available reef habitat and their palaeobiogeography and Miocene, Badenian, Acanthuridae migration routes are yet to be studied.

1. INTRODUCTION western slopes of Medvednica Mt. The surgeonfish tooth was Northern and north-eastern Croatia during the Miocene epoch discovered during recent field work (BOŠNJAK et al., 2014; TRI- were covered by the Paratethys Sea (Fig. 1). Abundant and diverse PALO et al., 2015). marine fossils can be found in sedimentary rocks of this age. Medvednica Mt. acted as an island near the SW coast of the Para 2. GEOLOGICAL SETTING tethys, and Miocene shelf deposits are exposed today in the form The investigated area is a part of the Miocene deposits belt sur- of a belt around the mountain core. rounding Medvednica Mt., and is situated on its south-western Middle Miocene marine environments of Medvednica Mt. slopes (Fig. 1). Geological and palaeontological features of this were divided into the ‘’Dolje’’ development in the south-western area were summarized in publications following the Basic Geo- part of Medvednica Mt., the ‘’Čučerje’’ development in the cen- logical Mapping, by ŠIKIĆ et al. (1977, 1979), and later by ŠIKIĆ tral part of Medvednica Mt., and the ‘’Zelina’’ development in (Edit.) (1995). the south-eastern part of Medvednica Mt., each characterized by During the Miocene, the whole area palaeogeographically different fossil assemblages (KOCHANSKY, 1944). belonged to the south-western margins of the Central Paratethys Miocene deposits of the south-western part of Medvednica (Fig. 1), and geotectonically to the Pannonian Basin System (PA Mt., considered to be a part of the ‘’Dolje’’ development, have VELIĆ, 2002). been studied by many authors (e.g. KOCHANSKY, 1944; KO- An approximately 16 metre high subvertical outcrop, with CHANSKY-DEVIDÉ, 1957; ŠIKIĆ et al., 1977, 1979; KOCHAN sedimentary rocks of ca. 11.5 m in thickness, is exposed at the SKY-DEVIDÉ & BAJRAKTAREVIĆ, 1981; VRSALJKO et al., Dubravica locality near the path leading to the Veternica Cave 2006, 2007; BOŠNJAK et al., 2014). (Fig. 2). Triassic dolomites in the base are transgressively over- The Dubravica locality (Fig. 1), records the first occurrence lain with dolomitic breccias, which gradually pass into breccias- of fossil Acanthuridae in Croatia, and is situated on the south- conglomerates, and are later replaced with floatstones containing ARTICLE IN PRESS

Figure 1. Location of the Dubravica locality and palaeogeographic map of Central Paratethys (after VRSALJKO et al., 2006; from BOŠNJAK et al., 2014). Geologia Croatica (e.g. KOCHANSKY, 1944; ŠIKIĆ et al., 1977, 1979;as ŠIKIĆ, 1995; fossil 2015). collected al.,2014; al., et The et NJAK TRIPALO a fishtooth from the familyAcanthuridae wasdiscovered (BOŠ layer (Fig.marl grey 2),light the sponge spicules.In and spines andfish bryozoans, teeth bones, echinoid ostracods, raminifera, layers (Fig. 2), juvenile containing molluscs, and fragmented fo analyzed three the of all in microfossilpresent assemblagerich wide and ca. 5 m high. con („ buildup reef coralgal a with replaced laterally are marls south, (Fig. horizon this 2). within occurs 0.5thick, mca. bivalves with gastropods, lowish and sandstone of intercalation molluscs. yelwith An thick, m3 ca. marl, grey fossils. byof The aupper part the light-iscolumn characterized contains molluscs, solitary Itlayer.corals, bryozoans,this overlies echinoids thick, and m trace 2.5 ca. marl, grey dark of posed bivalve moulds and dispersed dolomitic cobbles. A horizon com- the surgeonfish tooth is marked with arrow. light grey marl, 3: intercalation of yellow sandstone.etal., 2014). LocationNumbers 1, 2, and 3mark investigated of thelayers; horizon1: dark grey marl, 2: with Figure 2.

