SPHENOTHALLUS HALL, 1847 FROM CAM BRIAN
OF SKRYJE–TÝØOVICE BA SIN (BARRANDIAN AREA,
CZECH RE PUB LIC)
Oldøich FATKA & Petr KRAFT
Charles Uni ver sity in Prague, Fac ulty of Sci ence, In sti tute of Ge ol ogy and Palae on tol ogy, Albertov 6, 128 43 Prague 2,
Czech Re pub lic; e-mails: fatka@natur.cuni.cz; kraft@natur.cuni.cz
Fatka, O. & Kraft, P., 2013. Sphenothallus Hall, 1847 from Cam brian of Skryje–Týøovice Ba sin (Barrandian area, Czech Re pub lic). Annales Societatis Geologorum Poloniae, 83: 309–315.
Ab stract: Two con i cal tu bules from shales of the Skryje Mem ber (Buchava For ma tion) at the Skryje–Luh and Týøovice–“Pod hruškou” lo cal i ties, two of the key out crops of this strati graphic unit in the Skryje–Týøovice Ba sin (Barrandian area, Czech Re pub lic), are de scribed. One spec i men con sists of a small, com pressed tu bule with a very low ex pan sion an gle and wide and flat thick en ings. The sec ond, larger spec i men ex hib its in di ca tions of very nar row thick en ings of a more abruptly ex pand ing shell. Both spec i mens are as signed to the ge nus Sphenothallus Hall, but the lat ter only pro vi sion ally. Sphenothallus shows a world wide dis tri bu tion with nu mer ous spe cies, rang ing from Cam brian to Perm ian in age. How ever, re ported oc cur rences in the Cam brian are rel a tively sparse, in the form of rare spec i mens from the lower to mid dle Cam brian strata of Laurentia, East ern Gond wana and Eu ro -pean peri-Gond wana. Ac cord ing to ac cepted palaeo geo graphi cal re con struc tions, Cam brian Sphenothallus oc cu-rred in low as well as in higher palaeolatitudes.
Key words: Sphenothallus, mid dle Cam brian, Buchava For ma tion, Barrandian area Czech Re pub lic. Manuascript re ceived 5 August 2013, ac cepted 27 December 2013
IN TRO DUC TION
The ge nus Sphenothallus Hall, 1847 is a wide spread
Palaeozoic (Cam brian to Perm ian) ma rine taxon of prob
-lem at i cal, sys tem atic po si tion. It orig i nally was de scribed as
a ma rine plant by Hall (1847, p. 261), but since the twen ti
-eth cen tury it has been rec og nized as an in ver te brate, with
pre vi ous au thors var i ously clas si fy ing it as incertae sedis
(e.g. Neal and Hannibal, 2000), as a “worm” (Fauchald et
al., 1986), or as a hydrozoan or scyphozoan cni darian (e.g.
Van Iten et al., 1992, 2013). In the Barrandian area, a small
num ber of oc cur rences of Sphenothallus have been re ported
from Or do vi cian to De vo nian fos sil as so ci a tions of the
Pra-gue Ba sin (Brabcová and Kraft, 2003, p. 265), but un til now
there has been only one study, deal ing with these fos sils sys
-tem at i cally. In that re port (Fatka et al., 2012), rare spec i mens
of the smaller Sphenothallus kozaki Fatka et al., 2012 and the
gi ant ?S. kordulei Fatka et al., 2012 are de scribed from the
Jince For ma tion of the Cam brian Pøíbram–Jince Ba sin.
The pur pose of the pres ent pa per is to de scribe the first
oc cur rence of pre sumed con i cal phos phatic fos sils in the
mid dle Cam brian Buchava For ma tion of the
Skryje–Týøo-vice Ba
sin (Barrandian area; Fig. 1B), which is lo
cated
north of the pre
vi
ously men
tioned Pøíbram–Jince Ba
sin
(Fig. 1A); both bas ins strik ingly dif fer not only in their geo
-log i cal his tory, but also in con tained fos sil as so ci a tions (see
Geyer et al., 2008).
