Wima1lkirtlandicus Goniopholis lromthe forgotten crocodile Formation, SanJuanBasin, Kirtland NewMexico NIM Socorro, Resources, andMineral Paleontologist, NewMexico Bureau of lVlines Vertebrate byDonald L. Wolberg, Crocodilian bone, armor, and teeth are common in Late Cretaceous and Early Tertiary deposits of the San Juan Basin and elsewhere.In the Fruitland and Kirtland Formations of the San Juan Basin, Late Cretaceous crocodiles were important carnivores of the reconstructed stream and stream-bank community (Wolberg, 1980). In the Kirtland Formation, a mesosuchian crocodile, Goniopholis kirtlandicus, discovered by Charles H. Sternbergin the early 1920's and not described until 1932 by Carl Wiman, has been all but ignored since its description and referral. Specimensreferred to other crocodilian genera may actually represent G. kirtlandicus or other mesosuchians. A better understanding of the distribution of G. kirtlandicus may aid in determining.age relationships in Late Cretaceousdepositsof the San Juan Basin. Numerous studies related to Late Cretaceous vertebrate faunal assemblages of the San Juan Basin frequently mention crocodiles, but usually only in passing. Published work includes Gilmore (1916, 1919, 1935), Bauer (1916),Reeside(1924),Colbert (1950),Powell (1972, 1973), and Wolberg and LeMone (1980). Unpublished reports include Kues and others (1977) and LeMone and others (1977, 1979). Gilmore (1916) reported crocodiles and Brachychampsa sp. from the Kirtland and "crocodiles" from the Ojo Alamo Sandstone. Later, in 1919, he reported "crocodiles" from the Fruitland, Kirtland, and Ojo Alamo but noted in 1935 that "the Crocodilia and Pisces are too fragmentary to be of assistance" (p. 187). Reeside (1924) lists Crocodylus sp. and Brachychampsa? sp. for the Kirtland Formation. Powell (1972, 1973) lists Crocodylus sp. and Brachychampsa? sp. for the Fruitland and Kirtland Formations. Crocodylus sp. and Brachychampsa? sp. are also mentioned by Kues and others (1977), LeMone and others (1977, 1979), and Wolberg and LeMone (1980). Few detailed analyses of crocodilians have been done, and the Late Cretaceous crocodiles of the San Juan Basin have not been adequately studied. Early paleontology in San Juan Basin Charles H. Sternberg, a well-known fossil collector of the late 1800's and early 1900's, sold a number of reptilian specimenshe collected in l92l in the San Juan Basin to Carl Wiman, paleontologist at the Paleontological Institute of Uppsala University (Sternberg, 1932). Wiman (1930) describedPentaceratops posed that the narial cavities of Parasaurolophuswerevocalresonatingchambers' Apparently included with this material shippedto Wiman was a partial skull that Wiman describedas a new speciesof crocodile, Goniopholis kirtlandicus. Wiman publisheda descriptionof G. kirtlandicusin 1932in the Bulletin of the GeologicalInstitute of IJppsala. Notice of this specieshas not appearedin any Americanpublication.Klilin (1955)presented a descriptionand illustration of the speciesin French, but essentially repeatedWiman (1932). Localityinformationfor Goniop holi s kir t landicus The skeletal material referred to Goniopholis kirtlandicus includesmost of the right side of a skull, a squamosalfragment, and a portion of dorsal plate. The referral of the an interpretadorsalplateprobablyrepresents tion of the proximity of the material when found. Figs. I and 2, taken from Wiman (1932),illustratethis material. Wiman (1932,p. 181)recordedthe following locality data, provided by Sternberg: "Skull of Crocodile.Kirtland shalesa 100feet below the Ojo Alamo Sandstonein the blue cley. Barrel Springs Wash, 2.5 miles S. Trading Store (Post), Ojo Alamo Wash New Mexico.CH. H. STERNBERG1921."In his Wiman notesthat descriptionof the specimen, the bones are light gray externallybut are darker. almost btack ("fast schwarz"), "on FIGURE l-RecoNsrnucrroN oF DoRSALASPEcToF Got'ttopnotts KTRTLANDICUS,approximately % size. Abbreviations: Na, external nares; In, indentation for pseudo-canine of mandible; Pm' premaxillary; Mx, maxillary; N, nasal; La, lacrimal; Pfr, prefrontal; Or, orbit; Por, postorbital; Fr, frontal; Fts, superior temporal fossa; Pa, parietal; Sq, squamosal (after Wiman, 1932). fenestra, a ceratopsian dinosaur found by Sternbergin Kimbeto Wash.Sternbergfound a similar specimenin 1922and sold it to the American Museum of Natural History. Henry Fairfield Osborn, the well-known American paleontologist,publisheda