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Ordovician and Silurian chitinozoan biozones of western Gondwana

Article  in  Geological Magazine · July 2006

DOI: 10.1017/S001675680600207X

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Geot. Mug. 143 (4). 101M, pp . 509- 529. © 2006 Cumbridg(' University Press 50 9
dol:10. 101 7/S00 16756R0600207X Printed in Ihe United Kingdom

Ordovician and Silurian chitinozoan biozones of

western Gondwana
YNGVE G RA H N "
UERJ, Faculdade de Geologia, Bloco A - Sala 400 I, Rua Sao Francisco Xavier 524, 20550-013
Rio de Janeiro, RJ " Brazil

(Recei ved 23 November 2004; rev ised \'('1'.\' i OIl received 15 Septembe r 2005; accepted 5 December 2005)

Abstract - A formal Ordovician- Silurian chitinozoan biozonation for western Gondwana is

proposed. This palaeogeographic province includes South America, and was located in medium
to high latitudes during Ordovician and Silurian times. Ordovician chitinozoans are known from
northern Argentina. southern Bolivia. and Brazil. Silurian chitinozoans occur in Brazil. northern
Argentina. southern Bolivia and southern Peru. No published information is available about
Ordovician-Silurian chitinozoans from Ecuador, Colombia or Venezuela. Altogether more than
150 localities (including wells and outcrops) are included in this study. and 154 species have
been encountered. A biozonation based on the first occurrence of critical chitinozoan species is
introduced. Five biozones are defined in the Ordovician (zones of Desmochltina sp. gr. minor,
Conochitina decipiens, Eremochinna brevis, Lagenochiti na obelig is and Tan uchit ina anticostiensisi ,
and nine in the Silurian (zones of Beion echitina postrobusta, Spi nachitina ha rringtoni, Pogonochitina
dj almuicMu rg achitina margaritana - Salopochitin a monterrosae. Angochitina echina ta, Eisenackitina
gra nutato. Fungochitina kosovensis and the subzoncs of Sphaeru chi tina solutidina and Desmochitina
cf. D. " elisa). These biozones are compared with known graptolite. conodont, acritarch and spore
zones from the same area. and chitinozoan zones on a global basis.

Keywords: Gondwana, Ordovician. Silurian, Chitinozoa. biozonation.