GEO LOG I CAL SET TING
AND AS SO CI ATED FOS SILS
Both spec i mens were col lected from the Skryje Mem
-ber of the mid
dle Cam
brian Buchava For
ma
tion in the
Skryje–Týøovice Ba sin (Fig. 1C), which is well known for
its di
verse fos
sils (e.g., Barrande, 1852; Šnajdr, 1958;
Fatka, 2004). A rich fauna of trilobites, agnostids, hyoliths
and brachi o pods as so ci ated with rare echinoderms and mol-
luscs has been stud ied for nearly two hun dred years (e.g.,
Šnajdr, 1958; Chlupáè, 1999; Geyer et al., 2008).
Acritarchs and ichnofossils have been de scribed since the sec
-ond half of the twen ti eth cen tury (for sum mary, see Fatka,
1999; Mikuláš, 2000). Re cently, rare spec i mens of a poorly
biomineralized and soft-bod ied Bur gess Shale-type fauna
were stud ied by Chlupáè and Kordule (2002), Maletz et al.
(2005) and Fatka et al. (2011); some of these spec i mens are
pre served to gether with di min u tive ichnofossils (Mikuláš et
al., 2012). Re cently, the fodinichnial as so ci a tion rep re sent
-ing a spe cific type, termed “mortichnia”, was es tab lished by
Fatka and Szabad (2011) in the Skryje Mem ber.
Gen eral over views of the depositional set ting and stra
-tig ra phy of the Skryje–Týøovice Ba sin, in clud ing the
Bu-chava For ma tion, are avail able in Kukal (1971), Havlíèek
(1971), Geyer et al. (2008) and Fatka and Mergl (2009).
Fatka et al. (2011) sum ma rized all ear lier data, con cern ing
the lithostratigraphic sub di vi sions of the sed i men tary
sequence; three units, the Skryje, Slapnice and Mileè mem
-bers, are now dis tin guished within the Buchava For ma tion
(Fig. 1C).
De spite more than 150 years of in ten sive in ves ti ga tion
by both pro fes sional palaeontologists and am a teur col lec
-tors, only two spec i mens of Sphenothallus are known from
the Buchava For ma tion. Both were re cently re dis cov ered in
the col lec tion of the late V. Kordule, who care fully gath ered
Cam brian fos sils and brought to gether di verse non-tri lo bite
fos sils, col lected by him and sev eral oth ers dur ing the last
forty years. Sphenothallus sp. orig i nates from the Skryje –
Luh lo cal ity, while ?Sphenothallus sp. was col lected at the
Týøovice – “Pod hruškou” lo cal ity.
Skryje–Luh lo cal ity
The large nat u ral out crop (Fig. 2), known as the “Skryje
– Luh lo cal ity”, be longs to the clas si cal sec tions in the up
-per half of the Buchava For ma tion. In old col lec tions, this
out crop did not get a sep a rate name, and spec i mens from the
slope above the bridge were des ig nated as Skrey in the col
-lec tion of Joachim Barrande in the Na tional Mu seum
Pra-Fig. 1. Map show ing lo ca tion of dis cov ery site of new spec i men of Sphenothallus. A. Czech Re pub lic and Bo he mian Mas sif with Cam -brian rocks of Skryje–Týøovice and Pøíbram–Jince bas ins. B. De tail of dis cov ery area show ing dis tri bu tion of mid dle Cam -brian Buchava For ma tion with Skryje–Luh lo cal ity (1) and Týøovice–“Pod hruškou” lo cal ity (2). Ge ol ogy mod i fied from Mašek et al. (1997). C. Stra tig -ra phy of Buchava For ma tion with po si tion of Skryje–Luh lo cal ity (1) and Týøovice–“Pod hruškou” lo cal ity (2). St-ra tig -ra phy af ter Fatka et al. (2011)
gue (Chlupáè, 1999, p. 11), as well as in other in sti tu tional
col lec tions, in clud ing the Schary col lec tion in the Mu seum
of Com par a tive Zo ol ogy, Har vard Uni ver sity, Boston,
U.S.A. (O.F. per sonal ob ser va tion). Chlupáè et al. (1998, p.
169) thought that the “Skryje–Luh lo cal ity” was prob a bly
dis cov ered by Krolmus as early as 1832. This as sump tion is
based on an ob ser va tion by Sklenáø (1966), who re ported on
a let ter, in which Václav Krolmus, a priest at Zvíkovec, de
-scribes a tri lo bite-bear ing site on the right bank of the
Berounka River at Skryje. Tak ing into ac count all of the char
-ac ter is tics of ex po sures of Cam brian rocks, in clud ing their
ac ces si bil ity, dis tance from roads and the abun dance of fos
-sils, it can be in ferred that the first fos sils col lected in the
Skryje–Týøovice Ba sin re gion prob a bly orig i nated from this
out crop (XY in Fig. 1B and the poly gon in Fig. 2).
