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AN ANN
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Datel
HEAL
MEMOTRS
OF THE
CARNEGIE MUSEUM:
1904-1906.
VOL. IL.
W. J. HOLLAND, Editor ,
MAR 4 1907
1GR092
PITTSBURGH.
PUBLISHED BY THE AUTHORITY OF THE BOARD OF TRUSTEES OF THE
CARNEGIE INSTITUTE.
C
coh ee
[Pesala ee INIA.
The publication of the Memoir by Dr. Arnold E. Ortmann upon “The Craw-
p)
fishes of the State of Pennsylvania,” concludes the Second Volume of the Memoirs
of the Carnegie Museum.
The paper on “The Osteology of Haplocanthosaurus” by Mr. J. B. Hatcher
was the last paper from his pen to appear in the publications of the Museum. He
died on July 3, 1904, and the sole responsibility for the supervision of the Memoirs
and Annals of the Museum devolved upon the writer, who without any assistance
has endeavored to perform the combined task of proof-reader and editor of the pub-
lications, a task which, while in some respects pleasant, has laid a great burden
upon him, taxed as he has been by the care of several departments of activity in
the museum, and the general supervision of the whole.
The pleasurable and consolatory reflection always remains, that by these publi-
cations very important additions are being made to the sum of human knowledge.
The first Memoir in Volume IV. upon “ Early Chinese Writing” has been pub-
lished, and the first Memoir in Volume III. upon the Archeology of the Pacific
Slope of Costa Rica by C. V. Hartman is in press and shortly will be given to the
public.
W. J. Hoxanp.
December 1, 1906.
ill
‘a Nites
page ay
A aw
(Mey oy
For
TABLE OF CONTENTS.
PREFATORY NoTE
List or Fiaures 1n TExt
List oF PLATES : : A
MeEmorr I. Osteology of Haplocanthosaurus. J. B. Hatcher
Memorr II. Osteology of Baptanodon Marsh. C. W. Gilmore
Memorr III. Fossil Avian Remains from Armissan. C. R. Eastman
Memorr’ IV. Description of New Rodents. O. A. Peterson
Memorr VV. The Tertiary of Montana. Earl Douglass ‘
- Memorr VI. The Osteology of Diplodocus Marsh. W. J. Holland
Memorr VII. The Osteology of Protostega. G. R. Wieland .
Memoir VIII. New Suilline Remains from the Miocene of Nebraska. O. A.
Peterson
Memoir IX. Notes on the Osteology of Baptanodon. C. W. Gilmore .
Memorr X. The Crawfishes of the State of Pennsylvania. Dr. Arnold E.
Ortmann
Index
JEJSIE Oils" MUG Isa Ss) TOs) IMS CIe:
PAGE.
Diagram of portion of bone-quarry near Canyon City, Colorado, showing positions of Haplo-
canthosaurus priscus and H. utterbacki when found .............-.2.2 0 eee eects 4
Diagram of portion of bone-quarry near Canyon City, Colorado, worked for Carnegie Museum — 5
Sixth (?) dorsal of type of Haplocanthosaurus priscus seen from right side.......... 10
EhirsieWorsaluotelaponiscis seen tromericint sides a) 1 ae ciee era ae 10
Superior view of sacral, anterior caudal, and posterior dorsal, or lumbar, of Menopoma
CHG WG UCTISIS) 558 i) ol ocd eiteren cena upc See eee eee Sl ae ee tenn a a nadie eed ola ain 21
Superior view of sacrum, anterior caudal, and posterior lumbar or dorsal of Iguana tuberculata 21
Chevron of Haplocanthosaurus between caudals 8 and 9, side view........0....0.-.-055- 23
Chevron of Haplocanthosaurus between caudals, posterior view........ 0.020220 e eee eee Py)
Chevron of Haplocanthosauwrus between caudals 13 and 14, side view..............-..--- 23
Chevron of Haplocanthosaurus between caudals, posterior view... .....- 0+. 200+ 00e ress 23
Anterior view of second (?) rib of right side of Haplocanthosaurus.... 0.050000 0020 eee eee 24
Anterior view of fourth (2) rib of left side of Haplocanthosawrus.... 0.06.0. 0000 eee cee 24
Anterior view of thirteenth (?) rib of left side of Haplocanthosaurus. 0.0.0... 000 eee eee 24
Left femur of Haplocanthosaurus priscus, front view......... ++ 0+. . 002-022 ee 27
Sacrum of Haplocanthosaurus utterbacki, seen from right side........0...2. 20000 eee 36
Anterior view of disarticulated parapophyses (sacral ribs) of Haplocanthosaurus utterbacki.... 36
Rostenionaviewsotmlettiscapula oil anterbackinniaar in = see sane. title aen ere 42
Bcretna laviewaotslettyscapulaoimlarutterbackine tl. ys) a eee eae ae ae oar ee 42
external pviewaormohtreoracoidl ois utlerbackiness-)) ats yeasts as 2a een ee 42
Centrum of fourth sacral of H. utterbacki seen from right side........-....... 022-005 51
Dcapulecnole C chrosatmisitonguss Owens aetna cine aaa cet oo lemons sear -ee = ee TE ate 52
Comencl OF Ognonairis (Ongiay OVGis ann sopemeeeeunoeemadsasaatpedcebouobbgcass co 53
Photograph of footprint in Jurassic sandstone, near Canyon City, Colorado............... 61
Photograph of ripple-marks on surface of Jurassic sandstone, Big Horn Mountains, Wyoming 62
View of Atlantosaurus beds, entrance to Garden Park, near Canyon City, Colorado........ 63
The “ Nipple,” showing in foreground trench cut by Professor Cope in collecting Dinosaur
TRETTEDINNS 6c Gaara octet aa Orc thay ers Glee Pea ERE GE ERE Re ECE Sera MIC ammIN Ente With nlp ola! a5 8 64
Biss CEHmenirance toy COpexqUannky ete sce cies ccciae cca o 4a seals obennlen keys tuate es? cae nee Rei Rare 65
View from near Cope quarry with the “ Nipple” in foreground, Cooper Mountain in the dis-
ienaee, Cauclen letyAke logy tae CIN VO”G5e050ce00805G0c0000000G0C000GnG0000 He 66
Inaxhing aia! wlban, ow JDiyailocltenn. 66 code so cenoecsconcadnaaeeobnuos I teeny 73
Ep POsednclaviclesols Wiplodoctss: seni s ay shcneyes oi oeiyo/eie siete ala ch pene perenne ey eavaterey S/ ae ae LSE 73
Vill LIST OF FIGURES IN TEXT.
Cross-section of upper and lower jaws of Baptanodon........... 00000000 ceecevseceee RE
CRISES TOM Gt lone jer OF LeAMOC Ds o503ens0000008050000000550n05n00005 54558 95
Internal view of posterior end of left mandibular ramus of Baptanodon discus............ 97
Internal view of left articular of Baptanodon discus............-000- 0 esses eevee eee: 98
AmberiOrnview, Of lettanticulaniotesaptancdonsdiscus nner nna eten ene eee 98
JPOP tool Or Jxanomoclom CHSCUIS >o.c0050%9090300 c00 5000 00056000053040" sooo WY)
Cross=section\o1 toothvotws a/ptanod onmdisctiss = arn ety eee eae ee 99
IAN: WayrRoanall OF JRayneMOtlOn CHIMS. 0025000096 08o0%250s0000500 4001 0000 bos 00008 100
Anterior view of atlas and axis of Baptanodon discus..................0..0.00-.005- 102
Lateral view of atlas and axis of Baptanodon discus...................-+..+.-- eae 102
Posterior view of atlas and axis of Baptanodon discus ............ 0.00022. cece esse 103
Left half of paired neural arch of atlas of Baptanodon discus ............ 0.24 -2.200.. 104
Superior view of anterior cervical of Baptanodon discus ............... 0.00 eee esses 104
Anterior view of anterior cervical of Baptanodon discus .............-2-.22++2+--0-- 104
Cross-section of an anterior dorsal centrum of Baptanodon discus..........-....+-5-+-- 105
Cross-section of dorsal centrum of Baptanodon discus ............ 05+ I Mie ek Ratan 106
Cross-section of posterior dorsal or anterior caudal centrum of Baptanodon discus......... 106
Lateral view of centra from different regions of Baptanodon discus ............-.++.-.- 106
End and lateral views of an extreme caudal of Baptanodon discus............... REA vary 107
Posterior view of a right dorsal (?) of Baptanodon discus.......... .........:......... 108
Ventral view of pectoral girdle of Baptanodon discus as found in matrix................ 109
Interior view of interclavicle of Baptanodon discus .... 1.01... 2. eee ee ee ee 113
Superior view ot tore paddleot Baptanodon manshi.. 1.) 2.02 222 essa oe oo 114
Dorsal view of humerus\of Baptanodon marshi. 1.23.22 25-3 20.5222 ees ee 115
tefighinds(?)\spaddlevotssaptanodonidisctissee ir eee et eee ee re eee 115
Atlas, axis, and third and fourth cervical of Baptanodon marshi................0...55. 122
Crown view of superior grinders of Steneofiber fossor.........................002-0-- 141
Crown view of inferior grinders of ISLENCOPDERN [OSSOT ey arse can ese Se eel na ee aa 141
Fore-limb of Steneofiber fossor in position FOR Sea NG: SeAU RL RS MR EMA Re CaaS 158
SAAN 20) LOG POOR CONG ie ata 6. oti cao Me AE OIC He SIGHS HO Lovo OL GRGS OS © 50.0 '51S Hes SDIsO Gd 5 6 5s 166
Field sketch of a weathered rhizome containing type of Steneofiber barbourt ............. 185
Weathered portion of a Demonelix containing skeleton of Steneofiber fossor ............. 186
BlormistotDsemione lisccrje ee wes tnskene stesso od oe eae RCI a RCT er cee hota ne SE 187
Spiralwandspartrorechizomesotell) mone lien ye Pree er ere een ee ee ere 188
AN neryihy Cli lente Dios lNTARONY 40005500 cdc coe oD Dad E oo H SSDS FN DOUIS GLO UsOODbSDE 189
Diagrambotcastiotmanioldiburrowsot rane Doge rorn: Heer ere nee inEes ac se oe 189
iWndergroundifoxtresstotgimole Clalpanewnopaca) armprar ree eae ere ener ee ye ees 189
Skull and anterior cervical vertebrae of Diplodocus Curnegiei Hatcher as placed in restoration
at Britishe Miiseu msi). sil iia. Speese: hel sic eee ei et Re ote ean rere ae tee Rog eae 228
LIST OF FIGURES IN TEXT.
PAGE.
Sketch by Miss Alice B. Woodward showing supposable attitude of Diplodocus........... 229
Side ww oF skull ar JOWNeCClaGis ea. See pee ane etaone den oceans guioucooe uot cnn us = 230
Rosterionnview,ofsthelbackotskulltofmDiplodocusis acs se ae ey ieee ts ne raraee 231
Lateral view of posterior portion of skull of Diplodocus. ............-...-2.00-200-+-- 233
Superior view of posterior portion of skull of Diplodocus...........--+....+0020+-05- 235
Anterior view of posterior part of skull of Diplodocus.......... .........2...2+-... 236
Diagram showing relation of the bones at outer margin of supratemporal fossa of Diplodocus. 237
Inferior view of posterior part of skull of Diplodocus..................5...2..-0045. 241
Posterior view of intercentrum of atlas of Diplodocus................0 020.00 -- 0020s: 247
Anterior view of intercentrum of atlas of Diplodocus ...............2 0000 se eee eee eee 247
Inferior view of intercentrum of atlas of Diplodocus ...............+.2++- eee eres 247
External view of left neural arch of atlas of Diplodocus............................. 247
Internal view of right neural arch of atlas of Diplodocus.... .......+.+-++--++-.5.0-- 247
Imreriongvie wae te Odontordieenmyacwinenseyeyereree tiie cs eo sree -atsiel el ssane sd tricia Cate Tey es eee ae 248
Stperiornviewsote Od onto tacmesey ernie ecient seca ei tardeieitere el eaies sane alates sok ae eR Ieee 248
Rosterionnviews Ol" Odontordes jarvis sn: ctee ca ie chcl/acyalemcoe ts, susseav sce ci tay tye suas ecaaers cheneetamenen sre 248
AriterioraviewnotmOdontoid gece nase oeclae oye a riety velo eke ote @ ee yen eae 248
External, internal and inferior views of supposed rib of atlas of Diplodocus in American
MuscumyorNatural SEU Storyp ht crcescttse cucin cnet ea colons & lepchaiige 4: e st tne ates Meera 259
External, internal and inferior views of supposed ribs of axis of Diplodocus in American
Museumho te Na toralttistony irra eae tertcieiaey cra ts sieens ceshirsia earniciesccacecnia o eease eeee 250
Sketch of manner in which the two cervical ribs of Diplodocus may have functioned in con-
MACHO Kymdn HS CHES EIG! Ehdkonon bo dag oneGand oom ens Sola) Oboe Someone me oc 250
Outline drawing of a series of posterior caudal vertebrae of Cetiosawrus leedsi Hulke ...... 254
Vertebrie of posterior part of tail of Cetiosawrus leedsi Hulke, as exhibited in the British
IMI Se Uma spce eee ale cones heen rr Mncd tira euaget Visas eleaaian Biss sla eration col ostiae ter ues 254
Photograph of the two sternal plates and the supposed clavicles as provisionally and tempor-
arily placed by the author in the restored skeleton in the British Museum........... 257
Lateral view of the sternal plates and supposed clayicles at British Museum............. 258
Superior view of pectoral girdle of Crypotoclidus omoniensis............ 0000 ee eee 259
Anterior view of pectoral girdle of Cryptoclidus owoniensis ,..........0 0000 eee eee eee 260
Dice wossupposed: clavicles\ ote Mzplodocusiy:.\s cru ote 2 voi) e = 5) she's sino as = 2 eis) eeieea ciate 261
Sections of shafts of supposed clavicles of Diplodocus.....0 9 6... eee eee eee ee eee 262
Protostega gigas — superior view of skull with lower jaw and hyoid, right anterior border
Gimcarapace and aright torent pera rs: eur iparssr foley Geist. aicicie cer ete ey 64) soaks ener aya eee 282
Protostega gigas. Carapace less the nuchal and marginal series, inner view ............- 286
Dntanviowsotsaslarceshum enusyolgeenolostegalgigasier tre) -ine eae eke etait 287
ibiehieshoulderandsiipper, of erotosiegangigas 9. «5.2%. 22 se oes vole sun eo Oe 288
ich pelvicsoirdle andetlip per Otselenotosieg a gigaser ria Mec) acl ilies ec reeerre 291
xX LIST OF FIGURES IN TEXT.
PAGE
Wngualliphalangeslot -enotostegalgigass eee eee ener ek tee Rae 292
ILS aR ae hyo OF GemoGodys UHRA oocccccbvcoadovvab000c000000e00005005000% 293
Fore and hind flippers of Colpochelys kempi in the United States National Museum........ 294
Oblique internal view of left mandible of Thinohyus (B) subequans Cope ..........-..-- 307
External view of left lower mandible of Thinohyus (B) subequans Cope................ 307
Superior view of left lower mandible of Thinohyus (B) subeequans Cope.................. 307
Anterior portions of upper and lower jaws of Thinohyus siouwensis ....... Ee teen as 317
Internal view of the right mandibular ramus of Baptanodon natans ................--. 327
Cross-section of posterior end of left mandibular ramus of Baptanodon natans ........... 327
Cross-section) on scleroticiplatesiotaB. natansin an nae ace eine eee cae 0 O28
Lateral view of a section of the anterior part of rostrum of Baptanodon natams.......... 329
Cross-section near middle of snout of Baptanodon natans.... 0.2.0... 00. eee ee ee 329
Atlas, axis, third and fourth cervical vertebree of Baptanodon natans..............-.-+- 330
Lateral view of intervertebral disk of Baptanodon natans.........0 20.04.0022 0002 02s 330
View of the proximal end of humerus of Baptanodon natans........ 0... 2-00.00 0 ee ee 331
Wi natofeBaptanodoninatanss tas wc iiss e rors sei ses ees er suet ns cere nseaN eee iy ata ere 331
RadiustotmBaptanodon matansmrcer rae erie are hho aes cir ara in ae Aan sare eee ene 331
External view of acetabular end of ilium of Baptanodon discus........-....-.0.0-.005. 332
Atlas, axis, third, fourth, fifth and sixth cervical vertebree of Baptanodon robustus........ 33
Lateral view. of anterior or median dorsals of Baptanodon robustus ..............02045- 334
XXIV.
XXYV.
XXVI.
XXVIII.
XXVIII.
XXIX.
JbJSIE Qua PILLAI aS,
Presacral Vertebree of Type of Haplocanthosaurus priscus.
. Vertebre of Type of Haplocanthosaurus utterbacki.
. Nineteen Anterior caudals of Haplocanthosaurus priscus.
. Pelvis of Brontosaurus ; Pelvis of Diplodocus ; Pelvis of Haplocanthosaurus. All
seen from left side.
. Inferior, Anterior and Posterior views of Pelvis of Haplocanthosaurus priscus.
. Restoration of Diplodocus carnegiet.
. Diagram showing Position in which the Elements of the Specimen of Baptanodon
discus Marsh were found Imbedded in Concretion.
. Side view of Skull of Baptanodon discus Marsh.
. Skull of Baptanodon discus.
. Side view of Skulls of Baptanodon discus.
. Posterior Views of the Skull of Baptanodon discus.
. Anterior, Dorsal, and Ventral Views of Pectoral Girdle of Baptanodon discus.
. Taoperdix keltica Kastman.
. Taoperdix keltica Kastman. Restoration.
. Taoperdix pessieti (Gervais).
XVI.
XVII.
XVIII.
XIX.
XX.
XXII.
XXIT.
XXIII.
Taoperdix pessieti (Gervais).
Fossil Rodents from the Upper Miocene, Harrison Beds.
Fossil Rodents from the Upper Miocene, Harrison Beds.
Restoration of Steneofiber fossor.
Mounted skeleton of Steneofiber fossor.
Microscopic Sections of Cellular Structures in Deemonelix.
Lower White River Mammals. Ictops and Xenotherium.
Side view of Skulls of Diplodocus longus Marsh, Preserved in the United States
National Museum.
Top view of Skulls of Diplodocus longus Marsh, Preserved in the United States
National Museum.
Back view of Skulls of D. longus Marsh, United States National Museum.
Photograph of Skull of Diplodocus Preserved in the American Museum of Natural
History.
Posterior and Anterior View of Back Part of Skull of Diplodocus.
Superior view of Back Part of Skull of Diplodocus.
Posterior Caudal Vertebree of Diplodocus carnegie: Hatcher.
xi
Xil
PLATE.
OO:
XXXL.
XXXL.
XXXL.
XXIV.
XXXYV.
XXXV I.
XXXVI.
XXX VII.
A.
B.
XXXIX.
XI.
XUI.
XLII.
XLII.
LIST OF PLATES.
Restoration of the Skeleton of Diplodocus carnegiei Hatcher.
Protostega gigas Cope. Shoulder Girdles, Pelvic Girdle, Hyoids.
Protostega gigas Cope. Forearm, Hand, and Foot.
Dermochelys coriacea.
Thinohyus siouxensis Peterson. Right side of Skull and Lower Jaws and Superior
View of Mandible.
Superior and Palate Views of skull of Thinohyus siouxensis Peterson.
Side View of Skull of Baptanodon discus Marsh.
Superior and Inferior Views of Baptanodon discus.
Pectoral Girdle of Baptanodon robustus.
Cambarus obscurus, C. diogenes, and C. carolinus reproduced by the three-color
process.
Cambarus bartoni, C. bartoni robustus, C. limosus, and C. monongalensis, reproduced
by the three-color process.
Anatomical details of structure of Pennsylvanian Cambari.
Chel of Pennsylvanian Cambari and burrows of D. bartoni and C. carolinus.
Burrows of Pennsylvanian Cambari. .
Preglacial Monongahela River, and present range and distribution of species.
Map of Pennsylvania showing distributional areas of crawfishes.
OSTEOLOGY OF HAPLOCANTHOSAURUS,
WITH
DESCRIPTION OF A NEW SPECIES,
AND
REMARKS ON THE PROBABLE HABITS OF THE SAUROPODA AND
THE AGE AND ORIGIN OF THE ATLANTOSAURUS BEDS.
By J. B. Harcurr.
ADDITIONAL REMARKS ON DIPLODOCUS.
By J. B. Hatcuer.
[Reprinted from Mrmorrs CARNEGIE MusEUM, Volume II., No. 1, November, 1903.]
LAncasren, PA.
bok “stay iy I
Pe : i
——
Ag sy Rae
* z s aivel dee an
se
OSTEOLOGY OF HAPLOCANTHOSAURUS,' WITH DESCRIPTION OF A
NEW SPHCIES, AND REMARKS ON THE PROBABLE HABITS
OF THE SAUROPODA AND THE AGE AND ORIGIN
OF THE ATLANTOSAURUS BEDS.
By J. B. Harcuer.
The present paper is the third of a series of memoirs based on the fossil verte-
brata in the collections of the Carnegie Museum. ‘These memoirs, prepared either
directly by the curator of the Department of Vertebrate Paleontology, by his assis
tants, or others under his general direction, will continue to appear at irregular
intervals. ‘Their chief purpose will be to describe in detail and to illustrate with
fidelity some of the more important fossil skeletons in the collections. While in
every instance these papers will be based on material belonging to the collections of
the Carnegie Museum, for the sake of completeness, wherever other and supplemen-
tary material is accessible in the collections of other museums it will be utilized and
the fullest credit will, in all such instances, be given for such favors. The preca-
rious conditions attending the preservation, fossilization, and final recovery of the
skeletons of extinct vertebrates have necessarily been such as to render the occur-
rence of really complete skeletons conspicuously rare. This is especially true of the
gigantic Sauropoda, and notwithstanding the very large collections in several of our
leading museums, it is still possible to determine anything like the complete oste-
ology of the different genera only by selecting the best preserved skeleton of each as
a basis and supplementing this from material pertaining to the same genus but to
other skeletons and belonging to the same or other museums. Owing to the cordial
relations at present existing between the various museums of this country, aided by
‘Proce. Biol. Soc. Wash., Vol. XVI., 1903, pp. 1 and 2, and p. 100.
2 MEMOIRS OF THE CARNEGIE MUSEUM
the exceptional energy with which a few institutions are increasing their collections
our knowledge of the structure and relations of a considerable number of known
dinosaurian genera has been materially increased during the past few years, while
occasional discoveries of entirely new forms have been announced.
It appears somewhat remarkable however that a Sauropod dinosaur of such
gigantic size and showing such distinctive generic characters as does Haplocantho-
saurus should have been discovered so recently at the exact locality, near Canyon
City, Colorado, so long worked and rendered classic by the researches of the late
Professor Othniel Charles Marsh. This discovery may be taken as an indication not
only of the great wealth of this particular locality in the remains of the Dinosauria
but of the great diversity that existed in the reptilian life of this region in Jurassic
times. For since this single bone quarry, restricted in area to a few hundred square
feet and with the bone-bearing horizon not more than three feet thick vertically, has
already produced representatives of at least a dozen genera and species and twice or
thrice that number of individual skeletons it would seem difficult to overestimate the
wealth of the reptilian fauna of this region in Jurassic times or to exaggerate the
total number of genera and species that must have existed throughout the period of
time required for the deposition of the several hundred feet of sandstones and shales
that here constitute that formation and imbedded within which we may still hope
to find remains of additional genera and species pertaining to that peculiar but long
since extinct group, the Dinosauria.
For the material upon which the present paper is based we are indebted first
of ali to the generosity of Mr. Andrew Carnegie whose munificence made it
possible to carry on the excavations necessary for its recovery. To the skill,
energy and patience however of Mr. W. H. Utterback we are directly indebted
for its recovery from the hard, almost granitic sandstones in which the bones
lay buried beneath many feet of other sandstones and shales only a little less refrac-
tory than those actually containing the fossils. After these superincumbent sand-
stones and shales had been removed over a considerable area the actual and more
difficult work of developing and recovering the fossil bones was begun. These, as
has been stated above, lay buried in a thick stratum of heavily bedded and hard
sandstone. Not only was this sandstone for the most part extremely hard but it
was also considerably fractured in such manner as greatly to increase the difficulty
encountered in taking up the bones in a proper manner. All difficulties were how-
ever met and overcome by Mr. Utterback with commendable patience and inge-
nuity, and the different blocks were received at the paleontological laboratory of the
museum with all the vertebree and other bones in each block still in their original
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 5)
positions relative to one another. While aided by diagrams of the quarry, repro-
duced here in Figs. 1 and 2, and the proper marking of each block as it was taken
up, it is now easily possible to assign the different blocks to their proper position in
the quarry and thus to determine with accuracy the relative positions of all the
different bones as they lay imbedded in the rock.
In the laboratory the bones have been very carefully and skillfully freed from
the matrix under the direction of Mr. Arthur $. Coggeshall as Chief Preparator
assisted by Messrs. W. H. Utterback, L. S. Coggeshall and A. W. VanKirk.
When freed from the matrix the bones were all faithfully drawn by Mr Sydney
Prentice, draughtsman in the Paleontological Department of this Museum.
The type No. 572 of the present genus consists of the two posterior cervicals,
ten dorsals, five sacrals, nineteen caudals, both ilia, ischia and pubes, two chevrons,
a femur and a nearly complete series of ribs, all in an excellent state of preservation
and pertaining to an individual fully adult as is shown by the codssified neural
spines and centra.
Posrrion oF THE DirrErENT Bonks As THEY Lay IMBEDDED IN THE QUARRY.
The pelvis, sacrum, left femur and nineteen anterior caudals were the first por-
tions of the skeleton discovered. They lay in the position shown at A. 572 in the
diagrams of the quarry shown in Figs. 1 and 2. The ilia, ischia and pubes still oc-
cupied approximately their normal positions relative to the sacrum, and the femur
was directed backward and downward, with the head removed about two feet from
the acetabulum. The anterior caudal was displaced from its normal position rela-
tive to the distal extremity of the sacrum, but the succeeding eighteen caudals were
interlocked by their zygapophyses. ‘The two chevrons lay as shown in the diagram,
approximately in position, with caudals eight and thirteen. I personally assisted
in taking up this portion of the skeleton and am therefore somewhat familiar with
its appearance as it lay in the quarry.
At a distance of about twelve feet but on the same level as the pelvis and bones
above mentioned, were found the nine posterior dorsal vertebree shown at B. 572 in
the diagrams of the quarry. These were all interlocked by the zygapophyses and most
of the ribs were still in place. The last of this series agrees very well in size and
general appearance with the first sacral of the series found at A. 572, and there would
seem no good reason for assuming that the two series pertain to other than one and
the same skeleton, though, of course, this cannot be absolutely demonstrated, but
the characters exhibited by the two series demonstrate that they pertain to the same
species at least and I have little doubt but that they belong to the same individual.
4 MEMOIRS OF THE CARNEGIE MUSEUM
With the ninth from the posterior of this series of vertebrae there was an interrup-
tion, and the three vertebrae shown at C. 572 were found closely adjacent to the an-
Scalé about peel lo 1 inch
Fic. 1. Diagram of west end of that portion of bone quarry near Canyon City, Colo., worked by W.
H. Utterback, showing the positions in which the types of Haplocanthosaurus priscus (No. 572, A, B, C)
and H. utterbackii (No. 879) were found. The shaded bones pertain to a different genus. A. 572 femur,
pelvis, sacrum and nineteen anterior caudals ; B. 572 nine posterior dorsals ; C. 572 first dorsal and last
two cervicals.
terior of the nine dorsals mentioned. ‘These three vertebree were interlocked by their
zygapophyses and consist of the first dorsal and the last two cervicals. They evi-
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 2)
dently pertain to the same series as the nine dorsals and the differences in the spines,
positions of the rib facets, ete., demonstrate that a number of dorsals are missing
between this first dorsal and the anterior of the series of nine posterior dorsals ;
while the remains of a second skeleton pertaining to a different species of the
same genus fixes the number of missing dorsals at four. This would place the num-
ber of free dorsals in the present genus and species at fourteen instead of ten, the
probable number in Diplodocus and Morosaurus. It is possible, however, that in the
Dinosauria the number of dorsals may vary in different individuals within the same
species as is well known to be the case in numerous instances in the Mammalia.
The bones within the dotted lines in the upper left-hand corner of the first dia-
gram (Fig. 1) for the most part pertain to and constitute the type of a new species
of Haplocanthosauwrus, which will be described later in this paper. The shaded bones
Scale about 12 feet lo 1 inch
Fic. 2.— Diagram of that portion of bone quarry near Canyon City, Colo., worked by Mr. W. H.
Utterback for Carnegie Museum. The lower irregular line shows limit to which quarry had been worked
by the Jate M. P. Felch for Professor Marsh.
within the dotted lines and the scapula and coracoid beneath pertain to one or more
genera different from Haplacanthosaurus. The relative positions of these bones as
they lay imbedded in the sandstones are well shown in the diagrams and will be re-
ferred to in detail, when we come to describe the species of which they form the type.
The quarry from which these remains were recovered is the one long worked by
Professor Marsh. It is situated on the west side of Oil Creek (Four Mile Creek) at
the entrance to Garden Park and some nine or ten miles east by north of Canyon
6 MEMOIRS OF THE CARNEGIE MUSEUM
City, Colorado. The horizon is in the Jurassic” and some 100 to 150 feet above
the summit of the red Triassic? sandstones. It is I believe a decidedly lower hori-
zon than the dinosaur beds near Morrison, Colo. ; Como, Little Medicine Bow and
Sheep Creeks, Wyoming or Piedmont, South Dakota.
Tn the diagram of the quarry shown in Fig. 2 the bottom line shows the limit to
which the quarry had been worked by Professor Marsh while the area above this
line is that worked by Mr. Utterback for the Carnegie Museum.
DerscrRIPTION OF THE Type (No. 572) or HApLocanrHosaurus Priscus.
The Vertebree.
The Cervicals (Plate I., Figs. C15 and 14).— Only the last two cervicals were re-
covered. Fortunately these, together with the first dorsal, were still interlocked by
their zygapophyses and thus the actual position of these three vertebree in the ver-
tebral column can be definitely determined. They were somewhat crushed and
distorted, but considering the hard and fractured nature of the sandstone in which
they were imbedded they are in a very good state of preservation and remarkably
complete. These vertebree are rather low, broad and short for the posterior cervi-
cals of a Sauropod dinosaur of such dimensions as is indicated by the remains of
the present skeleton and suggest a reptile with a neck which, though, of moderate
length, was decidedly more abbreviated than was that of Diplodocus, a contempora-
neous but more highly specialized Sauropod.
The Fourteenth? Cervical (Plate I., Fig. C 14).— Assuming that there were the
same number of vertebree in the cervical series of Haplocanthosaurus as in Diplo-
docus, the first of the series of vertebree now under consideration would correspond
to the fourteenth cervical. It is not improbable, however, that the number of cer-
vicals in the present genus was less than in Diplodocus. Hence, I have interrogated
the numerical position of this vertebra in the cervical series, although as already
stated, there can be no doubt of its being the last but one of that series.
The centrum is strongly opisthoccelous and with the transverse diameter exceed-
ing the vertical, though these dimensions have doubtless been somewhat altered by
pressure. The sides of the centrum are invaded by long and deep pleurocentral
cavities® separated only by a thin median septum. These cavities are extended for-
ward into the base of the ball of the centrum while posteriorly they are only sepa-
rated by a thin plate of bone from the cavity for the ball of the succeeding vertebra.
* By some considered as Lower Cretaceous.
*For an explanation of the names applied to the various cavities, Jamine, etc. of the Sauropod ver-
tebrze, see the author’s paper on Diplodocus, Mem. Car. Mus., Vol. I., No. 1, pp. 16-19.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS i
In this manner the centrum of the vertebra is reduced to superior and inferior hori-
zontal plates united by a vertical median septum or plate. At the posterior ex-
tremity these plates expand into a deeply excavated disk which forms the cup for
the succeeding vertebra while at the anterior extremity they unite to form the ball
of the centrum. A cross-section of the centrum midway between the anterior and
posterior extremities is irregularly I-shaped and somewhat suggestive of that of an
I beam in structural materials. The inferior surface of the centrum is broad and
flat, much expanded posteriorly and moderately expanded anteriorly where at a
point a little back of the ball it gives rise to the cervical rib. The cervical rib is
firmly codssified both with the centrum below, through the intermedium of a para-
pophysis, and with the diapophysis above. There is a short anterior branch of the
cervical rib and a longer posterior one. The latter stops short of the posterior ex-
tremity of the centrum. It is proportionately broader and stronger than in Diplo-
docus carnegii but decidedly shorter and less robust than in Brontosawrus excelsus.
Seen in front this vertebra appears rather low, with broadly expanded cervical
ribs and prezygapophyses. There is a single supraprezygapophysial cavity and two
infraprezygapophysial cavities separated by a median septum formed by the union
of the horizontal laminze of opposite sides and supported below by the superior wall
of the neural canal. In the vertebra under consideration the greater portion of this
septum has been lost. It is restored in plaster, and in the drawings the restored
parts are indicated by broken lines in the shading. As shown in the drawings the
neural spine is also absolutely simple instead of deeply bifurcated as are the spines
of the vertebree of this region in all other known genera of Sauropod dinosaurs
wherever it has been possible to determine their character. The neural canal is
rather large as compared with that in Diplodocus.
Seen from the rear the neural canal is nearly circular and appears as if sunk into
the superior surface of the centrum. The postzygapophysial laminee each send for-
ward a broad thin plate. These unite with the neural spine and enclose a very deep
suprapostzygapophysial cavity while below as in front there are two small but deep
infrapostzygapophysial cavities separated by a median septum.
The diapophyses are only moderately expanded and they are braced antero-
posteriorly by the horizontal laminze and inferiorly by the inferior branches of the
diapophysial laminze which are very short and almost perpendicular. There is no
superior branch of the diapophysial lamina. The posterior branch of the horizontal
lamina runs obliquely upward and backward from the diapophysis to the posterior
zygapophysis, thus giving additional support to the latter element. Another lamina,
horizontal in position but homologous with one of the oblique lamine, runs directly
8 MEMOIRS OF THE CARNEGIE MUSEUM
backward from the diapophysis nearly to the posterior border of the centrum.
There are deep and well-defined post-, pre-, supra- and infradiapophysial cavities.
The Fifteenth ? or last Cervical (Plate I., Fig. C 15).— This vertebra differs from
the one preceding it in being a little shorter and with more widely expanded neu-
ral spine and cervical ribs. The pleurocentral cavity is less extended posteriorly
than in the preceding cervical and its bottom is interrupted by an oblique and an
intersecting lamina. There is a shallow infracentral cavity on either side of the
mec..in line on the inferior surface near the anterior end of the centrum. There is
a single infraprezygapophysial cavity. The neural spine is absolutely simple as in
the preceding cervical. The postzygapophyses are higher and the posterior branch
of the horizontal lamina consequently more nearly vertical than in the preceding
vertebra. The anterior branch of the horizontal lamina has the margin somewhat
expanded as shown in Plate I., Fig. C 15, indicating that this vertebra gave some
support to the scapula.
The First Dorsal (Plate I., Fig. 1). — Fortunately as has already been stated this
vertebra and the two preceding were still closely interlocked by their zygapophyses
when discovered in the quarry. They were taken up in a single block of the enclos-
ing sandstone and were received at the musenm still occupying their original posi-
tions relative to one another. In consideration of these facts there can be no ques-
tion regarding the exact position of these three vertebree in the vertebral column.
That the vertebra now under consideration was a dorsal is conclusively shown not
by the presence of tubercular and capitular rib facets showing that it supported on
either side a free rib, for there are in our collections of sauropods, skeletons of other
dinosaurs fully adult but, with the posterior cervical, bearing free cervical ribs
articulating by both tubercular and capitular facets as do the ribs of the dorsal re-
gion. ‘The character in this vertebra distinguishing it as a dorsal is the broadly ex-
panded external border of the anterior branch of the horizontal lamina. This ele-
ment has been thus modified in this and the succeeding dorsal, no doubt, as is
known to be the case in Diplodocus to give greater surface for the attachment of the
powerful muscles necessary for the support of the scapula. That this was the first
and not the similarly modified second dorsal is conclusively demonstrated by the
fact that it was found interlocked by its zygapophyses with the last cervical.
This vertebra is essentially complete, although the form of the centrum has been
considerably altered by crushing. In the accompanying drawings this distortion
has been eliminated as much as possible. The length of the centrum is noticeably
less than that of the last cervical and the antero-posterior diameter of the pleuro-
central cavity is greatly shortened. The floor of this cavity is interrupted by neither
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS by)
oblique nor intersecting laminee. There is no infracentral cavity. The capitular
rib facet is nearly circular and slightly pedunculate. Its position is beneath the
anterior border of the pleurocentral cavity.
The neural arch is decidedly higher than in the posterior cervicals. The dia-
pophyses are more widely expanded and support at their extremities small triangular
tubercular rib facets which face outward and a little downward. These rib facets
are not pendant as they are in this and the two succeeding dorsals in Diplodocus.
The anterior and posterior zygapophyses are both somewhat more elevated than the
diapophyses and they are supported laterally by the anterior and posterior blades
of the horizontal laminee which are subequal and unite at an obtuse angle to form
and give support to the transverse process or diapophysis. Throughout about two
thirds of its length the external margin of the anterior blade of the horizontal lamina
presents a greatly expanded rugose surface, which no doubt served for the muscular
attachment of the scapula. From below, the transverse process is supported by the
short, rather slender inferior blade or branch of the diapophysial lamina which runs
obliquely downward and forward to unite with the superior branch of the prezyga-
pophysial lamina, while an extended and powerful oblique lamina runs obliquely
downward and backward, uniting with the lateral wall of the neural arch and giv-
ing additional support to the transverse process. The pre-, infra- and postdiapo-
physial cavities are all deep and well enclosed, while the supradiapophysial cavity
is shallow and left open anteriorly.
Seen from in front this vertebra appears low with the transverse processes, zyga-
pophyses and neural spine greatly expanded. The neural spine is low and extremely
broad. The apex on one side is injured. It is quite simple, not at all bifurcated
and with a broad, rugose, median surface. ‘The anterior aspect of the spine is strongly
convex transversely throughout its entire length. The articular surfaces of the an-
terior zygapophyses are elliptical in outline, with the transverse diameter the greater.
Between the anterior zygapophyses there extends a thin lamina having the appear-
ance of a broad shelf or platform. Inferiorly the zygapophyses are supported by
the powerful inferior branches of the prezygapophysial laminze while the superior
branches of these laminze are rudimentary. The infrazygapophysial cavity is deep
and simple, the supra- is quite shallow.
Posteriorly there is a deep cup on the centrum for the reception of the ball of
the succeeding vertebra. The articular surface of the posterior zygapophysis faces
downward and outward.
The postzygapophysial lamine are branched, the internal and smaller of these
branches from the zygapophyses of the opposite sides meet in the middle line and
10 MEMOIRS OF THE CARNEGIE MUSEUM
form a widely open V. The supra- and infrapostzygapophysial cavities are very
deep and at the bottom the latter is subdivided into three unequal pockets by two
short, delicate laminze. On the posterior surface of the neural spine there is a
median rugose surface suggestive perhaps of a postspinal lamina.
As has already been remarked, the series of vertebree was interrupted at the
first dorsal and a number of the succeeding vertebree are missing from the series.
I have estimated the number of missing dorsals at four, the second, third, fourth
and fifth.
it is, as will appear later, the first dorsal of the series of nine mentioned above as
If this estimate is correct, and there are many reasons for believing that
pertaining to this skeleton would be the sixth of
the dorsal series. That these two series of verte-
bree. pertained to one and the same skeleton is
demonstrated beyond the possibility of a reason-
able doubt, not alone by their proximity to one
SN
EX
e )
another in the quarry where they lay imbedded in
the sandstone as shown in Figs. 1 and 2 at B. 572
and C. 572, but by the relative sizes of the verte-
bree, their color and texture and the entirely closed
sutures of the neural arches, indicating in each in-
stance an animal of identically the same age.
Figs. 3 and 4 are side views respectively of the
supposed sixth and the first dorsal. They are intro-
duced here for direct comparison with one another
>
Fie. 3. Sixth (?) dorsal of type of
572)
seen from right side, 5 natural size,
pA, lamina; 4, become more apparent after an examination of
horizontal lamina ; ol, oblique lamina.
Ronloceniiiocommnn prtccns ONO. and to show the great disparity in structure exist-
ing between these two vertebrae. These differences
postzygapophysial
Se ee ee ae plate A, where posterior and anterior views of the
IG. 4. ivs orsal Of Same, same
view ; ol, oblique lamina ; al, inferior
blade of diapophysial lamina ; ¢, tuber-
cular rib facet ; ¢, capitular rib facet ;
S, surface for muscular attachment of
scapula, 7; natural size.
the transverse processes and the height and form of the neural spine.
same vertebree are also given.
The greatest structural differences exhibited in
these two vertebree are to be found in the relative
height of the neural arches, the form and position
of the capitular rib facets, the form and positions of
All these and
many other differences of only less importance will become apparent as we describe
the sixth dorsal in detail.
Sixth ? Dorsal (Plate I., Fig. 6).— This vertebra is complete save a small
part of the anterior end of the centrum and a portion of the upper part of the
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS ile
neural spine. The entire length of the spine is represented, but the margins, except
the posterior, are weathered away and have been restored in plaster. It thus
happens that the drawing represents the top of the spine less complete than it
actually is.
The centrum was opisthoccelous with the cup moderately deep and the ball at the
anterior extremity rather more convex than represented in the drawings _ It is con-
stricted medially, both laterally and inferiorly, and the inferior surface presents a
broad median longitudinal ridge. There are no infracentral cavities. The pleuro-
central cavities are large, irregularly triangular in outline and very deep, with the
dividing median septum reduced to a thin lamina.
The neural arch is high, much constricted transversely and much shorter antero-
posteriorly than the centrum. On the anterior lateral margin of either side it sup-
ports an elongated, sessile, capitular rib facet situated midway between the anterior
zygapophyses and the superior border of the centrum. ‘This facet is quite unlike
that of the succeeding dorsals, it is very distinctive and is most like that of the sixth
dorsal in H. utterbackii to be described later, as compare Plates I. and II.
The transverse processes are high and directed obliquely upward and outward at
an angle of about forty-five degrees. At their extremities they bear tubercular rib
facets which face outward and a little upward. Inferiorly the transverse process is
supported by a powerful lamina arising from the posteroexternal border of the
neural arch and forming the greater portion of the broad posterior surface of the
transverse process. Although this lamina occupies a position identical with that of
the inferior blade of the diapophysial lamina in the corresponding dorsal of Diplo-
docus, nevertheless it is clearly homologous with the oblique lamina of the first dorsal
of this skeleton. In this vertebra the diapophysial lamina, only the inferior branch
of which is represented in the last cervical and first dorsal described above, has be-
come quite obsolete. There is, in Haplocanthosawrus, no division of the anterior
blade of the horizontal lamina into superior and inferior branches such as has been
shown to be the case in the anterior dorsals of Diplodocus carnegu. ‘This fact at
once distinguishes that lamina marked a/ and d/, in the first dorsal and last cervi-
cal as the diapophysial and that marked o/, as an oblique lamina, though in no
sense to be considered as homologous with the oblique lamina that in the middor-
sals of Diplodocus carnegui gives support posteriorly and inferiorly to the capitular
rib facet.
The anterior blade of the horizontal lamina is long and broad, the posterior
short and narrow. There is a short and narrow superior blade of the diapophysial
lamina invading the bottom of the deep supradiapophysial cavity shown at d/, in
12 MEMOIRS OF THE CARNEGIE MUSEUM
Plate I., Fig. 6, second column, and a little anterior to this is a second lamina ai,
which may be an anterior branch of the superior blade of the diapophysial lamina,
but which is interpreted as a branch of the prespinal.
Although the transverse process appears massive it is really very light and thin,
and save toward the base it is made up entirely of the anterior blade of the hori-
zontal lamina and the oblique lamina. ‘These two laminz meet at nearly right
angles so as to enclose a very deep trough which opens downward, forward and
outward and is confluent with the very deep infradiapophysial cavity. There is no
prediapophysial cavity and the postdiapophysial cayity is very shallow and incon-
spicuous.
Seen from in front, the neural arch appears high and constricted just beneath
the capitular rib facets but expanded in the region of the superior border of those
surfaces. The anterior zygapophyses are elevated and have the articular surfaces
elongated transversely and abbreviated antero-posteriorly. Beneath the anterior
zygapophyses there is a deep infraprezygapophysial cavity confluent with a deep
trough into which the neural canal opens. The supraprezygapophysial cavity is
shallow and separated from the one below by a short, stout, transverse plate which
gives support superiorly to the lateral borders of the neural arch. In cross-section
the neural spine is triangular with the apex of the triangle directed forward and
forming the rather broad and rugose prespinal surface.
Seen from behind, the neural spine is broad and rugose, though much narrower
than in the first dorsal. This is due to the greater development of the superior
blades of the postzygapophysial lamine, which are thin and expanded and con-
tinue to the summit of the neural spine. There are shallow supra- and infrazyga-
pophysial cavities and the latter is much elongated and inclosed laterally by the
inferior blades of the postzygapophysial laminze. These give support inferiorly to
a well-formed hyposphenal process developed on this and the succeeding dorsals
showing that in addition to the ordinary zygapophysial articulation of the preced-
ing vertebree the dorsals of this region had a hyposphene-hypantrum articulation.
Dorsals Seven? to Fourteen? Inclusive (Plate I., Figs. 7-14).— After the above
rather tedious detailed description of the posterior cervicals and dorsals one and six?,
the succeeding dorsals may best be described together. Since, in their more impor-
tant and general characters, they agree very well both with one another and with
the supposed sixth dorsal just described, the following description may very well be
limited to a general reference to those characters wherein they all agree, followed by
a special reference to the more important distinctive characters of each, wherever
such exist. As already stated, these vertebrae, together with the one last described
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 13
when discovered in the quarry near Canyon City, Colorado, were all closely inter-
locked by their zygapophyses. They were taken up in a single block of matrix
and were received at the museum still imbedded in the sandstone and in their
exact original position relative to one another. Thus whatever question there may
be regarding their position relative to the last cervical, there can be absolutely no
question regarding their position relative to each other, while the same can be said
with only a little less certainty regarding their position in relation to the sacrum,
for the supposed fourteenth or Jast of this series, although removed some ten feet
from the sacrum, agrees very well in size with the first sacral and has the posterior
extremity modified for articulation with that vertebra. There can be no reasonable
doubt but that these dorsals and cervicals formed part of the same skeleton as that
to which belonged the pelvis and caudals shown in Plates I1I., [V., V., and the femur
shown in text Fig. 14.
In the dorsals now under consideration, the centra are comparatively small, con-
stricted medially, opisthoccelous throughout, though less decidedly so in the posterior
region. They are subequal in length, with those of the posterior region a little
shorter than those of the anterior. The pleurocentral cavities are deep and sub-
equal in area. hey are all irregularly ovate in outline with the broader end di-
rected anteriorly. The neural arches are high and the neural spines short and stout.
There is a striking contrast in the proportionate length of the neural spines and
height of the neural arches in the dorsals of Haplocanthosaurus when compared with
those elements in the same vertebree of any other genus of Sauropod dinosaur known
to the present writer. This contrast is especially noticeable in Diplodocus and Bron-
fosawrus but is less marked in Morosaurus. The capitular facets are somewhat
pedunculate and gradually assume a more elevated position in the anterior dorsals
until the eighth is reached when they attain an elevation equal to that of the an-
terior zygapophyses. In the eighth and succeeding dorsals their position remains
constant. The transverse processes throughout the entire series of vertebrae now
under consideration are subequal in length and are directed upward and outward
at an angle of about forty-five degrees. The transverse processes of the posterior dor-
sals are somewhat more slender than are those of the anterior dorsals. Commenc-
ing with the eighth dorsal the superior blade of the diapophysial lamina becomes
very well developed and in this and the succeeding vertebra it unites, about mid-
way up the spine, with the superior blade of the postzygapophysial lamina to form
a single lamina giving lateral support to the neural spine. The posterior position
of the extremity of the transverse process in the eighth dorsal as shown in Plate I.,
Fig. 8, is due to distortion and is not the normal position of that element. In the
c
14 MEMOIRS OF THE CARNEGIE MUSEUM
seventh and succeeding dorsals the antero-posterior diameter of the neural spines
exceeds the transverse and the extremities of all these vertebree are somewhat ex-
panded and rugose. All the dorsals of this region exhibit the hyposphene-hypan-
trum articulation.
The Sacrum (Plates IV. and V.).— In the present skeleton as in all other fully
adult Sauropod dinosaurs to whatsoever known genus or species they may pertain,
there are five vertebree, codssified by their centra and functioning as sacrals by giv-
ing support to the ilia either by means of so-called sacral ribs or transverse processes
or by both these elements.. Whether or not all five of these vertebrae should be re-
garded as true sacrals must remain very largely a matter of individual opinion.
This matter has already been discussed at some length by the present writer in his
paper on Diplodocus which formed the first of a series of memoirs, which will con-
tinue to appear from time to time relating to the dinosaur remains in the collections
of this museum.
Whether the number of true sacrals in the Sauropoda be five or less it is evident
that in those genera of American Sauropods where the complete sacrum is known,
namely, Diplodocus, Brontosawrus, Morosaurus and Haplocanthosawrus, the number
of vertebree functioning as sacrals, that is giving support to the ilia, is constant and
is in no sense different in or diagnostic of the several genera as was supposed by the
late Professor Marsh. It frequently happens in the case of isolated sacra pertaining
to young individuals that one or two of the functional sacrals through not having
been firmly codssified with the three vertebree which, according to the present
writer's opinion, constitute the true sacrals, have become detached and lost and in
this manner the number of functional sacrals has been mistakenly reduced to either
three or four according as the number of detached vertebree was one or two. By an
unfortunate circumstance this proved to be the case with the sacra of Diplodocus
and Morosawrus first discovered and described by Marsh. ‘The sacrum of the first
of these genera was found detached and consisted of three codssified centra, while in
that of the second (the type of MZ. grandis) there were four codssified centra From
these circumstances Professor Marsh quite naturally concluded that the number of
functional sacrals in these genera was respectively three for the former and four for
the latter and proceeded to make those numbers diagnostic of the genera and fam-
ilies to which they pertained. Subsequent discovery of more complete material has
demonstrated beyond a doubt the number of functional sacrals to be five in each of
these genera as in Brontosawrus. The fragmentary sacra of Apatosawrus and Atlanto-
saurus figured by Marsh are evidently incapable of furnishing any definite proof as
to the exact number of sacrals in those genera, which should they finally prove to
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 15
be valid will doubtless also be found to be provided with five functional sacrals.
The same remark also applies to the recently described genus Brachiosawrus of
Riggs. On the other hand it sometimes happens in the skeletons of very old indi-
viduals that an anterior caudal or posterior dorsal becomes codéssified with the
functional sacrals. As an example of the latter the sacrum of the type of Bronto-
saurus excelsus Marsh may be cited. In such instances however there is no danger
of misinterpreting the additional vertebree since they never bear so-called sacral ribs
ribs or give any support to the ilia.
The sacrum in the present genus and species may be described in general as_be-
ing broad, low, with short neural spines and consisting of five vertebrae with
subequal, codssified centra. All five of these vertebrae bear parapophyses (sacral
ribs) and give support to the ilia through the intermedium of these and the dia-
pophyses. The parapophyses of the three median or true sacrals expand distally
and unite to form the inner superior border of the acetabulum as is well shown in
Plate V., Fig. 1.
Seen from below (Plate V., Fig. 1) the sacral centra appear subequal in length,
with the transverse diameter of the first and last exceeding that of either of the
three median or true sacrals. All five of these vertebree bear so-called sacral ribs
springing directly from the middle of the centra, save that of the first, which springs
from the superior internal border of the centrum. The excellent state of preserva-
tion in which this sacrum was found, firmly attached to the ilia of either side,
demonstrates beyond a doubt the fact that all five of the vertebrae bear those proc-
esses which have been called sacral ribs. As to whether or not the first of the
sacrals is homologous with that which in Diplodocus I have described as the last
dorsal, though there functioning as a sacral, | am as yet undecided. I believe, how-
ever, that it is, although since it is the neural spines of this and the two succeeding
vertebree that are codssified in the present sacrum, this fact might be considered by
some as tending to disprove this assumption, for in Dip/odocus it is the spines of the
three median vertebree, the true sacrals, that are codssified. However this may be, I
am inclined to the opinion that the first vertebra which in Diploducus gives support
to the ilia did in fact bear what has usually been interpreted as a sacral rib and
should therefore be considered as a sacral by those who accept the presence of this
element as distinguishing the sacrals. The imperfect condition of all the Diplodocus
sacra so far discovered precludes the possibility of determining this point with abso-
lute certainty in that genus. In the type of Diplodocus carnegu the right side of
this vertebra is present though in a somewhat imperfect condition and presents an
element which, though occupying a decidedly more elevated position than that of
16 MEMOIRS OF THE CARNEGIE MUSEUM
the so-called sacral ribs in the succeeding sacrals, does however spring from the
superior lateral surface of the centrum.. It may therefore be considered as homol-
ogous with those elements in the true sacrals. Its position with relation to the
vertebral centrum may be considered as evidence that this vertebra though function-
ing as a sacral is in reality a modified dorsal and that, contrary to Osborn’s asser-_
tion, the sacrum in the Sauropoda may have expanded by the addition of at least
one posterior dorsal.
The diapophyses of all the sacral vertebree send downward thin vertical diapo-
physial lamine. These unite at their extremity with the sacral ribs or as I prefer
to call them the parapophyses* of their respective vertebree to form thin partitions
separating the four large sacral foramina to be seen in the inferior view of thig
sacrum with ilia attached, shown in Fig. 1, Plate V. Internally these foramina are
enclosed by the sacral centra and externally by the extended iliac bar formed by the
expanded and coalesced distal extremities of the parapophyses (sacral ribs).
The parapophyses (sacral ribs) of the first and fifth sacrals are longer but rather
more slender than those of the three median or true sacrals. This is especially true
of the first sacral. In this vertebra this element springs from the superoanterior
surface of the centrum, continues outward for some distance as a strong bar when it
expands and divides into two branches enclosing a small foramen bounded externally
by the ium and shown in Plate V., Fig. 1. The inferior of these two branches abuts
against the base of the pubic peduncle, the superior unites with the diapophysial
lamina in giving support to the widely expanded anterior blade of the ilium.
In the posterior sacral the parapophysis springs from the middle of the centrum
at its anterior extremity and continues as a single bar, only moderately expanded
distally, where it gives support to the posterior blade of the ilium. Superiorly it is
united throughout its entire length with the diapophysial lamina. The diapophy-
sis branches distally and with the posterior blade of the ilium encloses the foramen
seen in Plate V., Fig. 3.
The parapophyses of the three median sacrals are all short and stout. They
differ from those of the first.and fifth sacrals in having their extremities expanded
and coalesced so as to form a strong iliac bar not only giving support to the ilia but
constituting the inner superior borders of the acetabula. These three vertebrae were
the first to become codssified. Throughout the entire life of the individual they
gave the chief, and during a certain period of its youth perhaps, almost the only
support to the ilia. It is for these reasons that I consider these vertebree as the only
true sacrals of which the sacrum in the earliest Sauropods was alone composed.
‘The homologies of these elements will be discussed more fully when we come to speak of the caudals.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 17
Should we ever be so fortunate as to discover representatives of the very earliest
Sauropod Dinosaurs it is not at all improbable that in these the sacrum will be
found to consist of only these three vertebree. This number is, however, a decided
advance over that which is supposed to have constituted the sacrum in the primitive
reptilia. This supposition, however, is at present purely conjectural though sup-
ported by considerable evidence. If we consider the three median vertebree as the
true sacrals the anterior might very appropriately be called a dorso-sacral and the
posterior a sacro-caudal.
In all the functional sacrals the parapophyses spring from the anterior extremi-
ties of the centra of the several vertebrae, but in the first and second frwe sacrals
there is in each instance some slight union between the posterior extremities of the
centra of these vertebree and the succeeding parapophysis as shown in Plate V., Fig. 1.
Viewed from above, the diapophyses of the sacrals in Haplocanthosawrus are each
seen to be formed by the union of two lamine. One of these springs from the
spine of that vertebra to which the process pertains and the other from the antero-
external margin of the spine of the immediately posterior sacral. These lamin
rapidly converge both inferiorly and exteriorly and unite in forming the diapophy-
ses or transverse processes. These are on a level with the superior border of the
ilium and a short distance before coming in contact with that element they expand
anteroposteriorly and present broad, rugose, superior surfaces.
The neural spines of all the sacrals are extremely short as compared with the
same elements in either Diplodocus or Brontosawrus and in this respect they more
nearly resemble the same elements in Morosawrus. Those of the three posterior
sacrals are directed upward and a little backward. ‘The spines of the three anterior
sacrals are coalesced and form an elongated bony plate. In Diplodocus and Bronto-
saurus itis the three (sometimes the two anterior in the former genus) true median
sacrals that have the spines coalesced. Superiorly and posteriorly the spines are
much expanded and they each present prominent lateral rugosities at the apex.
Seen from behind or in front the sacrum is considerably distorted by pressure.
In the drawings, Plate V., this distortion has been for the most part eliminated and
the sacrum appears low and very broad with the neural arches of only moderate
height when compared with those of the dorsals. In so far as I have been able to
determine there is in the present genus no unusual development of the neural canal
in the region of the sacrum.
The principal characters of the sacrum in the present genus are well shown in Plates
IV. and V., where in the former comparative views are given of the pelves of Bron-
tosaurus, Diplodocus and Haplocanthosawrus with their respective sacra in position.
(C M ih ns OF LE Dy IN EG LE: )y hi
18 MEMOIRS OF THE CARNEGIE MUSEUM
The more important dimensions of the sacrum of the type of Haplocanthosaurus
priscus are as follows:
mm.
Total length of the five coossified sacrals.......2....-..ceccsecrscereccerecsaceeesceene. 795
Greatest expanse of transverse processes of first sacral....................0.00005- 640
ce ee Wg Bg Va Sb poe erin caicoa emcee sence cen 700
Height of top of neural spine above bottom of centrum in first sacral......... 520
x ee oe a oe ue “s TAS Ui Mey lentes 485
Anteroposterior length of three codssified neural spines .......... ...........-.-- 398
Height of anterior neural spine above zygapophyses...................20e0e0s eee 252
i posterior ‘‘ rit ef Ee vinta ataaneuenncrnenctactonendice 180
The Caudal Vertebre (Plate III.).—Nineteen anterior caudals were found associated
with the present skeleton. Their position in the quarry relative to one another and
to the sacrum are shown in Figs. 1 and 2.
The centra throughout the entire series of nineteen caudals are remarkably short
when compared with the same vertebree in Diplodocus. They are somewhat con-
stricted medially and are slightly amphiccelous with the concavity of the anterior ex-
tremity more pronounced than that of the posterior. The centrum of the first caudal
is the shortest of the series. From this they very gradually and slowly increase in
length until the twelfth caudal is reached when they begin very gradually to de-
crease in length.
The neural spines throughout are comparatively short and directed somewhat
backward. They are compressed and with rugose extremities which are quite simple
throughout instead of being laterally expanded and emarginate as in caudals one to
eight in Diplodocus carnegi.
The anterior zygapophyses are slender and extended far forward in advance of
the anterior extremities of their respective centra. The posterior zygapophyses are
not extended beyond the posterior extremities of the centra.
The transverse processes even in the anterior caudals are quite simple when com-
pared with the same elements in Diplodocus and Brontosaurus. In the anterior
caudals they appear as simple, broad plates of bone springing directly from the
neural arches and the superior lateral surfaces of the centra. These bony plates are
nearly flat and thin. They are entire instead of being perforated as in Dzplodocus,
and their posterior and anterior surfaces are entirely destitute of that series of vertical
or radiating laminee seen in the anterior caudals of Diplodocus carnegu. The trans-
verse processes of the caudals decrease rapidly in size as we proceed posteriorly and
in the twelfth caudal they are reduced to a rounded knob of bone on either side of
the centrum near the superior border, while just above this on the middle of the side
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 19
of the neural arch there is a second prominence less pronounced, however, than that
on the centrum. In the thirteenth caudal the prominence on the centrum is only
faintly distinguishable. In the succeeding vertebre it has disappeared entirely,
while that on the neural arch continues on the thirteenth, fourteenth and fifteenth
caudals, but is wanting on the succeeding vertebree. Of these prominences or tuber-
osities the superior or that one situated on the neural arch doubtless represents a
rudimentary diapophysis, while the inferior or that situated on the side of the centrum
may be considered as homologous with the parapophysis. 1t would, therefore, appear
as though the transverse processes in the anterior caudals were made up of the coal-
esced diapophyses and parapophyses. Just what bearing this may have on the exact
homologies of the so-called sacral ribs in the Sauropoda it is impossible to say. It
would appear, however, that Osborn’s assertion that a “sacral rib is not a transverse
process’’° is open to criticism when that term is applied to these elements in the
dinosaur pelvis, or at least needs some further support, and that Marsh’s statement
that ‘each sacral vertebra supports its own sacral rib or transverse process’? may
not have been so far from correct as Osborn supposed it to be, though Marsh’s
assertion that the sacral vertebree in the Sauropoda were without diapophyses is
doubtless erroneous. If, as Osborn asserts: ‘“‘ The sacrum of Sauropoda (Cetiosaurs)
is reinforeed by the addition, not of dorsals, but of anterior caudals,” it would seem
quite evident that those elements which spring from the sacrals and give support to
the ilia are in reality only the modified transverse processes of the caudals, and since,
as has already been shown, the latter appear to have been formed by the union of
parapophyses and diapophyses, there would seem very good reasons for assuming that
the so-called sacral ribs which spring directly from the sacral centra are homologous
with the parapophyses, while the superior bar giving support to the superior border
of the ilium represents the diapophyses and that these two elements with the con-
necting diapophysial lamina together constitute the transverse process. According
to this interpretation the so-called sacral ribs become morphologically quite distinct
from those elements in the tailed Amphibians as described by Flower on page 66 of
his “ Osteology of the Mammalia,” and I am inclined to the opinion that, while the
articulation of the ilium with the sacrum in the Hell Bender (Menopoma) and other
allied forms is by means of a sacral rib interposed between the ilium and the trans-
verse process of the sacral vertebra in the Sauropoda as well as in all the other terrestrial
vertebrates requiring more or less rigidity in this region this interposed sacral rib, if
it ever existed, has disappeared altogether, allowing the ilium to come in direct con-
tact with the transverse processes of the sacrum. In Menopoma the transverse proc-
°>See Memoirs Am. Mus. Nat. Hist., Vol. I., Part V., p. 202.
20 MEMOIRS OF THE CARNEGIE MUSEUM
ess of the sacral is stronger not only than those of the preceding and succeeding
vertebrae, but it is stronger than its sacral rib, although the latter is more robust
than the movable ribs borne by the transverse processes of the immediately preced-
ing and succeeding vertebree. It would seem more probable, therefore, that the
smaller and more slender sacral rib would become obsolete than the stronger
and more robust transverse process. Whether this elimination was accom-
plished by the complete disappearance of the sacral rib or by its fusion with
the transverse process cannot be told. It by the latter process, however, the
so-called sacral ribs in the Sauropod sacrum would then be homologous with
the coalesced sacral ribs and transverse processes. But in the sacra of the
Sauropoda and other highly specialized terrestrial vertebrates, whether reptiles
or mammals, it would appear to be quite evident that in all those sacral
vertebree added to the primitive sacrum through the modification of anterior cau-
dals it is the transverse processes (united diapophyses and paropophyses) that have
been modified to give support to the ilia instead of true sacral ribs homologous with
the free ribs borne at the extremities of the transverse processes in the anterior cau-
dals of Menopoma, for in no instance are the transverse processes of the anterior cau-
dals of even moderately specialized terrestrial vertebrates known to have borne such
ribs. Even in the modern Iguana and in the crocodiles where the sacrum is still
exceedingly primitive consisting of only two ununited vertebre ,there are no movy-
able or other ribs on the transverse processes of the anterior caudals and none are
known to the present writer even among the earliest known Dinosauria. It does
not seem at all reasonable to suppose that these ribs were present in the primitive
forms in the caudal region, that they disappeared and then reappeared in the suc-
cessive caudals as these were added to the primitive sacrum more especially since
their presence would tend to produce instability rather than strength in that region
where rigidity is especially advantageous. In Figs. 5 and 6 are given superior
views of the sacra together with the immediately preceding and succeeding verte-
bree in Menopoma allegheniensis and Iguana tuberculata. A study of these figures
shows the marked difference in the structure of the sacrum in the two. In Meno-
poma the ilia articulate with the transverse processes of the solitary sacral through
the intermedium of sacral ribs, while in the Iguana this articulation is directly with
processes firmly fixed one on either side of the centra of each of the two sacrals.
Whatever the exact homologies of these latter processes may be it is impossible to
say with certainty, though embryology ought to offer some evidence. In general
form and in position, however, it is evident that they approximate much more
closely the transverse processes than true sacral ribs. If, however, they are homolo-
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 21
gous with the true sacral ribs as seen in Menopoma, which to the writer seems ex-
tremely improbable, it does not follow that they are ‘profoundly different from the
dorsal ribs” as has been stated by Osborn;° for an examination of a skeleton of
Menopoma will show the morphological identity of the sacral ribs with the free ribs
borne at the extremities of the transverse processes alike of the anterior caudals and
the entire presacral series, while the latter must be homologous with the dorsal ribs
6
Fig. 5. Superior view of sacral, anterior caudal and posterior dorsal or lumbar of Menopoma alleghen-
iensis, twice natural size. s, sacral; c, anterior caudal; d, posterior dorsal; ¢.p., transverse process ;
s.r., sacral rib; il, ilium; f.r., free rib.
Fic. 6. Superior view of sacrum, anterior caudal and posterior lumbar or dorsal of Iguana tuber-
culata, natural size. d, posterior dorsal; s.1, first sacral; s.2, last sacral; c, anterior caudal; t.p.,
transverse process except that on last dorsal which is a free rib ; J, ilium.
in the terrestrial vertebrates as will become apparent by a study of the skeleton of
Iguana where the transition from the short straight ribs of the dorso-lumbar region
to the elongated and curved ribs of the true dorsals is quite gradual.
In consideration of the characters just described as obtaining in the transverse
processes of the caudals of Haplocanthosaurus in connection with those already men-
tioned as pertaining to the sacrum in the various genera of the Sauropoda, it appears
to the present writer that the following characters relative to the structure of the Sau-
ropod sacrum as a whole and the homologies of the different elements with those
of the other vertebrre seem quite probable though not at present capable of being
demonstrated with certainty.
First. — That the Sauropod sacrum is composed of five coossified vertebre: which func-
tion as sacrals.
Seconp. — That the three median of these five functional sacrals alone composed the
sacrum in the primitive Sawropoda and may be regarded as the true sacrals.
5Vol. I., Part V., Mem. Am. Mus. Nat. Hist., p. 201.
to
bo
MEMOIRS OF THE CARNEGIE MUSEUM
Turrp. — That the number of sacrals in the Sawropoda has been increased to five by
the addition of a posterior dorsal and an anterior caudal.
Fourra. — That the sacrals give support to the ilia solely by means of the transverse
processes (diapophyses and parapophyses).
Frera. — That there are no true sacral ribs homologous with those elements in the
tailed anvyphibia and that the so-called sacral ribs are really homologous with the para-
pophyses or inferior branches of the transverse processes.
It is true that the parapophyses (sacral ribs) of the sacrals, as also the trans-
verse processes of the caudals in the Sauropoda are derived from centers of ossifica-
tion distinct from those which give origin to the centra, and this fact may by some
authorities be taken as proof that they are not portions of the transverse processes,
though I should not consider it as such.
The principal dimensions sometimes materially modified by crushing of the
several vertebree in the type of Haplocanthosawrus priscus (No. 572) are given in the
following table: In column 1 the greatest expanse of the transverse processes of
the diapophyses are given; column 2, greatest length of centra; column 3, trans-
verse diameter of centra at posterior extremity; column 4, height of neural spines
mim. 1 in mm 2 in. mm oe in mm 4 in
214. Cervical. | | 259 112 1D | 6} 320 128
215. aC | 247 91 150 5L 352 13%
1. Dorsal. | 403° | es 224 83 153 6 355 14
6. «“ | 420 16} 185 7h 135 5h 568 99-5.
Ws « | 458 18 173 63 145 5b 590 231
8 ‘i AS | IR 165 64 150 5L 551 215
9 “ | 457 18 185 7h 154 6 583 227
10 «“ 440 17} 164 62 153 6 582 297
11 ve 430 162 170 6: 161 62 597 234
12 “ 425 163 150 5t 178 7 607 237
13 ‘ 410 16} 146 5a 191 74 610 24
14, 410 164 125 42 208 8 615 241
1. Caudal. 416 162 103 4 195 78 510 20
2. & 390 154 82 31 178 7 471 183
3. os 320 123 91 38 190 7h 365 142
4. G6 353 133 86 BS Gs 63 375 143
5, te 314 122 94 Sires aele7 5: 62 396 153
6. a 230 9} 95 32 137 58 404 15:
Ws ? 2 105 41 ? 2 415 163
8. ‘ 216 83 Or | —B: 125 423 397 153
9 te | 100 4 | 115 43 ? >
10 114 AN TGS 63 345 13}
11 & 105 41 105 4h 338 133
12 ‘ 110 43 11@ | 43 314 128
183. “ 101 4 | 114 4} 298 118
14. « 100 4 | 109 Ai | GYBTL 10}
15 OG 100 32 | 107 A HRB 103
16 99 4 101 4 261 10}
17 “ 95 38 | 100 BR Dei 98
18 ‘“ 92 38 | 100 i 229 9
19 be 91 3g | 92 38 210 8}
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 23
above middle of inferior border of centra in presacrals and above inferior border
of posterior end in postsacrals.
The inconsistencies that appear in the above table of measurements are due to
the varying amount of crushing to which the different vertebrae were subjected
while entombed in the sandstones. In this connection it should be remembered
that these animals lived in a period long previous to that which witnessed the final
upheaval of the front range of the Rocky Mountains and that the bones, as well as
the sandstones in which they were imbedded, have been subjected to the enormous
pressure which effected the upheaval of that mountain range. Little wonder that
they are in many instances much crushed and distorted. It thus happens that the
measurements given above are of value only as giving a general idea as to the sev-
eral dimensions of the various vertebrae. In most instances they cannot be consid-
ered as representative of the exact measurements and therefore capable of being
compared critically with those of other skeletons.
The Chevrons (Figs. 7, 8, 9, 10). — Only two chevrons were found. One of these
(Figs. 7 and 8) was found in position between the eighth and ninth caudals. It
does not differ materially from the chevron of the
same region in Diplodocus or Brontosawrus. It is Y-
shaped with the open portion somewhat abbreviated
and the inferior portion elongated, compressed and
with spatulate extremity. The articular surfaces of
opposite sides at the proximal ends are not confluent.
The length of this chevron is 313 mm. When seen
to)
from the side, it curves less strongly backward at the Fries. 7 and 8, chevron between
: : ; caudals 8 and 9, side and posterior
distal end than does the same chevron in Diplodocus.- _. ss
views respectively, one-tenth nat-
The other chevron (Figs. 9 and 10) was found in ypya) size.
position articulating with caudals thirteen and four- Fics. 9 and 10, chevron between
feen. It differs from the one just described in its Ccavdalstéand 14 jside‘and posterior
smaller size and in the more elongated open portion fC ee eee
of the Y as compared with the closed inferior portion. avi
At the point where the two branches meet it is greatly constricted antero-posteriorly,
while distally it is much expanded in the same direction, but without the anterior and
posterior projections which are already quite prominent in the same and the preced-
ing chevron in Diplodocus. This chevron has a length of 184 mm.
The Ribs (Figs. 11, 12 and 13).—The ribs do not differ essentially from those of
other members of the Sauropoda. They increase in length and strength quite rap-
idly from the first to the fourth when they continue subequal in length until in
24 MEMOIRS OF THE CARNEGIE MUSEUM
about the region of the ninth or tenth. Posterior to these, they rapidly become
shorter and more slender. ‘The ribs of the anterior and mid-dorsal region are much
expanded proximally where they present a rather deep concavity on the posterior
surface, while the anterior surface in the same region is convex. Beyond this they
become subcircular in cross-section and somewhat spatulate at their distal extremi-
ies. The ribs of the posterior region are decidedly less expanded proximally and
in the middle they are semicircular in cross-section.
Figs. 11, 12 and 13 represent respectively anterior views of the supposed
second, fourth and thirteenth ribs. The second rib has a length of 911 mm., the
Fic. 11. Anterior view of second? rib of right side, one-tenth nat. size.
Fie. 12. Anterior view of fourth? rib of left side, one-tenth nat. size.
Fie. 13. Anterior view of thirteenth? rib of left side, one-tenth nat. size.
fourth 1,394 mm. and the thirteenth 710 mm. Compared with the size of the
animal as a whole the ribs of Haplocanthosawrus are neither long nor robust.
Throughout the entire series the capitulum and tuberculum are well separated.
The capitulum is pediceled while the tuberculum is sessile, save in the anterior ribs
where it is also pediceled. ;
The Pelvis (Plates IV. and V.).
All the elements of the pelvis were found approximately in position and in a
splendid state of preservation.
The Ilium (Plate IV., Fig. 3).—In general form the ilium resembles that of other
members of the Sauropoda. In the present skeleton both ilia were found attached
to the sacrum which lay imbedded in the sandstones with the spines directed up-
wards but reclining a little on its right side. It thus happened that these elements
received the pressure of the superincumbent rocks in a direction obliquely vertical
and from the left. This pressure was sufficient to accomplish considerable crushing
and the superior borders of the ilia have been considerably flattened and instead of —
describing the are of a circle as was doubtless the case before this distortion took
place, for a considerable distance along their superior borders they now present a
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 25
nearly flat surface. In the drawings this distortion has not been entirely elimi-
nated.
The ischiac peduncle is broad and sessile with the transverse diameter of the
articular surface for the ischium considerably exceeding the anteroposterior
diameter.
The pubic peduncle is elongate, extending far below the inferior border of the
ilium. It forms most of the anterior border of the acetabulum. The articular sur-
face for the pubis has the transverse diameter greater than the anteroposterior. At
its base the pubic peduncle is supported internally by the parapophyses of the dorso-
sacral and the first true sacral vertebra as shown in Plate V., Fig. 11.
The ilium is produced far in front of the pubic peduncle into a broad anterior
blade. At its extremity this anterior blade of the ilium is broad instead of pointed
as in Diplodocus and Brontosaurus. In superficial area the anterior blade of the
ilium constitutes nearly one half of that element. Internally it is supported by the
powerful and widely expanded diapophysis of the dorso-sacral.
The ilia are not produced far behind the ischiac peduncles and the posterior
blades are therefore short but rather broad.
The upper one-half of the acetabular border is formed by the ilium, the pubic
and ischiac peduncles and the acetabular bar formed by the united extremities of
the parapophyses of the three true sacrals.
The anterior extremities of the ilia of opposite sides are very widely separated
and the posterior extremities less decidedly so while in the middle, both superiorly
and inferiorly the ilia of opposite sides approach more nearly to each other. It thus
happens that the diapophyses and parapophyses of the three true sacrals are shorter
than those of the dorso-sacral and sacro-caudal.
The Pubis (Plate IV., Fig. 3,and Plate V., Fig. 2).—The pubis is proportionately
short and stout and greatly expanded proximally. At their distal extremities the
internal borders of the pubes were in contact for a short distance only. Above this
point of contact, when in position, the pubes were separated by an elongated aper-
ture 500 millimeters in length, while above this aperture they meet again and form
an elongated pubic symphysis about 300 millimeters in length. In either instance
the union between the pubes of opposite sides was ligamentary. The direction and
position of the superior of the two pubic symphysial surfaces is horizontal and ven-
tral rather than vertical. The pubis forms the antero-inferior one fourth of the
acetabulum. The pubic foramen is very large and somewhat elliptical in outline ;
just posterior to and above it there is an extended sutural surface for contact with
the ischium.
26 MEMOIRS OF THE CARNEGIE MUSEUM
The Ischium (Plate IV., Fig. 3, and Plate V., Fig. 3).—As compared with the pubes
the ischia were slender. Proximally they expand and form the postero-inferior one
fourth of the acetabular border. Beneath the acetabular border they present broad,
rugose, sutural surfaces for articulation with the pubes. Posteriorly the ischia con-
tract rapidly and form broad flat bars with broadly rounded external surfaces.
These bars converge and meet distally where they are codssified to form a symphysis
about 195 millimeters in length.
The form and principal characters of the different elements of the pelvis are well
shown in the figures in the plates accompanying this memoir.
The principal measurements of the different elements of the pelvis are as follows:
mm
Greatestrlenothiotiilimmitr iasassc-rencasten ach ae sel dasnaducetoraree nase cuclaetioc RG arene 827
Distance from inferior extremity of pubic peduncle to top of iliac crest...... 512
oe a HG ef ischiac ‘* ie i CS ee 332
Length of pubic peduncle below superior border of acetabulum............... 249
hixpanse) ofelliagatcn tenlonex trent yn eeecec reer Eer rere eeeee eee ce eee ee eee Ee eer eee 1140
oe Se IG MOA OS LOTION a taiei nt aes etch pisses ae hiaetercdee Sate e CREE MC EC ECE 810
a GE oil ony OL EXONUB GOI Ii sen oak atonchd aabimandaiynenb pa ddbeaLancnacasscee bop aledacors 786
ss HOSISCHIAC™ | pits NS cram aa anienn saavemelspa sets dea anagem a seeeace mtaamsttetatias 685
WBE M OCHO L! PWDIS wrsiagec cece sie siranstoeneaesies sepia ricer eesme cacti ae rae seer 693
Greatestibreadthyof pubis. eter ancsuscre atcmecneirece daca cuserenoenetaittace ee meets 432
Least SS Be A Gs aa rgbitens isin. tac aeemee ASIAN oe Sion e Salee No Reto ioneialaniasasaaraey an ata 165
Leneth of ischia from middle of acetabular border to distal end............... 790
Extent of acetabular border of ischium................0.00000 ceeeeseeeeescsseeeeeeeee 210
Breadth of ischium just above symphysis .................0:ccecccceeeeeeeeetceeeeeees 85
Depth ‘ ue os ue PESO TOL Tebaaiae cist ee sale seperate cla automate 50
The Femur (Fig. 14).
Unfortunately the femur is the only element preserved of either the fore or hind
limbs and this is not entirely complete, though sufficiently well preserved to show
most of the more important characters. As shown in the diagram it was found not
far removed from its normal position relative to the pelvis, so that there can be no
reasonable doubt that it pertains to the same skeleton. It does not differ materially
from the femur in other members of the Sauropoda although as compared with
the other portions of the skeleton it appears rather long and stowt. ‘There is a
low and elongated fourth trochanter on the postero-internal margin midway
between the proximal and distal extremities, and just external to this is a shallow
concavity with a markedly rugose surface. The external condyle is larger than the
internal and they are well separated by a deep interecondylar notch. The head is
large and hemispherical in form but without distinct neck. The articular surface
HATCHER: OSTEOLOGY
OF HAPLOCANTHOSAURUS
27
is very rugose and this rugosity is continued along the superior surface of the
greater trochanter to the external surface of the shaft. Both the internal and external
margins of the shaft of the femur curve very gently outward as shown in Fig. 14.
The principal dimensions of the femur are as follows:
mm
ome t emer rcs ru rh eee eens tuna Bae hie 1275
Transverse diameter at proximal end....................... 353
oe ue OG HIS{E Mig ies Caara gens ieee mea 309
ee ue Smid dllesoteshatimertceereernceeenes: 207
CONCLUSIONS.
When considered together the remains upon which the
present genus and species are based indicate an animal of
rather unusual proportions for a member of the Sauropoda.
The number of dorsals and the comparative length of the
individual dorsals indicate a thoracic region proportionally
longer than in Diplodocus, Brontosawrus or Morosauwrus.
While the cervical region appears somewhat abbreviated
and the caudal region must have been remarkably short as
is indicated by the reduced length of the individual verte-
bree, though this was probably made less apparent by an in-
crease in the number of caudals. Judging from the femur
alone the limbs were comparatively long, and the animal
proportionately high and short for a Sauropod dinosaur.
HAPLOCANTHOSAURUS UTTERBACKI sp. nov.
(No. 879.)
