Marcin Machalski & David M. Martill

Annales Societatis Geologorum Poloniae (2013), vol. 83: 99–104.

FIRST PTEROSAUR RE MAINS FROM THE CRE TA CEOUS

OF PO LAND

Marcin MACHALSKI1 & Da vid M. MARTILL2

1

In sti tute of Paleobiology, Pol ish Acad emy of Sci ences, Twarda 51/55, 00-818 Warszawa, Po land; e-mail: mach@twarda.pan.pl

2

School of Earth and En vi ron men tal Sci ences, Uni ver sity of Portsmouth, Burnaby road, Portsmouth PO1 3QL, United King dom; e-mail: Da vid.Martill@port.ac.uk

Machalski, M. & Martill, D. M., 2013. First pterosaur re mains from the Cre ta ceous of Po land. Annales Societatis Geologorum Poloniae, 83: 99–104.

Ab stract: The first re cords of pterosaurs from the Cre ta ceous of Po land are re ported, on the ba sis of frag men tary re mains from the ma rine Up per Albian (Lower Cre ta ceous) of the Annopol Anticline, cen tral Po land. The new ma te rial con sists of four bone frag ments, ten ta tively in ter preted as: 1) a por tion of wing pha lanx; 2) a me dial el e ment of fused skull bones (pa ri etal crest?); 3) a frag men tary car pal or tar sal; and 4) a dis tal pha lanx of the pes (or a very small frag ment of a long cer vi cal ver te bra). Pre vi ously, only the re mains of ma rine ver te brates have been re ported from the Cre ta ceous of the Annopol area. The pterosaur fos sils stud ied most prob a bly be longed to in di vid u als that died while over the sea. The pos si bil ity that they rep re sent re mains dropped from float ing car casses, in tro duced into the ma rine en vi ron ment by rivers, is re garded as less prob a ble, as there are no re mains of di no saurs or other ter res trial fauna in the Annopol de pos its.

Key words: Pterosauria, Albian, Cre ta ceous, Po land.

Manu script re ceived 24 Oc to ber 2013, ac cepted 25 No vem ber 2013

IN TRO DUC TION

Pterosaur bones are very frag ile due to their thin walls and a gen eral re duc tion of in ter nal trabeculae (e.g., Elgin and Hone, 2013). The best pre served pterosaur re mains oc -cur in fos sil Konservat-Lagerstätten (sensu Seilacher, 1970), where skel e tons may be fully ar tic u lated, uncrushed and have as so ci ated soft tis sues, in clud ing wing mem -branes, pre served. These de pos its are ex em pli fied by the litho graphic lime stones of the Ju ras sic Solnhofen For ma -tion in Ger many, the Early Cre ta ceous Yixiang For ma -tion of China and the Santana For ma tion of Brazil (e.g., Well-nhofer, 1991; Unwin, 2006).

Pterosaur re mains also oc cur in fossiliferous de pos its of the con cen tra tion type (Konzentrat-Lagerstätten), typ i fied by the mass oc cur rence of or ganic re mains. The best-known pterosaur-bear ing de posit of this lat ter type is the low er most Cenomanian Cam bridge Greensand of east ern Eng land. This Konzentrat-Lagerstätte has yielded hun dreds of

remanié pterosaur fos sils of Late Albian age which, though

disarticulated and frag men tary, nev er the less pro vide a valu -able source of data on pterosaur tax on omy, phy log eny and di ver sity (e.g., Unwin, 2001; Buffetaut, 2006; Martill and Unwin, 2012).

Here the au thors de scribe and il lus trate rare pterosaur ma te rial from an other highly-fossiliferous de posit of the con cen tra tion type, namely the phosphorite-bear ing Up per Albian sands of the Annopol Anticline, cen tral Po land (Fig. 1A, B). The new ma te rial is a part of a rich ver te brate as -sem blage col lected from the Annopol area by a team, led by the first au thor. These col lec tions were made dur ing sub ter -ra nean ex plo -ra tion of the aban doned phosphorite mine Jan 1 (Machalski et al., 2009) and screen wash ing of the sed i ment at the sur face lo cal ity of Kopiec (see Fig. 1B for lo ca -tion of both set tings and Popov and Machalski, in press, for a de tailed de scrip tion of the screen wash ing pro ce dures).

The ma te rial de scribed here, al though highly frag men -tary, rep re sents the first pterosaur re mains from the Pol ish Cre ta ceous. The ma te rial is now housed at the In sti tute of Paleobiology of Pol ish Acad emy of Sci ences, Warszawa, Po land (ZPAL).