Jahn (1896, fig ure on page 668) first used the des ig na
-tion “Die Schlucht oberhalb der Luher Fähre“ (the gorge
above the Luh ferry in the orig i nal Ger man de scrip tion) for
the out crop, named by all sub se quent au thors as the “Skryje
– Luh lo cal ity”. Jahn (1896, p. 727–734, fig. 7) pub lished a
de tailed de scrip tion of the sec tion on the right bank of the
Berounka River; the sec tion starts at Vosník Hill and ex
-tends from there to Šlovice vil lage.
Chlupáè et al. (1998) de scribed a sec tion ex posed in an
ex ten sive trench, ex ca vated through the cen tral part of this
out crop in 1997. Within the se quence stud ied, six in ter vals,
each de fined on the ba sis of fos sil con tent and li thol ogy,
were rec og nized.
How ever, the lack of field doc u men ta tion pre vents the
ex act de ter mi na tion of the in ter val, from which the spec i
-men of Sphenothallus sp. orig i nates.
Týøovice–“Pod hruškou” lo cal ity
The his tory of re search and a list of fos sils re cently
were sum ma rized by Fatka (2004).
As so ci ated fos sils
At the Skryje–Luh and Týøovice–“Pod hruškou” lo cal i
-ties, di verse and usu ally well-pre served fos sils have been
col lected since the first half of the nine teenth cen tury (see
Barrande, 1852; Šnajdr, 1958; Fatka, 1990; Chlupáè et al.,
1998; Chlupáè, 1999; Fatka, 2004). The fos sil as so ci a tion
col lected at these out crops in cludes com mon paradoxidid,
conocoryphid and solenopleurid trilobites, as so ci ated with
rarer agnostids (see Šnajdr, 1958; Fatka, 1990). Ar tic u lated
cinctan, eocrinoid and stylophoran echinoderms (Lefebvre
and Fatka, 2003), the usu ally small and rel a tively rare
bra-chiopods (Mergl and Šlehoferová, 1991), sim ple
ichnofo-ssils (Mikuláš, 2000), and di
verse prob
lem
at
i
cal fos
sils
have been es tab lished at sev eral strati graphi cal lev els.
Con i cal fos sils
Con i cal fos sils of vari able mor phol ogy, di men sions and
chem i cal com po si tion oc cur at nu mer ous lev els within the
Buchava For
ma
tion, in
clud
ing the Skryje–Luh and the
Týøovice–“Pod hruškou” lo cal i ties. At least ten spe cies of
heavily min er al ized con i cal fos sils have been as signed to
the class Hyolitha Marek, 1963 (Barrande, 1867; Marek,
1983; Valent, 2006; Valent et al., 2011, 2012, 2013; Valent
and Fatka, 2013, un pub lished ob ser va tions). Rarely oc cur
-ring tiny con i cal spec i mens have been clas si fied as
Volbo-rthella tenuis bohemica Prantl, 1948.
SYS TEM ATIC PALAE ON TOL OGY
Phy lum Cnidaria Hatschek, 1888
Class, Or der, Fam ily Un cer tain
Ge nus Sphenothallus Hall, 1847
Type spe cies: Sphenothallus angustifolius Hall, 1847
Dis cus sion: The ge nus is char ac ter ized by a flex i ble phos phatic or organophosphatic, slen derly con i cal (Wang et al., 2003) exoskele- ton with a smooth sur face and a rounded to subel lip ti cal trans -verse cross-sec tion. The shape of the tu bule is strongly in flu enced by the two mar ginal thick en ings, run ning from the apex to the ap er ture. The course of the thick en ings di rectly con trols the ex pan -sion an gle, be cause they are po si tioned at the ends of the long axis of the oval crosssec tion. The tu bules are usu ally com pressed, be ing ori ented by the lon ger axis of the crosssec tion par al lel to bed -ding. In this ori en ta tion, the thick en ings are po si tioned lat er ally and form the out line of the tu bule. Ad di tion ally, the thick en ings vary in ro bust ness and width, and it is some times very easy, in other cases very dif fi cult, to iden tify the thick en ings, de pend ing for ex am ple on their mode of pres er va tion, the li thol ogy of the host sed i ment and the de gree of flat ten ing of spec i mens. It is no less dif fi cult to dem on strate the phos phatic na ture of the tu bule in spec i mens, the exoskeleton of which has been dis solved. Char ac -ter is tic de for ma tion of the smooth and less rigid walls be tween the lon gi tu di nal thick en ings con firm the orig i nally flex i ble phos phatic na ture of the tu bules. Con versely, pres er va tion of an undeformed smooth sur face, lack ing the afore men tioned sec ond ary fea tures, can make it dif fi cult to de ter mine whether the spec i men orig i nally was organophosphatic. Thus, it is some times dif fi cult to iden tify the ge nus Sphenothallus un equiv o cally. That is why the au thors clas sify the spec i men show ing the di ag nos tic fea tures as Spheno-thallus, while the sec ond spec i men is as signed to the ge nus Sphe-nothallus only ten ta tively.
