Mitt. Mus. Nat.kd. Berl., Gcowiss. Reihe 4 (2001) 189-221 05. 1 I. 2001
An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England
David M. Unwin]
With 15 figures and 2 tables
Abstract
The Cambridge Greensand, a remaniC deposit that crops out in Cambridgeshire. eastern England, has yielded numerous. though fragmentary, late Early Cretaceous (Albian) vertebrate fossils including more than 2000 isolated pterosaur bones. So far. 32 species of pterosaur have been proposed in connection with the Cambridge Greensand material. but there has been and continues to be considerable confusion concerning the validity of these taxa, their relationships to each other and to other pterosaurs, and the material upon which they were established. A comprehensive systematic revision identified eleven valid species distributed among three families: the Ornithocheiridae (Ornithoeheirus sirnus and possibly a second. as vet un- named species of Ornithoeheirus. Coloborhynchus capito, Coloborhynchus sedgwickii, Anhangiiern cuvieri, and An1iririgiierrr jirtoni); the Lonchodectidae (Lonchodectes compressirostris, Lonchodectes rnachaerorhynchus, Lonchorlrcres niicrotiori and Lonchodecies plarystornus): and a species of edentulous pterosaur (Ornithostonin sedgwicki) that may represent the earliest record for the Pteranodontidac. It is possible that some of the taxa currently recognised represent sexual dimorphs (Colohor- hynchus cupito and Coloborhynchus sedgwickii, Lonchodectes compressirostris and Lonchodectes mrrc~i~ierijr~i~richrts),or dis- junct populations of a single species (Ornithocheirus sirnus and Ornithocheirus sp., Lonchodectes conipressirosrri.v and Loriclio- dectes microdon) and that there may be as few as seven valid species, but the Cambridge Greensand pterosaurs are too poorly known to demonstrate this at present. The Cambridge Greensand pterosaur assemblage is similar to a slightly youngel-. but much smaller assemblage from the Lower Chalk of England and shares some elements, such as ornithocheirids. in com- mon with many other late Early and early Late Cretaceous assemblages. It is distinguished by the absence of tapejarids and the presence of Lonchodectes which, so far, is only known from the Cretaceous of England. The disparity in taxonomic composition is possibly related to ecological differentiation, and might also reflect some provincialism in late Early and early Late Cretaceous pterosaur faunas.
Key words: Ptcrosaur, Lower Cretaceous, Cambridge Greensand, England, pterodactyloid, Ornithoeheirus.
Zusammenfassung
Der Cambridge Greensand, eine in Ostcngland aufgeschlossene RemaniC-Ablagerung, hat zahlreiche Wii-beltiere aus der obcren Unterkreide (Alb) geliefert. Darunter fanden sich mehr als 2000 isolierte Pterosaurierknochen. Insgesamt wurden aus dem Greensand bis zu 32 Flugsauriertaxa beschrieben, was zu einer betrachtlichen taxonomischen und nonienklatorischen Verwirrung gefuhrt hat, die bis heute andauert. Eine vollstandige Revision erkennt 11 Arten aus drei Familien an: (I) die Ornithochciridae (Ornithoeheirus sirnus und vielleicht eine zweite, bislang unbenannte Art von Ornitlzocheirus,sowie Colohor- kynchirs cupito, Coloborhynchus sedgwickii, Anhunguera cuvieri und Anhanguera fittoni); (2) die Lonchodectidae (LorIc/70- dectes cornpressirostri,r,Lonchodectes nzachnerorhynchw, Lonchodectes microdon und Lonchodecres platysromiis): und schlielj- lich einen zahnlosen Flugsaurier (Ornithostonin serlgwicki), der zu keiner der vorgenannten Familien gehbrt und sich als slratigraphisch altester Nachweis der Pteranodontidae erweisen konnte. Es ist nicht auszuschlieflen. dass einige der gegenwar- lig erkannten Taxa eher einen ansgepragten Sexualdimorphismus illustrieren denn taxonomisch distinkte Arten darstellen (Colohorl~~nt~hiasciipito und Coloborhynchiis scdgwickii, Lonchodectes cornprssirosfris und Lonchodectes ninchoeror-liy11ci,its) oder sogar lediglich Endpunkte einer intraspezifisch variablen Population (Ornithoeheirus sirnirs und Ornithocheirus sp.. Lori- choilectes conil,rc.ssirostris und Lonchodecies microdon). In dieser strengeren Fassung bestunden nur sieben gultige Artcn. cloch leider sind die Flugsaurier des Cambridge Greensand zu sclilecht bekannt, um diese Fragen zu beantworten. Die Flus- saurierfauna des Cambridge Greensand ahnelt jungeren kreidezeitlichen Faunen aus dein Lower Chalk von England. Weiter- hin cnthalt sie Faunenelemente, wie etwa Ornithocheiriden, die auch fur zahlreiche andere Faunen der hohen Unterkreide und tiefen Obcrkreide charakteristisch sind. Das Fehlen von Tapejariden und das Auftreten des anscheinend cndeniischen Lonchodecres sind weitere Kennzeichen des Cambridge Greensand. Die Zusammensetzung dcr Pterosaurierlaunen folptc olfenbar okologischen Differenzierungen und illustriert eincn gewissen Provinzialismus an der Grenze Unter-Oberkrcidc.
Schliisselwiirter: Pterosaurier, Unterkreide, Cambridge Greensand, England. Pterodactyloldea, Ornirhochcirii~.
' Museum fur Naturkunde, Zentralinstitut der Humboldt-Universitat zu Berlin, InvalidenstraRe 43. D-10115 Berlin. Ger- many. Received March 2001, accepted July 2001
( WILEY-VCH Verlag Berlin GnibH. 13086 Berlin. 2001 1335 lc)43/01/0411-0189$ 17 50 t iOi0 190 11nwiii. D. M.. Cretaceous pterosaui- assemblage from England
Introduction flattened skeletons found in many other deposits and lent itself to a series of important functional In the mid 1800s large scale exploitation of the studies (Hankin & Watson 1914, Bramwell Rr Cambridge Grecnsand. a basal Upper Cretac- Whitfield 1974. Watson 1974, Frey & Riess eous remanik dcposit that cropc out in Eastern 1981). Pterosaur material from the Cambridge England (Womam R. Taylor 1969). principall!. Greensand continues to be highly significant in around Cambridge. led to the recovery of many that although better preserved Cretaceous ptero- fossi 1 re in ai n s i ncl udins ni o re than 2 ()OO fragm e n - saw fossils have now been found, it forms the tary. but uncrushed bones of pterosaurs. At that b.cisis .'. for a number of important taxa that are time. pterosaJ1-s \vere still relatiisel! po~rl!~ also known from other Cretaceous deposits in- known and. ipart from sonic isolated bones cluding the Hastings Sands, Gault Clay and from the Wea den (O\ven 1846) and Chalk (Bo- Chalk ol' England, the Kem-Kern beds of Moroc- werbank 185 1. Owen IS5 1 a. 185 1 b) of England. co. the Crato and Santana Formations of Brazil these nere tht first substantial remains to be re- and many others (see reviews in Wellnhofer co\rered from the Cretaceous. The Cambrid~e 199la. LJnwin et al. 2000: tab. 1; Fig. 1). Greens anct p t t , rosa urs ~vere t h c subject of nia n!' The main problem posed by the Cambridge papers by three eminent British palaeontologists: Greensand pterosaurs. and it is just as relevant Richard Owen ( 185 1 b. 1 SS9a. 1 S59b. I ShOa. today as it was for Owen. Seeley and Hooky, is 186Ob. 1 861. 1574). Harry Seek!- (1864a. 1 8h4b. the nature of their preservation. The material 1 864~.1 865a. 1865b. 1866;). 18h9a. I S69b. 1870. consists ol' fragmentary bones. which. although 1871. 1876a. 1381. 189la. 189117. 1901) and Wal- uncrushcd. are rarely complete and, contrary to ter Hooley (15 14). These publications had a pro- Seele)*'s comments (e.g.. 1869a), were not (or found impact on our understanding of ptero- cannot now be shown to have been) preserved saurs. principally btith regard to their taxonomy. in association and must therefore be treated as system ati cs an il evolution a ry his t or!! ( t'. g.. Art h a - isolated elements. Consequently, the main chal- ber 1922, Plieiiinger 1930. Kuhn 1967. Wcllnho- lenge for all those who have studied these ptero- fer 1978. 1991 a). The tlirt.e-diriiensional preser- saurs has been to determine how many taxa are vation of the Cambridge Greensand bones also present and which bones belong to which taxa. stood in sharp contrast to the compressed and Fortunately. Richard Owen established the prac-
North + South Ameria Australasia
Lysaya Gora, Anhanguea, Lonchodectes Saratov. Russia Anhanguera Kern Kern beds, Morocco Azhdarchidae, Coloborhynchus, ionchode,:tes. Ornithocherrus, ?Pteranodontidae, Tapejandae Paw Paw Fm., Ornrthostcrna Texas, USA Coloborhynchus Toolebuc Frn., Australia Anhanguera Zuunbayan Svita, Lagarcito Fm., Khuren-Dukh, San Luis, Argentina Mongolia Pferodaustro Coloborhynchus Santana Fm., Araripe, Braz Anhanguera, Brasileodactylu Cearadactylus, Coloborhyncl Ornithocheirus, Tapejara, Tupuxuara
Crato Frn., Araripe, Brazil Arthucdactylus, ?Azhdarchidz Tapelara Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 4 (2001) 191
tice of basing new species on jaw remains, since, Second, difficulties are posed for taxonomic in his opinion (e.g., Owen 1860a), the skull and assessments of other Cretaceous pterosaurs. jaws were the most diagnostic part of the skele- especially those that appear to be similar to ton - a proposition that has been borne out by Cambridge Greensand forms. In the late 1800s. more recent studies of pterosaurs from other de- numerous, poorly founded species of Cretaceous posits (e.g., Wellnhofer 1968, 1970, 1975, 1978, pterosaur were assigned to Ornithocheirrrs. often 1987, 1991a, 1991b, Bennett 1994, 2001, Kellner with little or no supporting evidence, by taxo- & Tomida 2000). nomic revisionists such as Lydekker (1888). Re- Owen (1859a, 1859b, 1861) erected four spe- vision of these taxa has been difficult because of cies of pterosaur on the basis of the Cambridge the uncertainty surrounding Ornithocheirirs. Greensand material, three of which are still valid Moreover, in the last three decades. many new today (Unwin et al. 2000). Seeley published Early Cretaceous pterosaurs have been found many new names (principally Seeley 1869a, (summarised in Wellnhofer 1991a, Unwin et al. 1870), both for species and for higher level taxa, 2000), but, even when they show remarkable si- but with little regard for the fledgling taxonomic milarity to Cambridge Greensand forms, the ten- procedures of the 19th century. Not surprisingly, dency has been to assign them to new genera most of his taxa are highly dubious and the man- and species. For example, even though the corre- ner in which Seeley proposed and then sponding region of the skull of BSP 1987 I 46 (mis)treated the few names that are valid, most (Wellnhofer 1987: fig. 2) from the Santana For- notably Ornithocheivus Seeley, 1869a, confused mation of Brazil is strikingly similar to the holo- both contemporary and subsequent workers. type of Omithocheirw sinzits from the Cam- Hooley (1914) attempted to revise Seeley’s bridge Greensand, as some workers have noted work, but was only partially successful and some (Unwin 1988, Fastnacht in press), Wellnhofer of his proposals, such as the erection of the (1987) assigned this skull to a new genus and
genus Lonchodectes, though taxonomically valid, species: ‘Tropeognathus’ nzesemhrinirs ( E Or- have been largely ignored. During the latter half nithocheirus mesembrinus). of the 20th century various authors tried to Third, the uncertainty surrounding the taxo- make sense of the published material, but this nomic validity, systematic relationships and, con- generally led to further confusion. For example, sequently, the skeletal morphology of important. in the standard work on pterosaurs, published as but poorly known taxa such as Omirhocheinrs part of the Handbuch der Palaoherpetologie ser- has made it difficult for researchers to incorpo- ies, Wellnhofer (1978), like some earlier workers rate Cambridge Greensand pterosaurs into phy- (Khozatskii & Yur’ev 1964, Kuhn 1967), erro- logenetic analyses (see for example Howse 1986. neously cited Pterodactylus compressirostris (= Bennett 1989). This, in turn, has hindered recon- Lonchodectes compressirostris) Owen, 1851a as struction of the evolutionary history of pterodac- the type species for Ornithocheirus and the tyloid pterosaurs. adoption of this error by some authors (e.g., The account presented here represents work Kellner 1990, Olshevsky 1991, Mader & Kellner begun in 1984, under the supervision of the late 1999, Kellner & Tomida 2000) has further com- L. B. Halstead. A principal aim was to revise the plicated pterosaur taxonomy. taxonomy of Cambridge Greensand pterosaurs The confusion surrounding the taxonomy and and to utilise the results for reviewing the taxon- systematics of Cambridge Greensand pterosaurs omy and systematic relationships of other ptero- continues to hinder our understanding of ptero- dactyloid pterosaurs. Much of the work on the saurs in three important ways. First, while it is Cambridge Greensand pterosaurs was completed evident that the fossil material from the Cam- by 1990 and formed the basis of a PhD disserta- bridge Greensand represents a relatively diverse tion (Unwin 1991). The concurrent discovery and important Cretaceous pterosaur assemblage and description of relatively complete, articu- (Wellnhofer 1978, 1991a, Unwin et al. 2000), the lated pterosaur skeletons from the Santana and true diversity remains unclear, as do the relation- Crato Formations of Brazil (reviewed by Welln- ships of taxa within the assemblage to each other. hofer 1987, 1991a, 1991b, 1991c, Campos & Kell- Consequently, it is difficult to assess the signifi- ner 1985a, Kellner 1991, Martill & Frey 1998. cance of anatomical and functional studies based Kellner & Tomida 2000), often representing taxa on Cambridge Greensand pterosaurs and also similar to or synonymous with those from the difficult to set this pterosaur assemblage within a upper Lower and lower Upper Cretaceous of wider ecological or evolutionary context. England (Unwin 1988, Unwin et al. 2000, Fast- 192 lina.in. D. M.. Cretaceous pterosaur assemblarrc from England nacht in press). was most timely and helped to will provide detailed accounts of the taxonomy, resolve some key problems concernins the tax- anatomy and phylogenetic relationships of the onomy aiid systematic relationships of the Cam- Cambridge Greensand pterosaurs. bridge Greensand pterosaurs. It is my intention to publish the results of this uork in a series of Inslitutional abbreviations: BMNH. The Natural History MU- seuni. London. England: BSP. Bayerische Staatssammlung papers coveri 1g \'arious aspects the Cam- ffir Pal~cjntolc,~ieulld Gcologie. Munich, Germany: bridge Greens Ind pterosaurs including their pre- C'AMSM. Sedgivick Museum. Cambridge, England: GSM. servation. the history of research and the sv~- Geological Survey Museum. Kcyworth. England; MANCH, blanchester Museum. University of Manchester. Manchestcr. telnatics tk major groups. first k-'aper Engl;l11d: YORM. Ycjrkshire Museum. Yc>rk,England. presents important background material iriclud- Transliteration and taxonomic conventions: Transliteration ing the palaeoIltolosJ" of the Cam- of Russian and Mongolian names follows Bcnton el a[. (2000). When first quoted. junior synonyms are followed by bridge Greens the preservation and taphon- ihr scnior synonym cited in parentheses: a full listing of sy- @my of the ptcrosaur assemblage. a SUnllnar!- of non!m.; and valid tau is given in Table 1. Enclosure in sin- the taxonomic status of the Calnbi-idge GrecI1- gle quotation marks is used to indicate taxonomic namcs that nrr in\ aiid or of doubtful validity. Paraphyletic laxa are dis- sand taxa. anc a brief discussion of the general tinpui,,led hy marks, significance of the assemblage. Further papers
Trible 1 Checklist of \petit s names. in .ilplinhctical order.. hnsed on pterosaur material fr.oii1 the Cambridge Greensand of Cambridge- shire. England. Tlic oripinal tiuthor and the first occasion on \\hich the prlicular combination was cited is given in each case. Comments. \\ herc appropriate. 211-e gi1.t.n in the right hand column. Valid names arc shown in bold.