Lithothamnium semblages when compared with the available published data data published available the with compared when semblages temporary age. The visible part of this buildup is ca. 10 m 10 ca. is buildup this of part visible The age. temporary Marls and sandstones from the Dubravica locality contain a 15ca. ofsampled the section, towards the m vicinity the In Schematic geological column of the Dubravica locality (after BOŠNJAK limestone“, sensu KOCHANSKY, 1944) of 1944) KOCHANSKY, sensu limestone“,

ARTICLE IN PRESS - - from the investigated Dubravica locality, one tooth from the the from was also found. Acanthuridae tooth one locality, Dubravica investigated the from CHANSKY, 1944; ŠIKIĆ et al., 1979). Among other microfossils Lamna era, and itera, is strongly1970) related to diet (TYLER, (Fig. 3). al., 2013). Dentition can vary dramatically (SORENSON et within and acrossinvertebrates and gen detritus organic matter, mal tively wide variety of dietary resources, including planktonic ani TYLER, 1970). Acanthurus well denticulated on one side only; other genera ( small, minute or absent; slightlymostlyconicaldistallybutcompressed, denticulationswith variously MedvednicaMt., fossil teethof reflectingtheir solid construction. Fromthe Badenian deposits of Paratethys. Fish teeth are particularly common and well preserved Fish remains are common fossils in the Neogene deposits3. RESULTS of Central wastooth found on the 0.2 sieve. mm surgeonfish The discovered. were teeth fish fossil where tions those with mesh size of 0.2 and 0.125 mm as these were the frac 0.5, the 0.20.125 and sieves. given mm was to Specialattention peroxide. After twenty-four hours, samples were flushed through Soft marls were then crushed, soaked in the water with hydrogen ofDepartment Geology, Division of Geology and Palaeontology. the in Wetpared laboratory, at the Faculty of Science Zagreb, in (Fig. 2) were followinganalyzed wet sieving. Samples were pre Selected sampleslayers the from three of the Dubravica locality MATERIAL2. AND METHODS nian) age (BOŠNJAK et al., 2014; BOŠNJAK et al., 2015). Bade (Upper Miocene Middle of are deposits the that suggest KOCHANSKY-DEVIDÉ, 1981; VRSALJKO et al., 2006, 2007), PIKIJA in VELIĆ & VLAHOVIĆ, 2009; BAJRAKTAREVIĆ & Figure 3. Figure 2014). al., et (PELLISSIER influences biogeographic Neogene flects re largely and families fish reef other to similar is distribution al.,2013;SONet al.,2015).geographic et LUDT modern Their SOREN therein; BELLWOOD,references & and 1993 CELL and are one of the dominant groups of fish onthe coral reefs(PUR DALL, 2001). aggressive(RAN behaviour inter- in intra-specific used beand 1993),on the pedunclecaudal (WINTERBOTTOM, which may fishes) derivesfromthe presence of a scalpel-like modified scale The Acanthuridae includes fishes whose common name (surgeon FAMILY: Acanthuridae BONAPARTE, 1832 ORDER: (BLEEKER, Perciformes 1859) CLASS: Actinopterygii (KLEIN, 1885) Surgeonfishes are primarily herbivores,areSurgeonfishes primarily butconsume arela seas, subtropical and tropical in liveToday, all acanthurids sp., and Examples of teeth of the Acanthuridae family: family: Acanthuridae the of teeth of Examples ) – fixed teeth, compressed with well developed denticulations (after Mylobatis Ctenochaetus sp . have been observed to date (KO date to observed havebeen – numerous, elongated, moveable teeth Chrysophrys Paracanthurus, Zebrasoma sp., Geologia Croatica 69/2 Croatica Geologia Naso Odontaspis – fixed teeth, fixed – sp. and - - - ------, Geologia Croatica ------ - - - -