I. Introduction are not easily applied on a regiona l. and even less so

Gra ptolites and/or shelly fossils from Ordovician and
Silurian strata in the intracratonic basins of western
Gond wana (Fig. I) are present in north ern Argentina,
southern Bolivia and southern Peru. In Brazil and cast-
ern Paraguay these fossils arc limited to specific levels
in the stratigraphic column, and therefore are only
of restricted biostratigraphic value. Recent investig-
ations on chitinozoans, acritarchs and spores have made
possible a more detailed dating of the Silurian rocks .
It should be noted that such an important microfossil
gro up as conodonts is, except for the Ordovician
in northern Argentina, Llandovery in southern Peru,
early Wenlock in southern Bolivia, and Caradoc strata
in eastern Paraguay (Aconttodus sp. reworked with
other shelly fossils into Aeronian Vargas Pefia shales:
unpub. data), unknown from Ordovician and Silurian
rocks of southern Bolivia, Brazil and eastern Paragu ay.
Acritarchs and spores have been applied in biostrati-
graphy only on a limited scale. However, local Silurian
chitinozoan biozonations have been suggested for the
Parnaiba (Grahn, Melo & Steernans, 2005 ) and Parana
(Grahn, Pereira & Bergamaschi, 2000; Grahn, 2005b)
basins. The latter scheme has also been applied to lower
Siluria n strata in the Tarija Basin (northern Argenti na)
by Grahn & Gutierre z (200 I). These local biozones
• E-ma il: yngvcgrahn@hotmail.com
on a global, scale. The purpose of this paper is 10
present an Ordovician-Silurian chitinozoan biozona-
tion applicable for the intracratonic basins of western
Gond wana. and to compare this biozonation with
other zonations in well-known areas on a global sca le.
2. Previou s chitinozoan studies in th e Ordovician
and Silur ian of western Gondwana
Ordovician and Silurian chitinozoans from Brazil and
Paraguay have been discussed in a series of internal
reports from PETROBRAS and UERJ (Universidade
do Estado do Rio de Janeiro) by H. Muller (Internal
report, PETROBRA S, Salvador, 1962), R. F. Daemon
(Internal report , PETRO BRAS, Salvado r, 1964), Y.
Grahn (Internal reports, PETROBRAS , Rio de Janeiro,
1988a-<1, I989a--<:, I992c-f, and Internal reports ,
UERJ, Rio de Janeiro, 1997, 1998) and Y. Grahn &
1. H. G. Melo (Intern al report, PETROBRAS, Rio de
Jane iro, 1990), and they have also been the subject in
two doctoral theses (N. M. Costa, unpub. Ph.D. thesis,
Univ. dc Sao Paulo, 1972; L. P. Quadros, unpub . Ph.D.
thesis, Univ. Federal do Rio de Janeiro, 1985) and one
Masters thesi s (P. Mendlowicz Mauller, unpub. M.Sc.
thesis, Univ. do Estado do Rio de Janeiro, 2004) . Only
in the latter thesis has a formal biozonation (sensu
Grahn, Pereira & Bergamaschi, 2000) been used. Since
the papers by Lange (1967), Somme r & van Boekel
510
-.
N
O'
t o'
20'
PaCtricoc ean
Atranuc Ocean
J(}'
o and Vila Maria (Para na Basin) sedime nts as levels with
.
o 1000 km
80' 80" 40'
Figure 1. Silurian sedimentary basins in South America. Based
on Rubinstein. 1997. Asterisk symbolizes a sub-basin.
( 1963, 1965 ), Cos ta (1970, 1971a ,b, 1974), Quad ros
(1982, 1988), Wood & Mill er (1991), Gray et al.
(1992 ), Grahn ( 1992a ,h) and Grahn & Paris ( \ 992),
new geological information has been publi shed about
Silurian Chitinozoa from Brazil and Paraguay (Grahn
2005a,b; Grahn & Melo, 200 3; Grahn, Pereira &
Bergarnaschi , 2000 ; Grahn, Loboziak & Melo, 200 I,
2003; Grahn, Melo & Steemans, 2005 ; Lc Hcri sse
et al. 200 I; Azevedo-Soares & Grahn, 2005 ). In Grahn,
Pere ira & Bergarnaschi (2000) a regional chitinozoan
bio zonation was suggested for the Silurian of eastern
Paraguay, which was also applied to the lower Silurian
