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NOTA BREVE - SHORT NOTE

KOGIA PUSILLA FROM THE MIDDLE PLIOCENE OF TUSCANY (ITALY) AND A PHYLOGENETIC ANALYSIS OF THE FAMILY (ODONTOCETr, )

GIOVANNI BIANUCCI 't E \IALTER LANDINI '!'t

Receired. JanuarT 22, 1999; accepted JuQ 27, 1999

Key-oords: Cetacea, Kogiidae, Systematics, Phylogeny, Middle idae are very rare: only two almost complere skulls from Pliocene, Tuscany, Italy. late sediments of Mexico (Barnes, 1973,1984) and Peru (Muizon, 1988) have been described previous- Riasswnto. Viene preso in esrme un crrnio quesi completo di specimens cetaceo odontoceto provenienre dai sedimenti del Pliocene medio di ly. Those are the holotl'pe oI rhe extincr gen- Monte Voltra.io (provincia di Pisa, Toscana). Tale esernplare, in pessato era and species cedrosensis and Scaphokogia erroneamente attribuito alla famiglia Ziphiidae e descntto come cochlearis, respectively. Other presumed records of kogi- olotipo della specre Hyperoodon pusillus, viene qui riferito al genere ids are fragmentary and consist essentially of mandibu- pusiila è Kogla (farniglia Kogiidae). La specie ridescritta e messe lar fragments (Kel1ogg, 1929), isolated teeth (Matsumu- a confronto con le specie atruali K. brniceps e K. simus. to,1927) and/or auditory 1986a, 1986b, Da un punto di vista filogenetico, viene ipotízzart una sepa- bones (Pilleri, razione antica (ahneno nel Miocene inferiore) dei Kogiidae e Physe- 1987,1988; Pilleri et al., 1989). teridae (). Il mancato ritrovamento di reperti attribuibili The skull from the Pliocene of Tuscany described con sicurezza alla famiglia dei Kogiidae in sedimenti piìr antichi del here represents the third known significant record of Miocene superiore è probabilmente dovuto alla raritàL di quesri cetecei. this family. This specimen s/as referred erroneously to Lanalisi delle relazioni filetiche tra i Kogiidae è stata condotta pren- dendo in esame supposte apornorfie relative alla morfologia e/o esten- the Ziphiidae in the past. Capellini (1893) assigned it to sione del bacino sopracraniale che alloggia le sacche aeree e l'organo Placoziphiws and Pilleri (1987) described it as holotype dello spermaceti. of the species Hyperoodon pusillus. This fossil sku1l, already referred to the genus Kogia by Bianucci (1997) Abs*act. A partial skull of an odontocete cetacean from Middle and et (1998), is redescribed Pliocene sediments of Monte Voltraio (Pisa Province, Tuscany, Italy) Bianucci ai. in detail in this is exanrined. f'his fossil, erroneously referred to the fanily Zrphitdte paper. Kogia pusilla represents the ancesror of Kogia bre- and described in the past as holotype of the species Hyperoodon pusil- viceps and K. simus, the only two living species of this /as, is assigned here to the genus Kogia (family Kogiidae). The species family. Kogia pusilla is redescribed and compared to the living species K. bre- oiceps rnd K. simus. PhylogenericalÌy, an old separation (at least in the Lower Miocene) of Kogiidae and Physeteridae is suggested. The lack of sub- Systematic description. stantiated kogiid records until the Upper Miocene is probably due to the rarity of these cetaceans. Phyletic analysis within the Kogiidae is undertaken and supposed apomorphies in the morphology andlor the Linnaeus, 1258 extension of the supracranial basin that houses the large air sacs and Class Mammalia the spermaceti organ are considered. Order Cetacea Brisson, 1762 Suborder Odontoceti Flower, 1867 lntroduction. Superfamily P hys e t e r o i de a (Gray,1821,) G111,1.872 The Kogiidae are a family of small odontocetes (Cetacea) similar and closely reiated to the extant sperm Family K o g i i d a e (Gill, 1871) Mlller, 1923 whale (Physeteridae). !íell-preserved records of Kogi- Genus Kogia Gray, L846

'r- Museo di Storia Naturale e del Territorio, via Roma, 103, 5601 1 Calci, Pisa (Italy). E-mail: [email protected] 'r''t Dipartimento di Scienze della Terra, via S. Maria, 53,56126 Pisa (Italy). E-mail: [email protected] 446 G. Bianwcci f: W Land.ini

