Eocene relatives of cod icefishes (Perciformes: Notothenioidei) from Seymour Island, Antarctica

Geo log i cal Quar terly, 2013, 57 (4): 567–582DOI: http://dx.doi.org/10.7306/gq.1112

Eocene rel a tives of cod icefishes (Perciformes: Notothenioidei)from Sey mour Is land, Antarctica

Ma³gorzata BIEÑKOWSKA-WASILUK1, *, Niels BONDE2, 3, Pe ter Rask M�LLER4

and Andrzej GAZDZICKI5

1 Fac ulty of Ge ol ogy, Uni ver sity of War saw, ¯wirki i Wigury 93, 02-089 Warszawa, Po land

2 In sti tute of Ge og ra phy and Ge ol ogy, Uni ver sity of Co pen ha gen, Voldgade 10, DK-1350 Co pen ha gen K, Den mark

3 Fur Mu seum, Nederby, DK-7884 Fur, Den mark

4 Nat u ral His tory Mu seum of Den mark, Uni ver sity of Co pen ha gen, Universitetsparken 15, DK-2100 Co pen ha gen �, Den mark

5 In sti tute of Paleobiology, Pol ish Acad emy of Sci ences, Twarda 51/55, 00-818 Warszawa, Po land

Bieñkowska-Wasiluk M., Bonde N., MÝller P.R. and GaŸdzicki A. (2013) Eocene rel a tives of cod icefishes (Perciformes:Notothenioidei) from Sey mour Is land, Antarctica. Geo log i cal Quar terly, 57 (4): 567–582, doi: 10.7306/gq.1112

Frag men tary skull bones and ver te bra from the Up per Eocene La Meseta For ma tion on Sey mour (Marambio) Is land, Ant arc -tic Pen in sula have been de scribed as gadiform fishes, in for mally named “Mesetaichthys”. Here we de scribe jaws asMesetaichthys jerzmanskae n. gen. and n. sp., and re fer this taxon to the perciform suborder Notothenioidei. This group is al -most un known as fos sils. Sim i lar i ties to the liv ing, “prim i tive” nototheniid Dissostichus eleginoides are in di cated in thedentition. Gadiform evo lu tion in the Paleocene–Eocene is dis cussed, and the pos si bil ity of a cor re la tion be tween the or i ginand evo lu tion of notothenioids in con nec tion with the de te ri o ra tion of the cli mate in Antarctica dur ing the LateEocene–Oligocene is con cluded.

Key words: Notothenioids, cod icefishes, Eocene, Sey mour Is land, Antarctica.

INTRODUCTION

Some frag men tary fish fos sils from the up per La MesetaFor ma tion (Up per Eocene) of Sey mour Is land (IslaVicecomodoro Marambio), West Antarctica (Fig. 1), have beenre ferred to Gadiformes by Jerzmañska and Œwidnicki (1992).We in tend to show that these fishes are the al most only knownfos sil rep re sen ta tives of cod icefishes’ rel a tives, the perciformsuborder Notothenioidei. Large jaw frag ments are de scribed as Mesetaichthys jerzmanskae n. gen. and n. sp.

The palaeontological data pro vides new in sights into theAnt arc tic Eocene fish com mu ni ties, and the or i gin of codicefishes, and the op por tu nity to re late their evo lu tion to the cli -ma tic changes dur ing the Late Eocene to Pleis to cene, re call ingthat this group, to day is found dom i nat ing in the South ernOcean, and is clearly adapted to cold wa ters.

The stud ied spec i mens were col lected by A. GaŸdzicki andA. Tatur dur ing the Ar gen tine-Pol ish Field Party on Sey mour Is -

land in the aus tral sum mer of 1991–1992. The col lec tion com -prises 45 spec i mens and is housed at the In sti tute of Paleo -biology, Pol ish Acad emy of Sci ences, Warszawa un der cat a -logue num ber ZPAL P.V./1–45; all from the lo cal ity ZPAL 3,Telm7, up per most Eocene (Figs. 2 and 3).

We pro vide de scrip tions of the jaw frag ments (part den ta -ries and dermarticulars – “angulars” and premaxillaries), abasioccipital and iso lated ver te brae pre vi ously in for mally na -med by Jerzmañska and Œwidnicki (1992) as “Mesetaichthys”.There are four premaxillae (ZPAL P.V./1–4), 12 den ta ry frag -ments (ZPAL P.V./5–16), seven dermarticulars (ZPALP.V./17–23), a basioccipital (ZPAL P.V./24), and 21 ver te brae(ZPAL P.V./25–45).

GEOLOGICAL AND STRATIGRAPHICAL SETTING

The Eocene La Meseta For ma tion, which crops out in thenorth east ern por tion of Sey mour Is land (Fig. 1), is a 720 m thickse quence of richly fossiliferous, shal low-ma rine, deltaic and/orestuarine poorly con sol i dated clastic strata (Figs. 2 and 3), ac -cu mu lated within an in cised val ley (Sadler, 1988; Porêbski,1995; Marenssi et al., 1998; Marenssi, 2006; Tatur et al., 2011). The for ma tion pre serves an ex cep tional re cord of a shal -

* Corresponding author, e-mail: [email protected]

Received: March 8, 2013; accepted: June 3, 2013; first publishedonline: 8 July, 2013

https://gq.pgi.gov.pl/article/view/7727

low-ma rine eco sys tem (Feldmann and Woodburne, 1988; Stil -well and Zinsmeister, 1992; GaŸdzicki, 1996, 1998, 2001; Dzikand GaŸdzicki, 2001; Fran cis et al., 2006; Uchman and GaŸdzi -cki, 2006; Ivany et al., 2008; GaŸdzicki and Majewski, 2012).

Sadler (1988: fig. 1) iden ti fied seven num bered lithologicalunits (Telm1–Telm7) within the for ma tion, and this sub di vi sionis ac cepted here (Fig. 2). For dif fer ent sub di vi sion schemes and their strati graphi cal cor re la tion see Elliott and Trautman (1982), Marenssi et al. (1998: fig. 4) and Marenssi (2006: fig. 5). TheMesetaichthys-bear ing strata rec og nized within Telm7 (Figs. 2and 3) con sists of about 3 m thick fine grained green-gray sand -stone, mudstone and shell beds. These strata con tain the stud -ied fish Mesetaichthys, the isocrinid Metacrinus fossilis (Ras -mus sen, 1979; Baumiller and GaŸdzicki, 1996: fig. 2), and thebi valves Hiatella and Mya (Fig. 2). It is prob a bly the same ho ri -zon in which the fish ver te bra de scribed by Jerzmañska (1988:fig. 1) were found. In this part of the for ma tion were also foundtwo large “land-birds” (Case et al., 1987; Tambussi et al., 1994;Case, 2006; Cenizo, 2012), nu mer ous bones of pen guins (Sim -pson, 1971; Myrcha et al., 2002; Jadwiszczak, 2006, 2009,2010, 2012; Jadwiszczak and Mörs, 2011) and whale re mains(Borsuk-Bia³ynicka, 1988; Mitch ell, 1989; Fostowicz-Frelik,2003). Myrcha et al. (2002: fig. 2) and Tambussi et al. (2006: ta -bles 3, 4) sum ma rized the in ver te brates and ver te brates of theTelm7 (= Submeseta Allomember).