some

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(BASSO et al., 2008), al., et (BASSO (MEYER, 1842; SCHULTZ, 2003). SCHULTZ, (MEYER, 1842; Lithothamnion Mesophyllum roveretoi roveretoi Mesophyllum from the Badenian of Austria (BELLWOOD & Acanthurushaueri and Findings of acanthurid teeth in the Paratethys realm coincide coincide realm Paratethys the in teeth acanthurid of Findings pos the of discussion opens record surgeonfish Croatian The rotfishes (Labroidei: Scaridae) with Scaridae) a of (Labroidei: rotfishes Description a New Species Calotomus Wien, Museum Ann. Naturh. Austria.– of (Badenian) from the Miocene Middle 55−71. A, 92 Italy, Bolca, of Monte Eocene the from Teleostei) (Acanthuroidei, relatives their Gi Ric. St. forms.− recent and fossil both to formula Blot the of application with 13–92. 6, Bolca Terz. ac. ocene maërl: the Croatian ”Litavac”.– Geologia Croatica, 61/2–3, 333–340. doi: doi: 333–340. 61/2–3, Croatica, Geologia ”Litavac”.– Croatian the maërl: ocene 10.4154/GC.2008.25 10.1007/BF01626074 doi: 11−19. 15/1, Reefs, Coral assemblage.− fish innova functional and fish fossils, reefs: coral on herbivory of years million Fifty 1−8 281/1781, B, Society Royal the Proc.of tions.− The tooth from the Dubravica locality differs from the Aus the fromdiffers locality Dubravica the from tooth The The appearance of acanthurids in the Paratethys corresponds Paratethys the in acanthurids of appearance The it was controlled by distributionPalaeoecologically suit of careum times in association with bryozoans and, herbivo other sporadically,one Interestingly only 2006). al., (VRSALJKO et corals rous reef fish has been recorded from this region, theparrotfish Calotomus 1991). SCHULTZ, with the Miocene thermal maximum. canWe discuss why are acanthurid of their teeth and size small the scarce; so finds these rather bizarre shape may be the reason that palaeontologists do material. recognize registeror not themin sieved the posed of coralline algae: algae: coralline of posed BELLWOOD, D.R. & SCHULTZ, O. (1991): A Review of the Fossil Record of the Par the of Record Fossil the of Review A (1991): O. SCHULTZ, & D.R. BELLWOOD, BLOT, J. & TYLER, J.C. (1990): New genera and species of fossil surgeonfishes and and of surgeonfishes fossil and species New genera J. (1990): J.C. & TYLER, BLOT, ACKNOWLEDGMENT of College the from BELLWOOD David thankProf. authors The Marine and Environmental Sciences, techniques analyzing James tooth fish Cookin University,help for Australia, Townsville, and discussion support. wish We to thank the Division of Geo REFERENCES aMi of flora coralline The (2008): T. & GRGASOVIĆ, D. VRSALJKO, D., BASSO, reef coral earliest the Bolca; of Monte Fishes Eocene The (1996): D.R. BELLWOOD, O. (2014): & S.J. BELLWOOD, BRANDL, C.H.R., GOATLEY, D.R., BELLWOOD, 6. CONCLUSIONS 6. south-west the locality in Dubravica the from tooth fish small A surgeon of record first the represents Mt. Medvednica of part ern teethsimilar area, nearby the In Croatia. in (Acanthuridae) fishes to referred been have Basin Vienna the of deposits Miocene from typethe Sci of Faculty DepartmentGeology, of Palaeontology, and logy ence, University of Zagreb, and the Croatian for reviewers Naturalboth gratefulareto support.We their Historyfor Museum, theiruseful comments suggestions. and sible sible ancestral populations and fish migration routes, but such along contemporaryrocks other of reexaminationrequiresstudy more of discovery possible the and formations reef Paratethys the surgeonfish remains. trian samples in shape, but is of similar size. It may belong to an- to belong may It size. similar of is but shape, in triansamples position different a from but species, same the to or species, other withinjaw. the with the Miocene extent thermal biogeographic maximum, possible and and it populations opens ancestral the discussion about Acanthuridaethe acrossParatethys. of able benthicable habitats, in this case coralgal or coralgal-bryozoan buildups. reef ------