~~ ~ ~~ Species Status here
- Coloborhynchus sedgwickii = Coloborhynchus cupito = Lonchodectes plat-ystornus
= \'lllCl spccw
~ \ alld S~"c1e'
= \ ,Illd \pec~e\ = Anhungcreru cuvieri
~ \'Illci speclc5
~ Colohorhynchus cupito Omith och eirus sim us = Coloborhynchus sedgwickii
~ Ornijhocheirris sirnus = Lonchodectes platystornus - Coloborhynchus cupito = Ornithocheirus simus = Coloborhynchus sedgwickii
~ \alld 5pcc1es = \'llld species = \ dlld species = Lonchodecres microdon = \'llld species = Anhangrreru cuvieri Lorzchodectes comprestirostris
= Anhungueru cuvieri = Colohorhynchirs cupito -- iioiiien nuduni = Ornithocheirus sirnus - Anhaiigueru cuvieri
~ Coloborhynchus sedg w ickii
~ Anhungtiern cuvieri = Anhungueru cuvieri - 4nhunguera cuvieri = Anhungtiera cuvieri = Coloborhynchus cupito = 4nhunzguerufittoni = Lonchodectes inicrodon = Ltmchodectes muchaerorhynchus
~ nomen nuduni - Lonchodecres microdon Mitt. Mus. Nat.kd. Berl., Gcowiss. Reihe 4 (2001) 19.3
Species Status here
0rriithncIzcirit.s rmsiitLrs Seeley, 1 870a = Anhanguera fittoni 0rnithoclieirir.s oweizi Seeley, 1870a = Lonchodectes microdon 0rnithochcirri.s oxyrhinrrs Seeley. 1870a = nomen nudum 0rrzithocheirii.s plutyrhinris Seelcy, 1869a = nomen iiudum Orr irli odr eirr IS p luiy rh ini 1s See I e y, I 870a = Ornithocheirus sirnus 0mirhocheirci.splerijx fonius Seeley. 1870a = Lunrhodectes platystomus Ornithoclic~iriispolyodon Seeley, 1870a = Anhanguera fittoni Orniihocheiria rdiSeeley, 1870a = Coloborhynchus capito Ornithocheirus simus (Owen): Seeley, 1869a = valid species Omiihochc~irirssctrphor-liyrzchus Seeley, 1870a = Anhanguera cuvieri 0mithocheirri.s setlgwickii (Owen): Seeley, 1 870a = Coloborhynchus sedgwickii Ornithochcinis teniiivostris Seeley. 1 870a = Lonchodectes compressirostris Ornir/ioclreir~i.swoorlwardi (Owen): Seeley, 1870a = Coloborhynchus sedgwickii Orriirhoclieirii.s x-ypliorhynchus Seeley, 1870a = Anhanguera cuvieri
Ornithostoma sedgwicki Sceley. 189lb = valid species Ortiithostonin .seele.yi Lyde k k er, 1904 = Ornithostoma sedgwicki Prenoductylus hrachvrlziriiis Secley, 186Ya = nomen nuduni Ptenorincrylris capito Seeley, 1869a = nomen nudum PtenotluctjlLrs colorhinits Seeley. I869a = nomen nudum Pfenodtrctyliis crei.s.siden.s Seeley. 186% = nomen nuduni Premtkrctylus ci~vieri(Bowcrhaiik): Secley, 186Ya = Anhanguera cuvieri Ptenorluctyliis deniertiis Seeley. 186% = nomen nudum Pterzorlactylits enchorhynchris Seeley, 1869a = nomeii nuduin Pteriorltrctylit.~eiirygnathiis Seeley, 186% = nomen nudum Pretiotlac.ryh~s,fittorii (Owen): Seek y. 1869a = Anhanguera fittoni Ptenotierctyhis machaerorhynchits Seeley, 1869a = nomen nudum Premrluctylirs mercrorhinus Seeley, 186% = nomen nudum Pteriotluctylu, nricrodon Seeley. 186Ya = nomen nudum Ptenorluctyli~.~rzusirtus Seeley, 186% = nomen iiudum Ptcriotlerctyliis oweni Secley. 1869a = nomen nudum Ptenodrictylirs oxyrhiniis Secley, 186Ya = nomen nudum Pt~wocltrctylri.splaty.stomii.s Seeley, 1869a = noiiien nudum Ptcrzoclerctylus polyoelon Seeley. 1869a = nomen nudum Pteri odacty fils scapli orhyn ch I is SeeIc y. 1 869a = nomen nudum Prerzorlactyhis sedgwicki [sic] (Owen): Seeley. 1869a = Coloborhynchus sedgwickii Pic1~oducty1ir.stenriirostri,~ Sceley. 1869a = nomen nudum Ptenotlactj1ii.t n~ootlwnrdi(Owen): Seeley. 186Ya = Coloborhynchus sedgwickii
Pterotlac/jli4.s crrrreri Sceley. 1865b = nomen nudum Pteroderctyhr.~ciivieri Bowerbank: Sceley, 1 865b = Anhanguera cuvieri Ptc~rotlnctyliisfitroni Owen, 185917 = Anhanguera fittoni Ptrrodnctylirs hoykirisi Seeley, 1864b = nomen nudum Pteroritrciyiirs hirxleyi Seeky, 186Sb = nomen nudum Pterorlnctj'1ii.s machtrevorhychirs Seeley, 1864a = nomen iiudum Ptcwclcrctj,Iir.s oiverii Seclcy. 1864b = nomen nuduin Ptrrodrtctjltrs sedgwickii Owen, 1859b = Coloborhynchus sedgwickii Ptemlactylii.s .sirnus Owen. 1861 = Ornithocheirus simus Pterodrrct~~hiswoo(1warrli Owen, 1861 = Coloborhynchus sedgwickii
A brief history of the study saw intensive exploitation of the deposit from of Cambridge Greensand pterosaurs the 1850s to the 1870s, led to the recovery of many fossil remains including over 2000 ptero- The first pterosaur remains to be reported from saur bones. Most of the pterosaur bones were the Cambridge Greensand, three fragmentary acquired by the Sedgwick Museum (then the wing bones, were described by Richard Owen Woodwardian Museum), Cambridge, but impor- (1851b: pl. xxxii, figs 6-8). Although these tant collections were also built up in other Brit- bones are not now of any particular taxonomic ish museums including Brighton (Booth Mu- or anatomical significance they were important seum), Glasgow (Hunterian Museum), London in the mid-nineteenth century because they pro- (Natural History Museum), Manchester (Man- vided some of the earliest evidence for the exis- Chester Museum), Norwich (Castle Museum) and tence of large pterosaurs that reached wingspans York (Yorkshire Museum) and by the British of up to 4-5m. The great Cambridge Green- Geological Survey, now in Keyworth. Nottin- sand coprolite mining rush (Grove 1976), which ghamshire. Small collections also found their way I03 LJm\in. D M..Crctxeous pterosaur assemblage from England to local musc~umsin the U.K. and some large rj*rhiiiii.s'. Consequently, as Pterodactylus sinzus institutions oiitside thc U.K. including the Mu- Owen was the only available and valid species seum d'Histo re Naturellc in Paris. France. the assigned to the genus at the time of its publica- Bayerische St aatssamlung fur Palaontologie in tion, according to the International Commission Munich. Germany and the Peabody Museum. on Zoological Nomenclature (1999: Article 68.3), Yale. USA. it automatically became the type species by Initially. the pterosaur material ~vasstudied by nionotypy. Furthermore, Seeley later stated Richard Owen who published a series of papers ( 188 1 ) "The other genus, Ornithocheiriis, in- (Owen 1859a. 185%. 1860b. 1861) on more than cluded three species, and had for its type the 60 specimens loaned from the Sedgwick Mu- Prc.r.ontrc.r\.liis siriiiis of Owen . . .", an act that, seum. Owen described all the pterosaur bones according to the International Commission on under the xme Pterorfacr!-lirs. and identified Zoological Nomenclature (19%: Article 70.2), in- four new spc cies: Prei~otlucr!*lirs serlg\t'ickii (= validated subsequent attempts by Khozatskii and
Colohorli!>i 1 ch I is .serlg~c
(l876a) and later Lydekker (1888) correctly cvodon) Seeley, 1870, and Ornithochc’lrlis ’ 11tu- pointed out that ‘Criorhynchus’ was a junior sy- chaerorhyn ch 11s ( = Lonch odecres II i cr ch rr eror- nonym of Ovnithocheiuiis, but, unfortunately, the hynchrcs) Seeley, 1870 were assigned by Hooley former name was adopted by some subsequent to Lonchodectes which was characterised by workers, a practice that began with Zittel (1890), small, uniformly spaced teeth. Hooley also ar- and which has continued to the present day (e.g. gued, convincingly, that Secley’s 1901 concept of Kellner & Tomida 2000). Ornithocheiriis was incorrect in many respects. Throughout the late 1800s Seeley continued to such as the reconstruction of a large cranial describe aspects of pterosaur anatomy on the ba- crest, and that it was not Pteranodori but for the sis of Cambridge Greensand material (1876a, teeth, as Seeley and Williston had supposed. 1881, 189Ja, 1891b). Most importantly, during Hooley added to the taxonomic confusion. how- this period, he began using Ornithostornu as a ever, by retaining ‘Criorhynchiis’ and synonyniis- synonym of Ptercrnodon, an idea that was subse- ing Coloborhynchi~with the latter taxon be- quently adopted by Williston (e.g., 1895, 1896, cause he believed the differences could be 1897) who published numerous papers on the attributed to the extensive attrition which the American Niobrara pterosaurs, and who also ar- holotype of C. cluvirostris had undergone, an in- gued for a close similarity between Ornitho- terpretation which has not been supported by cheiriis and Pteranorlon. In his final and largest more recent studies (Lee 1994, Fastnacht in work on pterosaurs, “Dragons of the Air”, See- press). ley used Cambridge Greensand material to illus- The three-dimensional preservation of thc trate many aspects of pterosaur anatomy. This Cambridge Greensand bones proved far more work also contained the first reconstruction of suitable for reconstructing the functional mor- the skull of Ornithocheinis, which was, in effect, phology of pterosaurs than the heavily com- an outline of Prerunodon into which Seeley had pressed remains found at most other localities. shoehorned various skull fragments (often misi- Dennis (1 861) and Owen (1 861) both mentioned dentified or incorrectly oriented) from the Cam- anatomical features visible in the Cambridge bridge Greensand (Fig. 2). Greensand material that suggested to them that Hooley (1914) made a valiant attempt to sort pterosaurs were competent fliers, but the first de- out the tangled systematics of the Cambridge tailed biomechanical study was undertaken by Greensand pterosaurs. By treating the fossils as Hankin and Watson (1914), who made some im- isolated elements he took a more realistic posi- portant observations on the function of the tion than Seeley, but created some difficulties pterosaur forearm and wrist. This work was for himself by accepting all the species proposed further developed by Short (1914). Subsequently. by Owen and Seeley as valid taxa. The main Braniwell and Whitfield in their classic study of achievement of Hooley’s work was the recogni- Pterunodon (1 974) utilised Cambridge Green- tion of a distinct genus, Lonchodectes, that had sand material to gain insights into joint function. been noted, but left unnamed by Seeley (1901). while Frey & Riess (1981) also used Cambridge Various Cambridge Greensand taxa including Greensand specimens in their controversial re- Ornithocheirus microdon (= Lonchodectes rni- construction of the pterosaur fore limb, cspe- cially the position of the pteroid. After Hooley (1914), no first hand systematic studies of the Cambridge Greensand pterosaur material seem to have been attempted, and most workers in the pre-Second World War period adopted Hooley’s taxonomy to varying degrees. Arthaber is of special interest because he at- tempted to reconstruct the skulls of some of the / Cambridge Greensand taxa (Arthaber- 1922: figs 5-7). Unfortunately, these restorations, notably Fig. 2. ‘The skull of Omithocheina. as restored by Seeley the skull of ’Cuiorhynchus’siniiis (Fig. were (1901). The outline of the skull appears to be based on Pte- 31, rru~nrlouand, confusingly, the missing elements are shown as highly inaccurate, as Nopcsa (1924) and later shaded. Seeley identified a fragmentary cranium bearing the workers pointed out, and as has been demon- hose of ;1 large postcrodorsally-directed crest and incorpora- strated by the recent discovery of a complete ted it into this restoration. but never directly referrcd to the spccimen in print. Later. Hooley (1914) assigned this speci- skull for the closely related form ‘Troyrogrinrliir.5’ men (CAMSM BS4.406) to Orrii/hostonza. mesernbrinuh (Welln hofer 1987). 196 I!nwin. D. M.. Cretaceous ptcrosaur assemblage from England