Kogia pusilla (Pilleri, 1982) In dorsal view (Figs. 1,a1, la2) the premaxillae, the (Fig.1-aa) maxillae and probably rhe vonìer (covered by marrix) extend as far as the apex of the rosrrum, as in living 1893 Placoziphizs, V Beneden - Capellini, pp. 287-288. species. A simiiar exrension of the rosrral bones was 1987 Hyperoodon pusillus Pilleri, pp. 35-36, fig. 11, pl. 13. observed also in Scaphokogia by Muizon (1988). Muizon 1997 Kogia pusilla (Pillerí) - Bianucci, pp. 172-1,73, tig. 6b. (1991) considered this feature an apomorphy of the 1998 Kogia p wsilla (P rllerl) - Bianucci et al., pp. 1,24, 126, 127, Íigs. Kogiidae. 3t1,22 . Two asymmetricai superior nares are present at Emended diagnosis of species. A species of the genus Kogla the base of the rostrum as in all Physeteroidea (Physe- that differs from K simus and K. brmicEs in having a more elongated teridae and Kogirdae). In particular, the left naris is 14 rostrum, a less anteriorly extended antorbital processr a smaller mm wide and 10 mm long whiie the right is 6 mm wide lacrimal, a slighrly more prounonced dorsal asymmerry and a lesser elevation of the posterior region of the neurocranium. It differs from and 10 mm long. The nasals are missing, as in the other in irs sn'raller size and in having a more narrow sagittal Kogiidae. !r"?:"r*Ot The ieft premaxilla exrends posteriorly as far as Holotype. IGF1540V distorted and incomplete skull, lacking the postero-dorsal margin of the skull and medial part of the right side and of the ventral portion of the braincase. Audi- its tory bones, teeth, rnandible and postcranial skeleton are not presewed. margin joins the right maxilla ro form a very promlnenr The specirnen s.as donated by Lawley in L877 to the Museo di Paleon- sagittal crest, as in living species. This crest is bent left tologia of the University of Florence, where it is preseroed today. in its anterior porrion. It is delirnited by vertical walls Type locality. La Rocca locality in Monte Voltraio hill, near and is very narros/ ar the verrex (width: 13.3 mm), as in Volterre, province of Pisa, Tuscany (Italy). Age. Middle Pliocene: Globorotalia crassaformis crassaformis K. simus. The width of the sagirtal crest is a diagnostic subzone of Iaccarino El Salvatorini (1982), approximately between 3 character for the distinction of the two living specles M-A and 2.6 MA (Bianucci et al., 1998). (Handley, 1966). Ross (1979) observed that in 26 skulls of K. simus the minimum wrdth of this crest ranges from Description. The condylobasal iength of the only 5 to 20.5 mm while in 22 skulls ol K. breaiceps this n-idth known but incomplete skull is estimated to be about 270 ranges from 14.5 to 49 n-rm (from 23 to 49 mm exclud- mnr. This skull size is similar ro rhar ol Kogia simus and ing five young specimens). it is slightly smaller than that of an adult of Kogia bred- The right premaxilla forms a small elevated basin ceps. In fact, according ro Ross (1979), the condylobasal on the neurocranium, extending from the sagirr.ll crest length of 24 skulls of Kogia simws range between 201 on the left and to its lateral margin on rhe right. A sìrr- mm and 323 mm whiie the condylobasal length of 22 ilar depression is present in the living species skulls of K. breviceps range berween 237 mm and 467 of Kogia mm (less than 300 mm are young ). where the spermaceri organ (Kernan & Shulte, 1918; Raven & Gregory, 1933; Schenkkan Ec 1973) Compared to the living species, the rostrum has a Purves, resides. In contrasr, Praekogia, simiìar triangular shape from a dorsal view; but differs in in rhe lareral rìlargin of the right premaxilla being rnore elongate (Fig. a). Moreover, it has a well- is not protuberant and the r.igbr maxilla and developed dorsal concrrvity and a venrral convexiry rhar the right prerl-raxilla forrl a single craniai fossa that u.'e harve not observed in extant species. Post morten-r probably resulted in a more developed sper- deformation may have accenruared these last characters naceri organ rhan n KogiLl. Scaphokogia shows an even in the fossil specimen. more dissimilar dorsal cranill architecture than Kogia.In The cerebral skull has a sernicircular outline in fact, the skull of Scaphokogta h:rs no sagitt:r1 cresr ar all dorsal view. In this respecr the skull is similar to thar of (the facial crest is displaced near rhe left m:.rrgin of the K. breoiceps and K simus but differs from the anrero- neurocraniunr) rnd con\eqLrenrly it is likely rlr.rr the posteriorly elongated skulls of both Praekogia and spermaceti organ also invaded the left side of cerebral Scaphokogia. Nevertheless, this skull differs from those skull (Muizon, 1988). of extant Kogia spp. in having a more pronounced dor- The antorbital notcl-res penerrate the supracranial sal asyrnmetry (due to the displacement of the external basin as in irving species o[ Kogia and in Praeleogia,but nares tosrard the left side and to the displacement of the nor in Scapbokogia and in the other Physeteroidea. This sagitt:ìl crest tor,'ard the right side), and in hrving small- character is considered by Muizon (1988) to be an apo- er elevation of the posterior region of the neurocranrum. morphy of the (Kogia and Praelectgia) re1:rted to The first of these two fearures may have been empha- the large antero-lateral development of the supracrxnial sized by Dost mortem deformation of the skull. basin.