Sphenothallus sp.
Fig. 3A, B
Ma te rial: Sin gle, al most com pletely flat tened to low-re lief tu bule, Fig. 2. Clas sic nat u ral out crop of shales on the slope des ig nated as Skryje–Luh lo cal ity, type sec tion of Skryje Mem ber (see Fatka et al., 2011, p. 854). Por tion of out crop, from which Spheno-thallus sp. was col lected is in di cated by poly gon
pre served as an in ter nal mould in shale, stored in the Na tional Mu -seum in Prague un der the in ven tory num ber L42355.
De scrip tion: The nearly com plete spec i men is pre served in bluegreen shale and is ori ented par al lel to the bed ding. It is partly flat -tened near both the apex and the ap er ture and pre served in low con vex re lief in the mid dle part. The shell was flex i ble and orga
nophosphatic as in ferred from the typ i cal de for ma tion of the flat -tened periderm, com pa ra ble to the spec i mens with pre served walls (with striae or wrin kles of dif fer ent au thors; e.g., Neal and Hannibal, 2000; Zhu et al., 2000). The phos phatic mat ter had been re -moved, while the or ganic com po nent is pre served in the form of a thin film, bro ken into an ir reg u lar tile pat tern, con cen trated mainly along the lon gi tu di nal thick en ings, which ex hib ited the high est con cen tra tion of or ganic mat ter. The pre served part of the conch is 14.9 mm long and 1.35 mm wide at the ap er ture. The conch is slightly curved in the prox i mal por tion and straight dis tally. Its apex is bro ken and no at tach ment ap pa ra tus can be ob served. The ex pan sion an gle is very low and ranges from 3° and 4° along the mid dle and adapertural parts of the frag ment. The ap er ture, in com -pletely pre served and poorly dis cern ible, is straight and smooth with out any dis tinct struc tures. The trans verse cross-sec tion of the shell was ap par ently rounded in its outer out line. Lon gi tu di nal thick en ings are in dis tinct, be ing vis i ble in the form of flat folds at sev eral places along both mar gins. The thick en ings ap pear to ex -pand to ward the ap er ture and reach their max i mum width of 0.35– 0.45 mm in the flat tened ap er tural re gion (see Tab. 1).
Re marks: The strongly flat tened and wide thick en ings in di cate that the orig i nal skel e ton was very thin (i.e. al most flat) and wide. This com bi na tion of fea tures dif fer en ti ates it from other spe cies of Sphenothallus. This is also the case for S. kozaki, the only spe cies de scribed from the mid dle Cam brian of Bo he mia. That spe cies is char ac ter ized by a cir cu lar out line of the tu bule and poorly de fined in ter nal thick en ings. How ever, the thick en ings of Sphenothallus sp., de scribed herein, are ob vi ously wider by com par i son with the tu bule width and even thin ner. This is in di cated by their in ten sive flat ten ing. In the case of the ro bust thick en ings, they would be more dis tinct, sim i lar to the de formed spec i mens of Sphenothallus ruedemanni (Kobayashi), fig ured in cross-sec tions by Choi (1990, figs 3.3, 3.4). The typ i cal ex ter nal and in ter nal cross-sec tions and the struc ture of thick en ings seem to be im por tant di ag nos tic fea -tures. How ever, it is usu ally dif fi cult to ob serve or to re con struct this char ac ter is tic. In the case of ob ser va tions that are in suf fi cient for a re li able clas si fi ca tion, an open no men cla ture is an ap pro pri -ate tax on omy.