in northern Argentina (Grahn & Guti erre z, 200 I), and
in Grahn (2005h) the Silurian chitin ozoans of the
Parana Basin in Brazil are compared with tho se from
eastern Paraguay. Grahn, Mclo & Steem ans (2005)
also introduced a chitinozoan biozonation for the
Siluri an Tiangua Formation in the Parnaiba Basin. A
formal biozon ation for the Solimoes and Amazonas
basins has hitherto not been propo sed . Chitinozoa ns
of Ordo vician andlor Siluri an age have also been
describ ed or discussed from southern Bolivia (Cramer,
Diez & Cuerda, 1974; Suarez-Riglos, 1975; Liachenko ,
1994; Heuse, Grahn & Erdtm ann, 1999; Melo, 2000:
Grahn, 2(02 ). southern Peru (Davi la & Ponce de Leon,
197\ ) and northern Arge ntina (Volkheimer, Mele ndi &
Acesolaza, 1980; Ottone, Toro & Waisfeld, 1992 ;
Grahn & Gutierrez, 200 I: Grahn. 20(3).
Y. G RA HN
3. Notes on the Ordovician and Silurian
palaeogeography and palaeoenvironment of
western Gondwana
South Ame rica was a part of Go ndwana and located
in medium to high latitu des durin g Ordovician and
Silurian times. The Late Ordovician (Hirna ntian)
glacia tion was centred in North Africa, and can be
traced in coeval sediments as ice-striated quartz grains
in the continental (glacial outwash) sedimentation of
the Amazona s and Parn aiba basins (Caputo, 1998).
No glaciation took place in the area despit e a geogra-
phic position near the South Pole (Grahn & Caputo,
1992 ). The Hirn antian glacia tion can also be traced
in the Parana Basin in connection with a glacial
centre in the Cape Basin (Grahn & Caputo, 1992).
Dur ing Lland overy times, three glacial events (upper
Rhudd ani an-I ower Aeronian, uppermost Aeroni an-
lower Telychi an and upper Telychian-Iower Shein-
woodian) took place, whic h are reflected in the
Nhamunda (Amazonas Basin), Ipu (Parn aiba Basin )
diam ictites (Grahn & Caputo, 1992 ). During deglaci-
ation periods the intracratonic basins were in con tact
with the Iapetu s Ocean . The diamic tite levels can also
be found in eastern Paragu ay (Euse bio Ayala Forma-
tion), north ern Argentina (Zapla Formation), southern
Bolivia (Can cafiiri Formation) and southern Peru (Sa n
Gaban Form ation ). The geog raphic South Pole was
situa ted in ce ntral Braz il in Early Silurian times. Durin g
the Late Silurian, western Gondwa na was still situated
in the polar region, and the clim ate continued to be
cool, as indicated by the palaeogeographi c position,
lack of carbonate sediments and the pre senc e of or-
ganisms with an Afro-S outh Am erican/Malvinokaffri c
affinit y.
4. Chitinozoan hiostratigraphy
During the study I felt obliged to redeterm ine many of
the previous chitinozoandeterminations. The new exact
names arc shown in App end ix I. A list of chitinozoan
spec ies reported from western Go ndwa na and cited
in text and figures is presented in App endi x 2, and
a list of chitinozoa n spec ies not reported from western
Gondwa na but cited in the text is shown in App en-
dix 3.
a.a. Southern Peru
Davila & Ponce de Leon ( 197 1) menti oned a find of
Rhabdochitina sp. (possibly Conochitina proboscifenn
from the San Gaban Formation in the Punco Punco
sect ion northeast of Ayaviri. If correct, it indicates
a correlation of the San Gaban Formation with the
Ca ncafiiri and Zapla form ations in southern Bolivia
and north ern Argen tina , respectivel y (Diaz-Martinez
et al. 200 I) .
Ordo vician and Silurian chitinozoan biozones
-4.b. So uthern Bolh-ia
Ordovician and Silurian chitinozoans from 15 localities
in the East Precordillera and Subandean Belt of Bolivia
have been described in the literature. From the East
Precordillera north and nonheast ofCuipina, and at Ce-
mentario and Estancia Savinga Mayu, Hcuse, Grahn &
