Paleontology  of  the  Late  Oligocene  Ashley  and  Chandler
Bridge  Formations  of  South  Carolina,  1:  Paleogene
Pinniped  Remains;  The  Oldest  Known  Seal
(Carnivora:  Phocidae)
Irina  A.  Koretsky  and  Albert  E.  Sanders
For comparisons we used postcrania lmateria lfrom several
ABSTRACT living and fossi lrepresentatives o faquatic and semiaquatic car¬
The proxima lhalves o ftwo femora from the Chandler Bridge nivores ,including Mustelidae (the genera Lutra ,Enhydra ,and
and Ashley Formations (early Chattian, late Oligocene) near Potamotherium) ;Otariidae (the genera Zalophus and Allodes-
Charleston ,South Carolina ,provide the earlies tevidence to date mus, the latter the most controversial eared seal (Berta and
o ftme seals .They are clearly referable to the Phocidae and fiimish Wyss, 1994; Barnes and Hirota, 1994)); and Phocidae (for ex¬
informatio nregardin gosteologica al n dmyologica fleature sthat ample ,the genus Cystophora as the mos tspecialized represen¬
had evolved in early phocids by early Chattian time .Although not
determinat et oth egeneri cleve lt,hes especimen srepresen  attaxon tative o fthe family).
closely comparable to the mos tspecialized phocid ,the modem Acknowledgments. —The authors are grateful to James
ge Cnyusstophora. Mead and Charles Potter for access to modem comparative ma¬
terial, to Clayton E. Ray for access to fossil pinniped collec¬
tions ,to Victor Krantz and Steven Jabo for specimen photogra¬
Introduction phy  (all  National  Museum  of  Natural  History  (NMNH),
Smithsonian Institution, Washington, D.C.), and to Robert J.
The long-standing question of phylogenetic relationships Emry (NMNH), Daryl Domning (Howard University, Wash¬
among the Odobenidae ,Otariidae ,and Phocidae is stil lopen. ington ,D.C.) ,and Mieczyslaw Wolsan (Institute of Paleobiol¬
The debates abou tthe monophyletic or diphyletic origin o fpin¬ ogy, Warsaw, Poland) for their helpful comments on several
nipeds from a series of ancestors (mustelid- or ursid-like ani¬ version so fthi smanuscript.
mals) are not yet resolved .The new materia lreported herein, Material and Locality. —Proximal portion of right femur
the proximal parts of two femora from the late Oligocene of (ChM PV5712), from a ditch in the Irongate subdivision, ap¬
South Carolina ,now in The Charleston Museum (ChM) ,helps proximately 3.9 km (2.4 mi.) SW of Summerville, Dorchester
little in solving the problems of phylogeny ,but we have for in¬ County, South Carolina, collected by Vance McCollum, sum¬
vestigation the earliest and most primitive remnants of Pho¬ mer  1978;  cast  of  ChM  PV5712  (USNM  299831)  in  the
cidae. Herein we refer to the two partial femora from the late NMNH (which subsumed the collections of the former United
Oligocene as the “Oligocene seal.” The two new specimens ap¬ States National Museum). Proximal portion of right femur
pear to represent the same taxon ,but they are so incomplete (ChM PV5713), from a ditch on Trolley Road (County Road
that any taxonomic assignment below the family level would 199), approximately 3.0 mi. (4.8 km) SW of Summerville,
be mere speculation .Thi smateria lnonetheles sdoe sprovide in¬ Dorchester County ,South Carolina ,collected by Chris Kenney,
formation concerning some derived (=apomorphic )and primi¬ 6 December 1978; cast of ChM PV5713 (USNM 306508).
tive (=plesiomorphic )character so fthe Phocidae. Formation  and  Age.—C  hM  PV5712:  Chandler  BridgeFormation, early Chattian (late Oligocene). ChM PV5713:
Ashley Formation ,early Chattian (late Oligocene).
Irin aA K. oretsky R, esearc hAssociate D, epartmen ot Pf aleobiology, The Chandler Bridge Formation ,an unconsolidated noncal-
Nation aMl useum o Nf atur aHl istor yS,mithsonia nInstitution W, ash¬ careous marine unit rich in vertebrate fossi lremains ,occurs at
ington D, .C 2. 0560-0121 A. lber tE S. anders D, epartmen to fNatural
Sciences T,h eCharlesto nMuseum 3,6 0Meetin gStree tC, harleston, numerous disjunct localities in Berkeley, Charleston, and
Sou tCharolin 2a9430. Dorchester Counties ,South Carolina .It unconformably over-
179
180 SMITHSONIA CNONTRIBUTION T PSOALEOBIOLOGY
Figur eI.—Femor ao Of ligocen eseal s( x1 )R. igh ftemur o, rigina Cl hM P V5712 c,as Ut SNM 299831 f,rom
Chandle Brridg Feormatio ne,ar lCyhattia nla, t Oe ligocen e ac:,aud avliew  bc;,rani avliew R.igh ftemu ro,riginal
ChM P V5713 c,as Ut SNM 306508 f,rom Ashle yFormation l,owe Cr hattian l,at eOligocene c :c,auda vliew d; .