invalid taxa. Many authors also continued to re- cognise 'Criorhyrzchus' as a distinct taxon and even assigned it to its own family, the Crior- hynchidae. while Ornithocheiriis continued to act as a refuge for numerous poorly known Cretac- eous pterosaurs. 'Omithocheirus curtus', which is based on a fragmentary distal end of a wing-me- tacarpal (BMNH R1440) from the Wealden of Sussex (described by Owen (1870, pl. xix, figs 8-9) as the distal end of a tibia), is a typical example: this species is undiagnosable and the holotype cannot even be assigned to any particu- lar family (Unwin 1991). Other taxa, however (such as Loncfiotiectes and Coloborhynchus), are During the ast SO years. with the exception of based on distinct, diagnostic material and clearly the above mcritioned functional studies. most ac- cannot be accommodated within the same genus. counts and conments on the Cambridge Green- Kuhn (1967) and Wellnhofer (1978. 19Yla) sand pterosaurs have been based on the early gave the most detailed systematic accounts of literature (priricipally Hooky 1914). rather than the Cambridge Greensand pterosaurs (Table 2) on direct examination of the fod material. Sev- and these have formed the standard systematic eral gencral trends can be discerned. Most arrangement followed by most other authors authors (e.g.. Kuhn 1967, Wellnhofer 1978. over the last 20 years. Description of much more 1991a) recognised that some. perhaps many ot complete and well preserved remains from the the Cambridt e Greensand ptcrosaur species Santana Formation of Brazil (Wellnhofer 1985, were of doubtful validity. but. understandably. 1991a. 1991b. 1991~.Kellner 1984, Campos & were unable to discriminate between valid and Kellner l985b. Kellner & Tomida 2000) has
Table 2 Summar! of systematic assessment\ hy tiuhn ( 1967) and Wrllnhofer ( 1978. 1991a) of the Cretaceous pterosaur assemblage from ttic Cambrid :e Greensand. En@nd.
Kuhn 1967 Wellnhoi'er 1078 Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 4 (2001) 197
thrown some much needed light on the anatomy, posed by phosphate quarrying during the nine- taxonomy and relationships of the Cambridge teenth century, but in many cases these expo- Greensand pterosaurs. A critical find was the sures were reclaimed for agricultural use (Grove skull of ‘ Tropeognuthus’ mesernbrinus (Wellnho- 1976). Following the cessation of mining activity fer l987), which demonstrated that the holotype in the late 1800s the number of exposures stea- of Ornithocheirris simiis Seeley, 1869a consists of dily declined and they are now rare (Worssam & just the anterior tip of an upper jaw bearing a Taylor 1969, Norman & Fraser 1991). large crest. Other material from the Santana For- The Cambridge Greensand lies unconformably mation, such as the crania assigned to various on the Gault (Reed 1897; Fig. 5). It passes up- species of Anhangueru (Campos & Kellner 1985b, wards into what has traditionally been called the Wellnhofer 1991b, Kellner & Tomida 2000), is Chalk Marl, but is now referred to as the Porcel- strikingly similar to Cambridge Greensand speci- laneous beds (Morter and Wood 1983) of the mens, but the exact relationships of the taxa Chalk Formation. Lithologically, the Cambridge based on these fossils have yet to be clarified. Greensand has always been considered part of the Lower Chalk (White 1932, Worssam & Tay- lor 1969) though much of its larger clast compo- Geology and palaeontology nent is thought to have been derived from the of the Cambridge Greensand underlying Gault (Reed 1897). Distribution and stratigraphic rela- S e d i m e n t o 1o g y : The Cambridge Greensand ti on s h i p s : The Cambridge Greensand, a rema- is a thin unit of micaceous, glauconitic, silty marl. nie deposit at the base of the Chalk, crops out in with a basal lag of reworked phosphatic nodules, a narrow, 80 km long swathe across East Anglia remanie fossils and exotic clasts, often encrusted (Fig. 4). It is best developed in the region of in small oysters (Sedgwick 1846, Seeley 1866b, Cambridge, from which it extends as far north as Bonney 1873, Jukes-Browne 1875, Reed 1897. Soham in Cambridgeshire, and as far south as White 1932). The contact with the underlying Harlington in Bedfordshire (Reed 1897). Large Gault is an uneven and intensely burrowed ero- tracts of the Cambridge Greensand were ex- sion surface (Jukes-Browne 1875, Hart 1973:
Key
Alluvium a Chalk Cambridge Greensand
0Gault
Lower Greensand
Kimmeridge Clay
Corallian
Oxford Clay
0 Barrington @ Cambridge 0 Soham
I 5km 1
Fig. 4. A. Simplified geological map of eastern England showing the distribution of the Cambridge Greensand. B. Location of main map. C. Cambridge Greensand workings in the vicinity of Cambridge that have yielded pterosaurs Adapted from Grove (1976) and Worssam and Taylor (1969). 1 us [In\\ in. D M.. C'retnceous pterosaur assemblagc froin England
Fig. 5). The Cambridge Greensand reaches a Suberbiola & Barrett 1999 for historical re- thickness of a >out 0.6 m Lvithin erosional troughs \+ass). In fact. it consists of not one, but threc (though up to 1.5 ni has been pi-o\.ed in bore- interrelated problems: the age of the Cambridge holes (Worssam & Taylor 1969)). and thins to al- Greensand: the age of the remaniC fauna; and most nothing over topographic highs of the ero- the length of time represented by the non-se- sive surface (Reed 1597). The basal lag. largely yuencc. The deposition of the bed itself is gener- composed of ihosphate nodules. is concentrated ally thought to have occurred very early in the in depression:,. and while occasionall!. reaching Cenomanian. though until quite recently the up to 0.25 m is more usually about 0.15 m in onlv evidence for this was stratigraphic (Casey in thickness (White 1932). Ednionds & Dinliarn 1965). Cookson and The matrix consists of a fine chalk mud. Hughes (1963) made a case for an early Cemo- mainly composed of coccoliths ( Bonne\. 1573. manian age on the basis of three specimens of Sollas 1876). thouyh in some places it contains the ammonite Schloeizhachin varirrns, but their large yuantitit,s of Gault Clay (Reed 1897). The e1,idence was rejected by later authors (e.g., $auconitic gr,iins. w%ich 2ii.e the unit ;i p~enCasey in Edmonds & Dinham 1965, Morter & tinge. seem to be derived from the Gault and Wood 1983). Hart (1973), using what seems to prove in most cases to be internal moulds of for- he an autochthonous fauna of foraminifera, was aminifera (Soi las 1 873. 1876). They are concen- more successful. showing that the matrix is trated at the lme of the deposit and rapidly de- equivalent to the uppcr part of the Neostlingo- crease in size and abundance upw~rds(White. c'ern.s cfircitmeiiw ammonite assemblage subzone, 1932). Lvhich is earliest Cenomanian in age (Fig. 5). The The phosph ite nodules have a similar distrihu- most recent study (Morter & Wood 1983) ac- tion to the gl.iuconitic grains. They vary in col- cepted the likelihood of a basal Cenomanian our from olive black through brown to pale buff age. but argued that a very late Albian age could (Fisher. 1873) and are usually irregularly shaped. not be ruled out. This idea was first proposed by though roundd and tubular forms also occur Spath (1923-43) who believed that the bed was (Seeley l866b). They were originally misidenti- deposited in late Stoliczkain dispur zone times fied as coprol tes. which gave rise to one of the (= Mortoiiiccvns peririjlritrrm subzone) and is Cambridge GIe c n saii d 's ma 111- ;i 1t e 1-11at i \,c names. supported by Cooper and Kennedy (1977), who the Coprolite Bed. In fact. true coprolites. usual- noted that derived ammonites froni the upper I!, I!, of fish, arc rare (Seele!. 1Sh6b). Thin sections part of the S. tlispcir zone are lacking. reveal that mi ny of the nodules arc the remains It is widely accepted that the renianik fossils of phosphatis-.d sponges (Fisher 187-3. Sollas are late Albian and not Cenomanian in age 1873). though internal moulds of molluscs and (Spath 1923-43. Cookson & Hughes 1964. phosphatised calcareous mud are also common. Worssani 24 Taylor 1969. Morter & Wood 1983), Numerous exotic clasts exhibiting a \f,ide but there is some debate as to the duration of range of lithologies and reachiny up to 60 kg in the non-sequence. Both Spath (1923-43) and \veight. were clescribcd by early workers (Scclcy Owen (1979) reported ammonites from as low as 1 S66b. Bonney 1873. Sollas & Jukes-BroLvne the basal part of the Callihoplites clziritiis sub- 1873). rhough it n~islater sho\vn that not all zone. though Morter and Wood (1983) suggested came from the Cambridge Greensand (Reed that these remains might not be from the Cam- 1897. Havkes 194.3). These exotic clasts \\.ere bridge Greensand. A nuniber of authors have probably rafte'l to thcir final rcstiny place in tree proposed that downcutting only reached the top roots (Hawkes 1943) rather than on shore ice or of the C. mritil.s subzone (Fig. 5) and thus the icebergs as eirlier authors contended (Seeley nonscquence represents the entire S. dispar zone 1866b. Bonne! 187.3. Sollas & Jukes-BroLvne (Spath 1923-43. Hart 1973, Morter & Wood 1873). althouigh some might possibly be the 1983). Others suggest that downcutting pro- dragged anchc'rs of seaweeds (White 1933). The cecded only as far as the top of the A. siihstrtderi many rolled. Lvater Ivorn. phosphatised fossils subzone (= top of M. rostraritm subzone in that make up ihe rcmainder of the basal lag are Fig. 5) and thus only the vcry latest Albian, discussed be1o.v. equivalent to the M. perin,flatiim subzone, is niissing (Breistroffer 1940, Casey in Edmonds & Agc ot the Cambridge Greensand: The Dinhani 1965. Carter & Hart 1977). In summary, age of the Cainbridge Greensand has long been the pterosaur material can be fairly confidently problematic (4 :e White 1932. Hart 1973. Pcreda- dated as late Albian because there is no evi- Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 4 (2001) I 00
I KEY Chalk El
Siltstone with
u-1
Aucellina-rich mudstone
Inoceramid-rich mudstone
Phosphate nodules and pebbles .'.a0014...D.q Disconformities-
Belemnite floods I flv
I\' \/
4p
Fig. 5. Stratigraphic relationships of the Cambridge Grccnsand Lo other Cretaccous deposits in the region of Cambridge. Eng- land. Adapted from Morter and Wood (1983). Beds numbered according to Gallois & Marter (1982). 700 I!n\\.in. D. M.. Cretaceous pterosaur assemblage from England dence of a Ccnomanian macrofauna in the Cam- & Jukes-Browne 1881, Reed 1897) distinguished bridge Greensand and the carliest derived re- by differences in colour, texture and preserva- mains are no older than the C. rriiririi.7 subzone tion. These differences may, however, be attribu- in age. table to highly localised variations in preserva- tioiial conditions, such as the degree to which Deposition a 1 history an d pa 1 a e o g e o - remains were initially buried. Seeley (1876~) graph y : The Cambridge Greensand was depos- mentioned associated remains in which indivi- ited in the centre of the Anglian trough. dual elements exhibited considerable variation in bounded to the north by thc North Norfolk swell preservation. Hart (1 973) also suggested that dif- and to the south by the Berkshirc-North Kent ferences in nodule colour were of regional and swell (Carter & Hart 1977: fig. 6). all of which not temporal significance. This hypothesis is sup- form part of the Anglo-Paris basin. During the ported by the results of this study in that no cor- late Albian tk ere was a hiatus in sedimentation relation was found between variation in the pre- associated wit i uplift following mild earth niove- servation of remains and either their size or ments at the 2nd of the mid-Albian (Lott et al. taxonomic distribution. 1980). Curren winnowing led to the exhumation Early workers published voluminous lists of of fossil remains. exotic clasts and phosphate no- taxa (Seeley 1869a, Bonney 1873, Jukes-Browne dules that hacl been formed in the Gault Clay. 1875. Penning & Jukes-Browne 1881, Woods All three wcrc rolled. concentrated and became 1891. Reed 1897), that were subsequently added the basal la!: of the Cambridge Greensand to by White (1932) and Chatwin (1948). Regard- (Reed 1897. 