The present species is named for Mr. W. H. Utter-
back, its discoverer, and in recognition of his services to
vertebrate paleontology.
Char. Sp.: It is readily distinguished from H. priscus,
the type species of the genus by the character of the pos-
terior dorsal centra which are rather more opisthoccelous
than in the type species. The fully adult indiviual was
doubtless of larger size in the present than in the first
named species of the genus. But the most distinctive
character is to be found in the sacrum which, in the
present species, has the five neural spines normally codés-
sified. The first four are codssified throughout their
See
Left femur of
Fic. 14.
Haplocanthosaurus priscus,
front view, seen obliquely
from within (No. 572), 7; nat.
ural size. h., head; gt., greater
trochanter; ¢r., fourth tro-
chanter; 7. e., internal con-
dyle; e.c., external condyle ;
i.g., inter-condylar groove.
28 MEMOIRS OF THE CARNEGIE MUSEUM
entire length, forming a long bony plate. The union between the fourth and fifth
is limited to the extremities while medially they are separated by an elongated
foramen. In H. priscus only the spines of the three anterior sacrals are codéssified,
those of the first and second sacrals remaining free. This difference exists notwith-
standing that the type of the present species was scarcely adult, the sacral centra
neither being codssified with one another nor with their neural arches. By some
this character might be considered as of generic importance although I prefer to
consider it as of only specific value since in all other parts of the skeleton preserved
there are no distinguishing characters which could be considered as of generic value.
DEscRIPTION oF THE Typr. (No. 879.)
The type of the present species consists of a left scapula and right coracoid,
several ribs and thirty-five more or less complete vertebree distributed as follows :
Ten ceryicals, thirteen dorsals, five sacrals and seven caudals. For the most part
these vertebrae are complete, but in a few instances they are represented only by
isolated spines and neural arches without centra, or by centra without spines and
neural arches, and one anterior cervical, probably the axis or the succeeding cervical,
is represented only by a portion of the neural arch. The position of these bones
relative to one another as they were found in the quarry is shown within the dotted
line in the upper left-hand corner of the diagram shown in Fig. 1 where the positions
of the different bones are indicated as follows : .
1 = cervical 3, placing the number of cervicals at fifteen.
i SoA
Se Gs,
Mies 1. US
Fos 6 NO,
6 and 6’ = RRA
ih) Fok sores
8= Tamia iss
Qa Cale
10) = ‘““ 15, or last of cervical series.
11 and 11’ = dorsal 2.
IDS SB
3; eye! BY 7 SE
(4) gyno! 4
15 and 15’ =
loa &
o>
AI
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 29
17 = dorsal 8.
igs @ ©,
19= “ 10,
= 6 tl:
Pec 12)
22 =) 13)
ts — |
Sa =spines and transverse processes of sacrals; a, b, c, d, e€ represent
respectively the spines of sacrals 1, 2, 3, 4, and 5.
24 = centrum of sacral 4.
VAD) = (a3 (73 (a3 5.
27 = caudal 1.
28 = a YD,
DOE ay 3
30 = 8G 4
SNe — eee S)
se 1G
Be ey
34 = parapophysis (sacral rib) of first sacral.
30-39 inclusive are ribs.
S= left scapula.
¢ = right coracoid.
Shaded bones in diagram do not pertain to Haplocanthosaurus.
As will appear by a critical examination of the diagram the vertebrae of the
cervical and anterior dorsal regions were much scattered and displaced before finally
becoming imbedded in the sands which later become solidified into the sandstones of
almost granitic hardness in which they were found encased. In the following
description of the vertebral column the reader should bear in mind that save for the
third dorsal and the first and second caudals the centra were detached from the
neural arches. Owing to the age of the individual there was as yet only a sutural
union between the centra and the neural arches of the respective vertebree. In
most instances the centra, while not directly attached to their respective arches, were
either found in position or only slightly removed from their normal positions relative
to one another. In some instances, however, as with dorsals 2, 5 and 6, the centra
and neural arches were found separated by a distance of from two to four feet, while
a few vertebrie are represented by their centra or neural arches only.
30 MEMOIRS OF THE CARNEGIE MUSEUM
The Cervicals. Plate II., Series 3 and 4.
The Third (2?) Cervical (Plate II., Fig. 3, Series 4).— The most anterior vertebra
of the cervical series pertaining to this skeleton I have referred to the third although
it may pertain to the axis. Its fragmentary condition precludes the possibility of
determining the exact position with certainty. Its position in the quarry is shown
at 1 in the first diagram. It consists of the posterior portion of the neural arch with
the posterior zygapophyses and it could hardly have occupied a position posterior to
the third cervical although it may pertain to the axis.
The Fourth Cervical (Plate II., Fig. 4, Series 4). — A little to the right of the cer-
vical fragment just described the present vertebra, which I interpret as the fourth
cervical, was found. Its exact position in the quarry is shown at 2 on the diagram.
It is essentially complete and but little distorted, though as with all the cervicals of
this series the rib is disarticulated as was to be expected considering the age of the
individual. The posterior zygapophyses and transverse processes are widely ex-
panded. Near the anterior extremity and on either side of the centrum a strong
process springs from the inferior lateral border. At the extremity this expands into
a capitular facet for the articulation of the capitulum of the cervical rib. ‘These
processes as well as the similar, though less pronounced ones found on the succeed-
ing cervicals may possibly be homologous with the parapophyses. In the present
vertebra they are produced far below the inferior border of the centrum. The
pleurocentral cavity is deep and invades the base of the ball. It-is confluent with
a rather deep cavity found on the superior surface of the process which supports the
capitular rib facet. It is imperfectly divided into anterior and posterior cavities by
a low rounded ridge which may be regarded as an incipient oblique lamina. The
centrum is markedly opisthoccelous with the cavity of the posterior extremity sub-
circular in outline. The inferior surface of the centrum is broad and there are five
shallow infracentral cavities. One of these, the posterior, is medial, and the an-
terior four are lateral, arranged two on either side of the central line, one at the
base of and two posterior to the processes which support the rib facets. The cen-
trum is much contracted medially.
The Eighth Cervical (Plate II., Fig. 8, Series 4). — Between the vertebra just de-
scribed and the next in our series it is evident that a number are missing. I have
estimated the number of missing vertebree at three. ‘This would make the position
of this vertebra the eighth in the series, a position with which it agrees very well
if we commence with the last of the series and work forward, so that I have but
little doubt that this was its correct position. It is essentially complete and not
badly crushed or distorted. Save for its greater size in its general form it very
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 31
closely resembles the vertebra just described. The pleurocentral cavities however
are more completely divided into anterior and posterior moieties by the presence of
more pronounced oblique laminze. There is a single large infracentral cavity and
the cup is broader than deep. ‘The anterior zygapophyses are supported inferiorly
by short and rather slender inferior branches of the prezygapophysial laminze while
inferior branches of the diapophysial Jaminze give support to the broad diapophyses
which bear at their extremities the tubercular rib facets. The position of this ver-
tebra in the quarry is shown at 3 in the diagram.
The Ninth Cervical (Plate II., Fig. 9, Series 4).— This vertebra found at 4 in the
diagram of the quarry was not far removed from the preceding. It consists of the
centrum with the posterior and anterior zygapophyses still in position. It is much
crushed and distorted but in so far as it is possible to determine, it agrees fairly
well with what we should expect to find in the ninth cervical. It has been errone-
ously drawn as complete in Plate IT., Fig. 9.
The Tenth, Eleventh, Twelfth and Thirteenth Cervicals (Plate IL., Figs. 10, 11, 12, 13,
Series 3). — These four vertebrie are in each instance well-nigh perfect and they dis-
play such a gradation of progressive characters that there can be no doubt as to
their constituting a continuous series.
The neural spines and posterior zygapophyses become successively more elevated
as we proceed backward in the series. ‘The spines however show no tendency to
divide, there being scarcely an emargination at the summit even in the last of the
four. ‘The posterior zygapophyses become successively more expanded and the supra-
postzygapophysial cavities become deeper and broader. The position of these
vertebree in the quarry was as follows: The tenth was found at 5, the neural arch
and spine of the eleventh was found at 6, and the centrum at 6’, the twelfth is
shown at 7 and the thirteenth at 8.
The Fourteenth Cervical (Plate I1., Fig. 14, Series 3). — Only the centrum of this
vertebra was recovered; it was found at 9 on the diagram. It is considerably
crushed, especially anteriorly but there is no doubt that it was a cervical and that
its position was posterior to the thirteenth. Its size, length and general characters
indicate that it belonged immediately behind the thirteenth I have, therefore, re-
garded it as the fourteenth.
The Fifteenth Cervical (Plate I1., Fig. 15, Series 3). — This is represented by a well-
preserved neural arch and spine without centrum found at 10 as shown on the dia-
gram. ‘he difference between this spine and that of the thirteenth is such as to
preclude the possibility of its pertaining to the fourteenth or immediately succeed-
ing cervical. I have, therefore, assigned it to the fifteenth or last cervical, with
2 MEMOIRS OF THE CARNEGIE MUSEUM
iS)
which it agrees very well when compared with the spine of that vertebra in
H. priscus where there can be no question as to the proper position relative to the
dorsals. Moreover if the spine of the fourteenth cervical in H. priscus be interposed
between the present spine and that of the thirteenth cervical in the present skeleton
they are seen to form a well-graduated series leaving little doubt that the positions
assigned to the various vertebree of this region of the cervical series in the skeleton
under consideration are correct. The neural spine is faintly emarginate at the apex.
The depth of the emargination is 4 mm.
The Dorsals. (Plate II., Series 1 and 2.)
The Second Dorsal (Plate II., Fig. 2, series 2.) There is in the vertebral series of
the present skeleton no vertebra corresponding to the first dorsal in H. priscus.
That vertebra is apparently unrepresented in the present series. The neural arch
and spine found at 11’ fits fairly well on the centrum found at 11 and I have con-
sidered them as pertaining to the second dorsal. As to the neural spine and arch
there can be little doubt as to this determination, but as to the position of the cen-
trum, it is by no means certain that it does not pertain to the first rather than the
second dorsal. Indeed as regards the length and form of the centrum, character of
the pleurocentral cavities, and position of the capitular rib facet, it would appear to
more properly pertain to the first dorsal than to the second, while the widely separ-
ated position (about four feet) in which they (the centra and neural arch) were
found might be taken as an indication that they pertain to different vertebre. I
have associated this centrum and spine in the same vertebra for no other reason
than that when adjusted to one another they seem to agree fairly well. I believe
it quite possible, even probable, that the centrum pertains to the first dorsal. As
regards the neural arch and spine however, after comparing them with those of the
first dorsal in H. priscus, there can be no reasonable doubt but that they pertain
to the second dorsal. This position is indicated by the character of the anterior
branch of the horizontal lamina which is much less modified to give support to the
scapula than in the first dorsal of H. priscus. The articular surfaces of the post-
zygapophyses have assumed a more perpendicular position in anticipation of the
hyposphene-hypantrum method of articulation that obtains in the median and
posterior dorsals. The neural spine is passing from the widely expanded scoop-lhke
element seen in the posterior cervicals and dorsal one to the simpler form character-
istic of the median and posterior dorsals. The superior branches of the postzygapo-
physial lamina continue, however, confluent with the neural spine, extending to its
very apex and enclosing laterally a rather deep cavity which, nevertheless, is much
HATCHER: OSTEOLOGY OF HAPILOCANTHOSAURUS 33
less deep than in the last cervical of this series or the first dorsal of H. priscus. As
in the succeeding dorsals the distance between the anterior and posterior zygapoph-
yses is much abbreviated. There is a very faint emargination at the apex of the
neural spine with a depth of only 7 mm.
The Third Dorsal (Plate II., Fig. 3, Series 2).— This vertebra lay on end as
shown at 12 in the first diagram. Owing to the position in which it lay in the
quarry its centrum was much shortened by the pressure to which it was subjected,
the ball having been forced down into the pleuro-central cavities and the whole
centrum telescoped as it were. Fortunately the neural arch and spine are in a splen-
did state of preservation. They are still held in position with the centrum, though
the sutures are very distinct. The spine when compared with that of the preceding
vertebra is much modified in the direction of the conditions that obtain in the suc
ceeding dorsals. It has assumed a nearly vertical position instead of being inclined
forward as in the preceding dorsals and cervicals.
It is very much compressed antero-posteriorly and is still connected with the
posterior zygapophyses by the superior branches of the post-zygapophysial lamine.
In the present vertebra however this lamina does not run obliquely upward and
backward in a direct and straight line from the zygapophysis to the top of the neural
spine as in the preceding dorsals and the cervicals, but it extends backward, rising
but little until it reaches the vertical plane of the anterior surface of the spine when
it rises vertically as a thin narrow lamina ascending to the apex of the spine. The
degree of differentiation in the neural spines of this and the immediately preceding
vertebra is the most marked of any of the vertebree even in this region where the
characters of the different vertebree are seen to change so rapidly. The apex of this
spine is also faintly emarginate, the notch having a depth of 9mm. The position
of the capitular rib facet is at the supero-anterior angle of the pleurocentral
cavity.
The Fourth Dorsal (Plate I1., Fig. 4, Series 2).—'The centrum and spine of this
vertebra lay as shown at 13 and 13’ in the diagram. They are both well{preserved,
and the nature of the spine and transverse processes demonstrate beyond a reason-
able doubt that its position in the vertebral column was immediately posterior to
the vertebra just described. The spine is now quite perpendicular and more ele-
vated. It is much compressed antero-posteriorly but somewhat expanded trans-
versely. Its anterior surface is transversely convex, the posterior is concave, form-
ing a long, shallow trough or scoop not nearly so deep as in the preceding vertebree.
The transverse processes in this and the immediately preceding vertebra are assum-
ing a more elevated position, the neural arches are becoming higher and the trans-
>)
34 MEMOIRS OF THE CARNEGIE MUSEUM
verse processes instead of being horizontal are directed successively more and more
obliquely upward as in the succeeding dorsals.
The Fifth, Sixth, Seventh, Kighth, Ninth, Tenth and Eleventh Dorsals (Plate I1., Figs.
5-11, Series 1 and 2).— The neural arches and spines of all these vertebrae were
found interlocked by their zygapophyses as shown in the diagram from 14-20 in-
clusive. The centra of the fifth and sixth had become detached and lay as shown
at 14’ and 15’. The remaining centra were in position at the base of their respec-
tive spines. The neural spines, transverse processes, capitular rib facets, neural
arches, etc., form a regularly graudated series except that the capitular rib facet of the
sixth is much larger than in the other vertebree. The neural arch, spine and trans-
verse processes of the fifth were much injured, but the spine is nearly entire and it
is evident that it pertained to the vertebra immediately posterior to that just
described as indicated also by the centrum. The spine is still compressed antero-
posteriorly but decidedly deeper in that direction than the spine of the verte-
bra just described. In the spine of the succeeding or sixth dorsal the transverse
and anteroposterior diameters are subequal. A hyposphene-hypantrum articula-
tion begins with the sixth dorsal and continues throughout the remaining dorsal
series.
The Twelfth, Thirteenth, and Fourteenth Dorsals (Plate I1., Figs. 12-14, Series 1). —
These vertebree do not differ materially from the same vertebree already described as
pertaining to the type of H. priscus. They were found as shown at 21, 22 and 23
in the diagram, interposed between the series just described and the anterior ex-
tremity of the sacrum. As shown in the diagram the neural arches were in posi-
tion relative to one another but the centra were a little removed from their normal
positions. They are all in a nearly perfect condition.
If the reader has followed carefully the above description of the dorsals pertain-
ing to the present skeleton together with those which pertained to the type of H. pris-
cus and will examine the accompanying figures it will have become apparent that
the complete dorsal series in Haplocanthosawrus must have consisted of not less than
fourteen free vertebree while it is scarcely possible that there were more than four-
teen. This is a very marked increase over the number (ten) which is believed to
have formed the complement of free dorsals in Diplodocus, Brontosaurus and Moro-
suurus. Nor does this increase in the number of dorsals in the present genus seem
to have been made at the expense of the cervical series, for as near as we can judge,
Haplocanthosauwrus, like Diplodocus, was provided with fifteen cervicals. Our deter-
mination of the number of cervicals however does not rest on anything like so good
a basis as does our determination of the number of dorsals but there can be little
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 35
doubt but that the number of presacrals in the present genus exceeded that of the
same series in Diplodocus.
The Sacrals. (Figs. 15, 16 and 20.)
The sacrum of the present skeleton differs considerably from that of the type of
Hf. priscus. Its position in the quarry is shown in the diagram at Sa. Only the
neural spines and diapophyses, the parapophyses of the right side and the centra of
the fourth and fifth sacrals are preserved. ‘The neural spines of the first, second,
third and fourth sacrals are confluent and firmly codssified throughout their entire
length, while the spine of the fifth is coossified only at the top and the bottom
with that of the fourth sacral, and medially there is an elongated foramen between
these spines shown at fin fig. 15. The spines of the two anterior sacrals rise nearly
perpendicular, those of the third and fourth are directed somewhat backward, while
that of the fifth is again directed forward to meet at the apex that of the fourth
sacral. The spines of the first, second, third and fourth sacrals bear diapophysial
laminee, and at the summit these expand into prominent rugosities.
The diapophyses of the first and second sacrals are directed outward, forward and
a little upward, nearly parallel to one another. Those of the succeeding sacrals are
directed outward, backward and a little upward, parallel to one another, but that
of the third meets that of the second in an acute angle at the base of the diapophy-
sial lamina, forming a letter V with the apex directed toward the spine. The dia-
pophysis of the third sacral is formed by the union of branches from the diapophysial
laminee of the second and third sacral spines.
All the parapophyses (sacral ribs) of the right side are present and nearly com-
plete. They all show sutural surfaces for articulation with the sacral centra and
with the iium. The parapophyses are shown in position in the view of the sacrum
from the right side seen in Fig. 15, while comparative front views of the individual
parapophyses are given in Fig. 16, a, b, c, d, é, which represents the series from the
first to the fifth, respectively. Of the five parapophyses the first is the more slender.
It is triangular in outline, with an emarginate base describing a nearly complete semi-
circle. It may be described as composed of a horizontal and ascending branch. The
surface for contact with the centrum is not greatly expanded. There are two sur-
faces for contact with the ilium, one inferior, the other superior and separated by a
distance of about 215 mm. The inferior of these surfaces is more pronounced and
both show prominent rugosities. The anterior surface of this bone is convex, the
posterior concave. Between the inferior and superior surfaces for contact with the
ilium there was an elongated foramen enclosed externally by the ilium and in-
ternally by the ascending branch.
36 MEMOIRS OF THE CARNEGIE MUSEUM
In the parapophysis of the second sacral the horizontal branch has become very
strong and much expanded at either extremity for contact with the centrum and
ilium. The ascending branch is broad but very thin, and presents at its extremity
only a small rugosity for contact with the ilium. The foramen between it and the
ilium was broader than that separating the same branch of the preceding para-
pophysis. Immediately above the surface for contact with the centrum there is a
rugosity which doubtless gave support to the descending branch of the diapophysis,
Fie. 15. Sacrum of Haplocanthosaurus utterbacki, seen from right side, ;'> natural size. a, anterior
extremity ; p, posterior extremity ; 1, 2, 3, 4, 5, spines of first to fifth sacrals; f, foramen between spines
of fourth and fifth sacrals.
Fic. 16. Anterior view of disarticulated parapophyses (sacral ribs) of Haplocanthosaurus utterbacki,
ip hatural size. a to e, first to fifth respectively ; y, surface for contact with sacral centrum ; 2, inferior
surface for contact with ilium ; z, superior surface for contact with ilium.
but it would appear that the diapophysis and parapophysis were not in contact
throughout their entire length, but were separated for a considerable distance by an
elongated foramen. ‘The external extremity of the horizontal branch is greatly
expanded transversely so as to unite with the same element in the succeeding para-
pophysis, to form the acetabular bar and give additional support to the ilium.
The parapophysis of the third sacral differs from that just described in its shorter
horizontal branch and narrower and more slender ascending branch. The shorten-
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 37
ing of the horizontal branch is of course brought about by the inward curvature of the
ilium in the region of the acetabulum.
The fourth parapophysis differs from all the preceding in its widely expanded
ascending branch which curves very gently backward. The horizontal branch is
stout and longer than that of the third parapophysis.
The fifth and last parapophysis differs from all the preceding in the absence of
any ascending branch. The horizontal branch is greatly expanded vertically, and
the articular surface for the ilium is elongate so as to give support to the ischiac
peduncle and posterior blade of the ilium. The principal characters of the para-
pophyses are well shown in the accompanying figures.
Only the centra of the fourth and fifth sacrals are preserved. Fortunately these
are well preserved. ‘The parapophyses of the right side fit very well when adjusted
to their respective vertebrae. Neither of these vertebree show any marked enlarge-
ment of the neural canal. They are both constricted medially, and the distal
extremity of the fifth is considerably expanded for contact with the first caudal.
The centrum of the fifth sacral is decidedly heavier than that of the fourth.
The principal measurements of the different sacral elements in the type of the
present species are as follows :
Distance along crest of the five codssified sacral spines..
wo ay ue ‘¢ four anterior ‘ He
‘« from anterior zygapophyses of first sacral to posterior zygapophyses
Oe 1TH SEYEREA | con oasdosanassncnagondbandadoogoocbGon0onobadeSddoaqadbddesosoboconeaNcEd Ho 575
Hxpansevondiapophysiscot, thirdisacralernc.s.cmccseesccesesc cee ececeeeeeasteceece 375
Greatest length of horizontal branch of first parapophysis.....................0+- 213
He height of ascending ‘“‘ Hales Uo 8) “Qogacce00d0g00 conendeds 300
ub length of horizontal ‘‘ UE excaaysle4G7 5 . dbcodqaodsqanacnonosacaa 146
He height of ascending ‘“ Bene ee rian, 414) Oe ppanddanadoooduaSDséadoS 300
oe length of horizontal ‘ “ third Rb {= ~~ ‘booapasocadoc9a000G0080% 128
ue height of ascending ‘‘ menage Ue he ~sesgbgeaonoNdodbOBRsHEaS 277
es length of horizontal ‘‘ PpacolWudehe Wes <1 Nl opadaesscoososaqobnduads 150
he height of ascending =‘ sea HOS We ie aia nerd aera Meee 250
“ length of horizontal — ‘‘ “* fifth SO OT iS asee eee cisacereerec cee 225
Length of fourth sacral centrum’ .... 2.2.2... 50... ..cecceeeesseeesarscarnenseseesecseseieee 132
Depth ‘' ¢* ee a
Wal cts Sc “ ori ee M POR Area scaempadencaac cedar oo TannoTenesee 132
Length ‘ fifth ae SEAS ee Ui sbaddortocadacohocWbensauusdsdbaocese 120
Depth ‘* “ a eames ee EG a rnaacooracobasqaSnsTesapdoncccnéoon 182
Viol tcc « Le oe ne Wl Gn jadoabaosoobooanoGoondudcodenoades 181
38 MEMOIRS OF THE CARNEGIE MUSEUM
The Caudals. (Plate I1., Series 5.)
Only the seven anterior caudals are preserved in the type of the present species
and these differ very little from the same vertebree in H. priscus, except that the
transverse processes are less well developed and not so much expanded supero-
inferiorly. Owing to the age of the individual the neural arches and transverse pro-
cesses are not codssified with their respective centra although those of the first two
are still held in place. The neural spines are short and stout and very rugose.
That of the first is curved rather strongly backward. All the centra are very short
and biconcave. The anterior zygapophyses are acutely wedge-shaped and extend
well forward with the articular surfaces facing decidedly more inward than upward.
The posterior zygapophyses are only flattened surfaces at the bases of the neural
spines. The transverse processes are suturally connected both with the centra and
the neural arches.
The Vertebral Formula.
From the foregoing descriptions of those portions of the vertebral column pre-
served in the type of the present species and in that of H. priscws it will have become
apparent that we must await future discoveries to determine with accuracy the
vertebral formula of Haplocanthosawrus. The number of sacrals however may be
considered as being definitely fixed at five, while the number of dorsals could not
have been less than fourteen, thirteen of which are represented in the skeleton con-
stituting the type of H. utterbacki. In this skeleton it would appear that only the
first dorsal is missing, and fortunately that vertebra in the type of H- priscus was
found interlocked by its zygapophyses with the last cervical. Although the various
vertebree in the anterior dorsal region of the type of H. wtterbacki were for the most
part found in such a scattered and disarticulated condition as to afford little direct
evidence concerning the exact positions relative to one another which they occupied
in the skeleton during the life of the animal, yet a close examination and careful
study of the vertebrae has convinced me that there are no duplicates among the
thirteen dorsals described and that there can be no question but that all of the thir-
teen are dorsals and that they pertained to the skeleton of one and the same indi-
vidual. That the first dorsal is wanting in this skeleton is shown by a careful com-
parison of the neural arch and spine of the most anterior of this series with that of
the known first dorsal in H. priscus, from which, as has been shown in the descrip-
tions, it differs materially and in the direction of those characters which we should
expect in the succeeding or second dorsal. For these reasons I have referred this
spine to the second dorsal although the centrum which was found detached and
separated, but which in the description and figure I have associated with this spine
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 39
may, as I have already remarked, pertain to the first dorsal. I do not think it at
all probable that more than one dorsal is missing from the series in HZ. utterbacki and
it is with a feeling of considerable confidence that I place the number of dorsals in
this species at least, at fourteen. Fully realizing the character of the evidences |
upon which I have arrived at this conclusion | have spared no pains to present to
the student all the evidence furnished by the material at my command, both as
regards its anatomical characters and the position in which the different bones were
found in the quarry. Aided by the accompanying diagrams and with the type
material at his disposal the future student will be in full possession of all the evi-
dence in the case and will therefore be in a position to decide for himself as to the
worth of my conclusions.
In placing the number of free dorsals at fourteen I am fully aware that this is a
considerable advance over the number that has of late come very generally to be
considered as characteristic of other members of the Sauropoda (Diplodocus, Bronto-
saurus, Morosaurus). Considering however the less specialized nature of the present
genus and the great differences seen, in other important characters, when compared
with the genera just mentioned I do not consider this increase in” the number of
dorsals as at all remarkable, for it is not at all Impossible that the earlier ancestors of
Diplodocus, Brontosawrus and Morosaurus were provided with an equal number of
free dorsals and that the reduction to ten in each of those genera may be regarded
as a specialized character attendant upon and which took place along with that re-
markable specialization which, as is well known, they must have undergone in other
respects and which is most marked in that exceedingly complicated arrangement of
laminz and buttresses seen in the dorsal and cervical vertebree of those genera.
It will doubtless have been remarked that in describing the cervicals I have
placed the number of vertebrz of this region at fifteen, the number present in Dzplo-
docus. It must be admitted, however, that the material at hand does not afford a
very reliable basis for determining the number of cervicals and I should not be at
all surprised if the actual number of cervicals in Haplocanthosawrus should prove to
be one or two less than in Diplodocus. In placing the number at fifteen, as in the
latter genus, I assume that Haplocanthosawrus was provided with four more pre-
sacrals than was Diplodocus. While my estimate of the number of cervicals in the
present genus may prove to be too great, it is hardly possible that it will be reduced
by more than two. This would still give to Haplocanthosaurus two more presacrals
than are present in Diplodocus. It would thus appear that in the various genera of
the Sauropoda the number of presacrals differed and that the number of cervicals is
not entirely dependent upon an increase or decrease in the number of dorsals in any
40 MEMOIRS OF THE CARNEGIE MUSEUM
genus within the group. Still it is easily conceivable that in any genus or species
the presacral formula might vary, without increasing the total number of presacrals
in the individual, according as the exact position in the presacral series at which
the change from cervicals to dorsals took place, and I am inclined to the opinion
that as a rule in any given genus of the Sauropoda where marked specialization has
taken place, there has been a tendency to increase the number of cervicals at the
expense of the dorsal series, due to the gradual shifting of the pectoral girdle from
amore advanced to a more posterior position, by which process anterior dorsals
Have been transformed into posterior cervicals and the cervical region considerably
elongated at the expense of the dorsal. An extreme instance of this is to be seen in
Diplodocus carnegii where the disparity in length in the neck and dorsum, probably
due primarily to the increased number of cervicals and decreased number of dorsals,
has been still further emphasized by the proportionate length of the individual
vertebree in the two series.
If, as does not seem improbable, the total number of presacrals was ever de-
creased in any genus it would appear to have been more readily accomplished by
the successive elimination of the less specialized, or at least more simple, anterior
cervicals than by the disappearance of the extremely complicated dorsals. Nor
does it seem probable or even possible that such a decrease in the number of presac-
rals could have been brought about by the gradual shifting of the pelvic girdle to a
more anterior position. Such anthypothesis presupposes the addition to the sacrum
of successive posterior dorsals and the liberating of posterior sacrals as anterior cau-
_dals, an hypothesis which to the present writer appears unworthy of serious con-
sideration. As to the total number of caudals in the present genus we have nothing
upon which to base anything like a reliable estimate. From the character of the pos-
terior five or six of the series of nineteen anterior caudals in the type of H. priscus
we may judge that while the tail was relatively short the number of caudals was
considerable and probably not less than forty. ‘The increased number of caudals is
indicated in the first place by the character of the chevron found attached to the
thirteenth caudal and which, as already remarked, resembles in general form the
chevron of a caudal occupying a more anterior position in Diplodocus. Moreover, the
very gradual change which is seen to be taking place in the posterior caudals of the
series preserved in the type of H. priscws indicates a very considerable number of
posterior caudals as having intervened between the last of the series and the end of
the tail. The extreme shortness of the centra in the caudals of Haplocanthosawrus
may be considered as sufficient proof that the tail was proportionately rather short
as compared with that of Diplodocus.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 41
After a careful consideration of all the evidence ‘at hand the following is sub-
mitted as the approximate vertebral formula in the present genus. Cervicals 15 ;
Dorsals 14; Sacrals 5; Caudals not less than 40.
Below I give the principal dimensions of the several vertebree pertaining to the
type of the present species. Some of these dimensions have been materially altered
by crushing in such manner as to cause apparent inconsistencies. ‘They should only
be taken as representing in a general way the dimensions of the various vertebre.
In column | the greatest expanse of the transverse processes is given, column
2, greatest length of centrum ; column 38, transverse diameter of centrum at posterior
extremity ; column 4, hight of neural spines above middle of inferior border of
centra in presacrals and above inferior border of posterior end in postsacrals.
4. Cervical. .- 154 645 148 58 61 22 136 53
8. at 210 8} 220 88 108 4 192 73
9. ag ? ? ? ? ? ? ? ?
10. G6 192 74 243 93 102 4 196 73
11. ae Hi 8 238 98 ila 42 225 83
112, a¢ 234 aoe 264 104 113 48 255 10
13. a6 258 104 292 113 125 47 265 | 107%
14, é¢ 268 * 103 112 43
15. as 320 1 || | |
2. Dorsal. 390 154 200 8 125 me 368 143
3 be 374 143 120 43 127 5 499 16
4, be 395 153 150 5t 115 4} 435 174
i. of 390 152 135 54 124 43 483 19
6. bg Heeal50 5a 109 4} 492 193
7 a6 135 5h 116 ree O00, 193
8 66 | 170 68 102 4 550 | 218
9. 6 | 152 6 137 58 540 211
10. ag 148 53 145 5am 558 | «(22
hte ee | 143 58 175 62 585 |
12. & eS 58 169 63 Bie) || Bil
18, be J 35 5a 178 7 560, | 22
14. te Vaal 5 415 170 63 552 213
1. Caudal. 350 133 | 115 43 160 62 425 163
2. a 310 123 80 3h 166 63 410 16}
3. be iD) 3 165 64 393 15}
4, He 83 34 157 64 355 142
5. a 3 31 143 58 346 133
6. a 90 34 130 54 331 13
he “ 85 32 5h 313 12}
133
The Pectoral Arch. (Figs. 17, 18, 19.)
Only the left scapula (Figs. 17 and 18) and right coracoid (Fig. 19) are preserved.
The Coracoid (Fig. 19).— The external surface of this bone is regularly but
gently convex. ‘The internal surface is concave. The anterior and inferior margins
are for the most part thin, but at the antero-inferior angle the margin is thickened
and presents an elongated rugosity shown at a 140 mm. in length and 40 mm. in
42 MEMOIRS OF THE CARNEGIE MUSEUM
greatest breadth, probably for the ligamentous articulation of the sternum. The
surface for articulation with the scapula has a length of 150 mm. and a greatest
breadth, at its junction with the glenoid border, of 90 mm. The coracoid forms
about one half the glenoid cavity and the glenoidal surface meets the surface for
articulation with the scapula at an obtuse angle. Between the inferior margin of
the glenoid cavity and the inferior border there is a rather deep notch in the pos-
Fig. 17. Posterior view of left scapula of H. utterbacki ; g, glenoid surface, ;'5 natural size.
Fic. 18. External view of same; g, glenoid surface ; c, surface for coracoid.
Fig. 19. External view of right coracoid of H. utterbacki; s, surface for scapula; g, glenoid surface ;
1
a, rugosity for supposed contact with sternal, ;'5 natural size.
terior border of the coracoid. The foramen is large and is situated about 35 mm.
beneath the coraco-scapular suture. It is elliptical in outline with the vertical
diameter the longer. ‘The dimensions are 57 mim. for the vertical and 30 mm. for
the transverse diameter at the external opening. The distance from the glenoid
border to the sutural surface at the antero-inferior angle is 295 mm. The distance
from the postero-inferior angle to the anterior border just beneath the coraco-scapular
suture is 350 mm.
The Scapula (Figs. 17 and 18).— The scapula displays the extreme development
of that form shown in Morosawrus with broadly expanded extremities. I cannot
describe the characters better than to give the measurements. The breadth of the
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 43
scapula at the proximal end is 372 mm., at the distal ® 396 mm. and in the middle
only 137 mm. Between the spine and the coraco-scapular suture there is a broad
but shallow cavity on the external surface of the scapula. Above this ridge, how-
ever, the external surface of the scapula is convex in all directions. The total length
of the scapula from its superior or proximal end to the coraco-scapular suture is 800
mm. The accompanying figures show very well the form and principal characters
of this element.
From the foregoing description of the types of the two species of Haplocanthosau-
rus at present known and from the accompanying measurements and figures it will
appear that as compared with Diplodocus the present genus was represented by
animals with the thorax somewhat more elongated and with neck and tail relatively
shorter than those which obtained in representatives of the former genus. While,
judging from the femur, the only element of the limbs at present available, the ap-
pendicular skeleton was relatively strong when compared with the axial.
Distinctive Generic Characters.
The principal characters distinguishing the genus Haplocanthosawrus may be
summarized as follows :
Frrsv. — Neural spines short and simple (not branched) throughout the entire verte-
bral column.
Second. — Newral spines of the anterior sacrals codssified, forming an elongated bony
plate.
Turrp. — Height of newral arches in posterior dorsals exceeding length of neural
spines.
Fourtu. — Transverse processes of the anterior and mid-dorsal regions inclined wp-
ward and outward instead of directly outward.
The two species described above may be distinguished as follows by their respec-
tive sacra.
HAPLocANTHOSAURUS PRISCUS. — With newral spines of the three anterior sacrals
coossified, those of the two posterior remarnung free.
HAPLOCANTHOSAURUS UTTERBACKI. — With newral spines of the fowr anterior sac-
rals coossified throughout their entire length and with that of the fifth sacral at the top and
bottom.
Taxonomy.
All systematists will, I think, agree with me in placing the present genus
y ) p ENS § 8
among the Sauwropoda. The scapula, coracoid, pelvis and sacrum, as well as the
presence of large pleurocentral cavities in the presacral centra, together furnish
°T consider that end of the scapula which opposes the humerus as the distal.
44 MEMOIRS OF THE CARNEGIE MUSEUM
conclusive evidence as to its affinities with that group of the Dinosauria. Never-
theless there are a few characters, such as the elongated neural arches and compara-
tively short and absolutely simple neural spines of the dorsal and posterior cervical
series, the conformation of the transverse processes and position of the capitular rib
facets, which are different from any other known member of the Sauwropoda of North
America and are more nearly paralleled by the characters which exist in the verte-
bree of this region in some members of the Predentata, more especially in Stegosawrus,
where, as in Haplocanthosaurus, the neural spines are short as compared with the ele-
vated neural arches from the summit of which, in each case in the posterior dorsals,
the transverse processes spring and diverge from the bases of the perpendicular
spines at angles of about 45° instead of being directed horizontally as is the usual
manner in the Sawropoda. The presence of characters so similar as those just men-
tioned in representatives of the Sawropoda and the Predentata while certainly not
indicative of any very close relationship may perhaps be considered as evidence of a
remote common ancestry for the two groups. If this view be taken, these characters
possessed in common would be considered not as parallel or analogous characters de-
veloped independently in each instance, but as persistent primitive characters which
were present in their remote but common ancestors. As the development of the
two groups progressed and they became more and more differentiated, such charac-
ters proved advantageous and became more emphasized in the Predentata while in
the Sawropoda, where for some reason they were not particularly advantageous, they
were gradually eliminated and disappeared altogether in the more highly specialized
forms though persisting in the more primitive Haplocanthosaurus. Itis by the pres-
ence in common, among the Sawropoda, Theropoda and Predentata, of such charac-
ters as those just described, that the student of the Dinosauria will find the most
trustworthy evidence as to the actual relationships or want of relationships in the
three groups. Not until a considerable number of genera within each group are
known from a detailed study of the osteology of fairly complete skeletons will it be
possible to pronounce with any degree of certainty even upon the question as to
whether the Dinosauria is a natural group as maintained by Marsh or an entirely
unnatural one, without any right to existence, into which has been thrown three
distinct groups, totally dissimilar and with nothing in common, as was held by the
late Dr. George Baur. In the discussion of this question however there are several
points which should be constantly kept in mind by the advocates of either view.
Among these are:
Frrst.— Those who are opposed to considering the Dinosauria as a natural group
should bear in mind the great antiquity that must be accorded to that group when
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 45
considered as constituting a single group. Evidence of such antiquity is found not
alone in the great diversity exhibited by the three subdivisions into which the
group as a whole has been divided but by the diversity and specialization exhibited
by the different families, genera and species within each of these three subdivisions,
As yet we know comparatively little of the earlier Dinosawria and the group if in
reality it be a natural one is at present represented in our museums for the most
part only by the later and more specialized forms. Of the Sawropoda we know only
those forms which lived just prior to their extermination when they were already
highly specialized. Consider for a moment the enormous time interval which must
have been necessary for the development of a reptile like Diplodocus. Yet his
remains are found associated in the same quarry with those of Haplocanthosawrus.
the most primitive Sauropod known, and the entire range of the Sauropoda through-
out the geological column in North America so far as at present known is limited
to certain horizons in the Jurassic or Lower Cretaceous of some authors, with a ver-
tical thickness never exceeding a few hundred feet, and from the top to the bottom
of which there are always found forms which are highly specialized, conclusive
proof that the paleontological record is exceedingly incomplete as regards this
group.
Although the time distribution of the Theropoda and Predentata as we now know
it is more considerable than that of the Sawropoda yet it is by no means complete
and we know little of the earlier forms of either of these divisions. The wonder
therefore is not that the three divisions as we now know them should show so little
in common, but rather that, considering their great antiquity and early differentia-
tion, they should have continued to possess in common even such characters as they
do show.
Seconp.— Although due weight should be given to every marked and important
difference in structure it should nevertheless be borne in mind that every character
‘possessed in common by these three divisions or by any two of them should be con-
sidered as an evidence of relationship until definitely proved to be fortuituous or as
having been developed independently in each instance.
Turrp.—It isin the, as yet undiscovered, earlier and more generalized members
of these groups that we must look for those characters which will throw most light
on this question. If future explorations should be rewarded by the discovery in
the early or middle Trias of a considerable number of representatives of each of the
groups which we now refer to the Dinosawria, and if together they were shown to
possess many characters in common and to approach one another much more nearly
than do the Jurassic and Cretaceous forms, this evidence would be considered as
46 MEMOIRS OF THE CARNEGIE MUSEUM
strongly favoring the including of them all in a single group, the Dinosawria. If on
the other hand they were found to show little in common or were even more widely
separated than are the later forms from the Jurassic and Cretaceous then there
could be no reasonable grounds for considering the Dinosauria as a natural group
and it would become necessary to discard that term, at least in the sense in which
it is at present used.
Since however as has been shown in Haplocanthosawrus, the Sawropoda and Pre-
dentata do possess certain important anatomical characters in common and since in
this the most generalized genus of the group at present known the relationships
between these two groups are more apparent than in the more specialized genera,
Diplodocus, Brontosaurus, etc. It is reasonable to suppose that in the yet undis-
covered but still more generalized forms greater similarity in structure will be found.
Moreover from our present knowledge the relationships between the Sawropoda, Pre-
dentata and Theropoda, as has already been pointed out by Marsh, is indicated by
a number of important characters possessed in common such as:
“1. Teeth with distinct roots either fixed im more or less distinct sockets or in longitudi-
nal grooves, never ankylosed, no palatal teeth.
“2. Skull with superior and inferior temporal arches.
“3. Double-headed cervical and thoracic ribs.
“4. Sacral vertebre codssified and more numerous than in other reptiles, seldom less
than five.
“5. Tliwm extended in front of acetabulum, in the construction of which latter the ilium,
ischium and pubis take part. :
“6. Fibula complete.
“7. The reduction in the number of digits commences with the fifth.”
The present author is, therefore, of the opinion that the Dinosauria should be
regarded as a valid and distinct group for the exact definition and description of
which we must await further discoveries as also for definite proof that the different
groups now included in it are actually related.
Admitting that the Dinosauria do constitute a natural group we have next to
consider the rank that should be accorded to it in any general scheme looking to a
classification of the Animal Kingdom as a whole. Here again we find there has
been great diversity of opinion. Without reviewing the various opinions that have
at various times been expressed upon this subject it would appear to the present
author, that, in consideration of the diversity in form, structure and habit which are
found withing this group where some members are carnivorous and others herbiv-
orous, some quadrupedal and others bipedal, some heavily armored and others un-
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 47
armored and with all the many and diverse anatomical characters shown in their
osteology which might reasonably be expected from such diversity of habits, there
would seem good reasons for considering the Dinosauria as deserving of the rank of
a subclass of the Reptilia comparable for example with the Metatheria of the
Mammalia and divisible into three orders for each of which several names have
been proposed by various authors. Of all these, those proposed by Marsh appear to
the present writer to be the most appropriate, these are :
1. THe THeropopa ; Hmbracing all the carnivorous dinosarus.
2. THE Sauropvopa ; Hmbracing all the herbivorous forms in which the predentary is
wanting.
3. THE Prepentata; Hmbracing all the herbivorous forms im which the predentary
is present.
In accepting the terms Theropoda and Sauwropoda rather than Megalosawria and
Cetiosauria I do so out of regard for the more comprehensive nature of those terms
as used by Marsh. The latter terms as used originally by Fitzinger (Megalosawri),
1843, and Seeley, 1874, respectively, I consider of subordinal rank only. Predentata,
Marsh, is preferable to Orthopoda, Cope, because it is in no sense coordinate with the
latter but a much more comprehensive term. Cope’s Orthopoda and the Ornithopoda
of Marsh (not Huxley) are more nearly synonymous.
Some authorities have considered the Sawropoda of Marsh (1878) as a synonym
of the Opisthocelia of Owen (1859). But this appears to me quite unwarranted.
For the latter term, although haying priority, was never adequately defined by
Owen. It was originally proposed as a suborder of the Crocodilia’ and was char-
acterized as embracing members of that group with opisthoccelous dorsal and cervical
vertebrae. Owens’ original definition of the Opisthocelia was as follows: ‘The small
group of Crocidilia, so called, is an artificial one based upon more or less of the
anterior trunk vertebrae being united by ball-and-socket joints, but having the ball
in front, instead of, as in modern crocodiles, behind.” As is now well known, the
above character in no way distinguishes these dinosaurs from members of either the
Theropoda or Predentata, and on the same page, in defining the order Dinosauria,
Owen describes the cervical vertebree as being opisthoccelous in some species. It is
thus clear that Owen not only did not adequately define his proposed suborder
Opisthocelia, but that he did not recognize its real relationships as being with the
Dinosauria rather than the Crocodilia. The character given distinguishes it from
the Procelia or true Crocodilia, but should be considered as uniting it with, rather
than separating it from, the Theropod and Predentate dinosaurs, for as has already
7See Report 29th meeting Brit. Assoc, Ady. Sci., 1859, pp. 164, 165.
48 MEMOIRS OF THE CARNEGIE MUSEUM
been stated this character is possessed in common by members of both these groups.
While Cetiosaurus is an undoubted member of the Sawropoda (Opisthoccelia) as
determined by Owen, this fact does not serve to define properly the latter term which
remains a nomen nudum, while the Sawropoda, proposed and defined by Marsh in the
American Journal of Science for November, 1878, page 412, should be accepted as
the first adequately defined name for this group of dinosaurs.
In proposing the term Sauropoda for this group of dinosaurs in the paper just
cited Marsh adds:
“The most marked characters of this group are as follows:
“1. The fore and hind limbs are nearly equal in size.
“2. The carpal and tarsal bones are distinct.
“3. The feet are plantigrade, with five toes on each foot.
“4. The precaudal vertebree contain large cavities, apparently pneumatic.
“5. The neural arches are united to the centra by suture.
“6. The sacral vertebree do not exceed four, and each supports its own transverse
process.
ry
“7, The chevrons have free articular extremities.
“8. The pubes unite in front by ventral symphysis.
“9. The third trochanter is radimentary or wanting.
“10. The limb bones are without medullary cavities.”
Although the subsequent discovery of more complete material has shown that
No. 6 of these characters is erroneous, and that certain others are possessed in common
by some members of the Theropoda and Predentata, yet Marsh’s original definition
still remains fairly diagnostic of the group, and the term Sawropoda should, there-
fore, it appears to the present writer, be accepted.
Whether this group should be considered as of only subordinal rank, as originally
proposed by Marsh or as-of ordinal value as considered in his later publications, is a
question concerning which there is at present no unanimity of opinion. Each
student must, for the present at least, determine for himself the rank to be assigned
such groups, and such decisions will necessarily be determined by, and vary accord-
ing as certain characters are considered as of greater or less importance by the dif-
ferent investigators. Without going into an extended discussion of this question the
present author feels warranted in considering the Sawropoda as a distinct order, com-
parable for instance with the Ungulata among the Eutherian Mammalia, or the Dipro-
dontia among the Metatheria, according to Gadow’s “Classification of the Vertebrata.”
It now remains to discuss the relations of the genus Haplocanthosawrus to the
various genera and families of the Sauropoda that have already been proposed.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 49
This will be the more easily understood if we first notice briefly the principal char-
acters of the different families within that order.
Marsh has divided the Sauropoda into six families which he has named as fol-
lows: (1) Atlantosawride ; (2) Diplodocide ; (3) Morosawride ; (4) Plewrocelide ; (5)
Titanosauride ; (6) Cardiodontide.
As already stated in my memoir on Diplodocus, when discussing the taxonomy
of that genus, it is not improbable that the number of families recognized by Marsh
is too great and should be somewhat reduced. However it would seem premature
to attempt a revision of the genera and families of this group until the large and
splendid collections recently brought together by the Carnegie Museum, the Ameri-
can Museum and the Field Columbian Museum have been thoroughly studied. It
is safe to say, however, that no such reduction in the number of families as that
proposed in the second yolume of the English edition of Zittel’s ‘“‘ Text-Book of
Paleontology ” will become necessary. Nor will it be found necessary or desirable
to associate in the same family genera so different as are Brontosuwrus and Moro-
saurus as was done in the volume just cited.
From the foregoing description of the types of Haplocanthosawrus priscus and
H. utterbacki it will readily appear that the affinities of that genus are with the
Morosawride. The relationships with that family are shown by the expanded
superior extremity of the scapula; the general form of the different pelvic elements,
more especially the pubes and ischia; the simpler structure of the presacral verte-
bree ; the short spines of the dorsals and sacrals; the biconcave centra of the cau-
dals and in the relative breadth and height of the sacrum. While the relationships
with the Morosawridx are clearly indicated by the presence of these and other char-
acters of scarcely less importance, yet there are present certain characters even more
marked than most of those which at present serve to distinguish even the most
widely separated families of the Sawropoda now known. ‘These are the perfectly
simple neural spines of the anterior dorsals and posterior cervicals; the different
position in the sacrum of the sacrals with codssified sacral spines; the greater num-
ber of dorsal vertebree and the much simpler structure of the individual vertebree
throughout the entire vertebral column. Such differences as these will doubtless be
considered by some as of family or at least subfamily importance. Since, for the
most part at least, they are only such differences as we might reasonably expect to
find among the more primitive and less highly specialized members of that family
I prefer to regard Haplocanthosaurus as pertaining to the Morosawride and including
species the most generalized of any yet known in that or any other family of the
Sauropoda.
50 MEMOIRS OF THE CARNEGIE MUSEUM
It is easy to see how the shght emarginations at the summit of the neural spines
in the anterior dorsals and posterior cervicals of H. utterbacki might become succes-
sively more and more emphasized until they became actually bifid, at least to the
extent that obtains in Morosaurus. Nor is it impossible that*while specialization
was taking place along this particular direction, the centra, transverse processes, etc.,
may have assumed a more and more complicated structure. It is by some such
process as this that the present author conceives that Morosawrus was developed
from some earlier and more primitive form which was alike ancestral to that genus
and Haplocanthosaurus, the latter, however, being less progressive than the former,
became less modified and preserves more nearly the general form and character of
their common ancestral stock. a
Although representatives of both genera lived contemporaneously, as is shown
by the presence of undoubted remains of both in the Canyon City quarry, it is more
than probable that Morosaurus long survived the other since remains of that genus
are abundant near the top of the Jurassic in southern Wyoming while as yet no
remains of Haplocanthosauwrus are certainly known to have been obtained there.
RELATIONS oF HAPLOCANTHOSAURUS TO EUROPEAN AND SoutH
AMERICAN Forms.
As yet we have only discussed the relations of the present genus with North
American representatives of the Sauropoda. It now remains to notice briefly the
relations of this genus with certain forms from Europe and South America with
which it seems to show some relationships.
Chondrosteosaurus compared with Haplocanthosaurus.
The characters exhibited by the vertebral centra of Haplocanthosawrus are some-
what similar to those shown in Chondrosteosawrus gigas Owen, founded on the cen-
trum of an anterior dorsal or cervical vertebra from the Wealden of the Isle of
Wight and described and figured in Supplement (No. VI.) to the Monograph of the
Fossil Reptilia of the Wealden and Purbeck Formations ; pp. 5 to 7, Plates LI-V, of
Owen's British Fossil Reptiles, London, 1876.
This resemblance is especially apparent in Plate II., Fig. 2, and Plate V., Fig. 1
of the paper just cited. However the resemblances shown are not of a character
which would indicate a closer relationship between Chondrosteosawrus and Haplo-
canthosaurus than between the former of these with any one of several other genera
of the Sauropoda. Nor is it possible, owing to the fragmentary nature of the ma-
terial upon which that genus was based, to decide with certainty even as to just
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS oul
what family of the Sauropoda it pertained, although Owen’s statement that the
transverse diameter of the centrum exceeds the vertical together with the broad,
almost flat inferior surface and other characters indicated by the figures would seem
to fix this centrum as pertaining to a median cervical of some member of the Moro-
sauride.
Comparison of Haplocanthosawrus with Bothriospondylus Owen.
A comparison of the various characters exhibited by the species of Bothriospon-
dylus described by Owen in his Monograph on that genus published as part II., of his
Reptilia of the Mesozoic Formations, pp. 15-26, Plates III.-IX., will show many strik-
ing resemblances to Haplocanthosaurus and at first sight one might be led to believe
that they pertain to the same genus as that to which the material under discussion
pertains. A closer examination however will reveal several striking and important
differences certainly to be regarded as
of generic if not of family rank. Such
distinctive characters are especially ob-
servable in the sacral centra as will be
seen by a comparison of Fig. 20 show-
ing side views respectively of the fourth
and terminal sacral centra of the type of
Haplocanthosawrus utterbacki with the
figures on Plates III. and IV., of the
monograph just cited. As will be seen
Fig. 20.
by an examination of Owen’s figures and
text the median sacrals of Bothriospon- Fig. 20. 4. Centrum of fourth sacral of Haplo-
4 Y canthosaurus utterbacki seen from right side (No. 879).
dylusare provided with both anterior and 2 © )
5. Same view of fifth or last sacral centrum of same.
posterior parapophysial facets, while in goth + natural size.
Haplocanthosaurus as shown in Fig. 20,
there is but a single facet, which in the centrum of the fourth sacral is median in
position antero-posteriorly, but somewhat elevated above the median longitudinal
line. Furthermore the pleurocentral cavities so conspicuous in the sacral centra of
Haplocanthosawrus beneath the parapophyses are entirely wanting in Bothriospon-
dylus. The cavities in the sacrals of that genus shown at f, Plate III., Figs. 2 and
4, and Plate IV., 4 and 5, of Owen’s Monograph lie above the parapophyses and are
not homologous with the pleurocentral cavities. These characters as well as others
of only less importance are quite sufficient to distinguish Huplocanthosaurus from
Bothriospondylus,
52? MEMOIRS OF THE CARNEGIE MUSEUM
Comparison of Haplocanthosaurns with Cetiosawrus Owen.
Of all the British representatives of the Sawropoda perhaps the most striking
resemblances to Haplocanthosawrus are to be found in Cetiosawrus longus Owen, as
shown in remains representing a considerable portion of a single skeleton discovered
in 1868-70 in quarries of the Great Oolite of Enslow Rocks at Kirtlington Station,
SS
————
eS
Ba =
EZEZ—E ==
LIE
<=
——
——
<=
Right. Left.
Fig. 21. Scapule of Cetiosaurus longus Owen, after Owen, {5 natural size ; hh, surface for articulation
with humerus; c, surface for articulation with coracoid.
eight miles north of Oxford, and showing so close a resemblance to the type of
C. longus that it was used by Owen in his detailed description of that species in his
Monograph on the genus Cetiosawrus in Part I. of his British Fossil Reptiles of the
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 53
Mesozoic Formations, pp. 25-43, Plate X., text figures 1-10. The resemblances in
these two forms are very numerous and are to be seen in the scapulee, as compare
Figs. 18 and 21; the femur, the ilium and the vertebre as figured and described
by Owen in his Monograph. Indeed, if the vertebra described as an anterior dorsal
in the last paragraph on page 29 of his monograph is really an anterior dorsal
this resemblance would seem to be more than superficial, for according to Owen’s
description the neural spine seems to be quite simple and the diapophyses are de-
scribed as being directed upward and outward at an angle of 45° with the neural
spine, characters precisely like those already described as obtaining in Haplocantho-
saurus. Unfortunately Owen does not figure this vertebra, and were it not for the
fact that he describes it as being massive, one
might readily believe on the evidence of this ver-
tebra alone that it pertained to a genus closely
related to or identical with those remains which
I have made the type of Haplocanthosawrus.
However the vertebrae of Haplocanthosawrus can
by no means be considered massive when com-
pared with the vertebrae of other members of the
Sauropoda. Moreover, in Haplocanthosaurus the
vertebrae show numerous large intra-mural cavi-
ties instead of the close, though cancellous tex-
ture of these bones, resembling that which obtains
in the whales, which is present in the British
genus and which suggested the generic name Fig. 22, Coracoid of Cetiosaurus longus
Cetiosaurus. This difference in character would Owen, after Owen. {5 natural size. sc,
seem a very important one, if it were shown to SUT#ce for seapula; 4h, surface for
exist in those vertebrae of Cetiosaurus which are ™“™™"*
most cavernous in Haplocanthosaurus. ‘There are, however, other and quite striking
differences, notwithstanding the general similarity in the o:teology of these two genera.
The coracoid, according to Owen, is especially different, as will be apparent after a
comparison of Figs. 19 and 22. If Owen’s figure is correct the coracoid of Cetio-
saurus is without a foramen, a character which, if correct, is entirely unique, in so
far as am aware, among not only the Sauropoda but ithe herbivorous dinosaurs
generally. It appears to me quite possible that Owen’s figure is erroneous and that
the coracoid is so distorted or imperfect as not to show a foramen in the example
from which his drawing was made. It does not seem possible that such a striking
difference could normally have existed in the coracoids of two genera otherwise so
54 MEMOIRS OF THE CARNEGIE MUSEUM
closely related as these genera appear to be and I would suggest a reéxamination of
the British specimen by those who may have access to it. While Haplocanthosawrus
and Cetiosaurus are certainly generically distinct | believe they may pertain to the
same family.
Although there are undoubted close similarities in structure between Haplocantho-
saurus and the three genera of British Sauropods mentioned above, yet, if we can rely
upon the characters represented by a single vertebra, it isin South America that
there has been found the remains of a Sauropod dinosaur showing the closest
relations with this genus. I refer to the dorsal vertebra recently described and
figured by F. Baron Nopsca ° and provisionally referred by him to Bothriospondylus.
From Nopsca’s figures, it will be seen that from the material at hand it is not
generically distinguishable from Haplocanthosaurus the corresponding vertebra of
which it resembles very closely. Note for instance the simple neural spine, elevated
diapophyses, high neural arches, reduced centra, character and arrangement of the
various lamin, position and character of the tubercular and capitular rib facets,
all of which are characters similar to, indeed almost identical with, those found in
the median dorsals of Haplocanthosawrus. Without claiming that the two are
undoubtedly congeneric I wish to emphasize the very striking similarity in structure
which they exhibit.
From the above study of the material constituting the types of Haplocantho-
saurus priscus and H. utterbacki the present author feels justified in regarding that
genus as representing the least specialized member of the Sauropoda yet discovered.
Of the families of Sauropoda already proposed its closest affinities are undoubtedly
with the Morosawride and I prefer to include it in that family rather than to erect
for it a new family, although some will doubtless think it deserving of the rank of
a distinct family, the Haplocanthosawride. According to that classification of the
Dinosauria which it appears to me is most acceptable, considering our present
knowledge of the group, the taxonomy of Haplocanthosauwrus would be best expressed
by considering it as a well-marked genus of the family Morosauridx of the order
Sauropoda, subclass Dinosauria, class Reptilia.
Probable Habits of the Sawropoda.
Great diversity of opinion has been expressed by various authors regarding the
habits of the different genera of Sauropod dinosaurs. Owen, on page 39 of his
“ Fossil Reptilia of the Great Odlite,” speaks as follows of the probable habits of these
®Sitzungsberichten der kaiserl. Akademie der Wissenschaften in Wien. Mathem. naturw. Classe,
Bd. CXL., Abth. 1, Feb., 1902, pp. 108-114.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 5D
giant reptiles. ‘These enormous Cetiosauwri may be presumed to have been of
aquatic and, most probably, of marine habits. ...” Seeley® at one time con-
sidered representatives of the genus Bothriospondylus (Ornithopsis) as “clearly
ornithic” and this idea suggested to him the name Ornithopsis for those reptiles.
Osborn in his memoir entitled “ A Skeleton of Diplodocus” leans to the aquatic
habits of these reptiles, holding that the tail is especially modified to function as a
swimming organ and was provided distally with a “ vertical fin” ! He believes the
chief function of the tail to have been that of a propeller to aid the animal in swim-
ming and that it functioned secondarily as a balancing and supporting organ.
While holding that the Sauropoda (Cetiosaurs) are aquatic and quadrupedal, he
‘infers that they were capable of migration on land and of assuming both a bipedal
and tripodal position, the tail when in the latter position functioning as a third
support in conjunction with the hinder pair of legs.
Marsh was the first to advance the aquatic habits of Diplodocus, haying con-
sidered the position of the narial opening as suggestive of such habits. In his
memoir on Diplodocus the present author accepted an aquatic life as that to which
representatives of that genus seemed best adapted when considered from their anato-
mical structure as a whole. I remarked in that connection “That I was inclined to
the opinion that Diplodocus was essentially an aquatic animal, but quite capable of
locomotion on land.”
So similar in general form and anatomical characters are the different genera of
the Sauropoda that we may consider the group as a whole as a remarkably homo-
geneous one, with quite similar though not identical habits characterizing most if
not all of its representatives. It would seem therefore more advantageous to discuss
the probable habits and mode of life of the group as a whole than those of any
particular genus. In any such discussion there are several classes of facts from
which evidence more or less important can be obtained bearing upon the subject.
Among these may be mentioned the following :
1. The anatomical or osteological characters of the group.
2. The natwre of the other associated fossils vncluding vertebrates, invertebrates and
plants.
3. The condition in which the remains are found imbedded in the matrix.
A. The nature of the matrix in which the remains are found.
Let us next consider in the order enumerated above the evidences afforded as to
the life habits of the Sauropoda by these four classes of testimony.
First: As to the evidence furnished by the osteological and anatomical char-
*“ On Ornithopsis, a gigantic animal of the Pterodactyle kind, from The Wealden.’’ Annals and
Mag. of Nat. Hist., 1870, p. 279.
56 MEMOIRS OF THE CARNEGIE MUSEUM
acters of the Sauropoda. As already observed, Marsh has considered the elevated
position of the anterior nares in Diplodocus as evidence of an aquatic life. But this
evidence, although strongly presumptive, is by no means conclusive, for there are
among living terrestrial vertebrates similar conditions, more especially among the
mammalia accompanied by habits which are in no sense aquatic, while such essen-
tially aquatic or amphibious reptiles as the crocodiles, alligators and gavials have
the narial opening in its normal position at the distal extremity of the snout. In
like manner Owen’s remark that, “These enormous Cetiosawri may be presumed to
have been of aquatic and, most probably of marine habits, on the evidence of the
coarse cancellous tissue of the long bones which show no trace of medullary cavity ”
is not conclusive since some of the most strictly aquatic reptiia and mammalia, as
for instance the Manatee among the latter class have exceptionally dense and heavy
bones. However this exception is not so important, as it might at first glance seem
to be, for it is a well-known fact that in the Manatee the bones have acquired greater
density and increased specific gravity in order to aid these animals in retaining a
submerged position while feeding on the aquatic plants found growing on the bot-
tom of the shallow waters in which they live. It is clear that the feeding habits of
the Sauropoda required no such modification of the osseous skeleton, and that if
modified at all for aquatic habits, it was in the direction of a more open and cancel-
lous structure even than that which obtains in the Cetacea and calculated not only
to give greater buoyancy to these massive quadrupeds when in water but, in addi-
tion, to give the greatest possible surface for muscular attachment compatible with
the required rigidity and with the least possible weight. As Osborn, in speaking of
the axial skeleton of Diplodocus, has truly remarked ‘“ It 1s a mechanical triumph of
great size, lightness and strength.” Although the present writer cannot agree with
Osborn in considering the chief function of the tail as that of a swimming organ,
even in Diplodocus the most highly specialized member of the order, with the prob-
able exception of Barosawrus; and while there seems to be no evidence whatever of
the presence of a vertical fin on the tail of that genus as suggested by Osborn, I
nevertheless believe that all the Sauropoda were aquatic to the extent that they fre-
quented swamps, marshes, lakes and streams, that they were capable of swimming
and that when moving about by that method the tail was a very efficient propelling
organ. From the character of the modification of the chevrons of the mid- and
posterior caudal regions in all the Sauropoda, however, I believe that when in a
normal position, whether on land or in water, the greater portion of the tail rested
on the surface of the ground, and I am not prepared to say that its function as a
balancing, supporting, or defensive organ, was secondary to its function as a swim-
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 57
ming organ. In arriving at any reasonably accurate conception of the habits and
usual mode of locomotion of these dinosaurs the structure of the limbs would seem
to be of the very first importance although they have not as yet been given more
than a passing consideration in this connection. The limbs and feet are now pretty
well known in several genera of the Sauropoda and in so far as the Jurassic forms
are concerned their structure shows them all to have been strictly quadrupedal, with
massive and rather long limbs and feet both behind and in front, the fore limbs with
one or two possible exceptions being the shorter. In no instance yet discovered is
there any marked or even noticeable tendency toward abbreviating or subordinating
the limbs along the lines so universally characteristic of the aquatic or amphibious
recent reptilia or mammalia. The structure of the limbs and feet in all the Sauro-
pod genera, where those elements are known, furnish to my mind conclusive evi-
dence that they were first of all ambulatory organs whose usual and normal func-
tions were to give support to the body and enable the animal to walk about over
the dry earth or to wade in the shallow rivers, swamps, lakes and other waters
while in search of food. When engaged in the latter occupation their great size
aided by their long necks would enable them to essay waters of no inconsiderable
depth with little inconvenience.
Second: As to the character of the associated fossils.
Wherever remains of Sauropod dinosaurs have been found in this country, there
has been found associated with them in more or less abundance the remains of
Theropod and Predentate dinosaurs. The classic quarry near Canyon City, Colorado,
where the types of the two species of Haplocanthosawrus described above were found
has also yielded remains of Diplodocus, Brontosawrus, Morosawrus,, Ceratosawrus,
Allosaurus, and Stegosawrus, besides other dinosaurs, crocodiles, turtles, fishes and
diminutive mammals all from the immediate sandstones in which the dinosaurs
occur. While in the adjacent clays there are numerous small lenticular masses of
limestone abounding in fresh-water gasteropods and remains of small fresh-water
plants. And in the clays themselves at a few especially favored localities the shells
and casts of several species of Unio occur in great abundance. ‘The character of the
fauna and limited flora found associated with these dinosaurs, whether considered
individually or as an assemblage is not what would be regarded as strictly aquatic.
Such aquatic forms as are found like the bivalves, gasteropods, fishes, turtles and
crocodiles indicate for the most part shallow waters or at least streams and lakes of ©
only moderate extent and depth.
Third: The condition of the remains as they lay imbedded in the matrix will
afford some evidence as to the habits of the Sauropoda, for it is evident that if these
oe)
58 MEMOIRS OF THE CARNEGIE MUSEUM
ponderous reptiles were actually aquatic they would require bodies of water of no
inconsiderable dimensions and depth for their accommodation and it is not at all
reasonable to suppose that when overtaken by death either through disease, old
age or the attacks of their enemies they would seek other than their natural habitat.
Therefore if they lived and died in deep water, after death it is only natural to
suppose that their carcasses would sink to the bottom and become buried beneath
the accumulating sands and clays with the different bones of the skeleton still in
their proper positions relative to one another. For among the strictly aquatic
forms (crocodiles, etc.) that lived contemporaneously with them there are none
sufficiently large and powerful to disarticulate the skeletons of these gigantic
dinosaurs. Since in most instances the skeletons of these dinosaurs are found in
even more disarranged and dismembered conditions than were the two described in
the present paper it seems far more probable that, as a rule, they have met death in
or adjacent to shallow waters, or on land where their carcasses were accessible to
the terrestrial carnivorous Dinosauria, to the ravages of which the dismemberment
of the skeletons is partially due, as is sometimes evidenced by the tooth marks still
preserved on the bones: silent but unmistakable evidences of those prehistoric
feasts.
Fourth: The character of the enclosing matrix furnishes important evidence as to
the nature of the habitats of those animals whose remains it contains, especially if
considered in connection with the character of the surrounding and overlying sedi-
mentary rocks. If as is the case at the quarries near Canyon City, Colorado, the
bones are found imbedded in lenticular masses of coarse sandstone showing cross-
bedding it is evident that such deposits were laid down in comparatively shallow
waters. Furthermore, if as is the case throughout the Jurassic generally, wherever
important deposits of dinosaur remains have been found, massive, coarse, or finely
bedded sandstones with or without frequent examples of cross-bedding are found
replacing the finer, more evenly bedded clays and shales both vertically and later-
ally even at moderately frequent intervals, it is evident that such beds were not
deposited in deep and quite waters; that the immediate region, which by every
reasonable presumption should be considered the habitat of the dinosaurs, presented
the appearance not of a great sea or lake, but rather of a flat and open country
where streams were constantly shifting their courses and the smaller lakes and
bayous, though confined within more fixed limits than the streams were not entirely
stable. That the country was flat and open rather than mountainous is shown by
the absence for the most part, of coarse conglomerates.
Ot
Wo)
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS
Conclusions as to the Habits of the Sawropoda.
In discussing the probable habits of Diplodocus in his memoir on that genus, the
present author remarked : “JI am inclined toward the opinion that Diplodocus was
essentially an aquatic animal, but quite capable of locomotion on land.” I would
now after a more thorough study of the osteological characters of several Sauropod
genera in connection with more extended geological observations since conducted
and bearing upon the probable physiographic features, during Jurassic times, of the
regions in which Sauropod remains have been found in more or less abundance,
amend this statement of my opinion as follows, making it applicable to the Sauro-
poda generally. ‘
I believe: That the Sauropoda were essentially terrestrial reptiles with amphibious
habits, passing much, perhaps most, of their time in shallow waters where they were able
to wade about im search of food. That their natural and normal mode of progression
was ambulatory, as is abundantly evidenced by the structure of their feet and limbs, but
that they were quite capable of swimming when throwgh choice or necessity they essayed
the deeper waters of the larger lakes and streams, to which they must frequently have
been driven to resort for protection from their natural enemies, the contemporancous
carnworous Theropoda with habits probably still less aquatic than were those of the
Sawropoda.
Origin of the Atlantosaurus Beds.
I have elsewhere (An. Car. Mus., Vol. I., pp. 8327-341) described in some detail
the geology of the country in the immediate region of the dinosaur quarries near
Canyon City, Colorado. It may be of interest in this connection however to
describe in greater detail some of the more important quarries of this region, ren-
dered classic by the researches of the late Professors O. C. Marsh and E. D. Cope,
and to describe the character of the various strata which in this region constitute
that series of sandstones, limestones, shales and marls which together make up the
450 feet of supposedly Jurassic deposits lying between the ‘“ Red Beds” below and
the Dakota sandstones above. Dr. C. A. White in his article entitled: MHresh-
Water Invertebrates of the North American Jurassic published as Bulletin 29 of the
United States Geological Survey on pages eleven and twelve speaks as follows of
these deposits: “The character of the strata in which the fresh-water Jurassic
fossils were found, both at the Colorado and the Wyoming localities, in addition to
the character of the fossils themselves, is such as to indicate for them a lacustrine
and not an estuary or a fluviatile, origin; that is the rocks are regularly stratified
and have such an aspect and character as to indicate that they were deposited in
one or more large bodies of water. If the strata of the Colorado and of the Wyom-
60 MEMOIRS OF THE CARNEGIE MUSEUM
ing localities really contain an identical fauna, it may be regarded as at least prob-
able that they were deposited in one and the same lake. The distance between the
Colorado and the Wyoming localities indicates that the supposed lake was nearly
200 miles across; and, if the Black Hills fossils also belonged to the same contem-
poraneous fauna, the assumed lake was much larger. The existence of a fresh-water
lake of even the smaller size suggested makes it necessary to infer that there was
then in that part of the North America of to-day a continental area of considerable
size, for such a lake could hardly be other than a part of a large drainage system.
“ But aside from these considerations, the existence of such fresh-water faunas as
are represented by these collections whether in large or in small bodies of water,
indicates with hardly less clearness than the proved existence of one great lake
would do the synchronous existence of a large continental area. Indeed it seems
necessary to assume that in the fresh waters of a large land area alone, could faunas
of such a character as those which are represented by these collections be developed
and perpetuated.” I can fully agree with Dr. White as to the necessity of assuming
the existence in Jurassic times of a continental land-mass of the dimensions inti-
mated in his paper. But it does not seem to me at all necessary to presuppose the
existence of a Jurassic lake of even the smaller or more moderate dimensions
assigned by him. While I do not wish to be understood as denying the possibility
of the existence of a great lake in Jurassic times extending as Dr. White has sug-
gested from the Arkansas River in Colorado to the Black Hills of South Dakota, it
does appear to me that our present knowledge of the character of the faunas, both
terrestrial and aquatic (fresh-water) as well as of the lithogic and stratigraphic
features exhibited by the beds themselves is decidedly against such a presumption.
If I properly understand Dr. White he finds nothing in the character of the aquatic
mollusca to preclude the possibility of their having lived and developed in smaller
lakes. After a personal examination of the localities at Green River, Utah, at Grand
River in western Colorado, Canyon City and Morrison in eastern Colorado, Como
and Sheep Creek in southern Wyoming, at the Spanish Mines in eastern Wyoming,
along the Big Horn Mountains in central Wyoming, about the Black Hills in South
Dakota and in the country near Billings in southern Montana, in all of which
localities the Atlantosawrus beds are exposed and exhibit in more or less abundance,
the remains of those dinosaurs which are characteristic of them, I am convinced
that neither the character of the vertebrate fauna nor the facts of stratigraphy at
any one of these places can be taken as affording anything lke conclusive evidence
of the presence of a great body of water. At several of these localities, however, the
occurrence at intervals of sandstones showing frequent examples of cross-bedding,
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 61
ripple marks and even occasionally exhibiting footprints is conclusive proof that
such sandstones had not their origin in the midst of a great lake, while the presence
almost everywhere of the remains of terrestrial reptiles and less frequently of
mammals tells only too plainly of an adjacent land-mass. In all this region I know
of no locality where any considerable extent of the Atlantosaurus beds occurs, in
which remains of quadrupedal, terrestrial dinosaurs have not been found. To my
mind, this fact alone affords very strong presumptive evidence that in Jurassic times
Fa
Fic. 23. Photograph of footprint in Jurassic SINTON, near Canyon City, Colorado.
this entire region was the habitat of these dinosaurs, which it could not have been
had it been covered by a great lake, for the structure of their limbs shows con-
clusively that the Dinosauria were not aquatic. Nor can I conceive of the possi-
bility of the carcasses of terrestrial animals being carried out into the midst of so
great a lake as that presupposed above and left in such abundance as the numbers
of their bones in these deposits would indicate. An hypothesis, which it appears to
me is far more reasonable and more nearly in accordance with the facts as we now
know them, is to consider this region as presenting in late Jurassic and early Creta-
62 MEMOIRS OF THE CARNEGIE MUSEUM
ceous times the appearance of a low and comparatively level plain, with numerous
lakes, both large and small, connected by an interlacing system of river channels.
The whole, when covered over with luxuriant forests and broad savannas, made
possible by the supposedly tropical climate of those times, would form an ideal
habitat not alone for the large Dinosauria, but for the smaller reptiles and diminu-
tive mammals of those days and for the fishes, mollusca and other aquatic life as well.
In Figs. 25 and 24 respectively, are reproductions of photographs of a footprint
from the dinosaur beds near Canyon City, Colorado, and ripple marks from the
same deposits along the base of the Big Horn Mountains in Wyoming.
Fic. 24. Photograph of ripple marks on surface of Jurassic sandstone, Big Horn Mountains, Wyoming.
In Fig. 25 there is given the reproduction of a photograph by Dr. E. H. Barbour
of the locality near Canyon City, Colorado, where were located the quarries so long
worked by Professors Marsh and Cope. The dark area in the middle foreground
just back of the tent is the quarry so long and successfully worked by Professor
Marsh and recently worked with equal success by Mr. Utterback for the Carnegie
Museum. At A directly across the cafion and on about the same horizon was
located another quarry also worked with some success by Marsh. At B a little
above and on the same side of the small cafion, but in a slightly different horizon,
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 63
in a layer of arenaceous shales there occurs a bed of Unios from which were obtained
most of the species described by Dr. C. A. White as coming from this locality, while
the shales underlying the thick stratum of sandstone seen at the top of the escarp-
ment forming the cafon wall just to the left and in front of the tent contains
numerous small lenses of impure limestone filled with the silicified remains of
fresh-water gasteropods and the stems and seeds of small aquatic plants apparently
pertaining to some species of Chara. These limestone lenses are especially abund-
ant and quite fossiliferous at the locality marked C in the photograph and at a point
on the same horizon of this talus-covered slope a few rods in front of the extreme
foreground of the photograph and therefore not shown in the picture. The line of
trees just above and in front of the Marsh quarry marks the crest of the narrow
ridge that at this point separates the dry cafion in the middle of the picture from
Fie. 25. View of Atlantosaurus beds at entrance to Garden Park, eight miles northeast of Canyon City,
Colorado. From a photograph by Dr. E. H. Barbour.
Oil Creek on the extreme left. At this point this ridge is about 100 yards in width
from the brink of the cliff overlooking the bed of the creek and that of the dry
cafion. In the wall facing Oil Creek at the same horizon at which the bones occur
in such abundance at the adjacent quarry, dinosaur bones may be seen imbedded in
similar sandstones, showing that the bone-bearing horizon extends quite through
the ridge. From the great abundance in which the bones were found up to the
limits of the quarry as last worked and as shown in the accompanying diagrams, it
is only reasonable to suppose that many rare treasures await the explorer who has
the courage and funds necessary to remoye the 15 to 40 feet of sandstones and shales
beneath which they now le buried.
7
The isolated butte known as ‘‘Cottage Rock” seen at the head of the dry canon
in the middle background is capped with some fifty to one hundred feet of light-
64 MEMOIRS OF THE CARNEGIE MUSEUM
colored, massive sandstones generally referred to as Dakota, although neither here
nor elsewhere in this region in so far as I am aware, are these sandstones clearly
distinguishable from the Jurassic. Cottage Rock is situated about three quarters of
a mile north of the Marsh quarry and the top of the uppermost shales in this butte
which are clearly recognizable as pertaining to the Atlantosaurus beds is estimated
to be from 300 to 350 feet above the bone-bearing horizon at the Marsh quarry.
The isolated conical butte shown in Fig. 26, and locally known as the ‘“ Nipple’
is situated some 300 yards back and a little to the right of “ Cottage Rock.” It
?
Fie. 26. The ‘‘ Nipple’’ from the north, showing in the foreground the trench cut by Professor Cope
in collecting Dinosaur remains.
stands on the edge of the escarpment overlooking the valley of “Garden Park”
through which flows Oil Creek. This epee butte is composed almost entirely of
shales pertaining to the uppermost Atlantosawrus beds. It is capped with a mere
remnant of a former sandstone ledge belonging either to the top of the Atlantosawrus
beds or the base of the Dakota. About the base and over the slopes of this butte
fragmentary dinosaurian remains occur in considerable abundance and the locality
was worked to a considerable extent by Professor Cope. One of his abandoned
trenches may be seen on the left at the foot of the butte in the photograph.
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 65
Another quarry long worked by Professor Cope is shown in Fig. 27. This
quarry is situated about 500 yards West of the ‘“‘ Nipple” and the dinosaur bones,
belonging for the most part to Camarasauwrus, were found imbedded in a thick
stratum of chocolate-colored shales immediately beneath the light-colored, heavily
bedded, jointed sandstones seen at the summit in the figure and provisionally
Fic. 27, Eastern entrance to Cope quarry. Light-colored Dakota sandstone at top underlaid by
chocolate-colored shales with remains of Camarasaurus.
referred to the Dakota. Between this quarry and the ‘“‘ Nipple” there lies a com-
paratively level plain some 500 yards in width covered over with a growth of
juniper, pifon and other bushes characteristic of this region as shown in Fig. 28.
That quarry in this region which was perhaps worked with most success by Pro-
fessor Cope or men in his employ was situated about one mile north of that last
mentioned and at the same horizon, in chocolate-colored shales lying just beneath
the supposed Dakota sandstones. This last quarry I have never visited, but Mr.
Lucas, who was Professor Cope’s principal collector in this region, accompanied Mr.
Utterback to the quarry and explained to him how the bones were found. Accord-
ing to Mr. Lucas the more complete of the two skeletons of Camarasaurus supremus
which are now known to have been treated as one skeleton in Cope’s descriptions of
the species, was found at this last locality. The location of this quarry is about one
mile north of the “ Nipple” and on the edge of the escarpment facing Garden Park.
The above are the most important localities that have been worked for fossils in
this region although dinosaur remains have been found here at many other places
66 MEMOIRS OF THE CARNEGIE MUSEUM
but in no considerable abundance. It will be noticed from the above remarks
regarding the location of the several quarries worked in this region by Professors
Marsh and Cope, that the quarries operated by Marsh were in a distinctly lower
horizon than those from which Cope secured his material. While Professor Cope’s
material all came from near the summit of the Atlantosawrus beds, that of Professor
Marsh was derived from the lower members of those beds, certainly not more than
100 to 150 feet above the Red Sandstones. This difference in horizon, which can
be represented by scarcely less than 300 to 350 feet of sandstones and shales, must
of necessity represent an enormous time interval, much greater perhaps than is
ordinarily represented by sedimentary deposits of an equal thickness, for from the
Fic. 28. View from near Cope quarry with the ‘‘ Nipple’’ in the middle foreground and Cooper
Mountain in the distance. Garden Park lies in a depression about 600 feet below the ‘‘ Nipple’’ between
the crest of the bluff, indicated by the line of trees on either side of the ‘‘ Nipple,’’ and Cooper Mountain.
manner in which the sandstones and shales replace one another both laterally and
vertically, and from the frequent examples of cross-bedding and ripple-marked sur-
faces exhibited by the sandstones it is evident that the region was not one of con-
tinuous and universal deposition, but that degradation and aggradation were in
simultaneous operation and that while on the whole the latter agency predominated
there may have been and doubtless were considerable intervals during which erosive
agencies were the more efficient of the two. As should be expected the enormous
time interval which elapsed between the deposition of the sandstones of the Marsh
quarry and the shales of the Cope quarries, some 350 feet higher, was sufficient to
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 67
accomplish considerable changes in the dinosaurian fauna of this region, and these
changes are readily apparent in the faunas from these two horizons, though for
obvious reasons the present paper is not the place in which to discuss them. They
will no doubt be fully recognized and discussed by Professor H. F. Osborn in his
Monograph on the Sauropoda now in course of preparation for the United States
Geological Survey.