descriptionof this specimenand named it Pentacerotopssternbergii (Osborn,l92l). Wiman (1931)publisheda descriptionof still anotherdinosaur found by Sternbergin 1921,the hadrosaurParasaurolophustubicen. P. tubicenis characterized by the development of a cranialcrest(Lull and Wright, 1942)that may havefunctionedas a visual and acoustical displayorgan (Hopson, 1975).Wiman suggesteda similarideain his 1931paperand pro- oF rHE PALATAL FICURE 2-RrcoNsrnucrtoN ASeECToF Got'ttoptlotts KIRTLANDICUS,approximately % size. Abbreviations: Pmn premaxillary; Mx, maxillary; Pal, palatine; Or, orbit; ln, indentation for pseudo-canine of mandible (redrawn from Wiman, 1932,with modifications)' New Mexico Geology August1980 freshly fracturedsurfaces.He notesthat the enclosingmatrix consistsof a yellow-gray, fine-grainedsandstonethat provedto be very hard duringpreparation. The old Ojo Alamo Trading Post was situated along Alamo Wash in SE%NW% sec. 8 , T . 2 4 N . , R . l l W . a t a n e l e v a t i o no f approximately 6,340 ft (Clemens, 1973\. Sternberg noted that G" kirtlandicus was found approximately2.5 mi south of the trading post in Barrel Springs Wash, now known as De-na-zinWash. Sternbergprobably overestimatedthe distancesouth of the trading post. The specimenprobably came from nearthe middle of sec.20, T. 24 N., R. 1l W.: a distanceof 2.5 mi south of the tradingpost sitewould haveplacedSternberg in the extremesouthernportion of sec.20, out of De-na-zinWash(fig. 3). The stratigraphicdefinitionsof the uppermost Cretaceous and lowermostTertiaryunits in the San Juan Basinhavebeendiscussed at lengthby Baltz and others(1966),Fassettand Hinds (1971), and Clemens (1973). The boundary between the Kirtland and Ojo Alamo Formation, long disputedby various authors(Brown, l9l0; Sinclairand Granger, l9l4), lies within the area of the old Ojo Alamo Trading Post. Fassett and Hinds (1971)mappedas Kirtland the areadescribed by Sternberg'sdescriptionto Wiman. Sternberg (1932)seemscertainof his stratigraphic units; moreover, he reports lengthy discussionswith J. B. Reeside.who was also in the field in 1921; Reesideprobably provided Sternberg with stratigraphic information. Sternberg'sstratigraphy(1932)appearsto be coincidentwith interpretationspublishedby Reeside(1924\,and G. kirtlandicus must have beenfound in theKirtland Formation(fig. a). Gents Goniopholis Goniopholis was described by Owen on the basis of fossil material from the Wealden Cretaceous of England. The type species is Goniopholis crossidens. In addition to the type species and G. kirtlandicus, the following species have been recognized (Wiman, 1932; Kiilin, 1955): G. simus Owen, 1878, from the Wealden of England and Belgium, G. felix Marsh, 1877, from the Jurassic of Colorado, and G. gilmori Holland, 1905, from the Jurassic of Wyoming. With other genera that range in age from Late Jurassicto Cretaceous, Goniopholis is included in the family Goniopholidae. Romer (1966) suggested that Goniopholis and its allies were central to crocodilian evolution and formed the mesosuchianstock from which the eusuchians evolved in the Late Jurassic or Early Cretaceous. Goniopholrs was of typical crocodilian form with a broad and not overly long skull. The secondary palate was relatively more advanced than in other mesosuchians and extended farther back than in other forms. August1980 New Mexico Geology Oio Alano " T.P. z . *'."."tivl.q{'lu, ' x' fr* \),J-' .^ 'sl'l $(9 l->l $ \ .6 j_4i' ,. -"n*.(, On the basis of the observable morphology of G. kirtlandicus, the species is separable from other members of the genus noted above. lf additional specimens of G. kirtlandicus occur in existing collections, they should be separable from eusuchians, given adequatematerial. ,t Crocodilian phylogeny The crocodiles belong to the reptilian subclass, the Archosauria (the "ruling reptiles"), which includes the dominant animals of the Mesozoic Era, the dinosaurs and ptero' ,{' saurs. Along with other members of the sub,lot class, the order Crocodilia originated within !,a' the order Thecodontia sometime during the Triassic. The Crocodilia includes five 1F' suborders: Protosuchia, Archaeosuchia, Mesosuchia, Sebecosuchiaand the Eusuchia /L' (Romer, 1966). Since their appearanceduring t ,, the Triassic (fig. 5) members of Crocodilia have remained relatively generalized in structure and have retained many primitive characters, except for the development of a specialized palate. The crocodiles have