The Mesetaichthys-bear ing strata (ZPAL 3: GPS po si tion:S 64°14’13.197’’, W 56°38’12.014’’, ~157 m a.s.l.; see Fig. 3)can be cor re lated with the Submeseta Allomember(36.0–34.2 Ma – see Marenssi, 2006: fig. 5) and the Anthro -pornis nordenskjoeldi Biozone (Tambussi et al., 2006: fig. 2b;see also Jadwiszczak, 2006). The age of the top of the LaMeseta Fm. (Telm 7) is dated as 34.2 Ma, close to the Eocene- -Oligocene bound ary (Marenssi, 2006).

The geo chem i cal anal y ses made on the fos sil bi valve shells from the La Meseta For ma tion sug gest a cli ma tic cool ing eventdur ing the time of de po si tion of the up per most part of the for ma -tion (GaŸdzicki et al., 1992; Din gle et al., 1998; Ivany et al.,2006, 2008; Cione et al., 2007; Fran cis et al., 2009; B³a¿ejowski et al., 2010). This is cor re lated with the first Ce no zoic gla ci ationof Antarctica and with the open ing of the Drake Pas sage at theEocene-Oligocene bound ary (Zachos et al., 2001; Birkenmajeret al., 2005). The up per bound ary of the Telm7 is an un con -formity with the Late Mio cene Hobbs Gla cier For ma tion(Marenssi et al., 2010) or the post-Late Plio cene Weddell SeaFor ma tion (Fig. 2; GaŸdzicki et al., 2004).

NOTOTHENIOIDS FROM THE LA MESETA FORMATION

Notothenioids are the dom i nant and most di verse fishesaround Antarctica to day (East man, 2000). One would prob a blyex pect to find this group in the Paleogene, es pe cially in theEocene La Meseta For ma tion on Sey mour Is land, the onlyknown ma rine strata with fos sil fishes in Antarctica (Wood ward,1908; Welton and Zinsmeister, 1980; Grande and East man,1986; Feldmann and Woodburne, 1988; Jerzmañska, 1988,1991; Ward and Grande, 1991; East man and Grande, 1991;Long, 1991, 1992a, b, c; Jerzmañska and Œwidnicki, 1992;Cione et al., 1994, 2001; Crame, 1994; Cione and Reguero,1994, 1995, 1998; Doktor et al., 1988, 1996; Woodburne andCase, 1996; Reguero et al., 2002; Kriwet and GaŸdzicki, 2003;Kriwet, 2005; Kriwet and Hecht, 2008). Wood ward (1908) iden -ti fied some iso lated ver te brae as Notothenia, but their cur renttax o nom i cal sta tus re mains un clear (Grande and East man,1986; East man and Grande, 1989).

568 Ma³gorzata Bieñkowska-Wasiluk, Niels Bonde, Peter Rask MÝller and Andrzej GaŸdzicki

Fig. 1. Geo log i cal sketch map of the north ern part of Sey mour Is land (sim pli fied from Sadler, 1988) show ing the lo cal ity ZPAL 3 where spec i mens Mesetaichthys jerzmanskae n. gen.

and n. sp. were col lected

Eocene relatives of cod icefishes (Perciformes: Notothenioidei) from Seymour Island, Antarctica 569

Fig. 2. Strati graphic col umn of the Eocene La Meseta For ma tion on Sey mour Is land show ingli thol ogy and dis tri bu tion of in ver te brate and ver te brate fos sils

As ter isk shows the strati graphic level of the Mesetaichthys jerzmanskae n. gen. and n. sp.; mod i fied from Myrcha et al. (2002: fig. 2)

The most valid re fer ral of fos sils to the notothenioid group isthat by Balushkin (1994), who redescribed a part of a skull orig i -nally re ferred to Gadiformes by East man and Grande (1991),and in cluded it in the prim i tive notothenioid, Eleginopsidae Gill(see Balushkin, 1992). It was the first de ter mined fos sil noto -thenioid, which Balushkin (1994) named Proeleginops gran -deastmanorum. East man (2000) had been re luc tant to ac ceptthe mis iden ti fi ca tion, but later he ap par ently ac cepted it (seeClaeson et al., 2012). Arratia et al. (2004: p. 48) did not rec og -nize Balushkin’s re vi sion. Proeleginops grandeastma norumfrom lo cal ity RV 8200 in Telm5 (Case, 1992, 2006) which isMid dle Eocene, ca. 45 Ma in age (Case, 2006; Marenssi,2006), is the old est known notothenioid, some 10 m.y. olderthan the ma te rial de scribed here.

Grande and East man (1986) in a re view of all known Ant -arc tic ichthyofaunas, dis cussed the like li hood that jaw frag -ments and ver te bral centra from Up per Eocene of the LaMeseta Fm. could be notothenioids – a pos si bil ity the au thorsnat u rally ex pected. They found no pos i tive in di ca tions, that al -lowed “...a de fin i tive di ag no sis of this group...” al though they did stress dis tinct sim i lar i ties be tween some of the fos sil ver te braeand those notothenioid Dissostichus. Grande and East man(1986: fig. 5) fig ured two premaxillary frag ments and two den ta -ry frag ments from the Field Mu seum of Nat u ral His tory, Chi -cago col lec tion (FMNH), and de ter mined the for mer as “...agadiform type...”, while the den ta ries were de scribed as “...in de -ter mi nate tele osts...”. The lat ter, how ever, are now called �...de -fin i tively Merlucciidae...� by Claeson et al. (2012).

RECENT NOTOTHENIOIDS

Grande and East man (1986: p. 130) dis cussed the size ofthe Eocene fishes based upon the larg est of the ver te brae in theField Mu seum of Nat u ral His tory, Chi cago, which is “...gen er allysim i lar to those of the Re cent nototheniid Dissostichusmawsoni...”, the Ant arc tic toothfish. This spec i men is a cau dalcentrum 4.7 cm in di am e ter and 3.3 cm long. Grande and East -man (1986) com pared the size of this ver te brae to size of ver te -brae of D. mawsoni and know ing the size of D. mawsoni they es -ti mated that fos sil fish had a stan dard length (SL) of ca. 2 m. That was more than known at that time for D. mawsoni as the larg estnotothenioid in the Ant arc tic wa ters. This larg est fish caught byEast man and DeVries (1981) had a to tal length (TL) of 163 cm.To day the more north ern spe cies D. eleginoides – which is found along the coast of south ern South Amer ica from Peru to Uru guay

and at many of the South Ocean is lands – is known to reach a TL of 215 cm (Duhamel et al., 2005). This is the larg est notothenioid, prob a bly ex ceed ing 100 kg. In ter est ingly, this spe cies has justonce been caught off Green land (MÝller et al., 2003; see Fig.15), and it is the only notothenioid ever found north of the Equa -tor. Most of the notothenioids are much smaller reach ing lessthan half a metre in length (see Fig. 14).