Synonymy of the Atlantosawrus Beds.
Although these beds were first recognized, named and adequately described both
lithologically and faunally, by Professor Marsh they have received several different
appellations by subsequent authors. Scott has called them the Como-beds ; by Cross
they were referred to as the Morrison beds; Jenney named them the Beulah Shales
and this name was used by Darton. Considering the usual similarity of the faunal
and lithologic features of these beds wherever they are known to exist and the ease
with which they may be recognized even at different and widely separated locali-
ties, it would seem somewhat unfortunate that they have received so many names.
Since Marsh’s term the Atlantosawrus beds has priority, and has become well
known through long and general usage there would seem no good reason why it
should not be retained. Even should the reptilian genus Atl/antosawrus, as contended
by some but which has yet to be demonstrated, prove to be a synonym and have to
be abandoned, this would not invalidate the name of the formation. It would be
quite as reasonable to maintain that since Fort Union on the Missouri River from
which the Fort Union beds took their name, is no longer in existence that this great
formation should receive a new name. While the present writer is entirely in favor
of basing all new formation names on geographic names taken from the localities
where such formations are first studied or are best represented, it does not appear
desirable to make this rule retrogressive. Such retrogressive application of this
rule would not only work an injustice to many pioneers in American geology, but
what is of even greater importance, it would result in augmenting still further that
confusion which already exists in our geologic formation names. Surely from that
standpoint alone there is sufficient reason for deprecating any attempt to duplicate
such names. Nor does the plea advanced by some who have been most active in
giving new names to old and well known formations, that it is easier to give a new
name than to turn bibliographer and trace out the synonymy and priority of the
names already given by others, give promise of being justified by the results which
are sure to follow such a course. ‘To the present writer it would appear much the
better plan to accept formation names for formations already known, as we find them
having due respect for priority and general usage; to adopt as a general rule for our
68 MEMOIRS OF THE CARNEGIE MUSEUM
guidance in the conferring of new names on new formations the theory that each
such name should be derrved from the name of some locality at which the formation
is well displayed and may be easily recognized and studied. It might also be well
to remember in this connection that we are no more competent to legislate for
future generations than were our forefathers.
Age of the Atlantosawrus Beds.
There has been considerable difference of opinion regarding the age of the Atlan-
tosaurus Beds. By some they have been regarded as of Lower Cretaceous age and by
others as Upper Jurassic. When first discovered, these beds were referred by Pro-
fessor Marsh to the Cretaceous (see American Journal of Science, July, 1877, pp. 87-
88). In December of this same year Professor Marsh referred these same deposits
to the Upper Jurassic and in a note describing a new fish, Ceratodus giintheri, from
these deposits, published in the January number of the American Journal of Sci-
ence for 1878 he named them the Atlantosaurus beds. He ever after consistently
maintained their Upper Jurassicage. Cope and Hayden on the other hand referred
these beds, more especially as developed at Canyon City, Colorado, and at Morrison
to the Dakota, now generally recognized as pertaining to the lowermost member of
the Upper Cretaceous. The following paragraph from page 254 of the Proceedings
of the American Philosophical Society was written by Professor Cope and it ts sig-
nificant in this connection. It is as follows: ‘‘Dr. Hayden visited the locality of
Mr. Lucas’ excavations (near Canyon City) and informs me that the formation from
which the Camarasawrus was obtained is the Dakota. Professor Marsh has at-
tempted to identify what is, according to Professor Mudge, the same horizon, one
hundred miles north of Canyon City with the Wealden of England. Specimens
from the northern locality which I have examined render it certain that the horizon
is that of Mr. Lucas’ excavations. Of this I may say that there is no paleontolog-
ical evidence of its identity with the Wealden. The resemblance of the vertebrate
fossils to those of the English Odlite is much greater, but not sufficient as yet for
identification.” Ten years later however (American Naturalist, May, 1887, pp. 446—
447) Cope placed these beds in the Jurassic to which they had been previously
referred by both Marsh and King and which, from the paragraph quoted above,
would seem to have been the only course open to him. Indeed there is little doubt
that when Cope first referred these beds to the Dakota he did so entirely upon the
determinations of Dr. Hayden and regardless of the paleontological evidences
afforded by the fauna they contained, which, such as it was, as is shown by the quo-
tation above, he regarded as pointing to a decidedly greater antiquity even than the
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS 69
Wealden, with which Marsh had been in favor of correlating them and which was
-at that time very generally regarded as of Upper Jurassic age though at present
considered by most geologists as representing the lowermost member of the Creta-
ceous.
Professor Lester F. Ward, on page 377 of Part IL. of the Twentieth Annual
Report of the United States Geological Survey, in commencing his treatise on the
Jurassic cycads dismisses the age of these beds as developed in Wyoming with the
remark that there is no doubt as to their being Jurassic, and on page 384 he says
of the cycads from the Freeze Out Hills locality that “in some respects they resem-
ble the specimens from the Purbeck beds of the Isle of Portland.”
Professor Wilber C. Knight’ has remarked as follows concerning the age of the
Atlantosaurus (Como) beds. ‘There can be no mistake in assigning the Como stage
to the Upper Jurassic, but it seems quite possible that it is more closely allied to the
Purbeckian than to the Oxfordian.”
Darton” is not very clear as to just what age he wishes to refer these beds. In
his diagram at the top of page 387 of the paper just cited he refers them to the
‘Lower Cretaceous (or Jurassic)?”’ and immediately after on the same page in his
table of the thickness of formations, and again on page 393 in describing the charac-
ter and distribution of the Atlantosawrus beds (Beulah Shales) he refers them to the
Jurassic without a query. It would seem therefore that he also favored their
Jurassic age.
Osborn has I think consistently maintained the Jurassic age of these deposits.
On the other hand Scott and Williston have been in favor of placing them in the
Lower Cretaceous.
As already noticed Dr. C. A. White has regarded these beds as of Jurassic age
though apparently relying entirely upon the evidence afforded by the vertebrates
and remarking that the fresh-water invertebrates of the same beds are so modern in
type as of themselves to offer no suggestion of a greater age than Tertiary. And
again he adds: ‘Indeed so modern is the facies . . . that one is surprised to find
only a single type among them which is not common among American living fresh-
water species.”
In discussing the age of any geological horizon which is fossiliferous two classes of
evidence are of especial importance. First in importance is its stratigraphic position
and second the nature of its included fossils, vertebrates, invertebrates and plants.
The relative value of the different classes of fossils for purposes of correlation vary
” Bull. Geol. Soc. Am., Vol. 11, p. 387.
" Bull. Geol. Soc. Am., Vol. 10, pp. 387, 398.
70 MEMOIRS OF THE CARNEGIE MUSEUM
according as the beds in question are of marine or fresh-water origin. While marine
invertebrates and most terrestrial and aquatic vertebrates are as a rule safe guides
for purposes of correlation and second only in value to direct stratigraphic evidence,
fresh-water invertebrates, plants and certain vertebrates as for instance turtles, croco-
diles and some fishes are as a rule much less reliable guides.
Stratigraphic Position of the Atlantosawrus Beds.—As originally applied the term
Atlantosaurus beds refers to that series of sandstones and shales, some 450 feet in
thickness and containing the remains of dinosaurs, small mammals, ete., lying
between the red Triassic? sandstones below and the Dakota sandstones above on
either side of the cafion of Four Mile Creek (Oil Creek) near Canyon City, Colorado.
The dinosaur remains upon which Professor Marsh relied for the determination of
the age of these deposits at this locality all came from the lowermost 150 feet of the
series and it may therefore eventually prove advisable to limit the use of the term
to the lower one third of the series. Farther north in Wyoming and about the
Black Hillsin South Dakota similar dinosaur beds are separated from the Red Beds
by a series of marine shales and limestones named by Marsh the Baptanodon beds.
These latter beds are rich in the remains of marine vertebrates and invertebrates
and are universally regarded as of Middle or Upper Jurassic age, while the over-
lying dinosaur beds have as universally been referred to the Atlantosawrus beds
usually considered, as noted above, as of Upper Jurassic age. The marine Baptan-
odon beds throughout Wyoming and South Dakota are everywhere found accom-
panying and underlying the fresh-water Atlantosawrus beds though thinning out
toward the south and entirely disappearing as we approach the Wyoming and Colo-
rado state line. As already noticed they are entirely absent in the locality near
Canyon City, Colorado, the Atlantosawrus beds there resting directly upon the Red
beds and with at least apparent conformity. Nor does there appear to be any mate-
rial break in the conditions of sedimentation in this region from the base of the
Atlantosawrus beds to the summit of the Dakota. If this be true it would appear
that at Canyon City the lower members of the Atlantosawrus beds, those worked by
Marsh and by Mr. Utterback, are the fresh-water equivalents of the marine Baptan-
odon beds farther north, while the upper beds or those worked by Cope would
become the equivalents of the Atlantosawrus beds at Morrison, Colorado, and at
various localities in Wyoming and South Dakota. Such evidences of stratigraphy
as there are prove conclusively that the Atlantosawrus beds at Canyon City overlie
the Triassic and underlie the Dakota and that they are intermediate in age between
the two and are therefore of either Jurassic or Lower Cretaceous age or that they
represent, either wholly or in part, both those horizons. The latter seems to me the
HATCHER: OSTEOLOGY OF HAPLOCANTHOSAURUS ra
more reasonable conclusion when considered from the standpoint of stratigraphy
alone.
Hvidences as to Age Afforded by the Fauna and Flora.—As already noticed Profes-
sor Ward has regarded the cyeads from the Atlantosaurus beds of the Freeze Out
Hills, Wyoming, locality as indicative of a Jurassic age.
Invertebrate paleontologists have I think been unanimous in referring the marine
Baptanodon beds to the Middle and Upper Jura. hey have it appears been most
frequently correlated with the Oxfordian or lower member of the Middle Odlite.
By some however they have been placed in the Lower Oolite. Since, as has been
shown above, there is not a little evidence in favor of considering the lowermost
150 feet of the Atlantosawrus beds at Canyon City as the equivalents of these marine
beds in the north the age of the latter, as determined by its marine invertebrates,
may be taken as having a certain bearing on that of the former series. The verte-
brates of these marine beds appear to point to a somewhat greater antiquity than
the invertebrates, for Baptanodon, the most abundant and best known form, has its
nearest ally in the Liassic Ophthalmosawrus of Europe, and Mr. C. W. Gilmore, who
is engaged in a thorough and exhaustive study of the American forms, has recently
shown that the American form was not edentulous as had been supposed and that
it is scarcely distinguishable, at least generically, from the European Liassic genus
Ophthalmosawrus.
Turning now to the fauna of the Atlantosawrus beds, it is readily apparent that
the dinosaurs offer the best, indeed almost the only reliable paleontological evidence
as to their age. We have already called attention to the fact that Cope regarded
the dinosaurs of the uppermost of these beds as being most like those of the English
Odlite and we have shown that Haplocanthosawrus from the lower half of the series
resembles most closely Cetiosawrus from the Great Oodlite near Oxford.
Marsh was wont to correlate the Atlantosawrus beds with the Wealden which he
regarded as of Upper Jurassic age. On just what evidence he relied for this corre-
lation is not quite clear. Nor does a comparison of the dinosaurian faunas of these
two horizons seem to me to warrant such correlation. While from the fragmentary
nature of much of the material upon which the different genera and species are
based it is clearly impossible to make satisfactory comparisons in many instances
between the more closely related genera and species of American and European
dinosaurs, nevertheless when comparisons of the faunas as a whole are instituted
between the various American and European horizons most striking and important
resemblances and dissimilarities are at once apparent. Thus while in the Aflanto-
saurus beds the Sauropoda are the predominant forms both as regards size and the
72 MEMOIRS OF THE CARNEGIE MUSEUM
number of genera, species and individuals in the Wealden they are almost entirely
replaced by the Predentata and 'Theropoda. And the Iguanodontia so abundant in
the latter formation are quite unknown in the former. The same dissimilarity
though in a less striking degree is noticeable when the fauna of the Purbeck is com-
pared with that of the Atlantosawrus beds, and it is not until we get down into the
middle of the Odlite that we find a dinosaurian fuana comparable even with that of
the upper and middle Atlantosawrus beds.
In consideration of the evidences mentioned above it appears to the present
writer that the dinosaurian fauna of the Atlantosauwrus beds, as we now know it is
unmistakably Jurassic in type, but that these beds may in their uppermost mem-
bers represent a portion at least of the lower Cretaceous.
CarNEGIE Museum, April 15, 1903.
ADDITIONAL REMARKS ON DIPLODOCUS.
By J. B. HarcuHer.
Since publishing my memoir! on the osteology of Diplodocus additional discoy-
eries have thrown more light on the structure of these strange reptiles. It thus
becomes necessary to make certain alterations in the description and restoration then
given, especially relating to the structure of the fore limbs and feet. As stated in
the text of my memoir there were at that time in the collections of this museum
no representatives of the fore limbs or feet of Diplodocus and the brief descriptions
of those elements there given was based entirely upon the published descriptions by
Professor Osborn and upon photographs of the limbs kindly loaned by him. For-
tunate discoveries of the fore limbs and feet of Brontosawrus (No. 563) by Mr. C.
W. Gilmore and of the greater portion of a skeleton of Diplodocus (No. 662) by
Mr. W. H. Utterback have demonstrated two important errors in my previous paper.
These are :
First—The radial articulation at the distal end of the humerus is on the in-
ternal side and anterior to the internal portion of the ulnar articulation instead of
being external and anterior to the latter as stated in my memoir. When in position
the proximal end of the ulna entirely enclosed that of the radius posteriorly and
' Memoirs Carnegie Museum, Vol. I., No. 1, pp. 1-68, Pl. 1-13.
»
HATCHER: ADDITIONAL REMARKS ON DIPLODOCUS 73
externally, as shown in Fig. 1, and its articular surface is opposed to that of the
distal end of the humerus posteriorly throughout its entire breadth and presents a
broad and deep anterior projection enclosing the radius externally and articulating
with the anterior and external surface of the distal end of the humerus.
Fic. 2.
Fig. 1. a, front view of radius and ulna of Diplodocus (No. 662). 6, proximal end of same. Both
figures are one tenth natural size and show bones as seen when in position.
Fic. 2. a, front view of supposed clavicle of Diplodocus. 6, internal view of same (No. 662), one
tenth natural size.
The contact of the radius with the humerus is thus limited to the antero-internal
surface instead of the antero-external as erroneously shown in my original restora-
tion of the skeleton. The radius and ulna do not cross each other so completely as
supposed by Osborn and Granger, but occupied the position relative to one another
shown in Fig. 1.
Second.— The structure of the manus was entaxonic instead of mesaxonic as
erroneously represented in my original restoration where, as stated in the text, I
followed Osborn, having at that time no material upon which to base a restoration
of those elements. The manus was doubtless somewhat more plantigrade than I at
that time represented it. In the present restoration these errors in the structure of
the fore limbs and manus have been corrected. The principal characters of the
74 MEMOIRS OF THE CARNEGIE MUSEUM
latter are taken from the manus of Brontosawrus, a detailed account of the structure
of which was recently published by the writer.’
The Supposed Clavicles. —In my original description of Diplodocus carnegii I
figured and described a peculiar bone which I then considered as a clavicle, though
at the same time expressing some doubt as to its real nature. Fortunately we have
found associated with another skeleton (No. 662) of Diplodocus a second and more
complete clavicle? shown here in Fig. 2, a, b. The present specimen is somewhat
incomplete at the bifid extremity, the smaller branch having been broken away, the
opposite end is complete, somewhat expanded and spatulate as shown in the
figures. The spatulate portion has a length of 265 millimeters, a maximum breadth
of 75 millimeters and an average thickness of about 24 millimeters.
The entire length of the bone measuring along the are of that portion of the
circle which it describes is 620 millimeters. Between the expanded portion and the
forked extremity the bone is irregularly elliptical or subcireular in cross-section.
This bone is asymmetrical and is to all appearances a paired bone. In neither
instance have we as yet secured its opposite, though this is still possible with that
one now being considered, a considerable portion of the skeleton still remaining to
be unearthed. Just at the point where the rounded shaft passes into the flattened,
spatulate extremity there is on one side a shallow groove running obliquely across
the surface of the bone. This groove has the appearance of having been formed by
the overlapping edge of a coracoid or sternal. The flattened spatulate extremity pre-
sents aslightlyrugose, fibrous surface as though it had been imbedded in cartilagin-
ous or muscular tissue, and this together with the bifid nature of the other extremity
has suggested the possibility that the bone might be an os penis; in which case the
bifid extremity would be the distal end and the flattened the proximal extremity.
Against the probability of this assumption however, the marked asymmetry of the
bone offers a potent argument and I am still strongly inclined to consider it a clavicle
as which it might very readily have functioned. Although clavicles have not here-
tofore been recognized in the Dinosauria there would seem no good reason for sup-
posing that they were not present in some members of that group. A clavicle of
the size and form of the element under discussion, if attached to the anterior edge
of the broadly expanded sternals, coracoid and prescapula, could not have failed in
giving additional strength and rigidity to this portion of the skeleton.
The Anterior Cervicals. — In my former paper, owing to the incomplete nature of
ceryicals 3, 4, 5, they were figured as without cervical ribs; later discoveries (No. 662)
demonstrate that ribs were present on all these vertebree and they are so shown in
the accompanying restoration (Plate F).
*See Science, N. S., Vol. XIV., pp. 1015-1047; and Annals Carnegie Museum, Vol. I., pp. 356-376.
HATCHER: ADDITIONAL REMARKS ON DIPLODOCUS TD
The cervical vertebra figured by Marsh and reproduced as text Fig. 24 in my
memoir on Diplodocus, although referred by Marsh to Diplodocus longus, is now .
known to have pertained to a species of Brontosawrus instead, and hence is of no
value in distinguishing the different species of Diplodocus as I then supposed.
Puate I.
Puate II.
Puate III.
Puate LY.
Puate V.
Puiate VI.
Presacral vertebra of type (No. 572) of Haplocanthosaurus priscus, one tenth nat-
ural size. Series 1, as seen from right side; Series 2,as seen from in front;
Series 3,as seen from behind. (C14 and (15, cervicals 14 and 15; 1, first dor-
sal ; 6-14, dorsals six to fourteen respectively ; pz/, postzygapophysial lamina; oJ,
oblique lamina; //, horizontal lamina; dl, diapophysial lamina; az/, prezygapo- .
physial lamina; S, modified surface for muscular attachment of scapula ; t or ¢f,
tubercular rib facet; ¢ or cf, capitular rib facet, al, inferior blade of diapophy-
sial lamina in first dorsal and prespinal lamina in sixth dorsal.
Vertebrae of type (No. 879) of Haplocanthosaurus utterbacki, one tenth natural
size. Series 1 and 2, dorsals; 3 and 4, cervicals; 5, anterior caudals.
Nineteen anterior caudal vertebra of type (No. 572) of Haplocanthosaurus priscus,
one tenth natural size. Series 1, seen from right side; 2, seen from in front; 3,
from behind.
1. Pelvis of Brontosaurus excelsus (No. 563); 2. Pelvis of Diplodocus carnegii
(No. 94); 3. Pelvis of Haplocanthosaurus priscus (No. 572). All one tenth
natural size and seen from left side. i/., ilium; p. p., pubic peduncle; 7. p., ischial
peduncle; g.c., acetabulum; pb., pubis; is., ischium; a., anterior extremity ; p.,
posterior extremity; 1, 2, 3, 4, 5, spines of first, second, third, fourth and fifth
saerals.
1. Inferior view of sacrum of Haplocanthosaurus priscus with ilia attached (No.
572). a, anterior end; p, posterior; pp, pubic peduncle; is, ischiac peduncle ;
pfs foramen between ilium and parapophyses of first sacral.
2. Anterior view of pelvis of same with ischia detached. pp, pubic peduncle; p,
pubis; ps, pubic symphysis; pt, pubic foramen.
3. Posterior view of same, with pubis detached and anterior expanse of ilia not
shown. ip, ischiac peduncle: is, ischium. All one tenth natural size.
Restoration of Diplodocus carnegiti Hatcher.
From material in the collections of the Carnegie Museum, one thirtieth natural size.
Ree
LP al bg fee
PieAeane
y in
AWN
Yn)
y, |
i
’. SIZE. (No. 572.)
MEMOIRS CARNEGIE MUSEUM, VOL, II PLATE I.
Sxpxry PRESTicE DEL
PresacraL VertTEBR» oF Type or HAPLACANTHOSAURUS PRISCUS. 1, Sipe view; 2, ANTERIOR VIEW; 3, PosteRIOR VIEW. yy NAT. s1zE, (No, 572.)
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE II.
SS
Oo
J \
Gp
YE =
SYDNEY PRENTICE DEL,
VERTEBR® OF Typr or HAPLACANTHOSAURUS UTTERBACKII. 1, 2, Dorsats; 3, 4, CERvICALS; 5,
CaupaAts. =; Nar. size. (No. 879.)
& i =
‘ i mn
" a
- Si - J
7 2 5
[ : Res ; : =
ae : ic
7 aad SS = .
n ty Sy . i = =
7 1. S
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-
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o> = y
er. tr
j 2
er,
r F ‘ Veet o 5
If . a Xe
. a? Lae - A at
as ism 4 oa -
1 Cee 4
i ‘ 4 =
F ' i i
a4 ‘ ~ _ > =
DF \ ‘ ‘
: fy
oa * ny § ‘ A >
~ : = Sh
ca it ie : :
4 r
MEMOIRS CARNEGIE MUSEUM, VOL. II.
SYDNEY PRENTICE DEL.
NINETEEN ANTERIOR CAUDALS OF TYPE oF HAPLACANTHOSAURUS PRISCUS. 1,
PLATE III.
2, ANTERIOR VIEW; 3, PosTeRIoR view. =. Nav. size. (No. 572.)
La
S|
4a
Des tag
ta J
sey §
MEMoIRS CARNEGIE MUSEUM. VoL. II.
PLATE III.
JNETEEN ANTERIOR CAUDAL F TYPE OF P, T US P. C.
N AUDALS 0 yPE OF HAPLACANTHOSAURUS PRISCUS. 2, ANTERIOR VIEW; 3, PosTERIOR VIEW. jy NAT. SIZE No. 572.
’
ed i0 0 4 ( )
Pe ie ye See
& GRRMEGTE Srrawirtr, VOL sf.
EO Ee
: 4 Hse petite A
MEMOIRS CARNEGIE MUSEUM, VOL. II.
SSea
SS
SS
S
S
SSS
~
>=
=
SSS
=
} t S
>
=
yy), Fr
4 @
Re
[PLACANTHOSAURUS. ALL SEEN FROM LEFT SIDE. ;), NAT. SIZE.
MEMOIRS CARNEGIE MUSEUM, VOL. II
Sypyer Paesticr xi
1, Petyis op BRONTOSAUBES: 2, Pevis or DreLonoous: 8, Peuvis oF HIPLACANTHOSAURUS, AUL SEEN FROM LEFT SIDE, y'y NAT. sizk,
PLATE IV
ei ai We
| bisa AE ORAL ©
wir
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE V.
SY PS ~
\ h NS
HUE I r
lt \
\\9
i
S SS \
WAY Ny Ss —————
\ y Zz
‘\ UME =
Aig =
EE \ Mf WS Ow
SW | As =
g S y(t? = ul
GE SS Y | Wl ——=
a ~
/\\
GY oa
Ws
SYDNEY PRENTICE DEL.
Petvis or Typr or HAPLACANTHOSAURUS PRISCUS. 1, INFERIOR VIEW; 2, ANTERIOR VIEW; 3, PosTERIOR VIEW.
Aun 75 Nar. size. (No. 572.)
ME)
PLATE VI
]
MEMOIRS CARNEGIE Museum, VoL. II.
Restoration or DIPLODOCUS CARNEGII. 5 Nat. size.
PLATE VI
“4
Publications of the Carnegie Museum. Serial Nos. 30, 31, 32, and 35.
MEMOIRS
CARNEGIE MUSEUM.
VOL. II. Nos. 2, 5, 4 AND 5.
W. J. HOLLAND, Epriror.
2. OSTEOLOGY OF BAPTANODON MARSH.
By ©. W. GILMORE.
3. FOSSIL AVIAN REMAINS FROM ARMISSAN.
By C. R. EASTMAN.
4, DESCRIPTION OF NEW RODENTS AND DISCUSSION OF THE
ORIGIN OF DAMONELIX.
By O, A. PETERSON.
5. THE TERTIARY OF MONTANA, Parts I, IL.
By EARL DOUGLASS
PITTSBURGH.
PUBLISHED BY THE AUTHORITY OF THE BOARD OF TRUSTEES OF THE
CARNEGIE INSTITUTE.
Marcu, 1905.
MEMOLRS
OF THE
CARNEGIE MUSEUM.
\/ Olb, Ie INO), 7
OSTEOLOGY OF BAPTANODON (MARSH).
By C. W. Gru~More.
INTRODUCTION.
The discovery of the existence in the Jurassic of North America of a representa-
tive of the Ichthyosaurian reptiles was first announced by Professor O. C. Marsh in
1879." Although fossil remains of these animals are very abundant in Europe
where they have attracted the attention of naturalists for the past two centuries, with
one exception’ it was not until this late date that they were reported from this coun-
try. Notwithstanding that continued explorations in the Jurassic have brought to
light many representatives of this group, with the exception of a few short papers
nothing has been published concerning the skeletal features of these interesting
reptiles.
The skeletons are usually imbedded in exceedingly hard and refractory concre-
tions and the bone is so thoroughly impregnated with the matrix, which it closely
resembles in color and texture, that the work of preparation is exceedingly tedious.
The difficulty in preparing the material for study may in a measure account for the
apparent neglect of this group.
The Jurassic [chthyosaurian material preserved in the vertebrate collections of
this museum has been largely gathered by the different field parties which have
been operating for the past five seasons in the fossil deposits of the west.
Through the courtesy of Mr. J. B. Hatcher, late Curator of the Department of
Vertebrate Paleontology of the Carnegie Museum, this material has been placed at
the disposal of the writer for study and description.
1 Marsh, O. C., ‘‘ A New Order of Extinct Reptiles (Sauranodonta) from the Jurassic of North America,’’ Amer.
Jour. Sci. (3), Vol. XVII., pp. 85-86, January, 1879.
2 Leidy, Joseph, *‘ Notice of Some Reptilian Remains from Nevada,’’ Proc. Acad. Nat. Sci. Phila., Vol. XX., pp.
177, 178, 1868. A Triassic form.
77
78 MEMOIRS OF THE CARNEGIE MUSEUM
The present paper is based upon the remains of three individuals, Nos.* 603, 878
and 919, from as many widely separated localities. All have been entirely freed
from the matrix or worked out in relief by the writer and because of their remark-
ably well preserved condition several important osteological characters are shown
for the first time.
In studying this material it was found necessary to compare it with the types of
Baptanodon, preserved in the Yale Museum collections. Through the kindness of
the late Dr. C. E. Beecher the author was permitted to study not only the types
(B. natans and B. discus), but all of the Ichthyosaurian remains contained in the
collections of that museum, comprising parts of some nine or ten individuals. At
the American Museum of Natural History through the courtesy of Dr. W. D.
Matthew I was given the privilege of examining the quite complete Jurassic Ichthyo-
saurian skeleton in that collection. Through my former instructor, the late Dr. W.
C. Knight, I was accorded the privilege of studying the material at the Wyoming
University, Laramie, Wyoming, part of which I helped collect. Thus practically all
of the important Ichthyosaurian remains known from the Jurassic of North America
have been personally examined during the preparation of this paper.
My acknowledgments are especially due the late Mr. J. B. Hatcher for the
encouragement and kind consideration shown me in all matters pertaining to the
preparation of this study.
For the preparation of the drawings used the author is grateful to the skill and
patience of Mr. Sidney Prentice.
The photographs are by Mr. A. 8. Coggeshall.
OccURRENCE AND DiIstrIBUTION.
The Ichthyosaurian remains collected from the Jurassic of America have all
been found in the upper part of the marine division of that formation. ‘This marine
strata was designated by Marsh as the Baptanodon Beds,‘ and included all of the
Jurassic lying between the freshwater or Atlantosawrus Beds above and the red beds
or Triassic (?) below. The Hallopus Beds of Marsh, if at all recognizable, have never
been found in contact with the marine beds and their exact position in relation to
these beds is yet undetermined.
Knight® has proposed the name Shirley for the marine series without assigning
any place or giving any reasons for supplanting the old and generally accepted
term Baptanodon Beds, which has priority.
5 The numbers refer to the Card Catalogue of Fossil Vertebrates in the Collection of the Carnegie Museum.
‘Marsh, O. C., “‘ The Reptilia of the Baptanodon Beds,’’ Amer. Jour. of Sci. (3), Vol. 50, pp. 405-406, 1895.
5Knight, W. C., Jurassic Rocks of Southeastern Wyoming,’’ Bull. of the Geol. Soc. of America, Vol. XI., pp. 377-
388, May, 1900.
: GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 79
Dr. F. B. Loomis’ in a later paper proposes to confine the term Baptanodon
Beds to the single layer in which the remains of these reptiles are found. Hatcher’
has justly contended that the duplication by giving new names to old and well
known formations, “tends to augment still further the confusion which already
exists in our geologic formation names,’ and this would appear to be another
instance in which the old and generally accepted term should be retained in its
original meaning.
These beds have been briefly described by Knight as follows : 8 a Composed of
bands of shale, limestone, sandstone, and clay. The limestones are usually shaly.
The limestone beds are quite thin, but usually fossiliferous. The clays and shales
usually contain large concretions which contain both vertebrate and invertebrate
fossils. Septaria are common. ‘The invertebrate as well as the vertebrate faunas
are only partly known. ‘This has been largely due to the fact that the richest fossil
localities are where the concretions are well developed and until recently not
many of these were known.”
The following is a fairly complete list of the fossils described from the marine
Jurassic of this country.
INVERTEBRATES.
Astarta packardi, White. ~ Modiola sp.
Belemmnites densus, M.,& H. ; Ostrea engelmanm, Meek.
Cardioceras cardiformis, M. & H. Pentacrinus asteriscus, M. & H.
Cardioceras cardiformis, var. distans, Whitf. Pholodomya kingi, Meek.
Comptonectes bellistriata, Meek. Pleuromya subcompressa, Meek.
Comptonectes extenuata, M. & H. Pinna sp.
Gryphea nebrascensis, M. & H. Pseudomonotis curta, Hall.
Dentalium subquadratus, Meek. Pseudomonotis orbiculata, Whitfield.
Goniomya montanaensis, Meek. Tancredia cf. extensa, White.
Grammatodon inornatus, M. & H. Tancredia ef. inornata (M. & H.), Whitf.
Tima sp. Tancredia warrenana, M. & H.
Lingula brevirostris, M. & H. Thracia weedi, Stanton.
VERTEBRATES.
Baptanodon natans, Marsh. Megalneusaurus rex, Knight.
Baptanodon marshi, Knight. Plesiosaurus shirleyensis, Knight.
Baptanodon discus, Marsh. Pantosawrus striatus, Marsh.
Cimoliosaurus laramiensis, Knight.
5 Loomis, F. B., ‘‘On Jurassic Stratigraphy in Southeastern Wyoming,’’ Bull. of Am. Museum of Nat. History,
Vol. XIV., article XII., pp. 189-197, June, 1901.
THatcher, J. B., Memoirs of Carnegie Museum, Vol. I1., No. 1, November, 1903, p. 67.
8 Tit. cit., p. 385.
o)
(S
MEMOIRS OF THE CARNEGIE MUSEUM
Two fishes have been referred to by Knight as probably coming from these beds.
Amiopsis dartom, Eastman. Pholidophorus americanus, Eastman.
Hatcher ® gives the geographical extent of these beds as follows: ‘The marine
Baptanodon Beds throughout Wyoming and South Dakota are everywhere found
accompanying and underlying the freshwater Atlantosawrus Beds though thinning
out toward the south and entirely disappearing as we approach the Wyoming and
Colorado state line.” He has also shown that the lowermost 150 feet of the Jura
at Cafion City, Colorado, may be the freshwater equivalents of the marine Bap-
tanodon Beds farther north.
By far the largest number of skeletons of Baptanodon now known have been col-
lected from the exposures of southeastern Wyoming, though several individuals
have been discovered in the strata farther north. There is one specimen (No. 919)
in the collections of this museum from the marine beds of north central Wyoming.
Their remains are also reported from South Dakota.
Of the dozen or more specimens I have collected or helped collect every one was
found more or less enclosed in one of those concretionary masses spoken of by
Knight, the form and size of these concretions being dependent upon the shape
and position of the skeleton enclosed. When exposed to the atmosphere, as often
happens by the carrying away of the surrounding shale or clay, the concretion in-
variably cracks into an innumerable number of pieces.
It has been my experience that the anterior portion of the snout, the end of the
tail and tips of the extremities are not enclosed by the concretion, and when present
are found in the soil surrounding the rock. This will account for the poorly pre-
served condition of these parts in the material under discussion. Of the six skulls
and parts of skulls examined the tip of the beak is wanting in every instance, and
peculiar as it may seem, the anterior fourth of each protruded from the concretion
into the surrounding shale. The quite complete posterior caudal series preserved
in the collection of the University of Wyoming came from the clay. Four paddles
have been studied and although the bones of the proximal segments are retained
in their relative positions in the matrix, the distal part is wanting in every instance.
In one example, the type of B. discus (No. 1955)," quite a number of the smaller or
distal disks were preserved but their color and state of preservation indicated that
they had come from the soft clay surrounding the concretion which contained-the
skeleton proper.
Lit. cit., p. 70.
0 Catalogue number of the Yale Myseum.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 81
So far as I am able to learn there has never been more than a single individual
found in any one of these concretions.
Plate VII. shows the outline of the concretion with the bones of specimen No.
878 in their relative positions as they lay imbedded in the matrix. The skull was
lying on the right side and turned at a right angle to the vertebral column.
The snout projected from the concretion as explained above. The cervicals im-
mediately following the atlas and axis were crushed into the left orbit so completely
that it was deemed inadvisable to attempt their removal. In the course of prepara-
tion the skull has been separated from the block containing the other parts of the
skeleton. ‘Though slightly displaced the anterior 15 vertebre are well preserved.
The remaining six of the series are only represented by fragments and these are of
little value for purposes of study.
The coracoids and scapulee, as will be seen, were in their natural positions and
give an accurate idea of the manner in which they articulated. The clavicles and
interclavicle were somewhat removed from the remainder of the girdle, though the
relation of these bones is very well shown. There are many parts of ribs but none
are complete. ‘The proximal end of the right humerus is the only limb bone repre-
sented and this end was only slightly removed from the glenoid socket.
The bones shown in this diagram will be referred to later in the detailed descrip-
tion of the several parts.
This specimen (No. 878) was collected by Dr. J. L. Wortman and party during
the summer of 1899 on Troublesome Creek, Carbon Co., Wyo. It is the most com-
plete individual of the three considered in this paper, and is unique as being the
first to have the clavicles and interclavicle preserved. The parts of this skeleton
preserved consist of a good skull and lower jaws, with a series of 21 vertebree imme-
diately posterior to the skull, the complete pectoral girdle, with the proximal ex-
tremity of the right humerus, numerous parts of ribs and isolated paddle bones.
The second individual (No. 603)" is from the Baptanodon Beds of Sheep Creek,
Albany Co., Wyo. It was collected by Mr. O. A. Peterson and party during the
season of 1900. The parts recovered consist of a nearly complete skull and lower
jaws, numerous yertebree and pieces of ribs, with a few paddle bones.
(No. 919) is from the Red Fork of Powder River, Big Horn Co., Wyo., and was
collected by Mr. W. H. Utterback in 1902. The incomplete pectoral girdle and
numerous vertebrae are represented. ‘The dorsal and anterior views of the girdle are
shown in Pl. XII.
1 Hatcher, J. B., ‘‘ The Carnegie Museum Paleontological Expeditions of 1900,’’ Science, N. S., Vol. XII., No.
306, pp. 718-720, Noy., 1900.
82 MEMOIRS OF THE CARNEGIE MUSEUM
Considering our limited knowledge of the structure of the Jurassic Ichthyosau-
rians, I shall figure and describe this material in detail.
The following description of the several elements is based largely on No. 878
supplemented by such characters as are shown by Nos. 603 and 919. Unless other-
wise stated the material should be considered as pertaining to No. 878.
Tens Sy Owen, R., lit. cit.
“7 Seeley, H. G. Quart. Jour. Geol. Soc., of London, Vol. XXXVI.
90 MEMOIRS OF THE CARNEGIE MUSEUM
The matrix in this region has not been sufficiently removed to show fully
the inward extension of these plates. The surface of each plate is covered with
lines that radiate from the center of the plate inward and outward to either
lateral border.
The ring is nearly circular in form and almost fills the large orbit. The pupil-
lary opening measures about 100 mm. in diameter, the entire diameter of the ring
being 202 mm. The left orbit of No. 603 has the ring nearly as well preserved,
though the exact number of plates cannot be ascertained. The outer surface is
inclined somewhat from the center to the outer margin and does not present so flat a
surface as observed in a specimen (No. 877) of /. communis in the collection of this
museum. The sclerotic rings do not stand parallel to one another but are inclined
inward anteriorly.
Jugal (j.). — The jugal is a long curved bar that forms the lower boundary of the
orbit. This element in both of the specimens under consideration is unusually
strong. Anteriorly the end commences in a tapering point which is wedged in _be-
tween the lachrymal and maxillary. Medially it widens into a subcompressed bar
reaching its maximum width under the middle of the orbit. At this point supe-
riorly the surface is slightly concave but becomes flattened and compressed poste-
riorly terminating in a wedge-like end which is received between the postorbital and
quadratojugal.
Parietals (pa.). —In both skulls the median parietal sutures are entirely obliter-
ated. The lateral surfaces which form the outer walls of the brain case are smooth,
slightly concave antero-posteriorly and gently convex from the crest downward.
The matrix has not been sufficiently removed from the temporal fossa to show the
depth of the parietals or their relations to the lower bones of the brain case. The
brain case is narrowest just posterior to the middle region but expands both poste-
riorly and anteriorly. The parietal foramen or pineal eye appears to be bounded
posteriorly by the parietals and anteriorly by the frontals. Extending forward and
outward from the lateral boundaries of the foramen are the sutures (?) that unite the
parietals with the two small frontals. From the antero-internal angle of the tem-
poral fossa the suture between the postfrontal (?) and parietal extends inward and
forming an acute angle with the fronto-parietal suture.
The sagittal crest is indicated by a faint ridge posteriorly. The lower posterior
border of the parietal rests upon the underlying supraoccipital without sutural junc-
tion. ‘The posterior lateral extensions curve downward and backward descending
to the top of the exoccipitals, thus contributing to the upper and lateral faces of the
occipital surfaces. Seen posteriorly the median portion is slightly thickened, the
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 91
lateral extremities narrowing somewhat as they descend to oceupy the space between
the internal branch of the squamosal and the supraoccipital.
Frontals (fr.).— The frontals are, so far one may rely on the very unsatisfactory
evidence at hand, very small and in a general way resemble those elements observed
in the skulls of the Ichthyosawria generally. The sutures of this region are quite
indistinct being indicated by hardly more than fractures. These fractures however
having been produced by pressure would naturally have followed the line of least
resistance, which in this case is that of the sutural union of the different elements.
As these fractures have the position and direction of the sutures shown in the draw-
ings of Ichthyosaurian skulls I haye had them drawn in provisionally. See Plates
VIII. and IX. Because of the conjectural nature of the above evidence no attempt
will be made to describe this region in detail. This portion of the skull of No. 603
is so badly crushed that all sutures have been entirely obliterated.
Postfrontal (ptf.).— The posterior border of this bone is well shown, but anteriorly
as explained above the sutures are only indicated by transverse fractures. The
post- and prefrontals will be considered here as a single element extending from the
supratemporal behind to the rounded fracture forward where it unites with the elon-
gated nasals. Although in Ichthyosaurus Frass shows a suture immediately over
the eye separating the pre- from the postfrontal, there are no indications of such
a suture in our specimens. Reynolds* says “The exact position of the suture be-
tween the prefrontal and postfrontal is not known.” ‘This element as here considered
isa moderately long compressed bone forming the quite horizontal postorbital arch.
Posteriorly the bone is obliquely inclined so that the outer surface looks outward
and upward. Fore and aft the element is convex. The nearly square posterior
end unites with the supratemporal, the postero-superior margin with the anterior
branch of the squamosal, the latter being prolonged on the inner surface of the
postfrontal which is united by an extended suture. The outer or inferior border at
this end joins the upper border of the postorbital for about 63 mm., but more an-
teriorly this bone alone forms the upper boundary of the orbit, at this point assum-
ing a nearly horizontal position. Anteriorly this element acquires its greatest width
as it curves in front of the temporal fossa to meet the parietal and frontal. Though
how much of this wide portion if any is prefrontal we cannot determine at this
time. ‘The superior surface in front of the temporal opening is elevated into a
rounded transverse ridge which becomes narrower and somewhat depressed as it
approaches the junction with the parietal. Anterior to this ridge the surface is
slightly convex transversely and bends downward longitudinally to meet the
nasals.
28 Reynolds, ‘‘The Vertebrate Skeleton,’’ p. 196.
92 MEMOIRS OF THE CARNEGIE MUSEUM
Prefrontal ( prf.). —The shape of the prefrontal, or even the probable outlines, can-
not be determined from the present specimens. On the right side of the skull per-
taining to specimen No. 878 is a narrow portion forming the upper and forward
boundary of the orbit extending down to meet the lachrymal in front. The narrow
portion just cited is undoubtedly a part of the prefrontal as this is the position of
this element in all fully figured Ichthyosaurian skulls. Although Baur” says, ‘‘ The
lachrymal is free from the prefontal in Ichthyosawrus, as in many Lacertilians ; it is
united with this bone in Sphenodon.”
Nasals (na.).— The outline of these bones when viewed from above is triangular.
They are elongate antero-posteriorly and form an acute angle where the two ele-
ments pass beneath the overlapping premaxillee as shown by Owen” in I. camplyodon.
The anterior sutural boundaries of these bones are best shown in the skull pertain-
ing to specimen No. 603. Posteriorly and laterally the nasals are bent abruptly
downward and form the outer surface of the skull in front of the orbit and above
the nares. Anteriorly the outer surface is inclined less obliquely, gradually becom-
ing convex as the nasals extend forward. Above, the nasals are widest at their pos-
terior ends and in so far as one may judge from the damaged condition of this region
they were somewhat concave transversely. Posteriorly the nasals united probably
with the frontals and the prefrontals(?). The suture separating the nasal from the
premaxillary begins at the upper anterior angle of the nasal opening and extends
obliquely upward, forward and inward meeting its fellow medially at a point about
one third the distance from the anterior to the posterior extremity of the skull.
There is a small angular protuberance over the middle of the nares which I have
not observed in other Ichthyosaurian skulls. The suture separating the nasals is
plainly shown anteriorly in specimen No. 603. In the accompanying figure Pls.
VIII. and IX., the anterior sutures have been drawn from that specimen.
Lachrymal (la.}.—The lachrymal is a large subtriangular bone that joins the
prefrontal (?) above and the jugal below to complete the anterior boundary of the
orbit. The posterior border is concave. A spine-like prolongation extends back-
ward beneath the eye and unites by suture with the anterior third of the jugal, this
border more anteriorly meets the maxillary, terminating at the lower anterior border
of the nasal opening. ‘The superior process presumably, meets the prefrontal and
nasals above. he latter probably as in I. (zelandicus) quadriscissus sending down
a small projection posterior to the narine opening to meet it. Anteriorly the oblique -
superior border forms the lower boundary of the external nares. The facial surface
29 Baur, G., lit. cit., p. 838.
“Owen, R., ‘‘ British Cretaceous Reptiles,’’? Monograph, 1851.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 93
posteriorly is somewhat concave and looks outward and backward ; anteriorly it is
somewhat rounded, sloping in toward the nasal opening. The inferior side is nearly
straight though elevated obliquely from back to front.
Maxillary (ma.). — The anterior half of the left maxillary of specimen (No. 603)
was firmly fastened to a section of the left ramus by the intervening matrix and
being separated from its position in the skull we have a splendid view of the form
and structure of the anterior half of this bone. Laterally this element is subtri-
angular in outline. It commences posteriorly under the anterior border of the eye,
from which it is separated by the anterior portion of the jugal and the posterior
branch of the lachrymal. Anteriorly it expands into facial, alveolar and palatal
aspects, again tapering out to a slender pointed anterior end. The complete outer
anterior view is hidden by the enclosing premaxillary. The median superior
margin contributes a part to the lower boundary of the nasal opening. On the
median superior surface a long slender horizontal process is developed which fits
into a corresponding depression in the premaxillary. The matrix has not been
removed sufficiently from the inside to determine whether it articulated with other
bones beside the palatines and premaxillaries in that region. Nothing is known of
the upper part of the posterior extremity of the maxillary. Anteriorly the superior
or articulating surface is rounded, being covered with longitudinal markings.
The inferior surface anteriorly is concave and forms the deepest portion of the
alveolar groove.
Premaxillary (pma.).— The premaxille are characterized as in nearly all Ichthyo-
saurians by their great length. In B. discus the posterior termination is forked as
in many forms of this order. The lower posterior portion enclosed the anterior
portions of the maxillary and hides it from a lateral view. From the anterior
boundary of the nares the suture between the nasal and premaxillary extends for-
ward and upward to where the opposite branches become confluent and cover the
anterior extremities of the nasals. The anterior part of the snout narrows gradually
to the end and is composed of the premaxille alone. The tip as has been already
explained is wanting in all of our specimens. The sides are slightly flattened but
above are convexly rounded. The median suture is obliterated in both specimens
although the slipping of the two halves past one another in the skull belonging to
No. 878, see Pl. X., fig. 1, indicates that such a suture existed. The anterior half of
the premaxillary has a longitudinal channel along the side just above the alveolar
grooye, containing pits for the entrance of nerves to the teeth. Descending to the
alveolar surface it will be observed that this element forms the greater portion of
the upper jaw. ‘The inner alveolar plate of the premaxillary forms the main part
94 MEMOIRS OF THE CARNEGIE MUSEUM
of the arched roof of the upper dental groove. Medially the palatal portion is
developed as a long narrow vertical plate that extends downward meeting the same
plate of the opposite side mesially for a vertical distance of 12 mm. These narrow
alveolar plates posteriorly are separated by the intervention of the anterior extremi-
ties of the vomers and pterygoids, which eventually disappear under the premaxille.
Basisphenoid (b.s.). — In so far as I can ascertain, owing to the damaged condi-
tion of this region, the basisphenoid is a heavy subquadrate bone. Posteriorly it
articulates with the basioccipital by a deep vertical suture; laterally it is over-
lapped by the interior processes of the pterygoids; anteriorly the long slender pre-
sphenoid is given off. The suture between these bones is not shown nor have I
observed it in any of the skulls of Ichthyosawrus that are figured. The length of the
basisphenoid in specimen No. 603 is about 70 mm.
Presphenoid (prs.). —The presphenoid appears as a median anterior prolongation
of the basisphenoid. This long trihedral bar gradually tapers to a point as it
extends forward. The anterior extremity in No. 603 disappears under the dis-
placed pterygoids. This bone divides the long interpterygoid vacuity into two
parts posteriorly.
Pterygoid (pt.).—The pterygoid is an irregular elongate bone the most con-
spicuous element of the palatal region. The internal border posteriorly is concave
antero-posteriorly and forms the lateral boundary of the interpterygoid vacuity.
Posteriorly the inner margin is beveled off to a thin edge which laps over the lateral
inferior surfaces of the basisphenoid, and the truncated posterior interior angle prob-
ably reached and overlapped the anterior portion of the basioccipital. The posterior
end curves slightly upward where the superior surface meets the inferior lateral
border of the stapes. Seen from below the posterior border is slightly emarginate,
the lateral border posteriorly being deeply excavated. Between this excavation
and posterior emargination the pterygoid develops a process that extends outward
and downward and laps along the inner inferior one fourth of the quadrate. Ante-
rior to this lateral notch the bone again expands, the lateral extent of which is hidden
by the posterior portions of the lower mandible. More anteriorly this element is
again excavated by a suture that extends inward and forward, and it is by this
suture that the palatines (of modern nomenclature, transverse bones of Seeley and
Frass, the ectopterygoid of Owen), are united with the pterygoids. From this point
the latter extend forward and inward, meeting medially, the anterior tapering
extremities finally disappearing under and between the vomers. The surfaces of
the pterygoids are smooth, both of which dip at a slight angle toward the center of
the palate.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 95
Palatines (pl.). —'The palatines are next in size to the pterygoids of the bones of
the palate. They are flat plate-like elements that unite posteriorly with the ptery-
goids and laterally with the pterygoids and vomers. The outer and anterior extent
of these bones cannot be satisfactorily determined from our specimens.
Vomers (v.). — These elements are only shown in one specimen, No. 603. Those
parts interpreted as vomers are two narrow rod-like bones that begin along the outer
anterior margins of the pterygoids. Anteriorly they follow the outline of the
skull gradually converging toward one another finally meeting on the median line
just before they disappear under the premaxille.
There is no evidence of a bone between the vomer and palatine as shown by
Frass*' in Ichthyosawrus.
Mandible.— The mandible appears essentially like that of Ichthyosawrus. It
consists of five and probably six pairs of bones, the dentary, articular, angular,
Fria. 1. Cross-section of upper and lower jaws (No. 603), taken at the fracture just above the figure 2 seen in the
lateral view of the skull, Plate X. Onehalf natural size. ag., angular ; d., dentary ; mz., maxillary ; na., nasal ; pme.,
premaxillary ; pt. ?, pterygoid ; s.ag., surangular ; spl., splenial. rf
Fia. 2. Cross-section of the lower jaw of the same taken just anterior to the union of the two rami. One half
natural size. d., dentary ; spl., splenial.
surangular, splenial and coranoid (?) respectively. The element here called
splenial is the operculeer of Frass.
Dentary (d.). —The most anterior and largest of the bones of the lower jaw is the
dentary. It forms the greater part of the fore portion of the lower mandible. The
posterior extremity ends under the orbit, its posterior suture running obliquely from
below the maxillary to a little beyond the posterior end of the symphysis on the
lower border. The superior side forms the shallow alveolar channel in which the
teeth are supported. (See figs. 1 and 2.) Posteriorly, the thin inner wall of the
31 Frass, E., lit, cit., Pl. IL., fig. 2.
96 MEMOIRS OF THE CARNEGIE MUSEUM
grooye is supported by the quite heavy vertical plate of the splenial, but anteriorly
the latter element is supplanted by the higher and thicker inner wall of the dentaries
which here exceeds, the outer in height. In specimen No. 878 this inner wall, both at
the anterior and posterior parts of the channel, shows some faint vertical ridges, indic-
ative of alveolar compartments. Viewed from below the anterior portion of the den-
taries is evenly rounded. Just below the outer alveolar border is a shallow channel
into Which several vascular foramina open at the anterior end of the jaw.
Splenial (spl.).—The splenial in Baptanodon as in the other members of this fam-
ily is a long vertical plate that is applied to and forms the greater portion of the inner
surface of the posterior part of the ramus. The anterior portion extends below the
dentary a little and is visible from a lateral view of the mandible. Anteriorly it
meets its fellow medially where they unite and pass between and under the dentaries.
These appear to be ankylosed at the symphysis. (See fig. 2.) Posteriorly the lower
margin of the splenial is gradually confined to the inner side and rises obliquely for
a considerable distance. This suture at the posterior end of the jaw is obscure and
the boundary between the splenial and coranoid (?) cannot be determined, but the
latter element probably continues to the end of the ramus, lapping along the inner
side of the articular, assisting the angular and surangular in holding that element
in position. (See Pl. XL., fig. 2.) On the internal surface, just posterior to the symphy-
sis is an elongated oval foramen that probably represents the ‘‘internal mandibular
foramen” of the crocodile, though in this case it appears to be wholly enclosed by
the splenial while in the crocodile the splenial only forms the anterior border,
Posterior to this foramen, froma point on the lower inner side of the mandible, radi-,
ating linear impressions extend forward and upward to the posterior border of the
vacuity. They would seem to indicate a surface for muscular insertion.
Surangular (s.ag.).— Viewed laterally the surangular is a long slightly-bent bone
that forms the upper posterior margin of the ramus. On its inner posterior side this
bone develops a concave surface (see sq., fig. 3) which with the anterior end of the
articular forms the articulating surface for the quadrate. Just in front of this artic-
ulating surface is a node-like protuberance which Frass has pointed out as an
arrangement to prevent the dislocation of the lower jaws. (See fig. 3.)
The posterior part of the suture between the angular and surangular is not shown
by this miterial, though it is indicated by a dottel line, after the fractures on the
rami of No. 878 which were considered as taking the course of these breaks. Ante-
riorly the surangular gradually tapers to a point which from a lateral view disap-
pears between the dentary and angular just below the nares, although the elements
are continued still farther forward on the inside. (See s.ag., fig. 1.) ‘The lateral sur-
GILMORE : OSTEOLOGY OF BAPTANODON (MARSH) 97
face of the posterior end of the ramus in Baptanodon is composed of the angular and
surangular, the former contributing the greater share. In most of the Ichthyosaurs
the latter element contributes the greater part.
Angular (ag.). — The angular is a long curved bone that forms the lower posterior
boundary of the jaw. The anterior end tapers to a point similar to the surangular
and disappears between the splenial and dentary just in advance of the anterior end
of the surangular. (See Pl. VIII.) The suture between this bone and the splenial
posteriorly passes to the inner side. The angular then underlaps the splenial and
Fic. 3.. Internal view of the posterior end of the left mandibular ramus of Baptanodon discus (No. 603). One
half natural size. art., articular ; ag., angular ; cor., coranoid (?) ; s.ag., surangular ; s.q., articulating surface for the
quadrate.
coranoid (?) and more posteriorly develops an inner plate that encloses the lower
part of the articular.
Articular (art.). —The articular apparently is the least understood of any of the
elements composing the Ichthyosaurian jaw. Of all the literature to which I have
had access I have yet to find an adequate description of this bone. Fortunately the
mandible pertaining to specimen No. 878 has both of these elements preserved in
situ, while the right articular of No. 603 was found nearly in position but has been
detached for purposes of study. It is this element upon which this detailed descrip-
tion is based.
As in most reptiles the articular is placed far back in the ramus. In Baptanodon
it is held in position by the enclosing walls of the angular, surangular and cora-
noid (?). The angular entirely encloses the lower portion curving up on the inner
side, developing even more of an inner portion than is found in the Crocodile.
Viewed superiorly the articular is a short solid subrectangular bone (see figs. 4 and
5.) The upper surface is smooth and gently convex transversely, quite flat antero-
posteriorly with the exception of an elevated transverse ridge at either end. Poste-
riorly the end is subtriangular in form the longer side looking downward and out-
ward. ‘This end is somewhat convex from above downward, presenting a slightly
98 MEMOIRS OF THE CARNEGIE MUSEUM
roughened surface. The outer lateral surface of this bone is concave from above
downward (see fig. 5) and roughened for ligamentous attachment to the surrounding
bones. The inner surface is irregularly concave and roughened down to the inferior
wedge-like edge. Anteriorly the end resembles that of the posterior view though
somewhat dished from above downward, and more compressed transversely. It
was this end of the articular with the surangular that formed the concave articulating
surface (sq.), for the quadrate. There is no anterior process or elongation of this
bone and in this respect the articular of Baptanodon
is much shorter than the corresponding element in
the Crocodilia.
In the posterior position of the articular the man-
ner by which it is retained in the jaw and the part it
takes in forming the articulation for the quadrate it
Fig. 4. Internal view of left ar-
is most nearly approached by the articular of Chelon
ticular of Baptanodon discus (No. 603). f OK y/ Che Y
One half natural size. a., anterior end ; maidas.
p., posterior end ; sq., articulating sur- I have observed several figures of the lower jaw of
face for quadrate.
ye eee ee CU OS UUS which show the articular as a small
bone. One half natural size. d., dorsal triangular bone at the posterior end of the ramus
surface ; s.g., articulating surface for Jying between the surangular above and the angular
NL ak Aa case below and visible from a side view. In the speci-
mens considered here the articular cannot be seen from a lateral view. The
coranoid (?) apparently overlapped the inner margin above the angular.
Coranoid (cor.). — Of our knowledge of the extent and nature of the coranoid in
Baptanodon little can be said at this time. In our specimens it appears as a com-
paratively thin plate that extends along the inner side of the posterior end of the
ramus, lapping over the lower border of the articular, thus assisting the angular and
surangular in holding that element in position. Nothing is known of its extent
anteriorly.
Iam pleased to acknowledge my indebtedness to Dr. J. C. Merriam for the
identification of this element in the material under discussion. On his return from
Europe last year he examined the skulls and kindly gave me the benefit of his ob-
servations.
Dentition. —'The presence of teeth in the American species has long been sus-
pected by many paleontologists, but until recently” there has been no positive evi-
dence of their existence. The shallow dental grooves combined with the reduced
size and undoubtedly loose attachments of the teeth, will fully account for their
* Gilmore, C. W., ‘‘ Discovery of Teeth in Baptanodon, an Ichthyosaurian from the Jurassic of Wyoming,’’ Scienec,
N. S., Vol. XVI., No. 414, pp. 913-914, Dec., 1902.
‘f
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 99
absence in previously discovered specimens. The English form Ophthalmosaurus
icenicus shows about the same peculiarity in the retention of the teeth. Baur® has
observed small teeth at the end of the jaws of the (Baptanodontidee) Ophthalmosawrus
contained in the private collection of Mr. Leeds in Peterborough, England. Lydek-
ker™ also speaks of alveoli being present at the end of the jaw in this genus, although
the groove is known to extend posteriorly.
In the jaws of No. 878, although the alveolar channels are better preserved than
in specimen No. 603, there was not even a fragment of a tooth recovered, but we have
evidence of their existence at one time in the faint alveolar partitions preserved on
the inner wall of the right dental groove. (See Pl. VIII.)
As suggested in a preliminary paper in Science the final exhumation of the jaws
of No. 603 revealed more teeth, nine in all, six of which were in the groove of the
upper jaw, the remaining three being attached by matrix to the
alveolar surface of the lower mandible. With but one exception
all of the teeth were lying prone upon the surfaces of the grooves.
The largest and best preserved tooth found was cemented to the
lower jaw by the matrix just anterior to the forward extremity of
the left maxillary. The tooth as preserved (see fig. 6) measures 29
mm. in length, and undoubtedly represents the teeth of the pos-
terior part of the series. The base (see fig. 7) is somewhat anguiar
in cross section, but as the tooth rises it becomes more
rounded ending in a circular subacute apex. There is
no swelling of the base as may be observed in the teeth
Fia. 6. Poste-
rior tooth of Bapta-
of many of the Ichthyosawrs. A little more than the en a
nies cna eUPbes third is covered with enamel which is impressed Gross _ sec-
603). Twicenatural with fine longitudinal strize, between which are interven- tion of the
i 7 : 5 6 0 same tooth
size. The transverse ing depressions. ‘These grooves begin quite abruptly at
fracture represents (No. 603).
the point at whicn the base of the enameled surface and extend upward, Nearly
the cross section was gradually subsiding before reaching the apex which is twice nat
gree smooth. A cross section near the mid portion of the oie
cement covered base shows the tooth to have a somewhat flattened periphery at
either end of the line of its greatest diameter. Considering these surfaces as the
contact between the tooth and the outer and inner walls of the dentary groove, the
tooth would curve in slightly. But it is hard to understand how the shallow and
widely separated walls of this groove could ever have been in apposition with the
ch) Baur, lit. cit.
34 Lydekker, R., ‘* Catalogue of Fossil Reptilia and Amphibia in the British Museum.’’
100 MEMOIRS OF THE CARNEGIE MUSEUM
teeth it bore. The teeth of the median part of the snout were more slender, but
otherwise appear to be identical with the tooth described above. The two most
anterior found at a point just posterior to the tip of the rostrum were small, more
eone-like, but otherwise similar to those preserved posteriorly. It would appear
from the evidence before me that: DBaptanodon was well provided with comparatively
small, somewhat slender but functional teeth that extended along the full length of the
jaw; the most anterior ones being much reduced.
The teeth of B. discus differ from the one tooth known of B. natans in the per-
fectly smooth enameled surface of the latter.
Thyrohyal (th.). — Lying parallel to the posterior part of the right mandibular
ramus of No. 878 was a subcompressed rib-like bone which T have considered the
thyrohyal as described by Owen. There was no evidence of the corresponding bone
of the opposite side. Both ends are expanded, more especially the posterior which
Fia. 8. Right thyrohyal of Baptanodon discus (No. 878). One half natural size. 1, view of the inferior border ;
2, lateral view ; a., anterior end ; p., posterior end.
is almost twice the width of the anterior. PI. XI, fig. 1 (th.), gives the position of
the element as retained in the matrix.
An inspection of fig. 8 (1 and 2) shows this element as being curved from end
to end both vertically and laterally.
MEASUREMENTS.
No. 878. Greatest length of thyrohyal.....: SOS OSD DaGEABCeScBNCBEcdanSabdaeABuCoooND
“878. ‘¢ width of posterior end
v (eirtsh “s oy anterior ‘‘
Tur VERTEBRAL CoLUMN.
The material under discussion is not well adapted for a systematic study of the
several regions of the vertebral column, and although vertebre are preserved from
the different regions of the back bone, the vertebral series is imperfect in all of the
skeletons in this museum. The description of this part of the axial skeleton will
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 101
be supplemented somewhat by the more complete vertebral series in the collections
of the University of Wyoming.
With skeleton No. 603 there are thirty or more vertebre in various degrees of
preservation, but there are none arranged serially. However, every part of the
column is represented.
Specimen No. 878 has a series of twenty-one vertebree extending from the atlas
backward. (See Pl. VII.) The first fifteen are in a fair state of preservation but the
remaining six are very imperfect. The spinous processes are only preserved in a few
instances.
No. 919 which is a much larger individual has a series of ten cervicals com-
mencing with the atlas, a second series of eleven from the anterior dorsal region
posterior to the point where the diapophyses become distinct from the neurapo-
physial articular surface; a third section of eleven posterior dorsals commencing
just posterior to the first vertebra having the diapophysis and parapophysis united
to form a single node-like articulation. The fourth and last section contains parts
of twelve anterior caudals, these show the rapid decrease in the size of the centra
posteriorly as previously pointed out by Knight.”
The centra in all regions anterior to the extreme caudals are deeply biconcave.
These concave surfaces, with the exception of the anterior face of the atlas, begin
close to the periphery and slope in rapidly but evenly to the center. In this
respect Baptanodon may be distinguished from O. icenicus, as Lydekker * observes
of the vertebree referred to the latter genus. ‘The cervical region with the cupping
of the anterior face of the centrum confined to the central portion, and surrounded
by a flattened periphery.” There are some isolated centra of this character in the
collections of the Yale Museum, labelled as coming from the Baptanodon Beds of
the Rocky Mountains. From this slight evidence it would appear as though the
genus Ophthalmosawrus may also occur in this country.
The upper arches in this genus as in all previously described Ichthyosaurians
are free from the centra and were united to them by synchondrosis. The centra
are always short antero-posteriorly as compared with their breadth and height.
Atlas and axis (at. and ax.).— In Baptanodon as in nearly all adult members of
the Ichthyosauria, the centra of the atlas and axis are completely fused, so much so
that one would hardly suspect the existence of two vertebree if it were not for the
presence of the two sets of arches upon their dorsal surfaces (see figs. 10 and 26).
These vertebrae are represented in all three specimens, Nos. 603, 878 and 919. Those
35 Knight, W. C., ‘‘Some Notes on the Genus Baptanodon, with a Description of a New Species,’? Am. Jour. of
Sei. (4), Vol. XV., 1903.
36 Lydekker, lit. cit.
102 MEMOIRS OF THE CARNEGIE: MUSEUM
of No. 878 are in the best state of preservation and are the elements from which
the text figures have been drawn. Though not found in position (see P]. VII.) they
are readily distinguished from the succeeding vertebree by their more modified
centra and by their dorsal and transverse processes.
Seen anteriorly (see fig. 9) the centrum of the atlas is pentagonal in form, wider
Fic. 9. , Anterior view of atlas and axis of Baptunodon discus (No. 878). One half natural size. d., diapophy-
sis; ., paired neural arch of atlas; s., spinous process of axis; z., apophysis with which the first intercentrum
articulated.
Fie. 10. Lateral view of atlas and axis of Baptanodon discus (No. 878). One half natural size. d., diapophysis of
theatlas ; n., paired neural arch of atlas ; p., and p.’, parapophysis of the atlas and axis respectively ; s., spine of the
axis ; z., apophysis of the atlas ; z.’, apophysis of the axis.
than deep and considerably contracted inferiorly. The concavity for the reception
of the occipital condyle instead of sloping from the outer margin to the center as in
the centra that follow, has a narrow, flattened periphery, which is much enlarged
laterally and inferiorly. The cup of the centrum descends evenly, perhaps more
sharply, from the superior margin down to the center. The inferior surface presents
a slightly rounded subtriangular face which looks downward and forward, its
inferior extent reaching nearly to the middle of the coalesced centra. If present it
was with this face or apophysis that the first hypophysis or intercentrum articulated.
The superior lateral surfaces are quite extensive and curve backward to form the
“para- and diapophyses. The dorsal surface is about equally divided between the
a oie
‘that vertebra. The lateral margin below the
axis is deeply concave, the surface) slopes in
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 103
neural canal and the surfaces for the attachment of the pedicles of the neural arch,
the two surfaces being separated by narrow antero-posterior ridges.
The lateral surface of the combined centra (see fig. 10) is somewhat concave
antero-posteriorly. Superiorly the neurapophysial surface extends outward and
-coalesces with the heavy articular diapophysis. There is no indication of a dia-
pophysial surface on the axis such as is seen in the axis of B. marshi (fig. 26), or the
axis of I. longifrons, Owen. Just below the diapophysis and separated from it by a
non-articular tract is an antero-posterior ridge at either end of which arise articular
prominences which represent the parapophysis
of the atlas and axis respectively. The parapo-
physis of the atlas (p.) being the larger of the
two. The round tubercle like parapophysis (p.’)
of the axis is placed well posterior on the side of
parapophysis is concave antero-posteriorly even
down to the furthermost extent of the inferior
subtriangular part of the centra. I have ex-
amined the coalesced centra pertaining to six
different Ichthyosaurian skeletons from the Ju-
rassic of North America and have yet to find a
trace of the suture uniting the atlas and axis.
Posteriorly (see fig. 11) the centrum of the
gradually from the outer margin for half of the
radius, thence more sharply to the center. In-
feriorly the lower surface is produced into a sub- Fic. 11. Posterior view of the atlas and
triangular face which looks downward and back- axis of Baptanodon discus (No. 878). One half
ward, at a somewhat sharper angle (see fig. 10 Se ame une Spit cde ee
neural arch of the atlas; p., parapophysis of
than the apophysis of the atlas. This would be atias; s., spine of the axis; zyg., posterior
the articulation for the third intercentrum if ygapophysis of the axis; z.’, apophysis for
that bone still exists as a separate element in pia ee can a
this form. The axis of B. marshi, shows no such face, though the apophysis of the
atlas appears to still be present. The axes of both No. 603 and 919 appear to be
identical with that of No. 878. In these forms it would seem that at least two
intercentra exist as separate elements while the one between the atlas and axis has
become completely anchylosed to the centra. I haye examined the atlas and axis
of B. marshi (see fig. 26) and find here a still greater reduction in the number of
104 MEMOIRS OF THE CARNEGIE MUSEUM
the intercentra. There could not have been more than one retained and it may
be possible that all have disappeared. ‘This indicates a higher degree of specializa-
tion than has hitherto been found to exist among the Ichthyopterygia. The Trias-
sic form Shastasawrus,”” Merriam has five intercentra while in Ichthyosawrus there
are but three.
The paired neural arch is the only process supported by the atlas. They extend
upward and backward overlapping the lateral basal portion of the spine of the axis
and never uniting to form a spine. The left half of this arch was found displaced
though in an excellent state of preservation. Viewed from the side (fig. 12, b), it is
Fig. 12. Left half of paired neural arch of atlas of Baptanodon discus (No. 878). One half vatural size. «a, seen
anteriorly ; b, seen laterally ; c, view of distal or articular end.
Fic. 18. Superior view of an anterior cervical of Baptanodon discus (No. 608). One half natural size. a, anterior
d and « the combined dia- and neurapophysial surfaces ; n.c., neural canal ; p, posterior.
Fig. 14. Anterior view of anterior cervical of Baptanodon discus (No. 603). One half natural size. d and 2, the
coalesced diapophysis and neurapophysial surfaces ; n.c., neural canal ; p, parapophysis.
an irregular spear-shaped compressed bone that articulated with the centrum by an
expanded subtriangular articular end (fig. 12, c). An anterior view (fig. 12, a)
shows the sinuous curves made necessary for it to fit closely to the process of the
axis. ‘The function of these bones appears to be the same as the corresponding ele-
ments of I. longifrons as described by Owen, which they resemble somewhat in shape
and position. This paired neural arch does not develop zygapophyses. The pedicels
of the axis as they rise from the surface of the centrum converge and meet above the
neural canal to form a well-developed neural spine, the first of the vertebral series.
There are no indications of prezygapophyses on the spine of the axis belonging to
No. 878, though the process of B. marshi shows such articulating surfaces. ‘The pos-
terior portion of the spine is broken away so that its extent antero-posteriorly can-
not be determined accurately from our specimens, and it would appear after an
37 Merriam, J. C., ‘‘ Triassic Ichthyopterygia from California and Nevada,’’ Univ. of Cal. Pub., Vol. 3, No. 4, p. 75.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 105
examination of the axis of B. marshi that the restored spine in fig. 10 is too narrow
antero-posteriorly.
Vertebree Posterior to the Avis. —The succeeding cervicals are very similar in their
”?
general form which perhaps is best described as “shield shaped.” Fig. 14 is a
good example of the cervical centrum and the description given here may be con-
sidered typical of the vertebree of this region.
On the median superior surface is the smooth tract forming the basal boundary
of the neural canal (see fig. 13) on either side separated by slight antero-posterior
ridges are the shallow roughened neurapophysial surfaces for the attachment of
the pads of cartilage upon which the pedicels of the neural arch rested.
This roughened surface is placed largely on the anterior half of the cervical and
extends outward and downward becoming confluent with the diapophysis (see fig.
14). The parapophyses are node-like projections placed half way down and on the
anterior margin of the centrum, being separated from the diapophyses by a concave
longitudinal depression. The position of these two processes remains unchanged as
far back at least as the fifteenth vertebrae, as will be seen in Pl. VII.
The anterior vertebrae are broader than high and gradually increase in length
posteriorly. From the splendid series of forty-one precaudal vertebre pertaining
to the type of B. marshi, Dr. Knight has observed that the maximum length is
reached in the nineteenth from the skull.
Continuing back in the column (see fig. 15) it will be observed that the centra
remain about the same height but the transverse a
extent becomes considerably greater, also the artic-
ular surfaces of the diapophyses which are conflu-
ent with the neurapophysial surfaces in the cervical
region become separated. In Ichthyosawrus this
separation takes place on either the fourteenth or
fifteenth vertebra. Dr. Merriam has shown that in
one species of Shastasaurus it does not occur until
the thirty-fifth or later. The point at which this Bie by CESSES OF ein EBENIO?
dorsal centrum of Baptanodon discus (No.
transition is brought about in Baptanodon cannot FUE GROEN Peelers ch Ghereorsiw
be determined from the specimens under discus- sis; p, parapophys; x, neurapophysial
sion, although we have evidence that they do not sutface:
separate until the sixteenth or Jater.* (See Plate VIL.)
38 In the Am. Jour. Sci., Vol. 50, 1895, Marsh figures a vertebra of Baptanodon natans, designated as a cervical,
The position of the dia- and parapopbyses half way down on the side of the centrum at once shows this vertebra as
pertaining to the region posterior to the neck. This figure was published a second time in his U. S. G. S. Monograph of
the Vertebrate Fossils of the Denver Basin in 1897.
106 MEMOIRS OF THE CARNEGIE MUSEUM
From the cervicals back through the dors sal region the breadth remains greater
than the height but the shield-shaped form of the cervical centra gradually changes
to what has been called “pear shaped” (see figs. 16 and 17), in the regions here
(2
Fic. 16. Cross-section of a dorsal centrum of Baptanodon discus (No. 603). One half natural size. d, diapophysis ;
p, parapophysis ; z, neurapophysial surface.
Fic. 17. Cross-section of posterior dorsal or anterior caudal centrum of Baptanodon discus (No. 603). One half
natural size. jp, (?) parapophysis; x, neurapophysial surface.
considered as middle and posterior dorsals, and this form gradually assumes the
compressed elliptical contour seen in the caudals (see fig. 19).
The change in form of the centra brings a corresponding transition of the dia-
and parapophyses. These processes gradually move down the side of the centra,
until finally only one oblique process remains on the extreme lower side of the
centrum. ‘This type probably represents the posterior dorsal region (see fig. 17).
Fic. 18. Lateral view of centra from different regions of Baptanodon discus (No. 603). One half natural size. 1,
anterior cervical ; 2, anterior dorsal; 3, dorsal; 4, posterior dorsal or anterior caudal; d, diapophysis ; p, para-
pophysis. ‘
In Ichthyosawrus Owen described the reduction to a single facet as a union of
the two ’pophyses. He thought the diapophysis dropped more rapidly than the
parapophysis, thus meeting the latter with which it united to form the single oblique
process.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 107
The evidence to which I have had access is not conclusive but it has suggested
to me that in Baptanodon this is not a union of the two but a development of the
parapophyses with a consequent reduction and final passing away of the diapophyses.
(Compare 1, 2, 3, 4, fig. 18.) It will be noticed in the cervical (1) region that the
diapophyses presents the more robust articular surface of the two; the anterior dorsal
(2) shows the two facets nearly equal; the median dorsal (3) indicates an increase
in size of the parapophysis with a corresponding decrease of the diapophysis. It
will be observed that in all three regions the distance between the facets remains
about constant.
In the Triassic genus Shastasawrus Dr. Merriam has shown that the parapophyses
become obsolete in the anterior dorsal region with a consequent increase in the size
of the diapophyses.
The single process in Baptanodon gradually decreases in size posteriorly and
finally fades away at some point in the caudal region as is shown by fig. 19. Owen
has shown that this single process in J. communis disappears on the eightieth vertebra
and it is at this point that the downward bend or depression of the tail takes place.
Professor Knight has added some important information to our
knowledge of the tail of the American form as follows: ‘In speci-
men ‘TT’ in the same collection (University of Wyoming) there are
forty-six consecutive caudal vertebrae. ‘These are of the usual ichthy- Fig. 19. 1
osaurian type and represent an animal that had an extremely long and 3"¢ 2 end and
lateral views of
slender tail. The reduction in the size of the vertebree occurs very an extreme cau-
near the body and within a distance of a few inches, the vertebree dal of Baptano-
don discus. (No.
603). One half
reduction have reduced margins,” in fact in two of them the artic- natural size. ne,
decrease in diameter over one half. The vertebree in the area of
ulation nearly meets upon the side of the centrum. * * * Although ln!
caudal vertebree from at least a half dozen different animals have been examined,
no trace of chevrons has been observed, and the vertebree lack chevron facets.”
On the anterior vertebrze the neural arches are held together by well developed
zygapophyses. The anterior arches have paired zygapophysial facets which unite
to form a single median one at some point between the sixth and ninth cervicals.
This information is derived from the cervical region of No. 878. The zygapophysial
facets have their surfaces roughened indicating as shown previously by Dr. Merriam
in the genus Shastasaurus, the presence of considerable cartilage.
A number of the anterior cervicals of No. 878 (see Pl. VII.) have the arches and
39 It occurs to me that these vertepre with reduced margins may represent the point of divergence of the vertebral
column into the lower lobe of the candal fin.
108 MEMOIRS OF THE CARNEGIE MUSEUM
spines preserved nearly in place, the anterior zygapophyses of these look upward
and slightly inward, the planes of which if continued to intersection would meet at
an angle of 160° to 170°. The posterior zygapophyses look downward and some-
what outward, being separated by a slight vertical depression.
The neural arches as they rise from the centra are slightly inclined backward.
This inclination is continued in the spinous process at a still greater angle. The
spines of the anterior vertebree are greatly compressed laterally, being flattened into
a subquadrate plate of bone that is somewhat thickened above. Of the spinous proc-
esses of the posterior vertebrae nothing of importance is shown by the material under
discussion, though the articular bases of the pedicels must undergo modifications
corresponding to those that take place on the dorsal surface of the
centra of the several regions. Anteriorly the base is subtriangular in
form and somewhat protuberant, but upon the centra where the sep-
aration of the diapophysis and neurapophysial surfaces takes place
the articular end becomes long and narrow, and rests upon corre-
sponding ridges which rise from the dorsal surface of the centrum.
(See fig. 17, x.)
Ribs. — Though there are numerous parts of ribs preserved with
the different skeletons no complete ribs have been found. Those
pieces lying on either side of the anterior vertebra of No. 878 (see
Fig. 20. Pos- Pj. VII.) show the ribs of this region to have been very long and
terior view of a
right dorsal (?) ‘ : eae!
rib of Baptano- although both ends are wanting. This evidence appears to indicate
slender. On the left side of the column (7’) is about 725 mm. long
don discus (No. a very deep body cavity anteriorly. Corresponding to the form of
603). One half
aa the anterior vertebrae the ribs have a distinct tuberculum and capit-
natural size.
ulum. ‘he latter probably as in the other groups of the Ichthy-
opterygia disappears in the posterior dorsal region. As yet the abdominal splint-
ribs so numerous in Jchthyosaurus have not been observed in this genus.
Pectoral Arch. — Plate XII., Figs. 1, 2 and 3.
The pectoral girdle of Baptanodon is represented in two specimens, Nos. 878 and
919, the former having the most complete arch yet discovered of this genus.
Fig. 21 shows the inferior view of the elements as they were retained in the matrix.
The girdle pertaining to No. 919 though incomplete has been entirely removed
from the matrix and it gives us a good idea of the anterior and superior aspects of
this region. (See Pl. XII., figs. 1 and 2.) As mentioned previously this specimen
may prove to belong to a different species when more complete material is known.
It will be observed in fig. 21, that with the exception of the clavicles and inter-
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 109
clavicle the other elements have been retained in their mutual relations and estab-
lishes for all time the relative position of the scapulee and coracoids of this genus.
The position of the clavicles and interclavicle can be determined with a fair degree
of accuracy. An examination of the roughened surfaces (a and x), for ligamentous
attachment on the anterior borders of the scapulee shown best in Pl. XIL., figs. 1
and 2, No. 919 evidently indicates the place of attachment for the clavicles. The
firmly united clavicles evidently curved up along the anterior borders of the scapulze
and coracoids, being attached to the former by ligaments at 7 and «, to the latter
by the interclavicle, which fits into a groove on the posterior median side of the
Fic. 21. Ventral view of the pectoral girdle, as found in the matrix. Baptanodon discus (No. 878). One fifth
natural size. cl, clavicle ; co, coracoid ; 4, humerus ; 7.c/, interclavicle, sc, scapula.
clavicles and extends back until its upper excavated surface laps under the anterior
median union of the coracoids, the position in extant Lizards, thus forming a brace
between the two parts of the girdle. In both specimens the girdle is formed essen-
tially in the same manner as in Ichthyosawrus.