adapted to an aquatic environment by FIGURE 3-Mnp oF rHE oLD Oro AL.c.N,ro Tneuuc developing a powerful tail for swimming, parPosr enEe, T. 24 N., R. I I W. Sternberg's localtially webbed feet, and-most importantly-a ity data for Goniopholis kirtlandicus indicated secondary palate that separatesbreathing and that the specimen came from Barrel Springs Arswallowing functions. This last adaptation is royo (now De-na-zin Wash), 2.5 mi south of the obviously useful for an animal that takes at trading post. As can be seenfrom the map, Sternleast part of its food under water. berg must have overestimatedthe distance. Modern crocodiles representthe last surviving remnant of the archosaurian branch that DescriptionoI Goniopholiskirtlandicus dominated the Mesozoic world. These recent, Wiman (1932)describedG. kirtlandicusin still-surviving members of the Eusuchia are detail and suggested reconstructions for miss- actually a relatively late adaptive radiation ing portionsof the skulland palate(figs. I and that first appeared in the Lower Cretaceous 2). He further distinguishedthe morphology and includes two families, the Gavialidae and of G. kirtlondicus from the other recognized the Crocodylidae. The gavials are presently restricted to the species referredto the genus. In G. kirtlandicus, the skull is triangular in major rivers of India, Pakistan, and Burma shape(dorsalview)and lessthan twiceas long but earlier in the Cenozoic had a much greater as wide at the reconstructed base.The skull is distribution that included North and South moderatelybroad with the externalnareswell America, Europe, and Africa. - ) d a anterior and the orbits posterior and marginal. The specimen retains the right premaxillary,both nasals,the right lacrimal, both prefrontals,the frontal, the right postorbital, and most of the right squamosal.The superior temporal openingsmust have been well developed and roughly elliptical in outline,largerthan the orbital openings. Wiman's reconstruction of the palateshows a rather blunt snout region with ten teeth in the premaxillaries.The maxillariesbroaden posteriorlyand showspacesfor l8 teethalong the external margin of each maxillary. Wiman's palatal reconstructionis slightly broaderthan his dorsalreconstruction. The teeth in G. kirtlandicus are roughly conical but apically roundedand blunt (side view). Antero-posteriorly, the teeth are slightly recurved lingually with a distinct medial crestor ridge. Prominentapical-basal directedridgesor enamelrugaegivethe teetha wrinkled or corrugated appearance.The premaxillaryteethappearto be much smaller than the marginal maxillary teeth. The gaps for the pseudo-canines appearrelativelysmall in size. F-_--:+ I ml FIGURE 4-GENERALTZEDcEoI-oclc ueR showing units present in the area of Ojo Alamo Trading Post. Goniopholis kirtlandicus must have been found in the Kirtland Formation in the southern portion of T. 24 N., R. I I W. Abbreviations: Tn, Nacimiento Formation; Toa, Ojo Alamo Formation; Kk, Kirtland Formation; Kf, Fruitland Formation; Kpc, Pictured Cliffs Formation (after Fassettand Hinds, 1971; Clemens, 1973). The Crocodilidae includes the caimans, crocodiles, and alligators. The crocodiles, generally characterized by a narrower snout than the alligators, are relatively broadly distributed through tropical America, Africa, and southern Asia. The alligators, with a broader snout than the crocodiles. have a restricted range and occur only in the Western Hemisphere, except for a single species that survives in the Yangtze River of China. The Goniopholidae (and the included species Goniopholis kirtlandicus) belonged to the Mesosuchia. the crocodile suborder that appeared during the Jurassic. In overall appearance and probably in life habits as well, the goniopholids were similar to modern crocodilians. They attained substantial size, had broad but not especiallylong snouts, and substantial teeth. Their vertebrae, however, still retained primitive features and were amphicoelus (concave at either end). Although the palate was advanced in structure over earlier species,it was still more primitive than that in the Eusuchia. Towards the end of the Cretaceous the mesosuchians and many other reptilian lines of descent declined. Although mesosuchians survived into the Early Tertiary in some areas (Romer, 1966),competition with the expanding eusuchian lineage was most likely a factor in their decline. The mesosuchians were similar in form and probably in life