Osteological and phylo gen etic stud ies of liv ing notothe -nioids were pre sented by Balushkin (1984, 1992, 2000) andVoskoboinikova (1994, 1997, 1998), who also stud ied the on -tog eny of the skel e ton (Voskoboinikova and Bruce, 2001).Andersen (1984) stud ied the Nototheniidae and their clas si fi ca -tion and Iwami (1985) the Channichthyidae. How ever, pub -lished de tails con cern ing the skull anat omy, and teeth are dif fi -cult to find (only premaxillaries with teeth are shown by Balu -shkin, 1984). Greg ory (1933) had pub lished quite a few noto -thenioid skulls, in clud ing the prim i tive Eleginops that was re -ferred by Balushkin (1992) to its own monotypic fam ily. The mo -lec u lar ev i dence for their re la tion ship is dis cussed by Lecointreet al. (1997), Ritchie et al. (1997), DettaÎ and Lecointre (2004)and Near (2004) us ing the fos sil Proeleginops to cal i brate themo lec u lar clock for notothenioid or i gins.

EARLY GADIFORMS

Tra di tion ally gadiforms are sup posed to have orig i nated inthe north (Svetovidov, 1948). The old est known gadiform and the only Paleocene skel e ton as sign able to this or der is a com pletefish from the Danian of West Green land (Rosen and Patterson,1969; Co hen, 1984). The ear li est di ver si fi ca tion of the gadiformsis re corded by skel e tons from the ear li est Eocene Fur For ma tionof Den mark (Bonde, 1987, 1997; Bonde et al., 2008, 2011). Inthe Lon don Clay (late Early Eocene), there are two taxa of skel e -tal gadiforms, the merlucciid Rhinocephalus and the merlucciid- like teeth called Trichiurides (Casier, 1966; Rosen and Patter -son, 1969); there are also five otolith spe cies (Casier, 1966).

The gadiform otolith re cord is very dif fer ent from that of theskel e tons (Casier, 1966; Patterson, 1993). From the �CoralLime stone� of Faxe (Mid dle Danian, Early Paleocene, Den -mark) a few ju ve nile in de ter mi nate gadiforms are known(Schwarzhans, 2003). In the Dan ish Early Selandian (Mid dlePaleocene) as many as eight spe cies of gadiforms were found:three macrourids and five spe cies in four gadoid fam i lies. OneDan ish eulichthyid is known also from the Paleocene–Eoceneof the United King dom and Bel gium (Schwarzhans, 2003). Twomacrourids and two gadoid spe cies were found in Selandian,West Green land (Schwarzhans, 2004).

In South Aus tra lia and New Zea land the ear li est gadiformoto liths are from the Mid dle Paleocene (Selandian; a macrouridof the liv ing ge nus Nezumia, Schwarzhans, 1985), and 3 spe -cies are known from Early Eocene (Schwarzhans, 1980).Bregmacerotids and prim i tive Euclichthys-like ga doids werefound in Late Eocene. Macruronus oto liths are known from theEocene of South Aus tra lia (Schwarzhans, 1981).

Kriwet and Hecht (2008) pro vided a re view of early gadi -forms and de scribed a macrourid skull with oto liths in situ fromLa Meseta Fm (Telm3–5), an ex cep tional pres er va tion. Rattailskel e tons apart from that skull are un known from the Paleocene and Eocene. Kriwet and Hecht (2008) ac cepted the de ter mi na -tions as gadiforms by East man and Grande (1991) andJerzmañska and Œwidnicki (1992) for jaws from the La MesetaFor ma tion. Jerzmañska (in Doktor et al., 1996: figs. 11 and 12)has iden ti fied scales of gadiforms in the lower part of the LaMeseta Fm. (Telm2, Early Eocene). Balushkin’s ar gu ment(1994), that south ern gadiforms are un likely al ready in �LateEocene�, be cause they orig i nated in the north thus is irrevelant.


Fig. 3. View of the lo cal ity ZPAL 3 (as ter isk),Telm7 (= Submeseta Allomember), Late Eocene

Pho to graph by A. GaŸdzicki, Feb ru ary 1992

SYSTEMATIC PALAEONTOLOGY

Or der PerciformesSuborder Notothenioidei

Fam ily incertae sedisGe nus Mesetaichthys nov.

T y p e s p e c i e s. – Mesetaichthys jerzmanskae sp.nov., only known spe cies. La Meseta For ma tion, Late Eocene,Sey mour Is land, Ant arc tic Pen in sula.

D e r i v a t i o n o f n a m e. – From the La Meseta Fm.and Greek ichthys means fish. The name was used in for mallyby Jerzmañska and Œwidnicki (1992: p. 246).

D i a g n o s i s. – As for the type and only spe cies.S t r a t i g r a p h i c a n d g e o g r a p h i c r a n g e.

– La Meseta For ma tion (Telm7), Late Eocene, Sey mour Is land, Antarctica.

Mesetaichthys jerzmanskae sp. nov.(Figs. 4–11)

H o l o t y p e. – ZPAL P.V./5, a large frag men tary, butnearly com plete den ta ry (Figs. 4, 5 and 6F).

P a r a t y p e s. – ZPAL P.V./1 and 2, prox i mal frag mentsof premaxillae (Fig. 7).

M a t e r i a l. – Re ferred ma te ri als are dis cussed be low af -ter the rel e vant de scrip tions.

T y p e h o r i z o n. – Telm7 of the La Meseta For ma tion,Late Eocene.

T y p e l o c a l i t y. – ZPAL 3, Sey mour Is land, Ant arc ticPen in sula (Figs. 1 and 3).

D e r i v a t i o n o f n a m e s. – Named in hon our of thelate Pol ish palaeoichthyologist Prof. Dr. Anna Jerzmañska (see El¿anowski, 2003).

D i a g n o s i s. – The most char ac ter is tic fea ture is thedentition with strong, con i cal, slightly bent and pointed teethstand ing on large bony pedi cles (Figs. 4–8). Teeth mostly withan in ter nal cav ity with lon gi tu di nal ridges mir ror ing the ex ter nalridges and fis sures in the �enameloid� at the base of the teethproper. Teeth in one row, fang-like teeth in the premaxilla at thesymphysis, but teeth di min ish ing back wards. Small teeth at theden ta ry symphysis, but very large �ca nines� at least in the mid -

dle of the den ta ry. Wide open groove for the man dib u lar sen -sory ca nal (Figs. 4–6F).

D e s c r i p t i o n.

Holotype (The large den ta ry). The larg est den ta ry frag -ment (ZPAL P.V./5; Figs. 4–6F) is about 10 cm long, and a fewcenti metre are miss ing from the pos te rior end of the den taledge and the ven tral branch. The bone is about 1 cm wide at the den tal edge where strong, con i cal teeth are fused to high, bonypedi cles in a shal low groove (see Moy-Thomas, 1934; Fink,


Fig. 4. Left den ta ry, ZPAL P.V./5, holotype of Mesetaichthys jerzmanskae n. gen. and n. sp., dusted with am mo nium

A – lat eral, B – me dial, C – dor sal, D – ven tral view; pho to graphs by G. Dziewiñska

Fig. 5. Left den ta ry, ZPAL P.V./5, holotype of Mesetaichthysjerzmanskae n. gen. and n. sp.