Dis tri bu tion: Barrandian area, Skryje–Týøovice Ba sin, Buchava For ma tion, higher strati graphi cal lev els of the Skryje Shale, Skryje–Luh lo cal ity, Paradoxides (Eccaparadoxides) pusillus Biozone (Cam brian Stage 5, Drumian Stage, cor re spond ing to the Caesaraugustian Re gional Stage of West Gond wana, see Álvaro et al., 2004; Fatka et al., 2011; Fatka and Szabad, in press).
?Sphenothallus sp.
Fig. 3C
Ma te rial: A sin gle nearly com plete, partly flat tened tu bule pre served as an in ter nal mould in green shale and de pos ited in the Na -tional Mu seum in Prague un der the in ven tory num ber L42356. De scrip tion: The pre served part of the shell mea sures 20.8 mm in length and 4.1 mm in width at the ap er tural end. It is partly pre -served as an in ter nal mould and partly as an ex ter nal mould of the op po site wall. The in ter nal mould in the ap er tural por tion is
Ta ble 1
Bio met ric data and fea tures on mea sured spec i mens of Sphenothallus sp. and ?Sphenothallus sp. All di men sions in mm
Inv. number Preservation Length of shell Apertural width Thickening max. width
Expansion angle – apical
Expansion angle – distal
Sphenothallus sp. L42355 flattened to low
relief 14.9 1.35 0.35–0.45 4.7° 3.0°
?Sphenothallus sp. L42356 relief to low relief 20.8 4.1 ~ 0.1 8.5° 10.2°
Fig. 3. Sphenothallus from Buchava For ma tion, Skryje–Týøo-vice Ba sin, Teplá–Barrandian re gion, Czech Re pub lic. A. Spheno- thallus sp., flat tened to low re lief tu bule with clearly dis cern ible lon gi tu di nal thick en ings, L42355; Skryje – Luh lo cal ity. B. draw -ing of A. C. ?Sphenothallus sp., partly flat tened spec i men, coated with li mo nite, L42356; Týøovice – “Pod hruškou” lo cal ity.De pos -ited in Na tional Mu seum in Prague un der above given num bers
slightly con vex, with a nearly flat cen tral re gion and vaulted mar -gins. The cen tral part of the shell is rep re sented by an ir reg u larly de formed, partly cracked and nearly flat to slightly con cave in ter -nal mould. The api cal re gion is pre served as a con cave ex ter -nal mould. All sur faces are thinly coated with a yel low ish ma te rial, prob a bly li mo nite. This yel low ish ma te rial forms com pact pieces of fine-grained mat ter, which partly fill cav i ties in the fos sil. Such mat ter oc curs in the de formed cen tral por tion, es pe cially at its apical and ap er tural ends. The adapertural piece, lo cated ap prox i -mately in the mid dle of the shell, is sculp tured by shal low, but sharp lon gi tu di nal wrin kles (in its ap er tural part and along the mar gin) and by fine trans verse wrin kles (in its api cal part). Near the adapical piece, there is a short, very nar row and shallow groove, re sem bling the im pres sion of a thick en ing. It ex tends adaperturally on the right mar gin of the fos sil, but only along a length of 1.5 mm, be cause it is cov ered by rock ma trix closer to the ap er ture. The in ter -nal mould in the ap er tural por tion of the shell is cov ered by di ag o nal struc tures that are ac tu ally very thin, sharp ridges. Be -cause these ridges pro ject into the cav ity of the dis solved periderm, they rep re sent in fill ings of cracks in the shell walls. It ap -pears that some frag ments were slightly dis lo cated along these frac tures. The in ter nal mould in the ap er tural por tion is also cov -ered by much less dis tinct, low, nar row, trans verse ridges. These fea tures are ir reg u larly dis trib uted with re spect to their dis tinct -ness, and range from con spic u ous to in dis tinct. The ap er ture is not pre served be cause it has been sev ered transversally. This frac ture is al most straight, bent near the right mar gin and in ter rupted by a small cutout along a short lon gi tu di nal frac ture, on the left mar -gin. The ex pan sion an gle of the tube in creases slightly from 8.5° at the api cal end to 10.2° in the cen tral(?) part of the tu bule, but then in creases mark edly to 20.8° near the ap er ture (see Tab. 1). Re marks: The spec i men is here pro vi sion ally as signed to Spheno- thallus, be cause the pres ence of a pair of lon gi tu di nal thick en ings is not cer tain. Sim i larly, the phos phatic (or chitinophosphatic) na -ture of the dis solved periderm is un cer tain, be cause of the ab sence of any typ i cal de for ma tion. On the con trary, it seems that the shell walls were brit tle. Thus, al li ance of this fos sil with known spheno-thallids is open to ques tion.