habits to the eusuchians but lacked more progressive eusuchian features such as secondary palate development.In all likelihood, the eusuchians were better able to exploit the available habitats that both suborders occupied. A general cause for widespread extinctions may have operated late in the Cretaceous; certainly many archosauran lines terminated at the end of Cretaceous time. A combination of adverse environmental conditions and increasedcompetition from the eusuchiansmay have isolated mesosuchiansin ever-dwindling refuges,finally resultingin their extinction. Conclusions The recognition-intended by Wiman almost 50 years ago-of Goniopholis kirtlandicus as a component of Kirtland Formation (Late Cretaceous) faunas establishesthe presenceof both mesosuchiansand eusuchians in the San Juan Basin. Therefore, fossil collections should be reexamined to identify additional mesosuchian material. Material previously referred to Crocodyhls sp. or Brachychampsa? may actually represent Goniopholis, either G. kirtlandicus or other mesosuchian forms. Additional G. kirtlandicas material will enhance our understanding of the species.Similarly, other mesosuchian forms may be discovered. Whether mesosuchians also occur in the underlying Fruitland Formation and in the overlying Ojo Alamo Sandstoneis an important question. The highest occurrence of C. kirtlandicus in the San Juan Basin may have a significant bearing on the age relationships of the Ojo Alamo Formation and may be helpful in understanding the problem of Late Cretaceous extinctions. K u e s , B . S . , F r o e h l i c h , J . W . , S c h i e b o u t ,J . A . , a n d L u c a s , S . , 1 9 7 7 ,P a l e o n t o l o g i c a ls u r v e y , r e s o u r c e a s s e s s m e n ta,n d m i t i g a t i o n p l a n f o r t h e B i s t i - S t a r L a k e a r e a , n o r t h w e s t e r nN e w M e x i c o : A l b u q u e r que, Report to the Bureau of Land Management LeMone, D. V., and others, 1977, Applicant's p a l e o n t o l o g i c aal s s e s s m e natn d c o n t i n u i n g s t u d i e s for the proposed Star Lake coal mine: Chaco E n e r g y C o m p a n y , 1 5 2P . L e M o n e , D . V . , a n d o t h e r s , 1 9 7 9 ,P a l e o n t o l o g i c a l clearance assessmentand evaluation report, state lands, Bisti Project: Western Coal Company, 69 p. and appendices L u l l , R . S . , a n d W r i g h t , N . E . , 1 9 4 2 ,H a d r o s a u r i a n dinosaurs of North America: Ceological Society o f A m e r i c a , S p e c .P a p e r 4 0 , 2 4 2 p . O s b o r n , H . F . , 1 9 2 3 , A n e w g e n u s a n d s p e c i e so f Ceratopsia from New Mexico, Pentoceratops sternbergii: American Museum Novitates, no. 93, p H y L o c E N E r t c FIGURE 5-GerEnerRELATToN3p. s H I p so F c R o c o D r L r A N s u B o R D E R(sr e c t a n g l e sa) n d P o w e l l , J . 5 . , 1 9 ' 1 2 ,T h e G a l l e g o s S a n d s t o n e( f o r other archosaurian orders for reference. m e r l y O j o A l a m o S a n d s t o n e )o f t h e S a n J u a n Basin, New Mexico: M.S. thesis,University of A r i z o n a ,l 3 l p . AcxNowr-pocMENTS-This manuscript was 1973, Paleontology and sedimentation read critically by Stephen Hook and John m o d e l so f t h e K i m b e t o M e m b e r o f t h e O j o A l a m o Wright, New Mexico Bureau of Mines and Sandstone, ir Cretaceous and Tertiary rocks of Mineral Resources; Richard Holtzman, Mint h e s o u t h e r nC o l o r a d o P l a t e a u , J . E . 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P a p e r I 1 9 , 7 l p . 1 9 3 5 ,O n t h e r e p t i l i a o f t h e K i r t l a n d F o r m a t i o n o f N e w M e x i c o , w i t h d e s c r i p t i o n so f n e w s p e c i e so f f o s s i l t u r t l e s : U . S . N a t i o n a l M u s e u m , P r o c e e d i n g sv, . 8 3 , n o . 2 9 7 8 ,p " I 5 9 - I 8 8 H o p s o n , J . , 1 9 7 5 ,T h e e v o l u r i o n o f c r a n i a l d i s p l a y structures in hadrosaurian dinosaurs: Paleob i o l o g y ,v . l , n o . | , p . 2 l - 4 3 Kiilin, J., 1955, Crocodilia, in Traite de paleont o l o g i e , v . 5 , J e a n P i v e t e a u ,e d . : P a r i s , M a s s o n e t Cie, p. 695-784 Oil productionin New Mexico for the first threemonthsof 1980was 19,460,604 bbls,an increase of 38,873bblsoverthesameperiodin 1979.Gasproductionwas291,777,639,000 cu ft, an increase of 2,013,104,000 overthesame 1979 period (J. Ramey, Oil Conservation Division,N.M. EnergyandMineralsDept.). New Mexico Geology August1980 4l