A – lat eral, B – me dial, C – dor sal view; draw ings from Jerzmañska and Œwinicki (1992)

1981). Along the mid dle part of the lat eral face of the bone avery strong, broad ridge is run ning smoothly into the strongsymphyseal re gion, mak ing the bone al most 2 cm wide be lowthe last pre served tooth. Above this ridge along the laterodorsal mar gin there is a broad, dis tinct but rather shal low groove end -ing near the symphysis of the den ta ry, at two nerve fo ram ina,be low the stron gest ca nine tooth. The first nerve fo ra men is ca.1 mm in di am e ter, just pos te rior to and be low the pos te rior faceof this tooth, and the sec ond is 2 mm in di am e ter, be low the an -te rior mar gin of the tooth, it is run ning into a forwardly di rectedca nal (the men tal fo ra men). The pos te rior end of this groove islo cated at the lat eral in ci sion (Fig. 5A), the dor sal rim of whichseems to be the nat u ral thin edge of the bone (that tightly cov -ered the miss ing dermarticular’s an te rior pro cess), while the an -te rior and ven tral edges of this in ci sion are bro ken. The lat erallamina bend ing up at the pedicle be low the large tooth is slightly more frag mented now, when com par ing Fig ure 4A with Fig ure5A (from Jerzmañska and Œwidnicki, 1992), and like wise the in -te rior dor sal flange of the dor sal branch has a dam aged ven traledge slightly above the wide in ter nal in ci sion (see Figs. 4B and5B). The ven tral edge of the den ta ry (Fig. 4D) is very thin andal most straight and forms the in ter nal wall of a deep, opengroove for the man dib u lar sen sory ca nal. Seen from be hind(Fig. 6F) it is clear that the bone wall is quite thin around a wideman dib u lar (Meckelian) ca nal.

The in ter nal face of the den ta ry is al most smooth and plane,only bend ing very slightly in wards at the ven tral edge in the mid -dle of the bone (Figs. 4B, D and 5B). Be low the an te rior smallteeth there is a shal low de pres sion, pre sum ably for a mus cle orlig a ment near the symphysis. The lat ter seems to have its an te -

rior sur face al most in tact, about 15 mmhigh, and slightly bent in wards, in di cat ingthat the an gle be tween the two den ta riesmust have been a very ob tuse one. Thisin di cates a very blunt and rounded snoutwith out any fu sion be tween the twobones, but rather a thin car ti lage. The lat -eral sur face of the bone shows a weak�bulge� or con vex ity be low the interspacebe tween teeth num ber four and five, andfur ther an te ri orly there is a slight de pres -sion be low the three an te rior teeth (Figs.4A and 5A).

The sur face of this part and the strong lat eral ridge has a fine and some what ir -reg u lar or na ment of thin grooves andridges be com ing a lit tle coarser andstraighter on the pos te rior part of the lat -eral ridge (Fig. 4A). This or na ment, notshown in the draw ing (Fig. 5A), is the rea -son for as sum ing that the bones de -scribed here, e.g. the premaxillaries anddermarticulars are prob a bly from thesame spe cies. The or na ment is not well- de vel oped on the in side of the den ta ry(Fig. 4B), where there are only very faintand al most straight lon gi tu di nal groovesand ridges. The ex ter nal face of the ven -tral lamina is or na mented by much stron -ger and more reg u lar lon gi tu di nal gro oves and ridges (Figs. 4A and 5A). The opengroove for the man dib u lar sen sory ca nalis quite smooth and rounded, it is deep estat the pos te rior mar gin of the den ta ry andseems to dis ap pear into a fo ra men nearthe an te rior mar gin of the den ta ry (Fig.4D). The lat eral wall of this groove isformed by the thick ridge. There are nerve

fo ram ina for the sen sory �buds� in the ca nal at the top of the open groove (Fig. 4D), at least four can be seen (they are mis lead inglycalled �...ven tral sen sory ca nal pores...� in Claeson et al., 2012:fig. 4).

The three pos te rior large and very ro bust teeth pre servedare widely spaced. Be tween them are dis tinct de pres sions fromteeth hav ing fallen out. Prob a bly two teeth are miss ing be tweenthe two an te rior large teeth. Each tooth is con i cal, bent in wardsand it is con tin u ing the shape of its bony pedicle. The tooth's�shin ing� enamel-like cover has an or na ment of fine and thingrooves at the base dis ap pear ing to wards the tip of the tooth.These fine grooves were de scribed by Jerzmañska andŒwidnicki (1992: p. 245) as “...ra di at ing from the tip to half or atleast 1/3 height of the tooth...”, be cause ap par ently they did notrec og nize the base as a �bony pedicle� like de scribed here (andby Moy-Thomas, 1934; Fink, 1981). There does seem to be aclearly marked dis tinc tion be tween these pedi cles and the teeth proper with a dif fer ence of the sur face struc ture, as is clearly in -di cated in draw ings (Fig. 5) by Jerzmañska and Œwidnicki(1992). There are four teeth also on pedi cles be ing closelyspaced near the symphysis with bases a lit tle more oval. One isof �in ter me di ate� size, the three an te rior ones are small, theirtips are also in clined in wards, and at least three of them showthe same or na ment at their bases. These basal pedi cles allalong are al most about the same height as the teeth proper.

The tips of most of the teeth are slightly dam aged by post --mor tem breaks ex pos ing the thin outer and shiny layer be ing alit tle lighter col oured than the al most black in te rior, but none isbro ken at a level dis pos ing the in ter nal cav ity. In the interspacesbe tween the teeth the shal low groove shows ir reg u lar �spongy�


Fig. 6. Den ta ry frag ments of Mesetaichthys jerzmanskae n. gen. and n. sp

A–E – dusted with am mo nium; A–C – ZPAL P.V./9 in lat eral, me dial and dor sal views; D, E –ZPAL P.V./11 in lat eral and dor sal views; F – ZPAL P.V./5, the holotype in pos te rior view toshow the large in ter nal cav ity and the open sen sory ca nal; pho to graphs by P.R. MÝller

bone struc ture. No traces of re place ment teeth are seen. Theremight per haps have been two teeth be tween the fourth symphy -seal one and the an te rior large tooth, which in that case would bepo si tion sev enth. Marks on this tooth (Fig. 4B) are from sam plingfor iso tope anal y sis (see B³a¿ejowski et al., 2010).

This al most com plete left den ta ry (Figs. 4–6F) is slightly big -ger than that of the large Dissostichus (~180 cm long, ~70 kg;MÝller et al., 2003) from Green land. Hence it in di cates a fish ofabout 2 m length.

Other den ta ries. Jerzmañska and Œwidnicki (1992: fig. 4)de scribed a small frag ment (ZPAL P.V./6) from the pos te rior endof the den tal ridge with 3–4 closely set and di min ish ing teeth ofwhich the larger one clearly has a bony pedicle (their fig. 4). Thisis ac cepted here as prob a bly rep re sent ing Mesetaichthysjerzma nskae.

There are two other well-pre served den ta ries (ZPAL P.V./9and 11; Fig. 6A–E) show ing the symphyseal re gion. Both arefrom the left side and are 30 mm long and very close to 15 mmhigh at the symphysis as ZPAL P.V./5, but they are slightly shal -lower than the lat ter pos te ri orly to the symphysis. ZPAL P.V./9(Fig. 6A–C) is the better pre served end ing at a clear and nearlyver ti cal cut through the bony base of a tooth cor re spond ing tothe 6th or 7th po si tion in ZPAL P.V./5, which rep re sents the an -te rior large tooth.