Dis tri bu tion: Barrandian area, Skryje–Týøovice Ba sin, Buchava For ma tion, higher strati graphi cal lev els of the Skryje Shale, Týøo-vice–“Pod hruškou” lo cal ity, Paradoxides (Eccaparadoxides) pu-sillus Biozone (Cam brian Stage 5, Drumian Stage, cor re spond ing to the Caesaraugustian Re gional Stage of West Gond wana).
DIS CUS SION
Plot ted on the palaeogeographic maps of Cocks and
Torsvik (2002), Fatka and Mergl (2009) and Torsvik and
Cocks (2013), Sphenothallus may be seen to have been
pres ent at low as well as higher palaeolatitudes dur ing the
Cam brian (Fig. 4). This is sim i lar to the pat tern of dis tri bu
-tion, es tab lished for the Lower Or do vi cian (Van Iten et al.,
2013, fig. 5). The ev i dently flex i ble ma te rial of the skel e ton
of Sphenothallus sp. sup ports an epifaunal mode of life,
rather than a semi-infaunal or infaunal strat egy. De spite the
ab sence of any sphenothallid-like at tach ment discs, ei ther
iso lated or at tached to in ver te brate shells, the abil ity of this
an i mal to col o nize a muddy bot tom, as hy poth e sized by
Wang et al. (2003), is plau si ble.
As in the case of the clas si cal Cam brian of the Pøíbram–
Jince Ba sin (Fatka et al., 2012), the dis cov ery of
Spheno-thallus in the Buchava For ma tion of the Skryje–Týøovice
Ba sin is sur pris ing, for only two spec i mens are known af ter
a long and in tense field in ves ti ga tion, which yielded sev eral
thou sand spec i mens of other fos sils. Al though ex tremely
rare, this dis cov ery ex pands our knowl edge of or gan isms
with organophosphatic skel e tons in the Buchava For ma tion.
Ac knowl edge ments
Both spec i mens come from the col lec tion of the late V. Kor-dule and re cently were de pos ited in the Na tional Mu seum, Prague. The au thors are grate ful to J. Hannibal (Cleve land Mu seum of Na-tural His tory, U.S.A.) for re view ing the manu script. The pa per be-nefited from the help ful re marks and com ments of Heyo Van Iten (Hanover Col lege, In di ana, U.S.A.). Our re search was sup ported fi nan cially by the Pro ject PROVOUK P44.
REF ER ENCES
Álvaro, J. J., VizcaÎno, D., Kordule, V., Fatka, O. & Pillola, G. L., 2004. Some solenopleurine trilobites from the Languedocian (Late Mid Cam brian) of West ern Eu rope. Geobios, 37: 135– 147.
Barrande, J., 1852. SystÃme Silurien du cen tre de la BohÃme. Vol. I. [Pub lished by au thor], Prague, 935 pp.
Fig. 4. Palaeo geo graphi cal dis tri bu tion of ge nus Spenothallus Hall, 1847 in lower and mid dle Cam brian lagerstätten. Cam brian palaeo ge ogra phy mod i fied af ter Cocks and Torsvik (2002), Fatka and Mergl (2009) and Torsvik and Cocks (2013)
Barrande, J., 1867. SystÃme Silurien du cen tre de la BohÃme. Vol. 3. [Pub lished by au thor], Prague, 179 pp.
Brabcová, Z. & Kraft, P., 2003. Study of conulariid and re lated phosphatic con i cal exoskeletons from the Prague Ba sin (Czech Re pub lic). In: Albanesi, G. L., Beresi, M. S. & Peralta, S. H. (eds), Or do vi cian from the An des. INSUGEO, Se -ries Correlación Geológica, 17: 263–266.
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