(MEYER, ARTICLE IN PRESS IN ARTICLE Acanthurus haueriAcanthurus Acanthuridae fossil tooth, a) labial side; b) lingual side. lingual b) side; labial a) tooth, fossil Acanthuridae Acanthuridae gen. et sp. indet. Acanthuridaesp. et gen. Material: one partially preserved tooth with accreted sedi The rich fossil record indicates that acanthurids reached a Some fossil taxa of the Acanthuridae are known from the Miocene surgeonfishesPreviously only been have reported Miocene reefs and reef-like structures in Paratethys which As shown on Fig. 4, the exposed part of the small incisiform small the partof exposed the 4, Fig. on shown As ma the structure multidenticulate of typical a has tooth The 1842). They are of similar size as the specimensimilarthefrom Dubravica,as size of areThey 1842). but differ from it in shape. As we are lacking other parts of the skeleton, we can only speculate whether this is a different spe 5. DISCUSSION5. studied best the among surgeonfishes are Acanthuridaeor Fossil percomorph fishes. fossil significant diversityby the Middle Eocene(SORENSONet Bolca al., Monte the from surgeonfishes diverse and Abundant 2013). by studied were species, 16 and genera extinct 14 least at locality, 2000). BANNIKOV, SORBINI TYLER, TYLER, & & (BLOT authors several 1990; & TYLER b; 2005a, 1999, TYLER, b; 1998a, The Monte Bolca fish assemblage represents theoldest(BELL known assemblage fish reef coral a as defined be may that fauna and is deposit at a 1996) the fossil key birth WOOD, modernof Several genera share the 2014) et 2014). al., coral al., reefs (BELLWOOD et (BELLWOOD time this at form tooth multidenticulate suggesting that they may have been feeding on short algal turf 1993). BELLWOOD, & (PURCELL deposits (TYLEROligocene 2000; TYLER & SORBINI, 1998; TYLERMICKLICH,2011). & Vi the of Miocene fromthe and (TYLER, 1997) fromSulawesi enna Basin (MEYER, AustrianSCHULTZ, 2003). 1842; acan as described were teeth fish thurid spe same the to belongs tooth the that possible also is it but cies, cies, but had a different withinposition & the (SCHULTZ jaws 2004). BELLWOOD, acanthurids for habitats predominantly were suitable - com were Figure 4. 4. Figure Tripalo et al.: First record of Acanthuridae (surgeonfish) from the Miocene deposits of the Medvednica Mt. ment (Fig. 4) (Fig. ment tooth is almost black in Dimensioncolor. of the tooth (in mm) width. 1.13 height, 1.95 are: jority of acanthurids. One apical denticulation, two denticulations denticulations two denticulation, apical One acanthurids. of jority Den visible. are tooth the of side left the on three and right the on increase the both sides ticulations in on the base of size towards surface Its inward. curved slightly is tooth investigated The teeth. - Con 4). (Fig. side lingual and labial the both on smooth almost is sideringpartialthe withgeneratooth preservation, of andrange - pos not was level generic the at determination configuration, this sible. Geologia Croatica SORENSON, L., SANTINI, F., CARNEVALE, G. & ALFARO, M. E. (2013): A multi- SORBINI, L. & TYLER, J.C. (1998b): A new species of the Eocene surgeon fishgenus SORBINI, L. & TYLER, J.C. (1998a): A new genus and species of Eocene surgeonfish SCHULTZ, O. & BELLWOOD, D. R. (2004): SCHULTZ, O. (2003): The Middle Miocene Fish Fauna (excl. otolithes) from Mülbach RANDALL, J. E. (2001): Surgeonfishes of Hawai’i and the World.− Mutual Publish. and BOŠNJAK MAKOVEC M., KARAICA, B., SREMAC, J., VRSALJKO, D., HAJEK– BOŠNJAK, M., KARAICA, B., SREMAC, J., VRSALJKO, D., HAJEK-TADESSE, V., PIKIJA, M. (2009): Litavac i klastične naslage s vulkanitima (baden – M PELLISSIER,LEPRIEUR,L, F., PARRAVICINI, V., COWMAN, P. KULBICKI, M., PAVELIĆ,(2002):South-WesternD.The CentralBoundaryParatethys.–of Geologia MEYER, H. v. (1842): Iguana (?) Haueri, aus dem Tertiärgebilde des Wiener Beckens.− LUDT, W. B., ROCHA, L. A., ERDMANN, M. V. & CHAKRABARTY, P. (2015): Skip KOCHANSKY-DEVIDÉ, V. & BAJRAKTAREVIĆ, Z. (1981): Miocen (baden i sarmat) KOCHANSKY-DEVIDÉ, V.(1957): fauniO miocena tortonskomoi „šliru“ Medved KOCHANSKY, V. (1944): Fauna marinskog miocena južnog pobočja Medvednice (Za PURCELL, S. W. & BELLWOOD, D. R. (1993): A functional analysis of food procure