cra vniieawl .
lies the Ashley Formation, a highly indurated calcarenite un¬ Phocidae
derlying mos to fthe area encompassed by the aforementioned
counties (Sanders, 1980; Sanders et al., 1982). E. Martini has Genus and Species Indeterminate
found the nannoplankton of the Ashley Formation to be refer¬ Description.— The femora (Figure 1, Table 1) are similar in
able to zone NP24 (Sanders and Barnes, 2002). Sanders et al. size to those of the modem harp seal ,Pagophilus groenlandi-
(1982 )have estimated the Chandler Bridge Formation to be ap¬ cus. The shaft of the bone is flattened in the cranial-caudal di¬
proximately 28 million years old and the Ashley Formation to rection. The greater trochanter extends proximally slightly
be about 30 million years in age .Both units have yielded abun¬ higher than the head and is slender, although not well devel¬
dant remains of sharks; bony fishes; sea turtles {Carolin- oped .The trochanteric fossa is shallow and wide open ,actually
achelys ,Procolpochelys ,Syllomus ,two dermochelyids) ;three no treaching the middle o fthe greater trochanter .The intertro¬
nonmarine turtle taxa ;a large ,newly described estuarine croco¬ chanteric crest is not expanded. The lesser trochanter is very
dile ,Gavialosuchus carolinensis Erickson and Sawyer (1996); wel ldeveloped and is located on the posteromedia lside of the
marine birds (including a large pseudodontom with a wingspan bone at the same level as the distal border of the greater tro¬
of about six meters) ;sirenians ;and more than 20 taxa of primi¬ chanter. The cervix is short but narrow relative to the bone’s
tive cetaceans .Now a pinniped is added to the impressive fau¬ mass .The head is damaged ,but it clearly has an indentation for
nas o fmarine vertebrates from these two formations. attachment of the ligamentum teres in the center of the head.
numb 9e3r 181
Tab l1e.—Measuremen t(si nmm o )tfh leat Oe ligocen see aflemor farom the mur laterally .The actions of the gemell iand obturator intemus
Chandle Brridg Feormatio (nCh MPV571 2a)n tdh Aeshle Fyormatio (nChM are to abduct the femur and also to rotate it laterally.
PV571 D3)o,rches tCeorun Styo,u Ctharolina, Comparisons. —The Oligocene sea ldiffers from other car¬
Character nivores we examined (except for Cystophora and Zalophus) in
its shallow and poorly defined trochanteric fossa, from all ex¬
cep tCystophora by its sligh textension o fthe greater trochanter
above the head ,and from al lexcept Lutra and Potamotherium
in having a pit for attachment of the ligamentum teres.
Other distinc tdifferences between this sea land other genera
also exist. It differs from Cystophora (the hooded seal) in hav¬
ing a lesser trochanter (but this is always absent in females of
Cystophora)- a ,narrowe rand les sprominen tgreate rtrochanter;
and a more delicate ,not massive ,overal lbone stmcture .So far
as they can be determined ,the muscle attachments for the Oli¬
This insertion of the ligamentum teres is not a completely de¬ gocene sea lare in genera lthe same as for Cystophora ,but with
veloped fovea as it is in most terrestrial mammals. In phocids one exception; in female hooded seals ,which lack a lesser tro¬
thi sfove ai sabsent. chanter, the tendons of the pectineus and adductor cranialis
The slight difference in bone morphology of the two Oli¬ muscles are inserted on the same spot on the bone ,and the ili¬
gocene femora perhap scould be ascribed to sexua ldimorphism opsoas muscle inserts upon the medial tuberosity of the tibia
(Koretsky ,1987) ,but the limitations of the materia lpreclude a (Howe l1l,928).
mo rpereci sineterpretation. From Lutra this seal differs in the presence of a well-devel¬
According to authors who have studied the functiona lmor¬ oped and distinc tlesser trochanter and a much broader shaft .In
phology and myology of the limbs of different carnivores general ,there are no differences between these two taxa in in¬
(Howell, 1928, 1930; Savage, 1957; Mori, 1958; Mitchell, sertion of the described muscles ;gemelli ,obturator extemus,
1966; Pierard, 1971; Tarasoff, 1972; Sokolov et al., 1974), the and intemus muscles al linsert into the trochanteric fossa ,and
lesser trochanter is the point of insertion of the iliopsoas mus¬ the iliopsoas muscle inserts onto the lesser trochanter (Sokolov
cle. These authors also noted, however, that if the lesser tro¬ e tal. ,1974).