Spec i men ZPAL P.V./11 al though more bro ken (Fig. 6D, E),rep re sents an te rior part of den ta ry and is lack ing part of the me -dial sur face. It has seven tooth-po si tions pre served, the last ir -reg u larly bro ken bony pedicle cor re spond ing to the an te rior bigone of spec i men ZPAL P.V./5 (Fig. 4). Its small an te rior tooth ismiss ing from its po si tion very close to the me dial face di rectlypos te rior to the symphysis, but teeth num bers two to four arepre served as bony pedi cles cut near tops of pendicles, and teethtwo and four are show ing traces of the in ter nal cav i ties (Fig. 6E).The fifth tooth has left a deep, round groove, while the ratherstrong sixth is cut al most ver ti cally through the mid dle of its highbony pedicle and its in ter nal cav ity. The lat ter, as well as the base of the cav ity of sev enth tooth, shows weak traces of in ter nal, ver -ti cal ridges and fur rows, and the same is the case in the basalcav ity of the bony pedicle of one of the pos te rior pre served toothin spec i men ZPAL P.V./9. The lat ter is lack ing the small sec ondtooth, and all teeth be tween the third and pos te rior pre servedtooth, all leav ing deeper or shal lower pits, which show no tracesof the in te rior cav i ties (Fig. 6C). Teeth num ber one and threeclearly show fine ridges and fur rows at the base of the toothproper, and when ap proach ing the large pos te rior tooth the lat -eral edge of the den tal groove is as cend ing like in spec i menZPAL P.V./5. First tooth is the only tooth of M. jerzmanskae pre -serv ing the al most com plete tip, clearly with out a sharp �cap�.

Spec i men ZPAL P.V./9 ex hib its the an te rior end of the large man dib u lar (Meckelian) ca nal, the an te rior 22 mm of thesmooth groove for the man dib u lar sen sory ca nal, and the large, lat eral fo ra men be low the large pos te rior tooth. The bony sur -face shows the same or na men ta tion as spec i men ZPAL P.V./5, al though slightly stron ger lon gi tu di nal ridges mark the pos te riorpart of the me dial sur face of the frag ment (Fig. 6A).

There is no rea son to doubt that these two den ta ry frag ments rep re sent M. jerzmanskae. Very im por tantly, they show the in ter -nal cav i ties of the bony pedi cles with their faint ridges and fur -rows. The re main ing den ta ries, spec i mens ZPAL P.V./6–16 alsoseem to be long to this spe cies, but are more frag men tary.

Premaxillae. The better pre served premaxillae are spec i -mens ZPAL P.V./1, 2, de scribed and fig ured by Jerzmañska

and Œwi dnicki (1992: fig. 1; here Fig. 7), both from the left sidewith the prox i mal and ar tic u lar head of the bone, both here se -lected as paratypes. We fea ture in some de tail spec i men ZPALP.V./1 with two an te rior �fangs� be ing pre served (Fig. 7A, C,E–G). The short rounded as cend ing pro cess is well-pre servedand shows an ex ter nal or na ment sim i lar to that of the lat eralface of the den ta ries. This rather thin and flat pro cess has an in -ter nal face (Fig. 7F) with a coarse or na ment of straight, thin


Fig. 7. Symphyseal re gions of premaxillaries of Mesetaichthys jerzmanskae n. gen. and n. sp.

A, C – ZPAL P.V./1 in left lat eral and ven tral views; B, D – ZPALP.V./2 in me dial and left lat eral views; E–G – ZPAL P.V./1 in left lat -eral, me dial and latero-ven tral views; A–D – draw ings fromJerzma ñska and Œwidnicki (1992); pho to graphs by P.R. MÝller

ridges ra di at ing out from the me dial cav ity be tween the as cend -ing and ar tic u lar pro cesses. This me dial cav ity also has 2–3nerve fo ram ina �ra di at ing� in the same di rec tion as ridges of theas cend ing pro cess (also seen in ZPAL P.V./2; Fig. 7B). The as -cend ing pro cess is about 7–8 mm thick at its ba sis, but verythin, al most sharp at the dor sal edge. Only the an te rior part ofthe strong ar tic u lar pro cess is pre served (Fig. 7A, E, F). Spec i -men ZPAL P.V./2 shows the en tire ar tic u lar pro cess which isabout the same size as the as cend ing pro cess, but stron gerand slightly more nar row (Fig. 7B, D). It shows a lat eral, ver ti calridge in the mid dle of the pro cess, pre sum ably in di cat ing howfar for ward the ar tic u lar head of the maxilla would have cov eredthe premaxilla. Only a small part of the dentigerous branch ofthe bone is pre served in ZPAL P.V./2 cor re spond ing roughly tofour tooth po si tions but only the bony pedicle of the large an te -rior �fang� is pre served. In ZPAL P.V./1 the dentigerous branchof the bone is cor re spond ing to three an te rior teeth (Fig. 7E–G). Spec i men ZPAL P.V./1 has the two an te rior teeth and pedi clespartly pre served, the sec ond one bro ken in the pedicle show ingtraces of the in ter nal cav ity. The an te rior tooth is a very strong,con i cal and in wardly bent �fang� with the tip bro ken off to dis -close a large in ter nal cav ity (Fig. 7F). In clud ing nearly 15 mm as a bony pedicle, the tooth it self with strong, closely set ridges atthe base is about 7 mm high as pre served, and re stored withthe point would have been at least 2.5 cm high. The width of itsbase is about 8 mm, and the �enameloid� of the tooth has a lightbrown col our, as op posed to the much darker bone and thedentine in te rior of the tooth which is nearly black (Fig. 7F). Thepos te rior wall to wards the next tooth is slightly con cave giv ing itin fact a sub-tri an gu lar base (Fig. 7C, E, G). The sec ond tooth is bro ken at the top of the bony pedicle and as pre served is ca.12 mm high, also bent in wards, and with the in ter nal cav itymostly filled by spongy bone. Its base is 6 mm wide, but slightlywider per pen dic u lar to the jaw giv ing it an �ovaloid� shape, butwith a nearly flat front wall. The pos te rior wall is a lit tle con cave,ap par ently worn by the next and now miss ing tooth, which hasleft only a shal low cav ity (Fig. 7C, G). So the teeth are veryclosely set, de creas ing in size back wards, and or na mented likethe den ta ry teeth. They must have formed an �over hang ing� up -per jaw op pos ing the small teeth at the symphysis of the den ta -ry (Figs. 4 and 5). Spec i men ZPAL P.V./3 may also be from thisspe cies. Grande and East man (1986) fig ured sim i lar frag mentsmostly with miss ing teeth.

Spec i men ZPAL P.V./4 is a slim dis tal part of a premaxillawith three teeth and in ter ven ing shal low grooves of four miss ing ones (Fig. 8; as sum ing that this fish is much too ad vanced tohave a toothed maxilla). Only the bony pedi cles are pre servedof the teeth, and they have dark spongy bone in the in te rior andno cav ity. On its dor sal mar gin is a shal low groove which mayhave ac com mo dated the ven tral edge of the maxilla. We as -sume that all premaxillaries be long to M. jerzmanskae.