-l'uscar-ry). Fig. 1 Kogta pusilla (Pilleri, 1982) holotype; La Rocca Iocaìity, Monte Voltraio (province of Pisa, Almosr complete skull (lGF1540V). a1,a2, dorsal view; b1,b2, vcntral view cf, cranial fossa; Fr, fronral; L;r, lacrirnal; Mx, maxilla; n, naris; Pn-x, prerlaxillae; sc, sagittal crest; Vo, vomer. Kogia pusilla from Middle Pliocene of Tuscany 448 G. Bianucci & V/. Landini

10 cm

Fig.2 - Kogta pusilla (Pilieri, 1982) holotype; La Rocca localitS Monte Voltraio (province of Pisa,'Iuscanv). Almost con- plete s kull (I GF 15 40V). a1,a2, laterel view; b1,b2, pos- terior view. Fr, frontal; La, . lacrimal; Mx, naxilla; Oc, occipital; sc, sagittai crest.

The antorbital process is narrower than in living orly exìended, to a greater extent than in the other Kogi- species and does not extend anteriorly on the proximal idae. Muizon (1988) noted that the posrero-dorsal portion of the rostrum. As a consequence, the cranial process of the lacrimal is not present in the Physeteridae f ossae (antero-laterally delimited by the antorbital except in a young specimen of Awlophyseúer, and the processes) are less developed antero-posteriorly in this presence of this process in the Kogiidae is considered as fossil species. a plesiomorphic character. The anterior portion of left cranial fossa is almost The supraorbital process is thinner than in both completely covered dorsally by the lateral and sagittal Iiving species and Praekogia. crests, both being bent towards the fossa. Posteriorly, The lateral view shows also an elevation of the the cranial fossae are slightly more elevated than in K posterior portion of the cranium, smaller than rn living brear.ceps. This feature makes K. pwsilla very different species and in Praebogta. from K. simws, in which the posterior part of the craniai In posterior view (Figs. 2bi.2b') the sagirral crest fossae rises strongly (Handley, 1966; Ross, 1979). is high, narro'w and delimited laterally by vertical wal1s. The iateral view (Figs. 2ay 2a2) shows a larger In ventral view (Figs. 1b1, 1b2) the skull is badly antero-posterior exposure of the rostrum than in the liv- damaged and most of the ventral surface of the neuro- ing species. This condition is due to both the greater craniun is not preserved. ln particular, only the left elongation of the rostrum and the smaller anterior frontal, the left lacrimal and a small porrion of the left extension of the rrntorbital processes (in K. brer:iceps and squamosal are preserved. The medial groove of the ros- K. sìwws the antorbital processes cover the proximal part trum allows the vomer to be observed. Along the right of the rostrum) . preserved margin of the rostrum is a sma1l deep furrow The anterior portion of the lacrimal is smaller than but without distinct alveo1i. In this respect K. pusilla is in the living species and is comma-shaped as in the Phy- similar to the living species and differs îrom Scaphokogia seteridae, whereas it is triangular in the other Kogiidae. (with distinct but shallow alveoli). There is no evident lacrimal-maxilla suture. Figs" 2a2 and Comparisons. A peculiar kogiid feature of the only 3 present the probabie course of this suture: the pos- skuli known of l{ogia pusilla is the lack of both nasals. tero-dorsal process of the lacrimal is large and posteri- The presence o[ a supracranial basin and the strong Kogia pusilla from Middle Pliocene of Tuscany 449