GEO LOGIC BACK GROUND

A highly con densed sed i men tary suc ces sion, rang ing from the up per most Lower Albian to the Lower Turonian, is ex posed around the town of Annopol (Fig. 1A, B), along the

limbs of a small anticline (the Rachów or Annopol Anti-cline, see Samsonowicz, 1925, 1934; Marcinowski and Ra-dwañski, 1983; Walaszczyk, 1984, 1987; Machalski and Ken nedy, 2013). The suc ces sion re cords the early phases of the great Cre ta ceous ma rine trans gres sion (Marcinowski and Radwañski, 1983; Walaszczyk, 1987) and is ex tremely

thin (con densed) in com par i son to co eval strata in ad ja cent ar eas, due to its lo ca tion on a sub ma rine high (Cieœliñski, 1976).

With ref er ence to the transgressive-re gres sive model of Juignet (1980) and Han cock (1990), the Annopol suc ces sion is in ter preted as a se ries of transgressive units, num -Fig. 1. Geo logic back ground of the pterosaur re mains from the Annopol Anticline, A. Li thol ogy and stra tig ra phy of the midCre ta -ceous suc ces sion of the Annopol Anticline (mod i fied af ter Walaszczyk, 1984). 1. A quartzose sand stone re placed up wards by quartz sands trun cated by a bur rowed sur face. 2. Quartz sands with glauconite and sand stone nod ules near the top; unit 2 is capped by a phosphorite ho -ri zon. 3. Quartz sands with glauconite pass ing up wards into marly sands with glauconite and phos phates. 4. Quartzose marls with glauconite and phos phates in the lower part. 5. Glauconitic marls in fill ing bur rows in the un der ly ing unit, trun cated by a hardground with phos phate and glauconite im preg na tion. 6. Strongly glauconitic marls with bur rowed omis sion sur face at the top. 7. Lime stone with rare phos phatic nod ules and glauconite, trun cated by a hardground with glauconitic min er al iza tion. 8. Marls with bur rowed top. 9. Marls with black flint; b – bur rows. B. Geo logic map of the Annopol Anticline (mod i fied af ter Walaszczyk, 1987) with lo cal i ties stud ied and the lo ca -tion of the study area in Po land. C. Close-up view of the up per Up per Albian pterosaur-bear ing unit 3 at the lo cal ity Kopiec. LP – lower part of the Phosphorite Bed, UP – up per part of the Phosphorite Bed. Adapted from Popov and Machalski (in press) with per mis sion from Elsevier

bered 1–9, capped by lay ers of re worked phos phatic nod ules and clasts, hardgrounds and bur rowed omis sion/ero -sional sur faces, which re flect a se ries of re gres sion max ima (Fig.1A).

The pterosaur spec i mens re ported herein, four in to -tal, come from unit 3 as ex posed at Kopiec (Fig.1A, C). This unit cor re sponds to the up per part of the so-called Phosphorite Bed, dis tin guished in the Annopol sec tion by pre vi ous au thors (see e.g., Samsonowicz, 1925, 1934; Radwañski, 1968; Marcinowski and Radwañski, 1983, 1989).

Unit 3 (see Fig. 1A, C) is c. 50 m thick and is com posed of quartz sand with abun dant glauconite, pass ing grad u ally up wards into quartz-glauconitic marl. A dis tinct bed of pho- sphatic clasts and fos sils oc curs in the up per part of unit 3. Its up per bound ary is sharp, whereas the lower bound ary is dif fuse due to bioturbation (Marcinowski and Walaszczyk, 1985; Walaszczyk, 1987). The phos phatic clasts are c. 1 cm in di am e ter, but in the up per part of the layer they com -monly form larger ag gre gates, up to 12 cm in di am e ter. These ag gre gates are com posed of up to three gen er a tions of clasts show ing the same ma trix, but dif fer ing in in ten sity of phosphatisation, sug gest ing multi-phase intraformational re work ing of the phos phatic ma te rial.

The fos sil as sem blage from unit 3 is dom i nated by pho-sphatised sponges, calcitic-shelled bi valves Aucellina

gryphaeoides and Pycnodonte vesiculosa, and ros tra of the bel

-em nite Neohibolites ultimus. Shells of rhynchonellid and terebratulid brachi o pods, serpulid tubes and shrimp re mains are sub sid iary com po nents of the in ver te brate as sem blage. As so ci ated ver te brates in clude rel a tively com mon teeth and ver te brae of sharks and bony fishes, less com mon chimae-roid den tal plates (Popov and Machalski, in press), teeth, frag ments of jaws and ver te brae of ich thyo saurs and plesiosaurs, as well as re mains of ma rine tur tles. The most nu -mer ous and best pre served fos sils oc cur in the lower and mid dle part of the unit, those from its up per por tion are com monly poorly pre served, frag mented and over grown by phos phates.