Jaw ar tic u la tion. Sev eral strong dermarticulars werefound, and the two best pre served ones (ZPAL P.V./17, 18) arefig ured by Jerzma ñska and Œwidnicki (1992). The an a tom i callymore com plete frag ment (ZPAL P.V./18; Fig. 9C–F) is from theright side, ca. 3.5 cm long show ing the ar tic u lar sur face, whichex tends much fur ther down the me dial face than on the lat eralface. The length of the ar tic u lar or glenoid cav ity is ca. 15 mmmea sured across to the tip of the post-ar tic u lar pro cess. Thestrong post-ar tic u lar pro cess is ca. 13 mm high, and the ven traledge is in com plete where the retroarticular is miss ing. Be lowthe ar tic u lar sur face on the lat eral side is a wide open groove for the man dib u lar sen sory ca nal which has a small ridge over -hang ing it from the strong lat eral ridge of the der mal com po nent (Fig. 9C, D). The ar tic u lar sur face con tin ues for ward by the dor -sal as cend ing part of the bone at an an gle of about 30o with re -

in force ment along the dor sal edge. On the in side there is asmall cav ity up be hind the down-turned ar tic u lar sur face, andthere are two nar row cav i ties an te rior to the ar tic u lar sur face.There is a large cav ity in the bone be tween the strong lat eralridge (lat eral wall) and the me dial wall (Fig. 9E, F).

Spec i men ZPAL P.V./17 (Fig. 9A, B) from the left side isfrom a much big ger an i mal and is 4 cm long and al most 2 cmmea sur ing across the ar tic u lar sur face, and part of the bone isbro ken off an te ri orly. Its post-ar tic u lar pro cess is 25 mm high,and a tiny bit may be miss ing at the sharp ven tral edge. Roughand pit ted lat eral sur face shows where the miss ing retroarti -cular was su tured (Fig. 9A). The ven tral part of the bone be lowthe wide open groove for the sen sory ca nal is much higher thanpre served in spec i men ZPAL P.V./18, and the ven tral edgeseems bro ken and in com plete. The same three cav i ties orpock ets are seen be hind and in front of the smooth me dial�tongue� of the ar tic u lar sur face (Fig. 9B). The dor sal edgeslant ing up wards is not pre served. The lat eral sur face of thebones (Fig. 9A, C) has a struc ture rem i nis cent of that of theden ta ries (Figs. 4 and 6). Both bones are very in com plete infront with the pre sum ably long and pointed pro cess for ar tic u la -tion with the den ta ry bro ken off. We as sume that the de scribed(ZPAL P.V./17, 18) and the re main ing five dermarticulars(ZPAL P.V./19, 23) be long to M. jerzmanskae.

Basioccipital. This sin gle bone (ZPAL P.V./24; Fig. 10) is26 mm wide and 21 mm high at the con cave ar tic u lar facet,and it is 45 mm long at the midline, but bro ken at the front mar -gin. There are two deep ex ca va tions in the ven tral sur face(Fig. 10F) for thick pos te rior prongs of the parasphenoid with astrong ridge and in front a nearly 1 cm high, ver ti cal wall be -tween them (Fig. 10A). The dor sal in te rior sur face has a bluntridge in the midline on top of the men tioned wall, and the bonehas a flat slop ing sur face dip ping about 20° to wards the lat eraledge (Fig. 10A, D). The lat ter is bro ken at both sides (Fig. 10A, B), but rather thin, and the bone may not have been muchwider than about 3 cm. Com par ing with many skulls fig ured by


Fig. 8. Sup posed more dis tal right premaxillary frag ment ofMesetaichthys jerzmanskae n. gen. and n. sp. (ZPAL P.V./4)

A – ven tral view; B, C – draw ings from Jerzmañskaand Œwidnicki (1992) in me dial and ven tral views;

pho to graph by P.R. MÝller

Balushkin (1984) the lat eral wall may have been rather low,and there fore not much ap pears to be miss ing. Grande andEast man (1986) un for tu nately did not de scribe thebasioccipital of the fos sil skull.

Just in front of the dor sal edge of the facet for ar tic u la tion(Fig. 10C, E) is a sub-hor i zon tal ir reg u lar sutural sur face for theexoccipitals (Fig. 10D), in di cat ing that these were in a pos te riorpo si tion at the level with the facet, and they met each otherabove the facet. This sutural sur face also has paired and ratherdeep, blunt cav i ties which are 4 mm long and 3 mm wide, and in front and be low from those there are two much broader cav i ties1 cm across still filled with ma trix, and form ing the pos te rior wallof the ven tral part of the brain cav ity (Fig. 10A, D).

This ar range ment does not look at all like that ingadiforms, were the exoccipital ar tic u la tions are placed muchfur ther an te ri orly, and those two bones do not meet in themidline (see Greg ory, 1933; Svetovidov, 1948; Mujib, 1967;Rosen and Patterson, 1969). This crit i cism was made byBalushkin (1994) con cern ing the exoccipital-basioccipital po -si tions in the skull orig i nally de scri bed by East man andGrande (1991) as a �gadiform�, but shown by Balu shkin(1994) to be a noto thenioid. It is likely that in a fish of this size,in di cated by the iso lated basioccipital, the exoccipitals andbasioccipital would be fused to gether in a gadi form, whilenotothenioids tend to os sify very late (Voskobo inikova, 1994).

It is pos si ble that the basioccipital(ZPAL P.V./24) be longs to M.jerzmanskae, but this is un cer tain.

Ver te brae. The de scribed col lec -tion com prises 21 ver te brae withshort centra (ZPAL P.V./25–45; Jerz -mañ ska and Œwidnicki, 1992). Thecentra were mea sured by Jerz -mañska and Œwi dnicki (1992: ta ble2). Some have quite strong parapo -physes (Fig. 11A–D), as also in somenothenioids and many ga do ids. Many of the centra show sev eral lat eralbony lamellae (Fig. 11B, F; like theirfigs. 7 and 8 of ZPAL P.V./31, 32), afea ture found in both somenotothenioids (Fig. 12E) and somegadiforms.

Spec i mens (ZPAL P.V./29; Fig.12C, D and ZPAL P.V./30) hav inglarge, lat eral cav i ties for ribs, could befrom dif fer ent spe cies than the oth ers.They are un likely to be noto thenioids,as these have weak or miss ing ribs(Grande and East man, 1986: p. 128).Such centra were men tioned by Jerz -mañska (1988: p. 425) and spe cialsim i lar ity was found with the mer lucciid Macruronus. Some of the cen tra(ZPAL P.V./34–42) have been de ter -mined as some thing more spe cific byJerzmañska (dated 1992 on the la -bels), namely “Macruronus” (see Fig.12A, B). Jerzmañska (1988: figs. 7, 8)il lus trated very sim i lar centra and de -scribed them as gadiforms.

Gadiform ver te brae are not usu -ally short, but rather equally long andhigh. Sev eral of the centra are from

quite large fishes, prob a bly over one metre long. The ver te braecan not with cer tainty be re ferred to M. jerzmanskae, but largesize might in di cate, that some of them are quite likely from thistaxon.

O c c u r r e n c e. – La Meseta For ma tion (Telm7), LateEocene.

COMPARISONS

Grande and East man (1986) dis cussed the pos si bil ity thatup per and lower jaw frag ments they il lus trated from the LaMeseta Fm. were notothenioids, and they de cided that thepremaxillaries were prob a bly from gadiforms, and the den ta rieswere in de ter mi nate like the ver te brae. This opin ion was changedcon cern ing den ta ries (Claeson et al., 2012) when de scrib ing apoorly pre served bone with frag men tary dentition, and the de ter -mi na tion be came much more pre cise as �merlucciids�, fol low ingJerzmañska and Œwidnicki (1992). Clearly these jaw frag mentsare of the same sort as the more well-pre served ones de scribedhere, just from much smaller fishes.