'White 1932). The absence of evi- ing niicrofossils. Sobs (1872, 1876), Chapman dence for adult oysters encrusting clasts of the (1899) and Hart ( 1973) described the foramini- basal lag (Hart 1973) suggests relatively swift in- fera and Cookson and Hughes (1964) the dino- undation by srsdiment. indicating that the Green- flagellates and acritarchs. The only evidence of sand itself was deposited quite rapidly. The se- macroflora consists of dark coloured amber quence of events leading to the formation of the (Seeley 1866b, Reed 1897). Cambridge G -eensand seems to have been lar- The rich invertebrate fauna is dominated by gely controlled by variation in sediment supply cephalopods, consisting mainly of ammonites rather than ha rdpart input. corresponding closely (Spath 1923-43, Cooper 81 Kennedy 1977), to theoretical models such as Kidwell's (1986) some belemnites and a few nautiloids. Bivalves "Typc IV shell bed". which are based on this and brachiopods are common, while gastropods idea. are diverse, but not numerically abundant (Reed The Cambridge Greensand was deposited in a 1897). Echinoderms are scarce and, not surpris- shallow epeiric sea environment. to the north ingly. fragmentary. Other rare groups include and west of vrhich lay an archipelago (Bennison scaphopods, actinozoans and annelids. The cara- & Wright l97;3: fig. 14.4. Lott et al. 1980: fig. 15. paces of crabs and lobsters, some with evidence see below). 11 was from thcse islands that the of parasites (Bonney 1873), are not infrequent exotic clasts were presumably rafted arid upon and numerous phosphatised sponges have also which lived the various terrestrial vertebrates been described (Sollas 1873, 1876). subsequently recovered from the Cambridge This horizon has also yielded numerous, disas- Greensand. The shallow. Anglian trough sea pro- sociated vertebrate remains. Most, though usual- vided various habitats and feeding opportunities ly uncrushed, are poorly preserved, but on occa- for fish. marire reptiles. diving birds and ptcro- sion fine detail is exhibited. Fish make up most saurs. of the diversity with over SO reported species. Pa 1 aeon t o 1 o g y : In terms of both taxic diver- Sharks. represented by numerous teeth and sity and numtw-s of individuals the Cambridge spines. are common and include the ctenacanthi- Greensand is one of the single richest fossil ver- form Acrocliis and Hyhocliis, and the galeo- tebrate horizons in the British Isles. This is niorph s Hc.rat 1 ch z is, Lniiinn, Scupnouhy rich us, partly because the preservation potential was im- Sqii(i1icoin.x- and Synechotiiis. Chimaeras, such as proved by phosphatisation. and partly because of Ednpliocioi~ and Ischyodzrs, are also common. the intense cxploitation of the Cambridge Woodward (1893) listed a number of pycnodonts Greensand. which led to the accumulation of (Arlitndoti. Goelocliis, Aiionzaeodzis and Pycno- large collections (Jones in Darby 1938). The fos- tlirs). represented by well preserved tooth plates, sil remains ha\ e often bcen di\.ided into indigen- and also reported on scales of the semionotiform ous and deriv-d faunas (Bonney 1873. Penning Lcyitlorrs (Woodward 1895). Other fish include Mitt. Mus. Nat.kd. Bed.. Geowiss. Reihe 4 (2001) 20 I the pachycormid Protosphyraenn and teleosts mon (Seeley 1869a, 1879), though very fragmcn- such as the ichthyodectiforms Plethodus and tary. Six genera were recognised by Seeley Sartrocephalus, and the aulopiform Enchodus. (1879), but recent systematic reviews (Coonibs & Seven genera of turtles have been recognised. Maryaiiska 1990, Mclntosh 1990, Norman & Seeley (1 869a) proposed new species of Emys, Weishampel 1990, Pereda-Suberbiola & Barrett Trachydermochelys and Testudo, but all are no- 1999) indicate that most are nomina dubia. Saur- mina nuda and the remains on which they were opods are represented by the titanosaurian ‘Mcr- based probably belong to Rhinochelys (Lydekker criirosaurus semnus’ (Seeley 1869a. 1876d). con- 1904). Lydekker (1889a) erected new species of sidered by Le Loeuff (1993) to be a nomen Chelone, Protostegn and Lytolonm on the basis dubium, and based on two sets of caudal vertcb- of fragmentary mandibles, but it is doubtful rae that, though discovered some miles apart, whether any are valid (Buffetaut et al. 1981). were believed by Seeley (1876d) to have come The single remaining genus, Rhinochelys Seeley, from the same individual. An associated metatar- a small to medium sized chelonioid, is repre- sus (Seeley 1876d) might also belong to ‘Mci- sented by numerous well preserved skulls (See- crurosaurus’ (Mclntosh 1990), but this material ley 186%). Many species were established by has yet to be studied in detail. A nodosaurid an- Seeley (1869a) and Lydekker (1889a), but a sys- kylosaur, Anoplosaurus curtonotus Seeley. 1879. tematic revision by Collins (1970) indicates that represented by a partial vertebral column and only three are valid. The abundant postcranial appendicular elements including the shoulder remains of chelonians listed by Seeley (1869a) girdle and fore and hind limb bones, was re- are also probably referable to this single genus. cently redescribed by Pereda-Suberbiola & Bar- Seeley (1887) identified the proximal end of a rett (1999). These authors also identified other femur and a first sacral as belonging to a lizard, fragmentary remains of ankylosaurs, many of which he named ‘Patricosaiirus merocratus’. which are probably referable to nodosaurids. in- Though listed by Romer (1966), this dubious tax- cluding a partial foot and dermal plates. Other on was ignored by Estes (1983) in his compre- fragmentary bones appear to represent an inde- hensive review of lizards. Seeley (1869a, 1874) terminate iguanodontian (Norman & Weisham- erected two new species of ‘Crocodilus’ on the pel 1990). A tooth, first mentioned by Seeley basis of four procoelus vertebrae. Lydekker (1879) and later described by Lydekker (1888) (1 888) doubted they were correctly assigned to under the name of ‘Trachodon’ cantabridgiemis. genus and Buffetaut (in Buffetaut et al. 1981) is supposedly one of the earliest records for the noted that the material could only be deter- Hadrosauridae (Benton & Spencer 1995. Kirk- mined as Eusuchia indet. land 1998), although Head (1998) has recently Sauropterygians, including both plesiosauroids questioned this identification. and pliosauroids, and represented by numerous A single genus of bird, Enaliornis, represented teeth, vertebrae and a few limb bones (Owen by skull fragments, vertebrae and hind limb ele- 1851b, 1861, Seeley 1869a, 1876c), are relatively ments was first described by Seeley (1876b) and common. The various taxa proposed by Seeley the braincase material has recently been rede- (1869a, 1876c) and Lydekker (1889b) were lar- scribed by Elzanowski & Galton (1991) and Wit- gely dismissed by Welles (1962) and Persson mer (1990). Enaliornis appears to have been a (1963), but the material on which they were specialised, foot-propelled diver (Elzanowski 8: based would benefit from further study. The Galton 1991) and has often been allied with he- Cambridge Greensand also yielded many iso- sperornithids (e.g., Martin 1984), although, as latcd teeth, fragmentary rostra, vertebrae and shown by Elzanowski & Galton (1991), evidence some limb bones of ichthyosaurs. Three genera in support of this idea is not as strong as pre- were recognised by Seeley (1869a, 1873) and Ly- viously thought. dekker (1889b), but all the ichthyosaur material Pterosaurs, the most common tetrapods in the was subsequently assigned to Platypterygius by Cambridge Greensand, are discussed below. McGowan (1972). Bardet (1992) has cast doubt on this assignment, however, and has also ar- gued that all the species of ichthyosaur based on Taphonomy of the pterosaur remains Cambridge Greensand material are nomina du- bia. P r e s e r v a ti o n : Unlike pterosaur bones from Dinosaur remains, largely consisting of verteb- many other horizons, the Cambridge Greensand rae, limb bones and a few teeth, are quite com- remains are uncrushed, but, in the vast majority 202 Uiiwin. D. M.. Cretaceous pterosaur assemblage from England
Fig. 6. Asprctr 01 the pcsei-\,ation of ptcrosrur hones Irom the Cambridgc Greensand (Cretaceous) of England. A. Almosl perfectly prcwi-ve .I right proximal s!.ncai-pal (BMNH K229.7) in proximal \,ie\v. B. Fragment of right mandible (CAMSM B51.915) of an (-rnithocheirid in Iatcral \ie\\ \\ith a sniall phosphate overgrowth (ari-owed). C. Fragmentary rostrum (CAMShI Bi4.62.; ) of C'olohor./i\,ric111,( itrpito in right lateral \,ic\v \vith irregular pitting (nrrowed) produced by ?osteopeltid gastropod\. D. Fiagrientar! $enoid region of an ornithoclieirid scapulocoracoid (BMNH 35226) bearing an cxtensive dendri- tic praring pattei-n (arro\\ed) produced h! '!ostropeltid gastrqods. E. Distal end of ii left ulna (BMNH 35324) of an ornitho- chcirid in anterior \.ieu hearing attachment sites (arro\\-ed)of small oystcrs. Scale bar = 10 mm. Mitt. Mus. Nat.kd. Rerl.. Gcowiss. Reihc 4 (2001) 20.1
of cases, they are rolled, broken and abraded. from an ornithocheirid while the distal end is With only two or three exceptions, all long from a lonchodectid and in the second. the prox- bones are incomplete, though blocky and tabular imal end is from a right humerus and the distal elements such as vertebrae and carpals seem to end from a left. have fared better and one or two almost perfect Of greater significance. frotn a systematic examples are known (Fig. 6A). Most remains point of view, is a list of 33 supposedly asw- have suffered considcrable mechanical damage, ciated sets of remains, published by Seeley in the projecting processes are almost always broken “Index to the Fossil Remains” (1869a: 8-18), off and delicate features are usually, though not These sets were purchased from the Farren always, destroyed. Teeth are either missing from brothers by the Sedgwick Muscum, Cambridge their sockets or snapped off at the base. and, if genuine, as Seeley clearly believed they The remains vary in colour from buff to al- were, would be of considerable importance in cq- most black depending on the degree of minerali- tablishing the basic characters of Cambridge sation; the darkest tend to be the most heavily Greensand pterosaurs. phosphatised. The great ma,jority of specimens There is some circumstantial evidence that as- are entirely pervaded by phosphate and occa- sociated remains were occasionally found. Secley sionally even overgrown by phosphate (Fig. 6B). reports seeing associated remains of plesiosaur Phosphatisation probably played an important vertebrae (1879: 592) and hc also listed and de- role in increasing the resistance of remains to scribed associated remains of dinosaurs and mcchanical damage, but it should be noted that ichthyosaurs (186921. 1876~1879). Moreover. unphosphatised remains are also uncrushed. Pereda-Suberbiola & Barrett (1999) have re- Many remains exhibit evidence of attack from cently described an incomplete, but apparently bone feeders, possibly osteopeltid gastropods, associated skeleton of AIioplosLiiiriiJ and an a+ which rasp and bore away at exposed bone sur- sociated ankylosaur foot. It is possible that these faces (Marshal 1987). Three different types of relatively heavy elements resisted niovemeiit by feeding ichnitc have been identified. Irregular currents and remained in assocation. but this is pits up to 7.5 mm in diameter, for example on unlikely to be true for pterosaur bones. which CAMSM BS4.625 (Fig. 6C), and dendritic graz- were much lighter. Moreover, the pterosaur re- ing patterns, well exhibited by BMNH 35226 mains exhibit considerable evidence of mechani- (Fig. 6d) and CAMSM BS4.507 are common, cal damage (see above), which almost certainly particularly the latter, which occasionally covers could not have taken place without leadins to entire specimens. A third and much rarer type of disassociation of the skeleton. Seeley (1 879) ar- ichnite, exhibited by YORM 19831220, consists of gued that the damage resulted from maceration small circular pits 1-2 mm in diameter. Virtually of the bones as they lay on the sea bed. but this all specimens were originally encrusted by small fails to explain the broken teeth and other me- oysters, as numerous occurrences of their calcite chanical damage. attachment plates show (Fig. 6C, E). Most are Study of the components of the “associated fairly small, up to 10mm in diameter, but larger sets” removes any last doubt that they might be ones of 20 mm or more are occasionally found genuine. For example, three proximal ends of (e.g., CAMSM B54.415). Serpulid worms also humeri are listed in set 4 (Seeley 186%: 9). appcar to have attached themselves to some of More importantly, ten sets of bones (12-15. 18. the bones (c.g.. CAMSM BS4.507). 25-27 and 32-33) contain both large. short cer- A few specimens have been “repaired”, but vicals and vertebrae which were thought by See- study of the components reveals that in most ley (1869a, 1870) to be caudals, but that are now cases they are composites, made up of remains known to be cervicals of a small. long-necked from different individuals. For example, the base pterosaur (Padian 1984, 1986, Howse 1986). As- of the tooth borne by the holotype of Omithho- sociated skeletons have, as a rule. always com- cheiriis sirniis (CAMSM B54.428, Owen 1861: manded greater monetary value than singlc re- pl. i, figs 1-5) docs not correspond in shape to mains. It appears, therefore, that thc Farrens. the tooth socket, nor does its fracture surface who were probably well aware of this, put to- match the fracture surface of the tooth preserved gether “associated sets” which they sold for lar- in the socket; this specimen is almost certainly a ger sums than they would have received for in- composite. Furthermore, two apparently com- dividual remains, and that Seeley uncritically plete humeri, GSM 87870 and BMNH 34413, are accepted the purported associations of the composites. In the first case the proximal end is bones. 204 Unu in. D. M.. Cretaceous pterosaur assemblage from England
