Basin, Kirtland Formation, San

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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 Unpublished reports include Kues and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
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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 . F a s s e t t ,e d . :
nesota Geological Survey; Joseph Hartman,
F o u r C o r n e r s G e o l o g i c a lS o c i e t y , M e m . , p . l l l Department of Geology and Ceophysics, Unir22
R e e s i d e J, . B . , J r . , 1 9 2 4 , U p p e r C r e t a c e o u sa n d T e r versity of Minnesota; and Judith Schiebout,
tiary formations of the westernpart of the San
Louisiana State Universitv.
Juan Basin of Colorado and New Mexico: U.S.
G e o l o g i c a lS u r v e y ,P r o f . P a p e r 1 3 4 ,7 0 p .
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S i n c l a i r , W . J . , a n d G r a n g e r , W . , 1 9 1 4 ,P a l e o c e n e
History of nomenclature and stratigraphy of
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rocks adjacent to the Cretaceous-Tertiary bounAmerican Museum of Natural History, Bull., v.
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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
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