Erdtmann (1999) described upper Tremadocian
chitinozoa ns iDesmochitina sp. gr. minors from the
upper Cienguillas and Obispo formations, and lower
Arenigian species iCono chitina decipiens, Cono chitina
cf. e. havliceki and Conochitina sp.) from the base of
the Agua y Toro Formation. No chitinozoans have been
reported from younger Ordovicia n beds in Bolivia.
Ncar the pass of La Cumbre, about 20 km northeast of
La Paz, the Caneaiiiri Formation yields Belonechitina
sp.. Conoc hitina sp. and C yathochitina sp. 13, indicating
an Early Silurian age (Diaz-Maninez & Grahn,
2004) . Late Silurian (Ludlow- Pridoli) chitinozoan
assemblages arc known from the Subandean Belt
(Peru- Bolivia Basin), where they have been describ ed
from the Kirusillas Format ion (Cramer, Diez & Cuerda,
1974) at Cochabamba (Appendix I). Grahn (2002)
described chitinozoans from the Kirusillas and Tar-
abuco formations in the Rio Pilaya, Sella, Sobo-Sobo
and Tarabuco sections. Liachenko ( 1994) discussed
Upper Silurian and Lower Lochkovian chitin ozoans
from three sections in the Kirusillas and Tarabuco
formations along Rio Chimore. However, the quality of
her illustrations docs not allow reliable deter minations
(Appen dix I) . The chit inozoans have a Gondwanan
affinity in the Silurian.
-t,c. Northern Argentina
Ordovician and Silurian chitinozoans from eight local-
ities in the Tarija (=Peru- Bolivia Basin) and Chaco-
Parana basins arc known from the literature. From
Cajas and Los Colorados in the Tarija Basin, Ottone,
Toro & Waisfeld ( 1992) described an assemblage in
the lower and upper part of the Acoite Forma tion
of early-middle Arenigian age (Appendix I). Se-
quences at Arroyo Colorado (Cachipunco Format ion),
Arroyo Matos (Zap la and Lipeon format ions), Rio
Capi llas (Zap la and Lipeon format ions), and 9 de
Oetubre Mine (Lipeon Formation) yielded middle
Llandovery-early Wenlock chitinozoans ; at Angosto
Los Pereyras (upper Caehipunco Formatio n) occur
Ludlow-Pridoli chit inozoans (Grahn & Gutierrez,
200 1), and one core (EI Cabure- I) in the Chaco-Parana
Basin yielde d Late Silurian chitinozoans from the Copo
and Cabure formation s (Grahn, 2003) . Volkheimer,
Melendi & Aceso laza ( 1980) mentioned the finds of
chitinozoan fragme nts in the Mojotoro Formation of
probable Early Ordovician age . The Arenigian chitino-
zoans have a mixed Gondwana- Laurentian affinity,
while the Silurian assemblages display a Gondwanan
affinity.
5 11
-t.d. Eas tern Paraguay
Silurian chitinozoans are known from nine localitie s in
eastern Paraguay, all of them in the Itacurubi Group .
Grahn, Pereira & Bergamaschi (2000) described
chit inozoan assemb lages from the Shell-Pecten Asun-
cion I well and the Anschutz Minerals RD 11 5 and RD
116 wells, and the Minas Cue outcro p was described by
Grahn (2005b) . Two shallow borings (269 -R3 and 269-
R4) yielding Cyat hoch itinu sp, 13, Cing ulochitina cf.
e. serra ta (=e. aff. e. serratu s, Ancyro chitina sp. and
Angochitina sp. were mentioned by Gray et 01. ( 1992).
Two additiona l shallow borings (269 -R I and 269-R2),
yielding essentially the same fauna as described by
Grahn, Pereira & Bergamaschi (2000), were discus sed
by P. Mendlowicz Mauller (unpub. M.Sc. thesis, Univ.
do Estado do Rio de Jane iro, 2004) . Wood & Miller
(1991) described a chitin ozoan fauna (Plec tochitina
paraguayensis, Eisenackitina bej ui and Pterochitina
n.sp. = Ptero chitina deichaiii from the type localit y of
the Vargas Pefi a Formation. An additional assemblage
was discussed by Wood & Miller ( 1997).
-t.e. Brazil
Ordovician and Silurian chitinozoa ns from Brazil arc
well known and described from the Solimoes, Amazo-
nas, Parnaiba and Parana basins. Published infor mation