Dr. Knight’s observation that: “There was no evidence of an interclavicle, and
the peculiar union of the coracoids precludes an interclavicle of the regular Ich-
thyosaurian type. In consequence the interclavicle in Baptanodon must be consid-
110 MEMOIRS OF THE CARNEGIE MUSEUM
ered rudimentary or wanting,’ must have been based upon a comparison of the
coracoids of Bb. marshi with those figures of Ichthyosaurus which show the inter-
clavicle as lying between these elements anteriorly. The portion of a girdle figured
by him appears to be identical in form with the arches under discussion, one of
which, No. 878, has a well developed interclavicle making it appear that Knight
was mistaken in his interpretation of this region.
Coracoid (co.).— The coracoids are broad subquadrangular bones that join one
another medially by large elliptical facets. These facets are roughened and were
evidently united by a heavy pad of cartilage, the width of which is best shown in
Pl. XIL, figs. 1 and 2. :
The internal or superior surfaces (Pl. XII., fig. 2), are flattened, though both
elements are gently inclined toward the median line. The external or inferior sur-
faces are concave transversely and convex antero-posteriorly. The lateral borders
are especially thickened forming a heavy articular face for the scapulee and humeri.
This outer articular end is divided into two unequal faces meeting in an obtuse
angle. The more anterior and smaller one of the two is for the scapula, and looks
outward, forward and obliquely upward. The larger and posterior surface forms
the greater part of the glenoid cavity. It is very slightly convex from above down-
ward and covered with tubercle-like eminences indicative of aheavy pad of carti-
lage. This articular end is supported by a broad neck formed by a deep notch on
the anterior margin and a slight emargination on the postero-lateral border of the
coracoid. This anterior notch Seeley suggests, probably corresponds to the foramen
found in the coracoids of the Dinosaurs.
The anterior border from the inner notch thickens rapidly as it recedes poste-
riorly to form the intercoracoidal facet (see Pl. XII., fig. 1). The posterior part is
compressed to a thin plate, the border being rounded from the inner angle of the
emargination to the median union of the two elements.
The coracoids resemble those of the long-snouted species or Latipinnate form of
Ichthyosaurs of which J. tenwrostris is a good example.
MEASUREMENTS. ?*°
No. 878. Greatest length of coracoid antero-posteriorly.....................++5 253 min.
“ 878. cs width “ ss transversely ue
oS SyIG) sf eee ee U0; =) grousonobdaonanopocodsanouDEccte 200 ‘
“* 878. us sc“ girdle end to end of scapule...................06+ 1 On
Scapula (sc.). — The scapula is a moderately long bone, the upper half being nar-
row with nearly parallel sides, the lower or articular portion is broadly expanded
10 Measurements given of No. 878 are made from the right element, which appears to be the least distorted.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) ital
antero-posteriorly and is especially thickened on the posterior margins where it enters
into the formation of the glenoid cavity. The articulating face for the coracoid joins
this border at an obtuse angle.
The scapula extends outward, upward and forward, not backward as Seeley sus-
pected in his original description of the type of Ophthalmosaurus. Both arches before
me have the scapulee retained in their natural relations to the coracoids and their
position may be considered as absolutely determined. Viewed longitudinally the
inner surface of the scapula is slightly concave, as it curves up to lap over the side
of the ribs. On the upper free end the inner surface is quite flat, distally the articu-
lating end is concave antero-posteriorly. The anterior border above is gently rounded
but as it approaches the articular end it widens into a flattened oblique surface (x)
that looks forward and upward. (See Pl. XIL., figs. 1 and 2.)
Seeley * regarded a part of this border as the acromion process. ‘The oblique
surface mentioned above is roughened and I have interpreted it as being the place
of attachment for either the ligaments or cartilage which held the clavicles in posi-
tion. The position of these elements is beautifully shown by a specimen of J. quad-
riscissus, No. 6293, in the Stutgarter collection of which Frass* gives a figure.
The scapulee of our specimens are not notched to fit over the articulating sur-
faces of the coracoids as figured by Knight but unite by nearly straight articulating
faces. An examination of Dr. Knight’s specimen, the type of B. marshi, convinces
me that the articular ends of the scapulee of that species are not different than those
of B. discus, but in the former case were crushed so that their true shape and rela-
tions could not be accurately determined. The lower surface of the scapula is convex
from end to end. Antero-posteriorly the free end is gently convex, but somewhat
coneayve at the expanded articular end. Between the marginal ‘‘acromion process”
and the articulation with the coracoid the internal border is especially compressed
and remains free.
Seeley has best described the humeral articulating border as follows: “The
humeral articulation is an expanded triangular thickening of the bone, extending
posterior to what would otherwise be an extension of the parallel sides of the free
or distal end.”
MEASUREMENTS.
No. 878. Greatest width of free end of scapula. .......... 55 mm.
CONG: sf COE CATGICU AbD Paces eneeceettedanesrcecectecee sett 3
878. CO) Tenaya) CO READE penonsoonoosaancceesb00000 onobenscaddasnbodceHoaECes up
Clavicles (cl.). —So far as 1am able to learn specimen No. 878 is the first Ich-
“Seeley, H. G., Quart. Jour. Geol. Soc. of London, Vol. XXX., pp. 696-707, 1874.
“¥Frass, E., lit. cit., Plate 1V., fig. 2.
Ui MEMOIRS OF THE CARNEGIE MUSEUM
thyosaurian from the Jurassic of America to have the clavicle and interclavicle
bones preserved. The position of these elements as retained in the matrix is well
shown in fig. 21. It will be observed that the upper extremity of the right clav-
icle as well as the right transverse end of the interclavicle are wanting.
Viewed anteriorly if complete the anchylosed clavicles are bow-shaped, widest at
the middle, gradually narrowing as they turn up along the anterior borders of the
scapule. The ends must have been directed upward, outward and backward to the
extent that either clavicle would be opposed to the oblique roughened surface” on
the inner anterior borders of the scapulee which look forward and upward. The
outer or upper third must have been free from the scapula though lying parallel
with the free end of that element.
The left clavicle which is quite complete at its upper end is subcircular in cross-
section. The broken end of the right clavicle is somewhat angular in cross-section.
On the median posterior side of the blended clavicles is a deep longitudinal groove
for the reception of the transverse portion of the interclavicle. The latter is shown
in fig. 21 a little removed to the left from its normal place in the clavicular girdle.
Professor H. G. Seeley has pointed out four different ways by which the clavicles
unite in Ichthyosaurus. 1, clayvicles anchylosed or connate; 2, clavicles meeting in
the median line; 38, clavicles not meeting but joining by squamous union with the
extremities of the interclavicle ; 4, clavicles united by a long squamous suture. He
adds a fifth in Ophthalmosaurus, 5, clavicles united medially by an interlacing suture.
He considered these differences of generic value, and the fifth was one of the impor-
tant characters upon which he based the genus Ophthalmosaurus.
A careful comparison of Seeley’s figures and description of the clavicles of
Ophthalmosaurus with those of specimen No. 878 shows many similarities. Although
the presence of a suture and the interclavicle wedged in between the ends of the
clavicles in the former seems to indicate a distinct difference from the anchylosed
clayicles of Baptanodon which show no evidence whatever of a suture at the median
junction.
Interclavicle (i.cl.). — With the exception of the right end of the transverse bar
(see fig. 21), this element appears to be complete. It is of the usual “T’’-shaped
form, though possibly not quite so robust as observed in many members of the
genus Ichthyosawrus.
The inferior surface of the posterior stem is rounded transversely. The parallel
borders of this part as they extend forward diverge rapidly forming a wide triangular
*8Cuvier in ‘‘Ossemens Fossiles’’ as early as 1824 points out that the scapula of Ichthyosaurus has at its anterior
edge a prominence which supports the extremity of the clavicle.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 113
plate whose outer apices are produced to form the ends of the transverse bar. This
transverse portion lies in a groove on the posterior side of the clavicles and when
in position only the stem could be seen from an inferior view of the girdle. (See
PL. XIL,, fig. 3.) An examination of fig. 21 shows the interclavicle slightly removed
to the left from its natural position in the groove of the clavicles.
The upper posterior side of the stem is gently concave from side to side, indicating
that it probably underlapped the median ventral sur-
faces of the coracoids, thus giving support to the for-
ward part of the arch.
Nothing is known of the pelvic girdle at this
time.
Anterior Limbs.— Professor Marsh briefly described Fic. 22. Inferior view of inter-
the fore paddle (probably pertaining to specimen No, ©lavicle of Buptanodon discus (No. 878).
One fifth natural size.
1958) of Baptanodon as follows: “In the fore paddle
the humerus alone is differentiated. Below this the bones of the forearm, the car-
pals, metacarpals and phalanges are essentially rounded free disks implanted in the
primitive cartilage. The radius may perhaps be regarded as a partial exception, as
its free margin is nearly straight and somewhat thinner than the remaining border.
There are three bones of nearly equal size in the first row below the humerus. The
radius may be identified with certainty by its position. The next bone evidently
corresponds to the intermedium * and the third or outer one, to the ulna. In the
succeeding row there are four subcircular bones, and five in the next series. These
represent the carpals. ‘There are six metacarpals, and also six well-developed digits,
each composed of numerous phalanges, which are free and nearly circular in form.”
Knight has given additional information of the paddle bones in the following
lines: “The carpals, metacarpals and phalanges are compressed grooved cylinders
the most of which have slightly concave surfaces. The grooves are ornamented with
tuberosities for musular attachment. Along the margins of the limb the cylinders
have their exterior borders reduced to quite thin edges. Anyone finding the limb
of a Baptanodon for the first time scattered about in the field would surely try to fit
the ventral and dorsal surfaces of the metacarpals in trying to construct a digit.”
There is nothing to be added to this description by our material which consists
of the proximal end of a humerus (see fig. 21, h), and a few miscellaneous paddle
bones.
Humerus (h.). — Knight describes the humerus of B. marshi briefly as follows:
44 Now considered the ulna, while the succeeding element is identified as the pisiform. Dr. Williston proposes
the name ‘‘ epipodial supernumerary ’’ for the latter.
114 MEMOIRS OF THE CARNEGIE MUSEUM
“ Humerus about one third the length of the limb, with a stout twisted shaft that is
greatly compressed near the distal end. Planes passed through the articulate ends
of the humerus stand at an angle of 50°. The head is slightly rounded and is
almost identical with Ichthyosawrus. ‘There are three distal facets; but they are
not of equal size. The facet for the ulna (see fig. 24, w) is the largest, the one for
the radius (7) next in size and the one opposite the pisiform (p) is rudimentary, for
that bone was held in cartilage and did not articulate with the humerus. The
Fic. 23. Superior view of fore paddle of Baptanodon marshi, Knight. Reduced about 2} times. A, humerus;
p, pisiform ; 7, radius; uw, ulna; IL (?) second digit, first one wanting.
facets are elliptical in form, and those opposite the ulna and radius elongated in the
plane of articulation.”
MEASUREMENTS OF HUMERI PERTAINING TO ENGLISH AND AMERICAN FORMS.
| | | a
| | Width | Width |
| “greats | Proximal Distal | Remarks.
pb | i) le
100 mm.) 115 mm.| Type of species.
No. 1955. Specimen in Yale Museum, B. discus.....| =
|
|
WM, CRE ff ‘“ Univ. of Wyoming, B. marshi.| 190 1 eal
No. 878. ft “* Carnegie Museum, B. discus... — 115mm.) — | Distal end wanting.
| 130 | Type of the species.
IN, 1%, IB, “« British Museum, 0. icenicus...| 145 | 124
No. 47885. “ us ve | 160 | 145 |
Cotype O. icenicus in Leeds collection..... ........-++-+++| 164 115 _| 152 | Measurements given by Seeley. _
Posterior Limbs. — Professor Marsh in his original description of B. discus
describes and figures a beautifully preserved hind (?) paddle (see fig. 25) of that
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 115
species. Having made a personal examination of the extremity mentioned above
I find the figure to be a faithful reproduction of the limb, which clearly shows the
upper limb bone as having three facets, distally as originally described. However
Dr. Knight has observed that the distal extremities of the femora examined by him
Fig. 24. (a) dorsal view of humerus of Baptanodon marshi from type specimen. One half natural size. h, hu-
merus ; p, articular surface opposite the pisiform ; r, articular surface for radius; ¢, trochanter; w, articular surface
for ulna. (b) View of distal articular end of the same. One half natural size.
all have two facets only. This isimportant and raises the question as to the correct
determination of the paddle designated as the left pelvic limb of B. discws by Marsh.
This would be an important character for distinguishing the genus Baptanodon
from the closely allied genus Ophthalmosawrus, which is accredited with three facets
on the distal end of the femur. A photograph before me of askeleton of Ophthal-
te e) CO) @©000., 1
a — Y) @BOO0000,,
Oe Le) @O@Oo OO,
@@ O00000¢,,
9 (O09 Ooo¥
7) @ QO?
Fic. 25. Left hind (?) paddle of Baptanodon discus (No. 1955) seen from below. One eighth natural size. f,
femur ; ¢, tibia; fi, fibula; p, pisiform or epipodial supernumerary ; /, first digit ; v, fifth digit.
mosaurus shows the pelvic limb as being much smaller than the pectoral and this
would strengthen somewhat the idea that Professor Marsh was mistaken in his inter-
pretation of this extremity.
So faras I am able to learn there has not been a hind limb of Baptanodon found
intact and at this time it will be impossible to definitely determine the correct solu-
116 MEMOIRS OF THE CARNEGIE MUSEUM
tion of this problem, although from Knight’s evidence and the close resemblance of
this paddle to the fore limbs known, I am inclined to believe that Marsh was
wrong and that the limb figured and described by him as pelvic is really a pectoral
extremity. It will be noticed by an examination of Marsh’s drawing (see fig. 25)
that there is no contact between the proximal and distal portions of the femur (?)
which renders it somewhat more difficult to make accurate comparisons.
The additional finger developed in Baptunodon and the fact that some of the
Ichthyosaurs have as many as ten, is explained by Dr. 8. W. Williston as follows :*
“The additional fingers of the Ichthyosaurs may be also explained in a similar way.
The margin of the flipper has become hardened by fibro-cartilage, which by the
movement of the fingers was broken into segments, each of which finally took on
ossification. A division of the phalanges by segmentation would certainly have to
take place immediately in order to preserve the integrity of the paddle as an organ
of propulsion. If this explanation be correct, then the additional ossification in the
fore arm in the present form, as well as in the carpus, are not displaced elements but
new ones without homologies.”
Discussion OF THE GENUS BAPTANODON.
In the year 1879 Professor O. C. Marsh proposed and established the genus
Sawranodon * which being preoccupied was later replaced by the name Baptanodon.”
Since that time all Ichthyosaurian remains from the American Jurassic have been
referred to that genus. In his original description Marsh distinguishes Sawranodon
natans (the type species of the genus) from Ichthyosaurus by the absence of teeth,
of which he says: “The jaws appear entirely edentulous and destitute even of a
dentary groove.” ‘This statement is corroborated in a second paper * in the following
lines: ‘Since the first species of the present genus was described by the writer, eight
other specimens of the same group have been discovered and are now in the Yale
Museum. In three of these the skull is preserved, but there are still no indications
of teeth, so that we may consider these reptiles as entirely edentulous.”
_ While cleaning the skull pertaining to specimen No. 603 preparatory to study
the discovery was made that the animal had teeth,” two of which were found
45 Williston, S. W., ‘‘ North Amer. Plesiosaurs,’’ Part I., Field Columbian Museum, Pub. 73, Geol. Series, Vol. II.,
No. 1, 1903, p. 70.
4° Marsh, O. C., ‘‘ A New Order of Extinct Reptiles (Sauranodonta) from the Jurassic Formation of the Rocky
Mountains,”’ Am. Jour. of Sci. (3), Vol. XVII.
“7 Marsh, O. C., “ Note on Sauranodon,’’ Amer. Jour. of Sci. (3), Vol. XIX., 1880.
*© Marsh, O. C., ‘‘ Limbs of Sauranodon, with Notice of a New Species,’? Amer. Jour. of Sci. (3), Vol. XIX.
Feb., 1880.
“Gilmore, C. W., ‘‘ Discovery of Teeth in Baptanodon, an Ichthyosaurian from the Jurassic of Wyoming,”’
Science, N. S., Vol. XVI., Dec. 5, 1902.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) ntblG
between the jaws near the end of the snout. Subsequent to this first discovery the
jaws were partially separated from one another and seven more teeth were found
scattered at intervals along and between the mandibles. Though no teeth were
found in the dental grooves of the second specimen No. 878 rudimentary alveolar
partitions (see P]. VIII.) were present near the front together with a few faint
sockets at the back of the rostrum.
The finding of teeth in one specimen (No. 603) and evidence of their existence
at one time in a second individual (No. 878) led me to believe that dental grooves,
if not teeth, would be found in the type of the genus Baptanodon if that specimen
were thoroughly prepared. Only a little preparation was necessary to demonstrate
the existence of well-developed dental grooves on both upper and lower jaws in
both types, and just outside of the dental grooves imbedded in the matrix surround-
ing the rostrum of No. 1952” (B. natans, type of the genus) a small tooth” was
found.
The finding of dental grooves and teeth in the types as well as their existence in
two individuals preserved in the collections of this museum clearly demonstrates the
fact that American Ichthyopterygians of the Jurassic possessed teeth and were not
edentulous as originally described by Professor Marsh. Thus the one important
generic character which has for so long a time separated the American form from
Ichthyosawrus, and the closely allied European genus Ophthalmosawrus, has been
shown to be an erroneous determination.
Among the other distinguishing characters ascribed to the type B. (Sawranodon)
natans by Marsh, are the number and position of the sclerotic plates of the eye.
He says: ‘The sclerotic ring is composed of eight plates. . . . These plates are not
arranged in a nearly flat ring, as in Ichthyosawrus but form the basal segment of an
elongated cone, as in the eyes of some birds.” The badly crushed condition of the
skull of No. 1952 makes it impossible for any one to determine with any degree of
accuracy the precise number of plates composing the sclerotic circle. The right
orbit, which is the better preserved, contains only two plates with the impressions
of three others, the intervening space in the ring between the plates and impressions
being filled with matrix. The “cone-like”’ position of these plates is unmistakably
produced by crushing. The right orbit of No. 603 shows the sclerotic ring as hay-
ing been subjected to pressure antero-posteriorly, and here we find the plates assum-
ing the cone-like arrangement, though it is not so exaggerated as in the former case.
59 Catalogue number of the Yale Museum.
51 Gilmore, C. W., ‘‘ Discovery of Dental Grooves and Teeth in the Type of Baptanodon (Sauranodon) Marsh,”’
Science, N.S., XVII., No. 436, May 8, 1903, p. 750.
52 Type of the species, B. discus.
118 MEMOIRS OF THE CARNEGIE MUSEUM
The left orbit of the same specimen has the ring nearly circular (see PI. X., fig. 2)
and the plates quite flat, though as Frass has pointed out in life they were probably
arranged at somewhat more of an angle. It might be well to mention here that the
sclerotic ring in the skull of 1955” (B. discus) shows the same flat circular arrange-
ment of the plates observed among most of the members of the Ichthyosauria.
The specific characters given by Marsh for separating the two species of Baptano-
don are quite as superficial as the generic characters just reviewed. Size alone is the
only difference of importance, but as in most reptiles these probably continued to
grow throughout life. ‘The breadth of paddles,” “ elongation of the facial portion
of the skull,” “slender snout,” etc., are illusionary characters and would not serve
to distinguish the two species. For example in giving the distinguishing characters
between B. discus and B. natans (type of the genus) Marsh says: ‘The paddles, also
are broader in proportion to their size, than in the type species.” It is now definitely
understood that the extremities of B. natans areunknown. Howeverin the Catalogue
of Fossil Reptilia and Amphibia in the British Museum, fig. 5, p. 7, is a figure of
the left pelvic limb of B. natans (after Marsh from the Proc. Geol. Society). The
illustration shows the dorsal surface of the limb and is to all intents the reverse
view of the left pelvic (?) limb of B. discus and accidentally referred to the wrong
species.
In a more recent paper™ the late Dr. W. C. Knight has added much to our knowl-
edge of Baptanodon besides giving reasons for considering this genus distinct from
Ophthalmosaurus.
To correct some inaccuracies in the characters enumerated I will take up the
arguments advanced by him and make such comments as access to literature and
more complete and better preserved material render possible.
They are as follows:
“In comparing the limbs of 1. The absolute length of limb in
Ophthalmosaurus and Baptanodon one either form apparently unknown, but
should consider the following points : in so far as one may judge from illus-
1. In Baptanodon the humerus is trations the limbs of the two forms ap-
about one third the length of the limb.” pear to have about the same propor-
tions. Lydekker in the catalogue of
Fossil Reptilia and Amphibia in the
British Museum gives measurements of
53 Nicholoson and Lydekker have the same illustration in their ‘‘ Manual of Paleontology,’’ Vol. II., Fig. 1034, and
ascribed to the same species, though (after Marsh and Hulke).
54 Knight, W.C., ‘‘Some Notes on the Genus Baptanodon with a Description of a New Species,’’ Amer. Jour. Sci. (4)
Vol. XV.
GILMORE: OSTEOLOGY OF
2. “It has a twisted shaft which is
greatly compressed.”
3. The distal facets are all unequal
in size and one of them is merely rudi-
mentary, besides they are elliptical in
the plane of articulation.”
4. “There is also an abnormal
number of digits and the arm is much
more powerful and larger than found
in Ophthalmosaurus of equal size.”
5. “In comparing Baptanodon with
Ophthalmosawrus it will be well to con-
sider that in Baptanodon the interclay-
icle is either rudimentary or wanting.”
6. “The absence of the intercentra
between the second and third vertebree.”’
BAPTANODON (MARSH) 119
the humerus of O. icenicus which agree
essentially with the measurements of
the American forms.
2. A humerus pertaining to a spec-
men of Q. icenicus preserved in the
collections of the American Museum of
Natural History, shows the same char-
acters.
3. The humerus of O. icenicus has
three unequal facets on the distal end,
though the one for the pisiform could
hardly be considered as rudimentary.
The facets on the distal end of the
humerus mentioned above are elliptical
in the plane of articulation. — Dr.
Knight must have drawn his conclu-
sions from a comparison of the humerus
of B. marshi, with the humerus of 0.
cantabrigiensis a second species described
by Lydekker, which has three subequal
facets, the longer axis being vertical to
the plane of articulation. The former
species (QO. icenicus) however is the type
of the genus.
4. So far as known Ophthalmosau-
rus only has five digits and as Knight
has observed the limb of Baptanodon
was probably more powerful to the ex-
tent of being broader, though the other
proportions appear remarkably similar
5. It is now positively known that
Baptanodon has a well-developed inter-
clavicle. See description, this paper, p.
36.
6. The absence of this bone in
other species of Baptanodon besides B.
marshi has yet to be demonstrated, for
120 MEMOIRS OF THE CARNEGIE MUSEUM
it has already been shown that two
specimens in this museum, Nos. 603 and
878, have a surface on the infero-pos-
terior margin of the axis as if for the
articulation of such a bone. However
this element has never been found in
place.
7. “The development of large 7. The coracoids of O. icenicus ex-
facets upon the interior margins of the amined by me at the American Museum
coracoids.” of Natural History, New York, N. Y.,
show well-developed surfaces for artic-
ulation with one another medially.
Combining the characters given by Marsh and Knight with those shown by the
material under discussion the genus Paptanodon may now be distinguished by the
| following characters :
Gen. char.: Teeth present but loosely attached. Dentition somewhat reduced but ea-
tending the entire length of the jaws. Opisthotic reaching and partially enclosed by the
squamosal. Reduced number of hypocentra. Vertebral centra forward of the posterior
caudals uniformly biconcave. Ribs of the anterior part of the skeleton double-headed.
Clavicles firmly united. Well developed interclavicle present. Coracoids without pos-
terior notch and wniting im the median line by large elliptical facets. Humerus with
strongly developed trochanteric ridge on dorsal surface and having three wnequal facets
on the distal extrenuty, which articulate with three irregular polygonal elements in the
epipodial row. All remaining bones of the anterior extremities more or less rownded
and retained in persistent cartilage.
Three species of this genus have been described, of which a brief review is given
in the following pages.
Bapranopon NATANS, Marsh.
Sauranodon natans, Marsh, O. C., Amer. Jour. of Sci., Vol. X VII., 1879.
Baptanodon natans, Marsh, O. C., Amer. Jour. of Sci., Vol. XIX., 1880.
Ophthalmosauwrus natans, Lydekker, R., Geol. Mag., Vol. V., 1888.
Baptanodon natans, Lydekker, R., Cat. of Fossil Reptilia and Amphibia in the British
Museum, Part II., 1889.
This species, the type of the genus, was based upon a considerable portion of a
skull, a number of vertebree and ribs, and parts of one or more limb bones. As has
been shown previously hardly any of the original characters given for this species
can be considered valid.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 121
B. natans is typically the smallest species of this group. In fact until the type
specimen is properly prepared and redescribed this is the only way by which B.
discus may be distinguished from this form. The teeth when better known may
show specific differences. The one tooth discovered of B. natans has «a smooth
enameled surface as contrasted with striated surfaces of the teeth of B. discus.
The type No. 1952 was collected by Mr. W. H. Reed, in the vicinity of Como
Bluff, Albany Co., Wyoming, and is now preserved in the collections of the Yale
Museum, New Haven, Conn.
Bapranopon piscus, Marsh.
Sauranodon discus, Marsh, O. C., Am. Jour. of Sci., Vol. XIX., 1880.
Baptanodon discus, Marsh, O. C., Am. Jour. of Sci., Vol. XIX., 1880.
Ophthalmosaurus discus, Lydekker, R., Geol. Mag., Vol. V., 1888.
Baptanodon discus, Lydekker, R., Cat. of Fossil Reptilia and Amphibia in the British
Museum, Part II., 1889.
Microdontosaurus petersonii, Gilmore, C. W., Science, N. S., Vol. XVI., 1902.
This species was based upon a portion of a poorly preserved skull, numerous
vertebree including the coalesced atlas and axis, left coracoid, with the greater portion
of the left pelvic (?) paddle and numerous fragments of other portions of the skeleton.
It is with considerable trepidation that I assign the specimens considered in this
paper to the species discus, but after a somewhat superficial examination of the type
I was unable to find differences of sufficient importance to warrant the establishment
of a new species as was proposed in a previous paper. The study of the type was
superficial to the extent of trying to determine characters from material that has
been but little prepared. The parts of the skull exposed only show a small area
of the mid-portion of the rostrum and one sclerotic ring; the other parts are either
wanting or covered by the very refractory matrix. The characters shown by the
parts of the skull exposed together with the atlas and axis and left coracoid are the
principal elements upon which my comparisons were based. The great similarity
of these elements to the corresponding parts of oar specimens prompted me to the
present determination.
The type specimen No. 1955 is preserved in the vertebrate collections: of the
Yale Museum, New Haven, Conn. It was collected by Mr. W. H. Reed in
Wyoming.
3APTANODON MARSHI, Knight.
Baptanodon marshi, Knight, W. C., Amer. Jour. of Sei. (4), Vol. XV., 1903.
In the characterization of this species Dr. Knight distinguished it from all
others by the shape and arrangement of the bones of the fore paddle. The two
122 ' MEMOIRS OF THE CARNEGIE MUSEUM
important differences which he considered as typical are the consolidation of three
elements into one in the third segment and the development of the abnormal num-
ber of digits by a division of the third finger. Accompanying the description is a
drawing of the right pectoral limb showing the paddle as retained in the matrix.
A photograph of the same paddle is shown in fig. 23.
The paddle probably had six digits but the greatest transverse segmentation of
Fia. 26. Atlas, axis and third and fourth cervicles of Baptanodon marshi (‘‘S”’). Seen from the right side.
Type specimen. One half natural size. 1. Coalesced atlas and axis. 2and3. Third and fourth cervicles. at, atlas;
ax, axis ; d, diapophysis ; p, parapophysis ; n, paired neural arch.
the parts preserved only shows five elements, although in the original description
he says: “The limb that I have been studying and figured differs from the one
published by Marsh, inasmuch as the abnormal number of digits do not appear
until the phalanges are reached.” As the specimen does not show an abnormal
pumber of digits it would be assuming considerable, because of the peculiar arrange-
ment of the bones of the paddle in this form, to say definitely at what point the
extra digit, if it did exist, is developed. Moreover it does not appear justifiable to
compare this fore limb with the supposed hind paddle of B. discws and consider as
specific characters differences that are almost certain to exist in the structure of
anterior and posterior paddles.
After an examination of the type I should consider the large element (see Fig.
23) as due to a consolidation of two rather than three elements. An offset on the
distal border apparently indicates the point of union. This consolidation, instead
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 125
of being a normal character, may be due to pathological conditions as has been
shown to exist in the extremities of the Mososaurs.”
The species was based upon a considerable portion of the pectoral girdle, with a
part of the right fore limb and 41 consecutive vertebre extending from the atlas
posteriorly.
A comparison of the coalesced atlas and axis of B. marshi, with the same ele-
ments of B. discus appears to show differences that would serve for specific separa-
tion, 2. e., the reduced number of intercentra, shown by the absence of the apophy-
sis on the axis for the third incentrum, and the reduced size of the parapophyses of
the third and fourth cervicles. These were certainly not functional in this species.
Some other minor differences will be noted by comparing Figs. 10 and 26.
With these exceptions and the arrangement of the bones of the paddle the other
parts preserved do not differ materially from the corresponding elements of B. discus.
The type is specimen “S”’ preserved in the Jurassic collections of vertebrates of
the University of Wyoming, at Laramie, Wyoming. This specimen was collected
by Mr. W. H. Reed from the Baptanodon Beds of the Jurassic in the Northern part.
of Albany Co., Wyoming, in 1897.
DiscussIoN OF THE GENUS OPHTHALMOSAURUS.
In studying the genus Baptanodon it was necessary to review the literature per-
taining to the closely allied genus Ophthalmosaurus. Having this at hand and
because of the great similarity of the two forms it is believed that a brief review of
that genus would be appropriate at this time.
The type species is O. icenicus founded by Seeley” on the greater portion of the
pectoral girdle. |The important characters by which he distinguishes this genus
from Ichthyosawrus are the peculiar conditions found in the clavicular girdle, 7. ¢.,
the separated clavicles, the enclosing of the anterior part of the interclavicle in a
groove of the clavicles, and the embracing of the anterior margins of the coracoids
by the clavicles.
Further characters were made known by the description of a portion of the fore
limb of a second individual which is larger and better preserved than the corres-
ponding elements of the type. These differences are as follows: distal end of hu-
merus articulating with three bones, radius, ulna and olecranon (pisiform of modern
nomenclature) and the carpus with a row of four elements.
The characters cited above constitute all of the essential differences given. The
somewhat remarkable supposition that the clavicles encircled the coracoids is due
55 Williston, S. W., ‘‘ Univ. Geol. Survey of Kansas,’’ Vol. IV., PartI., p. 244.
56Seeley, H. G., Quart. Jour. Geol. Society of London, Vol. XXX.
124 MEMOIRS OF THE CARNEGIE MUSEUM
to Professor Seeley’s incorrect determination of the borders of the coracoid, as has
already been indicated by Lydekker,” “the one marked intercoracoidal being really
the glenoid cavity.” This correction is substantiated by the material under consid-
eration in this paper, and the clavicles instead of enclosing the coracoids (made im-
possible by the articulation between coracoid and scapula), curved up in front of the
scapulee as in Ichthyosawrus.
In 1888 in the “Catalogue of Fossil Reptilia and Amphibia in the British Mu-
seum,” Lydekker gave the following characterization of the genus :
“Teeth present but apparently small, and perhaps confined to the anterior por-
tions of the jaws. Humerus and femur with strongly developed trochanteric ridge
on dorsal surface, and articulating distally with three bones, as in Baptanodon, which
are of irregular polygonal contour. ‘The ulna being pentagonal. Clavicles (typically)
separate, with the interclavicle wedged in between them. Vertebral centra of the
general type of those of Campyldont subgroup of Ichthyosawrus. Coracoid without
posterior notch. Humerus and femur apparently (except at distal end) of the gen-
eral type of those of I. campylodon and relatively large in proportion to the vertebre.
In Baptanodon and probably also in this genus both pectoral and pelvic limbs rela-
tively wide (or that arising from the intermedium) contains two such rows, with the
consequent presence of two centralia.”
In 1898 Woodward® confined most of the observations made by Seeley and
Lydekker, and added important information concerning the pelvic region, 7. e.,
“The pubis and ischium are fused together, leaving a small obturator foramen.”
Besides, he figures a right pectoral limb which shows that member as being com-
posed of the normal number of five digits.
Combining the characters given by the different authorities the genus Ophthal-
mosaurus may be distinguished by the following :
Gen. char.: Teeth present but apparently small and perhaps confined to the anterior
portions of the jaws. Clavicles (typically) separate with the interclavicle wedged in between
them. Coracoids without posterior notch. Humerus and femur with strongly developed
trochanteric ridge on dorsal surface and articulating distally with three bones, which are
of irregular polygonal contour. Ulnais pentagonal. Pubis and ischiwm fused together,
leaving a small obturator foramen,
Two species of this genus have been described of which a brief review is given
The type specimens are from the Jurassic and Lower Cretaceous of England.
5 Lydekker, R., ‘‘ Cat. of Fossil Reptilia and Amphibia in the British Museum.’’ Part II.
5 Woodward, A. S., “‘ Vertebrate Paleontology,’’ p. 183.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 125
OPHTHALMOSAURUS ICENICUS, Seeley.
Ophthalmosaurus icenicus, Seeley, H. G., Quart. Jour. of the Geol. Soc. of London.
Vol. XXX., 1874, pp. 696-707.
This, the type species of the genus, was based upon quite a complete pectoral
girdle and portion of a fore limb, the latter pertaining to a second individual. The
type was found in the Oxford clay by Mr. C. Leeds, who later submitted it to Pro-
fessor Seeley for description. The remains of four or five individuals preserved in
the collections of the British Museum, and referred to this species by Lydekker
have all been found in the Kimmeridge clays which is considered Upper Jurassic,
while the type is from Middle Jurassic.
The following specific characters constitute the essential differences as character-
ized by Lydekker in the Catalogue of Fossil Reptilia and Amphibia in the British
Museum.
Sp. char.: “Larger than the type of the following species. The post-axial facet
smaller than that of the radius; and the antero-posterior diameter of the proximal ex-
tremity of the humerus less than that of the distal extremity. In the cervical region the
cupping of the anterior face of the centrum confined to the central portion and surrounded
by a flattened periphery.
OPHTHALMOSAURUS CANTABRIGIENSIS, Lydekker.
Ophthalmosaurus cantabrigiensis, Lydekker, R., Geol. Mag., Decade III., Vol. V.,
July, 1888, No. 7, p. 309.
This species was first proposed by Lydekker in a note to the Geological Maga-
zine, the description appearing later in the Catalogue of the Fossil Reptilia and
Amphibia in the British Museum. The material upon which this species is based
consists of a right humerus from the Cambridge Greensand.
The equal size of the three distal facets of the humerus, and their greatest elonga-
tion being vertical to the plane of articulation at once distinguishes this species from
Baptanodon. Lydekker in his original description adds, ‘This species may belong
to Baptanodon.” In reality the type of the genus (O. icenicus) is more closely
allied to that genus.
The type, No. 43989, is preserved in the British Museum.
Sp. char.: “ Typically of small size. The three distal facets of the hwmerus nearly
equal in size, and the antero-posterior diameter of the proximal extremity of the same bone
exceeding that of the distal.”
126 MEMOIRS OF THE CARNEGIE MUSEUM
RELATIONSHIP AND CLASSIFICATION.
In the general outlines of its structure, Bbaptanodon appears very similar to the
European genus Ophthalmosawrus, although the true relationship of the two genera
has long been an enigma to scientists.
In 1888 Lydekker® considered Baptanodon a synonym of Ophthalmosawrus after
Baur’s™ suggestion, that : ‘“ Ophthalmosaurus, Seeley. Vielleicht nicht verschieden
von Baptanodon,” a view which he abandons later in the Catalogue of Fossil Rep-
tilia and Amphibia in the British Museum.
In 1898 Woodward in his Vertebrate Paleontology, p. 183, observes: ‘‘ Baptano-
don from the Jurassic of Wyoming is remarkably similar to the European fossil and
perhaps generically identical.”
A number of equally eminent American paleontologists have verbally expressed
themselves as believing the two genera synonymous.
The presence of firmly united clavicles without suture, and the uniform bicon-
cave cupping of the anterior cervicals, together with the development of a sixth
digit in Baptanodon are characters which appear to be of sufficient value to warrant
the distinct separation of the two genera.
In the reduced size and loose attachment of the teeth, the arrangement of the
bones of the anterior extremities, and the retention of the disk-like elements of the
paddles in persistent cartilage, these two forms are very similar.
The characters enumerated above indicate a high degree of aquatic specialization
and Baptanodon, I believe, should be considered the most specialized of the known
Ichthyopterygia. ‘The closestaffinities of Baptanodon are with Ophthalmosaurus, next
approached by Ichthyosawrus. In some respects Baptanodon is but little more modi-
fied to purely aquatic conditions than is found in the earlier genus Shastasawrus
from the Triassic of California. The humerus of Baptanodon is not so broad as that
of Shastasaurus but it shows greater specialization in the development of three un-
equal facets on the distal end.
The reduction in the number of intercentra in Baptanodon marshi, as well as in
Baptanodon discus, appears to indicate the most specialized character of the genus.
The separated zygapophysial facets in the cervical region, which become single in the
vertebrxe more posteriorly, are approximated in the Ichthyosaurs.
If Baur’s classification of the Ichthyopterygia be accepted the family Baptanodon-
tidee proposed by Marsh would include the two genera Baptanodon and Ophthalmo-
59Lydekker, &., Geol. Mag., Vol. I1., p. 309.
* Baur, G., ‘‘ Ueber den Ursprung der Extremititen der Ichthyopterygia,’’ Bericht der XX. Versammlung der Ober-
rhein. geol. Ver., Vol. XX.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 127
saurus. The family would be distinguished by the peculiar arrangement of the
bones of the paddles, and the reduction of the number of intercentra.
The classification of the genus Baptanodon would then be best expressed by the
following :
Class, Reptilia; Order, Ichthyosauria; Family, Baptanodontids ; Genus, Bap-
tanodon.
Baur, G.
Bauer, F.
Cork, E. D.
Frass, F.
GILMoRE, C. W.
Harcuer, J. B.
Hu.ks, J. W.
Kyieut, W. C.
Loomis, F. B.
LYDEKKER, R.
Marsu, O. C.
BIBLIOGRAPHY.
On the Morphology and Origin of the Ichthyopterygia, Amer. Naturalists, Vol.
XXI., 1887, pp. 837-840; Ueber den Ursprung der Extremititen der
Ichthyopterygia, Bericht der XX. Versammlung des Oberrhein geol. Ver.,
Vol. XX., 1887.
Osteologische Notizen itiber Ichthyosaurier, Anat. Anzeiger, Vol. XVIII.,
1885, pp. 574-588.
On the Homologies of Some of the Cranial Bones of the Reptilia and on the
Systematic Arrangement of the Class, Proc. Amer. Assoc. Adv. of Sci., Vol.
XIX., 1870, pp. 194-247; A New Genus of Ichthyoptergyia, Amer-
Naturalist, Vol. XIII., 1879, p. 271.
Die Ichthyosaurier der Sitiddeutschen Trias und Jura Ablagerungen. Tiibin-
gen, 1891.
Discovery of Teeth in Baptanodon, an Ichthyosaurian from the Jurassic of
Wyoming, Science, N. S., Vol. XVI., 1902, No. 414, pp. 913-915 ; Dis-
covery of Dental Grooves and Teeth in the Type of Baptanodon (Saura-
nodon) Marsh, Science, N. S., Vol. XVIT., 1903, No. 436, p. 750.
Osteology of Haplocanthosaurus, Memoirs Carnegie Museum, Vol. II., 1903,
No. 1, p. 70-71.
Anniversary Address of the President Geol. Soc. of London, Quart. Jour.,
Vol. XXXIX., 1883, pp. 44-64.
Jurassic Rocks of Southeastern Wyoming, Bulletin of the Geol. Soc. of Aim.,
Vol. II., 1900, pp. 377-888 ; Some Notes on the Genus Baptanodon, with
a Description of a New Species. Am. Jour. of Science (4), Vol. XV., 1903.
On Jurassic Stratigraphy in Southeastern Wyoming, Bull. of the Am. Museum
of Nat. History, Vol. XIV., article XII., 1901, pp. 189-197.
Note on the Classification of the Ichthyopterygia (with a notice of two new
species), G'eol. Mag., Decade III., Vol. V., No. 7, 1888, p. 309-314 ; Cata-
logue of the Fossil Reptilia and Amphibia in the British Museum, 1889.
A New Order of Extinct Reptiles (Sauranodonta) from the Jurassic Forma-
tion of the Rocky Mountains, Amer. Jour. of Sci. (3), Vol. XVII., 1879,
pp: 85-86 ; The Limbs of Sauranodon with Notice of a New Species, Amer.
128 MEMOIRS OF THE CARNEGIE MUSEUM
Jour. of Sci. (3), Vol. XTX., 1880, pp. 169-171; Note on Sauranodon,
Amer. Jour. of Sei. (3), Vol. XIX., 1880, p. 491; The Reptilia of the
Baptanodon Beds, Am. Jour. of Sci. (3), Vol. L., 1895, pp. 405-406 ; Verte-
brate Fossils of the Denver Basin, Monograph U.S. G.8., Vol. XX VIL,
1897.
Merriam, J.C. Triassic Ichthyopterygia from California and Nevada. Univ. of Cal. Publica-
tions, Vol. IIT., 1902, No. 4, pp. 68-198 ; New Ichthyosauria from Upper
Triassic of California, Univ. of. Cal. Publications, Vol. III., 1903, No. 12,
pp- 249-263.
NicHoLoson AND LyDEKKER. Manual of Paleontology, Vol. II., 1889.
Owen, R., Monograph of the Fossil Reptilia of the Liassic Formations. Owen’s British
Fossil Reptiles, Vol. VII., 1881, part third, pp. 83-134.
SEELEY, H. G. On the Pectoral Arch and Fore-Limb of Ophthalmosaurus, a new Ichthyo-
saurian genus from the Oxford Clay (read June 24,1874). Quart. Jour. of
the Geol. Soc. of London, Vol. XX X., 1874, pp. 696-707.
Wiuuuiston, S. W. North American Plesiosaurs. Field Columbian Museum Publications. No.
73, Geol. Series, Part I., Vol. I1., No. 1, 1908.
Puate VII.
Puate VIII.
PLATE IX.
PLATE X.
PuatTE XI.
Prate XII.
GILMORE: OSTEOLOGY OF BAPTANODON (MARSH) 129
EXPLANATION OF PLATES.
Diagram showing the position of the elements of specimen No. 878, Baptanodon
discus, as they lay imbedded in the concretion. The irregular line surrounding
the bones represents the outline of the concretion. at., atlas ; co., coracoid ; cl.,
clavicle ; h., humerus ; 7.cl., interclavicle ; ., paired neural arch of atlas ; 0.0ce.,
occipital condyle; 7., ribs; s., spinous process of the vertebre ; sc., scapula.
Side view of skull of Baptanodon discus, Marsh (No. 878). One fifth natural
2.
bo
bo
co
size. Restored. ag., angular; d., dentary;j., jugal; la., lachrymal ; ma., maxil-
lary ; na., nasal; nar., narial opening; occ.c., occipital condyle; pa., parietal ;
pmx., premaxillary ; prf., prefrontal (?); ptf, postfrontal; pto., postorbital ;
qj. quadrato-jugal; qg.u., quadrate; s.ag., surangular;_ s.t., supratemporal ;
sel., sclerotic plates; stu., stapes; spl., splenial ; sq., squamosal.
. Top view of same skull. One fifth natural size. Restored. avt., articular ;
ex.occ., exoccipital ; fr., frontal ; na., nasal; nar., nares ; occ.c., occipital con-
dyle; pa., parietal; pin., pineal foramen; pinz., premaxilla; prf., prefrontal (2) ;
ptf., postfrontal ; s.ag., surangular; s.occ., suraoccipital ; s.t., supratemporal ;
s.t.f., supratemporal fossa ; sq., squamosal.
Inferior view of skull of Baptanodon discus (No. 603). One fifth natural size.
Restored. ag., angular; b.oce., basioccipital ; b.s., basisphenoid ; d., dentary ;
ipt., interpterygoid vacuity ; occ.c., occipital condyle; p/., palatine; prs., pre-
sphenoid ; pé., pterygoid; spl., splenial; v., vomer. The suture between ag.
and spl. and cor., is incorrectly placed posteriorly.
. Side view of skull of Baptanodon discus, Marsh (No. 878). One fifth natural
size. Seen from right side.
. Side view of skull of Baptanodon discus, Marsh (No. 603). One fifth natural
size. Seen from left side.
. Posterior view of skull of Baptanodon discus (No. 878). th., thyrohyal.
. The same, restored. One fifth natural size. ag., angular; art., articular ;
b.occ., basioccipital ; b.s., basisphenoid ; cor. (?), coranoid ; ex.oce., exoccipital ;
J.m., foramen magnum ; occ.c., occipital condyle ; op.o., opisthotic ; pa., parietal ;
pt., pterygoid ; q.j., quadrato-jugal; qu., quadrate; s.ag., surangular ; s.oce.,
supraoccipital ; s.¢., supratemporal ; sta., stapes; s.t,f., supratemporal fossa ;
sq., squamosal.
. Anterior view of pectoral girdle of Baptanodon (No. 919). One fifth natural
size. co., coracoids ; sc., scapule ; ~., surfaces for clayicles.
. Dorsal view of the same girdle. One fifth natural size. Lettering same as 1.
. Ventral view of pectoral girdle Baptanodon discus (No. 878). One fifth natural
size. Restored. cl., clavicle ; 7.cl., interclavicle ; h., humerus. Other letter-
ing same as figures 1 and 2.
MEMOIRS CARNEGIE MUSEUM, VOL ILI. PLATE VII.
DIAGRAM SHOWING THE POSITION IN WHICH THE ELEMENTS OF THE SPECIMEN OF BAPTANODON DISCUS Marsu
(No. 878) were Founp ImsBeppEep IN THE ConcrETION. + NATURAL SIZE.
6 “7 a , - ; g ofiane 4 ares
GayOLsayy “AZIS IWYALVN F (S18 ON) HSUV] SOOSTT NOGONV Ld FG 10 110g ao MATA Wag
g gE
/ ce g=_
YY
ALY,
IIA d ‘Il “TOA NNASNW YISENYVD SYIOWAW
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= til yy ———— —
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SAOSTG NOGONVId¥g 40 stTaxg
“YX FLVIg
‘Il “1OA ‘WO3SNW AION SHIOWaW
‘dad yOLsaY ‘G /ANOOY SV NUWIONAG ‘T ‘“AZIG, WUOLVN F (S18 ON) ‘SQOSTG NOUONFId¥g 40 TIONG AHL AO SMATA MOIUMISOg
My
x. op
WS
aii
Uff
N COT
8 :
yy
Y
lll
wy
Wy
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i WE
i
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TX aLvig TI “I0A ‘WNASN\ FIOANYVD SYIONS
MEMOIRS CARNEGIE MUSEUM, VOL, II. PLATE XII.
NN
SSW
: :
PrcroraL GIRDLE or BAPTANODON DISCUS. 1, ANTERIOR ViEW; 2, DorsaL View (No. 919); 3, VentRaL View
(No. 878). An + Natorat Size.
a
MEMOTRS
OF THE
Cra NEGIE MOST wir
VOI, Mk eee NO. 3.
FOSSIL AVIAN REMAINS FROM ARMISSAN.
By C. R. Hasrman.
The history of gallinaceous birds is traceable as far back as the Middle Eocene,
the earliest known genus being found in the Green River limestone of Wyoming.
This is known by a single, nearly perfect skeleton, and seems to be closely related
to the existing Ortalis. From the Upper Eocene of the Paris Basin have been de-
scribed two species of Palxortyx, which comprise small pheasant-like birds, with the
rostrum longer and less curved than in modern forms. Certain fragments occurring
in the Querey Phosphorites (Upper Eocene), and the calcareous marl of Vaucluse,
supposed to be of equivalent age, are likewise referred to Palxortyx, and several
species are also known from the Miocene of Southern France. The remarkable
and highly specialized Hoactzin, or Opisthocomus, of South America, whose habits
and certain points of structure suggest considerable affinity with the Cucwli, is rep-
resented by a closely similar genus (Fi/holornis) in the Phosphorites of Quercy.
The fresh-water limestone of Armissan, near Narbonne (Aude), by some geolo-
gists regarded as of Upper Eocene, by others as of Oligocene age, has furnished re-
mains of a single genus (Twoperdix), which agrees somewhat closely with modern
pheasants ; and Phasianuws itself occurs rather abundantly in the Middle and Upper
Miocene, and also in the Lower Pliocene of Pikermi, where it is accompanied by
Gallus. A species of fossil turkey (Meleagris antiquus) is known from the White
River Oligocene of Colorado, the same genus being also represented in the Pleisto-
cene of New Jersey, Pennsylvania and probably elsewhere in this country. Under
the name of Palxoperdix, three species of fossil pheasants have been described from
the Middle Miocene of Sansan (Gers). Gallus and Phasianus appear to be the only
known Pliocene genera, but from the Pleistocene and cavern deposits of various
132 MEMOIRS OF THE CARNEGIE MUSEUM
parts of the world a considerable number of gallinaceous birds have been brought
to light, the more important of which are enumerated in the following list.
TABLE SHowING GEOLOGICAL DisTRIBUTION OF GALLINACEOUS Brrps.
Middle Eocene.
Gallinuloides wyomingensis Eastman. Green River limestone; Uinta County,
Wyoming.
Upper Eocene.
Palxortyx hoffmanni (Gervais). Gypsum of Paris Basin, Montmartre.
s! blanchardi Milne-Edwards. Gypsum of Paris Basin, Montmartre.
ys sp. indet. Gypsum of Paris Basin, Montmartre.
FKilholornis paradoxa Milne-Edwards. Quercy Phosphorites.
‘ gravis Sele ges: e es
S debilis of i of es
Tuoperdix pessieti (Gervais). Lacustrine limestone ; Armissan (Aude).
3 (73 (73 ce
fs keltica Hastman.
Oligocene.
Meleagris antiquus Marsh. White River Beds ; Colorado.
Lower Miocene.
Palxortyx cayluxensis Lydekker. Phosphorites of Caylux (‘Tarn-et-Garonne).
s gallica Milne-Edwards. Lacustrine deposits of Allier, and Phosphorites
of Caylux.
as brevipes Milne-Edwards. Lacustrine deposits of Allier.
“ (2?) phasianoides “ is “ ;
«media 4 ee ‘““ (descrip. insuf.).
Middle Miocene. zZ
Palxortyx edwardsi Depéret. Saint-Alban-de-Roche (Isére).
Palxoperdix longipes Milne-Edwards. Sansan (Gers).
i prisca a ‘ 8
cc (?) sansamiensis “ ee 3
Phasianus alts of s ‘ and Saint-Alban-de-Roche (Isére).
medius es Sansan (Gers).
- desnoyerst a Orleannais.
Upper Miocene.
Phasianus altus Milne-Edwards. Lacustrine deposits ; Giningen, Switzerland.
EASTMAN: FOSSIL AVIAN REMAINS FROM ARMISSAN 133
Lower Pliocene.
Phasianus archiaci Gaudry. Pikermi (Attica).
e “ec (a9
Gallus xsculapii
Upper Pliocene.
Gallus bravardi Gervais. Ardé, near Issoire (Puy-de-Dome).
Pleistocene.
Tetrao tetrix Linn. Kent’s Hole Tavern, near Torquay, Devonshire.
“ —wrogallus Linn. Forestbed (?) of Norfolk.
Lagopus albus (Gmelin). Cavern desposits near Montauban (Tarn-et-Garonne).
“ — mutus (Montin). oo co ‘e ie my
Francolinus pictus (Jardine and Selby). Cavern deposits; Madras, India.
Coturnix novx-zealandix Quoy and Gaimard. Superficial deposits; New Zealand.
Tympanuchus pallidicinctus Ridgway. ie ot Oregon.
Pediocextes phasianellus Linn. “ as “e
“ nanus Shufeldt.
cc oe ce “co
lucasii
Polxotetrix gilli as “ fi a
Meleagris superbus Cope. i re New Jersey.
eg celer Marsh. te as i
‘sp. indet. Caves of Pennsylvania.
Gallus sp. Rare in Kuropean bone caverns.
IPG RIO ae i: “s
It will be seen from the foregoing that although gallinaceous birds are plentiful
in the late Tertiary, with the exception of the Pliocene, our knowledge of them in
the Eocene is confined to but four genera. Two of these, futhermore, are known
by a solitary individual each; and as Eocene bird remains are seldom well pre-
served, it is clear that the discovery of fairly complete skeletons from this horizon
is a matter of considerable interest and importance. The Carnegie Museum is there-
fore to be congratulated upon having recently acquired two tolerable skeletons of
Kocene birds from the lacustrine limestone of Armissan, near Narbonne (Aude), the
existence of which has not hitherto been made known.
These specimens form part of the famous Bayet Collection, presented by Mr.
Andrew Carnegie to the Pittsburgh Museum in 1903; and through the unfailing
courtesy of Dr. W. J. Holland, Director of the Museum, the present writer has been
generously entrusted with their description. For this privilege he desires to express
here his sincere thanks.
134 MEMOIRS OF THE CARNEGIE MUSEUM
It has not been possible, however, to offer descriptions of both of these speci-
mens in the following pages, owing to the fact that only one of them (fortunately
the better preserved of the two) has not been sufficiently disengaged from the matrix
to permit of its extended study. The specimen about to be described bears the
Carnegie Museum catalogue number 2023, and belongs unquestionably to the genus
Taoperdix, founded by Milne-Edwards' upon a unique individual from Armissan,
which had been previously described by Gervais under the name of Tetrao pessieti.*
Although agreeing with the type in its general characters, the Carnegie example
differs from it in its greatly reduced length of wing, and in the disproportion of its
limb bones; hence it may properly be regarded as constituting a distinct species. It
may be fittingly designated as Taoperdix keltica, in commemoration of the name be-
stowed by Aristotle upon the inhabitants of the country near Narbonne.’
TAOPERDIX KELTICA, sp. nov. (Plates XIII.—XI1V.)
Founded upon the crushed skeleton of a bird having approximately the size of a
ruffed grouse (Bonasa wmbellus), and differing from the type species of Tuoperdix in
the relative proportions of its limb bones, especially its much reduced humerus ; also
with shorter mandible. Upper Eocene ; Armissan.
Although the skeleton is considerably dismembered and confused, both in the
type of this species and in that of 7. pessieti, as may be seen from a comparison of
the plates, this circumstance must be regarded as rather fortunate than otherwise,
since it permits of a more precise examination of the several parts. It is also fortu-
nate that these two type-specimens should supplement each other in important
respects. For our knowledge of the cranium we must depend solely upon the speci-
men belonging to the Carnegie Museum, although the mandible is present in both.
Most of the limb bones, too, are better preserved in the new than in the older known
species; but the latter, on the other hand, alone exhibits the furculum, sternum and
pelvis in satisfactory manner. In the accompanying restoration of T. keltica, parts
which are wanting or not clearly recognizable in the actual fossil are represented in
outline after analogy with the type species or with recent pheasants, as the case may
be. hat is to say, when an epiphysis or articular condyle has become crushed or
otherwise obliterated in the fossil before us, these parts are restored according to the
' Oiseaux Fossiles de Ja France, Vol. IT., p. 225, pl. 127, Paris, 1871.
2 Comptes Rendus, Vol. LIV. (1861), p. 896.
3 Prior to the Roman conquest of Gaul, the whole of that country bore the name of Keltica ; but after its occupation
by the Romans the southern provinces were distinguished from the rest of Keltica by conferring upon them the name
of Gallia Narbonensis. An interesting description of the Narbonnaise is given by Strabo in the fourth book of his Geog-
raphy, and further accounts of this region are to be found in the well-known History of Polybius. Both of these ancient
writers mention among the wonders of this country the occurrence of so-called ‘‘ subterranean fish ’’ or ‘‘dug mullets,”’
which the inhabitants obtained by digging to a depth of two or three feet in marshy districts.
EASTMAN: FOSSIL AVIAN REMAINS FROM ARMISSAN 135
usual conditions in modern Phasianidx. All of the bones shown in Plates 1 and 2
are precisely as they occur in nature.
Passing now from these general considerations to an account of the several parts
of the skeleton, we note that the head is of average size, rather abruptly truncated
behind, with a maximum width of 2.2 em. across the occiput, the orbits being rather
posteriorly situated. ‘The mandible is slender, without apparent downward curya-
ture in front, approximately 3 em. in length. The sternum and pelvis, not clearly
indicated in the Carnegie example, but well shown in the type species, are regarded
by Lydekker' as suggesting affinity with Nwmidia and Meleagris. Itis an open ques-
tion, however, whether the lateral aspect of the sternum has not become widened
through mechanical compression of the matrix; at any rate the published figure
creates a suspicion that such has been the case. A small portion only of the sacrum
is preserved in the Carnegie specimen, the furcula is wanting, and the coracoids not
distinctly reognizable. The last-named elements in 7. pessieti are stout and straight,
without subclavicular process, and with broad sternal facets. In both species the
scapula is long and sabre-shaped, without being expanded posteriorly.
Important points of comparison are furnished by the limb bones, which are
excellently preserved. It is evident that the species under discussion possessed
but feeble powers of flight, the humerus being notably shorter than in T. pessieti and
most living pheasants. This boneexhibits a slight double lateral curvature, and has
a head of moderate size. ‘The remaining wing bones are proportionally abbreviate,
the brachium being of about equal length with the humerus. Nearly all of the
bones of the manus in both wings are well preserved, and the same is true of the pes,
The femur is relatively stout, of medium length, and with a shght forward curva-
ture, the head being concealed. ‘The tibio-tarsus is a stout bone, its length having a
ratio of 1.3 as compared with the femur, and 1.7 as compared with the tarso-metatar-
sus. The latter element is without spurs, so far as indications show, and appears to
have been more or less flattened from front to back. The trochleze are of moderate
size and widely separated. The phalangeals are slender and of medium length.
The relative length of the principal limb bones in this and other species is ex-
hibited in this table, the tarso-metatarsus being taken as a standard of comparsion.
Species. | Tarso-metatarsus. Tibio-tarsus. Femur. | Humerus.
Muawer die peasielisccssatp jesse. | 100 | 174 | 130 | 143
“ ‘keltica | 100 | 172 133 | 121
Paleortyx hoffmanni 100 | 172 124 | 141
The table given below is intended to facilitate comparison of the actual measure-
ments of various bones of the skeleton in both species of Taoperdix. As there exists
1“ Catalogue of Fossil Birds in the British Museum,” p. 139 (London, 1891).
136 MEMOIRS OF THE CARNEGIE MUSEUM
in some instances a discrepancy between the measurements given in the text of
Milne-Edwards’ work and those indicated in the plates, we have arranged the data
obtained fron both sources in parallel columns.
TABLE OF COMPARATIVE MEASUREMENTS.
Taoperdix keltica. Taoperdix pessieti.
Name of Parts. Length. | Data from Text. Data from Figure.
Tarso-metatarsusS...............-.. 3.3 cm. 3.5 em. | 3.7 em.
Tibio-tarsus......... 5.7 6.2 6.3
Femur........... 4.4 4.6 | 5.0
Humerus... 3.7 5.0 5.2 and 4.8
Wilnareee-s odbo 3.6 4.8 | 4.8
TRENG TRIS cencanda0nee590000500 868600060 3.3 ( Wanting ) | —
Metacarpus..........-..2...2.--.-+- 2.0 2.5 | 2.5
Coracoid ...........2ceceecereceeneees (Incomplete) 3.4 3.4
Sca pull ateencrenessecessesceeieceesect eh 5.0
Mandiblettercscssntesssceeessece: 3.0 — 3.8
TORRE Hoccnnononosacacnncas5ce0ses0c8 ( Wanting) — 3.8
No indications of plumage accompany either of the Armissan skeletons, although
isolated feathers, and even egg-casts, are not uncommon in the lacustrine deposits
of the south of France and in Switzerland. For references to the literature concern-
in the Sep-
?
ing detached feathers, one may consult an article on “ Fossil Plumage,’
tember number of the American Naturalist for 1904.
The following brief notice concerning the nature of the deposits at Aix and
Armissan, together with their principal fossil contents, is sufficiently interesting to
be reproduced in its entirety. It is taken from the second volume (pp. 551, 552) of
Milne-Edwards’ important work on the “Fossil Birds of France” :
“Les marnes d’Aix (Bouches-du-Rhéne), si riches en restes de Reptiles, de
Poissons et d’Insectes, en empreintes de feuilles, ete., n’ont pas encore fourni d’osse-
ments d’Oiseaux, mais on y a trové des ceufs et des plumes admirablement conservés,
dont quelques échantillons font partie du musée de Marseille. Ces marnes parais-
sent s’étre déposées a la méme époque que le gypse des environs de Paris. En 1886,
M. Coquand y a trouvé des restes de Paléothérium, d’aprés lesquels il établit ce
parallélisme qui ne fut pas adopté par Dufrénoy, mais que la plupart des géologues
actuels s’accordent 4 admettre.
“(est probablement aussi 4 la méme époque que se sont déposés les calcaires
lacustres d’Armissan (Aude), dans lesquels M. P. Gervais a fait connaitre l’existence
@un Gallinacé, le Tetrao Pessieti, que j'ai étudié récemment et rangé dans un genre
nouveau intermédiaire entre les Perdrix et les Paons, et que j’ai [re-]nommé Tao-
perdix Pessieti. Ces caleaires, toujours accompagnés de gypse, de rognons de soufre
et de silex pyromaque, contiennent des débris de Paléothérium, d’Anoplothérium,
des Reptiles, des Poissons d’eau douce, des Insectes, et une riche flore de végétaux
qui paraissent avoir péri sous l’influence des causes qui ont agi 4 Aix.”
EASTMAN: FOSSIL AVIAN REMAINS FROM ARMISSAN 137
SUPPLEMENTARY NOorTkE.
Since the above description was set in type, the task of working out the second
bird skeleton (Cat. No. 2022), referred to above on page 134, has been skilfully per-
formed by Mr. O. A. Peterson, one of the preparators of the Carnegie Museum. In
its original condition many of the bones were only partially visible, and portions of
them had been injured by the process of sawing or grinding to which the slab had
been subjected prior to its acquisition by the Museum. As the result of Mr. Peter-
son’s manipulation all of the preserved portions have been clearly exposed, thus
permitting a far more accurate knowledge of details than could otherwise have been
obtained. Fortunate as is this circumstance, it renders all the more conspicuous the
fact that many characteristic portions, such as the articular extremities of several
of the limb-bones, have been irretrievably lost through sawing the slab in two.!
Notwithstanding the considerable injury done to the remains, they still furnish
us with an important and very precious example of Eocene bird remains. The ex-
treme rarity of complete individuals, not more than four or five being known from
the Armissan deposits, increases the value of every item of information that is
obtainable from any one of them. In the case of the present specimen, the loss of
the head deprives us of perhaps the most significant information, concerning which
we would have eagerly sought enlightenment. It remains for us to make as search-
ing an inquiry as is possible from a decapitated body.
Fortunately we are able to satisfy ourselves in regard to a number of particulars
with approximate accuracy. The conformation of the pelvis, typically gallinaceous,
and the form and proportions of the limb-bones, all point to a very close relation-
ship between this skeleton and the type species of Taoperdix. Indeed, it appears
impossible to doubt that the present specimen and the type of T. pessieti are specifi-
cally identical, the differences between them falling well within the limits of indi-
vidual variation. This conclusion is supported, in our judgment, by the following
table of measurements, in which it has not been considered necessary to repeat the
measurements of T. keltica, owing to their relatively greater disproportion. The
same characters which enable us to discriminate between that species and the type
compel us to distinguish between it and the second Carnegie skeleton now under
discussion. A protographic reproduction of the new specimen (No. 2022), of slightly
less than the natural size, is given in Plate XV. of this Memoir. In Plate XVI.
several characteristic bones are shown as they occur in nature, except that in the case
1 The specimen is evidently a piece of pavement slab, which was only discovered to contain fossil remains after it
had been sawn. Editor.
138 MEMOIRS OF THE CARNEGIE MUSEUM
of the limb-bones, some of the articular extremities that have been injured are here
partially restored.
TABLE OF MEASUREMENTS.
Carnegie | | Carnegie
Type Specimen of Taoperdix pessieti. seer | Type Specimen of Taoperdix pessieti. | Ex aus
Species. || | Species.
Ne pay, | Data from | Data from || GS PS |Datalfroml| atattroma |i mee
Name of Parts. | Text. Bieare Length. | Name of Parts. | Text. Figure. | Length
a oy ha Oe LEE Gees ool ant plies ae eee ke =; ee | Sateen aed | PRL SS ae
Tarso-metatarsus ...... 3.5 cm, 3.7 em. ADEs |ll) WHE coosbodas ecco co5ne. | 48cm. 4.8 | 5.6
Tibio-tarsus. ........ 6.2 6.3 6.9 | Metacarpus. a 2.5 2.5 | Be
Femur ........... 4.6 5.0 5.0 | Coracoid .... | 3. 3.4 | ase
Humerus 5.0 5.2 and 4.8 5.5» || Scapula 0.0.0.0... [Nees tall eo Obie eet -0-=
EXPLANATION OF PLATES.
Puate XIII. Taoperdix keltica, sp. nov. Photograph of the specimen in its natural condi-
tion, x 4.
PLatTeE XIV. Taoperdix keltica, sp. nov. Restoration of skeleton, missing parts represented in
outline, x 4.
PuatE XV. Tuoperdix pessieti (Gervais). Photograph of the second Carnegie specimen (Cat.
No, 2022) in its natural condition, x +.
Prate XVI. Tuoperdix pessieti (Gervais). Lllustrations of the more characteristic bones of the
skeleton (Cat. No. 2022), some of the articular extremities slightly
All figures of the natural size. 1, humerus; 2, radius and ulna;
restored.
3, meta-
carpus; 4, phalanges of pes; 5, tarso-metatarsus ; 6, pelvis; 7, coracoid; 8,
femur ; 9, tibro-tarsus ; 10, scapula.
MEMOIRS CARNEGIE MUSEUM, VOL. IL. PLATE XIII.
TAOPERDIX KELTICA EAstMAN. PHOTOGRAPH OF THE TYPE IN THE CarNeGre Musrum.
rh
ee Med teea
MEMOIRS CARNEGIE MUSEUM, VOL. II.
PLATE XIV.
TAOPERDIX KELTICA Easrman. Resrorarion Naturan
SIZE.
Tne Misstng Parts SupeLtED IN OUTLINE.
PLATE XV,
MEMOIRS CARNEGIE MUSEUM, VOL. II
It
11h Musrum.
nC
NI
2 CAR
xINAL IN THEI
Ort
Q
4
PHOTOGRAPH OF THE
RVATS )
D)
STETI (Gt
RDIX PEs
>
4
"AOP.
iG
MEMOIRS CARNEGIE MUSEUM, VOL. IL. PLATE XVI.
TAOPERDIX PESSIETI (GERVAIS).
MEMOLTRS
OF THE
CARNEGIE MUSEUM.
WOE ar NO. 4.
DESCRIPTION OF NEW RODENTS AND DISCUSSION OF THE
ORIGIN OF DASMONELIX.
By O. A. PErerson.
One of the field parties of the Carnegie Museum with the writer in charge was
detailed by the Director to collect fossils in western Nebraska and eastern Wyo-
ming during the season of 1904. In the collection which was secured are some very
complete remains of rodents, belonging to two new species of Steneofiber not pre-
viously described. This material fully substantiates the views foreshadowed by
Professor Scott, Dr. Matthew, and others, that this genus is clearly distinct from
Castor. In the following detailed description of this new material it will be seen
that there are some features, especially in the cranium, which are similar to those
in the genus Castor, but these do not necessarily imply relationship. The skeleton
as a whole shows osteological characters very similar to those of Aplodontia rufa
with which I have compared it. Cynomys ludovicianus is also used for comparison
in studying the present fossil forms.
The material was discovered in the Upper Miocene sandstones, or Harrison
(Demonelix) beds, in Sioux Co., northwestern Nebraska, and also in Converse Co.,
Wyoming, immediately across the Nebraska-Wyoming state line, and in the same
general locality. Fossils are comparatively rare in this horizon. The list of genera
found is as follows: Promerycochwrus, ? Mesoreodon, a Peccary, Oxydactylus, and
other small species of camels not identified. The commonest fossils from this
horizon are the new species of rodents described in the following pages.
I take pleasure in acknowledging the valuable assistance of Mr. Earl Douglass
1A specimen of Aplodontia rufa (Col. U. S. Dept. Agri., No. 77975) was kindly furnished for comparison by Dr. C.
H. Merriam, Smithsonian Institution, Washington, D. C.
140 MEMOIRS OF THE CARNEGIE MUSEUM
in reading and correcting the manuscript and making kindly criticisms, of Mr.
Sydney Prentice for making the drawings, of Mr. O. E. Jennings for working out
the details in microscopic sections made from Demonelix and furnishing notes on his
observations, of Mr. A. 8S. Coggeshall for photographs and of Mr. A. W. Vankirk for
making the microscopic slides and assisting in cleaning up the material. To Direc-
tor W. J. Holland is due special acknowledgment for the privilege of describing
this interesting and complete material.
Steneofiber fossor spec. nov.
This species is quite common in the Harrison (Demonelix) beds in eastern Wy-
oming and western Nebraska. This horizon immediately overlies the Monroe
Creek beds and is probably equivalent to the upper John Day. There are fourteen
specimens of this species in the Carnegie Museum collections, which were found
inside of as many Demonelices. In some cases nearly complete skeletons were
discovered. The type (No. 1217) was found near the rounded end of a “rhizome,”
and consists of the cranium, the vertebral column, including eleven caudals, (the
extreme tip of the tail is missing), both clavicles, and a fairly complete set of ribs.
The right and left fore limbs are complete, except the superior part of the right
scapula and the feet. The pelvis and the posterior limbs are present, including the
greater part of the right pes. No. 1208 is used as a cotype, and has the skull, lower
jaws, eleven presacral vertebrae, the sacrum, both clavicles, the manubrium, and
several ribs. ‘The fore limbs are fairly well represented, and there are some frag-
ments of the hind limbs. In the following description of the osteology of S. fossor
use will be made of supplementary material, when, by so doing, additional light is
thrown on the subject. In each instance the museum catalogue numbers will be
referred to in connection with the specimens.
THE Supertor Dentirion.
The superior incisors are comparatively as large and strong as those of the recent
beaver; anteriorly the broad and flat surface is covered with a heavy coating of
enamel, which forms rather sharp angular edges laterally. The rounded lateral and
posterior faces have no enamel. The antero-posterior diameter is as great as the
transverse. The chisel-shaped gnawing portions of the teeth are long and terminate
anteriorly in a broad sharp edge.
The superior grinding teeth are rather small in comparison with the size of the
skull. They gradually decrease in size from p* to m?. Their position is nearly
parallel with the long axis of the skull, and they are placed about midway between
the anterior and posterior extremities of the skull.
PETERSON : DESCRIPTION OF NEW RODENTS 141
P+ is the only superior check-tooth that retains the internal enamel-fold after
the maturity of the animal. All the other teeth seem to have lost this character
early, the surrounding enamel-bands showing no folding.
In the cotype (PI. XVIL., Fig. 3) there is a slight dissimilarity in the enamel pat-
tern of the crowns of the premolars on opposite sides. The anterior fossette of the
right premolar is oblong, and is the continuation of the internal enamel-fold, while
on the tooth on the left side this fossette is smaller and rounded, and the anterior
island is apparently independent of the internal enamel-fold. On the external
enamel-band there remains a slight indication of the folding, which is shown only
in the cotype in our series of skulls. There is also a small rounded fossette near the
postero-external angle on both premolars, a character not found in the type.
Molar + is broader than long in the cotype, while in the type its diameter is
nearly equal in both directions giving it a subcylindrical form. There are three
fossettes on the grinding face, one antero-external and transverse, the other antero-
a
Le
A)) d
m3 m2 mi p#
Fic. 1. Crown view of superior grinders of Steneofiber Fia. 2. Crown view of inferior grinders of Stencofiber
fossor representing different stages of wear. a, type, No. fossor. a, type, No. 1217; b, cotype, No. 1208; ¢, an
1217 ; b, cotype, No. 1208; ¢, an old individual, No. 1207. individual showing p; im a nearly unworn stage ; d, in
Figures natural size. ternal view of same specimen as c, No. 453. Figures nat-
ural size.
internal and oblique, while the third isa large fossette occupying an oblique posi-
tion on the postero-external part of the tooth.
Molars 2 and 3. So far as the numbers and positions of the fossettes on
molars 2 and * are concerned they are like the premolars. The last molar in
the type is smaller than in any of the other skulls in the Carnegie Museum series.
THE Inrertor Dentirron.
The inferior incisor is as large and as strong as the superior, and is similar to
that of the recent beaver, excepting that in our fossil form, the anterior face of the
tooth is flat. This is characteristic throughout the entire series. The left premolar
is injured in the type, but the corresponding tooth on the right side is complete
The anterior fossette is still connected with the enamel band, while that on the oppo-
site tooth is free. This tooth (p,;) has the three transverse fossettes and the external
142 MEMOIRS OF THE CARNEGIE MUSEUM
folding of the enamel usually seen in this genus. In S. fossor the folding of the
enamel is somewhat similar in position, although not so deep as that in S. montanus.
In S. fossor the tooth has a slightly greater antero-posterior diameter than m,. It
has rounded anterior and posterior faces.
Molar 1.— With exception of the less rounded anterior face, and the deeper and
more oblique external enamel-folding, this tooth is like the one in advance of it.
In width and length the tooth has the same diameter.
Molar 2.—'This tooth is similar to My with a slightly less transvere diameter.
On the tooth in the left ramus the posterior fossette is divided in two portions,
while that of the opposite tooth is undivided.
Molar 3. —'The characters of the third molar are essentially those of the preced-
ing molars 1 and 2. The tooth is smaller than those in advance of it. The teeth
are quite prismatic, and terminate in short peg-like roots.
THE Skuut. Plate XVII., figs. 1, 2, 3.
The skull of S. fossor is comparatively large, with a rather low occiput, a low
sagittal crest, moderately long muzzle, and broad zygomatic arches. In general
outline the skull has a closer resemblance to that of Aplodontia rufa* than to any
other recent rodent. In studying the skull in detail there are many striking dif-
ferences between this and the recent genera. The nasals extend comparatively
farther back than in Aplodontia or Cynomys, so that the fronto-premaxillary suture
is more sinuous across the face and top of the skull than in these genera. Anteriorly
the nasals terminate rather bluntly, not overhanging the premaxillaries. The lateral
border of the nasal is supported entirely by the premaxillary, as in Aplodontia.
They are broad and flat anteriorly, forming the roof of a triangular anterior narial
opening.
The frontals are short, broad anteriorly, with a greatly constricted interorbital
region. ‘They terminate posteriorly in a wedge-shaped connection with the parietals.
The supraorbital rugosity is quite heavy in some individuals of this species, and the
temporal ridges take their origin at this swelling and rapidly unite opposite the in-
terorbital constriction. There is no postorbital process on the frontal. In this
respect the fossil is similar to Aplodontia.
In S. fossor the postorbital area of the frontal is more rounded than in Aplodontia,
and the postorbital constriction is greater. There is a distinct sagittal crest in the
ty pe.
1 The generic name first proposed in 1829 by Richardson was Aplodontia, and, while not satisfactory to a purist in
Greek etymology, has priority, and must stand.—Zditor.
PETERSON: DESCRIPTION OF NEW RODENTS 143
The parietals extend well forward and overlap the posterior part of the frontals.
Laterally, they unite with the squamosals by a suture, which runs antero-posteriorly
in a parallel line with the long axis of the skull. Posteriorly the parietals are wing-
shaped, on account of the large triangular interparietals. From the base of the
sagittal crest each ‘parietal is gently convex laterally. At the squamosal suture
there is a slight swelling, the temporal region of the squamosal being more
abruptly convex towards the zygomatic process than is the case in Aplodontia.
The brain case is rather flat, but rapidly expands back of the supraorbital con-
striction. ,
The interparietal is large, and is like that of the recent beaver. In this region
the skull bears a great resemblance to that of Castor, with a comparatively broader
occiput, and straighter lambdoid crest. Posteriorly, the interparietals are broad, and
they taper rapidly to an anterior point, thus occupying the V-shaped space in the
postero-superior border of the parietals.
The occipital surface is almost vertical, as in Cynomys, but is much broader
than in this genus. The supraoccipital occupies the greater portion of this area, as
the mastoid is but slightly inflated. The entire surface superior to the foramen
magnum is a vertical plane, and terminates superiorly in the lambdoid crest. In-
feriorly there is a shallow emargination, which forms the superior border of the
foramen magnum. At the point of contact with the exoccipitals in the lateral
margin of the foramen magnum the suture extends outward and upward, and con-
tinues in an almost straight line to the junction of the mastoid and the posterior
process of the squamosal. The sharp lambdoid crest takes its origin a little below
this point, and continues in a gently curved line to the sagittal crest.
The occipital condyles are of moderate size and are not greatly separated by the
emargination on the posterior face of the basioccipital. The paraoccipital process
is small and points directly downward, terminating in a less truncated end than in
the beaver. It is more nearly like that of Aplodontia. ‘The occipital condyle is
close to the otic bulla. The mastoid portion of the temporal bone is very little in-
flated. ‘The mastoid process is broad laterally, and compressed antero-posteriorly,
extending proportionally as low as that of Castor. It unites with the strong mas-
toid process of the squamosal, and is also fused together with the large external
‘auditory meatus. This opening is somewhat like that of the beaver, the latter hav-
ing a greater constriction just back of the opening of the tube. The whole region
back of the zygomatic arches is short in S. fossor, and has a general resemblance to
this region in Aplodontia. The tympanic bulla is inflated, flask-like, with a con-
stricted but rather large tube, which is directed outward and upward, not hori-
144 MEMOIRS OF THE CARNEGIE MUSEUM
zontally outward as in Aplodontia. The position of the tympanic bulla is a char-
acter which very strongly recalls that of the recent beaver.
In the type (No. 1217) the base of the skullis damaged, but the cotype (No. 1208)
supplements this region admirably. The basioccipital is entirely unlike that of the
beaver, and is more nearly like that seen in Aplodontia and Cynomys. Anteriorly
the basioccipital extends to opposite the anterior border of the otic bulla, in the type.
In this region the inferior face has a slight shallow groove with two faintly indicated
lateral ridges. These ridges meet 4 mm. back of the suture, and form a convex
elevation, which increases in width posteriorly. In the beaver nearly the entire
length and width of the basioccipital is deeply excavated inferiorly, which is a
constant character. The presphenoid of the fossil is apparently similar to that
of Castor.
The pterygoids are prominent and terminate in backward projecting alee, which
touch the otic bulle. Superiorly, these hamular processes are emarginated, so
as to form a large, oblong foramen, which leads into the large pterygoid fossa.
The outer process bounding the pterygoid fossa extends back to the antero-
external face of the otic bulla, and unites firmly with the floor of the brain case.
The foramen ovale pierces this outer process of the pterygoid near the extreme in-
ferior border. The alisphenoid cannot be outlined in either the type or cotype of
this species. Inskull No. 1212, however, the suture can be partly traced. ‘The bone
appears to be of large size, and similar to that of the beaver. ‘The posterior wing
extends to the tympanic, and forms a suture with it. The squamosal suture is im-
mediately below the lower border of the glenoid cavity, and extends anteriorly ina
sinuous line. The extreme anterior point cannot be ascertained, but may reach to
the posterior border of the orbit. There is an alisphenoid canal.
The squamosal occupies relatively a greater area superiorly, than in the beaver.
It has also a larger posterior portion, which joins the supraoccipital directly. In the
beaver the parietal has a descending lobe postero-laterally, which unites with the
mastoid portion of the squamosal. ‘The squamosal is thus separated from the supra-
occipital by this descending lobe in Castor. Anteriorly the squamosal extends rela-
tively farther forward in S. fossor than in the beaver. ‘The zygomatic process
of the squamosal is similar in size, shape, and position to that of Castor fiber. Pos-
terior to and continuous with the base of this process is a prominent and sharp
border, like that in S. peninsulatus, which according to Cope, enlarges the postglenoid
fossa more than in either C. fiber Linnaeus or S. Hsert v. Meyer (S. viciacensis Gervais).