locus timetree of surgeonfishes (Acanthuridae, Percomorpha) wit ture and supraneurals in acanthurids.− St. Ric. Giac. Terz. Bolca, 7, 21–34. Pesciarichthys ma.− St. Ric. Giac. Terz. Bolca, 7, 7–19. (Acanthuridae)from Monte Bolca, Italy, with similarities to the Recent Zebraso nodontinae).– Ann. Naturh. Museum Wien 105A, 287–305. the Lower and Middle Miocene of Central Europe (Osteichthyes, Labridae, differentTrigo parts of the same species seum Wien, 104 A, 185−193. am Manhartsberg and Grund near Hollabrunn, Lower Austria.− Ann. Naturh. Mu Bishop Museum Press Hawai’i. tus MAKOVEC,M.(eds.): International Scientific Meeting 100 development – Dubravica locality.– TADESSE,In: VRSALJKO,V., GRUBER,D., SREMAC,A., JEFTINIĆ, J. &S. &BOŠNJAK POSEDI, N. (2015): Miocene Dolje Students Geological Congress), Volume 8, p. 11. NW Croatia).– Acta Mineralogica-Petographica, Abstract series (5th Internationalblages and environments in the wider area of VeternicaGRUBER, cave A.,(SW JEFTINIĆ,Medvednica S.& Mt.,POSEDI, N. (2014): Middle Miocene fossil assem 1016−1019. doi: 10.1126/science.1249853 diversity.–Science,refugiapreservedQuaternary(2014):fish reefcoralD. 344, LITSIOS,OLESEN,G.,S., BELLWOOD, S.,WISZ,M. MOUILLOT,D.R.,& Croatica, 55/1, 83–92. doi: 10.4154/GC.2002.08 Beiträge zur Petrefacten-Kunde, 5, 33−34. 10.1016/j.ympev.2014.12.017 thuridae:Prionurus). MolecularPhylogenetics Evolution,and166–172. 84, doi: pingacross the tropics: Theevolutionary history of sawtail surgeonfishes (Acan 43–48. MedvednicaGebirgevon(Kroatien,Randes Jugoslawien)].− vjesnik,33, Geol. westlichen des Sarmat) und (Baden [Miozän Medvednice. ruba najzapadnijeg (1956), 39−50. 10 vjesnik, Geol. Gebirge)].− (Zagreber Medvednica von „Schlier“ tonischen nice(Zagrebačka gora). [Über die Fauna des marinen Miozäns und über den tor 2/3, 171−280. (Zagreber Gebirge)].− Geol. vjesnik Hrv. drž. geol. zav. Hrv. drž. geol. muz.,grebačke Vol. gore). [Miozäne marine Fauna des südlichen Abhanges der Medvednica Field trip Excursion Guidebook, 3−7. of Vandaof Kochansky-Devidé,ofMember Academy,Full 9 Zagreb, ment in two surgeofishes species 86−87. (GeologicaltheRepublicMapof Croatia).of Hrvatski geološki institut, Zagreb, I. & VLAHOVIĆ, I. (eds.): Tumač Geološke karte Republike Hrvatske 1:300.000 (Acanthuridae).− from Monte Bolca, Italy, with comments on caudal peduncle arma

Envir. Biology of Fishes, 37, 139−159. Acanthurus nigrofuscus Trigonodon jugleri Trigonodon oweni ARTICLE IN PRESS , a Chiseltooth Wrasse from and and th Ctenochaetus Birth Anniversary Asima jugleri are h revised family 4 ).− In: VELIĆ, th -11 th April,

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Mus. Civ. St. Nat. Verona, 24, 29–36. from the Eocene of Monte Bolca, Italy (Perciformes, Acanthuridae).− Gazolaichthysvestenanovae VE Arambourgthurus LIĆ, D., MIKNIĆ, M., BRKIĆ, M., KOVAČIĆ, M., HEĆI –

BasicGeological Map of SFRY 1:100000, Zagreb in Croatian].− Institut za geološka istraživanja, Zagreb (1972) Basic Geological Map of SFRY 1:100000, , a new genus of hypurostegic surgeon fish (Acan Marosichthys from the Eocene ofMonte Bolca, Italy Acanthurus gaudryi th Birth AnniversaryVandaof Ko , aputative, tetraodontiform, actu Geologia Croatica 69/2 Croatica Geologia Eonaso th -11 th , an Antillean fos April 2015, Ab April Naso De Zigno 1887

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