Taphonomi: history of the pterosaur purchasing policies are likely to have further re mains : Th,: preservation of the pterosaur rc- tnodified the original pterosaur samplc. mains bears witness to a complex taphonomic C o n c 1 us i o n : This brief account of preserva- history of whi:h only a brief account is possible tion and taphonomy can be used to draw a con- here. Followin,: death. vertebrate carcasses usual- clusion that is of special significance for investi- ly sink fairly rapidly through the water column gating the taxonomy and systematics of to the sea bed. Pterosaurs were relatively light. Cambridge Greensand pterosaurs. There is no however. and with the additional buoyancy pro- evidence to show that any of the pterosaur vided by the pneumatic system could probably bones were preserved in association. Moreover, float for some considerable time. Schafer (1962) even if there were one or two such cases, which showed that dead seabirds can float for up to 40 seems unlikely given the probablc taphonomic days or more. during which time they may drift history outlined above, they can no longer be considerable c istances. Assuming that pterosaur demonstrated. Consequently, the associations in- carcasses wen: capable of drifting for similar ferred by Owen (1859a, 1859b, 1861) and listed lengths of tin12 it is possible that some compo- by Seeley (1869a) must be ignored and each ele- nents of the Cambridge Greensand pterosaur ment must be treated as if it were an isolated fauna may have been derived from regions re- find. mote from whcre they finally came to rest. Remains arrived at the sea floor either as whole carcasses, or as parts of a carcass. A vari- Summary review of the taxonomy ety of processes occured at this stage. Soft tissue of Cambridge Greensand pterosaurs decayed awa) and the remaining skeletons. either associa .ed or perhaps already disasso- Thirtv-two species of pterosaur have been ciated to sore extent, were grazed upon by named in connection with the Cambridge Green- bone feeders. They also served as benthic islands sand (Table 1). Most of these species are based in the soft soupy mud, providing attachment on jaw remains (essentially the anterior ends of sites for bivahes and serpulid worms. It is prob- rostra and mandibular symphyses), but, in some able that in many cases remains became partially cases. the name was not attached to any particu- or completely buried in the Gault Clay. only to lar specimen and is thus a nomen nudum. Com- be exhumed by current winnowing or storm ac- parison of the jaw remains with each other and tivity. a cycle that may have been repeated a with other pterosaur material, principally from number of times and almost certainly led to the Wealden, Gault Clay and Lower Chalk of further damage and disassociation of the re- England and the Santana Formation of Brazil in- mains. Phosph;ttisation probably took place with- dicates that ten of the 32 named species are va- in the sediment. with the remains acting as preci- lid (Table 1) and there is possibly one additional, pitation nuclei, The origin of the phosphate is as yet unnamed, species of Ornithocheirrss. These not known, bL.t is most likely to have been or- 11 species are distributed among three families: ganic (Bonney 1873. Reed 1897. Norman & Fra- the Ornithocheiridae, the Lonchodectidae and ser 1991). Inirrediately prior to the deposition of the ?Ptcranodontidae. A brief systematic review the Cambridge Greensand. the pterosaur bones. of these families and their representatives from together with other vertebrate and invertebrate the Cambridge Greensand is presented here. remains. phosLJhate nodules and exotic clasts seem to have been concentrated and reworked Ornithocheiridae: More than 90% of the by storm eveits. which also scoured the sea identifiable pterosaur bones from the Cambridge floor. Any coniplete or articulated skeletons that Greensand can be assigned to the Ornithocheiri- had survived t iis far must have been broken up dae. an important family of medium to large- and disarticuLited at this stage. Final burial sized, piscivorous Cretaceous pterosaurs (Welln- seems to have taken place quite rapidly (Hart hofer 1991a, Bakhurina & Unwin 1995, Unwin 1973) followins resumption of sediment supply & Lu 1997. Unwin et al. 2000). The Ornitho- in thc early Cenomanian. cheiridae are diagnosed by the relativc propor- While not strictly part of the taphonomic pro- tions of the teeth in the anterior part of the den- cess it is worth noting that, in many cases. speci- tition (Unwin 1991, Bakhurina & Unwin 1995). mens were fu.ther broken up by the washing The first three teeth are relatively large, forming processes employed during the extraction of a terminal rosette, and show a marked increase phosphate. In addition. collecting practices and in siLe posteriorly. The fourth tooth pair is much Mitt. Mus. Nat.kd. Berl., Geowiss. Rcihe 4 (2001) 20.5
reduced in size and smaller than the first pair of teeth, when observed in lateral or anterior view, teeth. Proceeding posteriorly, there is a steady are perpendicular, or near perpendicular, to the increase in tooth size up to, typically, the ninth long axis of the jaw (Owen 1861: pl. i, figs 1-4. pair, which are of similar basal dimensions to the Wellnhofer 1987: fig. 2; Fig. 7). This is unlike largest teeth in the terminal rosette. Further pos- other ornithocheirids wherein the first three to teriorly, tooth size declines again. Consequently, four pairs of teeth are directed forward and out- in dorsal view, the rostrum has an expanded ward from the jaw, the degree of anterior and anterior tip that accommodates the large ante- lateral flare decreasing posteriorly (e.g., Welln- rior teeth, is narrowest in the region of the hofer 1991b, Bakhurina & Unwin 199.5: fig. 12. fourth or fifth tooth pair, and gradually widens Kellner & Tomida 2000: figs 10, 11). The crest posteriorly. The expansion of the anterior end of (crestless individuals are not known) is located the rostrum is most marked in large species and at the anteriormost end of the rostrum (Owen adult individuals, but may be practically absent 1861: pl. i, fig. 1, Wellnhofer 1987: fig. 2: in small species and juveniles. The mandibular Fig. 7A-C) and has a relatively flat anterior sur- dentition and symphysis show similar morpholo- face that rises vertically from the tip of the jaw gical patterns to the rostrum. (Fig. 7A), then curves backwards forming a high. At present, the Ornithocheiridae comprises thick, crescentic structure. The only other or- Ornithocheirus and at least two additional gen- nithocheirid with a crest located at the jaw tip is era: Coloborhynchus and Anhanguera (Bakhuri- Coloborhynchus (see below). In this case, how- na & Unwin 199.5, Unwin & Bakhurina 2000, ever, the anterodorsal surface of the crest is con- Frey & Martill 1994, Unwin et al. 2000). Other cave, not convex, when viewed laterally. taxa from the Santana Formation (Brasileodac- In the Cambridge Greensand Ornithocheirus is tylus, Araripesaurus and Santanadactylus) and represented by the type species, 0. simus, known the Crato Limestone Formation (Arthurdactylus) from fragments of 13 rostra and 5 mandibular of Brazil also appear to belong within the Or- symphyses, and possibly by a second, as yet un- nithocheiridae (Kellner 1984, Wellnhofer 1985, named species, represented by a single specimen 1991a, Unwin et al. 2000), but their taxonomic (CAMSM B54.890). The holotype of 0. sinzus status requires clarification because they are in (CAMSM B54.428, Owen, 1861; Fig. 7) repre- at least some (and quite possibly all) cases sy- sents a large pterosaur that, by comparison with nonymous with Anhanguera. the more complete skull of ‘Tropeognathus’ me- The Anhangueridae (Campos & Kellner sembrinus (Wellnhofer 1987) and other ornitho- 398%) were diagnosed in a similar way to the cheirid remains, is likely to have had a wingspan Ornithocheiridae and have almost the same in excess of 4m. Most other jaw remains of 0. taxonomic content (e.g., Kellner & Tomida sirnus are of a similar size to the holotype, 2000). Evidently, the Ornithocheiridae and the although one or two specimens (e.g., MANCH Anhangueridae are synonymous, so the senior L10832) represent somewhat larger individuals. name, Ornithocheiridae, is adopted here. Chronologically, the first genus of ornitho- In the Cambridge Greensand the Ornitho- cheirid to be described from the Cambridge cheiridae is represented by three genera: Or- Greensand was Coloborhynchus (represented by nithocheirus, Anhanguera and Coloborhynchus. Pterodactylus sedgwickii Owen, 1859a). The At present, ornithocheirid postcranial material genus name has a somewhat complex history. It cannot be assigned to any of these three genera, was first proposed by Owen (1874) for a new because, although there is some variation in the pterosaur, Coloborhynchus clavirostris, based on morphology of particular postcranial elements a highly distinctive anterior end of a rostrum such as the humerus and ulna (Hooley 1914), it from the Wealden of Hastings, Sussex (BMNH is not clear whether this variation is related to R1822). Hooley (1914) misinterpreted the jaw particular genera and, if so, how it is related. fragment, as Lee (1994) has pointed out, arguing The discovery and description of more complete that it had suffered considerable abrasion, and remains of ornithocheirids from the Santana and that originally it had the same morphology as Crato Formations of Brazil may eventually the holotype of Ornithocheirus simiis, wherein enable this problem to be solved. the teeth projected vertically from the palatal Ornithocheirus is one of the most common surface of the rostrum (Hooley 1914: pl. xxii, taxa in the Cambridge Greensand, and is distin- fig. 5). Apart from some enlargement of the den- guished from other ornithocheirids by characters tal alveoli, the rostrum is not abraded, however of the dentition. Notably, the first four pairs of (Lee 1994), and the unusual morphology where- 206 L'nniii. D hl.. Cretaceous ptero\aur awmblagc lrom England