There is a subsquamosal foramen present in the type. The glenoid cavity is an
antero-posteriorly elongated and laterally convex surface. The antero-posterior
PETERSON ;: DESCRIPTION OF NEW RODENTS 145
diameter is relatively greater than that of the beaver. The skull is broadest across
the zygomatic processes of the squamosuals.
The jugal is the most prominent bone in the zygomatic arch. ‘The anterior
process does not reach the lachrymal, and is proportionally shorter than in the
beavers. ‘There is a strong postorbital angle, similar to that in Castor fiber, S. vicia-
censis (see Cope, Tertiary Vertebrata, p. 841), and S. peninsulatus. The anterior part
of the jugal, across the postorbital process to the inferior margin of the arch, is the
widest surface of the bone. From this point backward, the jugal decreases rapidly,
and terminates in a somewhat enlarged end, which forms the exterior border of the
glenoid cavity. Asa whole the zygomatic arch is proportionally wider and stronger
than that of Castor, but otherwise is very similar.
The palatine plate of the maxillary extends posteriorly to the line between mt
and m*, and anteriorly to the posterior margin of the incisive foramina. In the
beaver, the palatine plate of the maxilla is shorter posteriorly. In the type the
posterior narial opening is well back of m#, and the front part of the palatine plate
is much more concave than in Castor. Anterior to the alveolar border is a promi-
nent ridge, extending from p* ina curved line on the side of the muzzle, terminating
abruptly below and in front of the infraorbital foramen. In the beaver, this ridge
is not so prominently extended to p*; it is heaviest at the infraorbital foramen, and
continues obliquely upward, across the maxillo-premaxillary suture nearly to the
top of the skull, and unites with the malo-maxillaryv ridge forming a deep trough-
like fissure for the masseter muscle. In S. fossor the malo-maxillary ridge extends
forward only a short distance and does not join the upward extended ridge men-
tioned in the skull of Castor. In the type (No. 1217), the infraorbital foramen is
small, which is characteristic of the genus. The zygomatic process of the maxillary
is very strong, vertically broad, and ends abruptly at the inferior Jjugo-maxillary
suture.
The lachrymal cannot be outlined in the type or cotype. In skull, No. 1207,
however, this bone forms an extremely small part of the facial region. The bone
is very small, perhaps even comparatively smaller than in the beaver, and, as
in that genus, is located immediately internal to the opening of the lachrymal
foramen.
The premaxillary is rather heavy to support the very large and powerful in-
cisors. Below, it is laterally convex, with a median ridge extending from the in-
cisive alveolar border to the anterior edge of the incisive foramen. ‘The latter is
relatively broader than in Castor. The maxillo-premaxillary suture in S. fossor,
may be traced from the posterior border of the incisive foramen in a slightly curved
146 MEMOIRS OF THE CARNEGIE MUSEUM
line to the root of the zygomatic process, thence obliquely back to meet the fronto-
nasal suture.
There is a strong swelling on the side of the muzzle, plainly indicating the direc-
8 § I
tion of the root of the incisor. The muzzle is broadest across this swelling. ‘The
naso-premaxillary suture converges backward, making the nasals narrower poste-
riorly than anteriorly.
THe Manprsre. Plate X VIL, figs. 1, 4.
The lower jaws are very heavy. ‘The ascending ramus occupies more than half
of the entire length of the mandible. The coronoid process takes its origin on the
exterior side of the ramus, opposite the posterior part of pg; thence ascending in a
gentle backward slope to near the summit, which is rapidly curved posteriorly, and
terminates in a thin, rounded, and transversely compressed point. There is rela-
tively a deeper and larger fossa separating the alveolar border from the coronoid
process than in the beaver. In the fossil, the alveolar border is abruptly elevated
above the diastema in front. Anteriorly, the jaws are united by a strong symphysis.
On the chin is a strong process, similar to that in the beaver. The external face of
the ramus is irregularly convex, the internal somewhat concave below the molar
series. The angle is greatly deflected outward, and the inferior portion descends
more below the border of. the horizontal ramus than it does in the beaver. In S.
fossor this angle terminates in a strong postero-lateral process. This process is very
similar to that in Aplodontia, with somewhat less inferior and exterior development.
The alveolus of the incisor terminates posteriorly in a heavy, rounded protuber-
ance on the external face of the ascending ramus below the condyle, similar to what
is seen in S. peninsulatus. The deep fossa above this protuberance is similar in the
two species, so far as can be judged from the type of S. peninsulatus. The form of
the condyle also agrees with Cope’s description: it ““is subglobular, and has con-
siderable more external than internal articular surface” (Tertiary Vertebrata).
With the exception of the less developed inferior process on the chin, the ap-
parently more rounded anterior face of the incisor, the comparatively heavier femur
and much longer tibia, the type of S. peninsulatus agrees closely with S. fossor. The
skull, which Cope associates with the lower jaw, and the hind limbs of S. peninsu-
latus, figured on Plate 68, figs. 18, 18a, 18) (Tertiary Vertebrata), is distinctly dif-
ferent from S. fossor in having less expansion across the zygomatic arches, and pos-
sessing apparently a much shorter tube on the otic bulla, and a somewhat longer
muzzle.
The skull figured and described as 8. pansus (Bull. Am. Mus. Nat. Hist., Vol.
ei
PETERSON: DESCRIPTION OF NEW RODENTS 147
XX., pp. 207-270, 1904) has the same general shape as the skull of S. fossor, but
the latter differs in having a relatively greater interorbital constriction ; the dias-
tema in front of the cheek-teeth is more highly arched; the palate is longer, and
the basioccipital is not excavated. In S. fossor the ascending ramus of the mandible
is not so vertical as that of S. pansus.
MEASUREMENTS.
Skull.
Type. Coty pe.
Greatest length mm. 76 mm.
Greatest width G6 62 OG
Greatest height, measured at m* to top of anterior part of sagittal crest ............. 30 es
Greatest width of occiput at external auditory meatus ..................0ceceee eee Pheer 48 te 48 5
Greatestawidthrofemuzzletantenonlys-sc-c.-cesocsescseasterorectnetectan tect tere ee anaes 12 a 15 “
Width of interorbital constriction “
ILE EANN. OL TACT Joconéonddo00005 008008d095de0ccb0 caso qn adadopcaqdooa0cecdoe oxecCoooDcEbOSoddboododcHoNeN we
Length of sagittal crest from union of temporal ridges to occiput ... 28 i
IRICEN OP CRED MB-codco: osesec000%= oosco6 onsocooq.bnqUdenCoODOEnEd Daas LeBsaoBOD bodeac aooceSnaedeeass 22 Oe
Antero posterior diameter of otic bulla........0. .......ccccsecesecoees cescecsescceveceseses 1B & behoulet at
Transverse diameter including tube and external auditory meatus...................+: 24 ss
Hxtremenwidthrof.occipitalecondylestcsssessarcsessseecce cere eee eee eee 16 tt
Spacesbebweenhncisorsjandeparnccan-cvesssscessdiccessereccetcr rece ee ateeec eestor eee eee 27 se 30 ue
Space from m# to and including condyle ..............:ceccccsecececsae cececeeeeeeeeecereeees 28 se
Width of palate at p4 Boa a 4 :
Wild thyotmpalatey bien seerececclncc-ccescelecsees ste sseeceteae ee assee es eaeece eae eater Enotes 6 a
Superior Dentition.
Length of incisor from the alveolar border to the cutting point..................---..6+- 21 ug 21 Oe
Antero-posterior diameter of incisor Baier Bile ot
Transverse diameter of incisor.......... 5 be 5 au
Antero-posterior diameter of grinding teeth........-....2..:2e.seceee seen ee eee eee eee eee rete 14 ee 14 ag
Antero-posterior GIETINGTIOR OF 7124 ceagocaascoonseessiooosanooxcaccannooDoGuGoaccdseadonSteds dooDaDeKC 4 oe 4 Be
Transverse diameter Of p+..............2.ccecceeccceeceecenereescr nese eceesecsesscecceeeeseceasees 4 ey 4 -
Antero-posterior diameter of m+ 3 ae Si aes
Transverse diameter Of mee... 2.2... -.c...-cncnnnscermerees oonndeees deceereesencereeesiserencrcseers 4 SF 4 oS
Antero-posterior diameter of m2 3 ‘ 3
Transverse diameter of m2 30 ** any
Antero-posterior diameter of m+ Ph 8 3 ue
Transverse! diameter! Of M2... cs. cecece cscs eer encissescrec comet roms oeneleserisrtiecceisdcssis-saeee= Pi) Oe 3 se
Mandible.
Greatest length of mandible including incisor.................. = 64 3
Length of mandible from process on angle to incisor —alveolar border.... ot 54 “
Greatest depth from angle to top of coronoid process...-......--.2-+:e-sseeeeeeeeeeeeee sees ue
TOYO IN 5 CUE SHOE pecocecac caogcceodoedeo DOLE cacorOnSo-aa care SoScenE pads sooaScaeneraacHoCAcaN coadoNecENaasIG: os 12 st
“ce 16 “cc
Height of coronoid’ process.........022.0.00+-c-ssercensecrnsere seecaretcencvereatserccrrerccseseas Way O56 “0
Antero-posterior diameter Of COndyle...........-.:-esescecnneeececereneeeecesseneeerceseeeeeeees Gira ee @G
148 MEMOIRS OF THE CARNEGIE MUSEUM
ransverse diameter Ol CONG YLe en sawcamece seman paisenccraementasisner oe ienenepts ese eases 5 mm. 5 mm.
ILGHYAAN OL CHESTENIE oonsoc osonsanbobp9opScocaDUaDORUBLAGED ScUBEODETCLSES sneacsscosseOsIbOCEeDIa09R: 16 uf 16
Inferior Dentition.
Length of incisor from alveolar border to cutting point..............22..10 0 cesses eee eeeees 25 uy 27 v7
Antero posterior diameter Of inCiSOr..............0+c002 ceeecs eee eeeeeteeeeee et eteeetetaereeees Dey Hineewia:
Mransversediam eters OfsinClSOl ese sreeeececsescreeeseetees i escece seks seeee ssa beres sear eeenee Ae Dia A Diets
Antero posterior diameter of grinding series...... ...... jeaanebosapatOs stobesbobbe Gop0onD.030 16 uy 16 oF
Antero-postenior diameter Of Piece... -ce nec er- cence sn esemslene eens ierineelerleleelsereseer ann AND Aig) 1S
TORN TS GHETNETEP OE ]Dzpsboscsdoouscada9s: avoq90 seconddaTbOS oEeqcEdeAEs, ceapSceedoEsaoRea9B02900R0C 4 fe 4 Me
Antero-posterior diameter Of myz...........-..-.0.cceecceeceeceer ere seepecceccnsecssereecessenners 4 oe 4 #
Transverse Wiameter Of Mz...............c0+se-eeenercnuerreee-eeinwscreserescrseense snpgoDons095000 4 re ARI Ties
Antero-posterior diameter, Of My....-..-....-- .-.--ceecereeeeer eer ecrmesnneernnes sermansannscess way 3H
SU IGNSD: CHETNEUWEE (1? Tig s-0n5sa5ee00du005 500000 paSd5u0090n5006000 oop agoancabedasOReDenoaEsgoqO0R 4 os 4 &
Antero-posterior diameter of m, 3 By
ADAIR, GINAINGID® O12 Mises snqsccdosecebcoacbSnosecn candusn00 Sona GecoBSHo.EREODoB =D AADeADceDeG HOD ae Of BH
VERTEBRAL FormuLaA. Plate XIX.
There were found, with the type (No. 1217), twenty-six presacral vertebree, the
sacrum, and eleven caudals. The vertebral column was not interlocked, vertebra
with vertebra, when found, but it seems reasonable, that the following formula may
be correct : seven cervicals, thirteen dorsals, six lumbars, five sacrals, and eighteen
caudals, assuming that the last seven vertebree are lost from the type. There are
probably not more than seven or eight missing, to judge from the gradual tapering
of the anterior caudal vertebree, which were found in position and continuous with
the posterior sacral vertebree.
The Atlas. —'The antero-posterior diameter of the atlas is smaller than that of
Aplodontia. ‘This is especially true of the superior arch. The inferior arch is a little
heavier than in the latter genus. The articulation for the occipital condyle is very
similar to that of Aplodontia; it is not very deep, and occupies, perhaps, more of the
area of the anterior face of the atlas than is seen in Aplodontia. The foramen for the
spinal nerve enters the superior arch internally, immediately above the cotylus, then
pierces the arch to the superior face, and again reénters it in a postero-lateral direc-
tion. There are two posterior openings, one at the base of the transverse process on
the lower side, and the other in the deep fissure above the articulation for the axis
and at the base of the transverse process on the upper side. The foramen is similar
to that of Aplodontia, but relatively smaller. In the type, the transverse process is
complete, and is remarkable for its short and heavy character. The general aspect
of the whole bone is similar to that of Aplodontia.
The Axis. — The axis is rather heavy, and is conspicuous on account of the high,
strong, and antero-posteriorly broad neural spine, which overhangs the neural canal
PETERSON : DESCRIPTION OF NEW RODENTS 149
in front. Posteriorly, the neural arch overhangs the superior part of the short
spinous process of the third cervical, so that the anterior part of the arch of the latter
is entirely within the arch of the axis. In the cotype (No. 1208), the third cervical is
entirely codssified with the axis (Pl. XVIII, Figs. 9,9a and 9b.) The articulation for
the atlas is extended well up upon the anterior border of the neural canal. The odon-
toid process is a heavy rounded peg. The delicate transverse processes are broken
off on the axis and the third cervical.
The Third Cervical. — The third cervical is charactistic in 8. fossor, on account of
its tendency to become coéssified with the axis. In the type, this vertebra is clearly
separated from the axis, but the encroachment of the latter bone on the third cervical
shows that in old individuals these two vertebrae may become codssified. In the
type the neural spine is not so high as in Aplodontia, and the centrum has the same
antero-posterior diameter as in that genus.
The Fourth Cervical. —'This vertebra is represented by the centrum and half of
the neural arch in the type specimen. The centrum is short antero-posteriorly, even
shorter than the centrum of the third. Transversely, the centra are broad, and de-
pressed vertically. The fourth cervical appears to have a heavier transverse process
than in Aplodontia.
The Fifth Cervical. —'The fifth cervical is almost identical in form with the fourth.
The neural arch and spine are somewhat heavier than in Aplodontia, and the trans-
verse processes appear stronger. The neural canal is large. The vertebrarterial
canal-is smaller thanin Aplodontia and Cynomys. ‘The prezygapophyses are directed
forward and downward, and postzygapophyses upward and backward.
The Siath Cervical. —The neural arch is injured, and the transverse processes of
this vertebra are lost. The antero-posterior diameter of the centrum is less than in
the preceding vertebra; otherwise there is not any marked difference between these
two bones.
The Seventh Cervical.—The neural arch is low and broad, similar to that in
Aplodontia, but heavier. The spine appears shorter. ‘The antero-posterior diameter
of the centrum is but very little greater than that of the preceding vertebra. The
transverse processes are heavy, and are directed horizontally outward from the cen-
trum, not postero-laterally as is the case in all the preceding vertebree. There is a
distinet facet for the first rib on the posterior face of the centrum. ‘This vertebra
has no vertebrarterial canal.
As a whole, the cervical region is comparatively short and broad. With excep-
tion of the shape of the third cervical, the neck is similar to that of Aplodontia.
Dorsal Vertebre. — There are thirteen dorsals more or less complete in the type
150 MEMOIRS OF THE CARNEGIE MUSEUM
specimen. The vertebra succeeding the thirteenth dorsal may or may not have
supported arib. This vertebra has no distinct transverse processes, and there are
no visible facets for the ribs on the sides of the centrum. In view of the fact that
Cynomys has apparently no distinct transverse processes on the first lumbar, I shall,
in this description, regard the vertebra under discussion as the first lumbar.
The First Dorsal. — This vertebra has a less robust spine than is seen in Aplo-
dontia, otherwise the neural arch is relatively as strong as in that genus. The short
centrum has a plane surface inferiorly. The transverse processes are rather short
and heavy, with a broad support for the tuberculum of the first ribs. The prezyg-
apophyses are, as usual, placed low down on the arches, at the base of the transverse
processes, while the postzygapophyses are higher up on the arch, in order to meet
the articular surface of the succeeding vertebra.
The Second Dorsal. —This vertebra is complete with the exception of the tip of
the neural spine. ‘The latter is crushed to one side, which gives it a somewhat
shorter appearance than the spine of the succeeding vertebra. The centrum is but
slightly concave antero-posteriorly, and its diameter is very little greater than that
of the first dorsal. The transverse process is strong, with a large tubercular facet
for the second rib. The prezygapophysis is even lower down on the anterior border
of the arch, than in the preceding vertebra, and does not extend so far beyond the
arch anteriorly as in Aplodontia and Cynomys. The postzygapophysis is large and
overhanging.
The Third Dorsal.— With the exception of a longer and somewhat stronger
neural spine, and the shorter transverse process this vertebra is very similar to the
one in advance of it.
The Fourth Dorsal.—'The fourth dorsal is complete, except the top of the
neural spine. The centra in this portion of the dorsal region gradually increase in
length proceeding backwards; they decrease in their transverse diameter, increase
in the vertical dimension, and acquire a more concave surface inferiorly. The
fourth dorsal has strong transverse processes, which point outward and upward from
their base. The rib contact is strong.
The Fifth, Sixth, Seventh, Highth and Ninth dorsals are so similar that the descrip-
tion of one will answer for the entire series.
The neural spines gradually become lower, the transverse processes shorter, and
the antero-posterior diameters of the centra greater. The inferior surface of the
centrum is more convex fore and aft in the eighth and ninth than in any of
the preceding dorsals. In these vertebree, and the one in advance of them,
the neural spines have lost their rounded form and gradually become laterally
PETERSON : DESCRIPTION OF NEW RODENTS 151
compressed. The tips of the neural spines of the sixth and seventh dorsals are
broken off.
The Tenth Dorsal. — In the type, this vertebra is only represented by the neural
spine, the prezygapophyses, and a small portion of the anterior part of the arch.
This vertebra has the most delicate spine in the dorsal series.
The Eleventh Dorsal. —'The eleventh dorsal is distinctly different from the pre-
ceding dorsals. This difference is chiefly found in the lateral convexity of the
postzygapophyses, and in the presence of well developed metapophyses. The trans-
verse process is only a small rounded knob, situated immediately below the base of
the metapophysis on the exterior face of the pedicle. ‘The spine of this vertebra is
much higher than on the corresponding vertebra in either Aplodontia or Cynomys.
The Twelfth Dorsal. —There is no transverse process on this vertebra. The
metapophysis is broken off. The neural spine is heavier, but is not so high as in
the preceding vertebra. The postzygapophysis is more rounded than in the eleventh
dorsal, and its articular face is directed more outward than downward. ‘The cen-
trum is less depressed than in the preceding vertebree.
The Thirteenth Dorsal. —Vhe right prezygapophysis and metapophysis are broken
off from this vertebra in the type. The top of the neural spine is also injured. The
centrum is much concave fore and aft, and is very little longer than that of the pre-
ceding vertebra. The large postzygapophysial articulation faces downward and
outward, but is not as convex as that of the eleventh and twelfth dorsals. The strong
metapophysis takes its origin at the base of the prezygapophysis on the superior part
of the pedicle and terminates in a rounded point in a parallel line with the postzy-
gapophysis. Its size, shape, and position are very similar to those of Aplodontia and
the prairie-dog.
The change of direction of the neural spine occursin the tenth dorsalin Cynomys,
while this change of position is only slightly noticeable in the tenth and eleventh
dorsals of Aplodontia. In Steneofiber fossor the change of direction of the neural
spine is gradual, similar to that in Aplodontia, but it takes place in the twelfth and
thirteenth dorsal (as in Castor), instead of the eleventh and twelfth. S. fossor also
differs from Aplodontia and Cynomysin having higher spines on the posterior por-
tion of the dorsal region.
The Lumbar Vertebrx. —'The three anterior lumbars and the thirteenth dorsal
were found articulated with one another by their zygapophyses. These vertebree
are complete except some of the delicate processes. As a whole, the lumbar
region in S. fossor is more nearly similar to that of Aplodontia than to that of
Cynomys or Castor.
152 MEMOIRS OF THE CARNEGIE MUSEUM
The First Lumbar. — The first lumbar vertebra is decidedly heavier than the last
dorsal. It is further characterized by the sudden increase of the antero-posterior
diameter of the neural spine. If the last dorsal and the first lumbar had not been
found in position, I would have been inclined to think that there might be a vertebra
missing at this point. This vertebra resembles the preceding in not having a trans-
verse process. Otherwise this bone is very similar to the following vertebra, which
has all the characteristic features met with in this region. ‘The inferior surface of
the centrum is slightly keeled, and has a greater diameter antero-posteriorly than
the last dorsal. The prezygapophysial articulation is a large surface, and is slightly
concave.
The Second, Third, and Fourth Lumbars are so similar to one another that the
description of one will answer for all. The antero-posterior diameter of the centra
gradually increases; their inferior faces are more keeled; and the transverse
processes become longer from the second to the fifth lumbar. The fourth lumbar
has apparently no metapophysis. In this respect it is similar to the same vertebra
in Aplodontia. The transverse processes are short and heavy, and are located on
the pedicle, and not directly in contact with the centrum asin Cynomys. In Castor,
these processes are high, but not as high comparatively as in S. fossor and Aplodontia.
The Fifth Lumbar. — This vertebra is represented by the complete neural arch,
the centrum having apparently become absorbed by plants that are so common
inside of Demoneliz. The most characteristic feature of this vertebra is the sudden
decrease of the neural spine. This is more apparent than in either Cynomys or
vplodontia. The transverse process is present on the left side, and is situated on a
horizontal line with the postzygapophysis.
The Siath Lumbar. — In the type, the left postzygapophysis and the end of the
right transverse process are missing from this vertebra. ‘The centrum has a greater
antero-posterior diameter, and is more keeled than in any other presacral. The
transverse process is prominent, and situated high as in the preceding lumbars.
The neural spine is somewhat more reduced than that in Aplodontia. Otherwise
this vertebra is very similar in these two genera.
The Sacrum. — There are five well codssified centra in the sacrum of S. fossor.
In Aplodontia there are six, in a young Cynomys three, and in Castor four. In the
cotype (No. 1208) there are five lumbar vertebree and a sacrum; the last sacral is
lost, but the centrum and the pleurapophyses on the preceding vertebra show, that
there was coossification between the fourth and fifth sacrals. The number of sacral
vertebrae in S. fossor consequently seems to be five, at least in two distinct cases.
The centrum of the anterior sacral vertebra in the type (No. 1217) is very nearly as
PETERSON : DESCRIPTION OF NEW RODENTS 153
heavy as that in the last lumbar. The centra of the posterior sacrals rapidly
decrease in size, corresponding to the moderately long tail.
The ilium is supported by the strong pleurapophyses of the three anterior sacrals.
In the type specimen, the neural spines are all separated in a way similar to those
in Cynomys. In:the cotype, however, the first and second spines are codssified. In
, Aplodontia, the whole series back of the first spine is coalesced into a thin plate of
bone, directed antero-posteriorly as also seems to be the case in Castor. In the type
of S. fossor, the neural spine of the third sacral vertebra was injured during the proc-
ess of cleaning up the specimen ; but the remains indicate, that it was perhaps the
heaviest spine in the sacrum. The spinous processes on the two last sacrals are
short, stout, and terminate in an enlarged rounded knob.
The sacrum as a whole presents the aspect of a narrow, long, and vertically deep
column of bones. In Aplodontia, the sacrum is of approximately the same length,
but has much less vertical depth, and decreases more rapidly toward the posterior
end, than in S. fossor. The ‘Sewellel” has a remarkably short caudal region,
while S. fossor has one about the length of that of Cynomys.
The Caudals. — As has been stated above, the tail of the type specimen consists
of eleven anterior caudals. There were seven in a continuous series and in connec-
tion with the sacrum. he four remaining vertebre are of about the right size to
‘continue the series from seven to twelve. Assuming that the caudal series in 8S.
fossor is similar to that of Cynomys, there would then be seven or eight vertebree
missing at the end of the tail.
The caudal region of S. fossor is heavier than in Cynomys, but in form it rather
resembles the latter genus than Castor or Aplodontia. In the type the centra are
short, round, and heavy. There is no neural canal back of the fourth caudal. The
antero-posterior diameter is nearly equal in the first and last caudal vertebree in the
specimen under discussion. In the five anterior caudals there are distinct transverse
processes. In Cynomys, these processes can be traced back to the ninth caudal,
while in Castor they continue to the end of the tail. From the sixth caudal to the
tenth, there are small protuberances on the inferior faces of the vertebrae, which may
or may not have supported chevrons. In the Trans. Amer. Philos. Soc., vol. 17,
1893, p. 77, Professor Scott states that the anterior portion of the caudal region was
provided with chevrons in S. montanus. The caudal vertebree present in the type
(No. 1217) indicate a heavy, moderately long, and round tail.
The Ribs, — Steneofiber fossor probably had thirteen pairs of ribs. Cynomys has
twelve, Aplodontia fourteen, and Castor canadensis fifteen.
In the type specimen, there is a fairly complete set of ribs. The first is much
154 MEMOIRS OF THE CARNEGIE MUSEUM
flattened, expanded distally, and provided with a strong contact for the cartilag-
inous rib of the sternum. The shaft of the rib decreases in width superiorly, and
is nearly round immediately below the heavy tuberculum. ‘The head is rather
small and rounded. The rib as a whole is different, especially distally, from that
of Cynomys and Aplodontia. The latter genera have the lower end of the rib more
rounded in cross section.
The ribs of the anterior half of the thorax are flat proximally, and rod-like dis-
tally. In the meso-thorax, and more posteriorly, they are rod-like throughout.
The Clavicle. —The clavicle in 8. fossor is heavier than in either Cynomys or Aplo-
dontia. In the cotype (No. 1208) both clavicles and the manubrium are present.
The latter is much heavier than in Aplodontia or Cynomys. Anteriorly, there is a
broad, gently rounded surface for the attachment of the clavicles. The attachments
for the first set of ribs are very large, and are located close to the anterior end of the
bone. Back of this rib-attachment is a long and gentle Jateral constriction. Infe-
riorly, the bone is slightly keeled ; supero-anteriorly itis concave. Posteriorly it
expands above and on the sides, indicating a rather large mesosternum.
Both clavicles in the type are complete, and are not so robust as those of the cotype.
The latter specimen represents a somewhat larger and more robust individual. In
shape, the clavicle is very similar to that of the beaver. The sternal attachment is
more enlarged than in Cynomys and Aplodontia. ‘The curve of the shaft in S. fossor
is similar to that in Aplodontia and Castor. The superior end is less flattened than
it is in the beaver and the prairie-dog, and does not terminate in an enlarged head
as in Aplodontia. ‘The clavicle of S. fossor gradually tapers from the large sternal
contact to the superior end, which has on the internal face a wide, shallow groove
at the contact with the acromion process of the scapula.
MEASUREMENTS.
Approximate length of the vertebral column including skull.........................eseeeeeeee 37 cm.
meng thyoficervica lar epion ssevee ses. veencccesasensensecie cern cee eee erecta see toeesee et neeeeee cesta 20 mm.
Greatest antero-posterior diameter of atlas..............0.:ccccseceecceceeenesec eee eeeeceeeeeeeeeseeees eee
Greatest transverse diameter Of atlas..................cccecceceeeeeteeeeeneereeeeeeeeesetenseceeseee sees OB} 8
Greatest diameter of cotyli for occipital condyle... ................ceceeeeeceeceeceeceeer eee eeeeeeeens Sins
Greatestsheich tioffatlastresssn-cscsecusste seine pees sotecceeseeciaeetcseceeceeactecreceeaecneerases Ie %
Antero-posterior diameter of centrum including odontoid process of axis g
Antero-posterior diameter of centrum of axis at the base of the odontvid process........... Bye Oy
Greatest transverse diameter of axis anteriOrly................cceceece sce eec eee eee eeeseseeneseeeeeees 122 66
Greatest height of axis including spine.......... .......0ccecceec eee eec eee eee eee eeeeecseueereceeeseeeees 1G
Greatest height of neural spine of axis. Gres
Antero-posterior diameter of third cervical acy
a sf se fourth ‘“ 33)
. ae ee fifth ee 3.5 ‘f
PETERSON : DESCRIPTION OF NEW RODENTS
Antero-posterior diameter of sixth Cervical. ............cc0ceecceeeeeenceeeeeees seeeeceeesecseeessceees
oc oe “ce
seventh ‘‘
Length of dorsal region
Antero-posterior diameter of first Corsal.............:ce0cceseececneeeeeeeeeeoeeees ceveeeeaseeeseceeees
Transverse diameter of centrum of first dorsal, posteriorly
Height of centrum of first dorsal
Height including neural spine of first dorsal '..
Antero-posterior diameter of centrum of fifth Uorsal............00.c00.c0eccceeceneeeeeeeceecceneses
Transverse a oH Gu 6
Vertical oe G6 ee
Total height including spine on fifth dorsal...............cesecceeeceeeeeesecesccccesaeseeaeeeueees eee
Total height of neural spine on fifth dorsal
Antero-posterior diameter of sixth dorsal...
Transverse st oe Si slolaleee asl iecieicirclsvistlctemuaial wens tea ok aaron hers sae vemeeracian
Heleht) ofscentrumyotesixthydorsalesaddeccue cect teere eececeeeteeaecereeee reer reeeceere eee
Total height including spine on sixth dorsal...............ceccseceeeecen scenes cents ee teeeceanes aes
Antero-posterior diameter of centrum of twelfth dorsal...... 2... ...02..cccecceeeeseeeeeeeeeeees
Transverse ie he be Ce soeaastee suls'cc qacecsuieee a ma oee oat eenet
Vertical diameter of centrum of twelfth dorsal............:s0ccccececcececeseeesecesneeeereutsaseeees
Total height including spine on twelfth dorsal....
Mengthiofplumbari ner iOntens.c-eose-ccsscsaceeecsesssde caciesacecemtesere
Antero-posterior diameter of centrum of first lumbar.........-.00.02. ceeeeeeeeeceeesenteeeee een ees
Transverse sf et YY i
Vertical oe i re oe
Total height including spine of first lumbar...................00002 cess veeceeceeaeeceeneeeeeceeee a
Antero-posterior diameter of centrum of third lumbar...................2.cceccecececeeeeeeeeeenes
Transverse diameter on posterior face of centrum of third lumbar...
Vertical diameter of centrum of third lumbar.............0...00ccceeceeeeeeceeeeeneceeeeeeeceeeeanees
Total height including spine of third Jumbat....................cecceec eee ceet ee ceeees cee eeneeneenes
Antero-posterior diameter of centrum of sixth Jumbar.................0:0cceceeeeeeeeeeec eee ee ees
Transverse diameter of centrum of sixth lurabar, posteriorly..............-....000eeeee eens eens
Vertical diameter of centrum of sixth lumbar
Total height of sixth lumbar including spine
Wenpthvotsacru less -eeceecesecsteaciestenceestercecscens
Greatest width of sacrum, anteriorly
OH " ‘¢ posteriorly
Antero-posterior diameter of first sacral ..............2.02 ceceeeceeceececee nesses teeeteueeeee tense netes
st cn BECOME ae ieee aU ae ea, cooat ae penemcceartntaan yee reec seen
“ ss BH Tre ses A eee st Peach, tie a oa Weed hee en ee Oe a
ms ee fourth ‘
“s oY fifth ‘* :
Height of sacrum anteriorly, including neural spine.................0c000 cceeeecseeee nee steer ees
uy ‘posteriorly, yy s Uo “goadoncos eoqsonscnas05a000000N0000 Fisetee oes
Total length of eleven anterior Caudals............:ccceeeceseeceeceeeeeeeeceecee ceeeuesetsesesseeseaees
Antero-posterior diameter of first caudal..............020:csceveececeeeeees ceseeeees cecees vereeceeeees
Transverse we £ OG | Qoacindasenypencoseacnndnmd doscusbosaseauoneiconanocHanaxobndaeL
Vertical ys a BOO Suen wanldeagetSnsunasnte sdevetecuscrteece eee eet
Total height of first caudal, including spine
1The top end of the spine is broken off in the type.
3
mm.
“a
«é
165 “
6 sc
a
5 “
“ee
“ee
7 ae
a
5 oe
1a
8.5
9 “ee
a5 ©
16.5 *
9 oe
9 oe
oe
ce
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6
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a &
Th oe
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Gra
ius
7B 2 6
6 cc
45
AU 08
9 oe
155
156 MEMOIRS OF THE CARNEGIE MUSEUM
Antero-posterior diameter of centrum of fourth caudal .............0..2.20.ce cee cee eee see eee eee eee 6.5 mm.
Transverse diameter of centrum of fourth caudal.............. .....eccc ccc cee ese eee ere eee ce eeees eee 4.5 ‘
Vertical fs ss fe yiliealeaeaocnea cho ssecomdank cane baacdangvosencHoaEaCuudoaon 45
Total height of fourth caudal, including tubercle.....................cccceecee eee eceeceeneeeeeeee Gr ee
Antero-posterior diameter of centrum of eighth caudal 6.5 **
Transverse es st ts ss ae
Vertical x ye 4 ~ ye De
Antero-posterior ‘‘ se eleventh ‘‘ yf
Transverse uf “ a of Rie
Vertical ue wee uy a Sai
Tue Fore-Lims. Pl. XVIII.
The left scapula in Steneofiber fossor is fairly complete in the type. In the cotype
the scapula is complete, and this one is figured together with that of the type (see
Pl. XVIII, fig. 2). This bone is conspicuous on account of the heavy and high spine.
The lower border of the latter arises abruptly on the external face of the scapula 5
mm. above the glenoid cavity, then descends again in acurved line to form the very
broad acromion process. ‘This process extends much below the glenoid cavity.
The spine forms a sinuous curve, and continues to the extreme superior border. It
divides the narrow blade so as to form a slightly greater prescapular than postscap-
ular fossa. The glenoid border forms almost a straight line from the glenoid cavity
to the suprascapular border. There is no constricted neck above the glenoid cavity
on the glenoid border. On a direct side view the spine is seen to greatly overhang
the postscapular fossa. The latter is concave the entire length of the bone. The
coracoid border of the scapula takes its origin at the base of the metacromion pro-
cess, and continues in a curved line to within one fourth of the distance to the supra-
scapular border, where the blade is broadest; then.it gently curves to the superior
end. The superior half of the coracoid border is turned out so that it forms a con-
cavity on the blade of the prescapular fossa. The suprascapular border is compara-
tively heavy. There is a heavy metacromion process, which is curved inward and
backward. The glenoid cavity has an oblong antero-posterior articulation for the
humerus, and is somewhat similar to that of Aplodontia and Cynomys. There is a
neck on the scapula above the glenoid cavity on the coracoid border similar to that
in the beaver. The internal surface of the blade, especially superiorly, forms an
open, reentrant, 3-shaped curve, like that seen in the beaver. The internal face
of the scapula in Cynomys has sharp ridges running parallel with the long axis of
the bone. ‘This is also seen, in a less degree, in Aplodontia. These ridges are not
sharp, and are gently curved in Stencofiber fossor, and are here more nearly like those
in Castor. In fact, the bone as a whole, with the exception of the well developed
PETERSON : DESCRIPTION OF NEW RODENTS 157
metacromion, and the broad acromion process, is very similar to that of Castor can-
adensis.
The Humerus. — Both humeri are represented in the type as well as in the cotype.
This bone (P]. XVIII, fig. 3) is comparatively short and stout, and has a great lateral
expansion distally. Proximally, the large head is well rounded, the antero-posterior
diameter being the greatest. ‘The tuberosities are fully as large as those of Aplo-
dontia. The shaft is triangular in cross-section, especially so superiorly. The deltoid
ridge is strong, with a transversely placed broad plate. This plate nearly equally
overhangs the ulnar and radial sides of the deltoid ridge ; it is entirely different from
that in Cynomys, Aplodontia, or Castor; in the latter it has a large tuberosity on this
ridge for the attachment of the deltoid muscle. In the type of S. fossor on the ulnar
side below the tuberosity there is a short, sharp ridge, turned slightly posteriorly.
As has been stated, the humerus is greatly expanded distally, the large internal epi-
condyle and the greatly expanded supinator ridge being the chief factors in this
expansion. As in Stencofiber montanus, the supinator ridge extends high upon the
posterior aspect of the shaft. The trochlea is relatively as broad and as shallow as
that in the beaver. Cynomys has the trochlea much deeper, and Aplodontia has it
about the same depth as S. fossor. The supratrochlear fossa isas deep or deeper than
in Aplodontia, and the anconeal fossa is approximately the same as in that genus.
The internal epicondyle is comparatively heavier than in the beaver, and is perfor-
ated by a large foramen, as in S. montanus and in Aplodontia. In the mole the
internal epicondyle is relatively heavier than in S. fossor, and is perforated by a
foramen.
radius and Ulna.—The right and left radius and ulna are present in the type
(Pl. XVIITI., fig. 4,4a@). The forearm is relatively shorter than in Cynomys or Aplo-
dontia. The shaft of the radius is a round and irregularly curved rod, and may be
in somewhat closer proximity to the shaft of the ulna than is represented in our
illustration. Superiorly the radius flares out abruptly into a large head to furnish
a good support for the trochlea of the humerus. Distally. the radius is even larger
than proximally, and sends down a heavy internal and a smaller external process
below the large articular facet for the carpals. In S. fossor this external process is
much better developed than in Cynomys, Aplodontia, and the beaver.
The ulna has a heavier shaft than the radius; it is compressed laterally, and is
broad antero-posteriorly, with a shallow groove extending from the sigmoid notch
to near the distal end of the bone on the external side. This groove is better defined
than in Aplodontia, which genus has the bones of the forearm in close contact
throughout. In Cynomys the radius and ulna are separated, as in S. fossor. In
158 MEMOIRS OF THE CARNEGIE MUSEUM
the fossil the sigmoid notch is wide internally, with a strong supporting rib-like
brace, which unites with the shaft below, and is relatively heavier than that seen
in Aplodontia. The olecranon process is more produced upward, and hasa stronger
muscular attachment in S. fossor than in either Cyncmys or Aplodontia. Tn the type
the shaft of the ulna is much less curved than that of the radius, and terminates in
a round knob, which articulates with the carpals.
Fic. 3. Fore-limb of Steneofiber fossor in position ; somewhat larger than natural size.
The accompanying figure (Fig. 3) represents the fore-limb and foot of S. fossor in
position. The limb presents a striking similarity to that of other fossorial rodents,
and approaches that of the mole in its position. The elongated and narrow scapula
of the mole, the heavy clavicle, the strongly built humerus, and the broad foot with
the long and powerful unguals, is rather suggestive of the habits of this animal,
which was probably fossorial to a considerable degree.
The Fore-foot. — Of the carpals there are, unfortunately, only the cuneiform, unci-
form, and trapezoid, the first and second metacarpals, and the first phalanx of the
PETERSON : DESCRIPTION OF NEW RODENTS 159
pollex represented in the type. Another individual of the same species, which is more
robust (No. 1204) has the phalanges, first, second and third metacarpals, and the
unciform preserved... A composite figure of the two specimens, the type, No. 1217
and No. 1204, represents the left manus of S. fossor on PI. XVIII, fig. 5. The
slight disparity of the two individuals is at once noticed in this illustration.
The Unciform in the type supports the fourth and fifth metacarpals. This bone
is rather broad laterally, and the articulation for the scapholunar and cuneiform, is
postero-radially a higher ridge than in Aplodontia, and is more nearly like that of
Cynomys. . Distally the articulation for the metacarpals is triangular in outline ; it
is concave fore and aft, and slightly convex transversely. The bone as a whole is
similar to that in Cynomys.
The Trapezoid. — The trapezoid is a small irregular bone, which supports me. IT.
The inferior articulation is obliquely concave antero-posteriorly. The superior sur-
face is nearly flat.
Metacarpal I.—The first metacarpal is fairly well developed, and is parallel
with the long axis of the manus. The first phalanx is strong, and supports an
ungual, which is perhaps moderately developed as in Cynomys, Aplodontia, and other
genera of this family. There is what appears to be a radial sesamoid in position
in a smaller species. ‘This species will be described later on in this paper.
Metacarpal IT. is shorter than in Aplodontia, and is of about the same size.
Metacarpals III, IV., and V. are present in the specimen No. 1204, and are
relatively short and heavy. Me. III. is broken off superiorly, but it indicates a bone
as long as the corresponding one in Aplodontia, and is much heavier. In the fossil,
me. III. is in comparison somewhat more expanded distally than that of Aplodontia.
Me. IV. is of the same length as that of Aplodontia, but is much heavier than in this
genus or in Cynomys. In our fossil, the proximal articulation is deep antero-pos-
teriorly. This is due to the large tuberosity on the plantar face of the bone. The
shaft is slightly constricted, and the distal end has a strong keel on the palmar face.
Metacarpal V.is extremely short, has practically no shaft, and is as heavy as me.
IV. The articulations for the unciform and the proximal phalanx are quite large
on the nodular-shaped bone. ‘There is a deep transverse groove on the plantar face. —
The proximal row of phalanges are depressed and transversely broad. The proxi-
mal articulation is a rounded pit with a moderately deep notch on the inferior
margin. The second row of phalanges are even more depressed than the proximal
row. Proximally, the phalanges are concave antero-posteriorly and slightly convex
transversely. The distal trochlea is broad, flat, and extends to a slight degree on
the anterior face of the bone.
160 MEMOIRS OF THE CARNEGIE MUSEUM
The ungual phalanges are heavy, long, and broad, with apparently no grooves
anteriorly. The unguals have more similarity to those of the beaver than to those
of Cynomys or of Aplodontia. Proximally, the unguals have a heavy plantar tuber-
osity. Distally the phalanges terminate in a sharp, depressed point. The plan-
tar surface is broad and flat, and is entirely different from that of Cynomys and
Aplodontia ; it is more nearly similar to that of Geomys.
MEASUREMENTS.
Greatest length of scapula including metacromion process..............0022050 -cseec eee eeeeeeeee seen erees 45 mm.
Greatest width of scapula 16 .
Greatestuherchtiofispinerofisca pull aresensmcsce ecco ie seeceecicesenciesiatsctine sheet see ises eraser eee eee see eeee 13 e
Length of acromion process...............-...1.+- sd ehee seat Riles ua os woh Sateen ee stclstieen se else elle oat aecee se eee 12 et
Antero-posterior diameter of glenoid Cavity...............0.c0.cecceeeee eee cecnee nec eeeeeeceeeseeseeceeeeesees 9 of
ransversedianietemoferlenold cally eenectesss sree sesse cece secede reise eeeisec eee eater eres na ee nes 5 iN
Greatestilensthiolwhumerusseeccsesecessecteees es ceeree es eee ca ence eee teeter Rerese etme 41 Re
Antero-posterior diameter of head of humerus... 9 &
Transverse diameter of head of HuMerus.............025....0.ceceecee eee eseeeccerececee ser cenereuse ees cesceeees Gh
Greatest transverse diameter of distal end of humierus....................02-0200ceeeeeceeceeseeeeeceeeeeeee 16 si
Greatest engthiofaradlussecetseseoterecscese recess cetera roretere acer erissseereccseece ret ercee aero 3215)
Transverse diameter of head of radius.... He
Transverse diameter of distal end of radius..............0.0.0.0cc cece eee e cece cee eee eee esse eerereeer sec eee 7 es
(CERNE MERE, OF WITEES cooconceacenhaq0n00. cen aoocadconuebooDasEqsqGedGeqcded07G00 d0b00 HoH SDasoaEABIAAODJaEecDoaNB0NE 46 &
Heishtrofolecranonsprocessssntsc.sosetecssceeteereee eee scarce reece ese eaetene sees a ceee eee ee ene 10 a
Aypproximatellengthvolsmans ssn. siscecn seas cose a iaesesveosieset a act ssecioce ear secon cieecteciasceersscieeeisecieee 43 su
18 SES) 08 GS2. TVR) Tit oma cbgonddooqdasoo obs Abo 968 nebanNeGanooodd bee sodkNS nSbeds os oun ELadESooDgohSoansondunHSoenboaGeDO ao 8} Me
airansverse\diameter, ofsuncilonmercc-cersecece cee eects eeascceas-ectecee sees eeceadaceeeeecrereeaeceeeceee sen tO sO mis
Motallengthroismetacar pall eleeseneeeeeceses eee ea seeese eee teen eee Reee eeeeee eeeecetoeecneieccecsnaces 5 “
MotaljlensthrofsmetacarpaleWcresnsresceeeeeereecree ss ceicecre testcase cneceetcescatter see sencerecees ee
Motalleng th) of metacarpal PV es. e cms sie saws se le celeste nt eelaieise aelee snleelneialseteleeisiemetelessielseisaseets pa uideeless se
Rotalblengthyotem etacarpalaViccssse esc ee eee ce rere ese ee ee eeee ene eee cee eee és
Length of ungual of fifth digit... Ke a
Weng throfsuncualvofsthindidigiteessescatee eee scteeee eer eeceeeeaeee eee eee eeesee eerereee tee eeeees oe
Length of ungual of second digit
* These measurements are taken from specimen No. 1204, which is a larger individual than the type.
Tue Hinp Limes. PI. XVIII.
The Pelvis. —'The pelvis is well preserved in the type, and presents considerable
similarity to that of Castor canadensis. The straight fore and aft diameter of the
ilium and ischium, the heavy ischial tuberosity, and the wide posterior border of the
thyroid foramen are especially similar in the two genera.
In Steneofiber fossor, the ium is trihedral throughout its entire length. The
supra-iliac border is not much expanded. ‘he sacral surface is moderately long,
and extends farther anteriorly than in Cynomys and Aplodontia. The external
surface is a concave fossa nearly the entire length of the bone. The ischial border
PETERSON : DESCRIPTION OF NEW RODENTS 161
extends well back. he acetabular border is much more prominent than in
Cynomys or Aplodontia. The pubic border is also quite prominent. The acetabu-
lum is deeper than in the recent genera referred to; the ischium and pubis are
- stronger, and the thyroid foramen is smaller. The pubic symphysis is short, like
that of Aplodontia, and a detached epiphysis was found in position in the type,
similar to that of Castor canadensis. . The vertical depth from the tuberosity on the
ischium to the pubic symphysis is greater than in Aplodontia and Cynomys. The
latter genera have the superior (anterior) part of the ilium turned outward (espe-
cially Cynomys) at a considerable angle, while S. fossor has not this feature. This is
due to the greater anterior extension of the ilium, beyond the sacrum, and in order
to give space for the posterior lumbar region in the recent genera; while in S. fossor
the attachment for the sacrum extends more anteriorly, the last lumbar being oppo-
site the supra-iliac border.
The Femur.— Both femora are well preserved in the type. This bone is some-
what more lightly constructed, and the neck below the head is longer than in S.
penimsulatus. Otherwise the similarity is very apparent in the two species. In 8.
fossor the head is well rounded. The greater and third trochanters are relatively
lighter, the whole bone longer, and not so flat as in the beaver. From Professor
Scott’s description (“The Mammalia of the Deep River Beds,” p. 77) it appears that
S. montanus has the third trochanter ‘ placed more proximally than in S. peninsu-
?
latus.’ In S. fossor the third trochanter is placed immediately above the middle of
the shaft, as in S. peninsulatus. In 8S. fossor the fossa between the condyles is nar-
rower and deeper than in S. peninsulatus. The condyles have the same relative
proportion, and the trochlea are somewhat deeper than in the John Day species.
In Aplodontia, the third trochanter is little developed, and is placed high up on the
shaft like that in Cynomys. In Castor, the third trochanter is placed nearly mid-
way between the proximal and the distal ends of the femur.
Patella. —The patella is present in the type, and its characters are nearly like
those in Cynomys and Aplodontia, but different from those in Castor. In the latter
genus, this bone is thick superiorly, and tapers to a trihedral point inferiorly, while
in S. fossor it has a more ovate outline, and is as heavy below as above. The bone
is larger than in Cynomys and Aplodontia.
The Tibia. —Vhe tibia and fibula are well represented in the type, being present
on both sides. The tibia is 4 mm. shorter than the femur. In the beaver, it is
over 20 mm. longer than the femur. In Fiber zibethicus the tibia and fibula are also
much longer than the femur. In the fossil, the relative length of the femur and the
shin-bone is more nearly similar to that of Cynomys and Aplodontia. Proximally
162 MEMOIRS OF THE CARNEGIE MUSEUM
the articular surface has apparently a greater antero-posterior convexity, indicating
perhaps a greater flexion of the limb than in Cynomys or Aplodontia. In the type
the median eminence, or spine, is well developed, showing strong attachment for an
intra-articular ligament. On the fibular side, near the edge of the articulation,
there is a small sesamoid in position, similar to that seen in Aplodontia on the
articular surface of the tibia. Proximally the shaft is strongly trihedral. The
cnemial crest is as strong as in Aplodontia and S. penimsulatus. In the latter species,
the tibia and the femur are of more nearly equal length than in S. fossor.’
Distally the shaft is more rounded in cross section. The astragalar facets have
a somewhat greater antero-posterior diameter than in Cynomys, and the posterior
downward process or malleolus is strong and not grooved as in the recent genus.
The external articular facet for the astragalus is higher up on the bone, has an oblique
position, and is much larger than the internal facet. There is a more distinct ridge
separating these two facets in S. fossor than in Cynomys.
The Fibula. — The fibula is relatively as large as that of Aplodontia and Cynomys,
and, as in these genera and the beaver, it is free. The proximal end is somewhat
injured in the type, but it indicates a large tuberosity on the external side for the
attachment of the lateral ligament, such as is found in Aplodontia and Cynomys.
In Castor, this tuberosity is produced into a prominent process, directed downward
and outward. In Fiber zibethicus this process is also quite prominent. The shaft
of the fibula, in the type, is more flattened superiorly than in Aplodontia, and is
more nearly like that in Cynomys. The distal epiphysis is slipped off so that its
character cannot be ascertained. The caleaneum indicates, however, that the fibula
may have touched the exterior face of the sustentacular facet.
The Tarsus. —The tarsus is represented in the type by the calcaneam, astragalus,
cuboid, all the metatarsals, and the phalanges of the third and fourth digits. The
second phalanx of the second digit is also present.
The tarsus and carpus of S. fossor have approximately the same relative size as
in Aplodontia and Cynomys, and are entirely unlike those in Castor, as the following
description will show.
Caleanewm. —'The caleaneum in S. fossor has, as in S. monianus, “a short de-
pressed, irregular, and club-shaped tuber.” The tuber is relatively much broader
than in the beaver, but the oblique exterior face seen in the latter genus, when the
bone is in its position in the pes, is also very apparent in S. fossor. The articulation
for the cuboid is a rounded, shallow pit. ‘The sustentacular facet is relatively as
1Piofessor Cope’s illustration (‘‘ Tertiary Vertebrata,’’ P]. 63, fig. 21) is incorrect. This illustration indicates a part
of the shaft missing, but the true contact between the two parts has been found, and the bone fitted together, in the
American Museum of Natural History.
PETERSON ; DESCRIPTION OF NEW RODENTS 163
broad as that in the beaver. As a whole, the caleaneum is perhaps more similar to
that of Castor than to that of Cynomys or Aplodontia.
Astragalus. —The principal difference, in the astragalus of S. fossor and Castor,
is found in the prominent tuberosity below the articulation for the tibia, on the
internal side near the sustentacular facet in the latter. This tuberosity is not present
in S. fossor. Cynomys has it slightly indicated. In Aplodontia it is like that of S.
fossor. In the latter, the constriction above the navicular facet, is relatively longer
than in Castor, and is similar to that in Cynomys. The articulation for the tibia is
regularly convex antero-posteriorly, and concave transversely, forming a rather shal-
low trochlea, similar to that in Cynomys. The navicular articulation is less spherical
in the latter genus than in S. fossor.
The Cuboid. —The cuboid is injured antero-superiorly, so that the articular sur-
face for the calcaneum is nearly lost, in the type. Judging, however, from the
rounded, shallow pit on the distal end of the calcaneum, the facet on the cuboid
would have a correspondingly convex, rounded appearance. In the beaver, the
ealeaneal articulation is a long, narrow, convex surface, which is placed obliquely
antero-externally and postero-internally on the proximal face of the bone. The
cuboid in the type has a more depressed appearance than in Cynomys, the distal
end being more expanded, with a larger articulation for mt. 1V., and no constriction
above this facet on the posterior face of the bone, as in Cynomys. There is a strong
contact for the ectocuneiform and the nayicular.
Metatarsal I. —(Pl. XVIIL., fig. 14.) The first metatarsal is somewhat more re-
duced in length than that of Cynomys and Aplodontia, and is not at all like the
greatly developed hallux of the beaver. The digit is parallel with the long axis of
the pes, as in Cynomys and Aplodontia. The proximal articulation is long and
irregular antero-posteriorly, and narrow transversely. The ascending postero-external
process, which reaches over and articulates with the plantar face of mt. II. in Cyn-
omys, is represented only by a short rounded knob in S. fossor ; it extends directly
backward. he articulation for mt. IL. is quite strong. The distal end is slightly
enlarged, and the shaft consequently constricted ; the transverse diameter is a little
greater than the antero-posterior.
Metatarsals IT. and ITIL, in the type, are of nearly equal length. Mt. II. has a
narrow proximal articulation, and not an expanded head like that in Cynonvys.
The shaft is flat and depressed, distally enlarged, and has a heavy rounded plantar
keel. The second digit in the beaver is comparatively lighter than in S. fossor; in
Cynomys and Aplodontia, this condition is more nearly similar to that in the fossil.
Mt. ILI. is the strongest metatarsal in the type; there is, however, not much differ-
. 164 MEMOIRS OF THE CARNEGIE MUSEUM
ence in the metatarsals II., III., and IV., as is also the case in Aplodontia and
Cynomys. The proximal articulation in mt. IIL. is slightly oblique, and extends
to the posterior end of the plantar tuberosity. The shaft is broad, slightly con-
stricted, depressed, and the distal articulation is moderately reflected on the anterior
face of the bone. There is a strong keel.
Metatarsal IV, is slightly lighter than mt. III., in the type. In the beaver, this
digit is a little the heaviest of the series. In the fossil the proximal end has some-
what similar interlocking characters with mt. III. and the cuboid, as in Cynomys
and Aplodontia. These latter genera, especially Cynomys, have more rounded and
much longer metatarsals than S. fossor. In Aplodontia, the metatarsals II., I].
and IV. are approximately of the same length as those of S. fossor, but they are
much lighter.
Metatarsal V. (PJ. XVIIL., fig. 14) is oniy half as long as mt. 1V. in the type. Its
relative length is much less than in Cynomys or Aplodontia. In the beaver the fifth
digit is relatively longer than in Cynomys. Metatarsal V. articulates with the outer
side of mt. [V., and has no direct articulation for the cuboid. In Cynomys the
superior end of mt. V. continues above the fourth metatarsal articulation in an up-
ward and outward direction, terminating in a strong tuberosity. This tuberosity is
developed in a much less degree in Steneofiber fossor, and is more like that of the
beaver. The shaft of the bone in the fossil is short, constricted, and trihedral. The
distal end is an enlarged, rounded knob, with distinct articulation for the first
phalanx.
Phalanges. —The first and second row of phalanges are broad and depressed,
similar to those in the manus. The articulation for the unguals is broad, plane, and
well rounded on the anterior face of the bone. The unguals are somewhat smaller
than on the fore foot, but of practically the same shape. The plantar rugosity is
not so prominent as that of the manus.
MEASUREMENTS.
Greatestilength of pelvis: ses: cece cer seston weiosncceicoa nals uatinn os re hoabne sfaelgseenene sae ae o ease eo Renecetene 66 mm.
Greatest width of superior border of ilium................2..cc ccc ee cece cence eerste cece este eeeeeeeceeeeeues 11 se
Length of ilium from acetabulum to supra-iliac border. ...............cccesecee cece ec eee eee eeceeee eee eienees :
Length of ischium from posterior border of acetabulum to ischial tuberosity....
Greatest width of ischium and pubis, posteriorly................0-.0cc.ceeeeeeeceeee eee ce ec erssee ness eeeeees 26 ‘
Vertical diameter of ischium and pubis at symphysis, when pelvis is in position. .................. 15
CNsE Hess NET PAIN Ohi TET oo: covsosoq90d09Ne 0006000 sono c9EenOseooDoHo0 SSaoDbaINCEDsD960000000 960050505 can000a7000000000
Transverse diameter of proximal end of femur...
Transverse diameter of distal end of femur.................ccccceceeceeeeecne eee ecenee eects eens eeeceeeteneneeteee us
Greatestilengthvotmtibiaerccssceenecescicm se oclsameciseaeeessseemdscercatrescreccinsteceerercicecceete cece encenes 51 es
Greatest antero-posterior diameter of tibia at proximal end...................-..066+ odso0scon00G000D0c00000 12 ef
PETERSON : DESCRIPTION OF NEW RODENTS 165
Transverse diameter of tibia at proximal end....................0.00000008- ccodoacoadeouneosusconcoqcoy ae0095000 12.5 mm.
Antero-posterior diameter of tibia at distal end...............:.2....s00scc20se+ sesececnscecessecuessecesenseee 7 oe
piransverseldiamenumonstibialatidistalye nc se cssseeseseeeeeteeers ae ee ee ae eee ee 7 ce
Createst#l ens throiccal caneuniis.cst.c.seassrscestosetrscrec steve sehen ee eee ee es 15 oe
Length of tuberosity of caleaneum Cesieeoee
Vertical diameter of tuberosity of calcaneum)... .........-22-c0cccccccseececeseeeeensereseseeseenessneraneees Uc)! ee
Transverse diameter of tuberosity of calcaneum............-.-:1:ccseecceeceececesteeceeeecescceeeaeenseeseeees 4 Ue
Width of sustentacular facet of calcaneum .............-..00.:.0cceeeeescececsecececesecnecersecesseneseesecees
Greatest width of caclaneum at sustentacular facet
Gireatestglenpthvohiastragalisisses- cs sesctseacheccus se cece ee saetecee ne secon ecise eee eee ee eee eeesteaees te
Greatestawidthyoffastragalushats...cs.ceccs se sctvese aoc one chcwn usecase sescnewen eee nace ane coat eeee DEAR me eee 7 ie
Antero-postenlon diameteriof euboide-essess-cs-ceeesesteese siete soe eeeseneeeseceeeeceeceeteceee eee ees erence 6 es
sliransverse; diam eter Of CabOi dss. so. cess accrediation ee 4 of
IRIGTANTE Oe GLOISYOIG Canaasonconeaeestopbeteet Beane ceaes ech aen chacooh prance socbbEaoademacmonaanad
Length of metatarsal I
eng thyofsm etatarsalellitaccs.2. cc decseace spacers ses te guaces da siescioeseen seea gues st aceevecttete oe seteeet nemeareonaaes 14 a
Beng thpolimetatarsa lille ceccat esos emesis Sooo ea eae Se SSE co BEE ERE Se eet
Length of metatarsal 1V..
WWenethyrofem eta tarsaley siecaccnaccendesance cote ee aoc esae een aa nee SESE CIES Sie cIR EE Hw eta Se inate ate inte fe
Henethrofsungualyphalanxoffonrbhidigiteessssceesssessesseeesscceecee ee ete eieest eae eee ce cne eee ease ere: 9 re
Length of ungual phalanx of second digit (approximately) .........0002.00ece.0cseeceeneenceceenteseeeees 10 ue
RESTORATION OF STENEOFIBER Fossor. Pl. XIX.
The type-specimen (No. 1217) is used as basis for this restoration. The scapula,
however, and four of the anterior ribs in the illustration are drawn from the cotype
(No. 1208). The greater portion of the fore-foot is reconstructed from the specimen
No. 1204, which has also been referred to in the foregoing pages.
The articulated skeleton presents a number of interesting features. ‘The most
noticeable are: The cranium, which is unusually large in comparison with the body ;
the short neck; the long and narrow scapula; and the powerful limbs and feet. The
thoracic cavity is quite large, equaling or perhaps exceeding, that of Cynomys or
Aplodontia. The lumbar region is short and strongly built, as in Aplodontia. The
pelvic region is even stronger than in either Cynomys or Aplodontia, and is in some
respects more like that of Castor. The tail is round, rather heavy, and of moderate
length.
The length of the fore limb of S. fossor, in comparison with the hind limb, is
similar to that in Cynomys, Aplodontia, and Castor. The latter genera have the fore
limb (the feet excluded) from 12 to 14 mm. longer than the hind limb; while S.
fossor has the fore limb 10 mm. longer than the hind limb.
The skeleton of S. fossor has many features similar to those of Cynomys and
Geomys, but, on the whole, it is perhaps more like Aplodontia. There are many
cranial characters, together with the longer and broader ungual phalanges, and the
much longer caudal region, which are very different from Aplodontia.
166 MEMOIRS OF THE CARNEGIE MUSEUM
The specialization of certain parts of the skeleton of S. fossor conclusively proves
that its ancestors had long been a divergent branch from the true forerunners of the
family Castoride. At the present time I cannot confidentially point to any living
rodent of which Steneofiber fossor might be the ancestor. That our fossil shows
highly developed fossorial characters, is clearly evident from the study of the
skeleton. It is plain that the beaver bears no close relation to it, so far as the
structure of the limbs is concerned. The long fore-arm, the short femur, the long
tibia, fibula, and pes, and the greatly developed hallux, in the beaver, are not sug-
gested in S. fossor.
Steneofiber barbouri' spec. nov.
This species is founded upon the greater part of a skeleton, which is worked out
in half relief (Fig. 4), and is still imbedded in its original matrix inside of a weathered
“rhizome” of a Demonelix. It was found in the same horizon, and only a few hun-
dred feet from where the type of S. fossor was discovered. To judge from Professor
Cope’s illustration (‘Tertiary Vertebrata,” Pl. LXIIIL., fig. 22), S. barbouri is of
approximately the same size as S. gradatus, but differs from this species in having
much more expanded zygomatic arches, more quadrate molars, larger tympanic
bullee and occipital condyles, and a much broader occiput.
The smaller size of the skull, and other characters, were at first regarded by the
writer as individual variations, or possibly sexual differences only, which did not
warrant a separation from S. fossor. However, a systematic study of the dentition
Gl ee) BC}
wy!
i
Ss
AY
Fra. 4. Steneofiber barbouri. } natural size.
and of the cranial characters has been made, which shows such marked differences
from the preceding species, that a specific separation was deemed proper. So far as
can be judged from the material at hand, the limbs and feet are apparently very
little different from those of S. fossor, except that they are of smaller size. The
! Named in honor of Professor Erwin H. Barbour, in recognition of his highly interesting work on Demonclix.
PETERSON : DESCRIPTION OF NEW RODENTS 167
dentition and the cranium furnish good specific characters, by which S. barbowri may
be distinguished from the species first described herein. They are as follows :
§ yi
SUPERIOR DENTITION.
The superior incisor is relatively shorter, more curved, and more abruptly
sharpened, with the anterior face somewhat more rounded, than in S. fossor. ‘This
tooth seems to agree more nearly with that indicated in S. pansus (Bull. Am. Mus.
Nat. Hist., Vol. XX., 1904, p. 250), than with that of S. fossor.
The cheek-teeth in S. barbowri have a relatively greater antero-posterior diameter
in the skull than they have in S. fossor, but as in that species they are placed nearly
parallel with the long axis of the cranium. The palate is also relatively broader
in S. barbouri. Very little abrasion has taken place in p+, so that the grinding sur-
face of the tooth has not attained its greatest diameter. The internal enamel invagi-
nations of all the superior cheek-teeth continue to the alveolar border of the maxilla.
This is not the case in S. fossor. Molar * is triangular in section.
Wena Srqgieit, Jel ROWIOL, Ie, @)e Jel, ROWIUUL, Ii, 7A0),
While the general outline of the skull of S. barbowri is similar to that of S. fossor,
a closer study reveals characters, which, besides the smaller size, distinguish it from
that of S. fossor.
The distance from p+ to the incisor is relatively a little less in S. barbowri than
in S. fossor. The palatine area of the premaxillaries, in S. barbowri, is very little
higher than the palate; while this region in S. fossor is greatly arched (see PI. XVIT.,
fig. 1,and Pl. XVIIL., fig. 20). Theinterorbital space in S. barbowri is comparatively
less constricted, and the temporal ridges are apparently less developed than in 8.
fossor. ‘Chere may or may not have been a sagittal crest ; the parietals are crushed
down in the type, so that a positive statement regarding this cannot be made, until
the discovery of more material. The anterior part of the zygomatic arch is rela-
tively less robust, and the tympanic bulla larger than in S. fossor.
The contour of the skull of S. barbowrt is not unlike that of the skull figured and
described as S. pansus (in Bull. Am. Mus. Nat. Hist., Vol. XX., 1904, p. 259). ‘The
excavated basioccipitals, (distinctly a character of Custor), the shorter palate, and the
more vertical and higher ascending ramus of the mandible in the specimen in the
American Museum, however, are clearly characters that distinguish it from S. bar-
bouri. The lattér species has the posterior nares back of m® ; the basioccipital is not
excavated, and the ascending ramus of the mandible is at an angle similar to that
in S. fossor.
168 MEMOIRS OF THE CARNEGIE MUSEUM
The Mandible. —(P]. XVIII., Figs. 18 and 19.) The inferior incisor has the rela-
tively short appearance seen in the superior tooth. The external enamel folding ex-
tends well down on all the cheek-teeth. Premolar z, as in the upper series, is little
worn, and presents two shallow internal, and one deep external enamel fold. The
configuration of the crown is naturally more complicated. This would by further
wear be changed. The first and second molars have about the same relative size as
the tooth in advance of them, which is also the case in S. fossor. The last molar is
proportionally somewhat smaller than in the latter species. The mandible as a
whole is, excepting in size, identical with that of S. fossor.
MEASUREMENTS. 2
Greatestilenothyotiskullteepecccstecsde east ccreceressessestoce cede secrete sere essa taeeerena esas aaeanaer 50 mm.
Greatest width of skull ss
AWA Oi? TTA rcccacoanasdchos906¢ 00060 qasc0n00sn0h09 6590200600, CEoddA nosENILbG0GD s09095 oe0b0s NAoDODSeDENSHENEGE sone £
Wadthofinterorbitalkconstrictlomsern-aeseeececsesceccscne-t-neceneeteesese eames naartl nearer erento ste seeee re acere 11 f
Distance MLromuincisOLojpsseeeees ase ease eee ee ese eee eee eee eee eee scene eecey eeteeer cere seer 16 a
Distance from m® to occipital cond yle..................ecceecee cece ce cec cece eeee eens cece es ceeeeeeeeceree ceeeeeees 19 fe
Width of palate at p* AN ee eae
Width of palate at m®... 6 a
Antero-posterior diameter of tympanic bulla........... 0.02.0... cccec eee eee ee cece este ener eeseneeesteeeeenes Wiley | SS
Transverse diameter of tympanic bulla, not including constricted tube........-..-...:065 ceeeeceeee 10 ss
Greatest trausverse diameter of tympanic bulla including constricted tube .........................-. 16 ns
Transverse diameter of occipital condyle ss
Vertical diameter of occipital condyle
Antero-posterior diameter of incisor Bit 4
Mransverse diam eterorsnclsonsamassssese seca sscereccsc ee csee ened eee acto ene eee e eee eee 3 a
Antero-posterior diameter of p*.........0......0ceeseeeeeeeeee ees BASE BOE CAREH CODE One Unio sac smenoodonG ssabHa CeOSeED Pa) 7 06
Transverse diameters ofip swiss tain seat se cee sce saisieatesn saan eiNe aches ec emincee Sees eee e eeee eaeeee DG
An tero-postenlor diam eter Olea seeeeacesecce ete ee Meee seer eee eee See eee eee ene ce ee eee eee ee ee eee PA yin eee
Transverse diameter of m4.............. .... Bas oe coo, - Bhi} SY
Antero-posterior diameter of m2 Px OC
Mransverse diam etenjol M2 c.csaa-ce mses pedyce ec eeesseeiensecteiseaee ae eecene che ose eee eee eat oeeee eee Bi
Antero-posterion diam eter Ole meee ceess cece esccnece eee ee a eee eee eee ee eee 2 or
Mransversediametersofome scree ces secce ore ee ao se rete cleaner ee ee eee Eee eee een Pe. Of
Mandible.
Greatestilenathtof mandibletsc.necs-pesssscsveesscsnsseeseceerceaencrecsceeccen tarece cere enema eee eee eeee oY
Greatest vertical diameter from coronoid process to inferior angle a
Heightioficoronoids processes: sacs. ocerosor ne es seer e ee oO ee ee eee s
Vertical diameter of jaw infront of p;, including process on the chin....................s0eeeeeeneeees OS ie tae
Vertical diameter of jaw at py..........--..0.65 oNdOSPEDBoCHONdEGoRDGEHOOGSSasCOONSS .sG0b CdeHaasHadsqnEONNS daeunoded 10.5 “
Distancesfromyincisor aly eolus\toypaeseescoorse Rees see te esseeeteereecoetectaacee ea eeeecenceeerese esters 10 ts
Wistanceifromim-tojextreme| backs ofiang] esate sees eee encase eee eee eee 14:5. ‘
PETERSON : DESCRIPTION OF NEW RODENTS 169
Inferior dentition.
Antero-posterior diameter of incisor... ................- BE See SHOE CSE TDOUEE BOE COBEABERRIco cer bocce LoocecucaRsadcKuaG 4 mm.
IETENCTED GHENT GR SHER Raed senode sosasczeosoodaadobtes oobeascooes50ssodacdacdevosensoan euiabaancednaseocoo>ucee 3 i
ANMIOIO HOSE COP CIE WTC G2 | Prrcccnenmboaceaddsacneeco5000 se ude coc suas osesocoOTOLoseCnnS doUsUceee oodosercengcconns 3:00
Transverse diameter of py......... 3 ie
Antero-posterior diameter of m; 3 we
MransverseiameterOlaM mn, DESCRIPTION OF NEW RODENTS 179
Euhapsis platyceps gen. & spec. nov.
Type, Plate XVII., Figs. 5, 6, 7, 8.
GENERIC CHARACTERS.
P-m;3. Teeth hypsodont. Premolar x has the transverse diameter very nearly as great
as the antero-posterior. The teeth in their worn condition have no external enamel inflec-
tions. The length of molars 7 and x together is very little greater than that of the preced-
ing tooth (pz). The zygomatic arches are much expanded. The length and breadth of
the skull is about equal. The occipital surface slopes forward. Parietals broad and
short. No postorbital processes on the frontals. The tympanic bulla is inflated and flask-
like. Bastoccipitals not excavated, but slightly keeled. Mastoid processes directed outward,
instead of downward.
SPECIFIC CHARACTERS.
Incisors strong, somewhat trihedral in cross-section, anterior face very little rounded,
and with a heavy band of enamel. Muzzle heavy and short. Infraorbital foramen
small. Orbit wrreqular and high. Interorbital region moderately constricted. The space
between incisor and p* on premaxillary very little arched. Basi-eranial region back of
cheek-teeth short. Otic bulla with long and greatly constricted tube. Occiput low and
broad. Two small foramina on either side of the median line of the basioccipital. Skull
broad and depressed. Animal somewhat smaller than Aplodontia or Cynomys.
The type (No. 1220) of this new genus and species consists of the skull and por-
tions of both mandibular rami, and was found in the Upper Monroe Creek beds,
near the head of Warbonnet Creek, Sioux Co., Nebraska. This horizon immediately
underlies the Harrison (Demonelix) beds. The specimen was found close to a nearly
complete skeleton of ? Mesoreodon, and in the same sandstone ledge, where the four
skeletons of Promerycochwrus were found, three of which are now on exhibition in
the Carnegie Museum. In the same horizon were also found remains of small car-
nivores of the family Canidx, small species of camels, and turtles, which are not yet
identified.
The general outline of the skull presents similarities to that of Mylagaulus
monodon (Mem. Americ. Mus., 1901, pp. 877-879), and Ceratogaulus rhinocerus (Bull.
Americ. Mus., Vol. XVI, 1902, pp. 291-300). The extreme width of the cranium
in comparison with its length, the wide and forward sloping occipital surface, and
the general depressed appearance of the skull, are especially suggestive of the Myla-
gaulide. A detailed comparison, however, shows that the animal was perhaps more
nearly related to Stencofiber.
180 MEMOIRS OF THE CARNEGIE MUSEUM
THE Superior DENTITION.
Unfortunately, only the incisors are present in the skull. The grinders have
all dropped out. The specimen is injured in the posterior part of the alveolar border
and the region of the pterygoids. ‘The superior incisors are nearly as strong relatively,
as they are in Steneofiber fossor, and are of practically the same pattern: a nearly flat
anterior surface, with heavy enamel. ‘The gnawing surface is gradually worn down
to a broad, thin, chisel-shaped point.
There are only two alveoli (for p*, m+) preserved on the left side of the maxil-
lary ; the anterior one (p*) is smaller than the succeeding one, and does not appear
— judging from its size—to have been occupied by a large tooth, such as is found
in Steneofiber. On the right side, there is no alveole for p+, and on excavating this
side of the maxilla, there was found no evidence of the presence of this tooth. This
may be an accidental character of this particular individual, and may have no
specific value.
oe Tue Inrertor DeEntiTIon.
The inferior incisors are both broken off close to the roots, and there are three
grinders in the left ramus. In cross-section, the incisors are subtriangular, similar to
the upper incisors, with a broad, smooth, heavily enameled anterior face.
The premolar (pz) has very nearly twice the antero-posterior diameter of the
succeeding tooth, which is contrary to what appears to have been the case with the
corresponding teeth of the upper jaw, judging from the alveolus. The width of py
slightly exceeds that of the succeeding two molars. ‘There are three irregularly
placed enamel lakes on the grinding surface of pg; two of them antero-internal and
transverse, the third oblique and postero-external. On the postero-internal enamel
border there is a minute groove. This groove may be the remnant of the third
internal enamel fold, asin Steneofiber. There isa slight evidence of the third enamel
lake as in the latter genus, but in /. platyceps its greatest diameter is antero-
posterior and it is not transversely placed.
The diameter of the first molar is one third greater transversely than antero-
posteriorly. It has two enamel fossettes ; the anterior one is transverse, and appears
to be the only remaining evidence of the internal enamel folding, the posterior one
is curved and oblique, similar to that.on pg. The second molar is similar to the
first in every detail. There is apparently no alveolus for mg, and it is question-
able if there was one. ‘The root of the incisor is very close to the alveolar border in
this region, so that, if the third molar had been present at any time, it must have
been small, and must have had a weak support. The teeth have roots similar to
those in the genus Stencofiber.
PETERSON : DESCRIPTION OF NEW RODENTS 181
THE SKULL.
The general outline of the skull, as has already been stated, is suggestive of the
Mylagaulids. It has, however, a number of characters similar to those of Steneofiber
fossor. There are important characters, which probably, when the superior grinders
are known, will show still more important generic diversity from Stencofiber. The
skull — minus the nasals and the cheek-teeth —is excellently preserved, and
deserves a somewhat detailed description.
The greatly expanded superior border of the premaxillaries indicates a broad
rostrum, which in width exceeds that of the interorbital space, although not to the
same degree as that which obtains in S. fossor. The nasals are comparatively
broader than in Steneofiber fossor, but do not extend any farther posteriorly. About
midway between the anterior and posterior ends, the nasals are broadest, then they
gradually taper to a serrated, rounded point at the junction of the frontals. The
fronto-premaxillary suture appears to be on a line with the posterior end of the
nasals, as is the case in Steneofiber, Cynomys, and Aplodontia. Thus the nasals in
Huhapsis are entirely supported laterally by the premaxillaries.
The frontals are rather short and broad anteriorly, with a heavy rounded supra-
orbital margin. The interorbital space is relatively as wide as that found in the
family Geomyide. The posterior extension of the frontal is also somewhat similar
to that in the latter family, with a slightly more pointed posterior process. ‘There
are no postorbital processes on the frontals.
The sutures in the posterior portion of the skull are not discernible, so that the
forms of the separate bones of this region cannot be ascertained. The parietal is
very wide, and necessarily short, on account of the forward slope of the occipital
surface. The median line is indicated by low ridges, which nearly meet to form the
low sagittal crest. The superior portion of the parietals have only a slight convexity
from side to side, giving a broad and depressed appearance to the skull. The inter-
parietal is not visible. The lambdoid crest, which takes its origin at the postero-
external point (mastoid process) of the squamosal, is highly characteristic in this
form. The extreme anterior slope of the occipital surface places the union of the
lambdoid and sagittal crests forward one fourth of the total length of the skull.
This forward slope is greater than in Ceratogaulus rhinocerus Matth. The latter has
a slope of 30° from a vertical position (Bull. Amer. Mus., XVII., 1902, p. 293),
while Huhapsis platyceps has a forward slope of about 55° from the posterior face of
the occipital condyles to the top of the crest. The lambdoid crest is moderately
high, but very sharp. The entire area of the occipital surface has a gentle con-
vexity from the base of the condyles upward and outward to the top of the
182 MEMOIRS OF THE CARNEGIE MUSEUM
crest, presenting a comparatively plane surface. The transverse diameter of this
surface is twice that of the vertical. The mastoid bulle are only slightly in-
flated. The supraoccipital cannot be outlined by the suture, but is undoubtedly
very large, to judge from the broad aspect of the skull in this region. The basi-
occipital is not excavated as in the beaver; it is somewhat triangular in shape,
with two small perforations, one on each side of the median line opposite the
posterior part of the tympanic bulle. Postero-laterally from these perforations
are the rather large condylar foramina. The occipital condyles are of moderate size,
not very greatly separated inferiorly by the deep triangular notch ; and they are
farther separated from the tympanic bulla than in Steneofiber fossor. The foramen
magnum is of large size and subtriangular in shape. The mastoid process is of
medium size, and has a unique outward and horizontal position, similar to that in
Aplodontia, but is more rounded. ‘The long, constricted tube of the auditory bulla
is supported by the mastoid process nearly to its outer portion — the external audi-
tory meatus. The latter opening is broken away in our specimen, but was perhaps
not of great size to judge from the greatly constricted tube. The tympanic bulla is
much inflated, chiefly laterally and antero-posteriorly ; it is depressed vertically, and
takes up a considerable area of the basicranial surface. The shape of the bulla is
flask-like, with an unusually long and much more constricted neck than in Aplo-
dontia. The genus Hntoptychus from the John Day Miocene has the constricted
neck of the bulla; but the general features of the skull in Huhapsis are entirely dif-
ferent from it, and bear a more general resemblance to Steneofiber.
The Squamosal has a considerable posterior process, which unites with the base
of the mastoid process from which the lambdoid crest takes its origin. In front of
this process, and immediately behind the zygomatic process of the squamosal is a
deep rounded emargination (the postglenoid notch), similar to that in Steneofiber,
Entoptychus, and the recent genus Aplodontia and the Geomyidx. In Huhapsis platyceps
the zygomatic process of the squamosal is of small size and rather short ; the jugal
and zygomatic process of the maxillary furnishing the greater part of the arch.
There is an obtuse, round, postorbital swelling, from which continues latero-
inferiorly a sinuous ridge, which is continuous with the anterior margin of the
zygomatic process. ‘The glenoid cavity is not as distinctly formed as in Fiber zi-
bethicus, and indicates a considerarable lateral motion of the mandible. The anterior
border of the squamosal cannot be accurately determined, since the suture is oblit-
erated at the posterior margin of the orbit. The parieto-squamosal suture is much
lower down on the side of the skull than in Steneofiber fossor.
In the region of the sphenoid bones the skull is damaged. The basisphenoid
PETERSON : DESCRIPTION OF NEW RODENTS 183
appears to be crowded especially posteriorly, where the tympanic bulla is encroach-
ing upon it. The posterior part of the basisphenoid sends a backward projecting
process on either side of the median line. These processes are fused with the internal
face of the otic bulle, and extend to a considerable distance back on these bones.
The pterygoid fossa is quite large, and the external wing of the pterygoid seems to
have reached well back, and is fused with the floor of the brain-case very close to
the antero-external face of the tympanic bulla. The foramina, ovale and rotundum,
seem to be coalescent, as in Aplodontia.
The Jugal is nearly vertical anteriorly, and is a comparatively heavy plate of bone.
In shape and size it is very nearly like that of Castor, the vertical portion just back
of the orbit being comparatively deeper than in the recent genus. At the extreme
inferior jugo-maxillary suture, the arch forms a heavy, rounded, tubercle-like angle.
At the supero-anterior portion the suture is not distinct, but I would judge that
the jugal forms a suture with the lachrymal; the suture of the latter bone is also
indistinct. ‘There is a Jarge lachrymal foramen in the orbit, similar to that of Castor.
The greatest width of the skull is obtained across the posterior part of the jugal.