,1 cm,

Fig. 3 - Kogia .pusilla (Pilleri, 1982) holotype; La Rocca localitv, Monte Voltraio (province of Prsa, Tuscany). Neurocrani- um ìn lateral view of skul.l (IGF1540V). Fq fronral; l.a, lacrimal; Mx, rnaxilla; Oc, occrpital; porp, posrorbital p roce \s i \q. : qur mo 'eì; r f. temporal fossa.

asymmetry of the skul1 are other characrers typical of all Moreover, as rhe extant species, K. pwsilla shows a srnall the Physeterida (Kogiidae and Physeteridae). Kogia basin on the cerebral porrion of the right premaxilla that pusilla is similar to K. breoiceps and K simus in the pecu- extends as far as the postero-dorsal margin of the skull. liar outline of the skull in dorsal view (the cranium is K. pusilla is more similar to K. simus than K. breuiccps in semicircular and the rostrum is triangular), and in hav- the relatively small size of the skull and in the narrow ing a vertical sagittal cresr on the cranium (Fig. a). sagittal crest at the vertex.

Fig. 4 - Comparisons of the skull in dorsal view of: a, Kogia pusilla (restoration of the skull based on holotype, IGF 1540V); b, Kogia brniceps (Laboratoire de Anatomie Comparée de Paris, 1833-483); c, Kogia simus (Accademia dei Fisiocritici di Siena). 450 G. Bianucci G \Y Landint

K. pwsilla differs from both K. brersiceps and K. Handley, 1966;Heyning, 1989; Simpson, 1945) consider simws in its longer rostrum, smaller lacrimal and perhaps that Kogia and Physeter belong to the same family Phy- grear.er asymmetry of its skull in dorsal view. seteridae. Others (Barnes, 1,973; Kaxrya, 1.973; Miller, 1.923; Muizon, 1991) think these rwo genera have such distinctive characters that their separation in two fami- Phylogenetic relationships. lies (Kogiidae and Physeteridae) is justified. This iast hypothesis was strengrhened by recenr studies on The genus Kogia shows evident affinities with cetacean mitochondrial DNA (Arnason et al., 1.993; Physeter. Common apomorphies of these ts/o genera Milinkovitch et aL.,1.994) that confirm a large divergence concern particularly the facial anatomy (Heyning, 1989) between Physeter and Kogia. and the concave shape of the dorsal surface the skull of We believe that the fossil and phylogeneric evi- (scaphidiomorphy) for housing the large air sacs and the dence reveal that the separation of these two groups may spermaceti organ. Consequently, the singie bones of the be relatively old. In particular, all fossil skulls of Physe- skull suff ered large modifications, in parricular, a teridae show a marked enlargement of the posterior por- marked asymmetry and the reduction of the right exter- tion of the right premaxilla, an apomorphy absent in nal nares. Kogia and in fossil taxa relared to this genus. As Even if there is consensus on the close affinities of observed by Muizon (1991) this feature is relatively less Kogia and Physeter, precise systematic relationships emphasized in living Pbyseter than in fossil physeterids. between these two genera are debated. Some authors This derived character is also present in the most ancient (Caldwell & Caldwell, 1989; Fraser & Purves, 1960; known skull of physeterid, assigned to Diaphorocetus

PHYSETERIDAE KOGIIDAE

QUATERNARY 1.8 MA

PLIOCENE

Kogia pusilla

MIOCENE

OLIGOCENE

Fig. 5 - Phylogeny of the Kogiidae. Dotted lines indicate no fossil or fragmentary record. Kogia pusilla from Mid,dle Pliocene of Tuscany 451