chanter is absent ,then the pectineus and adductor muscles in¬ The Oligocene seal differs from Enhydra in the shorter cer¬
sert on this place; that is true for al lcomparatively aquatic and vix and smaller size of the head compared with the rest of the
semiaquatic mammals .The iliopsoas muscle is a large ,heavy bone .Insertions o fthe muscles into the fossa trochanterica are
muscle connecting the lumbar spine and ilium with the femur. almost the same in Enhydra and Lutra, but the mass of these
The actions of this muscle are to flex and laterally rotate the fe¬ muscles is apparently greater in Enhydra .In contrast to the Oli¬
mur, tilt the pelvis forward, flex the thigh upon the pelvis, and gocene seal, Enhydra lacks the ligamentum teres (Sokolov et
adduct the femur (working as a synergist with the adductor al. ,1974 ;Howard ,1975).
brevis ()Gordon 1,983). Its differences from Potamotherium are the more media llo¬
The quadratus femoris muscle is absent in extant phocids .In cation of the lesser trochanter and the absence of the intertro¬
the otter ,this reduced and weakened muscle inserts on the dis¬ chanteric crest .In Potamotherium the tendon of the obturator
ta lpart o fthe greater trochanter ,on the cauda lborder o fthe fe¬ intemus did not merge with the common tendon of the obtura¬tor extemus and gemelli ;the quadratus femoris was located on
mur (Sokolov et al., 1974). In sea lions (otariids), the insertion th eintertrochanteri ccres t(Savage 1, 957).
o fthe m .quadratus femoris is along the entire laterocauda lbor¬ The Oligocene seal differs from Zalophus in having a more
der of the femur (Howell, 1928). The action of the quadratus medially located lesser trochanter and in lacking an intertro¬
femoris muscle is to extend the hip joint. chanteric crest .Presumably the insertion o fthe obturato rexter-
Howell (1928, 1930), Pierard (1971), and other researchers nus in the Oligocene seal is located distal to the trochanteric
described the trochanteric fossa as a place o fattachment o fthe fossa on the posteromedia lside of the greater trochanter; the
obturato rintemus and extemus muscles ,togethe rwith the two tendons o fthe two gemell imuscles were not joined together .In
gemell imuscles ,the superior and inferior .The gemell iand ob¬ Zalophus ,the tendon o fthe gemellus superior muscle does not
turator extemus muscles arise from the latera lborder o fthe ob¬ join with the tendon of the obturator intemus muscle; the ten¬
turator foramen and its membrane .The two gemell ijoin with don o fthe adductor brevis muscle inserts into the intertrochan¬
the obturator intemus muscle to form a common tendon for in¬ teric crest (Howell, 1928; Lyon, 1937); and the ligamentum
sertion .(Pierard at the same time (1971:68) mistakenly stated tere  isasbsent.
that the origin of the tendon of the obturator intemus muscle is From Allodesmus, the Oligocene seal differs in its smaller
a shallow groove in the middle one-third of the pubic edge.) and thinner greater trochanter and its lack o fthe intertrochant¬
The action of the obturator extemus muscle is to rotate the fe¬ eric crest. By analogy to the living otariids ,the adductor brevis
182 SMITHSONIA CNONTRIBUTION T PSOALEOBIOLOGY
Tab l2e.—Comparativ deiagnost iccharacte ro stfh feemor o atfh Oeligocen see aln sdom oethe mr embe ro sCfar¬
nivo r(+a=. charac tperrese n-^tc; harac taebrse n+/t;-=charac tvearriable.)
Character
muscle in Allodesmus inserted on the intertrochanteric crest insertion of the muscles that rotate the femur (Howell, 1928;
(Mitchell, 1966; Hirota, 1983), which is not present in the Oli¬ Tarasoff, 1972; Gordon, 1983). The Oligocene seal, with its
gocene seal .The fovea capitis ,and hence presumably the liga- greater trochanter only slightly expanded ,used its hind limb in
mentum teres ,is absen talso in Allodesmus. latera lmovemen tand with some flexion and adduction (similar
to Cystophora )for aquatic movement .This involved little rota¬
Discussion tion o fthe femur ,suggesting tha tthese Oligocene animals were
no tas good swimmers as later phocids.