The extraordinary width and strength of the zygomatic arch is one of the principal
characters of the skull, and recalls such recent forms as the Geomyide and Aplo-
dontia, and also the Loup Fork Mylagaulids.
The postorbital process on the jugal is fairly well developed in Euhapsis platyceps.
The orbit is imperfectly rounded, and is placed high.
The Maxillary. — The zygomatic arch of the maxillary is very similar to that of
Castor, but arises more posteriorly on the maxillary (opposite m+), than in the
beaver (opposite p*). The infraorbital foramen is small and almost entirely hidden
by a vertical ridge or projection from the maxillary like that in Castor. In fact the
skull in this region resembles the recent beavers, with the exception of the much
more produced angle at the inferior jugo-maxillary union on the zygomatic arch. '
The Premaxillaries. —'The premaxillaries are broad, short, and heavy. — Inferiorly
the palatal surface is comparatively broad and has not the long, gentle, and concave
antero-posterior sweep between the alveolar border of the maxillaries and the inci-
sors, which is seen in Steneofiber fossor and the recent beavers. In this respect
Euhapsis platyceps is more nearly like Arctomys monax, which has a continuous,
almost horizontal palatal surface from the posterior nares to the incisors. The
posterior limit of the premaxillaries in Huhapsis is just back of the long, narrow
incisive foramina; thence the suture ascends in an almost vertical line immediately
in front of the preorbital foramina to the superior borders of the zygomatic proc-
esses, and across the face in a slight posterior obliquity to meet the posterior
184 a MEMOIRS OF THE CARNEGIE MUSEUM
process of the nasals. The anterior narial opening appears to have a greater trans-
verse than vertical diameter. The nasals are not present in the type.
Tue MANDIBLE.
The angle and the posterior part of the mandible are broken off. The fragment
shows that the horizontal ramus is rather short, which is in keeping with the short
cranium. ‘The symphysis is long and heavy, inferiorly it terminates in a downward
projecting process, similar to that in Steneofiber fossor and the recent beavers. ‘The
alveolar border is nearly parallel with the long axis of the jaw. The base of the
coronoid process is present and indicates an exceedingly outward pointing direction
of this process, which naturally corresponds to the widely separated glenoid cavities
in the skull. The angle, perhaps, had similar characters to that in Aplodontia.
MEASUREMENTS.
Greatestilensthiofeskuallerecpcnccesadesecereeocea se eccsrseee eters ttes eeeesece metee teee ees eet res ceee es eecinse 60 mm.
Greatest width of skull ............ 0002: ceccseeeeeeeseeeeeeenee oe
Greatest width of occiput at mastoid processes MS
Greatesthwidthiofsmuzzlesnccrssscraconccsecececcsicecccnsecedseer cect eccce rece e eee ee coerce eee eee ees 14 oe
Vertical thickness of skull including occipital condyle to top of sagittal crest............... 2626... 19 os
Vertical thickness of skull including tympanic bulla............0...c0ccccsseeceeececeneceeeeesteneeseueeeens 22 ne
Vertical thickness of muzzle, approximately...............0:c:secceceeeeceeeeceeeceeueeecuecsateceeeneeseneess 12 oa
Length of muzzle from zygomatic process to anterior nareS...............02...0ecececeeseee eee eeeeeeeeees 15 or
Distance frombincisorstOy pz ewan. cacceseweeeeee eccecececseetes ace cote ere eee se cee eee Cena a ee eee eteeeesaes 18 uP
Distance from and including p*, to and including occipital condyle......................cc000eeeeeeee 39 3
Greatest width of occipital Condyles...................ceeee ceceeeneccneceeeeeeeeeeceneeeeeeeeeeueccecsseeeaeeess 15 nS
Greatest vertical thickness of condyles......................sec0eceeeeeeeeeeeee Do ete duisunc eset uncermonteansenaee 4 3
Greatest antero-posterior diameter of incisor.. 45 “
Greatest lateral diameter Of inciSOr......................cceeecseeen esc seeseeceeeecegceeseeneieceesseeceeeeeeeeeeees 5 ee
Mandible.
Total antero-posterior diameter of the three grinders...................0c.c0seceeencceeeeeeeeeeeeeeenee esses 10.5 “
Total antero-posterior diameter of pz..............ssecseceeeeeeeeen cee es $9099 O99 9000060600" Do=9aGecoGaNEEe900s5 5 &
Lotalitransverse diameterOlhpara.so- is larger than in I. montanus but it is a simple, one-rooted, laterally compressed
tooth. The diastema between it and P* is the same. P¥* is about the same in both
specimens. P#* and P* are longer in the present specimen and P* is wider. As in J.
montanus the anterior cusps of the molars are higher than the posterior ones. The
molars have nearly the same antero-posterior diameter in the two species, but in
P. tenuis the transverse diameter is much greater, and the inner portion of the
tooth is very narrow. Taking Matthew’s figure of J. thompsoni as a basis for measure-
ment the teeth of the present specimen are even wider in proportion to the length
than in that species. The last molar is absent from both sides.
The snout is very slender as compared with the size of the skull. The skull
is very broad at the zygomatic arches, but narrows rapidly at their anterior portions
and just in front of the orbit. The palate is broad behind and narrows gradually almost
to a point in front. The posterior narial opening is in the plane of the last molars.
There is a median point at the posterior portion of the palate. Though the skull
was larger, the depth of the malar portion of the zygomatic arch is about the same
asin I. montanus. The depression on the face at the anterior root of the arch has
about the same depth but is smaller, leaving a heavier ridge between that and the
orbit. The nasals are long and slender and are broadest at the anterior portions.
The skull is broader on top than in any other specimen of Jctops that I have seen.
The Mandible. — The greater portion of the left side of the mandible is present.
The coronoid process and the condyle are gone, and the angle is probably not quite
complete.
Portions of the first incisors, canine, and second and third premolars are pre-
served. -- ae
‘8
Fic. 29. The two supposed clavicles lying side by side with their curves approximately parallel. The figure on
the left is that of the bone recovered with skeleton No. 662, that on the right is the bone recovered with skeleton No.
84, a-b, bifid extremities of bones; c, broad flattened ends of bones; d-e, direction assumed by flattened end of
bone from No. 662 ; f-g, direction assumed by flattened end of bone from No. 84.
bones are regarded as belonging to the male copulatory organ then it becomes plain
that the position held by this organ in the two specimens must have been wholly dif-
ferent. Accepting for sake of argument the view that the flattened end of the bone
represents the portion of the os penis which was located in ligamentary attachments,
proceeding from the corpus fibrosum, with its plane placed vertically after the anal-
ogy of Lutra and other animals, then its distal extremity lay in nearly the same
plane pointing downward, with the convex side of the shaft on the right, and the
flattened side of the shaft on the left. Assigning to the specimen from No. 84 the
same position, so far as its flattened, supposedly proximal end is concerned, its distal
262 MEMOIRS OF THE CARNEGIE MUSEUM
extremity must have pointed upward and strongly to the right. If, however, leay-
ing the direction of the flattened ends of the bones out of sight, we place the two
with the flattened sides of their shafts in one position, so that the shafts occupy the
same relative position, then the bone from No. 662 points downward, while the
bone from No. 84 points upward and strongly to the left. It is wholly inconceivable
to the writer that such absolutely dissimilar arrangements should exist in the case
of the penis bone of any animal. Sectional drawings of the shafts of these bones
also show that they are very different from each other. (See Fig. 30.) The bone
Fic. 30. Sections one third of natural size of shafts of supposed clavicles. The upper figures represent sections
taken four and one half inches from the bifid end of the bone, the lower figures represent sections taken nine and one
half inches from the same point. The outlines of the bones themselves are reduced much more than one third and are
wholly diagrammatic. That on the left is from skeleton No. 662, that on the right from skeleton No. 84.
taken from skeleton No. 662 is very rib-like, the shaft having a flattened surface on
one side and a convex surface on the other. The bone taken from skeleton No. 84
has the same flattening on one side and a convexity on the other, though not so
strongly developed. The bifid extremity of the bone from No. 662 shows that a
small portion of one of the branches has been broken off, but irrespective of this
fracture the end does not agree with the bone from skeleton No. 84 either in the
shape or direction of the surfaces of the bifureating extremities. While Mr. Hatcher
was inclined to the view that both bones represented specimens taken from the same
side of the animal, it appears to the writer that they may very well be bones from
opposite sides. Not only are these bones, therefore, asymmetrical, but they differ
in a marked manner from each other to such an extent as to suggest that they did
not occupy the same place in the skeleton, but were most probably from opposite
sides.
Finally, against the theory advanced with great hesitation and rejected by Mr.
Hatcher, but which Baron Nopsca has undertaken to defend, that these bones
might have functioned as ossa penis, is not only the fact of their asymmetry and
HOLLAND: THE OSTEOLOGY OF DIPLODOCUS MARSH 263
the marked differences which exist in the two specimens, which are so great as
to make it appear, that, wherever located in the skeleton, they must have held
opposite, or, at least, very different positions, but the fact, that, so far as is
known to the writer, there is no record in any museum, or in all of the literature
of the subject, of the existence of an os penis among any of the reptilia, living or
extinct, whereas clavicles are found in many reptilian genera. The similarity of
these bones to the os penis of Lutra, which is pointed out by Baron Nopsea, is
curious, but entirely fails to carry conviction with it to my mind, and more partic-
ularly since I have carefully reéxamined the original specimens which are in my
custody. The fact of the bifidity of the penis of Struthio, which is pointed out by
Baron Nopsca, does not appear to the writer to possess great weight. The tracing of
resemblances between the struthious birds and the dinosauria appears to the
writer, as he knows it does to others, to be in danger of being greatly overdone.
Bifidity in the penis is characteristic of the organ in many widely different groups
of animals.
There is another thought or suggestion which has presented itself to the mind
of the writer during his studies, namely, that these bones may possibly have been
sternal ribs connected in some way by strong cartilaginous or ligamentary attach-
ments with the roughened and thick ends of the sternal plates, or imbedded in
cartilaginous or fibrous muscular tissues which do not exist in a fossil state in our
specimens. In this connection reference may be made to the sternal ribs obtained
with a skeleton of Brontosaurus, which Professor Marsh has figured in his work
upon the Dinosaurs.’ It is worthy of note that the length of the longest and most
attenuated of these bones is almost identically that of the supposed clavicle described
by Hatcher. It is furthermore inconceivable to the writer that there should have
been no sternal ribs in Diplodocus. ‘There must have existed a system of central
supports for the lower part of the wall of the huge thoracic cavity.
The attempt. to assign these bones to a position in which they may function as
clavicles is not wholly satisfactory to the writer. To regard them as ossa penis is to
the writer a far more thoroughly unsatisfactory hypothesis, as it was to his col-
league, Mr. Hatcher, who first suggested it. The conclusions of my friend Baron
Nopsea, reached in a labored argument based upon seven propositions, the first five
of which bear only indirectly upon the subject, and the last two of which are posi-
tively incorrect, are in the judgment of the writer untenable. The true position of
these bones is still in doubt, and having left the reproduction of them for a few
days in the position to which I had tentatively and experimentally assigned them
7“*Pinosaurs of North America,’’ p. 171.
264 MEMOIRS OF THE CARNEGIE MUSEUM
in the model of the skeleton, I removed them and turned them over to Dr. Arthur
Smith Woodward, requesting him to keep them until with the progress of discovery
we come to a point where we may be better able to tell what was the role which
they actually played in the osseous system of Diplodocus.
In concluding this brief paper upon the osteology of Diplodocus the writer cannot
forbear making passing reference to the interest which was manifested by the public
in the restoration, which was formally presented to the Trustees of the British
Museum by Mr. Carnegie, on May 12, 1905. He is informed that the number of
persons resorting to the Museum in South Kensington, after the announcement had
been made that the restoration was on view, exceeded the attendance at the institu-
tion at any time since the building was first thrown open to the public. It hap-
pened that at the time of the presentation Parliament was in session, and it was
exceedingly interesting, as well as amusing, to observe the manner in which the
fancy of the knights of the quill and brush seized the work of the paleontologist to
aid them in the field of political caricature. A score of amusing cartoons bearing
upon the political events of the day, in which the Diplodocus was made to do ser-
vice, appeared in the daily papers of England. In the field of commercial adver-
tisement the great reptile has been used, and the writer has discovered a number of
advertisements in which rude representations of the creature have been given, in
order to attract the attention of the public to wares which are described below the
cuts. Not only has the Diplodocus been pressed into service by the caricaturist and
the advertising agent, but the modeler has employed its form for decorative pur-
poses. “ Diplodocus vases,” bearing on their sides figures of the beast in high relief,
have been placed upon the market in London by one of the best known firms en-
gaged in the manufacture of majolica.
Thus the huge reptile, which a score of years ago was unknown, has become
more or less familiar, although there remain a number of doubtful points as to its
osteology to be solved by future study and research.
EXPLANATION OF PLATE XXIII.
The upper figure represents the left side of the skull which bears Professor Marsh’s number
1922 (U. 8. N. M., No. 2673). The mandible on the right side is dislocated and crushed down.
The lower figure represents the right side of the skull which bears Professor Marsh’s number
1921 (U.S. N. M., No. 2672). The specimen is badly broken and crushed in part, but the occip-
ital region is in part better preserved than is the case in the specimen represented in the upper
figure. The drawings of the skull of Diplodocus published by Professor Marsh, as well as other
drawings, which he caused to be made, but which he did not publish for reasons which are plain
to the critical student, show that he utilized both of these skulls in preparing his descriptions and
published figures.
266
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXIII.
Sipe View or SKULLS OF DIPLODOCUS LONGUS MARSH, PRESERVED IN THE UNITED
Srates NatTionat Museum.
EXPLANATION OF PLATE XXIV.
The figure on the left is a view from above of the skull of Diplodocus bearmg Professor
Marsh’s No. 1922 (U.S. N. M., No. 2673). The photograph shows that the right side of the
skull has been more exposed-to crushing than the left.
The figure on the right isa view from above of the skull of Dlplodocus bearing Professor Marsh’s
No. 1921 (U.S. N. M., No. 2672). Only the right half of the anterior portion of the skull is
preserved in this specimen. But it shows the posterior parts of the skull in some respects to better
advantage than they are shown in the skull catalogued by the U. 8. N. M. as No. 2673.
268
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXIV.
Tor View or SKULLS oF DIPLODOCUS LONGUS MARSH, PRESERVED IN THE UNITED
States NatronaAL Museum.
EXPLANATION OF PLATE XXyV.
The figure on the left is a view of the back of the skull of Diplodocus bearing Professor Marsh’s
No. 1922 (U.S. N. M., No. 2673). The basioccipital and the paroccipital processes of the ex-
occipitals are broken and for the most part missing, only the articulating surface of the latter
remaining in situ on the left side.
The figure on the right isa view of the back of the skull bearing Professor Marsh’s No. 1921
(U.S. N. M., No. 2672). The occipital condyle, the outline of the foramen magnum and the
mode of the articulation of the bones forming the back of the skull is revealed more clearly in
this specimen than in No. 2673, but there is much distortion and crushing.
270
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXV.
Back Virw oF SkuLis or DIPLODOCUS LONGUS MArsuH, PRESERVED IN THE UNITED
Srates NarionAt Museum.
pits
a owing
SA
EXPLANATION OF PLATE XXVI.
The upper figure gives a view of the left side of the skull (A. M. N. H., No. 969).
i
EXPLANATION OF PLATE XXVIII.
Fie. 1. Superior View of the Back Part of the Skull of Specimen No. 82, Carnegie Museum
Catalogue of Vertebrate Fossils.
Fic. 2. Superior View of the Back Part of the Skull of Specimen No. 694, American Museum
of Natural History Catalogue of Vertebrate Fossils.
276
“SQ00GO0TdI, (0) STTOMG do
vd NOV 70 MATA wolusang
THAXX aLV1d
TI "I0A ‘WhasnW aIDaNUV) SYHIOWA
EXPLANATION OF PLATE XXIX.
oF
The Plate gives an outline of the caudal vertebra (Nos. 37-73 inclusive) of the tail of a
Diplodocus discovered by Mr. W. H. Utterback on the Red Fort of Powder River lying in serial
order. The figures on the Plate are reduced to one fourth the natural size.
278
MEMOIRS CARNEGIE MUSEUM, VOL. II.
PosTerRIoR CAUDAL VERTEBR® OF JIPLO]
pai Ae
|MEMOIRS CARNEGIE MUSEUM, VOL. II, PLATE XXIX.
| Posrerion CAUDAL Verrenre oF Dirronocus CaRNEGIEI Hatcner, Nos. 37-73 ixcwvstve, rouxp anticuLaTeD ox Ren Fork oF Powner Riyer, Wyomixo, ny W. H. Urrernack,
:
|
i
1q
’ .
VAAN CD OF reat aM nee
UPON THRESEARCHES OF J. B.
HATCHER /
PLATE XXX.
MEMOIRS CARNEGIE MUSEUM, VOL. II.
OOO er eA
RESTORATION OF THE SKELETON or DIPLODOCUS CARNEGIEI Harcurr.
(THE DRAWING IS BASED UPON THAT GIVEN
LATE W ) SEDES. )
yey Vou. IL, P E, WHICH IT NOW SUPER
J. HoLLAND, MADE SUBSEQUENTLY TO THE PUBLICATION OF THAT FIGURE,
>}
J. B. Harcuer anp W. J.
IN THE MeEmorIrs OF THE CARNEGIE Mus M, nua 5
S$ OF
VI., WITH MODIHCATIONS BASED UPON THRESEARCHE
MEMOIRS
OF THE
CAIRN IDEA AiUhS a Ue
VOL. LE NOP.
THE OSTEOLOGY OF PROTOSTEGA.
By G. R. WIELAND.
The first mention of ancient gigantic marine turtles from America was made by
Cope in 1871 ina letter to Professor J. P. Lesley containing an account of a journey
in the valley of the Smoky Hill River in Western Kansas. This letter, as subse-
quently published in the Proceedings of the American Philosophical Society,’
includes a preliminary notice of the huge Niobrara Cretaceous turtle, Protostega
gigas. ‘The type specimen was collected by Cope himself on a bluff near Butte
Creek in the vicinity of Fort Wallace. It was secured in a more or less fragmentary
condition after the manner of the vertebrate collecting of the earlier days of western
exploration, for, in the more extended description given in the Cretaceous Vertebrata
of the West, Cope says this fossil is made up of more than eight hundred separate
fragments.
The various parts of Protostega gigas (type), although mostly in situ, were con-
siderably removed from their natural position. The original specimen is now in the
collection of the American Museum of Natural History. It includes much crushed
eranial elements with portions of the lower jaw, ten nearly free ribs, several verte-
bree, various plastral elements, the shoulder girdle, a humerus, radius, and ulna, and
several metacarpals and marginals, as well as parts of uncertain position. Owing,
however, to the manner in which the ribs lay athwart the rather imperfect plastral
plates, the latter were supposed to be dorsal, and to represent a very primitive con-
dition of carapacial development with large fenestree. The radius and ulna were
1 Vol. XIL., p. 175.
279
280 MEMOIRS OF THE CARNEGIE MUSEUM
not distinctly recognized as such although clearly figured. Owing also to a palpable
numerical error in the measurements of the cranium (‘.50 M. = 242 in.”) the total
length of the original animal was estimated at thirteen feet, and is so referred to in
text-books! The actual length is far less, as will clearly appear in the present
description of a much completer cotype almost exactly the size of the original Cope
specimen. ‘This error was, however, in a sense prophetic, as some of the turtles of the
related Dakotan genus Archelon, discovered by the writer twenty-four years after the
first Protostega, did actually reach, or possibly exceed, the enormous size of thirteen
feet in length.
Protostega remained a very vaguely known turtle until Baur?’ pointed out that
it must in its main characteristics agree with the Cheloniide, and that the plates
Cope supposed were dorsal must be plastral. That such was the fact was later more
definitely shown by Hay,’ who figured the nuchal and hyo- and hypoplastron of
another Niobrara specimen.
The next contributions to our knowledge of the osteology of the Protostegine
were made by the writer, after his discovery of gigantic turtles east of the Black
Hills in the Fort Pierre Cretaceous in the summer of 1895. In the communica-
tions,‘ which soon followed, the structure of the greater part of the carapace and
plastron was made known from remarkably preserved specimens. All the larger
limb bones were also determined and figured ; for the first time, indeed, in the case
of any extinct sea-turtles of America. There immediately followed these papers the
important contribution of Case,’ which, in addition to a careful discussion of the
systematic position of Protostega, added more particularly to a knowledge of the
cranial characters, as based on the description of various crushed, but otherwise well
defined disarticulated elements. The pelvis was also made known.
Further facts concerning the general cranial type in the Protostegine were next
given by the writer® in a paper describing the splendid skull of Archelon, now on
exhibition in the Yale Museum. Later an attempt was made by Williston’ to
restore the tarsus and give the organization of the hind flipper of Protostega, and by
2(a) ‘‘Die Systematische Stellung von Dermochelys Blainville,’’? Biolog. Centralblatt, IX., 1889. (b) ‘‘On the
Classification of the Testudinata,’’ American Naturalist, XXIV., 1890.
3“*On Certain Portions of the Skeletion of Protostega gigas,’’? Field Columbian Museum Publication, No. 7, 1895.
4(a) “ Archelon ischyros, a New Gigantic Cryptodiran Testudinate from the Fort Pierre Cretaceous of South
Dakota,’? Am. Jour. Science, December, 1895. (h) ‘The Protostegan Plastron,’’ Jbid., January, 1898.
5 **On the Osteology and Relationships of Protostega,’’ Journal of Morphology, Vol. XIV. (This publication bears
the date 1897, but did not appear until some time in June, 1898, its presumptively true date. )
®**The Skull, Pelvis, and Probable Relationships of the Huge Turtles of the Genus Archelon from the Fort Pierre
Cretaceous of South Dakota,’’ Am. Jour. Sci., Vol. IX., April, 1900.
7**On the Hind Limb of Protostega,’’ Ibid., Vol. XIII., April, 1902.
WIELAND: THE OSTEOLOGY OF PROTOSTEGA 281
Wieland* to show the carpal organization of Archelon ; but, as will be shown below,
both these efforts are almost entirely in error. From the foregoing review of the
slow progress of our knowledge of the Protostegine it is seen that the third of a
century which has elapsed since Cope’s discovery of Protostega gigas has not sufficed
to bring forth an entirely complete restoration of any single individual of these great
sea-turtles. How welcome then has been the discovery during the past two years
by Mr. Charles Sternberg in the Niobrara Cretaceous of Western Kansas of the
nearly complete specimens of Protostega gigas which permit the present descrip-
tion of the organization of the limbs, the most important of the parts yet unde-
scribed, as well as the least likely to be recovered in complete form. For happily
the elements of the first-secured and completer of these exceptional specimens,
though somewhat crushed, were found altogether, or nearly, in their naturally
articulated position, a condition imperatively necessary to a satisfactory description
of the flippers.
This rare fossil turtle was first briefly mentioned in Science by Professor Osborn ®
* a complete skeleton of Protostega which lay on its dorsal surface with the
fore limbs stretched out at right angles to the median line of the carapace, measur-
ing six feet between the ungual phalanges.” Afterwards it was secured for the
Carnegie Museum by Mr. J. B. Hatcher, who, though he crowded the brilliant
work, which might well have crowned the efforts of a long life, into a short one,
leaves this ripe fruition mixed with a sorrow surely not lessened by the fact that
the hardships of the plains of the Northwest and the Patagonian deserts had all too
plainly left their mark upon him.
Having expressed in conversation with Mr. Hatcher much interest in these more
recent discoveries of Protostega, I was invited to make a study of the newly acquired
material, this arrangement being concurred in by Dr. W. J. Holland, Director of the
Carnegie Museum. But early in July, 1904, when I visited the Carnegie Museum for
the purpose of doing this work, to my extreme sorrow I found, that, although I had a
brief word from Mr. Hatcher shortly before, he was so seriously ill that there could
be but little or no hope of his recovery. And indeed, as everyone feared, it was
but a few days before he passed away. However, it was under such circumstances
a relief to be busied, and Dr. Holland very kindly arranged for and furthered the
initial study of the material on hand.
Furthermore, during the past summer Dr. Holland has added to the collections
first obtained much additional material, also collected by Mr. Sternberg, including
8** Notes on the Cretaceous Turtles Toxochelys and Archelon, with a Classification of the Marine Testudinates,”’
Tbid., Vol. XIV., August, 1902.
®N.S., Vol. XIX., No. 470, p. 35, January, 1904.
282 MEMOIRS OF THE CARNEGIE MUSEUM
one very important specimen, a large Protostega, represented by a quite complete
cranium and lower jaw, accompanied by a humerus, radius, ulna, wrist, and palmar
bones — all in place on the same slab. The acquisition of this specimen has been
as timely as fortunate; for it has removed all doubt as to the carpal and tarsal
organization of Protostega, and made possible the avoidance of errors in the descrip-
tion of the completer specimen, most of the elements of which had been dissociated
from their matrix, and the position of some of them rendered doubtful through the
mistaken zeal of their collector. It will hence greatly simplify our description of
the limb organization of Protostega as based on the free elements of the original and
completer specimen, No. 1420, Carnegie Museum Catalogue of Vertebrate Fossils, if
the more recently acquired and less complete Protostega with its parts In approxi-
mately natural position, be considered first. ;
Protostega gigas Cope (Cotype).
Specimen No. 1421 (Carnegie Museum Catalogue of Vertebrate Fossils). —'This fine
fossil is from the Niobrara Cretaceous of Hackberry Creek, Gove County, Kansas.
The ex situ portions of the original skeleton, which had weathered out and were
secured in more or less complete condition, include the left humerus, radius, ulna,
: & /
Fic. 1. Protostega gigas. Carnegie Museum Specimen, No. 1421, from Niobrara chalk of Hackberry Creek, Gove
County, Kansas. Superior view of skull with the lower jaw and a hyoid, the right anterior border of the carapace and
most of the right fore flipper, all in nearly natural position and still partially imbedded as collected on a single slab of
the chalk matrix. t.
Skull. —pm., premaxillary ; m., maxillary ; n.0., external nares; p.f., prefrontal ; fr., frontal ; 0., orbital border ;
p., parietal; s.o., supraoccipital ; 7., jugal ; q.j., quadratojugal ; sq., squamosal ; q., right quadrate.
Lower Jaw. ae he dentalium ; a., angular; sr., surangular; sp., splenial; (h., hyoid).
Carapace.— N., portion of right ala of nuchal ; 2r., second rib; Zm., 2m., first and second marginalia.
Flipper.— H., humerus; R., radius; U., ulna; r., radiale(?); 7., intermedium ; w/., ulnare; p., pisiform; 1-5,
first to fifth carpalia respectively ; I.—V., first to fifth metacarpalia.
WIELAND : THE OSTEOLOGY OF PROTOSTEGA 283
ulnare, radiale, and pisiform, with fragmentary phalanges, and many broken pieces
of the plastron. The 7 sitw portion consists of the right anterior part of the skele-
ton, and was secured on a single slab of the chalk matrix, in which it still remains
intact, as shown in the accompanying drawing, Fig. 1, by Mr. Prentice. It includes
the following : the lower jaw in oblique inferior view, with the two larger hyoidal
elements ; the skull in superior view, less the left squamosal and with the right quad-
rate disarticulated and its anterior surface up; the right humerus, radius, ulna, wrist,
and palmar elements in superior view, and all in place, except the radiale which
has slipped across to the posterior border of the wrist ; the right ala of the T-shaped
nuchal with the first and second marginals, and the distal half of the second rib, all
in place and resting over the proximal extremity of the humerus.
Although the various bones are more or less crushed, after the manner of most
fossils from the Kansas chalk, the contours are exceedingly good. In the skull, as
simply crushed down in the vertical direction with little distortion of parts, there
appear in clear preservation and united by distinct sutures the premaxillaries,
maxillaries, prefrontals, frontals, postorbitals, and parietals, with the right squa-
mosal, jugal, and quadrato-jugal. The cervicals have in some way been dissociated,
but the skull les in a normal position with respect to the anterior border of the
carapace, from under which the right fore-flipper projects in the normal position,
the hand being bent back pronately over the position of the third marginal, part of
which appears in view.
It will at once be seen what exceedingly satisfactory information is furnished by
the present specimen, as compared with all other examples of Protostega hitherto
found. ‘The more important measurements afforded are the following :
em.
HO wennave-e bxtreme)leng thyoteramuss:-seerernnedoessesrcsienccnccreececsrs stern cere iee se omer 37
Extreme length of median symphySis..............2...cseecee:secesseeeeeceeeeeeeerseeees es 16
Cranium: Extreme length (as crushed flat) measured from end of beak to posterior ex-
NIN? Oi CLONAL OES Rnosaccossoccceccosone caekBe gAcES sconbUeoeDdceR obdocaadHaecaRooRC 58
Medianglengthyofimanialiopeningiec.sscesc-ee-nere=c sen edee carte ceestatseeenelienses eres 7.5
Greatest width of narial opening..............-...0..jcceeceeeeeereseeseeeeeecsceeeeesasee: 5.5
Orbital border of the prefrontal
Antero-posterior length of orbit
Humerus}: Tzenpth ‘across head) :..<.scc5 ee scivesevsecncesssseerceet seccenactesoesee oy seesena cde sceusmees ss
Wadth of distalend i ct. f2 sates seus ccresis so naivesseaneesese
RAC TUS Hs PILEN PED eos c.sseccscs vc cevandeaervesdcatecoectesecseeaecn sce enatenere rn censrocne
Ulna: Length
Carpalia : Width across the five carpalia in place...............0--02cesceerereaneesecneeeseneeseesers 18
Metacarpalia : (Length of 1-5 resp.) ; 7( + or — ), 10.5, 12, 12.5, 10 ( + or —) em.
The above cranial and hand measurements have not hitherto been obtained in
Protostega, whence it is of interest to note, that, allowance being made for the crush-
284 MEMOIRS OF THE CARNEGIE MUSEUM
ing of the specimens of the Niobrara chalk, no wide differences in proportion are
evident on comparison with the much larger turtles of the genus Archelon from the
Fort Pierre Cretaceous. ‘The lack of codéssification of the mandibular rami and
especially the very low radial crest of the latter form will, however, serve to dis-
tinguish it from the fossil before us. But of far greater interest than any question
of generic values that may arise is the fact that the bones of the hand are, with
the sole exception of the radiale, indisputably in place, the most striking fea-
ture being the immense ulnare, and the strong contact of the intermedium with
the first carpale. With these remarks we may pass on, basing our main de-
scriptions on the earlier secured and in some respects better specimen with its several
elements freed from their matrix.
Specimen No. 1420 (Carnegie Musewm Catalogue of Vertebrate Fossils). — The
present specimen of Protostega gigas is more nearly complete than any other as
yet discovered. As originally imbedded in its matrix of chalk, nearly every ele-
ment was present in an exactly or approximately natural position, and recovered
as follows:
1. Portions of the skull, with the crushed lower jaw and two large and well
preserved paired hyoids.
2. The Ist, 4th, 5th, 6th, 7th and 8th cervical, the dorsal, sacral, and eight anterior
caudal vertebrae.
3. The nearly complete series of ribs, including the anterior pair of sacral ribs,
also the neurals, and various marginals.
4. The left shoulder-girdle and proximal half of the humerus, with the right
shoulder-girdle and complete front flipper in place, except the ungual phalanx of
the fourth and the second phalanx of the fifth digit.
5. The pelvic girdle and hind limbs complete and in place, less only the fibu-
lare (if ossified), the ungual phalanges of the right, fourth, and fifth digits and of
the left fifth digit.
6. A nearly complete though very much crushed plastron.
Unfortunately the collector of this surprisingly complete fossil, in an attempt to
remove and separate the bones from their matrix of chalk, mis-marked some of them,
and also made it virtually impossible to either replace more than a very few of the
marginals, or to determine the outlines of any of the plastral elements with exact-
ness. Hence it is not worth while, in view of the marked crushing undergone, to
give figures of any of these parts now, although they will aid in the restoration,
As will be evident to any student of the fossil vertebrates the removal of the fossil
from its matrix in the absence of the necessary knowledge, training, and equipment,
WIELAND : THE OSTEOLOGY OF PROTOSTEGA 285
was ill advised. Such work is difficult enough in the best equipped labora-
tories. However none of the bones of the limbs are broken, and Mr. Sternberg
redeemed himself by discovering and securing in such excellent condition specimen
No. 1421, as just related. The position of the following parts is independently
determinable :
1. The fourth to the eighth cervical, the dorsal and sacral centra, with the
proximal caudal vertebrae and most of the ribs as shown in part in Fig. 2.
2. The shoulder-girdles and elements of the fore-arm ; in particular (as can be
determined by actual close articulation, or by such intervening matrix as the col-
lector permitted in much too small part to remain), the intermedium, first carpale,
metacarpale and its phalanges, and also carpalia 3-5.
3. The pelvic girdle and elements of the hind flipper, in particular the inter-
medio-centrale and tarsalia 1-3, and 4 and 5 (fused), as well as several metatarsals
and phalanges.
It should be stated here that it is claimed that, as figured, no element of either
flipper is actually misplaced, although the extent of the crushing undergone makes
it quite possible that in one or two instances dorsal surfaces have been mistaken
for ventral ones, or vice versa, and that the ends of several phalanges may have
been reversed. Withal it is deemed important to be thus explicit in giving the
manner in which the accompanying text-figures and photographs have been
obtained, before taking up more detailed description. (Because of the compression
undergone it was not thought needful to figure either the lower jaw, cervicals, or
caudals. The dorsal view of the carapace can better be given after the restoration
now being made.
I. Tae Carapace. (Fig. 2.
fo}
With the exception of the T-shaped nuchal and the marginals with dactylate
interior borders so peculiar to the Protosteginee in distinction from all other known
marine turtles, the main features of the carapace are very well represented in Fig.
2. The very thin, almost paper-like series of neuralia is present, though much
erushed down on the underlying neural arches. The ribs are free through at least
the distal two-thirds of their length. The first pair is comparatively slender, it not
being clear as to whether, or not, their distal extremities rested on the lateral alee-
like expansions of the nuchal. The shield contour is, instead of elongate as once
supposed, relatively broader than in either Thalassochelys or Chelone, the ratio of the
length of the dorso-sacral series to the breadth of the carapace being as three to four.
286 MEMOIRS OF THE CARNEGIE MUSEUM
ae * ~
Fic. 2. Protostega gigas. Carapace less the nuchal and the marginal series. Inner view. 1-10, the first to tenth
ribs respectively ; s, the first sacral. Ten dorsal, the two sacral, and the first of the caudal centra are shown. (A trial
drawing preparatory to mounting of specimen. About one-eighth natural size. )
This greater breadth is quite characteristic of Cretaceous turtles, and a distinct ap-
proach to a quite orbicular form like that of Lytoloma.” It is a form, moreover, that
well accords with the powerful and widely expanded flippers now to be described.
I]. Tur Front Fiiprer.
(Plates XXXI. and XXXII., with Figs. 3 and 4.)
The shoulder-girdle of Protostega is robust but presents no strongly marked
peculiarities, save the elongate coracoid which extended back to the pubis as
in Hretmochelys, but not either Chelone or the long-bodied Dermochelys. The
10 Wieland, “Structure of the Upper Cretaceous Turtles of New Jersey : Lytoloma,’’ Amer. Jour. Sci., Vol. XVII.,
September, 1904.
WIELAND: THE OSTEOLOGY OF PROTOSTEGA 287
features of the humerus are, however, highly characteristic, as is shown more
particularly in Figs. 3 and 4, which disclose certain approximations to the
humerus of Dermochelys not so readily discernible in the figures hitherto pub-
Lgl,
‘i ay i he Gi
j Wi ay yy
i } i I) y i if
Fia.3. Protostega gigas. Ental view of alargehumerus. Actual length, 34cm. «, head ; b, radial crest ; c, ulnar
condyle; d, ectepicondyle ; ¢, ectepicondylar foramen ; f, ectocondyle ; g, entocondyle,
lished. It must be noted, however, that crushing has been such as to greatly dimin-
ish the distal breadth and render it uncertain as to whether there is an ectepicon-
dylar groove. Case speaks of a foramen in describing his specimen," though his
figure suggests a groove. I suspect that there may be present a true enclosed fora-
men rather than a deep groove as in Archelon. In the latter the groove is much
further back from the anterior border and the ectepicondylar process correspond-
ingly larger, while the radial crest is not nearly so prominent as in Protostega. (See
Fig. 3.)
1! See foot-note 5.
288 MEMOIRS OF THE CARNEGIE MUSEUM
The carpal and finger organization of Protostega has been hitherto wholly un-
known, and, in fact, the only fossil marine turtle from America in which these
parts have been described is Toxochelys.'’’ In discussing the carpal organization it is
of foremost importance to recall that the itermedium, carpale 1, and metacarpale 1,
as well as carpalia 3-5, are still naturally articulated as in life, and in all particulars
agree precisely with these same elements in specimen No. 1421. Nor is there any
Fig. 4. Protostega gigas. Right shoulder girdle and flipper. 1. Niobrara Cretaceous. SS, scapular ; PC, pro-
coraco-scapular ; C, coracoid ; H, humerus; R, radius; U, ulna; r, position of the radiale ; 7, intermedium ; w, ulnare ;
c, centrale ; 1-5, first to fifth carpalia ; p, pisiform ; m, metacarpal 1; p, phalanx 1; I.—V., first to fifth fingers and un-
gual phalanges. The view is dorsal with the procoraco-scapular rotated into the plane of the paper.
further doubt concerning the identification of the other elements and agreement im
toto with that specimen. As so clearly shown in the figures and photographs, the
triangular-shaped centrale articulates strongly with carpale 1. I have supposed
this was not the condition in Archelon,” but must have been in error. No radiale
was recovered. Bearing in mind that the several carpal elements are somewhat
crushed, no further detailed description of them appears necessary, except that it
should be remarked that the ulnare is of sub-hexagonal outline and relatively very
large, being of the general form seen in Thalassochelys, and especially Colpochelys,
rather than Dermochelys. The ulnare of Archelon is of nearly the same relative size,
12 Wieland, ‘‘ Notes on the Cretaceous Turtles Toxochelys and Archelon, with a classification of the Marine Testudi-
nata,’”’ Am. Jour. Sci., Vol. X1V., August, 1902.
Loc. cit., Wieland, ‘‘ Notes on Toxochelys and Archelon,’’ etc.
WIELAND: THE OSTEOLOGY OF PROTOSTEGA 289
but of rounder and more regular contour. The nearly ovate pisiform is distinctly
intermediate in development between that of the existing Dermochelys and the
Cretaceous congener Toxochelys in which the pisiform is smaller than in any other
distinctly marine turtle.
The first metacarpal is broad, and the first finger short and robust asin Towochelys
(see Fig. 8) and other members of the Cheloniide. Finger disparity is pronounced,
the second finger being little elongate as in Toxochelys, with the third and fourth
fingers of medium and nearly equal elongation, and the fifth fully as elongate as the
second. ‘The first to third fingers bore free claws, but not the fourth and fifth.
In its general features the front flipper of Protosteya agrees much more closely with
that of Toxochelys (Fig. 8) than with that of other forms, as one might well expect.
The existing Cheloninee do not present so close a likeness, because of the peculiar
elongation of the radius and dependent carpal variations, although the boundaries
are much the same, the centrale in particular being in contact with carpale 1 in both
cases. With Dermochelys, in which carpale 1 is small and excluded from contact
with the centrale, the points of likeness are more obscure, although there is no dis-
tinct suggestion in the carpal organization, that the former belongs to an utterly dif-
ferent race. Pisiform development is also more like that of the primitive forms
than in Dermochel Ys.
The relative size of the front flippers as compared with the carapace is great,
since they are not only robust, but have a spread equal to about 33 times the length
of the dorso-sacral series of vertebree. In the existing carnivore Thalassochelys this
ratio is nearly as great, being equal to about three, but falls to two to one in the
algaphagous Chelone. In the carnivore Dermochelys the ratio is about 23 to 1, a
result of comparison rather unexpected in view of the very great length of fingers in
the latter, and accounted for by the great length of the clipper-built Dermochelan
body. There is in the comparison just made the very strongest suggestion that Pro-
tostega, more distinctly than any marine turtle thus far known, hunted prey, which
swam actively, and, bearing in mind other features, was perhaps even powerful.
In my deseription of the front flippers of Toxochelys™ I presented an interesting
tabular comparison of the relative percentages of length of the several elements of
the front flipper and humerus of various Testudinates, which exhibited the general
trend of change since the Jurassic in the development and variation of flippers from
amore generalized limb-type. I now include Protostega in this comparison. The
humerus is in each case considered as having a length of 100, and the finger lengths
as reduced to the same ratio, but including arbitrarily for convenience the meta-
carpals, as follows :
14See Footnote 8.
290 MEMOIRS OF THE CARNEGIE MUSEUM
|
|
|
|
|
|
|
|
|
|
|
|
|
|
mis se se 53 3
I = 5 =o zs =e eS of) ee =
ie 4 = #2 | Sa | 22 aS ee me
Size} bir! Sirs os aS
TB | 4B | FB ee |e
Dermochelisrrrene- teense. eee ease 100 43 39 127 180 209 1738 86 | 23
J BRA TTROB IANS 20000096000020008000005000 100 53 Aue 49 89 12& 105 44 12
Golpochelystimemeren neces eee 100 57 51 56 92 131 107 44 13
WPEGLOSLEQ Ota nano tec ccaise mentees suk 100 60 50 50 86 128 11@ | SO | Wy
PAT CRELO Me deatceaee os eels anes oh 100 54 51 — — “= — — —
Roxochelasreneswese noses.) ceeeeiscost sacs 100 58 | 50 | 51 73 100 104 G® —|- alil
(ONGUTEE Gs: ‘Goonce. ene! conodonsaueasosee6e 100 52 53 50 72 73 55 50 | small
Acichelys = (Eurysternum). ...... 100 57 51 40 54 63 66 51 17
Inspection of the above table shows :
L. Strongly marked radial and ulnar decrease in length.
2. Marked tendency to radial elongation as compared with the ulna, with con-
siderable variation in the length of the radius and ulna as compared with the
humerus.
5. Nearly static length of the first finger in the Cheloniidx, with a sharp increase
in Dermochelys.
4. That elongation of the second finger appeared slowly.
5. Early and persistent increase in the length of fingers three and four.
6. More or less variable tendency to elongation of the fifth finger, with a sharp
increase in Dermochelys, and a suggestion that this finger may have first elongated
in some forms, and then undergone decrease in length.
7. Great and persistent pisiform increase, which began relatively early.
8. Increased finger disparity is mainly codrdinated with depression of the radial
crest.
Ill. Tur Hrinp Fiirrer. (Fig. 5.)
The presence of an entire pelvis and the complete hind flippers, save only the
ungual phalanges of the right second digit, and of both fifth digits is quite all that
the most exacting anatomist could wish. Although as the result of compression in
the matrix the bones are of slightly broader outline than in life, their proportions
are, as in the front flippers, fairly well retained. Likewise, as is so often and so
fortunately the fact in the fossils from the Kansas chalk, most of the articular
facets and surface characters are, as in the other parts of this excellent specimen,
clearly indicated. :
Regarding the pelvis it is only necessary to note that the obturator foramen
must have been completely enclosed by the close contact of the ischium and ento-
pubis on the median line as in Archelon.” The figures given herewith would not of
15 Wieland, luc. cit., ‘‘ The Skull and Pelvis, ete., of Archelon,’’ p. 247.
WIELAND : THE OSTEOLOGY OF PROTOSTEGA 291
themselves be quite clear upon this point were it left unnoticed. The ilia are of
course shown in the lateral view; this, as the result of compression, being the only
view remaining in fairly exact proportion.
Similarly the left fibula, as shown in Plate II., is turned partly around with a
portion of the plastron adhering, the facet for contact with the distal end of the
tibia being thus directed upwards. There can be but the barest doubt that the posi-
tion of all the elements of the hind flippers as assigned in the accompanying Fig.
5 is correct. The presence of paired parts afforded a means of so exactly checking
ACA peas
This;
(hi Wh i
lif
U
WY
Mi
Waly
i jf
Fic. 5. Protosteya gigas. Right pelvic girdle and flipper. ><+. The dorsal or ectal view of flipper with the corre-
sponding ental view of the pelvic elements show in the plane of the paper. P, pubis; F, fibula; ¢, tibiale intermedium
and centrale fused, or caleano-astragalar element ; 1-3, first to third tarsalia ; 4, fourth and fifth tarsalia fused ; m, meta-
tarsal 1; p, phalanx 1; I.—V., first to fifth toes and ungual phalanges.
the determinations as to exclude all possibility of error. Moreover, nearly all the
parts, save perhaps the two ulnares, were imbedded in their matrix in a closely
articulated natural position. And it should be noted that the tibiale, all the tarsalia,
and the first metatarsal and some of the phalanges of both flippers remain so articulated,
the evidence as to their position hence being incontestable. Also metatarsal V.,
though not articulated, or otherwise joined by remaining matrix, is of such char-
acteristic form as would testify amply to its position, even though isolated. ‘The
same may be said of metatarsal 1. The development is in both cases much the
same as in all the marine turtles. Unfortunately, however, no element referable to
a fibulare, which appears to have been present, is now determinable. Nor is a
292 MEMOIRS OF THE CARNEGIE MUSEUM
fibulare represented in the drawing, it being difficult to determine what may have
been its outline.
The first toe is short, the second, third, and fourth of nearly similar moderate
elongation, and the fifth somewhat elongate, as set on the strongly bowed and en-
larged metatarsal V., which is similar to that common to all sea turtles.
The first toe bore a heavy and free claw, and I believe the claws of toes II.-IV.
were also free and clawed, as indicated by their form and the curvature of the an-
tero-lateral or ungual ridges. Reference may be made to Fig. 6, showing the
lateral view of all the ungual phalanges which were all dissociated.
Allusion has been made to Professor Williston’s restoration of the hind limb of
Protostega," and, before pass-
ing on to a general comparison
with other forms, it is neces-
sary to deal with this restora-
tion more specifically. Willis-
ton’s figure is that of a form
so different from Protostega and
all other turtles, that, if cor-
rect, it would indicate the ex-
istence of a new and hitherto
unknown genus, or even fami-
ly, of marine turtles. But this
cannot be; for the form and
proportions of the individual
elements is throughout essen-
Fic. 6. Protostega gigas. Ungual phalanges. f, fingers; h, toes.
Lateral view. 4%. (Trial series not found to necessarily be the exact tially thesame as in Protostega.
pugerand tpeiseries:) The fact is that the arrange-
ment of elements given by Professor Williston affords no approximation, and no
direct hint as to the tarsal organization in Protostega. Metatarsal V. is placed in the
position of metatarsal I., and the bone supposed to be metatarsal V. is probably from
another animal, or else, the metatarsal of the other flipper is much altered by the
compression to which nearly all the turtles from the Kansas chalk have been sub-
jected. The other tarsals are difficult to adjust, although it is in particular likely
that the element placed in the fibular position is a pisiform; for in both form and
proportions it agrees with the pisiform of both of the present specimens, and differs
very markedly in these respects from the true tarsal elements. The phalangeal
16 See foot-note 7.
WIELAND : THE OSTEOLOGY OF PROTOSTEGA 293
formula of 3, 3, 3, 3, 0, as proposed by Williston, is not that of Protostega, in which
the true formula is 2, 3, 3, 3, 3.
The strongly marked angular tubercle of the proximal end of the fibula, as
figured by Williston, reminds one of the similar and similarly situated process on the
femur of the Dinosaur, Camptosaurus. It is not present in any of the specimens of
Protostega known to me, and must at least indicate a new specific form. Should
further examination confirm the presence of this feature, the specimen in the collec-
tion of the University of Kansas, which shows it, should be known as the type of a
new species. And surely its most appropriate name would be one honoring the dis-
tinguished paleontologist who first studied it, and who rendered a very distinct ser-
Fic. 7. Toxochelys latiremis. Left front flipper. Dorsal view. a, head; 6, radial condyle ; c. ulnar condyle; e,
ectepicondylar groove; g, entocondyle; R, radius; U, ulna; Jn, intermedium; Ul, ulnare; C, centrale; 1-5, first
to fifth carpalia ; P, pisiform ; I.—V., first to fifth metacarpals and fingers.
vice by figuring it as best he could in the absence of further material or means of
checking results.
Structurally, aside from the more primitive clawed condition, there is no hiatus
between the tarsal and other features of the hind flipper of Protostega and the exist-
ing sea-turtles, the comparison with Dermochelys being quite as close as any other, in
some respects closer in fact than with a somewhat aberrant although primitive form
like that of Colpochelys (Fig. 8). In Dermochelys and Protostega the relative size and
development of all the tarsal elements is suggestively alike. It is indeed increasingly
difficult to believe that the former genus belongs to an utterly remote and primi-
tively separated line. But unfortunately there is not yet known amongst the fossil
marine turtles a single hind flipper which is sufficiently well preserved to afford
294 MEMOIRS OF THE CARNEGIE MUSEUM
further detailed comparison with Protostega. The writer has, however, pointed out
the fact that the littoral, or semi-marine, turtles of the Cretaceous of New Jersey,
especially Osteopygis and Lytoloma, present many interesting primitive features,
amongst these being relatively longer hind limbs, the inference being a quite clear
one, that in the evolution of the limbs of the marine turtles, in correlation with cara-
pacial and other changes, the hind limbs underwent shortening, and the front limbs
a compensatory elongation. In fact in Osteopygis, a form now known to be very
closely related to the ancestors of the Chelonine, the femur is distinctly longer than
Fic. 8. Colpochelys Kempii Garman. >< 3 Fore and hind flipper of a specimen in the United States National
Museum (ratio of fore to hind flipper, 1.25). :
Fore Flipper. H, humerus with ectepicondylar perforation set well in from the anterior border: R, radius in
natural position revolved somewhat beneath the ulna (U) ; r, radiale, bounded by the radius, the intermedium and first
carpale ; 7, intermedium, wu, ulnare; p, pisiform ; c, centrale bounded by the intermedium, ulnare, and first to fourth
earpalia; I., V., first and fifth fingers.
Hind Flipper. 7, tibia, F, fibula; L., V., first and fifth toes. The tibia and fibula support a single heel or astrag-
alo-calcaneal element, (a) theoretically formed by the fusion of four elements corresponding to the radiale, intermedium,
ulnare, and centrale. The first toe is borne on tarsale I., the second on tarsale II., and the third to fifth on tarsalia IIT.—
V., fused. Metatarsal 1 (m) is the exact equivalent of metacarpal 1, and metatarsal V. has a pisiformoid development.
the humerus, as is also the case in Chelydra, a tortoise which is in many respects sug-
gestive of specialization in some direction similar to that doubtless followed by the
direct ancestors of the marine turtles. In Lytoloma, an upper Cretaceous and lower
Eocene genus closely related to Chelone, the femur appears to have undergone some
WIELAND : THE OSTEOLOGY OF PROTOSTEGA 295
relative shortening. But though a nearly complete carapace has recently been de-
scribed,’ in the absence of further description of the Belgian material, now in process
of elaboration by M. Dollo, we know very little of the limb development in this
form, except byinference. In the case of Towxochelys, which is near to Lytoloma, we
fortunately know the organization of the front flipper within close limits (cf. Fig.
7), while I have but recently described the carapace of a most interesting new
species, 7. Bawri."* In Toxochelys we find a markedly primitive form of flipper, in-
deed the most primitive known, although the humeral contour and finger disparity
indicate an animal capable of navigating the open seas. Also, the carapace of all
the Toxochelyds known has very large fontanelles, suggesting a marine life quite as
decidedly as the flipper; for though the laws of carapacial reduction are as yet only
surmised, it is only in the marine members of the Cryptodira that prominent
pleuro-marginal fontanelles occur.
Because of the facts given it does, however, prove decidedly interesting to find
that even in Protostega of the Upper Cretaceous, the hind limbs yet remain rela-
tively long, though fully developed as flippers. For in Hretmochelys and Chelone as
well as in Dermochelys, the fore flippers vary from about 1.55 to 1.60 times the length
of the hind flippers; whilst in the orbicular bodied Protostega, with a tremendous
spread of front flippers as already pointed out, this ratio falls to 1.30, which is, how-
ever, a little in excess of Colpochelys. In the latter genus which has in some respects
the most primitive flipper organization of any existing Cryptodiran the fore flippers
are but 1.25 times the length of the hind flippers. Doubtless there are many
further interesting numerical relationships not yet discovered. Though the rela-
tively strong pattern and great size of both fore and hind flippers in Protostega and
the related Archelon, taken with the broad body and shortness of the dorso-sacral
series, may not indicate extreme and sustained swiftness, in conjunction with the
cranial features these proportions do go far to confirm the opinion already expressed
that these turtles, so powerfully equipped for both swimming and attack, may well
have hunted actively swimming prey. The Protosteginee plainly included the larg-
est and the most rapacious turtles which ever existed.
"Wieland, ‘‘Structure of the Upper Cretaceous Turtles of New Jersey ; Lytoloma,’’ Am. Jour. Sci. September,
1904.
18 Wieland, ‘‘A New Niobrara Toxochelys,’? Am. Jour. Sci., November, 1905.
296 MEMOIRS OF THE CARNEGIE MUSEUM
IV. Sysremaric Posrrron oF PROTOSTEGA.
CHELONIODEA (Baur).
(Superfamily of the Cryptodira.)
A parieto-squamosal arch ; palatine foramen and free nasals sometimes present,
(Desmatochelydinee); fourth cervical biconvex, with the centra of the sixth to
eighth much more modified in recent than in most Cretaceous forms.
' (A) DeRMOCHELYDID™.
No descending parietal processes; no palatine foramen ; other cranial and limb
characters not remote from those of the Cheloniide ; carapace represented by the
nuchal only, and body enveloped in a leathery hide with an osteodermal mosaic;
no claws. Genera: Dermochelys, Psephophorus, Hosphargis.
(B) CHELONITDA.
Skull with descending processes of parietals, so far as known ; palatine foramen
sometimes present; vomero-premaxillar union usual but not constant; a normal,
though often much reduced carapace and plastron ; nuchal with or without articular
process on under side; claws one, two, or more.
1. Protostegine (Wieland): No free nasals, and no palatine foramina ; obturator
foramen small and enclosed by ischio-pubie contact on the median line, as in many —
land forms; nuchal T-shaped; neuralia and pleuralia thin and investing the ribs
but slightly ; marginalia usually spiniferous on interior borders; plastral elements
of medium development with numerous digitations on both outer and inner border
of the hyo- and hypoplastron ; epiplastron of Trionychoid and Acichelysoid form ;
body enveloped in a leathery hide (?); claws three or more.
(a) Protostega Cope. — Cranial elements nearly as in Archelon ; mandibular rami
coossified ; radial process of humerus strong; front flippers very broad and of a
more distinctly marine type than those of the Toxochelydine (Toxochelys); centrale
in contact with carpale 1; hind flippers relatively very large, and tarsal region most
like that of Dermochelys. Species: P. gigas Cope from the Niobrara Cretaceous of
western Kansas :
Ratio of spread of front flippers to length of the dorso-sacral series. .... 3.2
Ratiovoftront=toybindMilipperssscss sean eer ore aces ace 1.3
First to third fingers free, clawed.
First to fourth toes free, clawed.
WIELAND: THE OSTEOLOGY OF PROTOSTEGA a 297
V. M@rasuREMENTS OF PROTOSTEGA GIGAS.
(All based on the single individual illustrated in Text-figures 2-7 and Plates I.
and II. As most of the elements are much flattened lengths only are given, these
doubtless remaining much the same as in life.)
(A) The Lower Jaw.
Length on median line (estimated, the tip being broken away )......-......-...:cesseeeereees 26 em.
ILGEHO AIO MEE TATA TIS., nosoooqcedno06069 ba00G0 ZooseeooEbosnoB0doe9DDUEDOEBEseEEDoL OGDGBOODeH cao0De Se: 30 ag
(B) The Carapace, Ribs and Dorso-sacral Centra.
Total length of the tem dorsal Centra...............002ceceeees veces teseeseteceeeeccsesetensecrens esses 68 Se
Total length of the three succeeding centra... ae
Length of the dorso-sacral series a
Extreme width of the carapace (exclusive of marginals) as crushed out flat and meas-
ured from tip to tip of the fourth ribs, both of which are complete and are the
NOC EITH Cl WO slo) SOPIES-coccoosaedonsedcan scconsoes toneadocn oacodens sobaed “coqoon secdsquadDoDUDEEETEE 106 Mi
Length of the first to tenth dorsal centra respectively, 5.5, 7.5, 8, 8, 9, 8, 7.5, 6.5, 4, 4 o
Lengths of the first and second sacral and first caudal centra respectively, 3, 3.5, 3.5 ae
IL CTA OF NG) WEL Tell) scccodcaco cosede one oocdcb cgnoadocadodonqode aoasbosuEouGanbedso0eKy eogngoSbdoadoDCBeS ue
ay CO" SACOG THOS coonegssocds voccancedans coumeas Sdakodbaaraaba sbabaquaecad voseod debatbaneogapesead fe
Re UC) HOLA 810 es saacee aoc tes daebaon Gootecoe neanecaasesabon Acco te ae
oa BG") HIRAM Talle) cscnoo edooa0, 6booneaoboccodéde aosde0 gsansa sooDoodoabobcoUBoNGDecHOdpaGdDHoDaDd booed tf
a BG: GISs(H0) TIN): ganna eqeson soe saacaboaccoaaoR one4eodtetedho, Seobbe p9s0n# bo bosUEdoDN pacHabEocGoBUES te
= SOs SASS VEN blot bipeseaacacse scares Naas aries camer cans cchecae. rehedeemt ame ne cease seekers 4
Length of first sacral rib (in place) if
(C) The Shoulder Girdle. \
Distance apart of the two distal extremities (the scapular and the precoracoidal) of
HES OCU EKG OREO DUE re, ce nqbocucbeEboedo pteebbone n960900R bogcoqeID900000 ocobo5 pobBNn voRHEN oRdceDconed 30 oe
Length from interior bow of procoraco-scapular to the glenoid cavity............... 00080550 13 ig
JOA VATE NS CORO. os cocoen sooooonbobeGseedasnoeoc8bn oSsbapqq0qdG0ce cdooHa cdeboducEcttiagendanas coaB008 40 ey
_D) The Pelvic Girdle.
Length of pubis (outer exterior border, on straight line)..........0...6.scceeceee seeeeeeee cece 18 sf
IEF GE THOU ossocosaqcocso0, bydede00 sone nobondebd edaouc baonen sobocaHeNq00HEdG00 HEEHODNAD HoORODESSCCNE SED ees
ILGEPAIN Or TTT, oscospeqnconssnus sooodoess aasbaacoasoscaboUNEEE0Nde DecsDUdeNDOND deopasoetadaeipooud seouG9e00 18 sy
Length of pelvis on median line a EGG)
Width across pubis at broadest point........ ........c..ceecscecscceercseeerecesen teense cecsteeecaeces 48 me
(E£) The Limbs.
The greatest spread of the front flippers (as based on measurement from the approxi-
mate median line of the shoulder girdles to tip of longest finger) Sadacon saaseees boneUaueG 250 ts
Greatest spread of hind flippers uP
(1‘ The Front Flipper.
Length from glenoid cavity to tip of finger IID...............0..-1ece eee nneee eee eee neeee ee eeeeee 106 UK
HOSEN GI! INTERES ossonn08 sees phebs uocoodod pacacebun sdorreaecach yesdonocEOkusacaabequanSseaqee edobonnades 34 Ha
TLSayet ts Oe NAGTIE) opecnconed oeceooe sodes vous obovodanee vonsagoas hasooDotDeosocegacbosean canepnicanenocodsoctiageoN Ct
Length of ulna... a
Length of intermedium.. ................ Jade dhe socacodaso syenguiane ioccbagoadeadacane ssnoaegoecosaLo Sna86s Uigoy. eee
WWF INOS FS AEN Ui eon oben cpacosusonunes ceoeoooconscbe Seat Bbet ecaSRcRR ARE Ve oa me cenapRenD edadseete ahi)
Length of carpale 1..............6.2.006.- Dondeus Boenddo0 _cedubecd ono ses Satsee seneisdosenooseRoconSeaD canetis 5 a
cs
Length laterally across carpalia 3-5, which remains as recovered naturally articulated. 10.6
bo
MEMOIRS OF THE CARNEGIE MUSEUM
ILS MEAN GS WVTENES concooncapcoqssassc0H90009000 HaosoaaDeDDaNHOOdaAIDGDODODDSCOONECEENAD-oSodEDEDoGONDSSERDOECS 10
Greatest width of ulnare............... ....+ : 7
Length of pisiform.............. 02.020 csseee00: cooobbasod00dd0n600004000 00000 aspc0B0des9G000000 s09000000000 6.1
Length, inclusive of metacarpals, of
LETITIELS TAYE) P5.o5000000860060020 260000 snopog acd SsecdapsnsGecoq cadoaonc0B00000000000000000000 600900009600020000 18
Second finger ¢
Third finger.............
(2) The Hind Flipper.
TROUT) NEE Re5 GoooGoon os600000005000000 acoaco0n9 spadoD0000G0050000 so4DOdECaSCORHSG0DG0000000600000000 39
TPT TYEE co3c500. aanedoaon boSEHe .cad000no 006600 DsonDD |dEEnDIADNGoGoANGDDEDSAS: EDsa0dDZ>050HO0o0AGRaG00K0. 30
Length from acetabulum (or head of femur) to tip of finger IIT...............0...:.cceeeeeee 85
IL EMEGAN OF NETNWTRS.c6005 co0050 chons5000 edodonogadeaKas osDbdsaDBBoSOOUGEUDSORDOROSODSOGODOS AEGON 30000 600000000 27
$0" ~ SCO BID Bice sicus sosweuaae sens test ecmesaienceue etree de cloaacaigee net oveteaiten ceaeutectemensceueeeremee geese 20
BO JE OND alliasaeaceous anewcteeraenek cae Seca acne aisione BE CEE Roda GRA CB RCHEABONA cone pan coqsoaaauucceadees 20.5
Gi eeieieeen. 1S) t:) (oh eanenaancandbercokaccborsnd sad deabaner clon adcSercnorad SasdeR eaabeS bce se roseoddnaccoacnecaden rai
Greatest length of the first to fourth tarsalia as they yet remain naturally articulated
(On Mh GORDES OME) sosooondo adoods sooade uoosseannans cosacaeance0s0600Ge0000200600 080400 eee erat 11
Length, inclusive of metatarsals, of
First toe. 18
Second toe 27
Third toe. . 30.5
Fourth toe 31
EXPLANATION OF PLATE XXXI.
Protostega gigas. (Carnegie Museum. Cat. Vert. Fossils, No. 1420.)
(a) Right and left shoulder girdles, with right front flipper and the proximal half-of the left
humerus. :
(6) Pelvic girdle with both hind flippers.
(h) The accompanying hyoids.
The elements are shown as photographed in superior view in approximately the position in
which they were imbedded and crushed out flat in the matrix of chalk. All the elements belong
to a single individual from the Niobrara Cretaceous of western Kansas.
300
‘adO) SPOID FOULSOLOLTL
TXXX ALVTd I] "IOA WNESNWN AIOANUVD SHIOWS
EXPLANATION OF PLATE XXXII.
Protostega gigas. (Carnegie Museum. Cat. Vert. Fossils, No. 1420.)
Enlarged views of forearm and hand (upper photograph), and the foot (lower photograph).
Observe in the upper photograph that the intermedium, carpale 1, and metacarpale 1, as well
as carpalia 2—4 are still articulated as in the matrix ; and in the lower photograph that the tibiale
with tarsalia 1-5 and metatarsal 1 are likewise articulated as found in their normal position in the
matrix.
302
‘ad0OD SPOID FOALSOLOYL
TIXXX aLV1g
EXPLANATION OF PLATE XXXIIT.
Dermochelys coriacece.
(@) Shoulder girdle with front flipper articulated, and the procoracoscapular thus shown
foreshortened. iat ee
: (6) Hind flipper. Superior view.
I., V., the first and fifth fingers and toes respectively.
304
WIXXX aLVTd
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‘TI “1OA ‘NWNASNIN SISANUVD SYTIONAW
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MEMOLRS
OF THE
CARNEGIE MUSEUM.
VOL. 1. oe NO. 8.
NEW SUILLINE REMAINS FROM THE MIOCENE OF NEBRASKA.
By O. A. PETERSON.
In a paper laid before the American Philosophical Society on February 17, 1888,
Professor Cope proposed a new generic name for some Peccaries from the John Day
formation. From his statement it appears that these Oregon forms agree in certain
cranial characters with Hyotheriwm yon Meyer (Palxochewrus Pomel) of the Miocene
of Europe.
The material of this group known at present is still too imperfect, and in some
eases the descriptions are too brief, to determine the validity, or non-validity, of the
generic separations of Thinohyus Marsh! and Bothrolabis Cope.” From the brief
descriptions and figures which Marsh gives of the two species Thinohyus lentus and
socialis from the John Day, the writer infers that that genus and Bothrolabis are
very closely related, if not the same. It is, however, best to await the discovery of
more complete material, before positively expressing an opinion as to the relations
between the different genera, which have been proposed by various authors. In
dealing with these suilline animals, Thinohyus being the prior name used, I propose
provisionally to treat Bothrolabis as a synonym.*
The object of the present paper is (1) to fully describe the principal characters of
1 Amer. Jour. Sci., [X., p. 248, 1879.
2 Proc. Amer. Philos. Soc., Vol. XXV., p. 66, 1888.
3W. J. Sinclair in a recent paper (Bull. Dept. Geol., Univ. Cal., Vol. IV., p. 135, 1905), has already arrived at
similar conclusions.
305
306- MEMOIRS OF THE CARNEGIE MUSEUM
the porcine remains, which have been found in the Loup Fork Miocene of Nebraska
by the field parties of the Carnegie Museum, working from time to time in that
formation since 1901 up to the fall of the present year (1905) ; (2) to carefully com-
pare the osteological characters of the new material with those already known to
occur in the different species of the same genus from the. John Day Miocene of
Oregon ; (8) it is intended, if possible, to more closely correlate and confirm the idea
of Mr. Hatcher as to the relationship of these two formations.* By careful com-
parison and deduction it would seem that at least a portion of the Upper John Day
is represented in the lower horizons of the Miocene sandstones of western Nebraska
and eastern Wyoming.”
TurinoHyvs (B.) suBmHQquANS? Cope. (Figs. 1, 2 and 3.)
See Cope, Proe. Am. Phil. Society, Vol. X VIII., pp. 374-375, 1879; Vol. XXV., pp. 67-70, 1888.
_ This species is represented in the paleontological collections of the Carnegie
Museum by a left mandible, No. 918. The specimen is doubtfully referred to the
above species. It was found in the lowermost Nebraska beds on the Niobrara
River, Sioux County, Nebraska. This horizon is undoubtedly of a much later age
than the uppermost John Day of Oregon. The ramus is not in good preservation,
but enough is present to determine its generic position. The following detailed de-
scription answers closely to that of Bothrolabis trichexnus® Cope, but as there is no
diastema back of p. it is well to temporarily associate the specimen with Bb. swb-
equans. If subsequent study of better material proves this determination to be
erroneous, the name 7. brachyceps may be substituted.
The alveolar border for the incisors is entirely wanting, as are also the canine,
the first, (@f there was a p,), and second premolars. Pg, is damaged, p, and
m, are complete. M, is slightly broken, and mj, has lost its internal face. The
angle of the jaw is broken off. The alveolus for the canine indicates a very
robust tooth which corresponds perfectly to Cope’s description. The alveolar
border is damaged just back of the canine, so that the absence or presence of
pi cannot be certainly determined. Cope states that this tooth in B. subequans
has one root, and has diastemata before and behind it, but the writer is inclined
to think, in view of the considerably later age of the geological horizon in which
the Nebraska specimen was found, that this tooth may in this specimen have
* Proc. Am. Philos. Society, Vol. XLI, p. 118, 1902.
5 In a paper now under preparation by the writer more complete data summing up the paleontological evidence on
this question will be published. (See Annals Curnegie Museum, Vol. 11I., Part 4.)
5 Proc. Amer. Philos. Society, Vol., XXV., pp. 74-77, 1888.
PETERSON: NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 307
been discarded in the process of modification. P, is separated from p, by a short
diastema, and the tooth had two roots. Pgs indicates a tooth with a compressed
simple crown. There is a considerable posterior heel and a slight cingulum on the
antero-internal face of the tooth. P, has two main cusps, proto- and deuteroconids,
well appressed, forming a cross-crest anterior to the middle of the crown, as in
Fie. 1. Oblique internal view of left mandible of specimen No. 913, Carnegie Mus. Cat. Vert. Fossils
3 nat. size.
Fia. 2. External view of left lower mandible of specimen No. 913, Carnegie Mus. Cat. Vert. Fossils. 4 nat.
VA
Cope’s description of Bothrolabis trichenus. Anteriorly on this tooth there is also a
small basal cusp and the cingulum on the postero-exterior angle of the tooth is
thrown into a short fold forming a minute tubercle. The elevated ridge on the
posterior heel, the metaconid, is perhaps more in the middle longi-
tudinal line than, according to Cope, seems to be the case in his
Bothrolabis trichenus. M, isa much worn tooth; the wear of the
triturating face, however, indicates the usual quadri-tubercular
style of tooth. The antero-posterior diameter is but slightly greater
than that of P,. The anterior half of m, is damaged internally
and externally. The hypo- and entoconids are well separated on
the broad grinding face, and just back of these, at the posterior
border of the tooth, there is a small median cusp, the hypoconu-
I a >)
IL lid. In the valley between the anterior and posterior cusps is a
é swelling of the worn surface which indicates a small median conu-
Fig. 3. Superior j- 6 F 3 E z 0
5 lid. On the internal margin opposite to this cusp is a basal pillar
view of left lower
mandible of specimen Which fills up the bottom of the transverse valley. No other
No. 913, Carnegie evidence of a cingulum is present on the tooth. The antero-pos-
Mus. Cat. Vert. Fos-
: : terior diameter of m, is slightly greater than that of m,. The
sils. 4 nat. size. q
increase of the antero-posterior diameter of all the teeth is very
gradual, as p. isonly 5mm. shorter than m,. Ms corresponds to that of Bothrolabis
trichenus which Cope described as haying “two pairs of cusps and a large heel.”
The paraconid can also be outlined on the tooth near the anterior cingulum. ‘This
308 MEMOIRS OF THE CARNEGIE MUSEUM
cingulum is interrupted on the antero-internal angle by a separate minute conulid.
No other evidence of a cingulum is present on the internal side of this tooth. The
external face is broken off. The enamel is slightly mammillated on the internal
margin of the posterior heel ; otherwise the enamel is smooth. The different tuber-
cles of the heel are quite solidly fused together, leaving a broad triangular cross-
valley back of the two posterior cusps.
The symphysis of the lower jaw is quite heavy, and indicates that the mandibular
rami in this region were very broad. On the whole the ramus is rather short
antero-posteriorly and deep vertically. The masseteric fossa does not extend below
the line of the dentition and is indicated on the jaw fragment to be rather deep.
MEASUREMENTS. -
Total length of jaw fragment................cccsccccseeesceeeeeeseceneeeccaececseeeeesceseones 129 mm.
Depthyolpjawilatpeouwetecccecdcsscececctc ree ceeececeecce en aaccecce eee ncres teas 30) ae
Po MoT. Gea nC Eansscerena co bedeapgoddecundaSecon pond bec gascancecoSu cance onuvcobnachoabbarasbocda 43“
Length of molar-premolar series, approximately .............:0c0sceeeeeeecseeeeeceeeeeens 100‘
+) “premolar series, approximately sense) acne -eeeeccccoeeeeee seen = -arcecsesceerss 46 “
CG SO TAAL TNO EY? ES 8IGS) caseoons0cos condo GoaconpbennHo500n000sb4000080 coason sboncooasasecbE009 54s tt
Antero-posterior diameter Of P3..............ceecceeseeceecceeeceetseceeeeeeeevenseeeeeneeeenees U2
; § antero-posterior 14.“
Diameters of P4
Uitransversestence cesta sedate aaa eee ee ECR la I@ 9
Ee ‘Mt { antero-posterior RonbAnacrnaccodscsodn 6xcacondd easSadssdece suapsapEsESoecdbos 1165, 68
cha UiatraTiS Verse ieee ncaa ciare sk eae os eee ere erent ewe ee eee Ts
* © M, § EXTRINSIC eopnade cogon6 onosG0. 56005000 dncoce cosncnangaddocs aqnagsobooE Leen
Uetransversere aes: Sie veetrsue cee cosas eee ete ree octane eae 183 8%
‘ ieMia) san tero-posteni On wenca-cestscce saaecencee esas tees tone oaeeeaeaerene Bp
Thinohyus (Bothrolabis) siouxensis, n. sp.
The type of this species is based on a nearly complete skull and lower jaws (No.
1425, Carnegie Museum Catalogue of Vertebrate Fossils). The specimen was dis-
covered by Mr. J. Grim, Jr., and was presented by him to the writer. It was found
in the upper part of the Harrison horizon on the upper Niobrara River, Sioux
County, Nebraska. he specific name indicates the type locality from which it
came.
Some characters of the specimen might be regarded as of generic value, viz., the
absence of P,*, but it is thought best to await the discovery of more complete skele-
tal material before a final decision regarding its true affinities is had.
*The position and shape of the glenoid cavity and the posterior narial opening are similar to those in Platygonus
leptorhinus Williston.
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 309
PRINCIPAL SPECIFIC CHARACTERS.
T3, Ct, Ps, M3. Occiput high. Region in front of the orbit elongated, with even
slope of the skull, from the inion to the tip of the nasals ; frontal region flat between
orbits, with deep grooves leading from supraorbital foramina to very nearly the end of the
nasals. The posterior margin of the orbit lies immediately in front of a line drawn verti-
cally from the anterior border of the glenoid cavity ; the infraorbital foramen is large,
placed obliquely, and situated above the posterior part of P*. The posterior narial orifice
has an extreme posterior position. P1 single-rooted and quite close to the canine, and
there is a space separating it from P?. P? is in a continuous series with the teeth back
of it. Py is absent.
CRANIUM.
The general contour of the cranium is quite similar to that of the peccary, Dico-
tyles tajacu (Linneeus), especially the region anterior to the orbits. ‘The comparatively
small size of the brain cavity, the high, sharp, and evenly sloped sagittal crest,
together with the simple premolars, are characters which are striking, and at once
separate this genus from the recent peccaries. The occiput is high and has a spoon-
shaped excavation above the foramen magnum. ‘The lateral occipital crest is very
prominent. From the junction with the sagittal crest it descends outward and
downward for a short distance, then almost vertically downward nearly to the con-
dyle, diminishing in prominence in the latter direction, so that the surface of the
exoccipital is comparatively smooth 10 mm. above the base of the occipital condyle.
The posterior temporal ridge forms a weak junction with the lambdoidal crest half-
way between the occipital condyle and the inion. The strong -posterior temporal
ridge points more strongly towards the conditions in the recent peccary and the hog
than to those seen in the John Day species, according to Cope’s description of the
latter. :
The base of the skull is injured. The occipital condyle is present, but the con-
tact is destroyed.’ The condyle is rather small and exhibits characters very similar
to those in the peccary. There was probably a moderately large-sized foramen
magnum, which separated the condyles proportionately less than in the peccary
and the hog. The baso-cranial axis has a much greater angle than is seen in Dico-
tyles tajacu. ‘This is due to the position of the posterior nares, which are relatively
much farther back in Thinohyus siowxensis than in Dicotyles.
The sutures in the cranium under discussion are entirely obliterated so that the
outlines of the elements cannot be traced. ‘The basioccipital is broken posteriorly.
7 The condyle is restored in its approximate position with plaster.
310 MEMOIRS OF THE CARNEGIE MUSEUM
Anteriorly it apparently narrows rapidly, and there is a small rough tuberosity with
a narrow groove through it in the median line. The basi- and presphenoids eyvi-
dently have much less lateral extent than in recent genera, as the space between
the tympanic regions is quite narrow. The foramina opticum and rotundum are
close together ; the latter is quite large. Inferiorly the sphenoids send out strong
wing-shaped lamine. These alee are firmly joined to the squamosal at the internal
base of the glenoid cavity and extend forward and downward, and obliquely out-
ward, to again form a strong contact with the posterior part of the maxillary and
the pyramidal process of the palatine. Some of the John Day species have a sim-
ilar structure in this region according to Cope.* The region of the posterior nares
of Thinohyus siowxensis is one of the chief characteristics of the species. The pos-
terior exit of the orifice is between the anterior part of the tympanic bulle, and its
almost vertical direction is indicated by the large swelling on the sphenoids at the
posterior boundary of the orbital fossa. In the peccary and Platygonus leptorhinus
this orifice is located well posteriorly, but not so far back as in Thinohyus siouxensis.
The posterior narial border is divided anteriorly into three deep triangular
grooves ; one in the middle and one on either side of an osseous septum. The outside
grooves are bounded externally by the deep wall of the inferior alee of the sphenoids
referred to above. ‘The foramen ovale and lacerum anterius are located at the an-
terior base of the tympanic bulla and are hidden from view by the extreme for-
ward extension of the latter. The entire occipital region of Thinohyus siouxensis 1s
narrow in comparison with that of the recent peccary and the hog. The supraoc-
cipital continues upward apparently to the top of the occiput, uniting with the
parietal as in Sus scrofa Linneeus and Phacochewrus xthiopicus Linneus. In the
peccary the supraoccipital does not extend so high and is met by the downward
curve of the parietal bones. ‘The parietals in the fossil are irregularly convexo-
concave. Superiorly, they terminate in the sharp sagittal crest; posteriorly they
are much extended, to help form the greatly overhanging occiput. The zygomatic
arch of the squamosal is very robust, especially behind. Below there is a well-formed
glenoid cayity, similar to that in Platygonus leptorhinus and the peccary, displaying an
oblique saddle-shaped surface like the glenoid cavity in some carnivora. Supero-pos-
teriorly the arch terminates in a high, transversely broad, and thin point, with the
apex gently rounded. At the base of this process (the origin of the posterior temporal
ridge) on the posterior face of the arch is located the rather small external auditory
meatus. Postero-laterally, the frontals show sharp and prominent temporal ridges.
These ridges terminate anteriorly in the rounded and somewhat elevated supercili-
8 Proc. Amer. Philos. Soc., Vol. XXV., p. 72, 1888.
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 311
ary border. The postorbital process is more prominent than in the recent forms
and is trihedral in section. It terminates inferiorly in a sharp point somewhat
posterior to the postorbital process of the jugal. ‘The orbit is open posteriorly by a
space of 12 mm. between the apices of the processes on the frontal and jugal. The
posterior half of the frontals is flat and they are surrounded by the superciliary
borders laterally, and by the prominent temporal ridges posteriorly. Anteriorly
they gradually become convex to meet the maxillaries and nasals. The supra-or-
bital foramina are close together and the deep furrows which lead from them ex-
tend very nearly to the end of the muzzle.
The jugal has a considerable depth below the orbit. ‘The postorbital process is
strongly developed. The latter is heavy antero-superiorly and tapers gradually on
the external and internal faces, forming a rather thin posterior edge. The process
terminates in a subacute point. The orbit is oblique, irregularly oval, and of con-
siderable size. On the anterior border the lachrymal tubercle divides the border
into two emarginations. The one above the process is shallow, but the one below
it is deeper, especially in the external face of the lachrymal. In this rounded
emargination is also located the lachrymal foramen. The zygomatic arch is not so
abruptly terminated at the lachrymal as in the hog, but continues in a gentle
sweep forward to meet the superior border of the maxillary. The temporal fossa
is proportionally much deeper than in the recent genera, while the orbital fossa is
of about the same depth. The spheno-maxillary fossee in Thinohyus siowxensis are
extremely deep, and divided by a thin septum of bone on the median line of the
cranium placed antero-posteriorly and vertically. The posterior opening of the
infraorbital foramen is large. The facial region of the skull shows no sutures.
The alveolar border is low, and does not extend as far back as in the peccary
and the hog. The deep fossa in front and below the lachrymal which appears
in Dicotyles tajacu is much less pronounced in Thinohyus siowxensis. ‘The fossa
in Thinohyus siowxensis is further forward. The infraorbital foramen is large,
obliquely placed, and situated above the posterior part of p*. The alveolus of
the canine presents, on the external face of the maxillary, an abrupt prominent
swelling. Back of this eminence is a faint horizontal ridge, which is an indication
of the much stronger ridge in the recent peccary. Immediately anterior to the
canine alveolus is the deep groove for the reception of the inferior canine.