in the palatal surface is reflecled uplvard to form 11701~0~~~~~~12sis.This fragment is virtually identical a blunt triangi dai- surface terminatins the antc- to the holotype of C. worlleighi Lee 1994, conse- rior tip of the rostrum. through which projects quently 'Sirocc,optc.r.),.r'is treated here as a junior the first pair of teeth. is original and unique to synonym of Colohorli~nc.liirs.Whether Colobor- this pterosaur Lee 1994. Fastnacht in press). Co- l?!,/?c/ii/.sriioroccwisis can be distinguished from lohor-/?!~/?c/z us' i 5 a 1 so distinguished from 0 n i if/?o - other species of Colohorhyizc.hri,s, including the c/ieiri~~by thr: orientation of the second and almost coeval taxa from the Cambridge Green- third pairs of 1 eeth in the rostrum which are di- sand. has yet to be clarified. Fastnacht (in press) rected forwarc s and outwards. forming a tooth has reported on a coloborhynchid from the San- grab that is bounded anteriorl!~ b!. the first pair tam Formation of Brazil. referring an associated of teeth. Crested forms of Colo/)oi./i!./i(./ii(.s(one rostrum and mandibular symphysis to 'Tropeog- Cambridge GIeensand species is crestless) are IILIIII~I.~'rohsrz/s Wellnhofer. 1987 and assigning further distingiiishcd from Oi.iiitl7ocllcil.ll.,~cl~~~i/.i/.~by the this species to C'olohor-li?tncliirs. Finally, a semi- shape of the crest. as discussed abo\e (see also corn 13 1e t e s k e let on of Coloho rhy nch r is, also from Lee 1994, Fastiiacht in press). the Saiitana Formation, is currently under study Un t i 1 re ce n tl !r. Colo ho r/i!,r icli I LS \va s tho ug lit to 13). Veldmeijcr (19%). be represented by only a single species. C. cirr1,i- Colohor.Ii!./ic.Illrs is represented by two species rosiris. known from a single specCinien fi-om Sus- in thc Cambridge Grccnsand: C. cnpito Seclcy, sex (Owen lS"4). In 1994 Lee described a nc'u 1870 and C. srrfgivickii Owen, 185%. Colobor- species. C. it~r/~~leiglii.from the Lo\vcr Cretaceous Ii\~/icli~/scnpito. a crested form, known from four (Albian) of Texas. again based on a single fray- fra gmen t a r y 1-0s t 1-21, in c1 udi n g CAMS M B 54.625, mcntary rostrum and in 1999 Mader and Kellner the holotype (Fig. 8), and possibly two fragmen- described anot ier incomplete rostrum from nun- tary mandibular symphyscs, is similar to the type marine Cretaceous (Albian-Cenomanian) beds species C. cl(/t*iro.stris,but distinguished by the of MOI-OC~~ii ; the holotypc of 'Si,.ot,c,oi)rc.r:\..\. much thicker and taller sagittal crest that, in lat- Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 4 (2001) 207
B
Fig. 8. Anterior end of the holotype rostrum (CAMSM B54.625) of Colo- horhynchils crrpito (Sceley. 1870) in (A) right lateral. (B) anterior and (C) ventral (palatal) view. Abbrcvia- tions: p, pitting caused by '?osteopel- tid gastropods: rc. rostral crest: 1-5. positions of first five dental alveoli. Scale bar = 10 mni. era1 view, has a distinctly concave rather than of the rostrum, including the holotype CAMSM flattish anterodorsal margin. Coloborhynchus ca- B.54.422 (Fig. 9), and possibly four fragments of pito was a large pterosaur: comparison with a the mandibular symphysis. The shape of the ros- relatively complete skeleton of a similar ptero- trum and the size and arrangement of the teeth saur from the Lower Cretaceous of Mongolia in C. sedgwickii is similar to that seen in C. cayi- (Bakhurina & Unwin 199.5) indicates that the ho- to, and the only clear distinction between the lotype represents an individual that was prob- two species is that C. sedgwickii lacks a rostral ably over 5 m in wingspan, while BMNH R4S1, crest. In other pterosaurs the presence or ab- the largest known individual, is likely to have sence of cranial crests has been interpreted as been well over 6 m. sexual dimorphism (Bennett 1992), thus it may Coloborhynchus sedgwickii Owen, 18.59a is well be that C. cupito and C. sedgwickii are sex- known from eight fragments of the anterior end ual dimorphs of a single species. Synonymy of
Fig. 9. Anterior end of the holotype rostrum (CAMSM BS4.422) of Coio- borhyrzchus sedgwickii (Owen, 1859a) in (A) anterior, (B) left latcral and (C) ventral (palatal) view. Abbrevia- tions: mr, midline ridge: 1-7. positi- ons of l'irst seven dental alveoli. Scale bar = 10 mm (Modified from Owen 185%). 208 Unu in. D. M.. Cr'ctaccous pterosaur assemblage from En~land these two species should wait. however. until we tinguished from other species of Anhangicem by have a better understanding of the remaining the relatively evcn width of the rostrum cranial and postcranial anatomy of these tam (Fig. 10A. B) and mandibular symphysis which and other coloborhynchids. and of the signifi- lack the marked expansion of the anterior end cance of cranial crests. seen in A. hlirrerstlocfi (Campos & Kellner A third geniis of ornithocheirid from the Cam- 198%). A. srrrzrtirzne (Wellnhofer 1991b) and A. bridge Greemand is identified here as Aii/7~iii- 'piscotor' (Kellner & Tomida 2000). The large giiern. This geiius was first established on the ba- series of jaw remains of A. czivieri from the sis of a skull from the Santana Formation of Cambridge Greensand includes some juvenile in- Brazil which \:as made the holotype of Aid7rm- dividuals. and numerous sub-adultsladults, typi- giieirr hlirrrr-stl irf/i (Campos & Kellner 198%). cally of about the same size as the holotype spe- This. and otht r nominal species of Anhrrngirertr cimen. which. on the basis of comparisons with (A. .sIitirmiic. A. .piwtror.'). exhibit typical or- more complete remains of Anhctnguern from the nithocheirid features such as the development of Santana Formation (Wellnhofer 1991~Kellner & a terminal rost tte of enlarged teeth. thus Aiili(iii- Tomida 2000) had an estimated wingspan of gii r rtr be 1o ngs within the Orn i t hoc h e i r i d ae. The about 3.5m. Some specimens, such as CAMSM problem with .4ii/itiiigirei.a is that it may be sy- B54.431 (Fig. 1OA. B), represent substantially nonymous with previously described Santana larger individuals that may have reached wing- Form at ion tax; I such as B rosileo rlir crjsli IS ( Ke I1n c r spans of up to 5 ni. 1984) or. as Bennett has noted (1994: 22). with The second species of Anhanguera, A. fittoni Strnrrrnu~rtc~r~~lzis(de Buisoiije 1980). However. (Owen 1KSC)b) is represented by five fragments until this and c ther taxonomic problems concern- of the anterior end of the rostrum (Fig. 10C-G), ing Santana pterosaurs have been resolved I pre- including the lectotype (CAMSM B54.423, Owen fer to retain the name A/iharigiter.rr for niatcrial lK59b: pl. i. figs 3a-c; Fig. lOF, G) and possibly assigned to A. blitrersriorffi'. and other nominal two fi-agnients of the anterior end of the mandib- species currently included in this genus (e.g.. ular symphysis. Arihnnguern fittoni is distin- Kellner & Tor ida 2000). guished by the broad, flattened and spatulate Aiihungiirro seems to be distinct from Colo- jaw tips that are blunter and rounder, with rela- hor/zyd~iisin that while the palatal surface is tively more widely separated tooth rows than in also reflected ipward at the anterior tip of the other species. Assuming that the skeletal propor- rostrum. the r2flection is at a relatively gentle tions wcre similar to other species of Anhcm- angle (about 45'. rather than 90 in Colohor- giier~r then. typically. individuals of A. fi'ftoni /z~nc/ziis)and not so abrupt. MoreoLZer. in skulls were probably about 3-33 m in wingspan. that bear a crest. this is located somewhat pos- terior to the jaw tip (Kellner & Tomida 2000: Loncliodectidae: This is a poorly known fa- figs 4. 5. 62, 60-69), rather than at the anterior mily of small to medium-sized (probably about end of the jam as in Colohoi.h?.iic./iii.l (see 1-2 ni wingspan) relatively non-derived ptero- above). In othx respects. however. such as the dactyloids recorded, so far, only from the late size. spacing and orientation of the teeth. these Lower and early Upper Cretaceous of southern two genera arc remarkably similar and further England (Fig. 1). Lonchodectids are distin- work is needcd to determine whether thcy are yuished by teatures of the jaws and dentition in- truly dislinct (sx also Fastnacht in press). cluding: dorcoventrally flattened jaw tips; distinc- Tbvo species of Arilinrigilr/n. A. cu\,ieri (Bo- tive straight. narrow, parapet-like dental margins; wcrbank 1851) and A. ,firroni (Owen 185%). are and small. round, subequally-sized dental alveoli currently recognised from the Cambridge Green- with margins that are raised into a low collar so sand. The holo ype of A. ciri,irri (BMNH 39409). that the teeth appear to be pedicellate (e.g., a well preserved rostrum. was collected from the Owen 1851b: pl. xxxi, figs 1-7, 1874: pl. ii, figs Lower Chalk of Susses and originally described 1-8. Hooley 1914: 535). undcr the nam: Pterotiacryliis ciri,icri by Bower- Lonchodectids have elongate cervical verte- bank ( 1 H51 ). i\ri/iarzgriero ciii,icri is by far the brae with low neural spines (Owen l860b: pl. i, most common species of pterosaur in the Cam- figs 35-37. 1861: pl. ii, figs 13-16) and a rela- bridge Greensand kvhere it is represented by 23 tively underived pterodactyloid humerus with a fragments froin the anterior end of the rostrum straight dcltopectoral crest (Seeley 1870: pl. iv, and 19 fragme its from the anterior end of the figs 1. 2. 7-1 1). Thus they are certainly not or- mandibular syniphysis. Atilratrgiirro cri\*ieri is dis- nithocheiroids which have short, deep cervicals Mitt. Mus. Na(.kd. Rerl., Geowiss. Reihc 4 (2001) zoo
A B
mr
C
EL
mr F G
Fig. 10. Anterior end of the rostrum (CAMSM BS4.431) of a largc individual of Anhringirera cirvieri in (A) lcft lateral and (B) vcntral (palatal) view. Anterior end of the rostrum (CAMSM B54.556) of Anhnnguera fiftoni in (C) right lateral. (D) ventral (palatal) and (E) dorsal view. Anterior end of the lectotypc rostrum (CAMSM BS4.423) of Anhnngwni ,fifioiii in (F) right lateral and (G) ventral (palatal) view. Abbreviations: mr, midline ridge: 1-11. positions of first eleven dental alveoli. Scale bar = 10min. with tall neural spines and a twisted deltopector- for the present, they are placed in an unresolved a1 crest on the humerus (Padian 1986, Bennett trichotomy with Ctenochasmatoidea and Dsun- 1989, Unwin & Lu 1997). The relationships of garipteroidea + Azhdarchoidea (Fig. 14, sec be- the Lonchodectidae to the major non-ornitho- low). cheiroid clades (Ctenochasmatoidea, Dsungarip- Currently, the earliest record for the Loncho- teroidea and Azhdarchoidea) are not clear and, dectidae is ‘Pteroductylits’sngittirostris (= Lonch- 210 L'iiwiii. D. M.. Cretaceous ptcrosaur assemblage from England otlectes .sogittijnstri.s). based on a sinple. inconi- crests (Owen 1851b: pl. xxxi, figs 1-7). pletc lower jaw from the Hastings Sands of Sus- Although it has been assumed that L. giganteus sex (Owen 1874). Lonchodectids are best known had short. deep jaws (e.g., Wellnhofer 1978), this from the C;imbridge Greensand uhich has species prchably had an elongate rostrum and yielded nine jLiw remains. representing. at most. mandibular symphysis as in other lonchodcctids. four species (--able 1 ). and a series of postcranial The holotype of L. conipressirostris (BMNH bones. iticludiiig fragmentary vertebrae and fore 39410) appears to have a narrow, blade-like tip and hind lirnl, elements (Hoole!. 1914. UnLvin to the rostrum (Owen 1851b: pl. xxviii, figs et al. 2000). 8-10). unlike the rather spatulate jaw tip of Lonchodect ds have also been found in the other lonchodectids. The anterior portion of the Lower Chalk of Kent. En~land.ivhci-e they are rostrum is heavily conipi-essed, however, and ori- re prese 11t ed b;,' two speci cs : L oric.1iodrc.tc.s gig[//I - ginally was probably much flatter and broader. [U/.Y (Bo\vurb::nk 1846) and L. L'orizi,r.r.s.sii.o.st/.i.\- In the Cambridge Greensand Z~~~i~chodc~ct~s (Owcn 185 1 a 1. 1,oric~liorlcc~rc.s gigriiiteirs is the coiiiI~~es.sir.ostri.sis represented by a single frag- onl!. lonchodi ctid kno\vn so far in udiich the ment of the rostrum (CAMSM BS4.584; upper and lev, er jaws are preserved in associa- Fis. 11A. B) that is identical to thc holotype spe- tion and has xoininent rostra1 and mandibular cimen from the Chalk. A second Cambridge B mr \ ' ' ' '-7 1 --.--.v ...-..__..-_.... -_ I \-- //,,,//,r,, I/ h ...... -... ----;r .