The osteology and interpreted myology of these proximal As the antagonistic muscle to those latera lrotators ,the iliop¬
femora indicate that these remains are definitely more closely soas muscle ,located on the cauda lside of the femur ,becomes
related to the Phocidae than to other carnivores (Table 2). In very much reduced and weakened .In paralle lto the decrease in
addition, the functional morphology of even these proximal its stress on the lesser trochanter ,the latter is reduced and fi¬
femora can suggest whether this Oligocene sea lused its hind nally disappears .In those mammals tha tdo no tbring their hind
limb predominantly for propulsion in water or to some degree
fo rterrestria lmovemen ta swell. limbs forward ,the lesser trochanter is less and less developed.We can observe this progressive reduction from Lutra ,to the
For example, the diaphysis of the femur of Lutra is almost Oligocene seal, to Cystophora (where it is small in males and
circular in cross-section and is better prepared for terrestria llo¬
comotion than for aquatic movement .Increasing the surface absent in females) ,and finally to Phoca ,where it is completely
area for insertion of the adductor muscle makes the shaft of the absent .The latera lrotation of the lower limb results in turning
femur broader .The enlarged adductor muscle also helps pre¬ the hind foot so that it can move in a horizonta lplane.
vent rotation and movement of the femur .The diaphysis in the The anatomy of Enhydra and Zalophus shows that the feet of
Oligocene seal is flatter and wider than in Lutra, Potamothe- these animal sare adapted predominantl yfo rlatera ml ovement,
rium ,and Enhydra ;similar to that o fAllodesmus ;and narrower but that vertica land horizonta lmovement also is possible for
than in Cystophora and Zalophus. body support and locomotor activity on land. On the other
The literature and observation o fliving animals both indicate hand, the anatomy of the pelvic limb of Lutra is similar to that
that terrestrial locomotion in Lutra (and presumably in Pota- o fcompletely terrestria lcarnivores .Phocids have undergone
motherium) is only slightly impaired. The increased surface major morphologica lchanges tha tare closely correlated with
area o fthe femur suggests that Enhydra ,Zalophus ,and the Oli¬ thei rspecialized methods o faquatic locomotion .In its patterns
gocene sea lhave more limitations on land .Cystophora has lost of aquatic locomotion ,Enhydra shows analogy to both Lutra
the ability to turn its hind limbs forward on solid ground, as and Cystophora and to the Oligocene seal .Moreover ,Enhydra
have other modem Phocidae, in contrast to the Otariidae and has a convergent similarity to the seal in its elongated hind
Odobenidae .When the anima lis on land ,the ligamentum teres limb as the major organ o fpropulsion.
allows rotational movement of the femur and helps to fix the The primary result of this study is that ,from the viewpoint of
head of the femur to the acetabular fossa .This ligament disap¬ adaptation to the water ,the numerous modifications o fthe Oli¬
pears when the animal spends less time on land (Tarasoff, gocene sea lcorrespond most closely with Cystophora .From a
1972) and has greater freedom of movement of the femur. In taxonomic viewpoint ,the relationship between the Oligocene
the Oligocene sea lthere is morphologica levidence for a higher sea land other phocids is supported both osteologically and ,by
degree of terrestriality than in Cystophora and Zalophus but inferenc me,yologically.
much less than in Lutra and Potamotherium. Within the range o ftaxa compared herein ,the Oligocene seal
The distribution among carnivore taxa of osteological fea¬ is closer in several characters (see Table 2) to the semiaquatic
tures associated with lower-limb propulsion can assis tin inter¬ Mustelidae than to the representative o fthe eared seals .These
preting the specific method of locomotion in both aquatic and characters are primitive (e.g. ,the lesse rtrochanter) ,indicating
terrestria lmovement .The main effect of the flattening and ex¬ its derivation from land carnivores. At the same time, the Oli¬
pansion of the greater trochanter is thus to increase the area of gocene sea lhas derived characters (for example ,the much en-
numb 9e3r 183
larged greater trochanter) that place it within the Phocidae sea lwas wel ladapted to a marine environment.
rather than as an ancestor of this taxon. The morphology of Savage (1957:235) concluded that the Phocidae “evolved
these partia lfemora ,and the functiona lmyology interpreted from a lutra-like mustelid in the Oligocene. The Otariidae and
from the osteology ,shows tha tthe locomoto radaptations o fthe Odobenidae may have had a separate origin.” The material of
Oligocene seal were much closer to the pinniped type (espe¬ this Oligocene seal is not sufficient to either support or falsify
cially the phocids) than to that of the semiaquatic mustelids. the hypothesis of mustelid ancestry, but it is sufficient to con¬
This ,along with its occurrence in marine deposits ,associated clude that the ancestor of Phocidae must be sought in deposits
with othe rfully marine vertebrates ,suggests tha tthe Oligocene olde rthan the late Oligocene.
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