The premaxillaries are heavy and greatly produced in front of the canine. The
anterior palatine foramina are separated by a strong bony ridge; they are round
and of considerable size. The posterior palatine foramina are close to the alveolar
border and are opposite the anterior part of m.. The palate is long and narrow,
312 MEMOIRS OF THE CARNEGIE MUSEUM
very slightly arched, and has a lightly rugose surface. There is practically no dif-
ference in the width of the palate from i* to back of m?®.
The tympanic region is strongly supported by the postero-internal portion of the
squamosal. ‘The tympanic bulla is of large size, filled with cancellous tissue, and is
closely appressed to the postero-internal angle of the postglenoid process. Anteriorly
the bulla overhangs the posterior nares in a peculiar manner, entirely hiding
from view the lateral borders of the orifice. The postglenoid foramen is situated on
the posterior edge of the postglenoid process, between the latter and the tympanic
region. ‘The foramen lacerum posticum and the condylar foramen are close together
and are situated, the former internal to, and the latter immediately back of, the base
of the paroccipital process. The foramen lacerum posticum is at the posterior
boundary of the tympanic bulla.
MANDIBLE.
The lower jaws of the type are in good preservation, except the posterior part of
the angle which is broken off and lost. The rami are completely codssified leaving
no trace of the symphysis. The latter is long and curves evenly with the backward
sweep of the horizontal ramus. The latter as a whole, is rather long and slender,
diverging only slightly posteriorly,.and has a short constricted area back of the
canine alveole. The alveolar border is nearly horizontal back of the diastema. The
latter has a considerable drop below the line of the border, causing on the
external face of the ramus a constriction similar to that in Dicotyles tajacu. The
external face of the alveolar border is reinforced by a rugose ridge extending nearly
the entire length of the border. This feature is also met with in the peccary. The
internal face of the horizontal ramus is slightly convex. The external surface is
divided posteriorly by a prominent rounded ridge, which disappears below p,, being
replaced by a smooth surface. The inferior border of the ramus forms a sinuous
line, as the angle has a considerable internal flexure. ‘There are three mental
foramina almost in a horizontal line, and situated below and in front of pe.
The ascending ramus is strong, its anterior border, or base of the coronoid proc-
ess, has a prominent ridge extending well forward on the external face of the ramus
as stated above. The temporal fossa is deep ; it is of considerable antero-posterior
dimension, but does not extend below the line of the alveolar border of the jaw.
The top of the coronoid process is broken off, but the base at the sigmoid notch indi-
cates a rather broad antero-posterior surface. The mandibular condyle has a small
antero-posterior diameter, while the transverse diameter is considerable. ‘The
coronoid process rises close to the anterior base of the condyle, forming a small
superior sigmoid notch.
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 3138
MEASUREMENTS.
Length of skull from occiput to end of premaxillary bone.............-.2:-..:0ee-see sadacccno00
‘« from occipital condyle to end of premaxillary bone, approximately...
i ‘“ Jambdoidal crest to postorbital process on the frontal.............. css
we ‘« postorbital process on the frontal to end of premaxillary................+/-.00+ Piles 9%
Total length of alveolar border including premaxillary .........-..--02:.:+ssseeeees eee ceeee eee 156 “
Length of palate from posterior narial orifice to anterior end of premaxillaries ............ PI oS
Mit ML Saag ‘« anterior border of posterior nares to the end of the premaxillaries.. 160 **
Greatest transverse diameter of occiput at lambdoidal crest. .......-...-..2+-seeseeeeeeee eee eeeees 44“
a tt O06 GP DYETIINETED coca nb0a65000 eooHoAeabeb on00B5000 aho00ab0EGaGdaCoDDGSNODDOD Age
WY a ‘¢ of frontals at postorbital processes... .............:..seeeeseeeeeees Coan
yy ot ‘¢ of muzzle at lachrymal bone 85
ie re BG= | SEG. CIEII@s 1100741 noobasudosearonaunbes sedemigessahonnioceeneadece scebestce Sie
e a ‘¢ of one condyle...............+ Lee aa aida Stas Sa beanie ee acl e te Dias
Greatest diameter of cranium at tympanic region...............cc0e6 cecee scene eee eneeeneeeesenens 69 *
Transverse diameter at glenOid Caviby............6..seecnceesees coeccaeecenseeeersecneceersceteecenceeers tig) 98
se ef of palate at m’... op)
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Greatest length of mandible, approximately
Length of mandible from incisor alveolar border to base of coronoid process back of m,.. 148
ILGHERHN Ge Gh NUS Elscodcocensceds6ces ocsacane9dbe boap0b sebon=0N8 o8ADs. Ooadbeoos céuDKE obtlooatnSaon uceceoHODODE og}
IDEN Ot TELUS THA JERS. O81 copes oaccocosascoons sande ascabocoEsdoDDGGdnovSomeboGADbonUOOHoDDY GGen600ed 25 aaa
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SUPERIOR DENTITION.
Puatrs XXXIV.-XXXV.
Unfortunately the incisors, canines, and premolars one and two are lacking in
the type. The alveoli are all preserved and indicate a large pair of median and
smaller lateral incisors; a heavy canine with the antero-posterior diameter greater
than the transverse, the alveoli being oval in outline. ‘The alveolus for p? is
suboyate, with the greatest diameter antero-posteriorly ; it is close to the canine
alveolus and on a line with the internal border of that tooth. P* was undoubt-
edly single-rooted. P? is separated from p* by a diastema of 5 mm. and has
two roots as is indicated by the double alveolus. It had very nearly the same
antero-posterior diameter as p®. The latter tooth is firmly implanted in the maxil-
lary by three roots. The antero-posterior diameter is a little greater than the trans-
verse and the tooth has an oblique position. It is sub-triangular in outline with the
apex antero-internal. The protocone is prominent and the posterior cingulum is
modified into a broad ledge with the greatest diameter postero-internal. The tooth
314 MEMOIRS OF THE CARNEGIE MUSEUM
is entirely surrounded by a heavy cingulum. The enamel is smooth. P* is wider
than long, as in Thinohyus pristinus Cope. The triturating surface of the tooth is
worn down, but indicates a single external cusp, the protocone. The deuterocone
is also strongly developed, and there are heavy anterior and posterior cingula. There
is an external cingulum, while the tooth is smooth on its internal side.
The first and second superior true molars are much worn. ‘The grinding surface
of the former is in its worn condition a large basin surrounded by heavy enamel.
The cones are entirely obliterated, but the external remnant of the transverse valley
indicates a quadritubercular crown. The cingulum on the postero-external lobe is
present, while that on the antero-external lobe is wanting. The internal face has
no cingulum. M?° has strong external and internal remnants of the transverse
valley, extending well in on the triturating surface presenting nearly an antero-pos-
terior 8-shaped basin. There is an anterior and posterior cingulum. Externally
the cingulum is weakly developed and internally the tooth is smooth. M2? has the
greatest antero-posterior diameter of all the teeth in the upper jaw. ‘Phe para- and
protocones of m® are well worn, but indicate that they were well separated and of
considerable size, especially the paracone. ‘The meta- and hypocones are of smaller
size and are closer together. The abrasion on this tooth indicates the former pres-
ence of a metaconule. The anterior half of the grinding face is too much worn to
justify the statement that a protoconule was present. ‘There is a heavy anterior
cingulum and a strong posterior heel. There is no internal cingulum, while the
external is faint. The tooth is broadest anteriorly and tapers rapidly on the exter-
nal face from the paracone to the evenly rounded heel. The internal border is an
almost straight antero-posterior line.
INFERIOR DENTITION.
Puate XXXIV.
All the incisors of the lower jaw and also the canine on the left side are want-
ing. |The right canine is present, but with the top broken off. This is a heavy and
almost vertically placed tooth, very deeply imbedded in the mandible. Its antero-
posterior diameter is greater than the transverse. There are shallow grooves on the
tooth, one on the internal and one on the external face. The anterior border is
narrow and evenly rounded, while the posterior border is broad and more angular,
thus presenting a sub-triangular cross-section. There is a deep oblique abrasion on
the posterior face caused by friction with the superior canine. P, is absent. There
is a long diastema from the canine to py. The latter is in a continuous series with
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 315
the teeth back of it and is implanted in the ramus by two fangs. The single pro-
toconid is greatly developed, taking up the principal part of the crown. Antero-
internally there is a minute cingular conulid ; while posteriorly the cingulum is quite
strong. The external and internal cingula are wanting or only faintly indicated.
Ps, has, as in pg, only the large protoconid with the anterior cingulum heavier and
the posterior heel much better developed than in the tooth in advance of it. The
external cingulum is strong, and internally there is no cingulum, except on the pos-
tero-internal angle, which, together with the posterior ledge, forms a small basin on
this part of the tooth. On py, the proto- and deuteroconids are closely fused, form-
ing a cross-crest, which is much worn in the type. ‘There is a strong anterior base,
and the metaconid is situated almost on the middle of the posterior base. No ex-
ternal or internal cingula appear on this tooth. M, is too much worn for accurate
description. There was evidently no external or internal cingulum on this tooth
M, has the proto- and metaconids equally developed, while the paraconid is a
minute element. Posteriorly the tooth is nearly as broad as anteriorly, the hypo-
and entoconids being well separated. ‘The posterior and anterior cingula are heavy.
The bottom of the transverse valley is filled up externally by the cingulum ; there
are no other external or internal cingula on this tooth. Molars one and two in the
mandible have a quadrate appearance, while ms; has a long sub-triangular form,
which is due to the greatly extended posterior heel. The proto- and metaconids of
the latter tooth are more elevated than the hypo- and entoconids. The postero-in-
ternal angle of the heel is again elevated, so as to form a broad transverse valley
back of the hypo- and entoconids. The heel is solidly fused. The open transverse
valley between the proto- and hypoconids is taken up medially by the hypoconulid,
as is indicated by the wear on the triturating surface. There is a heavy anterior
cingulum. No other cingula are present on this tooth.
MEASUREMENTS.
SUPERIOR DENTITION.
Totalilength ofthe upper!dentition -2.--..:1-..cs.s--se--ctseeiecereceseeeseecesbosatesscee
Antero-posterior diameter of the three incisor alveoli.
y i ( anterO-posteriOr........-2.:--scsssneseseeee senses Pee eeetcetes
Diameters of the canine 1 t
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Antero-posterior diameter of molar-premolar serieS ................cccecseeeceueseeees
cb eG “cc
“ the premolars
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i ; ‘
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(ictransverse:eicrn ty teacamete ne Rh aries Sua chars i 06
316 MEMOIRS OF THE CARNEGIE MUSEUM
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3 cc m3
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INFERIOR DENTITION.
Total length of the lower dentition)...... 0.22.0... .cscesse-00 \eorsseersseeorounoaseneernas= 146 =“
Antero-posterior diameter of incisor alveoli on one side, approximately......... 1125
; .__ f antero-posterior ...... ....-.ee cece eee cece eee cee teres 1s 289
Diameters of canine i 4 pe:
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3
l transverse 1A
In the paleontological collection of the Carnegie Museum is another specimen,
No 1418, Carnegie Museum Catalogue of Vertebrate Fossils (see Fig. 4), which I
refer to the same species described above. The specimen was found on the Agate
Spring Stock Farm, Sioux County, Nebraska, by Harold J. Cook, Jr., and was pre-
sented by him to the writer. The specimen, having (although badly worn)
incisors, a canine, and premolars in the upper jaw, and canines, and all the pre-
molars in the lower jaw, supplements the type in important respects. Although
the skull is distorted, it gives the complete length of the premaxillary and the
nasals, which are damaged in the type. Anteriorly the nasals appear to be narrow,
and terminate in a short subacute point, which slightly overhangs the anterior
nares. In spite of the crushed condition of the skull it is possible to recognize char-
acters like those encountered in the type of Thinohyus siouxensis. In No. 1418, the
specimen under discussion, the excavations for the lower canines are deeper, and
the anterior palatine processes of the maxillaries, immediately anterior to the
canines, appear to be somewhat more constricted, than in the type. The distortion
may possibly be in part the cause of this appearance.
The animal was an old individual, as is plainly indicated by the extremely worn
condition of the teeth. The crown of the median incisor in its worn state has the
appearance of a large, short, subcircular cylinder, with a strong curved fang deeply
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 317
imbedded in the heavy premaxillary. The triturating surface is convex in all direc-
tions by wear. The external margin has received the greatest abrasion. I? is less
worn, is much smaller than i', and has a basal cingulum on the postero-internal
angle. I* is the smallest of the series. This tooth. is more ovate in cross-section,
the antero-posterior diameter being the greatest. The crown is surrounded by
enamel which terminates posteriorly in an acute angle. There is no cingulum.
The principal wear is located obliquely on the anterior half of the crown. All
the incisors are divided by short diastemata. ‘The canine of this species is very
heavy, rivaling that of Dicotyles tajacu. In the fossil the cross-section of the tooth
is a more rounded oval than in the recent genus, but, as in the latter, the posterior
border is narrower than the anterior. ‘There is a long gradual abrasion caused by
friction with the inferior canine on the anterior face, which terminates in a rather
( |
Fig. 4. Anterior portions of the upper and lower jaws of specimen No. 1418, Carnegie Mus. Cat. of Vert. Fossil.
} nat. size.
blunt point, with traces of additional wear on the external face. ‘Twelve mm.
above the point of the right canine, on the external face, is a peculiar broad open
abrasion, extending quite across the tooth antero-posteriorly. On the left tooth is a
corresponding abrasion not nearly so well defined. These rubbed surfaces may indi-
cate the habits of the animal, at least I cannot account for the wear, except by sup-
posing that the canine was used for rooting. P! is separated from the canine by
diastemata, a very short one in front and a longer one behind. This tooth is one
of the chief characteristics of this species. It is a small tooth with the single pro-
318 MEMOIRS OF THE CARNEGIE MUSEUM
tocone, which has received very little wear on account of the absence of p;. There
are no cingula, except on the anterior and posterior angles on the internal side.
The tooth is implanted in a single alveolus. The two roots are closely coalesced,
with a groove indicating the separation of the roots in the species of earlier Tertiary
times. The premolars and molars back of p* are so badly worn that they present
no characters capable of description.
The fragment of the lower jaws which belongs to this specimen (No. 1418)
indicates an animal somewhat larger than the type. The ramus is somewhat
deeper and the diastema back of the canine is longer than in the type specimen.
The diameters of the teeth are practically the same as in the type. The principal
features of the fragment are the solidly fused chin and the absence of p,, showing a
correspondence to the tvpe. The canine reveals a long abrasion on the antero-
external angle which has removed part of the enamel, forming of the remaining
enamel a diamond-shaped surface at the.summit of the tooth.
MEASUREMENTS.
Length of skull from supraorbital foramen to end of premaxillaries ............. 200 mm.
oe“ “~~ infraorbital foramen to end of premaxillaries
«« “alveolar border from m! to end of premaxillaries..... ...............206+-
Antero-posterior diameter of the incisors ................2..22seceeseeee cece ee cece eeu eeees
Di aa antero-posterior
iameters 0
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. ce
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DISCUSSION OF RELATIONSHIP.
It will be safest to await the discovery of more perfect material representing this
group of Miocene peccaries before expressing a positive opinion as to their affinities.
There are two types of the genus Thinohyus, a dolichocephalic and a brachycephalic.
Professor Cope separated Thinohyus decedens, generically from other John Day
forms, giving as the principal character the absence of p?, which is found to be
erroneous.” Judging from the description by Cope and Sinclair, and also from the
°Sinclair, Bull. Dept. of Geol., Cal. Univ., Vol. IV., p. 135, 1905.
PETERSON : NEW SULLINE REMAINS FROM THE MIOCENE OF NEBRASKA 319
illustration '° by the latter author, it would seem that, aside from the erroneous defi-
nition of dental characters by Cope, this species is strongly characterized by its
short skull and its inflated facial region, and may, when additional material is
known, justify its retention in the separate genus Chenohyus proposed for it by the
latter author.
Thinohyus siouxensis is easily distinguished from other known species by the
single-rooted p', which has a short diastema in front and a somewhat longer dia-
stema behind; the unbroken sequence of p? with teeth back of it; the extreme
posterior position of the posterior narial orifice; the large and anteriorly projecting
tympanic bulle ; the posterior position of the infraorbital foramen, 7. e., above back
part of p*, and the absence of p,. The skull of 7. siowxensis is of approximately the
same size as that of 7. rostratus Cope, but the latter species differs from the former
in important characters, viz., the two-rooted p', which is separated from the canine
and p” by diastemata; the latter tooth also has diastemata in front and behind.
The molar-premolar series is relatively shorter, and the infraorbital foramen is
placed more in advance (above the middle of p*). In Thinohyus pristinus Cope p,
is small and as in T. rostratus, two-rooted, ‘the anterior root nearly reaching the
posterior edge of the canine alveolus, . . . p*
11
is wider than long and has but one
external cusp.” M? is longer than wide, which is due to the large heel. The
infraorbital foramen is located above the middle of p?, which is another character
showing similarity to 7. rostratus. According to Sinclair 7. pristinus is farther
characterized by diastemata in front and behind p,. T. trichenus Cope has p* as
12
long as wide, “and [the tooth] has a sub-quadrate base. P? is small, one-rooted,
and separated from p? by a short diastema. Premolars one and two in the mandible
are each separated by diastemata in front and behind. In Thinohyus subequans
Cope the infraorbital foramen is nearly as far back as in T. siowxensis, 1. e., above the
middle of p* in the former, and above the posterior part of that tooth in the latter
species. . swbequans is further characterized by the small two-rooted p’, which is
17 13
“almost entirely within the superior canine. P* is wider than long, as in T.
pristinus and T. siowxensis. In T. subequans there are short diastemata anterior and
posterior to p;." In Thinohyus lentus Marsh” “p, is separated from the canine and
10 L. c., Plate XVI.
"Cope, Proc. Amer. Philos. Suc., Vol. XXXY., p. 73, 1888.
2D. ¢., p. 7.
3 Cope, 1. c., p. 69. (The tooth is located close to the postero-internal angle of the canine—O. A. P.)
™ The lower jaw No. 913 in the Carnegie Museum Collection which is provisionally referred to this species is clearly
that of a short-faced type such as 7’. decedens.
‘5 T. socialis Marsh is imperfectly known. It isa John Day form of rather small size with mammillated posterior
cingulum on m*(?). (See illustration, Am. Jour. Sci., Vol. XLVIII., p. 271, Fig. 25, 1894).
320 MEMOIRS OF THE CARNEGIE MUSEUM
Pe by diastemata of respectively 8 mm. and 10 mm.” Thinohyus osmonti Sin-
clair is characterized by the long diastema in front of p,, which is double-rooted
and in an uninterrupted series with the teeth back of it. ‘“ P? is a single-crowned,
double-rooted tooth, separated from the canine and p? by diastemata.” The infra-
orbital foramen appears from the photographic reproduction given by Sinclair to be
above the posterior part of p®.
SUMMARY.
The study of the remains described above may be summarized as follows:
1. The animals represented by the specimens in the Paleontological Collection
of the Carnegie Museum are more modified than those representing the John Day
Miocene in other collections, and the former may, when more completely known
be regarded as new genera.
2. The horizon in which the remains were found is the upper part of the Harri-
son horizon, which is regarded by Hatcher” as filling the hiatus between the Upper
and Lower Deep River Formations. The uppermost Arikaree, or the Monroe Creek
horizon, which is regarded by the same author as equivalent to the Upper John
Day, has not as yet yielded any remains of peccaries. When such remains are
found in the lower horizons of the Miocene in this locality, they will undoubtedly
reveal characters more closely allying them to the John Day forms, thus differing
from Thinohyus siowxensis.
The writer wishes to express his thanks to Dr. W. J. Holland, Director of the
Carnegie Museum, for kind suggestions and assistance in the preparation of this
manuscript for the press.
The illustrations are from drawings made by Mr. Sydney Prentice.
CARNEGIE MUSEUM, November 24, 1905.
1’Sinclair, Bull. Dept. of Geol. Cal. Univ., Vol. IV., p. 110, 1905.
7 Sinelair, /. c., p. 139.
18 Proc. Amer. Philos. Soc., Vol. XLI., p. 118, 1902.
EXPLANATION OF PLATE XXXIV.
Upper figure. Right side of skull and lower jaws of Thinohyus siouxensis. (Type) No.
- 1423, Carnegie Museum Catalogue of Vertebrate Fossils.
Lower figure. Superior view of the mandible Thinohyus siouwensis. (Type) No. 1423.
Figures 3 natural size.
322
MEMOIRS CARNEGIE MUSEUM, VOL. II. : PLATE XXXIV.
on i
THINOHYUS SIOUXENSIS PETERSON.
EXPLANATION OF PLATE XXXYV.
Figure on left. Superior view of skull of Thinohyus siouxensis. (Type) No. 1423, Car-
negie Museum Catalogue of Vertebrate Fossils.
Figure on right. Palate view of same skull. (Type) No. 1423. Figures 3 natural size.
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXXYV.
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THINOHYUS SIOUXENSIS PETERSON.
MEMOIRS
OF THE
C AORN JEG JU) MNOS Ue
WOlbe JOG IN@, (8)
NOTES ON OSTEOLOGY OF BAPTANODON.
Wire A Description oF A NEw SPECIES.
By CHarites W. GILMORE.
During the season of 1903, while in the service of the Carnegie Museum, the
writer collected from the marine beds of the Jurassic in the Freeze Out Mountains
in Carbon County, Wyoming, quite a complete Baptanodon skull (No. 1441") and
lower jaws associated with other parts of the skeleton.
Through the courtesy of the Director, Dr. W. J. Holland,’ the writer was accorded
the privilege of studying this specimen, thinking perhaps it would give some infor-
mation regarding the obscure points in our knowledge of the anatomy of this
interesting reptile.
The skeleton like nearly all of the American Jurassic Ichthyosaurians was
enclosed in a very hard limestone concretion® much seamed and cracked by expo-
sure to the elements.
Since publishing a paper on the Osteology of Baptanodon?* additional discoveries
have thrown new light on the structure of this animal. It thus becomes necessary
'Card catalogue number, Department of Vertebrate Fossils of the Carnegie Museum.
2 My acknowledgments are especially due Dr. W. J. Holland, and I take this opportunity to express my appreciation
of the many courtesies extended during the preparation of these notes. The text-figures were drawn by Mr. H. W.
Hendley, of the U. S. National Museum, Washington, D. C.
5 A recent letter from W. H. Reed, of the Museum of the University of Wyoming, informs me that during the past
summer he collected a very complete Ichthyosaurian skeleton, which was quite free from the refractory matrix men-
tioned above. He regards this specimen as coming from a lower horizon than those found in the concretionary layer.
‘MEMOIRS OF THE CARNEGIE MuSEvM, Vol. II., No. 2, August, 1905.
325
326 MEMOIRS OF THE CARNEGIE MUSEUM
to make certain alterations and additions to the description and drawings then
given, especially those relating to the elements of the skull and lower jaws.
The parts of specimen No. 1441 recovered consist of a skull and lower jaws
lacking a median section of the rostrum, the codssified atlas and axis, third and
fourth cervicals, besides vertebree in various stages of preservation from other parts
of the vertebral column. With the exception of the proximal end of one humerus,
the ulna, and a few paddle bones, all other elements are missing.
Tar SKULL.
The cranium, although subjected to severe pressure, which has somewhat dis-
torted the outlines of the skull, appears very similar to those previously described
in the paper above mentioned, with the exception that it pertains to a somewhat
smaller individual.
The superior view shows the transverse displacement between the anterior end
of the postfrontal and the posterior end of the nasal, and, as breaks occur in identi-
cally the same place, and take the same transverse direction in two other crania
(Nos. 603 and 878) in the collections of the Carnegie Museum, the writer now as-
sumes this displacement as following the line of suture, and in the corrected draw-
ing (See Plate XX XVII, Fig. 1), they are indicated as absolutely determined. The
suture separating the premaxille at the anterior third of the rostrum is plainly
shown in this specimen. Another point of interest is the wide prefrontal developed
in this individual.
Viewed laterally this skull sheds new light on the posterior termination of the
premaxillary. In Baptanodon the posterior end of the premaxillary is somewhat
forked as in many forms of the Ichthyosauria. The lower posterior branch extends
back under the nares, laps over and hides from a lateral view most of the anterior
prolongation of the maxillary. (See Plate XXXVI.) The longitudinal channel
along the side of both the premaxillaries and dentaries, just above and below the
dental grooves (See Fig. 3), has many more nerve pits for the entrance of nerves to
the teeth than was indicated in the first restoration of the skull of Baptanodon. Marsh was mistaken in his identification of the limb in the
discus (No. 603), one i : f : =
half natural size, a, type of B. discus (1955) ° as a posterior extremity, a question dis-
articular end; (2), cussed in my previous paper. ‘This element resembles somewhat
view of articular end “7: as 3 5 : :
the ilium of Yoretocnenus californicus, a ‘Triassic Ichthyosaurian
of same; (3), cross sec- f t
tionofthebrokenend. described by Dr. J. C. Merriam.
M&EASUREMENTS.
No. 603. Greatest width antero-posteriorly of acetabular end...............-. 45 mm.
* 603. gs & a “« fractured end ..............-.- 27 mm.
Baptanodon robustus sp. noy.
The type material of this species includes a fairly well-preserved pectoral girdle,
a series of ten cervical vertebree beginning with the atlas; a second series of eleven
vertebree from the anterior dorsal region commencing back of the point where the
diapophysis becomes distinct from the neurapophysial articular surface ; a third sec-
tion of eleven posterior dorsals beginning just back of the first vertebree having the
diapophysis and parapophysis united to form a single node-like articulation for the
single headed ribs of this region. The fourth and Jast section contains parts of
twelve anterior caudals. These show the rapid decrease posteriorly in the size of
the centra, which has been previously pointed out by Knight.’ :
This specimen, No. 919, is from the Red Fork of Powder River, Big Horn County,
° Catalogue number of Museum of Yale University.
‘Knight, W. C., “Some Notes on the Genus Baptanodon with Description of a New Species,’? Amer. Jour. of
Science (4), Vol. XV., 1903.
GILMORE: NOTES ON OSTEOLOGY OF BAPTANODON 300
Wyoming, and was collected by Mr. W. H. Utterback from the marine beds of the
Jurassic in 1902. It is now in the collection of the Carnegie Museum.
The large size of all of the vertebrae and the long straight border of the external
ends of the coracoids for articulation with the humeri appear to indicate a new
species for which I propose the name Baptanodon robustus. Although the material
at hand is insufficient for a satisfactory diagnosis of this form, yet the dimensions
of the parts preserved, particularly the vertebree and ribs, show this to be the largest
member of the Baptanodontide. A right coracoid, No. 1953, in the Museum of Yale
University shows the same long straight border on the external end and may per-
tain to this species.
Vertebrx. — The coalesced atlas and axis are very large, and this is the first indi-
vidual I have examined which shows the suture separating these vertebree. (See
Fig. 12, 8.) On the lower border the atlas is 835 mm. wide antero-posteriorly, the
axis being 830 mm. ‘The great disparity in length antero-posteriorly between the
Fic. 12. Atlas, axis, third, fourth, fifth, and sixth cervical vertebrae of Baptanodon robustus. Type specimen, No.
919. One half natural size. 1, Coalesced atlas and axis. 2, 3, 4, and 5, third, fourth, fifth, and sixth cervicals d.,
diapophysis ; p., parapophysis ; s., suture between the atlas and axis ; z., hypapophysis.
lower and upper parts of the combined centra appears to be distinctive of this species.
The hypapophyses on these vertebrae are very indistinct and no attempt has been
made to indicate them in the drawing. (See Fig. 12.)
On the third and fourth cervicals the parapophysis appears to be indicated by
hardly more than a slight antero-posterior swelling on the side of the centra. In
this respect it resembles somewhat the corresponding elements of B. marshi. The
succeeding vertebrae have well developed parapophyses on the anterior half of the
centra. ‘The diapophyses on all of the vertebrae preserved are very robust and con-
fluent with the articular surfaces on the dorsal surfaces of the centra. The fourth
99
oo- MEMOIRS OF THE CARNEGIE MUSEUM
to the ninth cervicals have a vertical ridge connecting the di- and parapophyses on
the anterior margin of the centra. This character may also prove to be a feature
of this species.
The anterior or median dorsals have two well defined apophyses (See Fig. 13)
placed well down on the anterior margins of the centra. The diapophysis is the
Fic. 13. Lateral view of anterior or median dorsals of Buptanodon
robustus (No. 919). Typespecimen. Onehalfnaturalsize. d., dia-
pophysis ; p., parapophysis; s., spinous process; a.zyg., anterior zyga-
pophysis ; p.zyg., posterior zygapophysis.
larger of the two. In these verte-
bree the pedicels of the upper arches
are extended antero-posteriorLy,
being nearly as wide as the centra
upon which they rest. The spines
are somewhat compressed later-
ally, and very high. The arches
in what is considered the anterior
dorsal region are held together by
single zygapophyses, which are of
considerable extent obliquely.
The pedicels do not project later-
ally asin some of the Ichthyopter-
ygians but form a smooth surface
with the upper lateral surfaces of
the centra.
The series of posterior dorsals
appear very similar, except in size,
to those from this region of B. dis-
cus. The single apophyses pos-
teriorly recede to the lowermost
border of the centra as in the other
species.
The parts of caudals preserved,
besides showing the rapid decrease
in size posteriorly, appear to have
parts of several chevrons retained in the matrix near them and furnish the first evi-
dence of these bones in Baptanodon.
All of the vertebree are deeply biconcave as in other members of the genus.
GILMORE : NOTES ON OSTEOLOGY OF BAPTANODON BiB)
MEASUREMENTS OF VERTEBR&Z OF NO. 919.
Atlas and| :
Cervicals | axis | 3d 4th 5th | 6th 7th | 8th 9th 10th | 11th |
Greatest length of centrum in mm. 57 34 34 30 35 35 35 35 b — | =
Greatest width of centrum in mm. 100 100 | 100 90 80 70 60 60 — = =
Anterior dorsals | 1 2 Sal 4 5 6 g | 8 9 1@- || mi
Greatest length of centrum in mm. — | 4) 45 | 45 43, 43 43. a) 3a | 48a | 42a |
Greatest height of centruminmm.| 1006 | 1206) 110 | 1106; 110 | 110 | 101 | — | —~ | — Vires
Greatest height with spinein mm.) 2106 | 245a) 245a) 2106) — | — _ —- | —}— ] =
Posterior dorsals 1 2 3 4 5 6 7 8 9 10 11
Greatest length of centrum in mm. 37 37 38 40 40 3 45 45 50 48 50
Greatest height of centrum in mm. 100 104 | 104 | 104 | 104 | 106 | 106 | 102 99 99 99
a@ = approximate, b = broken.
Tue PrcroraAL GIRDLE.
The elements of the pectoral girdle of No. 919 are finely preserved and entirely
free from distortion. The scapule lack about half of their upper free ends and the
coracoids their posterior borders. (See Plate XX XVIII.) These missing parts have
been restored from the pectoral girdle of B. discus, No. &78 in the Carnegie Museum.
The coracoids are broad subquadrangular bones which join one another medially
by large elliptical facets. [The superior surfaces (Plate XX XVIII., Fig. 2) are flat-
tened, although both elements are gently inclined toward the median line. The
inferior surfaces are concave transversely and convex antero-posteriorly. The lateral
borders are especially thickened forming a heavy articular face for the scapulee and
humeri. This outer articular end is divided into two unequal faces meeting in an
obtuse angle. The more anterior and smaller one of the two is for the scapula, and
looks outward, forward, and obliquely upward. The larger and posterior surface
forms the greater part of the glenoid cavity. This border is much straighter and
longer antero-posteriorly than on the coracoids of B. discus. It’is very slightly
conyex from above downward and is covered with tubercle-like eminences. This
articular end is supported by a broad neck formed by a deep notch on the anterior
margin. The anterior border from the inner notch thickens rapidly as it recedes
posteriorly to form the intercoracoidal facet. (See Plate XX XVIII., Fig. 1.)
The scapule are moderately long bones the upper halves being narrow with
nearly parallel sides, the articular end is broadly expanded’ antero-posteriorly and
thickened on their posterior margins where they enter into the formation of the
glenoid cavity. The articulating face for the coracoid joins this border at an obtuse
angle.
The scapule extend outward, upward, and forward. Viewed longitudinally
the inner surface is slightly concave as they curve up to lap over the side of the
ribs. The articular ends on the dorsal surfaces are concave antero-posteriorly.
336 MEMOIRS OF THE CARNEGIE MUSEUM
The anterior border of the parts preserved is slightly rounded, but, as it approaches
the articular end it widens into flattened oblique surfaces which look forward and
upward and probably represent the points of attachment for the ligaments which
held the clavicles in position. (See P]. XX XVIIL., Fig. 1.) The ventral surface of
the scapule antero-posteriorly at the expanded articular ends is somewhat concave.
Between the point of attachment for the clavicle and the articulation with the
coracoid the internal border is especially compressed and remains free.
The clavicles and interclavicular bones were not found with this specimen.
MEASUREMENTS.
No. 919. Greatest width of coracoids, transversely...... ...-..cccseseeeceseeeeee 430 mm.
BS Bil, sf length of articular surface for scapula....................0..- 85
«919. ae “of straight external border....................0-sseeee INO),
35 Og, width of scapule, antero-posteriorly.................2::000+ il7f5j, 40
6 Og, ee length of glenoid articular face
ComMENTS ON Recent LITERATURE.
In discussing the age of the Baptanodon Beds Dr. 8. W. Williston® speaks of the
relationship of Baptanodon and Ophthalmosaurus and makes the statement that the
two species of Ophthalmosaurus are probably not congeneric and “it is also apparently
quite true that Baptanodon seems to be as closely allied to the Cretaceous as to the
Jurassic species.” In a previous paper’ I have shown that O. icenicus (the Jurassic
form) and Baptanodon are very closely allied, while O. cantabrigiensis is not similar
in any respect.
The discovery the past season (1905) by Mr. W. H. Reed of an Ichthyosaurian
in the Benton Cretaceous is of considerable interest as showing the occurrence in
North America of this group at a much later period than hitherto supposed.
Dr. J. C. Merriam, who has examined the fragmentary specimen collected by
Reed, says: “The centrum is very thin antero-posteriorly and in this respect some-
what resembles the corresponding centra in Baptanodon discus. . . . When more
material is available it will be interesting to learn whether this form really repre-
sents a true Ichthyosaurus or possibly a more highly specialized form of Baptanodon
than those we know from the Baptanodon beds. Should it be Baptanodon, it will
* Williston, 8. W., ‘‘ Hallopus, Baptanodon and Atlantosaurus Beds of Marsh,’’ Jour. of Geology, Vol. XIII., No.
4, May—June, 1905.
*Gilmore, C. W., ‘‘Osteology of Baptanodon,’’ MmmMorrs OF THE CARNEGIE Museum, Vol. II., No. 2, August,
1905.
Merriam, J. C., ‘“ The Occurrence of Ichthyosaur-like Remains in the Upper Cretaceous of Wyoming,’’ Science,
N.S., Vol. XXII., No. 568, pp. 640-641, November 17, 1905.
GILMORE: NOTES ON OSTEOLOGY OF BAPTANODON orl
probably show some extreme specializations, as the time separating the Benton from
the Baptanodon beds is considerable.”
Moreover, it would be of interest to note the presence of this group in the Cre-
taceous of this country, which has already been shown to be the geological range of
the closely allied European genus Ophthalmosaurus.
U.S. NatTionaL MusEuM,
February 6, 1906.
EXPLANATION OF PLATE XXXVI.
Side view of skull of Baptanodon discus Marsh (No. 878). One fifth natural size. Restored.
ag., angular; d., dentary; j., jugal; /a., lachrymal; ma., maxillary; na., nasal; nar., narial
opening; oce.c., occipital condyle; pa., parietal; pmx., premaxillary ; prf., prefrontal; pff.,
postfrontal ; pto., postorbital ; q,j., quadrato-jugal ; qu., quadrate ; s.ag., surangular ; s.¢., supra-
temporal ; sc/., sclerotic plates ; sta., stapes ; spl., splenial ; sq., squamosal.
we)
co
oo
MEMOIRS CARNEGIE MUSEUM, VOL, II.
PLATE XXXVI.
S=>
7 =__ Sy
SSF
Stipe View or Skuty or BAPLANODON DISCUS Marsu (No. 878).
SOME SUTURES DRAWN AFTER No. 1441.
ones
Natura Size. Resrorep.
EXPLANATION OF PLATE XXXYVII.
1. Top view of skull of Baptanodon discus Marsh(No. 878). One fifth natural size. Restored. _
art., articular ; ex.oce., exoccipital ; fr., frontal ; na., nasal ; nar., nares; occ.c., occipital condyle ;
pa., parietal ; pin., pineal foramen ; pmx., premaxilla; prf., prefrontal ; ptf, postfrontal ; s.ag.,
surangular ; s.occ., supraoccipital ; s.¢., supratemporal ; s.t,f., supratemporal fossa ; sq., squamosal.
2. Inferior view of skull of Baptanodon discus (No. 603). One fifth natural size. Restored.
ag., angular ; art., articular ; b.oce., basioccipital ; b.s., basisphenoid ; cor., coronoid ; d., dentary ;
ipt., interpterygoid vacuity ; occ.c., occipital condyle; pl., palatine; prs., presphenoid ; pt., ptery-
goid ; spl., splenial ; v., vomer.
BA()
MEMOIRS CARNEGIE MUSEUM, VoL. II.
PLATE XXXVII.
SS
ISS WQS
GIN WAS SS
AWN
LQG
a
Zz
——— oP
SKULL OF BAPTANODON DISCUS. 1, SupERIOR View (No. 878); 2, Inrerior View (No. 603). Bora 4 NaturRAL Size.
SOME SUTURES DRAWN AFTER No. 1441.
EXPLANATION OF PLATE XXXVIII.
1. Anterior view of pectoral girdle of Baptanodon robustus (No. 919). Type. One fifth
natural size. co., coracoids ; sc., scapule; x., surfaces for attachment of the clavicles.
2. Dorsal view of the same girdle. One fifth natural size. co., coracoids ; sc., scapule ; @.,
surfaces for attachment of the clavicles.
342
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXXVIII
Pectoral GIRDLE OF BAPTANODON ROBUSTUS (TYPE SPECIMEN). 1, ANTERIOR ViEW; 2, DorsaL VIEW
(No. 919). Born + Narurat Size.
5)
Publications of the Carnegie Museum.
Verge No. 10.
W. J. HOLLAND, Eprror.
THH CRAWFISHES OF THE STATH
OF PENNSYLVANIA.
By A. E. ORTMANN, Pa.D.
TABLE OF CONTENTS, AND INDEX OF VOL. IL.
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| MEMOTRS
OF THE
CARNEGIE MUSEUM.
VOL. II. ei ee NO. 10.
THE CRAWFISHES OF THE STATE OF PENNSYLVANIA.
By ArnoLtp E. Ortmann, Pu.D.
I. [yrropuction.
The present Memoir is a continuation of, and an enlargement upon, the prelimi-
nary paper published some time ago in the Annals of the Carnegie Museum (Vol.
III, 1905, p. 387 et seq.) under the title “The Crawfishes of Western Pennsyl-
vania.”’ The object of these publications is to furnish the student with an account
of the crawfish-fauna of the state of Pennsylvania as complete as possible, not only
from the morphological and zoégeographical, but also from the biological, ecolog-
ical, and economic standpoint. It is now believed that it is possible to present an
approximately complete report upon this important branch of the fresh-water fauna
of the state, and in the prosecution of the studies of the author a number of ques-
tions were raised, the solution of which proved to be highly interesting.
It may be well at the beginning to give an outline of the work done. At the
outset the writer resolved to go over the whole state, and to collect specimens in as
many different localities as possible. Very soon, however, it was discovered that
the different parts of the state are of unequal interest. Large tracts, located chiefly
in the central, northern, and northeastern parts of the state, proved to be rather
uninteresting, only one species of crawfish being present in them, while the western,
and chiefly the southwestern, and again the southeastern sections offered more
variety. Thus it became necessary to pay more attention to the latter areas. The
uninteresting regions were entered only in a few cases, but a good deal of work was
done around their edges, in order to trace their limits as accurately as possible.
The location of the writer in Pittsburgh was advantageous, being central within
that section of the state which offered the greatest number of problems. Most of
343
344 MEMOIRS OF THE CARNEGIE MUSEUM
the collecting excursions were undertaken with Pittsburgh as a base. However, on
three oceasions the base was shifted. Visits were twice made to the eastern part of
the state, where the writer spent several weeks in September of the years 1904 and
1905 in Philadelphia and its environs, and once to the eastern central part, where
several days were spent in Harrisburg in June, 1905. he latter visit was marred
by rainy weather.
The work of collecting was done for the Carnegie Museum by the writer in con-
nection with his duties as Curator of Invertebrate Zodlogy, and all the necessary
expenses were paid by the Museum. In order to give an idea of the amount of
field-work done, a few statistics may be interesting.
Altogether one hundred and thirty-eight days were spent in the field, counting
only those days on which actual collecting was done: four days in 1903; sixty in
1904; and seventy-four in 1905. A few additional records were obtained in 1906.
The distances covered in travelling were as follows :
Total.
Byarall inl 904 aeeomeessosnsecscenseteeceeeceoescceesccece 3238 miles.
Senta ecu ree NOOR ace. a aoc MRR) 10,817 miles.
By team, in 1904 .. oo Ao 12 miles.
pees! 1 OD irecachuaectcatecataninecn vansaceanetateneeares OY} = SC 38 miles.
Onifoob pine O0SPerccecceccese ee ercoeee cone eee eaceeen ee 3 miles.
porte teas SSA OA Malt SecuenctecSen oan saaentnesciacamececene seas 17.
CU Sao dle KS ta hanna ners Suse Roe BR SASS once NOro ATC eM ESC ae 209‘ 385 miles.
GrandiDotalyscso ences coh eaenase oheee coe aarencosneaees 11,240 miles.
Collections were made at about one hundred and fifty-six different localities,
most of them in the state of Pennsylvania. Of the sixty-seven counties of the state
thirty-nine were visited.’ Besides, visits were made to a number of localities situ-
ated in neighboring states, namely : in Camden County, New Jersey; in Allegheny
and Garrett Counties, Maryland; in Morgan, Mineral, Tucker, Preston, Monon-
galia, Pleasants, Wetzel, Marshall, Ohio, Brooke, and Hancock Counties, West Vir-
ginia; and in Harrison, Carroll, and Stark Counties, Ohio.
The material secured on these excursions belongs to and has been deposited in
the collections of the Carnegie Museum, and comprises 303 entries in the Cata-
log, including 1869 specimens. But this does not represent the entire number of
specimens collected, since large sets, which have not been cataloged, have been set
aside as material for exchange, study, ete.
‘Material was secured, seen, or was previously known from fifteen additional counties, so that only thirteen are
not explored, namely : Carbon, Juniata, Lackawanna, Lebanon, Mifflin, Monroe, Montour, Pike, Schuylkill, Snyder,
Susquehanna, Union, and Wyoming. All these belong to the central and northeastern section of the state, where only
one species of Cambarus (C. bartoni) is to be expected, with the exception of those localities which are in the immediate
vicinity of the main branches of the Susquehanna Riyer, where also C. limosus may be present (Juniata, Montour,
Snyder, and Union Counties),
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 345
Practical experience gradually revealed to the! writer the best method of col-
lecting crawfishes. At first the writer was rather inexperienced, and did not know
where to look for certain species. But the necessary knowledge and skill were soon
acquired.
To collect the species living in streams, rivers, and ponds requires no special
effort ; it is only necessary to wade into the water-course to be investigated, or to
walk along its edge, and to discover the hiding-places of the crawfishes, which is
done by turning over stones. Mr. McConnell collected a newly moulted male with a soft shell, of the first form, at Bloomsburg, Columbia
County, on July 19, 1905, and during August he has several dates for these males: August 10 (Reading) ; August 18
(Marion) ; August 21 (Greenpark) ; August 22 (Landisburg).
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 479
Andrews (1904, p. 166) places the normal mating season in the months of February,
March, and the beginning of April, and says that there seems to be an autumnal
pairing (October, November) “in place of or in addition to” the spring pairing. I
believe, however, that the autumnal pairing is the normal one, which may be
extended through the winter. Indeed I have observed it in January, but only in
specimens kept in captivity. Since Andrews’ observations were made in the labora-
tory, it appears probable that the mating may be continued or repeated under excep-
tional conditions such as are offered in captivity, but that this is not normal. In
C. obscurus I am positive that under natural conditions copulation does not take
place in March and April.
The time of spawning, as observed by Andrews (1904, p. 176) agrees well with
our records (end of March and April), also the time of hatching (/. ¢., p. 187), late
in May.
As to sexual maturity, Andrews did not gather facts to show that females are
mature and oviposit at the end of the first year (1904, p. 206), although he observed
copulation at the end of the first summer. I observed, on November 4, 1905, copu-
lation taking place in specimens less than 45 mm. long, and found females “in
berry” of the size of 50 and 45 mm. (Cumberland, May 9, 1905). Since the same
fact has been observed in the case of C. obscwrus it is certain that males as well as
females are sexually mature at the end of the first summer, and that the sexual
union is effective, the females spawning the following spring. However, in such
small females the number of eggs is generally very small (fifty or less).
Thus it seems that C. limosus agrees perfectly with C. obscwrws in its life-history,
and that the only marked difference from Andrews’ account concerns the mating
season. ‘This is however apparently due to the fact that Andrews’ observations were
made in the laboratory. The explanation for this is very likely to be sought in the
temperature conditions. ‘The water used in tanks in laboratories has generally a
rather uniform temperature throughout the year, while under natural conditions the
temperature of ponds, rivers, and streams varies considerably in summer and winter.
I made a few observations with reference to C. obscwrus. In January, under the ice,
the water is near the freezing point, say about 35° F.; on April 6, when females
were found spawning, the temperature of Thorn Creek, Butler County, was 45° F.
On May 1 the temperature of Grave Creek, Marshall County, West Virginia, was
66° F. and spring moulting was going on. In midsummer I observed a temperature
of 82° F. in Bates Fork, Greene County, on July 24, and a temperature of 78° F. in
the Ohio, at Ambridge in Beaver County on August 24.
This gives a range of from about 35 to 80 during the year, and I have no doubt
480 MEMOIRS OF THE CARNEGIE MUSEUM
that the nice restriction of certain periods in the seasonal history is primarily due to
differences of temperature. It is only natural that an equalizing of the temperature
must tend to efface the seasonal periods.
4. Cambarus diogenes.
According to my observations, which extend over the period from March 22 to
November 17, this species also agrees in the main features of its seasonal develop-
ment with C. obscwrus.
I have the following records for females with eggs: April 6, 1905; April 19,
1905; May 2, 1904; May 14, 1899, (collected by Atkinson, Graf, and Williamson) ;
May 21, 1906; May 22, 1905; May 27, 1904. On May 21, 1906, and June 2, 1905,
I found several females with newly hatched young under the abdomen. In no
other part of the year has this been observed, and thus the spawning and hatching
season is well fixed (April, May, and the beginning of June), and is found to be
identical with that of C. obscwrus.
The number of eggs is considerably less than in the case of C. obscwrus, and
generally falls considerably short of one hundred. The spawning does not take
place outside of the burrows, but inside of them, and this was most evident in a
female collected on April 6, 1905, (Renfrew, Butler County), in which the eggs were
quite fresh, with traces of the “apron” still visible. This female was dug out of its
hole, as were all the rest with eggs or young ones.
After hatching the young remain a short time under the abdomen of the mother.
But soon they leave her, yet remain in the same hole. I have repeatedly found
young specimens in the same hole with their mother, namely, on June 13, 1904;
June 15, 1905; July 6, 1905; July 19, 1905. The smallest were about 10 mm.
long. These young specimens generally occupy a separate part of the burrow, and
are often found near to and inside of the mouth of a closed chimney. They remain
in the hole until they attain a length of 20 mm., which happens toward the end of
July. Then they leave the hole of the parent crawfish and begin to build their own
little holes and chimneys. I observed this on July 26, 1904, at Derry, Westmore-
land County, when I discovered a female 20.5 mm. long in a small hole of its own.
At the same date I found a larger one, 30.5 mm. long, which may have belonged to
the same generation. On August 4, 1904, at Francis Mine, near Burgettstown,
Washington County, I found numerous young specimens between 20 and 29.5 mm.
long, all in their own holes. On August 22, 1905, at Squaw Run, Allegheny County,
I discovered two young specimens 22 and 28 mm. long. On August 26, 1905, at
Baden, Beaver County, specimens 31.5 to 42 mm. long were found apparently under
ORTMANN : THE CRAWFISHES OF THE STATE OF PENNSYLVANIA A81
the same conditions. As late as September 5, 1904, (Smithfield, Fayette County),
and October 6, 1905, (Kittanning, Armstrong County), I found two very small speci-
mens (24 and 20.5 mm. long) in small holes. Never after July 19 have I found
young ones in the hole of the mother, so that it is quite sure that at the end of July
they invariably shift for themselves when they have attained a length of about 20
mm. ‘The largest found in the hole with its mother was 18.5 mm. in length, on
July 19, 1905. Since young specimens found in the same hole, apparently being
brothers and sisters, often have a different length (15 to 18.5 mm. in the case just
mentioned), and since, as said above, specimens of only 20.5 mm. in length are found
as late as October, the rule is established in the case of this species also that the in-
dividuals of the same litter grow up at a different rate.
With regard to the presence of males of the first form, the same conditions seem
to prevail as in the case of C. obscurus. These males are frequent in spring. I
found them at the following dates: March 23; April 2, 6, 15, 16, 24,30; May 2, 3,
13, 14, 21, 22, 27, 29; June 2; 15. Then follows a gap of over a month to July 20.
Within this period I made observations upon the following dates: June 16, 18, 26,
27; July 6, 16. At none of these dates did I discover a male of the first form. It
is true that the material in this species is less abundant, a dozen specimens collected
on one day representing a rich haul; but it is nevertheless remarkable that during
the period just mentioned, in which particular pains were taken to get males of the
first form, none were secured. But after this they again appeared regularly, namely :
on July 20; August 7, 8, 22, 26; September 5, 7, 15, 19, 21; October 6, 9, 11, 18,
24; November 5. This makes it evident that in early summer (end of June and
beginning of July) there is a time when no males of the first form are present.
However, males of the second form are found at any time in the year as fre-
quently as those of the first form. In this respect C. diogenes seems to differ from
C. obscurus. This seems to be due to the fact that C. diogenes attains sexual
maturity at a later age than C! obscuwrus and the river species in general. The
smallest male of the first form ever found measures 55 mm. in length (August 22,
1905, Montrose). It is hardly possible that this individual should belong to the
generation born in June of the same year, since the latter are known to be at that
time about 30 or at the utmost 40 mm. long. We may assume that C. diogenes,
like C. obscwrus and C. limosus, may attain at the end of the first summer a length
of about 40 or 50 mm., but these individuals do not then assume the first form as
the river species do. The same is true of the females. The smallest seen in copula-
tion (or associated with a male) was 63 mm. long, and the smallest female with
eggs was 62 mm. long.
482 MEMOIRS OF THE CARNEGIE MUSEUM
Young specimens less than 30 mm. long have not been found in spring, with
one exception, when a female of 25.5 mm. in length was found on May 27, 1904, at
Squaw Run. ‘This, however, apparently was an exceptionally delayed individual
of the generation of the previous year. It was found under unusual conditions,
under a stone on the banks of the creek, evidently removed from its original habi-
tation by winter or spring floods. No additional specimens were seen in close
proximity.
I cannot say much in the case of C. diogenes in reference to the regular moult-
ing periods which were observed in C. obscwrus to take place in spring and fall. I
have repeatedly found soft shells, and on April 24, 1904 (Nine-Mile Run, Pitts-
burgh), at a place where a large colony of this species was present, I picked up
numerous cast-off claws,” which would indicate an early spring moult. But these
claws may have accumulated during winter and spring. The rate of increase at a
moult was measured in one instance. A female, 52 mm. long, captured on April
6, 1905, was kept in captivity, and moulted on July 16. After this process it was
54 mm. long. This cannot be regarded as entirely normal, since the specimen was
kept under unfavorable and artificial conditions.
The copulating season of this species also falls in the autumn. I have only
twice observed a male and female in copulating attitude, but in both cases they let
go when disturbed. This was on November 5, 1904 (Nine-mile Run, Pittsburgh),
and on October 24, 1905 (Branchton, Butler County). The first couple was found
in water inside and near the mouth of a comparatively simple hole. The male was
70 mm., the female 81 mm.long. The second couple was found a little deeper, but
not over a foot, also in water. The male was 64 mm., the female 63 mm. long.
Mr. F. E. Kelly reports a similar observation made by him on November 14, 1904.
Besides on two other occasions I found males and females associated in couples in
the same hole. Three cases were observed on September 5, 1904, at Smithfield,
Fayette County, and two cases on August 26, 1905, at Baden, Beaver County.
Since it is an absolute rule that under ordinary circumstances only one specimen
occupies a hole, these finds are significant, and, inasmuch as in all these cases it
was always a male of the first form which was associated with a female of good size
(over 65 mm. long), it is evident that this association was connected with the mating
process. Whether the male visits the female, or vice versa, 1 do not know. In
every case the pair was easily captured, being lodged not far from the entrance of
the hole. In some of these cases I was struck by the simple character and small
depth of the burrow, and it may be that the couples dig out small, temporary holes
7° After moulting the shell is generally eaten up, with the exception of the big claws.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 483
for the mating time. This, however, needs further investigation, and possibly, if
found to be the case, may, nevertheless, not be the general rule.
In no other part of the year were similar observations made, and this fixes the
mating season for the months of August, September, October, and November,” agree-
ing with what we have observed in the case of C. obscwrus.
Thus we see that the seasonal cycle in the life of C. diogenes corresponds closely
to that of the river-species. ‘The only difference is in the time when sexual ma-
turity is reached, and it seems that in the case of C. diogenes this does not occur
earlier than at the end of the second summer. Whether this influences the dura-
tion of life is not known. Nevertheless the fact. that this species frequently, or even
regularly, reaches a size superior to that of C. obscwrus, specimens of over 90 mm. in
length being quite often found, suggests that this crawfish may live more than three
years, possibly four or five.
The resemblance of the life-history of this burrowing form to that of the river
species is due, I believe, in large part to the similarity of conditions of temperature.
As has been stated, C. diogenes lives near stagnant water and swamps, in places
where there is generally not much fresh and cool water, although such places are
not strictly avoided, and where the temperature of the water is subject to consider-
able seasonal changes. In winter and spring the water in the holes is rather cool
(43° Fahr. on March 23, 1905, in Nine-Mile Run), while in midsummer it becomes
when stagnant, almost lukewarm.
The above observations are in part at variance with those made by previous
writers on the same species. Girard (1852, p. 88), near Washington, D. C., found
females with eggs in March and April, which agrees with our dates, making allow-
ance for the difference of climate between Washington and western Pennsylvania.
Girard also noticed the fact that as a rule only one individual was found in each
hole and mentions as an exception that in one burrow a male and a female were
found together. However, he neglects to tell the exact date of this find (his obser-
vations were chiefly made in spring). In one case, he says that a male was seen
walking over the surface of the ground, as he believes, in search of the female.
But in this instance also no date is given.
Tarr (1884, p. 127) never found male and female together (in May, near Wash-
ington), and always only one individual in each burrow, and he never found speci-
mens outside of the holes.” He further believes that the burrowing crawfishes re-
™ It possibly extends further into the winter, asin the case of the other species discussed.
78 Although I have myself never seen a specimen of (C. diogenes walking over the ground, this must sometimes oc-
cur, for males and females must come together in the mating season, and the holes do not communicate underground.
According to Williamson (1901, p. 12), C. diogenes and C. monongalensis are nocturnal, and that they come out of their
holes at night is shown by the fact that Mr. Rhoads captured some of them in traps set out over night for rodents.
484 MEMOIRS OF THE CARNEGIE MUSEUM
treat to the streams in the winter, and in spring construct holes for the purpose of
rearing their young, and that impregnation takes place after the winter has passed.
These ideas are not supported by any evidence, and are, as we have seen above, in-
correct. His opinion that the same burrow is not occupied for more than one year
is also not supported by our observations. The time of hatching of the eggs is given
as about the middle of May (p. 128), which agrees with our dates. Faxon (1885a,
p. 74) reports that according to Mr. P. R. Uhler the female during the period of
incubation goes into pools, ditches, ete. This, however, is contrary to the observa-
tions of Girard, Tarr, and myself. All these particulars refer to the eastern form of
C. diogenes, on the coastal plain, and it seems that with regard to the spawning sea-
son and the spawning habits this form agrees with that of western Pennsylvania,
always considering the slight difference in climate which makes this season begin a
little earlier in the Atlantic lowlands.
The observations made on the western form show more marked differences.
Bundy (1877, p. 171): reports the discovery of a female with eggs nearly ready to
hatch, near Mechanicsburg, Henry County, Indiana, on January 1, 1875. Hay
(1896, p. 491) found that the breeding season in Indiana is in early spring, and ob-
served copulation on April 2, 1892. At this time the specimens leave their bur-
rows, and are frequently found in open ditches and streams. The eggs were laid
from April 18 to April 30. He also repeatedly saw females with well grown young
in small streams. According to Harris (1890, p. 267) a female with eggs was found
in Kansas on May 38, 1891,” apparently in an open ditch, as is shown by the sub-
sequent sentences.
With the exception of Bundy’s record these dates show April and May to be the
normal spawning season of the western form also. The observations of Hay and
Harris, that C. diogenes frequents open ditches in spring, and that it copulates in
spring, are, however, entirely at variance with the habits of this species in western
Pennsylvania. ‘This is not the case here, and I have never seen specimens outside
of their holes in spring. My observations began as early as March 23, at a time
when the frost was hardly out of the ground.
That this is also not the general rule in northern Indiana is shown by notes sent
to me by Mr. E. B. Williamson from Bluffton, Wells County, Indiana. Mr. Wil-
liamson writes to me: “As to C. diogenes congregating in ditches and the like in
spring I have no evidence here. ‘The large males of diogenes can be expected in
almost every little stream. ‘They move about on the bottom restlessly, not lying
concealed. Often the current catches them and they roll over and over, but they
79 Sic. Note the discrepancy between this date and the date of publication.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 485
always keep moying. Diogenes also wanders across country at this time, in day-
light as well as at night. I have found several crushed in wagon roads. Under
these circumstances I have yet to find a female.” In another letter Mr. Williamson
says pertinently that the specimens taken by him in spring in open ditches are all
old males, ‘in which the death instinct had developed.”
Thus it seems evident that the western form does not agree with the form found
in Pennsylvania in so far that in early spring the specimens seem to habitu-
ally leave their holes. Whether it is only old males when about to die (analogous
to what we observed in the case of C. obscwrus) which wander about, or whether the
females with eggs also are found in open water, and further, whether copulation
normally takes place in spring, are assumptions which remain to be proved. The
observations of Bundy, Hay, and Harris are surely correct, but it remains to be
ascertained whether they represent exceptional cases, or whether they are the rule.
Moreover it is not improbable thatin the western form the seasonal cycle is slightly
different, since it lives under somewhat different surroundings. Hay (1896, p. 491)
reports that during the dry months of the summer C. diogenes seems to lie at the
end of the burrow (which contains hardly any water) in a sort of a stupor. I never
observed anything like this in Pennsylvania, the holes of C. diogenes being always
well filled with water at the bottom, and the crawfishes being very lively.
That observations on the habits of this species should always be considered care-
fully with reference to all accompanying facts is evident from the following case :
Dr. D. A. Atkinson found a number of specimens on April 20, 1905, in open pools
near Westview, Allegheny County, Pennsylvania, in a region where this species is
abundant. These pools were in the course of an old, abandoned mill-race, which
dried out late in summer. All these specimens, seven in number, were young,
measuring from 33 to 52 mm. in length, and consequently belonging to the genera-
tion of the previous year. Now, bearing in mind the fact that the late summer and
fall of 1904 and also the winter of 1904-5 were characterized in our region by an
extreme lack of precipitation so that all streams were exceptionally low till the
middle of March, 1905, when a flood (March 20 to 25) restored the normal condi-
tions, it is very likely that this mill-race was dry in the summer and fall of 1904,
when these young specimens began to make their own burrows. ‘They selected this
place as a favorable one, and remained there all through the winter, a few smaller
floods, one on January 13 and another on March 9, not disturbing them, till the big
flood filled the mill-race again for a longer time, Such conditions, however, do not
suit this species, and consequently the specimens came out of their holes, and were
found, at least for a time, in the open pools, till they had selected more convenient
486 MEMOIRS OF THE CARNEGIE MUSEUM
locations in the neighborhood. Thus this case must be regarded as exceptional, not
as a regular or normal episode in the life of the species.
5. Cambarus barton. and Cambarus bartoni robustus.
In all the species discussed so far we have found a regular seasonal period in the
life-history, marked chiefly by a distinct mating-season in fall, a spawning-season in
spring, and a season in early summer when no males of the first form are present.
But it is entirely different in the case of C. barton. In this species none of these
seasons is recognizable.
As to the mating period, I have observations on only two dates. On May 27,
1904, I found a couple 7m copula in Squaw Run. Here I was able to make a close
observation. The act of copulation is similar to that in the case of C. limosus, as
described by Andrews (1904), but the male does not take hold of the anterior walk-
ing feet of the female with its chelee, and its second|pereiopods are clasped around
the carapace of the latter, lying in the cervical groove, and almost touching each
other on the back of the female. In this case it was the fifth pereiopod of the left
side, which was stretched across the sternum in order to elevate the copulatory
organs. The male of this couple was 67 mm. long, the female 73 mm. long. The
other observation occurred on October 6, 1905, when I found two couples together
at Weskit, near Kittanning. The male of the first couple was 63.5 mm., the female
59mm. long. In the other couple both male and female were 63 mm. long. Both
couples separated when captured, and thus I cannot give particulars.
These two dates are so far remote from each other that it seems hardly probable
that they belong to one and the same breeding season. It is possible that one of
them is exceptional, but I have no means of deciding this. On the other hand, as
we shall see presently, spawning takes place at such different times of the year
that very likely the mating-season is also irregular.
Females with eggs have been found on the following dates: July 6, 1905; July
10, 1905 ; July 20, 1904; July 29, 1905; August 1, 1905; August 9, 1904; August 10,
1905. The number of eggs was between seven and one hundred and thirty-three,
the smallest number being found in the smallest individual, 59 mm. long. In ad-
dition I took a number of females with young under the abdomen. ‘The following
records are at hand. At Princeton, New Jersey, in February, 1898. The exact
date is not recorded, but it was toward the end of the month. The length of the
female was 48 mm., the number of the young was ten.” Further: March 31, 1905;
6° This number is unreliable, but represents as many as were secured. In some cases quite a number of the young
dropped off when the mother was captured.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 487
length of mother 71 mm., ten young; August 18, 1904, length 59 mm., ninety-two
young ; September, 20, 1905, two cases, one 53 mm., long, with thirty-nine young,
and another 55 mm. long, with thirty-five young ; November 8, 1905, 73 mm. long,
with one hundred and eleven young; November 22, 1905, two cases, one 84 mm.
long, with seventy-five young, and another 67 mm. long, with sixty-eight young.
This extends the spawning season over the following months : February, March,
July, August, September to November. Since young were found in February and
November, these must have been in the egg-stage at least a month before they were
captured, and this would add January and October. Thus we have only interrup-
tions in December and from April to June. The gap in December may easily be
filled, and be due only to the incompleteness of our investigations in winter, but
the gap in April, May, and June may be real.
If there is any spawning-season in C. bartoni it would cover nine months of the
year, from July to March. This, however, is entirely different from what we have
seen in the river species, where the spawning season falls exactly in the months
where no spawning has been odserved in C. bartoni. And besides, this gap may be
partly filled in C. bartoni, for I have found very young specimens (between 10 and
2U mm. long; the newly hatched young are 9 to 11 mm. long) on the following
dates: May 16, 1905 (13 to 14 mm.); May 25, 1905 (11 mm.); June 2, 1905 (about
17 mm.); June 12, 1905 (14 mm.); June 17, 1905 (15 mm.); August 22, 1905 (10
to 11 mm.).
The conclusion is that very likely C. bartont has no defined spawning-season,
but may spawn at any time of the year, and that accordingly the mating-season is
also not restricted to a particular part of the year. The latter is further substantiated
by the fact that males of the first form are found practically all the year round. I
have the following dates: March 21, 28; April 19; May 7,9, 17, 21, 25, 27, 30;
June 2, 3, 6, 12, 13, 16, 23, 24; July 10, 12, 18, 26, 29; August 1, 10, 18, 22, 26;
September 11, 16, 20, 21, 30; October 5, 6, 10, 12, 17, 24, 31; November 8, 22;
December 25. The only two months missing are January and February, when no
collecting was done. On the other hand males of the second form are also abund-
ant all the year round, and were found, with the exception of January and February,
in every month.
Under these circumstances it is impossible to say anything about the life-cycle
of the single individual, since different generations cannot be traced. But one
thing should be mentioned. The males of this species do not seem to attain sexual
maturity as early as the river-species. The smallest male of the first form ever
found in eastern Pennsylvania is 49 mm. long, and in western Pennsylvania 50
488 MEMOIRS OF THE CARNEGIE MUSEUM
mm. long. The smallest female with eggs or young is from New Jersey (Princeton),
and is 48 mm. long. From the eastern part of our state I have seen none smaller
than 55 mm. long, and in the western part the minimum is 59 mm. in length.
This is considerably above the minimum size of sexually mature specimens of C.
obscurus and agrees better with C. diogenes.
In one case I have been able to observe the increase in size which takes place
upon moulting. On July 11, 1905, I found at Tionesta, Forest County, a female
in the act of shedding, and succeeded in keeping her alive till the new shell was
hard enough to be measured. ‘The old shell was 32 mm. long, and the new one 36
mm. in length. In this case the crawfish withdrew from the old shell through a
crack that appeared on the dorsal side between the carapace and the abdomen.
We have seen above that the regular seasonal cycle observed in the river-species
is probably due to the regular and considerable changes of temperature taking place
in the rivers. C. barton lives in small streams, which generally are much cooler in
summer than the larger ones, and this apparently explains the difference in the
seasonal history. The temperature conditions under which C. bartoni is found, are
more uniform throughout the year, and consequently no regular seasonal periods
in the life are observed.
No previous observations on this species have been published, except William-
son’s note (1899, p. 47), that this species was found with young under the abdomen
on March 28, 1899, at Columbus, Ohio. This lack of information is rather singular,
considering the extreme abundance of this form in the eastern part of the country.
Cambarus bartoni robustus very likely is identical in its life-history with the
typical form. I have made observations at only a limited number of dates, but
they tend to show that there are no marked seasonal periods.
The following dates for the capture of males of the first form are at hand: May
27, 1904; July 11, 1905; August 22, 1905; September 18, 1900 (Atkinson collec-
tion); September 30, 1905; October 4, 1904; October 6, 1904; November 14, 1903
(Mus. Oberlin). The smallest male of the first form measures 63 mm. in length.
Males of the second form were taken in the months of May, June, July, August,
September, and October. They were abundant in every case, considering the number
of specimens secured.
Copulation was never observed. A female with eggs was found on July 11,
1905, at Spartansburg, Crawford County. It was 84 mm. long, and the number of
eggs was 228, more than twice the number of those usually observed in the typical
C. barton. Young specimens, less than 20 mm. long, were taken on May 27, 1904
(18 mm.); and were numerous in a lot collected by Miss G. Kinzer on August 27,
1905 (9 to 16 mm.).
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 489
For the months of December, January, February, March, and April, no records
are at hand.
6. Cambarus monongalensis and Cambarus carolinus.
The temperature conditions under which these two species are found are similar
to those observed in the case of C. barton, and they are even more uniform, con-
sidering the fact that both are exclusively restricted to spring-water, avoiding even
small streams. In 1905 I took the following measurements of the temperature of
the water in the holes of C. monongalensis (May 16 at Morgantown, the rest at
Edgewood Park); March 18 (ground still frozen in places); 39° F.; May 16, 58° F.;
July 8, 63° F.; August 18, 68° F. The range is considerably less than that given
for C. obscwrus (35° to 80° F., see above, p. 479). We consequently should expect a
similar irregularity in the seasonal history as in ©. bartoni, differing markedly from
the third chimney-builder, C. diogenes. This is indeed the case.
My observations are rather complete with reference to C. monongalensis, covering
the time from March 18 to December 26. During no period within this time were
males of the second form absent or scarce, but males of the first form were also
almost regularly found ; the following are the dates for the latter: March 18; April
4,21, 24; May 1, 6, 9, 16, 21, 24; June 3, 30; July 6, 8, 20, 24; August 7, 13, 18;
September 10; October 9, 10; November 8.
Copulation was not observed; but in one case, May 6, 1904, (Fern Hollow), a
male of the first form anda female (55 and 72 mm. long respectively) were found
together in the same hole. The smallest male of the first form ever found was 53
mm. long.
Females with eggs were obtained on June 25, 1906 (O. T. Cruikshank); June
28, 1905, (three specimens); June 30, 1904, and July 20, 1905. These few obser-
vations would tend to restrict the spawning-season to the months of June and July,
but this conclusion is not admissible, since newly born young are found at various
other parts of the year. I have such (less than 20 mm. long) taken from the hole
of the mother, at the following dates: April 4, 1995 (13 mm. long); April 29, 1905
(18 mm. long); May 1, 1905 (17 to 20 mm. long); May 2, 1905 (16 to 21 mm.
long); May 6, 1904 (19 mm.‘long); June 11, 1904 (19 to 20 mm. long); August
18, 1905 (14 mm. long); September 24, 1898 (19 mm. long and above, collected by
Rhoads and Williamson); October 29, 1904 (19 mm. long). This extends the
spawning-season considerably, but it has the appearance of being interrupted
during the winter.
The smallest female with eggs is 58.5 mm. long. The number of eggs is between
thirty-eight and seventy-nine, which is considerably less than in the river species,
and also on the average slightly less than in C. bartoni and diogenes.
490, MEMOIRS OF THE CARNEGIE MUSEUM
In this species the young of one litter seem likewise to grow at a different rate.
Thirteen young found with the mother in the same hole on May 2, 1905, were
between 16 and 21 mm. long. ‘Ten young found on June 16, 1904, were from 20.5
to 32.5 mm. long. Twenty-two young, found on July 20, were from 22 to 27 mm.
long. Twenty-four specimens dug out with the mother by Rhoads and William-
son *t on September 24, 1898, are from 19 to 29.5 mm. long.
Thus it seems that C. monongalensis agrees well with C. barton. No well-marked
spawning-season is present. At any rate the spawning-season extends over a very
large part of the year, and, correspondingly, no well-marked mating-season can be
distinguished. Males of the first form may be found at any time, and also males of
the second form. Sexual maturity is delayed, males turning into the first form
comparatively late, and the females also are not mature before they have reached a
larger size than the river species. In all these respects, except sexual maturity, C.
monongalensis differs from C. diogenes.
There are further differences from C. diogenes in the development of the young.
We have seen that young C. diogenes remain in the hole of the mother for some time
after they have left the abdomen of the latter. When hatched they are about 9 or
10 mm. long, and leave the mother very soon, since free individuals have been found
only 10 mm. long. When they have grown to about 20 mm. in length, they leave
the hole of the mother.
In C. monongalensis, however, they stay considerably longer in the hole of the
mother. The exact time cannot be ascertained, but we can draw conclusions from
their size. Free young specimens from 13 to 25 mm. in length are always found
with the mother. The smallest specimen which had begun to make a hole of its
own was 26 mm. long (August 7, 1905, Fern Hollow). Another was 29 mm. long
(October 28, 1905, Edgewood Park). Specimens over 80 mm. long generally have
built their own burrows. But there are exceptions. As we have seen, young up to
32.5 mm. long have been found with the mother; and further, on July 24, 1905
(Deer Lick), I took out of one hole fifteen young, measuring from 27 to 33 mm. in
length, and out of another hole three young measuring 37, 39.5, and 40 mm. in
length. In these cases the mother was also in the hole. Although in the last two
cases conditions seem rather abnormal, it is certain that the young of C. mononga-
lensis remain longer in the hole of the mother than those of C. diogenes. While the
latter begin to shift for themselves when about 20 mm. long, young specimens of
C. monongalensis do not do so before they reach 25 or 80 mm. in length, and may
even postpone this step till they have attained a larger size (maximum 40 mm.).
1 Williamson (1901, p. 12) says that there were forty-seven young ones: only twenty-four are now in the collec-
tion of the Carnegie Museum (Cat. No. 74. 25). Possibly this discrepancy is due to a misprint.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 491
I once observed the change of a male from the second form to the first form.
The specimen was 58 mm. long, and was dug out of its hole on August 18, 1905, at
the type locality, Edgewood Park. It was kept in a jar in water, and had moulted
on August 30. I had been away on an excursion on the two preceding days, but on
August 27 it had not yet shed. When the fact was discovered it had eaten the
larger part of its old shell, only the claws remaining, which were also eaten up sub-
sequently, all but the finger-tips, by September 4. The new shell measured 61 mm.
in length. When captured this specimen was of the second form. After moulting
it was of the first form.
On account of the irregular spawning season it is impossible to trace the life-his-
tory of one and the same individual, and consequently we cannot draw conclusions
as to the duration of life.
The few observations on C. carolinus entirely agree with those made on C.
monongalensis.
Males of the second forms were found in May, June, July, August, and Sep-
tember. For males of the first form I have the following dates: May 17, 1905;
June 24, 1904; August 2, 1905; August 11, 1904; August 12, 1904 ; September 5,
1905 ; September 7, 1904 ; October 16, 1905. The smallest male of the first form
measures 56 mm. in length.
Two females with eggs were secured on July 12, 1904. One was 80 mm. long,
and had only three.eggs ; the other was 77.5 mm. long, and had seven eggs. These
numbers seem strangely small, and apparently are not normal, for on August 1, 1905,
I found a female 69.5 mm. long, with twenty-two young under the abdomen. But
even this number is below the average of C. monongalensis. While these cases seem
to indicate a spawning season in July, the finding of very young ones in the hole of
the mother at other dates considerably extends this period. I have found such on
May 17, 1905 (17 to 21 mm. long); June 13, 1905 (14.5 to 21 mm. long); August
1, 1905 (18 to 25 mm. long); August 2, 1905 (28 mm. long); August 9, 1904 (17
to 25 mm. long); August 11, 1904 (19 to 29 mm. long).
The largest young remaining with the mother were 29 mm. long, while the
smallest in a hole by itself was 30.5 mm. long. Thus the time of leaving the hole
of the mother is about the same as in C. monongalensis.
The above observations are not at all sufficient to show that C. carolinus agrees
entirely with C. monongalensis, but since both species are alike in so many particu-
lars, morphological and ecological, and since the above dates do not show any differ-
ences, we may safely assume that the life-history of both species is similar.
The seasonal history is rather well known in four of the species above discussed,
492 MEMOIRS OF THE CARNEGIE MUSEUM
C. obscurus, C. bartoni, C. monongalensis, and C. diogenes. These are the species
found in Allegheny County, and they are most complete, since I had the best chance
to study them, three of them being found in the immediate vicinity of my residence
and the fourth (obsewrus) within a few miles and within easy reach.
We are able to distinguish two main types of life-history, which I should like to
call for convenience the warm water and the cool water types. ©. obscwrus and
diogenes represent the first, and agree with each other in having well marked mating-
and spawning-seasons, and in early summera period when no males of the first form
are found. They differ, however, in the fact that in C. obscwrws sexual maturity is
reached, as a rule, at the end of the first summer, which does not seem to be the
case in C. diogenes. Of the other species, of which no complete series of dates are at
hand, the river-species, C. limosus, C. propinquus, and C. propinquus sanborm, very
likely agree with ©. obscwrus, for the comparison of the dates does not reveal any
differences.
The cool water type is represented by C. bartont and C. monongalensis. Both are
characterized by the absence of well marked mating- and spawning-seasons. They
may be expected in any stage of development at any part of the year, even winter
making no exception. C. carolinus probably belongs also to this type, although the
observations are too scanty to positively establish the fact.
One thing in conclusion should be especially emphasized. The life-history and
the habits of different species of the genus Cambarus are by no means similar. On
the contrary they differ considerably, and the differences may be accounted for pri-
marily by the different ecological conditions under which they live. Consequently
it is inadmissible to generalize from facts observed in one species only, and further
it is to be expected, if other species are studied, that additional types of life-history
will be discovered.
VI. ECONOMIC VALUE.
1. Popular knowledge of Crawfishes.
The crawfishes of this state are generally well known to the population. They
are abundant and large enough to attract the attention even of the casual observer.
But it is chiefly the small boy who is interested in them. Three popular names are
employed for them, crab, crayfish, and crawfish. ‘ Crab” obviously is a misnomer,
belonging originally to the marine Brachywra, but it is largely in use all over the
state, and chiefly so in the cities. The word “crayfish” is used the least. In my
experience I heard it mostly in the mouths of such people as had a certain amount
of schooling and had acquired some knowledge of natural history. This word is
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 493
preferred by teachers generally, very likely in consequence of its use in one of the
standard works on these creatures (Huxley, ‘“‘The Crayfish”). The third word,
“crawfish,” is the proper American name. I found it commonly in use in the rural
communities where ‘“‘crayfish”’ and “crab” were often entirely unknown. This is
chiefly the case in the southwestern section of the state and in West Virginia. In one
or two cases in Fayette and Somerset Counties I heard a distinction made between
“erab’’ and “crawfish.” The former name was used for the river and brook
forms, C. obscwrus and C. barton, the other for the chimney-builders. All three
words go back to the same root, Old German krebis, from which is derived on the
one hand the modern German Krebs, and the English crab; on the other hand the
French éerevisse, the English crayfish, and the American crawfish. The latter form,
being typically American, and being exclusively known to the natives of a large
part of the country (the farmers), I have decided to use it in preference to the other
two forms.
In literature “crawfish” was used by Say (1817), Harlan (1835), Hagen (1870),
Abbott (1873), Hay (1896). “Crayfish” was used by Abbott (1884 and 1885),
Faxon (1885, 1890, 1898), Hay (1898, 1899), Andrews (1895, 1904), Shufeldt (1896),
Osburn and Williamson (1898), Harris (1900), Williamson (1901, 1905). Thus
“crawfish ” has the priority.
Other names have been given incidentally. Rafinesque (1817) calls C. limosus
“mud lobster,’ (I heard this name once in Delaware County). Say (1817) and
Harlan (1885) call C. bartoni “freshwater lobster,” and Williamson (1899) uses the
abbreviation ‘“‘cray.”
2. The use of crawfishes as food and bait.
Although well known, crawfishes are not much used as food by the population
of Pennsylvania; but this is generally the case in the United States. In some of
the larger cities of the United States they are found more or less regularly on the
market (see Ortmann, 1900, p. 1260), C. limosws being one of the species which is
principally used for food. I have, however, never heard that this is the case in our
own state, but it may be found in the markets of Philadelphia.”
Nevertheless crawfishes are eaten in this state, but not regularly. I have heard
sometimes from boys that they had tried them, but only in ‘“‘sport,” and only excep-
tionally have I met persons who had eaten them repeatedly and were fond of them.
Generally, this source of food is unknown to the masses in this state. Yet a dish of
crawfishes is not to be despised. It is true, our species never attain the size of the
S?Rafinesque (1817, p. 42) says of C. limosus at Philadelphia, tbat it is ‘‘ good to eat.’?
494 ; MEMOIRS OF THE CARNEGIE MUSEUM
highly esteemed European forms, but I know from my own experience that, as
regards quality, the former are not inferior to the latter. Young specimens (and
chiefly soft shells) may be fried in butter and eaten shell and all, while the abdom-
inal muscles of older ones, when boiled in water, are very good.
Of course, it is hard to create a taste for crawfishes among the masses, but I do
believe that it would be worth while to try. Crawfishes are so abundant in certain
parts of the larger rivers, C. limosus in the Delaware, and OC. obscwrus in the Ohio
drainage, that it is easy to get any amount of them. It also would not be difficult
to raise them, for instance in ponds, and to supply the market regularly and judi-
ciously. And further, I do not see, why the “tails” (abdomen) could not be used
for canning, exactly like the tails of shrimps and prawns.
Beyond this, crawfishes are used only as bait by fishermen. ‘This use is quite
general, and crawfishes form an important part of the fisherman’s outfit especially in
western Pennsylvania. ‘They are most valuable in fishing for Black Bass (Microp-
terus), since these fishes seem to be very partial to this bait.