C D d C h a
...... _I I . .;._ ..
dm E F I Mitt. Mus. Nat.kd. Berl.. Cieowiss. Reihe 4 (2001) 71 1
Greensand species, Lonchodectes microdon, re- dectes. Consequently, even if crests are a sexually presented by fragments from both the rostrum dimorphic character it seems likely that at least and the mandibular symphysis (Fig. 11C-F), is two species of Lonchodectes are present in the probably synonymous with L. compvessirosfris, Cambridge Greensand. although this synonymy cannot be satisfactorily demonstrated on the basis of existing material. ? P t e r a n o d o n ti d a e : The principal represen- A third Cambridge Greensand species, L. mn- tative of this family, Ptevanodon, is well known chaevorhynchus, is represented by a single, frag- from the Smoky Hill Chalk Membcr of the Nio- mentary mandibular symphysis (CAMSM brara Formation and the overlying Sharon BS4.855) that is similar to material assigned Springs Member of the Pierre Shale of western to L. microdon, but distinguished by the pre- North America and has been extensively re- sence of a prominent triangular median crest viewed by Bennett (1992, 1993. 1994, 2001). (Fig. 12D, E). A fourth species. L. platystomus, Ptcranodontids are distinguished by their unu- which is also known from the English Gault sual skull morphology, with long. slender. (Owen 1874: pl. ii, figs 5, 6; Fig. 12C), is repre- pointed edentulous jaws, a crest that projects sented by fragments of the rostrum (Fig. 12A, B) posterodorsally from the fronto-parietal region and mandibular symphysis, both bearing median of the skull, and many other cranial and postcra- crest\. The crests arise from the anterior tips of nial characters (Bennett 1994: 24). Nyctostrui-its, the jaws and are rather different in shape and another edentulous pterosaur from the Upper proportions to those of other species of Loncho- Cretaceous of the New World (Wellnhofer
B dm
1 3 4
C
mc' D E dm, mg, r------', ', \ \
abc d
mc
Fig. 12. Anterior cnd ol a rostrum (YORM 19831113F) of Lorichorlrctes pltrtysiomris in (A) left lateral and (B) ventral (pnla- tal) vicw. Anterior end of a mandibular symphysis (BMNH 43074) of Lonchorlrctes plutysionzirs in (C) left lateral view. An- tcrior end of the holotype mandibular symphysis (CAMSM BS4.SSS) of Larzclzodecies mnchrirrorliyndllrs in (D) left lateral and (E) dorsal view. Abbreviations: a-d. positions of four dental alveoli; dm, dental margin: mc. mandibular crest: mg. mid- line groove; mr, midline ridge: rc. rostra1 crest: 1-4, positions of first four dental alveoli. Scale bar = 10 mm. 212 Unwin. D. M.. Cretaceous pterosaur assemblage from England
1991a) has often been included in the Pterano- the Sedgwick Museum, Cambridge led to the dis- dontidae (e.g.. Wellnhofer 1978). but recent stu- covery. in 1986, of two edentulous jaw fragments dies (Bennett 1989. 1994. Unwin 1995. Unwin & and it seems likely that these are the original Lii 1997) suggest that it may not be as closely specimens referred to by Seeley. Hooky (1914) related to Preranocion as some toothed taxa such also assigned various postcranial bones (frag- as ornithocheirids. While most workers (e.g., mentary notaria, scapulocoracoids, type 'B' hu- Wellnhofer 1978) have confined the Pteranodon- meri and ulnae, and group 1 femora) from the tidae to Ptercirzodor? and N\..ctosniiriis (though Cambridge Greensand to Omithostoma, but see above). Padian (1986) and subsequently Ben- there is no evidence to support this opinion. nett (1989. 1904) expanded this family to include Ornithostonin sedgwicki was a medium-sized ornithocheirid:, (Anl?arigrrerri. Sar?tniiricl~ict!~liis. pterosaur with elongate, edentulous jaws and and possibly 'c~riorh!~iichiis'. ' Tropeogr intliiis ' and probably no more than 2-3 m in wingspan. The Ornithocl?eiriis) and I.Ytiodcrct!tlit.s (formerly Or- rounded. triangular cross-section of the rostrum nithodesnziis, sze Howse et al. 2001). This group- (Fig. 13B). which also has low, rounded, marginal ing is equivalmt to Ornithocheiroidea (see Un- ridges (Fig. 13C) clearly distinguishes Omithosto- win 24 Lu 1997) and here the Pteranodontidae is 1110 from azhdarchids wherein the rostra have restricted to Ftera17odon and another apparently concave lateral surfaces and lack marginal ridges closely related form. Ornitlzostoma. on the jaws (Wellnhofer & Buffetaut 1999: The name C~rriithostomnwas first proposed by fig. 4). Ornithosromn is also distinct from Tape- Seeley (1871) for a fragment of toothless jaw jnrcr and Rrpiixiinra, which have prominent mid- (CAMSM B54.485. Owen 1859b: pl. iv. figs 4-5: line crests on the rostrum and the mandibular Fig. 13A-D) from the Cambridge Greensand. syniphysis (Kellner & Campos 1988, 1994, Kell- This fragment had been mentioned in the 'Index' ner 1989. Wellnhofer & Kellner 1991). By con- (Seeley 186%) as possibly from a toothless pter- trast. as Seeley (1901) noted, Ornithostomn osaur. predatir'g the first description of the eden- shows some similarities to Pternnodon, especially tulous jaws of Prwnrzorlon (Marsh 1876) by some in the cross-sectional proGle of the rostrum and seven years. 3cCambridge Greensand speci- the presence of marginal ridges (Bennett 2001: men was later made the holotype of Ol-nirliosro- figs 2. 3). although the latter are low and MZCI .~edgwicki(Seeley 189la). In his 1871 paper rounded in Ornithostomn, rather than narrow Seeley also ni 2ntioned two other specimens re- and relatively tall, as in Pteranodon. For the pre- presenting a toothless pterosaur. but these were sent. Ortiithoston7a is tentatively assigned to the not figured cr described by Seeleq. or later Pteranodontidac. This determination appears to authors. A thcrough search of the collections in be supported by the presence, in the Cambridge B
mr
C D Fig. 13. Fragmentary holotypc rostrum (CAMSM BS4.485) of Ornithostotizu seclgwicki in (A) right lateral, (B) an- tcrior crosb section, (C) ventral (palatal) and (D) dorsal view. Abbreviations: mr. marginal ridge. Scale mr bar = 10 nim. Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 4 (2001) 213
Greensand, of a fragmentary cranium bearing 14). This total is inflated by taxonomic oversplit- the base of a tall, posterodorsally directed crest ting and the retention of invalid taxa: when (CAMSM B54.406, Hooley 1914: pl. xxii, figs these are taken into account, the list is reduced 1-3), supposedly a diagnostic character of Ptera- to only ten species (Unwin et al. 2000: table 1) nodon (Bennett 1994) although, interestingly, and true species diversity may have been even this type of construction is evident in a new, as lower. Partly because of its taxonomic richness yet undescribed ornithocheirid from the Crato and the large number of individuals recovered. Formation of Brazil (Frey quoted in Viohl 2000: and partly due to its age and geographic loca- 30, see also pl. ix, fig. 4). tion, the Cambridge Greensand pterosaur assem- Summary: Systematic revision of the Cam- blage has played, and continues to play, a promi- bridge Greensand pterosaur assemblage shows nent role in the reconstruction of the that of the 32 species named so far, only 10 are evolutionary history of pterosaurs. Two aspects, the history of particular clades, and the ecology valid, although there may be one as yet un- named species (Table 1). This probably repre- and palaeobiogeography of Cretaceous ptero- sents an overestimate of true diversity because, saurs are discussed here in the light of our cur- in one case (Lonchodectes compressirostris and rent understanding of this assemblage. L. microdon), the two species are probably sy- nonymous, although this cannot yet be demon- History of Cretaceous pterosaurs: Re- strated because of the lack of directly compar- cent discoveries, for example in South America able material. In a second case (Ornithocheirza (Santana Formation, Crato Formation, Lagarcito simus and Ornithocheirus sp.), the two species Formation) and China (Tugulu Series, Yixian may represent disjunct populations of a single Formation), coupled with older records from species in which intermediate forms are not yet Europe (Purbeck Limestone Formation, Hastings known. In a number of other cases (Loncho- Sands, Cambridge Greensand) are informative dectes compressirostris and L. machaerorhynchus, regarding the history of pterosaurs in the Early Coloborhynchus cupito and C. sedgwickii) taxa Cretaceous (see Wellnhofer 1991a, Unwin et al. that are distinguished only by the presence or 2000; Figs 1, 14). By contrast, the Late Creta- absence of crests may represent sexual dimorphs ceous pterosaur fossil record is more poorly of a single species, as has been proposed for known: there are no major assemblages such as Pteranodon and other pterosaurs (Bennett 1992). those found in the Cambridge Greensand and Should all these putative synonymies eventually Santana Formation and, apart from a few re- be demonstrated this would leave only seven cords such as that of Azhdarcho from the Beleu- species in the Cambridge Greensand. A further ta Svita of Uzbekistan (Nesov 1984, Bakhurina reduction in species diversity is unlikely, how- & Unwin 1995), little is known about late Ceno- ever, because the remaining taxa can be clearly manian-Santonian pterosaurs (Fig. 14). Conse- distinguished from one another on the basis of quently, the Cambridge Greensand and closely dental and other characters. related, but slightly younger, Lower Chalk ptero- saur assemblages of England are of special inter- est as they provide the last 'view' of several Early Cretaceous pterosaur clades, and possibly General significance the first view of an important Late Cretaceous of the Cambridge Greensand pterosaurs lineage, the Pteranodontidae (Fig. 14). The Cambridge Greensand and Lower Chalk The Cambridge Greensand has yielded one of of Kent yield some of the youngest records for the most diverse pterosaur assemblages yet the Ornithocheiridae, a relatively long lived known (Wellnhofer 1991a), a statistic that re- clade that is certainly known for almost the en- mains true even if there was a minimum of se- tire Early Cretaceous (Fig. 14) and probably ori- ven species. The only Cretaceous locality that ginated in the Late Jurassic. The Cambridge has produced a similar taxonomic diversity is the Greensand contains possibly the youngest known Lower Cretaceous Santana Formation of Brazil record for Coloborhynchus, which extends back (Wellnhofer 1991c, Kellner 1991; Figs 1, 14), to at least the Valanginian. Coloborhynchus nior- which in a recent paper (Kellner & Tomida occevlsis from the Kem Kem beds of Morocco 2000) is accredited with 17 species (this does not may represent an even younger record, but only include taxa from the Crato Limestone, which is if the age of the Moroccan deposits is accepted a distinct and somewhat older unit - see Figs 1, as Cenomanian, and not Albian. Ornithocheirrrs 211 I'nnin. D. M.. Cietaccoua pterosaur assemblage from England
1 Javelina Fm. Maa v) z a -. 3 m a n 2 c c1 0 Two Medicine Fm. -0 2" 0 eC za
sari , Niobrara Fm. I -8 -8 Con
Tur - Cen Lower Chalk Kern Kern beds -Cambridge Greensand
A'b Santana Fm.
Crato Fm. Apt
- Bar Yixian Fm.