AGE OF THE PTEROSAUR MA TE RIAL

The pterosaur-bear ing unit 3 of the Annopol suc ces sion is dated on the ba sis of spec i mens of the ammonite

Morto-niceras (Subschloenbachia) sp., pre served as at tach ment scars on the left valves of some oys ters (Machalski and Ken nedy, 2013). These spec i mens show the clos est af fin ity with those of Mortoniceras (Subschloenbachia) which are char ac ter is tic of the up per Up per Albian Mortoniceras

perinflatum Zone (for a de tailed dis cus sion and zonation of

the rel e vant part of the Albian Stage see Machalski and Ke-nnedy, 2013, ta ble 1).

There is no ev i dence of any biostratigraphically older fos sils within unit 3, and the late Late Albian is taken to be the age of the pterosaur ma te rial de scribed herein. This con -trasts with the sit u a tion from the low er most Cenomanian Cam bridge Greensand as the pterosaur re mains con tained in the lat ter unit are thought to rep re sent older, Late Albian, ma te rial (e.g., Martill and Unwin, 2012).

DE SCRIP TION AND COM PAR I SONS

Spec i men ZPAL V.38/390 (Fig. 2A–C) is the most di -ag nos tic of the pres ent ma te rial. It measures16 mm in length, is 10 mm high and 5 mm wide. It has an ul tra thin (thin nest 0.3 mm – thick est 0.6 mm) bone wall, and the typ i -cal (for pterosaurs) lamellar struc ture can be seen in SEM (Fig. 2C). There are two prom i nent trabeculae vis i ble, the re main der of the crosssec tion sur face be ing filled with sed -i ment (a pneu mat-ic cav -ity dur -ing l-ife) (F-ig. 2B, C). The crosssec tion is an oval with two op pos ing, slightly flat -tened sides and is typ i cal of the cross-sec tion found in non-azhdarchid pterodactyloid pterosaur wing-fin ger pha lan ges. As sum ing this iden ti fi ca tion to be cor rect, in dor sal view (as though the wings were out stretched) there is a gen tle ex-pansion (slightly more prom i nent on one side than on the other), sug gest ing this is a por tion of the pha lanx diaphysis close to ei ther the dis tal or prox i mal ter mi na tion.

FIRST PTEROSAUR RE MAINS FROM THE CRE TA CEOUS, POLAND

101

Spec i men ZPAL V.38/273 (Fig. 3A1–A4) is 15 mm in

its max i mum di men sion. The thin na ture of the bone wall of this frag ment iden ti fies it as pterosaurian. It has two fea tures that might al low a rea son able at tempt at iden ti fi ca tion of the el e ment. There is a suboval articulatory sur face with a slight depression on one mar gin. The bone is bro ken in what might be its dis tal part. It is a rather tab u lar el e ment and there is a slight in fla tion on one side. It is pos si bly a car pal or tar sal.

Spec i men ZPAL V.38/816 (Fig. 3B1–B4) is 25 mm in

its max i mum di men sion. The thin bone wall of this tri an gu -lar frag ment iden ti fies it as pterosaurian. It has a strongly de vel oped ‘me dian’ ridge, from which fas cia de scend sym met ri cally to de fine an an gle of 90 de grees. The ridge de -creases and dis ap pears to ward the apex, and the (pre sumed) left mar gin ap pears en tire, while the cor re spond ing mar gin of the other side is slightly dam aged. Nev er the less, the bone prob a bly had a tri an gu lar ter mi na tion in life. It may be a me -dial el e ment of fused skull bones, such as a pa ri etal crest. This iden ti fi ca tion is ex tremely ten ta tive.

Spec i men ZPAL V.38/1195 is not il lus trated here, be

-cause of its very poor pres er va tion. This small frag ment (c. 10 mm in its max i mum di men sion) is also very thin walled, and most likely a pterosaur bone. There are few fea tures to di ag nose it, but what ap pears to be a grooved articulatory sur face sug gests that it is pos si bly a dis tal pha lanx of the pes. How ever, it has a flat face and slightly ex panded ter mi -na tion, and as such, could also be a very small frag ment of a long, slen der cer vi cal ver te bra.