K. breviceps gia elongata (emended of Miokogia elongatum), a genus K. simus and species created on rhe basis of three teeth from Bal- K. pusilla ringen (Piileri, 1986b). In fact, all teeth referred to this Praekogia presumed kogiid are similar in shape and size to those of the physeterid Scaldicetws Scaphokogia bellunensls frorn the Early Miocene Diaphorocetus of Belluno (North Italy) (Dal Piaz, 1977).In Squalodon accordance with orher authors (Barnes, 1976; Muizon, 1988) we attribure a 1ow systematic value to isolated teeth and therefore consider Fig. 6 - Maxinun-parsimony cladogram (cosistency index : 100) nr.tmina dubia borh senus conputed from the data in Tab. 1 using Henning 86, ver- and species names of Miokogia elongata. sion 1.5 (Farris, 1988). Considering other described kogiid fossils, only two skulls were referred correctly to this family. One, from the Late Mìocene of Mexico (Barnes, 1973,1984), powcheti (Early Miocene from Argentine; Moreno, 1892; was named Praekogia ced.rosensis, and the other, from the Lydekker,1894) Late Miocene of Peru (Muizon, 1988) was named The primitive state of Kogiidae, regarding the Scaphokogia cochlearis. Praekogia and Scaphokogia share shape of right premaxillae, suggesrs rheir origin no is wrth Kogia the loss of nasal bones, an apomorphy of the younger than the beginning of rhe Miocene (Fig. 5). The family Kogiidae. Nevertheless, the rwo fossil genera dif- lack of of kogiid records prior ro the Late Miocene may fer from Kogta by having a different architecture of the be due frequency to their low in the ancient sea. supracranial basin. In parricular, in Praekogia the basin Flowewer, some isolated auditory bones and reerh from [or the spermaceri orgrn exrend\ ro rhe entire right por- the Lower Miocene of Piedmont (North Italy) and Ba1- tion of the dorsal surface of the cerebral skull that is tringhen (Austria) and Middle Miocene of Visiano formed by the premaxilla and maxilla. As observed by (North Italy) were referred ro this family by Piileri Barnes (1.973), this genus lacks a distinct right maxillary (1986b, 1988) and Pilleri et al. (1989). Nevertheless, fossa that is present in Kogia. The basin for the sperma- because these elements (periotics and teeth) are rhe only ceti organ in Scaphokogia takes up rhe entire dorsal sur- remains of kogiids found in the sediments, their conclu- face of neurocranium and only a large cranial fossa is sion must be viewed with caution. As in Kogra, some present. Because of this peculiar shape of the supracra- periotics show an articular surface for the tympanic flat nial basin and other peculiar characrers, Scapholeogia was and parallel to the general plane of the periotic and not assigned to the subfamrly Scaphokogiinae by Muizon ventrally oriented; nevertheless, on the whole rhey dif- (1988). Apparently Scaphokogia shows affinities with fer substantially from Kogia periotics and show also Pbyseter in having a large cranial basin for the spermaceri affinities with those of some fossil physeterids. The dis- organ. Nevertheless, rhe cranial basìn of Scapbokogta tinction of the two families on rhe basis of rhese struc- does not extend on the rostrum as in Pbyseter. A cranial tures is problematic, considering also that skulls of fos- basin more anteriorlyextended than rn Scapholeogia is sil physeterid and kogiid genera with preserved auditory present also in Kogia and Praekogia, both having the bones are rare. The isolated teerh are referred to Mioho- dorsal depression that exceeds the external nares. More-

Taxon Character

I 2 7 + 5 6 7 8 9 10 11 12 l3 I4

Siua,lodon 0 0 0 00 0 00 0 000 ;--T--" ; t; Diaphorocetus 1 1 UI 0 00 0 ò o-- UU^-^ Scaphobogia 2 ,T- 0 , : 0 nnln olo >:0 _f Praekogia 2 1 o OO:i ::0 T ll K"4o p"tr\ 1 2 00 1 01 i. 110 r 00 simus 1 - Kogia 2 00 1 01 1 111 0 f- T Kogia breoiceps 1 2 00 1 01 1 11. 1 ll