The jaw frag ments de scribed by Jerzmañska and Œwidnicki(1992) were only com pared with gadoid fishes (8 spp. from 3–4fam i lies). Seven fea tures were used (two from premaxillae,


Fig. 9. Dermarticulars of Mesetaichthys jerzmanskae n. gen. and n. sp.

A, B – ZPAL P.V./17, left side, lat eral and me dial views; C–F – ZPAL P.V./18, right sidein lat eral (C, D) and me dial (E, F) views; pho to graphs (A, C, E) by P.R. MÝller;

draw ings (B, D, F) from Jerzmañska and Œwidnicki (1992)

three from den ta ries and two from dermarticulars), and of thesetraits six were sim i lar in Merluccius, while the rest (in clud ing themerlucciid Melanonus) only matched one or two char ac ters.The con clu sion was �gadiform� re la tion ship for the fos sils, al -though the match with the ma jor ity of the ma te rial was poor, and they in di cated (Jerzmañska and Œwidnicki, 1992: p. 248) thatthe dentitions are quite dif fer ent. Claeson et al. (2012) took thede ter mi na tion a step fur ther as �merlucciid� (al though very lit tlesim i lar ity with Melanonus), and they tried to in voke also thedentition as an ar gu ment. We find es pe cially the lat ter en tirelyun con vinc ing, be cause Merluccius has very dif fer ent dentition.

Notothenioids like Chaenocephalus (Fig. 14E) have pre -maxillary as cend ing and ar tic u lar pro cesses not un like those ofthe fos sils which like in most gadiforms are low and rounded,not with the as cend ing pro cess higher and slim mer than the ar -tic u lar pro cess, as is the case in by far most perciforms and also nototheniids like Dissostichus (Fig. 15D; see also Balushkin,1984: fig. 15; Greg ory, 1933). The �trend of de vel op ment� innotothenioids is the re duc tion in rel a tive size of the as cend ingpro cess of pre maxilla (Balushkin and Voskoboinikova, 1995).

We rec og nize that the fos sil den ta ries in di cate a broad androunded snout. The dentition is very un like that of gadi forms, as none of those have �fangs� at the front of the premaxillary, nei -ther gadi forms have small teeth at the den ta ry symphysis.

The dentition of the gadoid Mer luccius (see Figs. 13 and14C, D) with sharp, pointed, rather strong teeth in two rows isnot sim i lar to the dentition of Mesetaichthys: the teeth are slimand smooth (also in side) and there are no �fangs�, they areabout the same size all along the jaws. They also have a smallsharp �cap� (Fig. 14G, H) on each tooth, which ad mit tedly couldhave been bro ken off in the Mesetaichthys jaws show ing onlyone of the small symphyseal teeth with the point pre served. The only sim i lar ity be tween the dentition of Mesetaichthys andMerluccius is that the teeth are bent in wards (Figs. 5C, 7F and13) and some are hol low (Fig. 14H).

One fea ture which could be a gadi form one is the large,open groove for the man dib u lar sen sory ca nal (Figs. 4A, D; 5Aand 6F), as also stressed by Claeson et al. (2012). Mostgadiforms have such open grooves (Fig. 14D), while this is veryrare in perciforms (ex cep tions e.g., per coids-like Howellidae;see Prokofiev, 2007), and we have seen none in notothenioids,nei ther in our few skel e tons (Fig. 14) nor our X-rays (Fig. 15).The open sen sory man dib u lar ca nals are un known in noto -thenioids (ac cord ing to per sonal in for ma tion from Dr. Balu -shkin, April 2012; see also Claeson et al., 2012). We are, how -ever, struck by some de tailed sim i lar ity in the dentition of thesefos sils and liv ing notothenioids, spe cif i cally that of the hugeDissosti chus (Fig. 15). Mesetaichthys has two strik ing sim i lar i -


Fig. 10. Basioccipital, sup pos edly from Mesetaichthys jerzmanskae n. gen. and n. sp. (ZPAL P.V./24)

A – an te rior, B – right lat eral, C – pos te rior, D – dor sal, E – pos te rior, F – ven tral view;pho to graph by M. Dziewiñski, dusted with am mo nium (A) and by P.R. MÝller (B–D);

draw ings (E–F) from Jerzmañska and Œwidnicki (1992)


Fig. 11. Ver te brae of ?Mesetaichthys jerzmanskae n. gen. and n. sp.

A–D – ZPAL P.V./31 in an te rior, right lat eral, ven tral and pos te rior views; E–H – ZPAL P.V./32 in an te rior, right lat eral, ven traland pos te rior views; pho to graphs by P.R. MÝller (C–D) and by M. Dziewiñski (G–H, am mo nium dusted); draw ings (A, B, E, F)

from Jerzmañska and Œwidnicki (1992)

Fig. 12. Ver te brae

A, B – ab dom i nal of so-called“Macruronus”, ZPAL P.V./35, ?an te riorand ven tral views; C, D – ZPAL P.V./29, an te rior ab dom i nal with large, deep ribcav ity from Teleostei indet; E, F – ab -dom i nal ver te bra of re cent Noto thenia;G–J – Wood ward’s so-called Noto -thenia sp., fore most ver te bral cen trumin an te rior, pos te rior, left lat eral andven tral views (from Wood ward, 1908);pho to graphs by P.R. MÝller (A, B) andby M. Dziewiñski (C, D dusted with am -mo nium)

ties to this liv ing rather prim i tive nototheniid in the pat tern of thedentition and in the de tailed struc ture of the teeth. Dissostichusalso has fanglike teeth at the pre maxillary symphysis, verysmall teeth at the den ta ry symphysis, fanglike teeth in the mid -dle of den ta ry, and the sin gle teeth fur rowed on the base at thepedicle, and the cav ity is ridged to cor re spond to the ex ter nalstriation.

There is only lit tle over lap in basioccipital (Fig. 10) and theskull roof and neurocranium of the notothenioid (Mid dleEocene) from the La Meseta For ma tion, Proeleginops grande -a st manorum Balushkin (1994), but, in fact, the lat ter has abasio ccipital which is not re ally like a gadiform one. There forewe do not know if the two taxa could be syn on y mous, and in that case the skull roof only a few cm long would clearly be from a ju -ve nile spec i men.

Iso lated centra from Sey mour Is land were de scribed byWood ward (1908: fig. 5) and re ferred to Notothenia (Fig.12G–J). It seems cer tain that there is more than one type of ver -te brae pre vi ously re ferred to “Mesetaichthys” and ac cord inglymuch doubt about which centra may be long to the skull bonesof M. jerzmanskae, and this raises doubt also about the re fer ralof the basioccipital. It seems quite likely that some centra arenotothenioid, and that the larger ones could be long to M.jerzmanskae.