Be cause of the highly frag men tary na ture of the ma te rial, com par i sons with other pterosaurs are dif fi cult. Nev er -the less, an af fin ity with non-pterodactyloid monofenestra-tans, Rhamphorhynchidae, and all pre-Mid dle Ju ras sic pte-rosaurs can be ruled out, on ac count of the Cre ta ceous age of the new ma te rial. The thin wall of the bones also rules out af fin i ties with Dsungaripteroidea in which there is a ten -dency to bone-wall thick en ing (Martill et al., 2013). Thus the af fin i ties of the new Pol ish ma te rial prob a bly lie within Ornithocheiroidea and/or Azhdarchoidea, all of which are known from the Cre ta ceous of Eu rope (Unwin, 2001; Martill et al., 2013).

Fig. 3. Pterosaur re mains from the Annopol Anticline, A. Spec i men ZPAL V.38/273 (a frag men tary car pal or tar sal) in four dif fer ent views (A1–A4). B. Spec i men ZPAL V.38/816 (me dial el e ment of fused skull bones, pa ri etal crest?) in four dif fer ent views (B1–B4)

DIS CUS SION AND CON CLU SIONS

In gen eral, the re cord of pterosaur re mains in Po land is poor. Prior to the pres ent study, the sole pub lished bodyfos sil ex am ple was an iso lated tooth from the Up per Ju ras -sic (Up per Tithonian) Corbulomina ho ri zon of the Kcynia For ma tion ex posed in the Owadów-Brzezinki quarry near the town of Tomaszów Mazowiecki, cen tral Po land (Kin et

al., 2013). Foot prints at trib ut able to pterosaurs were descri-

bed from the top most part of the Up per Ju ras sic (Lower Ki-mmeridgian) Oo lites and Platy Lime stones (in for mal litho-strati graphic unit) ex posed in the Wierzbica quarry, south of the town of Radom (Pieñkowski and NiedŸwiedzki, 2005).

The fos sils de scribed here, al though frag men tary, are suf fi ciently di ag nos tic to dem on strate the pres ence of ptero- saurs in the Pol ish Cre ta ceous, and are the first re mains of Cre ta ceous pterosaurs to be re corded from this coun try. One may hope that fur ther palaeontological ex plo ra tion of the Up per Albian de pos its in Annopol anticline and other parts of Po land will yield more di ag nos tic pterosaur re mains.

The Late Albian pterosaur re mains doc u mented here are the only nonma rine el e ments in the ver te brate as sem -blages re corded from the Annopol anticline (Samsonowicz, 1925, 1934; Marcinowski and Radwañski, 1983; Machalski

et al., 2009; Popov and Machalski, in press). In view of the

ab sence of any non-ma rine strata (or their rem nants) in the Annopol sec tion, the au thors sug gest that the pterosaur re -mains be longed to in di vid u als that died while over the sea. The pos si bil ity that they rep re sent re mains dropped from float ing car casses in tro duced into the ma rine en vi ron ment by rivers is re garded as less prob a ble as there are no di no -saur or other ter res trial fauna re corded from these de pos its. A land area in south ern Po land, sep a rat ing the Pol ish epi-con ti nen tal ba sin from Tethys in Late Albian (e.g., Jasko-wiak-Schoeneichowa, 1978) may have been a hab i tat for the pterosaurs de scribed here.

Ac knowl edge ments

The pres ent work was sup ported by the Pol ish Na tional Cen tre of Sci ence grant no N N307 529238 “Taphonomy of ver te -brates from con densed mid-Cre ta ceous de pos its of the Annopol anticline (north-east ern bor der of the Holy Cross Moun tains)” to Marcin Machalski. Wies³aw Liwiñski, Mayor of the town of An-nopol, is thanked for en thu si as tic sup port and as sis tance dur ing field work at Annopol. Agnieszka Kapuœciñska, Adam Zaremba, Maciej Duda, Maciej Ma³ysiak, Krzysztof Nejbert, Micha³ An-dziak and the fire men of the OSP Sucha Wólka are thanked for their par tic i pa tion in field work at the lo cal ity Kopiec. Gra¿yna Dziewiñska is ac knowl edged for pho to graphs of the spec i mens, and Aleksandra Ho³da-Michalska for com puter pro cess ing of the fig ures. Danuta Olszewska-Nejbert is thanked for dis cus sions on the Annopol phosphorites and Da vid Hone and Mariusz Kê-dzierski for crit i cal re views of the ear lier ver sion of this pa per.

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