Tab. 1 Data matrix used in the cladistic analysis. 't= missing characters. Characrer states: 1) Supracranial basin: O, no; 1, yes; 2) Strong rsym- metry: 0, no; 1, yes; 3) Lack of nasals: 0, two nasalsl 1, one nasal; 2, nasals absent; 4) VomeS premaxillae and maxillae reach the apex of rostrum; 5) Enlargement of posterior portion of right premaxillae: O, no; 1, yes; 6) Antorbital notch penetrating rhe cranial fossae: O, no; 1, yes, 7) Supracranial basin formed by only one hemispherical depression posteriorly developed and anteriorly closed (see Muizon, 1988, 1.991.,for more detail on the bone modifications): 0, no; 1, yes; 8) Craniurn half-circle shaped: O, no, 1, yesl 9) Rostrum triangular: 0, no; 1, yes; 10) Small basin on the cerebral portion of right premaxilla: O, no; 1, yes; 11) Very prominent sagittal crest: O, no; 1, yes; 12) Very large antorbital process and lacrimal; 0, no; 1, yes; 13) Rostrum verv shorr: 0, no; 1, yes; 14) Relarively broad sagittal crest: 0, no; 1, yes. 452 G.Bianucci&WLandini over, in these last two genera, the cranial basin takes up Kogiidae older (at least Early Miocene) than that sug- the antorbital processes and consequently the antorbital gested by the paleontological data (Late Miocene) is notches penetrate the cranial fossae. This last character suggesreq. was considered as an apomorphy of the Kogiinae (Kogia In accordance with Muizon (1988, 1991), the and Praekogia) by Muizon (1988). Kogiid,ae are separated into two subfamilies on the basis Considering interrelationships among the three of different architecture of the supracranial basin: the species of the genus Kogia, apparently K. brez,iceps and Kogiinae (Kogia and Praekogia) and the Scaphokogiinae K. simws are more derived than K. pusilla in having more (Scaphokogia). The morphology of rhe supracranial anterioriy extended antorbitai processes (and conse- basin of Kogia pusilla is similar to that o[ K. breaiceps quently more elongated cranial fossae) and a shorter and K. simws eyen if the basin of the fossil species differs rostrum. A parsimonious interpretation would suggest in having a smaller anterior exrension. that the wide sagittal crest of K. brer.ticeps is an apomor- phy, considering the primitive status of the narrow crest Acknouledgements. observed in K. pusilla and K. simus. are very grateful to the following us The characters discussed belovr are used to create 'We persons for grrnting access to cetacean collections and for their assistance during our vrsirs a maximum-parsimony cladogram of the relationships at the museums: P. Mazza and F. Cozzini (Museo di Geologia e Pale- of Kogia pwsilla using Henning 86, version 1.5 (Farris, ontologia, Università di Firenze); D. Robineau (Laboratoire de 1988) (Tab. 1, Fig. 6). Anatomie Comparée de Paris); F. Cancelli (Accademia dei Fisiocririci Regarding other fossils referred to the family di Siena). \7e also wish to thank C. de Muizon (Museurn Natinal d'Histoire Naturelle de Paris) for critically reading the rnanuscnpr. Kogiidae in the past, we agree with Barnes (1973) and The photographic plates were prepared by the euthors and M. Gini Muizon (1988) on the attribution of Kogiopsis floridanus (Dipartimento di Scienze della Terra, f Inir.ersità di Pisa) from the Miocene of Florida (Kellogg, 1929) as a prob- able physeterid and in considering the name of the species Kogia prisca (Miocene?, from Japan; Matsumoto, RE,FERENCES ll t1l/ ) a nomen au0tum. Pilleri (1986a, 1982) identifies the living species A.r,rror Ú., Grllb..g A. 6c \X/idegren B. (1993) - Ceraceen Kogia simus and K, breoiceps in the Pliocene of Tuscany mitochondrial DNA control region: sequences of all on the basis of isolated teeth and periotics. \fe consider extapt baleen whales and two sperm r.vhales..species. these fossils unpublished specimens as and other Mol. Biol. Eool.,v. 10 (5), pp. 960-97A, Chicago. belonging to the Kogiidae and most probably to the Barnes L.G. (1973) - Praekogia cedrosensis, a new genus and gentts Kogia. Nevertheless, we do not agree with Pilleri species of fossil pygmy sperrn whales from Isla Cedros, in referring these f ossils to the two living species Baja California, Mexico. Contr. Sci., nat. Hist. Mus. Los because the periotics show different characters and the Angeles C., v. 247, pp. 1-20, Los Angeles. teeth are not diagnostics at this systematic level. It is Barnes L.G. (1976) - Outline of eastern Noth Pacific fossil possible that most of these specimens belong to the fos- cetacean assemblages. Syst. Zool., v. 25 (a), pp.321-343, sil species Kogia pusilla. New Haven. Fossil kogiids were reported recently from Barnes L.G. (1984) - Fossil odontocetes (Mammalia: Cetacea) Pliocene sediments of the Yorktown Formation (Vhit- from the , Isla Cedros, Mexico. Pale- more, 1994) and northern Appennines (Cigala Fulgosi, obios,v. 42, pp. 1-46, Berkeley. 1996), but they were not described in detail. Bianucci G. (1997) - The Odontoceti (Mammalia, Cetacea) from Italian Pliocene. The Ziphiidac. 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