3. Crawfishes as scavengers. Thew food. Their enemies.
The indirect economic value of crawfishes is best expressed by saying that they are
scavengers, as decapod crustaceans in general. They dispose effectively and quickly
of any decayed matter, animal or vegetable, coming within their reach. They also
eat living creatures. This was known previously. Abbott (1873, p. 83) calls them
(C. limosus and bartoni) “ omnivorous,” and “scavengers,” and says that they eat
water-weeds, and seize young Cyprinoid fishes. Andrews (1904, p. 175) fed C.
limosus in the laboratory on raw and cooked meat, raw eggs, pieces of earthworms,
and on Chara and Hydrodictyon. Williamson (1901, p. 12) reports that C. monon-
galensis was caught in traps baited with raisin and oatmeal. I used for my speci-
mens in the laboratory all kinds of meat, and since I am especially fond of smoked
sausage, I let them often partake of it when I had it for lunch. They also eat earth-
worms and green vegetable matter, for instance seedlings of several weeds (Galin-
soga, and Rumex acetosella), grass, and water-weeds (Vallisneria). In nature they are
often found at carcasses and other animal refuse lying in the water. They eat in-
sects. For instance I have seen C. bartoni taking grasshoppers used as bait while
fishing for trout (Tub Mill Run, Ross Furnace, Westmoreland County). In the
case of the chimney-builders vegetable matter seems to be largely resorted to, not
only fresh plants, but also decaying vegetation being used. In digging them out of
their holes I repeatedly found masses of decaying leaves and the like lodged in some
side branch of the hole in such a position that they could not have fallen in acci-
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 495
dentally, but must have been brought in by the crawfish. In one case (October 9,
1905, Nine-Mile Run), I found in a side-pocket of a hole of a female C. monongalen-
sis anumber of ripe fruits of Crategus, about a handful, which under no circum-
stances could have fallen into the position where they were found. The hole was
under a large Crategus bush.
Thus it seems that any vegetable or animal matter, either fresh, or decaying,
serves as food for crawfishes, and although some species may prefer certain classes of
food on account of taste or necessity, they all take readily to any kind, as is seen by
the fact that in captivity they eat everything that is offered to them without dis-
crimination. If nothing is given, they eat one another.
Crawfishes in turn serve as food for many animals, chiefly those which are
aquatic. Among mammals we know that raccoons hunt for them. As has been
mentioned above, birds eat them, and the kingfisher and other equatic birds do so
quite regularly. The report of Audubon, (see Ortman, 1900, p. 1250), that the White
Ibis captures the chimney-builders by throwing fragments of the chimney into the
hole, and watching for the crawfish to come up, does not seem strange to me. At
Ohiopyle I was told that a domesticated turkey kept upon the grounds of the hotel
had the habit of watching the holes of C. carolinus, and that frequently he captured
this species. I have myself seen this turkey standing motionless before a hole, but
I did not observe the actual capture. I do not entertain the slightest doubt that
this and other birds are able to catch crawfishes in this way, and do not think that
it is necessary to drop dirt into the hole, since the crawfish comes up frequently on
its own account, when it may be seized.
Crawfishes constitute an important part of the diet of certain snakes, more par-
ticularly of the water-snakes, Natrix sipedon and leberis. I have seen the latter dis-
gorge C. obscwrus when captured. (See also Atkinson, Ann. Carn. Mus., I. 1901, p.
149, 150.) On two occasions I have found garter snakes, Hutznia sirtalis, in holes of
C. monongalensis; two specimens of this snake in one hole on October 18, 1904,
(Fern Hollow), and one snake in a hole on October 28, 1905, (Edgewood Park).
However, whether the snakes were after the crawfishes, or whether they simply were
using the holes for winter quarters, remains doubtful.
Professor H. A. Surface writes to me that Cryptobranchus allegheniensis and Nec-
turus maculosus are among the chief enemies of the crawfishes, and, indeed, these
two salamanders are generally found at places where crawfishes abound. (Compare
Eydeshymer, American Naturalist, XL, 1906, p. 128.)
They are, however, most valuable as food for the fish-fauna of our waters. As
has been mentioned above, crawfishes are good bait for certain fishes, and it is very
496 MEMOIRS OF THE CARNEGIE MUSEUM
likely that many of our freshwater fishes depend largely upon crawfishes for nutri-
ment. It would be interesting to investigate how far this mutual correlation
between fishes and crawfishes holds good in our state. The presence of a river-
species in our western streams, and its absence in any drainage systems in the cen-
tral parts is very remarkable. Indeed C. bartona is found in rivers, but only occa-
sionally, and in small numbers. My own observations are not sufficient to give an
approximate idea as to these relations, since I did not pay much attention to the
fish-fauna, and the latter has decidedly deteriorated, at least in quantity, and the
fish have become rather scarce in most of our streams. Possibly the decrease in the
number of fishes has caused an increase in the number of crawfishes.
4. Crawfishes as obnoxious creatures.
For the river-species hardly a point can be mentioned which would tend to show
that they are obnoxious to human interests, except the fact that they occasion-
ally capture young fishes. It is different with the burrowing species, which often
become troublesome. In regions where chimney-builders are abundant I have
repeatedly heard complaints about the chimneys, and chiefly so in the case of C.
carolinus in Somerset County, Pennsylvania, Garrett County, Maryland, and Preston
County, West Virginia. Here the mud-piles may hamper farming operations by
interfering with the harvesting machines, clogging and ruining them. At Selbys-
port, Maryland, I was told that conditions were so bad that the farmers tried to
exterminate the crawfishes by throwing unslacked lime broadcast over the fields,
which operation was partly successful, the crawfishes coming out of their holes by
hundreds in a dying condition. I was told that this treatment, repeated several
times, had considerably reduced the numbers of the red crawfish in this neighbor-
hood. At no other place did I hear of attempts made to kill these crawfishes,
although farmers were unanimous in denouncing them as a nuisance.
At a few places another complaint was made, namely, that the chimney-builders
were cutting off and eating up sprouting crops. This was affirmed with reference
to ( carolinus at Reedsyille, Preston County, West Virginia, where a farmer told
me that this species had cut off the largest part of a crop of buckwheat, so that
practically nothing was harvested. At Parson, Tucker County, West Virginia,
complaints were made that the same species had damaged sprouting corn; and at
New Martinsville, Wetzel County, West Virginia, I heard that C. diogenes was
charged with eating up all kinds of sprouting crops, corn and beans being especially
named.
I do not doubt that these complaints are justified, and that the burrowing species
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 497
actually eat and damage crops to a considerable degree. As I have observed, in
captivity C. monongalensis and C. diogenes eat young plants, and they surely do so
when not in captivity, young sprouting corn, buckwheat, ete., being rather succulent
and attractive to them. If sown ina place where crawfishes abound these crops will
surely be attacked.
This being the case, and besides the chimneys being also a nuisance, it might
be desirable to exterminate the crawfishes in a given locality, or at any rate to
reduce their numbers. For this purpose unslacked lime, the means employed by
the farmers at Selbysport, might be used. But I am in no position to vouch for
the efficiency of this remedy, having no personal experience (with the exception of
the one case mentioned aboye, p. 346). I simply report what was told me.
Another way might be to drain the places where crawfishes are plentiful. But
this hardly will be as efficient a means as desired. Drainage only lowers the level
of the groundwater, and in the case of C. carolinus, which is the chief offender, we
know that it digs down sometimes over three feet to reach the groundwater. In
Rainier Park at Ohiopyle this species used to be very abundant, but the draining
of the park has reduced its numbers. Still it is present there, and the chimneys
are thrown up all over the lawn, where the holes must in places go down at least
three feet before reaching water. Thus, although a decrease in numbers may be
brought about by drainage, a complete extermination by this method must not be
expected.
Another form of damage done by chimney-builders is known. They are reported
to burrow into and to do damage to the dams on ponds, reservoirs, and rivers.
(The levees of the Mississippi. See Ortmann, 1900, p. 1262.) No instances of this
kind are known to me in Pennsylvania. In one case, at the reservoir of McGee
Run, at Derry, Westmoreland County, I saw holes of C. diogenes not only along the
banks, but also in the dam. ‘The specimens were all young, and the holes small,
since this reservoir has existed only for a few years. But it would not be astonish-
ing if the crawfishes should gradually work deeper into the dam, finally causing
serious damage.
Vil. BEARING OF THE ABOVE STUDIES ON THE THEORY OF
EVOLUTION.
Our observations on the Pennsylvania crawfishes, morphological, ecological, and
geographical, serve to illustrate certain phases of the process of evolution, and certain
theories propounded in connection with them. Naturally they do not elucidate
this process in its fullest scope. Thus I shall only pick out a few points upon which
my observations may have some bearing.
498 MEMOIRS OF THE CARNEGIE MUSEUM
1. The Mutation Theory of De Vries.
The latest fashion in evolution theories is the so-called “mutation theory” of
De Vries (De Vries, 1905). It is much discussed at present, and the general trend
of opinion is that, although De Vries’ idea of the origin of species may not hold
good in all cases, he certainly has demonstrated at least one way by which species
may be formed. It is generally maintained with emphasis that his experiments
are beyond doubt and that the facts demonstrated by him cannot be denied.
This indeed is the case, and it would be lamentable if any of the statements pre-
sented by De Vries as facts should prove to be unreliable. I am decidedly of the
opinion that the statements are correct, but I also hold that De Vries was not the
first to bring the facts forward. They belong to a class that was known long ago.
But furthermore, I believe that the conclusions drawn by De Vries from these
facts are entirely wrong.
I recently have devoted several articles to demonstrate this, and shall not again
go into detail here (see Science, May 11, August 17, and November 30, 1906).
However, I shall discuss here a special part of De Vries’ theory, which concerns
the distinction he makes between “ fluctuating variation” and “mutation.” The
latter is said to be characterized by ‘“‘sudden leaps,’ while the former is said to be
by “small steps.” Although De Vries sometimes does not lay much stress upon
this distinction (see Copeland, 1904, p. 421), this difference is often regarded as
paramount in his theory (see MacDougal, in Popular Science Monthly, vol. 39,
1906, p. 207). And since De Vries believes that species are formed only by muta-
tion, it should. be expected that the morphological differences between existing
species should at least frequently exhibit signs of ‘sudden leaps.” If such leaps
are observed in our species of Cambarus, this would tend to support this part of De
Vries’ theory ; if not, the theory that mutations are always or generally marked by
discontinuity of variation, should be dropped.
2. Species, Varieties, and Variations among the Pennsylvania Crawfishes.
I have distinguished in the systematic part of this monograph seven species and
one variety among the Pennsylvania crawfishes. Besides I have discussed another
extralimital variety. This means that the characters distinguishing these forms are
different in their taxonomic value, and the reasons for thus estimating them should
be given. .
The seven species of Pennsylvania belong to two subgenera, Faxonius and
Bartomus, which are distinguished by very sharp differences in the male copulatory
organs.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 499
The subgenus Faxonius is represented in our state by three species: CO. limosus,
OC. propinquus, and C. obscurus. The first is geographically, as well as morpho-
logically, separated from the other two ; and here again it is the shape of the male
organs which serves as the chief distinguishing feature. Besides there are other
characters, such as the shape and the spinosity of the carapace, which make it pos-
sible to recognize C. limosus at a glance. No transitional forms being present, the
standing of C. limosus as a “ good species” is beyond doubt.
It is different with C. propinquus and obscwrus, and the extralimital form C.
sanbormi. These three resemble each other very closely, and it is hard, indeed
impossible, without close examination to distinguish them. ‘They also live under
similar ecological conditions, and their ranges together form a unit, so that it is evi-
dent that they are closely allied genetically. The differences of C. obscurus from the
other two forms are furnished by the ‘‘shoulder”’ of the male organ and the tubercles
of the annulus of the female, together with the complete lack of the median keel of
the rostrum. Other differences, such as sculpture and spinosity of the chelipeds,
are of secondary value and not entirely reliable. But it must be emphasized that
within the established range of C. obscuwrus, from Fish Creek in the southern part of
the Panhandle of West Virginia to the upper Alleghany and the Genessee Rivers
in McKean and Potter counties, and from Cheat River at the West Virginia state-
line, to the upper Shenango River in Crawford County this species is remarkably
uniform in the characters mentioned. No specimens have been found within this
area which show the slightest tendency toward C. propinquus.
Thus, with reference to this form, the postulate that a species should be sharply
and constantly separated from the coexisting allied forms is fulfilled (see Ortmann,
1896, p. 191) and accordingly I regard C. obscwrus as a good species.
As regards C. sanborni, matters seem to be slightly different. It agrees in the
shape of the sexual organs with C. propinquus, and differs only from the latter in
the lack of a rostral keel and some minor features in the armature of the chelipeds.
In the lack of a rostral keel it approaches C. obscwrus, but always may be distin-
guished by the shape of the sexual organs. Its relation to C. propinquus remains
doubtful. My observations do not cover the region in which possible transitions
might be expected (northern and western Ohio), and thus I must leave this question
open, and I follow Faxon in regarding C. sanborni as a variety of C. propinquus.
But it should be possible to settle this question by proper investigation, and I would
not be astonished if it should be finally discovered that C. sanborni actually is a
good species, sharply and constantly separated from C. propinquus.
The subgenus Bartonius contains four species in Pennsylvania. One of them,
500 MEMOIRS OF THE CARNEGIE MUSEUM
C. bartoni, differs from the rest ecologically as well as morphologically. It is dis-
tinguished by a number of characters, and there is no possibility of morphologically
intermediate forms, so that C. bartoni not only is a good species, but also belongs to
a different section of the subgenus.
C. bartoni possesses in Pennsylvania a variety, C. bartoni robustus, which, accord-
ing to my experience, is constant, and never runs into the typical form. It also
seems to occupy a slightly different territory, although often found associated with
the latter. ‘These facts would justify us in regarding it as a good species. I have
not done so in the systematic part, since the facts at hand are too meagre to finally
decide this question. The range of C. robustus in Pennsylvania is only a small part
of the area occupied by this form, and in the states of Ohio, New York, and in
Canada, the conditions are entirely unknown. Furthermore a form similar to our
robustus, although, as it seems to me, not entirely agreeing with it, has been reported
from Virginia, Maryland, and Kentucky, and before particulars about the relation
of this form to C. barton and to our robustus are known, we cannot judge as to the
taxonomic position of C. robustus. Therefore I have refrained from modifying the
position hitherto assumed, that this form is a variety of C. bartoni.
The other species of the subgenus Bartonius in Pennsylvania are C. carolinus, C.
monongalensis, and C. diogenes. ‘They belong to the diogenes-section, and all three
are closely allied. C. carolinus and monongalensis are more nearly related to one
another than to C. diogenes. ‘The latter apparently is a more advanced form.
C. carolinus and C. monongalensis are distinguished by rather insignificant mor-
phological characters, discovered in the shape of the rostrum and the armature of
the chelipeds. But the difference in color is so striking that it is impossible to con-
found them in the field. Other characters also, although slight, hold good accord-
ing to my experience, and I never have seen intermediate specimens. Moreover
the distribution of these two forms is very characteristic, they being sharply sep-
arated topographically, and never being found associated at the same locality. Thus
all requirements leading us to pronounce them good species are met. Of course this
applies only to conditions in Pennsylvania, Maryland, and northern West Virginia ;
whether they are the same or different farther south remains to be seen.
C. diogenes is more sharply separated from the species just discussed, and there
is no possibility of mistaking this species, more particularly as the color is markedly
different. But the morphological characters are also very nicely expressed, so that
in a case of a red (albinistic) specimen of this species I was not a moment in doubt
that I had to deal with C. diogenes, and not with C. carolinus, although the latter
was found associated with this form at this particular locality (Dunbar). There is
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 5OL
no doubt that C. diogenes is a good species, and even when discovered in company
with C. monongalensis or C. carolinus we found no intermediate forms which might
render the identification uncertain.
As has been demonstrated above, there are two races of C. diogenes in Pennsyl-
vania, an eastern and a western. They never have been distinguished before, and
indeed are very similar, so that it is hard to tell them apart. But I think Iam
able to do so. The differences are very slight, but I never observed intermediate
forms, and their existence is improbable, the ranges of the two races being widely
distant from each other. The constancy of the differential characters being the
only criterion of specific difference, while the amount of difference is of no con-
sequence at all,* we might regard the eastern form as a different species from
the western. This may prove to be the correct view, and then the eastern form
should be called C. diogenes Girard, and the western possibly C. obesws Hagen.
I have not taken this course in the systematic part, since our knowledge of C. dio-
genes is by no means complete. I know only the conditions in this state, but the
eastern range of this species extends over large parts of the coastal plain, while the
western occupies a vast territory reaching to the Rocky Mountains and the Gulf.
It is also not impossible (although improbable) that the eastern and western areas
are connected somewhere, (in Virginia’). Before this question is finally settled, and
before we know more about the conditions under which C. diogenes occurs in the
extralimital parts, it is best to refrain from expressing a positive opinion. Never-
theless it is quite possible that there is a tendency in diogenes to split into varie-
ties and species. A variety has been distinguished in Louisiana.
We see that in certain forms my studies have led to a positive decision as to their
taxonomic position. In other cases my observations must be completed and sup-
plemented by additional evidence to be gathered in other parts of this country before
a final opinion can be reached. The fault is not with the material at hand, but
with the insufficiency of our knowledge of the extralimital parts.
As to variations, that is to say, occasional aberrations from the typical form, we
have seen that such are extremely rare among the Pennsylvania crawtishes, and
have in most cases the character of freaks. Some of them, however, are interesting
from certain points of view.
No variations were discovered among one hundred and nineteen individuals of
C. limosus. With reference to C. propinquus in Erie and Crawford Counties, I have
pointed out that there is a certain amount of variation in the development of the
§3De Vries (1905, p. 127) talks of ‘‘an old rule in systematic botany, that no form is to be constituted a species
upon the basis of a single character.’’ This rule is entirely unknown to me in botany as well as in zodlogy.
502 MEMOIRS OF THE CARNEGIE MUSEUM
keel of the rostrum, and in that of the spines and the carpopodite and meropodite
of the chelipeds, and we have also seen that there is sometimes a notch on the
anterior margin of the male organ. All these characters mark a certain inclination
toward C. obscwrus which will be discussed below.
The six hundred and eighty-seven specimens of C. obscwrus at hand are, as has
been seen above, very uniform in their characters. A tendency has been observed
toward an increase of the spines of the meropodite of the cheliped in a direction
from the northeast toward the southwest within the range of this species. Here we
have apparently the first step toward the formation of a variety: a variation
becomes more frequent in the southwestern part of the range, possibly in conse-
quence of hereditary transmission, and begins to ‘“‘breed true.” But it is only the
beginning of it, the varying form not being found to the exclusion of the original,
and thus it remains ‘‘ variation” only.
Other variations (mentioned p. 375 and 376) are very likely due to injuries re-
ceived during life*, and again others are of the character of freaks, namely the two
cases of apparent hermaphroditism. One of these is rather interesting (Pl. XX XIX,
Fig. 7° and 7°). Here the male sexual organs do not at all correspond to the typical
form of this species, but approach in shape to that known in the limosus-section. It
seems to me that we have to deal here with a case of atavism. The /imosus-section has
been regarded as the most primitive type of the subgenus Maxonius, on account of the
very slightly separated tips of the copulatory organs. The propinquwus-section is next
to it, but here the tips are separated for a greater distance. It is quite probable
that the latter section descended directly from the former, and it seems that in the
instance discussed the sexual organs have reverted to the original /imosus-type, and
thus the assumption that the propinquwus-section is a descendant of the limosus-
section gains additional strength.
In the seven hundred and twenty-five specimens of C. bartoni we again have to
emphasize the great uniformity of the characters. ‘The variations discussed are
rather insignificant, and consist chiefly in the shape of the rostrum and the size.
A single individual has been observed in which one lateral spine of the carapace
was present, apparently an atavistic feature. Other variations are of the hermaph-
roditic type.
No remarkable variations have been found in ©. carolinus, and a few insignifi-
* Variations due to injuries are most frequently observed in the case of regeneration of the chelipeds. I did
not mention them in the systematic part, since they are very common. If the claws are lost they are replaced by new
claws, which differ from the old ones not only in size, but alsoin shape. The fingers are proportionally longer, and the
palm proportionally shorter than in normal claws. This difference in shape remains even if the claws, after repeated
moults, again attain a good size. Regenerated claws may always be recognized by the short palm and long fingers.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 503
cant ones in C. monongalensis. The same is true of C. diogenes, leaving out of
account the differences between the eastern and western forms. The most important
variation is that of the width of the areola. In this there is a tendency toward
regional restriction, but it is not complete. The wide areola, being a more primi-
tive character, does not represent the variation, but the original condition, which is
retained only in a small part of the range and is even there not general. It is a
character that has a tendency to disappear and may be classed under atavism. Impossibility of crossing, due to any cause, mechanical, physiological, or ecological, see Eimer, 1895, p. 14.
Gulick (1995, p. 95) calls this ‘‘ Impregnational Isolation.’? Under this head falls also Romanes’ ‘‘ physiological
isolation.”’
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 507
Even if it should be the case that C. obscwrus may cross with C. propinquus and
C. propinquus sanborni, this does not invalidate its standing as a species, for we have
numerous examples in nature in which true species form hybrids.
Thus we see that these three species, the origin of which as species belongs to
the beginning of the Glacial time, have come together again; but each seems to
remain in its original area, and where they come into actual contact the one species
is hardly able to oust the other. To a small degree hybridization seems to be pos-
sible at the points of contact. The assumption that C. sanborni might be a hybrid
between C. propinquus and C. obscwrus is rendered impossible by the exclusive
presence of C. sanborni all over its range (excepting Fishing Creek), without any
trace of the two other species.
It remains to consider the question what the relation of the specific characters
to isolation may be. We see that in the case of C. obscurus it is chiefly the “shoulder”
of the male sexual organ which distinguishes this species. This shoulder is found
at a place where an external stimulus acts upon this organ, namely, just where it
is touched by the fifth pereiopod in the act of copulation. A similar shoulder is
found in many other species of Cambarus of different groups and even subgenera,
and thus it is highly probable that it is this external stimulus which induces the
development of this feature. But this does not afford us an explanation why this
shoulder did not develop in other species, especially in C. propinquus. At present
I am unable to answer this question. The fact remains that we have to deal with
a specific character, which is clearly due to an external stimulus,” and I have
always held the opinion that every variation is invariably caused by a reaction of
the organism to some external influence. (See Ortmann, 1896, p. 188, and 1898,
p- 157.) But the view that acquired characters are transmissible is not fashionable,
although now admitted by its chief adversary, Weismann. In consequence of the
modern tendeney to deny the effect of external causes upon variation, at any rate
to deny the possibility of the hereditary transmission of such variations, not much
attention has been paid to the mutual relations between external stimuli and the
reaction of the organism upon them. But here I think much room for investigation
is left. In the present case the reaction of the organism upon the external stimulus
caused by the contact of the fifth pereiopod with the sexual organ is to form at the
point of contact a notch or angle (shoulder) on the sexual organ.
This reaction may be slightly advantageous, but it is not absolutely necessary,
for we see that there are many other species in which this reaction has not taken
place, even among the most closely allied forms, which are nevertheless well off and
8i Under ‘‘ pressure of the environment,’’ as Merriam puts it (1906, p. 244).
508 MEMOIRS OF THE CARNEGIE MUSEUM
flourishing. In other words, the ‘“selectional value” of this character is practically
at the zero-mark. This demonstrates again that the conception of “ natural selec-
tion ”’ as “selection of the fittest” isincorrect. With regard to fitness there are many
characters which are entirely indifferent, and this is one of them. The absence or
presence of a rostral keel, and of tubercles in the case of the female annulus, the
other specifie differences of these forms, belong to the same class. We thus see that
natural selection has played no part in the development of these characters of these
species. But this does not imply that selection has had nothing to do with the evo-
lution of these species, on the contrary this factor has always acted, and if these
characters had not been fit to survive, the species would not have been able to sur-
vive. Natural selection (in the modified sense, according to Pfeffer, see Ortmann,
1896, p. 176). resulted in the fact that the propinquus-group, such as it actually is,
is able to live and to flourish, but it is not responsible for the splitting up of this
group into two or three species.
The latter fact is entirely due to isolation. In the present case the isolation was
in effect only during a short period in the past, but it was enough to differentiate
several species. At the present time there is a tendency to undo this effect. These
species are beginning to mingle again. But this process has not yet progressed far,
and for several reasons will very likely be slow in future. It is hard to say what
the outcome will be, whether we shall have a hybrid form, or whether one will sup-
press the others. (C. obscwrus is the most advanced form, and also seems to be
slightly more vigorous than the others. Thus it may finally overrun them and
crowd them out, unless it is in turn conquered by a still more vigorous from, C.
rusticus, advancing from the southwest.
From the above discussion we see that whatever may have been the processes
of variation and of natural selection, or independently of what we may think
of the possibility of the inheritance of acquired characters, the fact that the
propinquus-group has split up into species is solely due to isolation, which in this
case is strictly topographical. We have here three forms with identical ecological
habits, in which topographical isolation is evident, illustrating the rule that “closely
allied species occupy neighboring areas.” (See Ortmann, 1905), p. 127, Jordan,
Science, Nov. 3, 1905, p. 546, and Merriam, 1906, p. 248, et seq.)
(c) C. bartont.
This species is morphologically well isolated from the other Pennsylvanian
species, and also has peculiar ecological habits. Being found all over the state it
necessarily comes into contact with all the other species and is often found associated
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 509
with them. This is preeminently the case with the river forms, C. limosus, C.
obseurus, and C. propinquus.
Here we have an instance in which at a given locality two species may be found
side by side. This, however, is due to secondary processes. Originally each of the
two species had a different center of radiation, and thus we again see the action of
isolation. The center of C. bartoni lies in the mountains of the Appalachian sys-
tem; the common center of C. limosus and the propinquus-group is in the central
basin of the Mississippi, and the special center of C. limosus in the coastal plain,
and that of the propinquwus-group in the Erigan and Lower Ohio drainage.
Nevertheless these species came together (see Ortmann, 1896, p. 186), but the
migration was in different directions, the river species coming up the rivers, while
C. bartoni migrated down stream. Although living side by side there is no danger
of hybridisation, since their morphological differences are such that kyesame-
chania exists. he different shape of the sexual organs of C. barton: from that in
the subgenus Faxonius precludes any idea of their being able to cross. Such cases
do not offer anything remarkable, since the occupation of and the association at the
same locality of different forms coming from different directions, and not being
closely allied, is the general rule in any ecological community (biocenosis).
Conditions are slightly different in the cases where C. barton is found in close
proximity to the chimney-builders. Here there is closer affinity, but also it seems
here that these species are so far separated morphologically that kyesamechania
exists, although the shape of the copulatory organs is similar. Moreover, wherever
C. bartoni comes into contact with the burrowing species it generally occupies situa-
tions slightly different from those preferred by the chimney-builders. It favors
running water in open streams, while the burrowers are found in holes at a certain
distance from the streams. Nevertheless, C. bartoni is sometimes found in burrows
and in springs close to the one or the other of the burrowers (it is even found in the
holes of the latter, see p. 414), but in such cases we have again the same conditions
as above: different species coming from different centers occupy the same
locality.
Yet as a rule C. bartoni occupies a different habitat from the burrowers, even if
found elose to the latter. A fine illustration of this is in Nine-Mile Run, near
Pittsburgh. Here three species, C. bartomi, C. monongalensis, and C. diogenes., are
found together upon a space hardly more than twenty feet square. The locality is
a pile of talus swept down into the valley of Nine-Mile Run by a small stream.
The stream comes through an insignificant ravine, and spreads out over the talus,
forming a kind of a delta, rendering the lower parts of the pile of talus rather
510 MEMOIRS OF THE CARNEGIE MUSEUM
swampy. At the upper end of the talus, in the outcrop of sandstone rock, and not
far (about fifteen feet) from the bed of the spring, is a copious spring, the water of
which runs directly into the clay and humus of the pile of talus, in a large part
underground. ©. bartoni is found in the small stream under stones; C. monon-
galensis is found at and immediately below the spring referred to; and C. diogenes
is abundant all over the pile of talus down to the bottom of the valley. At the
upper end of the pile of talus is the place where all three species come close to-
gether, but each is subject to different ecological conditions. :
Similar conditions have been frequently observed, and we thus have here the
occupation of the same localities by closely allied species, which differ ecologically,
that is to say, topographical isolation is not observed here, but the isolation 1s ¢co-
logical, and the differentiation of the chimney-builders from C. bartoni very likely
is connected with and largely due to the latter.
(d) C. carolinus and C. monongalensis.
We have seen that these two species are very closely allied, but that the distin-
guishing characters are constant. Ecologically they are similar, so that hybridisa-
tion might occur when they come together. The latter case, however, has never
been observed, at least in Pennsylvania, Maryland, and northern West Virginia.
The western escarpment of the Chestnut Ridge forms a sharp boundary between
them. This case corresponds to that observed in the western river-species (prop-
inquus-group). Two species identical in their ecological habits are separated topo-
graphically. But in this case the barrier separating them is of a different character.
W hat the essential feature of this barrier is, is hard tosay. Chestnut Ridge in many
respects forms a boundary. Altitude seems to play a part, but whether it is para-
mount is doubtful. Absence of extensive deposits of clay on the western side of
this ridge on account of the destruction of the Old Tertiary base-level by subsequent
erosion, may also be of importance. Further studies in West Virginia surely will
lead to a solution of the question, but this much is certain, that these two species
again illustrate the rule that “closely allied species occupy neighboring areas,” and
further they illustrate the fact that specific differentiation is-due to isolation, which
is topographical in this case.
What are the actual causes of the difference of the specific characters (color,
shape of rostrum, and sculpture of chelipeds), that is to say, what external influ-
ences are responsible for them is even more obscure, as it is in the case of the pro-
prngu USs-Zroup.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 511
(e) C. diogenes.
C. diogenes is sharply separated from the other chimney-builders, but resembles
them ecologically to a certain degree. In Pennsylvania it comes into contact with
them, but in the case of C. carolinus this has been observed only once, while it is
more frequent in the case of C. monongalensis. However, intermediate forms have
never been observed, so that we must assume that kyesamechania prevents crossing.
In both cases, with reference to C. carolinus as well as C. monongalensis, it is to
be remarked that whenever one of these is found associated with C. diogenes it
is always only a contact, not a real mixing of both forms. This is best observed in
the case of C. monongalensis and C. diogenes. All over the range of C. monongalensis
in southwestern Pennsylvania C. diogenes is also found. But as has been stated
(p. 417 and 458), although they frequently dwell at the same localities they do not
occupy the identical locations, C. diogenes belonging to a lower level than C. mononga-
lensis. ‘Thus we see again a separation, which is primarily expressed in the difference of
altitude. Whether the latter is most important seems doubtful. It has been stated
that C. monongalensis prefers spring-water, while C. diogenes lives mostly in swamps,
where the water is more or less stagnant and not so cool in summer. (Compare the
instance from Nine-Mile Run given above.) But, whatever may be the essential
feature which separates both species, it is clear that it is an ecological factor, and,
when these two species are found together, it is at a place where the ecological con-
ditions favorable to them come together.
That C. diogenes depends on different ecological laws from C. monongalensis is
also evident from the fact that the former has, outside of Pennsylvania, an entirely
different range.
Thus we have here a case similar to that of C. bartoni when it associates with the
burrowing forms. Two allied species occupy (in Pennsylvania) almost the same ter-
ritory, and are not separated topographically, but their ecological separation is evi--
dent, and very likely is connected with their specific differentiation.
In the two races C. diogenes, the eastern and western, we again see the influence
of separation. According to our theory that the area of C. diogenes was a unit in
Preglacial times, and that it was separated by the advancing ice into an eastern and a
western section, which subsequently remained separate, we must expect, if isolation
effects specific differentiation, that the eastern and western form of C. diogenes should
show at least a tendency to develop differential characters of specific value. This is
indeed the case, as we have seen above (p. 401 et seq).
Isolation, or Habitudinal Segregation, as the factor forming species, is thus
clearly seen in every case discussed. We may condense the results obtained in the
following sentences.
ile MEMOIRS OF THE CARNEGIE MUSEUM
1. The normal case is when two closely allied species, possessing identical or
nearly identical ecological habits occupy separated areas, which lie close together but
do not overlap. (Examples: propinquus-group ; C. carolinus and C. monongalensis.)
2. Whenever allied species are found in one and the same locality (overlapping),
isolation becomes apparent in the following forms.
(a) The two species have different centers of origin, that is to say, they were
separated formerly, but occupied the same territory subsequently. In this case, if
very closely allied, hybridization may be possible (C. obscurus and C. sanborni at
New Martinsville, and C. obscurws and C. propinquus in the Lake Erie drainage),
if no kyesamechania exists. If the latter is present, which always means that the
two species in question are less closely allied, the two species may actually live
side by side under identical conditions (C. bartoni and the river-species), or one may
conquer and suppress the other. No instances of the latter kind are known in
Pennsylvania, but may possibly occur in southwestern Ohio and in Indiana, between
C. rusticus and C. propinquus.
(b) If the centers of origin are more or less identical (absolute identity is hardly
possible), the two species always differ ecologically, and although living at the
same localities, prefer different surroundings. In this case they are not so closely
associated, and they generally remain ata certain distance from one another,
although their general areas are overlapping. Under such conditions hybridisation
might occur, but it has not been observed in Pennsylvania, and the species existing
under such conditions are probably separated by kyesamechania. (Example: C.
diogenes and monongalensis.)
Case (a) and (b) may be combined, that is to say, two species living together may
have different centers of origin and may be ecologically different. This is seen in
the example of C. bartoni and the burrowing species.
I believe that in every case where closely allied species overlap in parts of their
ranges a close investigation will reveal that one or the other of the above cases is
realized. Isolation is, in my opinion, a necessary factor in the differentiation of
species, and I do not think that a case ever will be discovered where two closely
allied species possess precisely the same distribution. But in order to ascertain this
a mere superficial knowledge of the species in question and their range is insuffi-
cient, and every case should be investigated as exactly as possible, in a manner
similar to the above studies.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 5138
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ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 7!
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Unperwoop, L. M.
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518
Fie.
Fic.
Fie.
Fic.
Fie.
Fic.
Fic.
Fic.
Fie.
oo bo
bo
la.
1d.
le.
If.
MEMOIRS OF THE CARNEGIE MUSEUM
EXPLANATION OF PLATES.
Puate A.
Cambarus obscurus Hagen. Male of the first form, natural size.. Collected by the
writer, Sept. 7, 1905, in the Alleghany River, Sandy Creek, Allegheny County.
Cambarus obscurus Hagen. Female, natural size. From same locality.
Cambarus diogenes Girard. Male of the first form, natural size. Collected by the
writer, Aug. 26, 1905, at Baden, Beaver County.
Cambarus carolinus Erichson. Female, natural size. Collected by the writer, Sept. 5,
1905, at Rainier Park, Ohiopyle, Fayette County.
PLATE B.
Cambarus bartoni (Fabricius). Male of first form, natural size. Collected by the
writer, Aug. 7, 1905, in Fern Hollow, Pittsburgh.
Cambarus bartont robustus (Girard). Female, natural size. Collected by Miss G.
Kinzer, Aug. 27, 1905, at Sixteen Mile Creek, Northeast, Erie County.
Cambarus limosus (Rafinesque). Female, natural size. Collected by the writer, Sept.
10, 1905, in the Schuylkill Canal, Manayunk, Philadelphia County.
Cambarus monongalensis Ortmann. Female, natural size. Collected by the writer,
Aug. 18, 1905, at Edgewood Park, Allegheny County.
PLaTE XXXIX.
Cambarus barton (Fabricius). Rostrum. All figures 4.
Female, 70 mm. long. Collected by the writer, June 3, 1904, in North Versailles
Township, Allegheny County, opposite Stewart. Catalogue number 74.327. Shape
very broad, margins parallel. Not rare in western Pennsylvania.
Female, 52 mm. long. Collected by the writer, Aug. 22, 1905, at Squaw Run, Alle-
gheny County. Catalogue number 74.626. Shape typical; very frequent.
Male, first form, 63 mm. long. Collected by the writer, Sept. 16, 1904, at Valley
Forge, Chester County. Catalogue number 74.413. Shape typical, and character-
istic of eastern specimens, but also found in the west.
Young male, second form, 34 mm. long. Collected by the writer, Aug. 22, 1905,
in Squaw Run, Allegheny County. Catalogue number 74.626. Usual shape in
young specimens.
Young female, 21 mm. long. Collected by the writer, June 25, 1904, in Jacob’s
Creek, Laurelville, Fayette County. Catalogue number 74.356. Slightly longer
than usual, but not rare in young specimens.
Male, first form, 78 mm. long. Collected by the writer, May 27, 1904, in Squaw Run,
Allegheny County. Catalogue number 74.320. Unusually short and strongly taper-
ing, with exceptionally thick margin.
Fic.
Fia.
Fie.
Fia.
Fic.
Fie.
3a.
4b.
6b.
Te.
ORTMANN: THE CRAWFISHES oF THE STATE OF PENNSYLVANIA 519
Cambarus bartoni robustus (Girard). Rostrum. 4,
Female, 89 mm. long. Collected by the writer Oct. 4, 1904, in Temple Creek, Albion,
Erie County. Catalogue number 74.435. Normal shape.
Young female, 18 mm. long. Collected by Miss G. Kinzer, Aug. 27, 1905, in Sixteen-
Mile Creek, Northeast, Erie County. Catalogue number 74.630. Normal shape in
young specimens.
Cambarus carolinus Erichson, Rostrum. 4.
Male, first form, 66 mm. long. Collected by the writer, Sept. 5, 1905, at Ohiopyle,
Fayette County. Catalogue number 74.640. Normal shape.
Male, second form, 30 mm. long. Collected by the writer, June 11, 1904, at Indian
Creek, Fayette County. Catalogue number 74.365. Shape exceptional. Most ex-
treme case as regards convergence of margins.
Cambarus monongalensis Ortman. Rostrum. 4.
Male, first form, 63.5 mm. long. (Type.) Collected by the writer, May 21, 1905, a
Edgewood Park, Allegheny:County. Catalogue number 74.316. Normal shape.
Female, 73 mm. long. Collected by the writer, Oct. 12,1904, at Hill, Westmoreland
County. Catalogue number, 74.449. Shape unusually broad, and margins almost
parallel. Most extreme case in this direction, standing rather isolated.
Cambarus limosus (Rafinesque). Left first pleopod of male, first form. 3.
Inner view. Collected by the writer, Sept. 19, 1904, in Marcus Hook Creek, Mar-
cus Hook, Delaware County. Catalogue number 74.423.
Posterior view of same.
Cambarus propinquus Girard. Left first pleopod of male. 3.
Inner view, male, first form. Collected by the writer, Oct. 4, 1904, in Temple Creek,
Albion, Erie County. Catalogue number 74.439.
Inner view, male, second form. Collected by the writer, June 7, 1904, in a tributary
of Conneaut Creek, Conneautville Station, Crawford County. Catalogue number
74.336.
Cambarus obscurus Hagen. Left first pleopod of male. 3.
. Inner view, male, first form. Collected by the writer, Aug. 24, 1904, in the Ohio
River, Ambridge, Beaver County. Catalogue number 74.401.
. Posterior view of same (horny tip of outer part hidden behind inner part).
Inner view, male, second form. Collected by the writer, June 24, 1904, in the Loyal-
hanna River, Crisp, Westmoreland County. Catalogue number 74.352.
. Inner view, hermaphroditic specimen (type of male, first form). Collected by Atkin-
son, Graf, and Williamson, May 14, 1899, in the Ohio River, Neville Island, Alle-
gheny County. Catalogue number 7.436. (See text, p. 376.)
Posterior view of same.
Fie.
Fie.
Fic.
Fie.
Fic.
Fie.
Fie.
Fic.
Fig.
Fic.
Fie.
Ne)
Iie
or
8a.
MEMOIRS {OF THE CARNEGIE MUSEUM
Cambarus bartoni (Fabricius). Inner view of left first pleopod of male, first form. 3.
Collected by the writer, Oct. 6, 1905, at Weskit, near Kittanning, Armstrong
County. Catalogue number 74.665.
Cambarus carolinus Erichson. Inner view of left first pleopod of male, first form. 4.
Collected by the writer, Sept. 7, 1904, at Dunbar, Fayette County. Catalogue
number 74.410.
Cambarus monongalensis Ortmann. Inner view of left first pleopod of male, first form
(cotype), #. Collected by the writer, Oct. 10, 1903, at Edgewood Park, Allegheny
County. Catalogue number 74.182.
Cambarus diogenes Girard. Inner view of left first pleopod of male, first form. 3.
Collected by the writer, September 5, 1904, at Smithfield, Fayette County. Cata-
logue number 74.406.
Pate XL.
Cambarus obscurus Hagen. Upper view of right chela of a male, first form, 77 mm.
long, natural size. Collected by the writer, Sept. 30, 1905, in the Alleghany River,
Twelve-Mile Island, Allegheny County. Catalogue number 74.663.
Cambarus bartoni (Fabricius). Upper view of right chela of a male, first form, 82 mm.
long, natural size. Collected by the writer, Nov. 22, 1905, in Fern Hollow, Pitts-
burgh, Allegheny County. Catalogue number 74.681.
Cambarus bartoni robustus (Girard). Upper view of right chela of a male, first form
98 mm. long, natural size. Collected by the writer, July 11, 1905, at Spartansburg
Crawford County. Catalogue number 74.596.
Cambarus carolinus Erichson. Upper view of right chela of a female, 77 mm. long,
natural size. Collected by the writer, Oct. 16, 1905, at Dunbar, Fayette County.
Catalogue number 74.669.
Cambarus monongalensis Ortmann. Upper view of right chela of female, 71 mm. long,
natural size. Collected by the writer at Edgewood Park, Allegheny County, April
4, 1905. Catalogue number 74.495.
Cambarus diogenes Girard. (Eastern form.) Upper view of right chela of a male, first
form, 83 mm. long, natural size. Collected by the writer, Sept. 21, 1905, at Ridley
Park, Delaware County. Catalogue number 74.654.
Cambarus diogenes Girard. (Western form.) Upper view of right chela of a male,
second form, 93 mm. long, natural size. Collected by the writer, April 15, 1905,
at Millvalle, Allegheny County. Catalogue number 74.507.
Burrow of Cambarus bartoni (Fabricius). In spring on hillside, west of Spruce Run,
Avalon, Allegheny County, opened by the writer, July 2, 1904.
Side view (section); 86. Upper view. ‘mp, pile of mud consisting of clay, sand, and
small stones; d, ditch; x, place where crawfish, female, 52 mm. long, was found.
At & springwater was running into the hole ina strong flow, and running out through
the hole, over and past the pile of mud into a ditch.
Fic. 9.
9a.
Fie. 1.
Fie. 2.
iGo
Fic. 4.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 521
Burrow of Cambarus carolinus Erichson, located in a swampy place in stiff yellow clay,
at Listie, Somerset County. Opened by the writer Aug. 12, 1904.
Diagram of disposition of piles of mud seen from above. 96. Section of hole along line
A-D. 9c. Section of hole along line A~B-C. A. Open chimney. B. Closed
chimney. D. Closed chimney, hole filled up a good distance below surface of ground.
C. Open hole, without pile of mud, situated under the edge of a large fiat stone (s).
wl, water level ; x, place where the crawfish (male, first form, 61 mm. long) was found.
PLATE XULI.
3urrow of Cambarus bartoni (Fabricius). Located in the sand and gravel of the dry
bed of a small stream, Edgewood Park, Allegheny County. Opened by the writer,
Oct. 10, 1903. mp, pile of mud, consisting of mud, sand, and gravel; s, large slab
of stone, lying imbedded in sand and gravel ; wl, water level (the stream was dry for
long stretches, only here and there pools of water were left); x, place where crawfish
(female, 63.5 mm. long) was taken.
Burrow of Cambarus monongalensis Ortmann, Located in yellow clay (mixed with
humus), at a springy place on the bank of small stream, near Monongahela City,
Washington County. Dug out by the writer, June 16, 1904.
Diagram of burrow and chimneys, seen from above; 26, section of hole along line
A-B_-C’; 2c, section of hole along line C-D-E. A, hole opening laterally, with
one-sided pile of mud in front, keeping up the level of water; B and D, closed
chimneys ; C, open, large, and regular chimney ; w/, water level ; st, stream ; 2, places
where the old female (mother, 65 mm. long), and ten young (20.5 to 32.5 mm. long)
were fouud. Water, in a weak flow, was running in at “and wasrunning out at A.
Burrow of Cambarus monongalensis Ortmann. Located in yellow clay, at a springy
place on the bank of a small stream, Edgewood Park, Allegheny County. Dug out
by the writer, May 9, 1904. The burrow is of a type similar to the one figured in
Fig. 2, but less complex. a, hole opening laterally, with one-sided pile of mud keep-
ing up the level of the water ; b, closed chimney ; w/, water level ; st, stream ; x, place
where the crawfish (female, 63 mm. long) was taken.
Burrow of Cambarus monongalensis Ortmann. Located in black muck, at a springy
and swampy place at the bottom of the upper part of Fern Hollow, Pittsburgh, Alle-
gheny County, opened by the writer, Oct. 18,1903. Type of a hole in level ground,
with the water near the surface. No adults and only four young were found in this
hole, but possibly the hole had additional branches, which were not discovered, the
high stage of the water and its icy coldness rendering investigation difficult. About
1.50 m. from this hole another was opened, which contained a female C. diogenes.
a, closed chimney ; 6, one-sided chimney in front of hole opening obliquely ; w/, water
level ; x, places where young specimens (11.5 to 16,5 mm. long) were found.
Fic.
Gane
Fie.
Fie.
Fie.
nie?
5 Se
6.
8.
(J)
MEMOIRS OF THE CARNEGIE MUSEUM
Burrow of Cambarus diogenes Girard. Located in stiff blue clay, in a ditch on a road-
side, Nine-Mile Run, Pittsburgh, Allegheny County. Opened by the writer, Nov.
5, 1904. The season had been very dry, and not much water was in the hole.
Pebbles were lying on the bottom of the hole. a, old chimney, leveled down by rain,
probably built in spring; 6, fresh mud, brought up recently (beginning of fall
activity) ; wl, water level ; x, place where the specimen (female, 77 mm. long) was
taken.
Burrow of Cambarus diogenes Girard. Located in yellow clay and humus, at a springy
and swampy place in woods on the side of a wagon road, upon which water was
standing (after a heavy thunder-shower on the previous day), at Squaw Run, Alle-
gheny County. Dug out by the writer, May 27, 1904. a, chimney, consisting of
’ in the mouth of the chimney, distinctly differing from the
yellow clay ; 6, “stopper ’
chimney, the material being yellow clay mixed with blackish mud and leaf-mould ;
wl, water level ; 7, road, with mud-puddle upon it ; w, place where the crawfish (male,
first form, 76 mm. long) was found.
Burrow of Cambarus diogenes Girard. Located in yellow and blue clay, on the border
of a swampy place, Schenley Farm, Pittsburgh, Allegheny County. After a sketch
furnished by Mr. F. E. Kelly, Nov. 14, 1904. sw, swamp; be, blue clay ; ye, yel-
low clay ; a, one-sided chimney, consisting of yellow clay (probably made in spring
and summer) ; 6, new chimney, consisting of blue clay (fall activity, reclaiming of
old burrow at ¢) ; ¢, old burrow, filled in (during summer) with blue clay, taken or
washed in from near the mouth of the lower entrance (a) of burrow ; w/, water level ;
x, place where the crawfish was taken.
Burrow of Cambarus diogenes Girard. Located in blue and yellow clay on the bank
of a small stream, Schenley Farm, Pittsburg, Allegheny County. After a sketch
drawn by Mr. F. E. Kelly, Nov. 15, 1904. s, stream ; bc, blue clay ; yc, yellow
clay ; a, new chimney, consisting of yellow clay, evidently coming from the newly dug
shaft going down vertically ; 6, upper end of ascending branch of hole, without open-
ing (possibly originally open, but sealed up, and the pile of mud overgrown and
obliterated by vegetation) ; wl, water level ; x, place where the crawfish was taken.
The chimney at @ shows fall activity, and the vertical shaft is being built by the
crawfish in order to get deeper down into the ground.
PuatE XLII.
Preglacial Monongahela River, after Leverett (1902, p. 89, fig. 1).
Present range of Cambarus obscurus Hagen and C. propinguus Girard. (Including
variety sanborni (Faxon)).
Distribution of Cambarus propinquus Girard, propinquus sanborni (Faxon), and C.
obscurus Hagen.
(For further explanation, see legend on map, and text, p. 433-446).
Til.
IV.
ORTMANN: THE CRAWFISHES OF THE STATE OF PENNSYLVANIA 523
Puate XLITI.
Map of Pennsylvania, showing distributional areas of crawfishes. (See Legend on
map, and text, p. 465-466).
CONTENTS.
PAGE.
Inrropuction: Scope or Work; Mretruops; ACKNOWLEDGMENTS ............- 343
Histortcan REVIEW OF ouR SysTEeMatTic KNOWLEDGE OF THE CRAWFISHES OF
BENING WV TaVAAINUAG eee cine ai nen Bese cise teraiace cas URDU REE Deh ira Mnent heval ty tt Uk wat Rai 348
MorPHoOLoGy AND CHOROLOGY OF THE PENNSYLVANIA SPECIES ............. 350
PAR G enreralmvemlarkswt nesses scence ese eee alee aise sees BAS) ses SEGUE SUE SURE Toe Sleere ELSE MER LO 350
Beeyatortheennsylvanianspectesuerrie is arene ei ae eine oho eiaeic 351
C. Description and Distribution of the species ......... 0.0.00. eee ee eee eee 352
Pa C amibarisi(Haxonins)plimosus) (vats) perrere irs ere erie ete crue an tener 352
4; (Canmlorins (Meow) joey Cite. gaboseseohaecccsuoconnded non cod 358
2a. Cambarus (Faxonius) propinquus sanborni (Fax.) (extralimital) ........ 365
3) Chimlaiis (EMins) Costin BEG 5 oop ocogoecocoup bo GG occoDodCenpoS 369
aio (Crumilogietisy (sencionyins)) logit (NS) ooonn oso odes odes b0cucdaeo6sau0sc005 377
4a. Cambarus (Bartonius) bartoni robustus (Gir.)........... sxc Sea ts aieh pious ea 388
H, Camloyanis (acio@mns) corals Wis covoceboodootasocpodsccvosooDenoos 394
6. Cambarus (Bartonius) monongalensis Ortm...................-065 ewitnieGg Ao)
7. Cambarus (Bartonius) diogenes Gir. ............+4.- ER een eas edu iaades A02
EcoLoGy AND GEOGRAPHICAL DISTRIBUTION................-: demesne ifn 3 66 0 410
PASE COLODayaay Sasi stale faye feet Lbeiai ays) cisions) Sts a) nl sees fenay stele aca ayes Aer eet Gi \ ios cat nner ee 410
1. The River-Species : C. limosus, propinquus, obscurus .................... 410
2. The Mountain-Stream Species: C. bartoni..............-...02- seeeee- 413
3. The Burrowing Species: C. carolinus, monongalensis, diogenes............. 416
Gem General Ealoibati anne dese ee Monks gehen auc rela ety cies SeLSLUS ES Reiieth yr ince aU ee 416
Ge ShaperorBurrows iy als ocular ees ol elergistisroseiey cueieps aos Weekes eee et eae 418
CHi@onstructroncole Burrowssc oeneaeeeiee ek Gone Oe ene eee 419
BrGecoorap ical Distrib ations yoo sje si als iau+ ayers olsi 285) peo Savi ayes aeere neem Ted 425
iP Cambarussiimosuseepieerenr ein ee TAEDA ee Huieoseecee emia eID)
CPMOUTOTN AY Off LACUS ete ye cere ay hee castes yt nee eee ele eTe eR OR eager ea 425
GreOrioinioteGistributlonyic%.crates-c erie wok erc tr) aces ated eae oes aereanee ne een 425
2. Cambarus propinquus, sanborni, and obscurus ........... Rene ase eae o6 433
Gr, SMM? OGG 5 s56cgodoccad¢ocnde SG eitelie ipuprese sca ar nets tres 433
GomOricoimuot CishribMPtON rieececbs vs yc scissile uslivonsenehe aps geye te ce ls coy ee Mepem ats vee. 404
Ja OAM HALUSHOALLOML Sies terete ce aa ars Ace Seetoes col ees louse Bn ere ad Re ee RO 447
524 MEMOIRS OF THE CARNEGIE MUSEUM
braOriginvofadistribution wre yesimn, cos tran ae a tore ie seer yee AAT
Au Cambarusabartoniprobustuswasisecesmis SoCo ee oer kote eee en 449
GE TSUMIMALy FOL MACtS weeerg sas ee ae Chan mee ees i ned Tee eee 449
beOrioimmoiidistribubonveere sie eae ene cee ete os oor lite emote ee omneree 449
Hey Cambarusicarolimuse seta secatstoe cn koa ays ae eine covecoee weiner cae ane camepe eet mS 451
Ge SUM MAL Yes OL LACUS a ep syee ce ee hese oss cya ene RCo cee omscer ee Relic a Ie eer egos 451
bvOrioingotadistribution sc seese se eee eee ea ae ee 452
6seCambarusmmonongalensismmei eemee aero ae iene 453
GC POuMMAary, (Ol factSin cami sists a sree crews Sie apt recta a are een eros 453
bss Origin zotadistributionce nc. cers seuss ono oso are eA a eC 454
(2 Cambarus diogeness aia aihorcrreeioset ois oe CAG eG oun AOR ee meena Oe 457
GQ OUM Mayle actsyc mest Petals cucesien eis) sire sera Aaa Te re ese eR eae 457
eOricinvofedistributione ss: aces ay aes race eee ete eons 459
Se jummanyaofestudvestonudistributioneen ae eee ere eee ae ese orc 462
Vem errs ETT SMO RAY eresesye vey ape yous auc date) at taped auch -acrrtcbalg pam et Splat ten erearaar ale sure re Ree eg 469
ie Cara bavustOWSCULUS|s nieratccuns, arsine suchas aero aer reas Merc neR ee oan oie eerie oe 470
2. Cambarus propinquus and C. propinquus sanborni...............-..-...- 476
S53 Caml barus li mMOSisicsc. isos co sive: seuss deccuebetercasney aio cdo So ce coer ere Neca eee weno AT7
Al Cambarus: Gio gemes|scs fe % ico hn yas 35 vies Bs wien a) ARR os outs sree ty ere terete 480
5. Cambarus bartoni and C. bartoni robustus............-.2+.0..00...--. 486
6. Cambarus carolinus and monongalensis.............--......+++-++++--:: 489
AV ANies A CON OMG AW CAUG WEN fara feran cae) cuca 08 So, sulsie nats aa aReee ame aI oo SSA OP ome eet aseee 492
ipeeopularsknowled seo ficrawtish esac arec hirer re teeter erm on ae 492
Jpeulthewuselotacrawtlishes asi oodsands bait ae enoee ernie cc eee 493
3. Crawfishes as scavengers ; their food; their enemies...............-+.... 494
Awe @rawiisheslasro bnoxious\creaturess-m-pe ereineaeeicr eee eee 496
VII. BEARING OF THE ABOVE STUDIES ON THE THEORY OF EYOLUTION...........--- 497
jp uhherVintation—heonyoteDeclViriesseeeee ene cece eer 498
2. Species, Varieties, and Variations among the Pennsylvania Crawfishes...... 498
3. Formation of Species by Isolation, as exemplified by Pennsylvania Crawfishes. 504
PTB TE TOG RA Payor oe rs ts ite Menhees aaa oes iselfere ATS ete a ee eae St ee ene eee 513
EXP ANATION: O Rick TPAC DES ate, ‘<5 teicnaceits, rncan peaerelste ee oats ein egal ceo Ue TCU Reames 518
PLATE A,
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MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XXXIX,
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MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XL.
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scale: 50 cm.
CHEL® Or Cambarus, Fras. 1-7. Burrows or Cambarus, Fries. 8-9.
MEMOIRS CARNEGIE MUSEUM, VOL. II. PLATE XLI.
scale: 50 em.
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Burrows or Cambarus.
MEMOIRS CF THE CARNEGIE MUSEUM, VOL.I PLATE XLII
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Fig 1 Preglacial Monongahela Fig. 2. Range of C. obseurus& propinguus
Fig. 3 Distribution of C. propinquus, propinquus canborni, obscurus
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(A separate table of contents is provided for the article on the “The Crawfishes of the State
of Pennsylvania” by Dr. A. E. Ortmann on p. 523.)
Abbott, J. DeB., 419
Academy of Natural Sciences of Philadelphia,
347
Aceratherium, 174
Acichelys, 290
Additional Remarks on Diplodocus, 72-75
Agate Spring Stock Farm, Sioux Co., Ne-
braska, 316
Agriculture, Department of, Harrisburg, 348
Agriocheerus, 205, 211
minimus, 220
Albion, Erie County, Pennsylvania, 361
Alleghany County, Maryland, 344
Allosaurus, 57.
American Jurassic Ichthyosaurians, 325
Amiopsis dartoni, 80
Andrews, Dr. C. W., 225, 244, 259
Apatosaurus, 14
Aplodontia, 142, 143, 144, 148, 149, 151-
153, 156-165, 178, 181-184
rufa, 139, 142
Archelon, 280, 284, 290
ischyros, 280
Astacidee, 350
Astacine, 350
Astacus affinis, 348, 352, 377
bartoni, 348, 352, 377
(cambarus) carolinus, 394
ciliaris, 377
fossor, 348
limosus, 348, 352
pusillus, 377
Astarte packardi, 79
Atkinson, Dr. D. A., 413, 416, 450 473, 476,
485
Atlantosauridse, 49
Atlantosaurus, 14, 232
Oo
OL
Atlantosaurus, beds, 63, 64, 68-70, 72, 78,
80
Aux Plains River, Illinois, 363
Bad Lands, White River, Dakota, 212
Baptanodon Beds, 70, 78, 80, 81, 336, 337
Baptonodon, 126-128
discus, 78-80, 82, 84, 98-106, 108, 113,
114, 118, 121, 123, 330, 332, 335,
336, 338, 340, 342
marshi, 79, 103-105, 110, 113, 114, 119,
121-123, 330, 33
natans, 78, 79, 117-121, 327, 331
robustus, 330-334
Barbour, Dr. E. H., 62, 63, 185
Bartonius, 350, 351
Battle Creek, Michigan, 363
Bauer, Franz, 82
Baur, Dr. George, 44, 86, 127
Bayet Collection, 133
Belemnites densus, 79
Benton Beds, 33
Cretaceous, 336
Beecher, Dr. C. E., 78
Beulah Shales, 67
Bighole River, 209, 211
Big Horn Mountains, Wyoming, 62
Black Tail Deer Creek, Montana, 176
Bothriospondylus, 51, 55
Bothrolabis, 305, 308
tricheenus, 306
Boulenger, Dr. G. A., 225
Brachiosaurus, 15
Brachylophus fasciatus, 254
Braeburn, Pennsylvania, 380
Brontosaurus, 14, 17, 18,523, 25, 27, 34, 39,
46, 49, 75, 255, 263
526 INDEX.
Brontosaurus excelsus Marsh, 7, 15
Bunotheria, 213
Camarasaurus, 65, 68
supremus, 65
Cambarellus, 350
Cambarus bartoni, Description of, 377
Distribution of, 381
Remarks on, 386
earolinus, Description of, 394
Distribution of, 396
Remarks on, 397
diogenes, Description of, 402
Distribution of, 405
Remarks on, 407
limosus, Description of, 352
Distribution of, 356-358
Remarks on, 358
monongalensis, Description of, 398
Distribution of, 400
Remarks on, 401
obscurus, Description of, 369
Distribution of, 372
Remarks on, 374
propinquus, Description of, 358-362
Distribution of, 362-863
Remarks on, 364-365
robustus, Deseription of, 388
Distribution of, 390
Remarks on, 391
Key to the Pennsylvanian species of, 351
bartoni, 344, 349, 393, 413-415, 418-
440, 443, 451, 452, 465, 466,
469, 476, 486-490, 498, 494,
496, 508-512
bartoni robustus, 349, 351, 377, 388,
393, 416, 518-520
diogenes, 352, 402, 416, 509-511,
518, 522
monongalensis, 351, 398, 510-512,
518-521
blandingi, 431, 469
carolinus, 346, 394, 404, 416, 419, 420,
451, 452, 457, 464-466, 489, 495-
497, 500-502
diogenes, 349, 414, 415-420, 423, 424,
453-463, 465, 480-485, 489, 490,
49.7, 500, 501
dubius, 349, 394, 398, 402
indianensis, 503
limosus, 344, 351, 352, 380, 410, 415,
425, 426-432, 445, 461, 463, 464, 467,
469, 470, 473, 475-478, 481, 486,
492, 499, 501, 508, 505, 509, 519
monongalensis, 346, 349, 414, 416, 417,
419, 420, 423, 424, 453-461, 465,
466, 489-492, 495, 497, 500, 501,
508, 509-512, 518-521
montanus, 377
obesus, 402, 501
obscurus, 349, 351, 361, 362, 366, 368,
369, 380, 410, 412, 483-448, 463-
468, 470, 474, 477-485, 489, 492,
495, 499, 502, 505-509, 512, 518-520
pealei, 352
pilosus, 419
propinquus, 349, 350, 351, 358, 359,
366, 368-371, 380, 410, 433, 434,
436, 438, 439, 441, 442, 445, 465,
476, 492, 499, 501, 505-507, 509,
512, 519
propinquus sanborni, 350, 365, 368, 433,
434, 436, 439, 446, 476, 477, 492,
499, 505, 506, 507, 522
robustus, 450, 500
rusticus, 349, 369, 438, 466, 508
Camden County, New Jersey, 344
Camptosaurus, 293
Canide, 179
Canis, 174
Caiion Ferry, Montana, 212
Canyon City, Colorado, 4, 5, 6, 138, 57, 58,
61, 63
Cardioceras cardiformis, 79
Cardiodontide, 49
Carnegie, Andrew, 2, 133, 226, 264
Carroll County, Ohio, 344, 367
Castor, 139, 143, 144, 145, 152, 153, 156, 157
canadensis, 153, 176
fiber, 144, 145
Ceratogaulus rhinocerus, 179, 181
Ceratosaurus, 57
INDEX. 527
Cetiosauria, 47, 48, 55 Crawfishes of the State of Pennsylvania, Arnold
Cetiosaurus, 52, 53 E. Ortmann, 343
leedsi, 254 Popular knowledge of, 492-493
longus, 52 river species, 410
Chelone, 289, 294 species, varieties, and variations among
Cheloniid, 280, 296 the Pennsylvanian, 498—504
Chelonoidea, 296 the use of, as food and bait, 493-494
Chelydra, 290, 294 Crocodilia, 47
Chondrosteosaurus, 50 Cruikshank, Dr. O. T., 450
gigas, 50 Cryptobranchus allegheniensis, 495
Cimoliosaurus laramiensis, 79 Cryptoclidus, 260
Cockerell, Professor T. D. A., 407 oxoniensis, 259
Coggeshall, A. S., 78, 140, 226 Cryptodira, 295
Coggeshall, L. S., 3 Cuvier, R., 82
Colodon, 211 Cylindrodon, 211
Colpochelys, 290, 293, 295 Cynomys, 151-154, 156-170, 178, 181
kempi, 294 ludovicianus, 139
Como Beds, 67
Comptonectes bellistriata, 79 Deemonelix, 139, 177, 185-190, 202
extenuata, 79 Davison, Professor A. E., 425
Conotton Creek, Ohio, 367 Dentalium subquadratus, 79
Cook, Harold J., Jr., 316 Dermochelydide, 296
Cope, E. D., 59, 66, 68, 82, 86, 127, 175, 174, Dermochelys, 286, 288, 290, 293
185, 213, 305, 310 coriacea, 304
Cosoryx, 174 Derry, Pennsylvania, 380
teres, 174 Description of New Rodents and Discussion ot
Coturnix noyee-zelandiz, 133 the Origin of Demonelix, 139-191
Crawfishes as obnoxious creatures, 496, 497 Desmatochelydinz, 296
as scavengers, 494—496 De Vries, Hugo, 503
Bearing of the study of, on the theory of Mutation theory of, 498
evolution, 497 Dianthera americana, 412
Bibliography of, 513-517 Dicotyles tajacu, 309, 311
Collectors of, 347, 348 Didelphys, 213
Ecology and Geographical Distribution Dinosauria, 5, 20, 44, 45, 46, 47, 74
of Pennsylvanian species of, 410—469 Diplodocide, 49
Economic value of, 492-497 Diplodocus, 5, 6, 9, 14, 17, 23, 25, 27, 34, 35,
enemies of, 494 39, 45, 46, 49, 55, 57, 59, 73, 75, 226,
food of, 494 227, 233, 235, 236, 239, 245, 246,
formation of species by isolation, 504 249, 264
life history of the Pennsylvanian species carnegiei, 7, 11, 15, 18, 40, 74, 75, 228,
of, 469-492 250
morphology and chorology of the Penn- longus, 75, 251
sylvanian species of, 350-410 vases, 264
mountain stream species, 413 Diprodontia, 48
historical review of our systematic knowl- Dogtown Mine, Montana, 212
edge of, 348-350 Dolichorhynchops osborni, 85
528 INDEX.
Dollo, M., 295
Douglass, Earl, 139, 213
Tertiary of Montana, 204—223
Eastman, C. R., Fossil Avian Remains from
Armissan, 131-158
Echidna, 208
Edward VIL., King of England, 226
Elotherium, 209
Entoptychus, 182
Eosphargis, 296
Eretmochelys, 286, 290, 295.
EKuhapsis platyceps, 179-183
Eumys loxodon, 174
Eutheria, 208
Eutherian Mammalia, 48
Exposition Society of Western Pennsylvania,
226
Faxonius, 350, 351
Felch, M. P., 5
Fern Hollow, Pittsburgh, Pennsylvania, 388
Filholornis, 131
debilis, 132
gravis, 132
paradoxa, 132
Fishing Creek, West Virginia, 368
Fort Pierre Cretaceous, 284
Fort Union Beds, 67
Fossil Avian Remains from Armissan, 131-138
Fowler, H. W., 478
Francolinus pictus, 133
Frass, E., 82, 127
Freeze Out Hills, 69, 71
Freeze Out Mountains, Carbon Co., Wyoming,
325
Galinsoga, 494
Gallia narbonensis, 134
Gallinuloides wyomingensis, 132
Gallus, 131
esculapi, 133
brayardi, 133
sp., 133
Garden Park, Colorado, 5
Geomyide, 181
Gera, H., 478
Gidley, J. W., 225, 239
Gill, Dr. Theodore, 225
Gilmore, C. W., 71, 72, 98, 116, 127, 225,
239, 336
Gilmore, Charles W., Notes on Osteology of
Baptanodon, 325-337
Osteology of Baptanodon (Marsh), 77-129
Goniomya montanensis, 79
Grammatodon inornatus, 79
Great Oolite, 52, 71
Green River, Edmonson County, Kentucky,
363
Green River limestone, Wyoming, 131
Grim, J., Jr., 308
Gryphea nebrascensis, 79
Hackberry Creek, Gove Co., Kansas, 282
Hallopus Beds, 78
Hancock County, West Virginia, 344
Haplocanthosauride, 54
Haplocanthosaurus, 2, 5, 6, 14, 17, 21, 24, 39,
40, 44, 45, 46, 48-54
priscus, 4, 6, 10, 22, 27, 28, 32, 34, 35,
38, 43, 49, 54, 75
utterbacki, 4, 11, 27, 28; 36, 39, 42, 43,
49-51, 54, 75
Harrison Beds, 139, 202, 308
Harrison County, Ohio, 344
Hatcher, J. B., 77, 78, 79, 81, 127, 204, 225-
227, 246-252, 255-257, 260-263, 306,
320
Hatcher, J. B., Additional Remarks on Diplo-
docus, 72-75
Osteology of Haplocanthosaurus, with
Description of a New Species, and re-
marks on the probable habits of the
Sauropods and the Age and Origin of
the Atlantosaurus Beds, 1-72
Heer, Oswald, 185
Helodermoides, 205, 211
Hendley H. W., 325
Hesperomys loxodon, 174
Hippotherium, 174
Hoactzin, 131
Holland, Dr. W. J., 133, 140, 204, 281, 320,
325, 348
INDEX. 529
Holland, Dr. W. J., Osteology of Diplodocus
Marsh, With Special Reference to the
Restoration of the Skeleton of Diplodocus
Carnegiei Hatcher, Presented by Mr.
Andrew Carnegie to the British Museum,
225-264
Hollidays Cove, Hancock Co., West Wirginia,
387
Hulke, J. W., 127
Hyzenodon, 205, 211
Hyotherium, 305
Hyracodon, 205, 211
Ichthyopterygia, 82, 126-128
Ichthyosaurus, 82, 84, 85-87, 94,98, 99, 116—
118
campylodon, 124
communis, 90
longifrons, 86, 105
quadriscissus, 86, 89, 92
Ictops, 210, 211
acutidens, 213, 214, 218, 222-224
bicuspis, 213
bullatus, 214
dakotensis, 212, 214, 223
didelphoides, 213
intermedius, 217
major, 205, 219-224
montanus, 214, 217-224
tenuis, 217-221, 224
thompsoni, 214, 219, 222, 223
Iguana tuberculata, 20254
Iguanodon, 246
bernissartensis, 244
Insectivores, 208
Ischyromys, 205, 211
James, Joseph T., 185
Jennings, O. E., 140, 441
Notes on the Vegetable Tissues in Deemone-
lix, 190-191
John Day Formation, 183, 305, 306, 310,
319, 320
Jurassic, 45, 46, 61, 64, 117, 325
of Colorado, 6
fossils, 79
Jurassic, of North America, 77
Kelly, F. E., 423, 482
King Edward VII., 226
Kinzer, Miss Grace, 488
Kirtlington Station, England
Knight, Wilbur C., 69, 78, 80, 118-121, 127,
332
Lafayette College, Easton, Pennsylvania, 347.
Lagopus albus, 133
mutus, 133
Lake Douglass, Michigan, 363
Lankester, E. Ray, 410
Leeds, C., 125
Leidy, Joseph, 77
Leptictide, of the Lower White River Beds,
208, 210
Leptictis, 213
haydeni, 212, 214
Lesley, J. P., 279
Lima sp., 79
Limnenetes, 205, 211
anceps, 220
platyceps, 220
Linesville, Crawford County, Pennsylvania,
440
Lingula brevirostris, 79
Little Medicine Bow, Wyoming, 6
Loomis, Dr. F. B., 79, 127
Loup Fork Miocene, 306
Lower Madison Valley, Montana, 177
Lower Pliocene, 131
Lutra, 261, 263
Lutraria senna, 185
Lydekker, R., 99, 124-127
Lytoloma, 286, 294, 295
Mammoth Caye, Kentucky, 380
Manatee, 56
Marsh, Othniel Charles, 2, 5, 6, 14, 44, 47,
48, 49, 56, 59, 62, 66, 67, 68, 77, 113,
116, 127, 225, 227, 229, 232, 238-243,
250-252, 266, 268, 270, 272
Marshall County, West Virginia, 344
Marsupials, 208, 213
Matthew, Dr. W. D., 78, 225
McCarty’s Mountain, 204, 214
530 INDEX.
McConnell, W. R., 358, 426, 441, 474, 478
Megalneusaurus rex, 79
Megalosauria, 47
Meleagris antiquus, 131, 132
celer, 123
superbus, 133
sp. ind., 133
Menopoma, 19, 51
allegheniensis, 20
Merriam, Dr. J. C., 98, 104, 128, 332, 336
Merrill, Dr. George P., 225
Merychyus, 174
Mesodectes, 213
Mesohippus, 211
latidens, 220
Mesoreodon, 139, 179
Metamynodon, 209
Metatheria, 47, 48
Micropterus, 494
Middle Miocene, 131
Island Creek, West Virginia, 367
Miocene, 131
Middle White River, 210
Mineral County, West Virginia, 344
Modiola sp., 79
Morgan County, West Virginia, 344
Morosauride, 49, 54
Morosaurus, 5, 13, 14, 17, 27, 34, 39, 42,
49
grandis, 14
Morrison Beds, 67
Multituberculates, 208
Mylagaulide, 179
Mylagaulids, 172, 181
Mylagaulus monodon, 179
Natrix sipedon, 495
Necturus maculosus, 495,
New Hagerstown, Ohio, 415
A New Monotreme-Like Mammal, 204
New Suilline Remains from the Miocene of
Nebraska, 305-320
Niobrara Cretaceous, 296, 300
Niobrara River, Nebraska, 308
Nopsca, Baron F., 54, 262
Notes on Osteology of Baptanodon, 325-33
Notes on_the Vegetable Tissues in Demonelix,
190-191
Oberlin College, Ohio, 355
Oil Creek, Colorado, 5
Oligocene, 131
Ophthalmosaurus, 71, 111, 112, 126, 128, 336,
337
cantabrigiensis, 119, 125, 336
discus, 121
icenicus, 119, 120, 123, 336
natans, 120
Opisthocelia, 47, 48
Opisthocomus, 131
Oreodon Beds, 209, 210
Ornithopoda, 47
Ornithopsis, 55
Ornithorhynchus, 208
Ortalis, 131
Orthopoda, Cope, 47
Ortmann, Dr. Arnold E., The Crawfishes of
the State of Pennsylvania, 343
Osborn, H. F., 18, 55, 67, 69, 72, 225, 227,
228, 244, 247, 252, 253, 255
Osteology of Baptanodon (Marsh), 77-129
Osteology of Diplodocus Marsh, with Special
Reference to restoration of Skeleton of Dip-
lodocus Carnegiei Hatcher, Presented by Mr.
Andrew Carnegie to the British Museum,
225-264
Osteology of Haplocanthosaurus, with Descrip-
tion of a New Species, and Remarks on the
Probable Habits of the Sauropoda and the
Age and Origin of the Atlantosaurus Beds,
1-72
Osteology of Protostega, 279-298
Osteopygis, 294
Ostrea engelmanni, 79
Owen, R., 82, 89, 128
Oxydactylus, 139
Paleictops, 213
Paleocheerus, 305
Paleolagus, 211
Paleoperdix, 131
longipes, 132
INDEX. 531
Paleoperdix, prisca, 132
sansaniensis, 132
Paleortyx, 131
blanchardi, 132
brevipes, 132
cayluxensis, 132
edwardsi, 132
gallica, 132
hoffmani, 132
media, 132
phasianoides, 132
Paleotetrix gilli, 133
Paleotherium, 209
Panhandle of West Virginia, 372
Pantosaurus striatus, 79
Paracambarus, 350
Parastacine, 350
Paris Basin, 131
Paulmier, F. C., 467
Pedioceetes lucasii, 133
nanus, 133
phasianellus, 133
Penns Manor, Pennsylvania, 355
Pentacrinus asteriscus, 79
Peratherium, 211
Perdix sp., 133
Peterson, O. A., 81, 137
Description of New Rodents and Discus-
sion of theOrigin of Deemonelix,139-191
New Suilline Remains from the Miocene
of Nebraska, 305-320
Phacocheerus eethiopicus, 310
Phasianide, 135
Phasianus, 131
altus, 132
archiaci, 133
desnoyersi, 132
Philadelphia, Pennsylvania, 344
Pholodomya kingi, 79
Pholidophorus americanus, 80
Piedmont, South Dakota, 6
Pilsbry, H. A., 427
Pinna sp., 79
Pipestone Springs, Montana, 215, 214
Pittsburgh, Pennsylvania, 343, 347
Plates, Explanation of, 129, 138, 194, 196,
198, 200, 202, 224, 518
Platygonus leptorhinus, 308, 310
Pleistocene, 131
Plesiosaurus shirleyensis, 79
Pleuroccelide, 49
Pleuromya subcompressa, 79
Polychrus marmoratus, 254
Potamobiide, 350
Potamobius, 350
Potomac River, 355
Prairie Dogs, 188, 189
Predentata, 44, 45, 47, 48, 72
Prentice, Sydney, 3, 78, 140, 204, 283, 320
Preston County, West Virginia, 344
Procambarus, 350
Procamelus, 174
Proccelia, 47
Promerycocheerus, 139, 179
Protohippus, 174
Protostega, 290, 292, 293, 294, 295
gigas, 279-292, 300, 302
Prout, Dr. Hiram, 209
Psephophorus, 296
Pseudomonotis curta, 79
orbiculata, 79
Purbeck, 72
Quercy Phosphorites, 131
Rathbun, Miss M. J., 350
Red Fork of Powder River, Big Horn Co.,
Wyoming, 81, 333
Reed, W. H., 123, 325, 329, 336
Rocky Mountains, 23
Rosebud Beds, 174
Rough Run, Butler County, Pennsylvania, 391
Rumex acetosella, 494
Sauranodon, 117
Sauropoda, 16, 19, 21, 24, 39, 40, 43-47, 49,
51, 55, 57, 59
Sauropterygia, 259
Schenley Park, Pittsburgh, Pennsylvania
Sciuromorphs, 170
Scott, W. B., 69, 139 -
Seeley, H, G., 89, 128, 259
532 INDEX.
Septaria, 79
Shastasaurus, 104, 105, 107
Sinclair, W. J., 305
Smoky Hill River, Kansas, 279
Sphenodon, 92, 245
Spirophyton, 185
Stark County, Ohio, 344
Stegosaurus, 57
Steneofiber barbouri, 166, 167, 169, 171, 175
178, 185, 194, 196
complexus, 177
eseri, 144
fossor, 140, 141-144, 147, 150-154, 156—
159, 170, 175, 177,178-187, 194, 196,
198, 200
gradatus, 166, 173, 178
hesperus, 176
montanus, 157, 162, 175, 176
nebrascensis, 172
pansus, 146, 147
peninsulatus, 144, 145, 172, 175
platyceps, 194
viciacensis, 144, 145, 172, 177
Sternberg, Charles, 281
St. Louis Fur Company, 209
St. Mary’s Lake, Michigan, 363
Struthio, 263
Surface, H. A., 495
Sus scrofa, 310
Talpa europea, 189
‘Tancredia extensa, 79
inornata, 79
warrenana, 79
Taonurus, 185
Taoperdix, 131, 134
keltica, 132, 134, 138
pessieti, 132, 134-138
Templeton, Pennsylvania, 387
Tertiary of Montana, 204—223
Tetrao pessieti, 134
tetrix, 133
urogallus, 133
Theropoda, 44, 45, 46-48, 72
Thinohyus, 305
brachyceps, 306
Thinobyus, decedens, 318
lentus 4, 319
osmonti, 320
pristinus, 314, 319
rostratus, 319
siouxensis, 308, 309, 310, 316, 319, 322,
324
socialis, 319
subeequans, 306, 319
Thracia weedi, 79
Three Forks, Montana, 220
Titanotherium, 205, 209, 211, 220
Titanotherium Beds, 204, 213, 214, 217
Todd, W. E. C., 440
Toretocnemus californicus, 332
Torresdale, Pennsylvania, 355
Toxochelys, 281, 288-290, 295
bauri, 295
latiremis, 293
Triassic of Colorado, 6
Trigenicus socialis, 220
Trigonias, 209
Troublesome Creek, Carbon Co., Wyoming, 81
Tucker County, West Virginia, 344
Twelve Mile Island, Pennsylvania, 371
Tympanuchus pallidicinctus, 133
Tyranosauridee, 49
Uinta, 299
Ungulata, 48
Unio, 57
Upper Eocene, 131
Upper John Day, 306, 320
Upper Miocene, 139, 177
Utterback, W. H., 2, 3, 4, 5, 6, 62, 65, 72,
81, 227, 252, 253, 255, 278, 333
Uhler, Dr. P. R., 393, 484
VanKirk, A. W., 3, 140
Varanus niloticus, 254
Warbonnet Creek, Nebraska, 179
Ward, Lester F., 69
Wealden, 71, 72
Weskit, Armstrong County, Pennsylvania, 387
White River Beds, 209, 211, 213
INDEX. 533
White, Dr. C. A., 59, 63, 69
White River, Indiana, 363
Oligocene, Colorado, 131
Wieland, G. R., 295
Osteology of Protostega, 279-298
Williamson, E. B., 376
Wills Creek, Ellerslie, Maryland, 372
Williston, S. W., 69, 85, 116, 123, 128, 280,
2925293, 336
Woods, J. G., 189
Woodward, Miss Alice B., 229, 230, 254, 259
Dr. A. S., 82, 124, 225, 254, 256, 259,
264
Dr. Henry, 23
Wortman, Dr. J. L., 81
Xenotherium, 211
unicum, 204, 224
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