NOTOTHENIOID EVOLUTION ANDDETERIORATION OF CLIMATE

The old est rep re sen ta tives of notothenioids, P. grande -eastmanorum Balushkin, 1994, (Telm5) and M. jerzman -skae n. gen. and n. sp. (Telm7) are rec og nized in the LaMeseta For ma tion. In the for ma tion a grad ual de te ri o ra tion ofcli mate is re corded, with the ev i dence of de creas ing tem per a -tures through the later Eocene cul mi nat ing with the first Ant -arc tic glaciations at the Eocene-Oligocene bound ary (GaŸdzi -cki et al., 1992; Din gle et al., 1998; Birkenmajer et al., 2005;Ivany et al., 2006, 2008; Tatur et al., 2006; Fran cis et al., 2009; see also Cantrill and Poole, 2012). Most pos si bly, the or i gin of

the notothenioids is linked to this cli ma tic event, as was hy -poth e sized al ready by Regan (1914) and men tioned byGrande and East man (1986: p. 134).

The ear li est di ver si fi ca tion of notothenioids with anti-freezeliq uids in the blood (AFGP of Near, 2004) ac cord ing to the mo -lec u lar bi ol o gists took place in the ear li est Mio cene (~24 Ma;


Fig. 13. Merluccius merluccius (Jrn. 215, dry skel e ton), gapeshow ing the strong, slim, pointed teeth with glit ter ing acrodincaps in two rows in the jaws (and strong vomerine teeth arebarely vis i ble)

Pho to graph by P.R. MÝller

Fig. 14. Dry bones of re cent fishes

A, B – Notothenia microlepidota (ZMUC uncat., 7495 –dry skel e ton), left lower jaw and right premaxilla, both inlat eral view; C, D – Merluccius merluccius (fish shop,Co pen ha gen, ZMUC uncat.), views as above; E, F –Chaenocephalus aceratus (ZMUC uncat., 7741 – dryskel e ton), left lower jaw and premaxilla; G, H – Mer -luccius merluccius tooth, SEM photo show ing smoothpulp cav ity, cut ting edge, also on the dis tinct cap/tip(uncat. spec i men); pho to graphs by P.R. MÝller (A–F)and C. Bonde (G–H)

Near, 2004). The prim i tive nototheniid branch with Dissostichushas a min i mum age of about 14 Ma (Near, 2004), so that the35 Ma old Mesetaichthys could be in a stemgroup po si tion tothe AFGP notothenioids.

CONCLUSIONS

Jaw frag ments of tele ost ean fishes and pos si bly abasioccipital and some ver te brae from the Late Eocene of theLa Meseta For ma tion (Telm7) of Sey mour Is land are con sid -ered to be notothenioids (cod icefishes rel a tives amongperciforms). Pos tu lated �gadiform� fea tures are shown to be du -bi ous, apart from the open groove for the sen sory ca nal in thelower jaw. This fea ture is seen in a few perciforms, but is notknown in any liv ing notothenioid.

The most well-pre served lower jaw frag ment (Figs. 4, 5 and6F) was se lected as the holotype for Mesetaichthys jerzman -skae n. gen. and n. sp., a notothenioid incertae sedis – withpremaxillary tooth bear ing frag ments as pararatypes. The di ag -nos tic fea tures are pres ent in the dentition (which is re mark ablysim i lar to that of the big, pre da ceous nototheniid Dissostichus)in com bi na tion with the open groove for the sen sory ca nal. TheMesetaichthys jerzmanskae n. gen. and n. sp. de scribed hereas notothenioid and a 10 m.y. older skull of Proeleginopsgrandeastmanorum Balushkin, 1994 de scribed ear lier alsofrom the La Meseta For ma tion are the only fos sil notothenioidsrec og nized up to now.

COMPARATIVE MATERIAL

Re cent Notothenioids, Nat u ral His tory Mu seum of Den -mark, Uni ver sity of Co pen ha gen, SNM.


Fig. 15A–C – large teeth from the mid dle of left den ta ry and near the premaxillary symphysis ofthe huge Dissostichus eleginoides (ZMUC P632150, 180 cm, 70 kg, wet spec i men) in lat eral view show ing fis sures and the pedicle like base, in sec tion show ing the “ridged” soft in te rior pulp,and from the base look ing into the cav ity with ridges on the wall; D – X-ray of the snout and jawsof a small D. eleginoides ex pos ing large fangs near the premaxillary symphysis and in mid dlepart of the den ta ry, which has smaller teeth at the symphysis (ZMUC P6341, 28 cm SL); E – jawsof Dissostichus eleginoides (ZMUC P632150, 180 cm) show ing dis tri bu tion and size of teeth

Pho to graphs by P.R. MÝller

Nototheniidae: Notothenia microlepidota ZMUC uncat.(7495 – dry skel e ton; Figs. 12E, F and 14A, B); Dissostichuselegino ides ZMUC P632150 (155 cm SL; Fig. 15; MÝller et al., 2003: fig. 1), ZMUC P6341 (28 cm SL, X-ray; Fig. 15).

Chaenichthyidae: Chaenocephalus aceratus ZMUCuncat. (7741 – dry skel e ton; Fig. 14E, F).

Eleginopsidae: Eleginops maclovinus ZMUC P63275(84 mm SL, clear and stain); ZMUC CN 2 (365 mm SL, X-ray).

Re cent Gadiforms (skel e tons in Nat u ral His tory Mu seumof Den mark, Uni ver sity of Co pen ha gen, SNM).

Merlucciidae: Merluccius merluccius Jrn. 215–217 (Figs.13 and 14C, D, G, H).

Gadidae: Arctogadus glacialis ZMUC P371663, Boreoga -dus saisa CN 16, Brosme brosme Jrn. 28, Gadus morhuaZMUC P374478, P372971, Lota lota Jrn. 30 x, Melano -grammus aeglefinus Jrn. 301, Merlangus merlangus Jrn. 348,Molva molva Jrn. 58, Pollachius pollachius Jrn. 835, Triso -pterus esmarkii ZMUC P3711, Trisopterus luscus Jrn. 9.

Phycidae: Phycis blennoides Jrn. 41, Urophycis brasilien -sis Jrn. 30(I).

Moridae: Antimora rostrata ZMUC uncat. (Ingolf exp.), Moramora Jrn. 76.

Macrouridae: Coryphaenoides rupestris ZMUC uncat.(Skagerrak), Macrourus berglax Jrn. 15, Trachyrincus murrayiZMUC P375081.

Re cent Lophiiforms. Lophiidae: Lophius piscatorius ZMUCJrn. 62.

Ac knowl edg ements. Dis cus sions with Dr. A.V. Balushkin(St. Pe ters burg) and Dr. J. Niel sen (Co pen ha gen) on the anat -omy on notothenioids are very much ap pre ci ated. We are verygrate ful to re view ers Prof. J. Kriwet (Vi enna) and Dr.P. Jadwiszczak (Bia³ystok) whose com ments have led to im -prove ments of the manu script. NB are very thank ful to the FurMu seum for fi nan cial sup port to study-trips to the In sti tute ofPaleobiology PAS, Warszawa. AG wishes to ac knowl edge thelo gis tic sup port of the Instituto Antártico Argentino and FuerzaAérea Ar gen tina dur ing the field work in Antarctica. Prof. A.Tatur (Warszawa) ac com pa nied AG dur ing field col lec tion ac -tiv i ties. K. Gre gersen, Zoo log i cal Mu seum (Co pen ha gen) made skel e tons of the notothenioids.

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Documents

Eocene relatives of cod icefishes (Perciformes: Notothenioidei) from Seymour Island, Antarctica