Hau
Val Hastings Sands __ I I I I I Ber .I Purbeck Lst. Fm. 3 Ornithbcheiroidea I Pterodactyloidea ...... _._....:...... I
Fig. 1 Stratigrap iic disti-ih~iti~not' important C'rc.tacco~i~pterowiir assemblages and their relationship to thc temporal histo- I!. of the main cl.ides ot Crt'taceous ptcrouurb. Ph! I( i! ha\ecl 011 I,'n\\in (l095). Unwin XC Lii (1997) and Unwin et al. (2000). and incluil :\ only thow clacles that h;i\.c a C'rrt OLI~rccortl. 'l'hick. solid lines indicate the known stratigraphic range of ii partic~ilnrclam le. although fossil recoi-ds ma! not ha\ e hern reported for all stqes within that range. Thick. dashed lincs indicate ;I pos4hle I-angc extension hascd on an unconfirnietl I-ccortl. 01- records. Thin solid lines indicate lineages prcdicled by the ph! IoFenrtic liypothesis adopted here Thin dashrcl line$ indicate tincertainty concerning the relationship of a clade to oilier taYx Solid circle5 indicate ii confirmed I-ecord tor ;I particiilai- assemhlage. circles enclosing a cluestion mark indicate an unconfirmcd recoi d. Ahhri.\ iations: 41h. Alhian: Apt. Aptian: Bar. Bai-remian: Rer. Bcrriasian: Cen. Ccnomanian; Con. (lo- iuacim: Ciiip. C'ai-ipanian: Fin. Formation. Hau. 1 inuteri\ im Lst. 1.iniestonc. hlaa. Maastrichtian: San, Santonian: Tur. Turo- nian: Val. \'al:iiicii ian Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 4 (2001) 913
simirs from the Cambridge Greensand is the Ecology and palaeobiogeography of youngest record for the genus, which is other- Cretaceous pterosaurs: Comparison of the wise only certainly known from the Aptian. An- Cambridge Greensand pterosaur assemblage hangiiera is well represented in the Cambridge with other late Early and early Late Cretaceous Greensand and persisted into the Cenomanian pterosaur assemblages (Figs 1, 14) provides sonie where A. aivieri, from the Lower Chalk of Kent, insights into the ecological diversity of Creta- is the youngest record both for this genus and ceous pterosaurs. The Cambridge Greensand as- for the Ornithocheiridae (Fig. 14). Various mid semblage is dominated by medium to large prob- to Late Cretaceous pterosaurs have been as- ably piscivorous ornithocheirids, but also signed to the Ornithocheiridae (see Wellnhofer contains another relatively small, probably pisci- 199la), but none of these can be certainly de- vorous form, Ornithostoma, and lonchodectids, monstrated to have belonged to this family. small to medium-sized seemingly non-specialised Moreover, although the Late Cretaceous ptero- pterosaurs. The Cambridge Greensand and un- saur record is relatively poor, as mentioned derlying Gault Clay are shallow marine deposits above, the absence of any definite post Cenoma- and it seems likely that the Cambridge Green- nian records for the Ornithocheiridae contrasts sand pterosaurs inhabited shallow shelf seas. with the growing number of records from the feeding on fish and other aquatic prey caught Early Cretaceous and it seems likely that or- from the surface of the water. The slightly nithocheirids became extinct early in the Late younger Lower Chalk of Kent, England contains Cretaceous. a smaller, but almost identical assemblage to the The presence of Lonchodectes sagittirostris in Cambridge Greensand (Figs 1, 14). This sc- the Hastings Sands of Sussex, England, demon- quence also represents a shallow shelf sea. con- strates that the Lonchodectidae were established sequently the pterosaur assemblage from this de- by the Valanginian (Fig. 14), while records from posit would seem to represent a continuation, the Gault Clay, Cambridge Greensand and Lower into the early Cenomanian in the West Eur- Chalk show that this family persisted throughout opean region, of a pterosaur community typically the rest of the Early Cretaceous and survived into associated with a shallow marine environment. the Cenomanian. The apparent absence of any The Kem Kem beds of the Ksar es Souk Pro- younger records suggests that the clade became vince, Morocco (Fig. 15). interpreted as non-mar- extinct sometime in the early Late Cretaceous. ine possibly deltaic deposits (Sereno et al. 19%). The presence of Ornithostoma seclgwicki in the are approximately coeval with the Cambridge Cambridge Greensand is of special interest be- Greensand-Lower Chalk of England (Fig. 14). cause, if it is indeed a pteranodontid, as sug- The Kem Kem assemblage shares some elements gested above, it is the earliest record for this in common with the English pterosaur assem- clade. Pteranodontids are otherwise only cer- blages, notably the presence of ornithocheirids tainly known from the Coniacian-Campanian in- and perhaps pteranodontids (Mader & Kellner terval (Bennett 1984, 2001), thus the Cambridge 1997, 1999, Wellnhofer & Buffetaut 1999; Fig. 1). Greensand record represents a considerable Interestingly, the one identifiable ornithocheirid range extension. This discovery, which is also from the Kem Kem beds, Coloborhynchus. has supported by the tentative identification of a generally been reported from deltaic (Paw Paw pteranodontid jaw fragment from the Cenoma- Formation, Hastings Sands) or lagoonal (Santana nian of Morocco (Wellnhofer & Buffetaut 1999), Formation) deposits, unlike other ornithocheirids is not unexpected because, if the Ornithocheiri- which have often been found in marginal marine dae is the closest known sister group to Pterano- or fully marine deposits (Unwin et al. 2000). tlon (Bennett 1989, 1994, Unwin 1995, Unwin Lonchodectids have not, so far, been reported et al. 2000) the existence of this clade in the from the Kem Kem beds, but the latter have Early Cretaceous (Fig. 14) implies that the line- yielded an azhdarchid (Kellner & Mader 1996. age leading to the Pteranodontidae must also Wellnhofer & Buffetaut 1999) and a species of have existed by this time, or possibly even ear- Tapejara (Wellnhofer & Buffetaut 1999), neither lier. Ornithostoma appears to have been much of which have been found so far in Albian-Ceno- smaller than Pteranodon (wingspans of manian deposits of England. Tapejara has been 4-6.25 m, Bennett 2001) which suggests that the interpreted as a frugivore (Wellnhofer & Kellner evolution of large size within Pteranodontidae 1991) which, if correct (a detailed analysis has occurred independently from the same trend in yet to be carried out) means that this taxon in- other clades such as Azhdarchidae. habited continental environments. Azhdarchids 216 1 ii\\ in. D M . Crct'iceous Dterosdur assemblage from England have frequen ly (though not exclusively) been et al. 2000) are not known from the Cambridge reported froni continental deposits (Wellnhofer Greensand. while Lonchodectes is so far un- 199la). but their fccding ecology: scavcnp. \va- known from the Santana Formation. In some der or piscivoc is still unclear (see Martill 1997. cases. such as Trrpjora, this difference might be Prieto 1998 for discussion). attributed to ecological differences, in that the The diffcrerices in the composition of the Ens- Santana basin seems to have been a land-locked, lish and Morc ccan assemblages are most readily or largely land-locked marine-brackish water ba- interpreted as the result of ecolosical d sin (Martill 1903). into which putatively conti- tion. The Ken1 Kern beds yield tasa. such ax EI- nental forms such as thc supposedly frugivorous pejjc/rr/ and az,idarchids, that are typically found tapejarids may have fallen. In the case of Lon- in Continental or coastal deposits and that have cliorlecte.~.howcver, its absence from the Santana been argued by some workers (Lawson 1975. Formation and indeed from any other Lower Langston l98:1. Wellnhofer &: Kellner 1901) to Crctaceous deposits other than those of England be adapted tc continental habitats. B!, contrast. might reflect a degree of endemism. the absence o ' any large land masses in the vici- In summary. at present it seems that the taxo- nity of Englaild in the late Albian (Fig. 15) is nomic distinctions apparent between the main consistent with the presence of typically marine late Early and early Late Cretaceous ptcrosaur forms such as the ornithocheirid Aiil1rrrigirci.rr assemblages cannot be attributed to evolution and the putati ie pteranodontid 0rnif1iost01nr1. Lvithin clades. or to clade origin or extinction as The pterosaur assemblage from the Santana nearly all the main clades persist throughout this Formation is :.ornewhat similar to that from the interval (Fig. 14). Rather, the differences can be Cambridge Greensand in that it too is domi- attributed to palaeoecological differentiation, nated by orniihocheirids (Figs 1. 14). Species of although the restricted distribution of Loncho- 0rn it11 o ch eiri I:, Colo h o r/ij*ii cl I i I .T and A I i I7 01ig i I r IYI tiecws also susgests the operation of some de- have all been reported from this deposit (Unwin srec of endemism. Previously, Unwin et al. et al. 2000) aiid. in most cases. are remarkably (2000) outlined evidence for palaeoecological similar in size and morphology to the Canibridy differentiation in Early Cretaceous pterosaurs, Greensand tar a. By contrast. other Santana tasa principally between marine and continental such as thc t: pejarids 7irpc.jrrrrr and Tqi~~/r1ru.forms. The observations prescnted above support and the ctenochasniatid Crni.rrritrct~~Izi.s(Unwin this conclusion.
Fig. 15. P~il~icogi.c.~i-apliicili\trihution of late Earl!. and earl! Late C'i-ctaceous ptcrosaur assemblages. Taxonomic composition of ai\cmhlage\ \I1 )\vn on Fiz. 1. Palacogrogi apli! hadon Smith et al. 1994. Ahhi-evialions: 1. Cambridge Greensand. Eng- land: 2. Louc'r Cl-alL. England: 3. Ztiunba!m S\ itii. Khtircii-Dukh. klongolia: 4. L!wya Cora. Saratov. Russia: 5, Kern Kern red he&. Lloroccc 1: 6. Pa\\ Pa\\ Foi-mation. Te\;ij. LSA: 7. Laprcilo Formation. Snn Luis. Argentina: 8. Santana and Crato For mat io 11\. Cc n 11 . B ra7i I: 9. Tool c I) tic Formal io 11. Qucc nsland . ,\u? t rali ;i . Mitt. Mus. Nat.kd. Berl., Gcowiss. Reihe 4 (2001) 217
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Cai, Zhejiang Museum of Natural History, Hangzhou, China. Buffetaut, E., Cappetta, H., Gayet, M., Martin. M., Moody. I thank N. Bakhurina, S. Bennett, M. Benton, M. Fast- R. T. J., Rage, J. C., Taquet, P. & Wellnhofer, P. 1981. Les nacht, N. Fraser. E. Frey, G. Hazelhurst, s. Howse, A. Kell- vertebrks de la partie moyenne du CretacC en Europe. - ner, W. Langston Jnr., D. Martill, K. Padian, C. Walker and Cretaceous Research 2: 275-281. P Wellnhofer. for many helpful discussions of the Cambridge Buisonje, I? H. de 1980. Santanadactylus hrasilensis nov. gen. Greensand pterosaur assemblage. The Department of Zool- nov. sp., a long necked, large pterosaurier from the Ap- ogy, Reading University, Dept of Earth Sciences, Bristol Uni- tian of Brasil. - Proceedings Koninkl Nederland Akade- versity, the Royal Society and the British Council are mie Wetenschappen, B 83: 145-172. thanked for funding aspects of this work. A. Jenkins and S. Campos, D. de A. & Kellner, A. W. A. 198Sa. Panorama Powell assisted with the photography, T. Bergmann, S. of the flying reptiles study in Brazil and South Ameri- Grimm, C. Finkenwirth and A. Schulz very kindly helped ca. - Anais da Academia Brasileira de Cihcias 57: with the maps and line drawings and R. Schoch translated 453-466. the German abstract. I extend my best thanks to M. Romano - 198Sb. Um novo exemplar de Anhangcicra hlitrervtlorffi (Sheffield University) for his advice and encouragement in (Reptilia, Pterosauria) da formaqiio Santana, Creticeo In- the early stages of this project. J am very grateful to N. Ba- ferior do Nordeste do Brasil. - Boletim de Resumos. 9 khurina, P. Barrett, s. Bennett, M. Benton, D. Martill, D. Na- Congress0 Brasileiro de Paleontologia: 13. ish and l? Wellnhofer for their constructive criticism of ear- Carter, D. J. & Hart M. B. 1977. Aspects of mid-Cretaceous lier versions of this paper, to N. Bardet and N. 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Note added in proof relatively small). Moreover, it is consistent in other respects with ornithocheiroid anatomy (e.g. wing-nictacarpal of simi- lar length to the humerus). Haopterus appears to be the Nuopterus grricilis, a relatively small (1 35 m wingspan) ptero- smallest adult ornithocheirid known so far, and one of the dactyloid was recently described by Wang and Lu (2001) earliest representatives of this clade (cf. Fig. 14). and may in- from the Yixian Formation (Lower Cretaceous: Barremian) dicate that the large sizes achieved by some Cambridge of western Liaoning, China and assigned to the Pterodactyli- Greensand ornithocheirids occurred independently from that dae. Haoprerus lacks apomorphies of Ctenochasmatoidea in other ornithocheiroids (Dtiodactylris. pteranodontids) or (Pterodactylidae + Ctenochasmatidea, sensu Unwin et al. other pterodactyloid clades. 2000). such as an almost horizontal quadrate and associated complex of characters (Unwin & Lii 1997), but exhibits two ornithocheiroid apomorphies (coracoid longer than scapula, Wang Xiaolin & LLi Junchang 2001. Discovery of a pterodac- metatarsal <25% length of humerus) and a character com- tylid pterosaur from the Yixian Formation of western plex diagnostic of ornithocheirids (first three pairs of pre- Liaoning, China. - Chinese Science Bulletin 46 (13): maxillary teeth incrcase in size posteriorly. while the fourth is 1112-1117.