is available from more than one hundred localities. In
the Solimoes Basin, Ordovician chitinozoan s from the
Benjamin Constant Formation have been repon ed by
Grahn (l992h) in three deep drillings (PETROBRAS
2-BT- I-AM , I-JD- I-AM and I-JT-2-AM wells ), and
Silur ian chitinozoans from the lower Jutai Form ation
in PETROBRAS I-JD-I -AM well reponed by Grahn ,
Loboziak & Melo (2003 ). The Ordovician-Silurian
chitinozoan assemblages are well doc umente d from the
Amazo nas Basin. An Ashgill (Rawtheyan) fauna was
described from the upper Autas-M irim Formation in
PETROBRAS I-AM-I -AM well by Grahn (I 9920,h).
Silurian chitinozoans illustrated from nearly 70 local-
ities in the Pitinga and lower part of the Manacapuru
form ations by Lange ( 1967) have subsequently been
described by Grahn & Melo (2003), and in 33 additional
localities by Grahn (2005h) and Azevedo-Soares &
Grahn (2005) . Sommer & van Boekel (1963, 1965)
described chitinozoans from the shallow well 130m
Jard im 56 at Itaituba (Append ix I). Costa ( 1970,
19710) described chitinozoans from Igarape da Rainha
(sample 4-65-68 of Grahn, 20050), a tributary o f Rio
Tapajos (Appendix I). Costa (1974) investigated Early
Silur ian chitinozoans from Madame Island below the
Viramundo waterfall. Chitinozoans from this loca lity,
designated as AM 76 by PETROBRAS, were also
described by Grahn (20050) . The Silur ian chitinozoan s
of the Trombetas Group (Pitinga and lower part of
the Manacapuru formations) were also discussed by
Grahn & Paris (1992 ). In the Parnaiba Basin, Silurian
chitinozoans are known from 13 localities in the upper
512
lpu, Tiangua and lower part o f the Jaicos form ations
(Grahn, Lobo ziak & Melo, 200 I). Quadro s ( 198 2)
discussed Silurian Chitinozoa (Appendix I) from three
of the se localiti es (PET ROBRAS I-MA - I-PI, 2-SL -
I-MA and 2-PM -I-MA wells). Add itiona l species
were described by Grahn in Le Heris sc et al. (2001 ),
and commented upon by Grahn , Loboziak & Melo
(200 1). A complete list of Silu rian chitinozoan s from
the Parnaiba Basin is published by Grahn, Melo &
Steemans (2005) . Silurian chitinozoa ns from the
Parana Basin are restricted to the shales of the Vila
Maria Formatio n, and only known from one local ity
for certain (Grahn, 1992a, 2005b ). The chitinozoan
fauna was desc ribed by Grahn, Pereira & Bergamaschi
(2000) and Grahn (2005b).
5. Ordovician and Silurian chltinozoan zones
of western Gondwana
It should be noted that no publi shed information
abou t O rdovician and Silurian Chitinozoa is available
from Ecuador, Colombia and Venezuel a. The proposed
biozo nes arc in gene ral interval range zones defined
from the first occ urre nce biohorizon (FOB) ofthe index
species to the first occ urrence of the index species
in the following zone. Altoge ther, 154 species have
been encoun tered, of which 16 are in the Ordovician
sequences. No global chi tinozoan biozona tion has been
suggested for the Ordovician, but good biostratigraphic
schemes exist for the Welsh Borderland (Jenk ins,
1967) , North Gondwana (Par is, 1990), Quebec and
western ewfoundland (Achab, 1989), Baltoscandia
(NOlvak & Grahn, 1993; Nolvak, 1999), Brabant
Massif (Samuelsson & Verniers, 2000) and mid -
continental U.S.A. (Grahn & Bergstrom, 1984). Some
regional bios tratigraphic schemes have been publ ished
for the Silurian, such as Baltoscandia (Laufeld, 1974;
Nesto r, 1994 ), Ukra ine (Laufe ld, 197 1; Nestor, 1994;
Paris & Grahn, 1996), Brabant Massif(Verniers, 1982),
North Africa (Jardin e & Yapaudjian, 1968), Bohemia
( Dufka, 1992; Kriz et al. 1986), Spain (Sc hwe ineberg,
1987). Saudi-Arabia (Pari s et al. 1995; AI-Hajri &
Paris, 1998), Quebec (Asse lin, Achab & Bourque,
1989; Soufiane & Ach ab, 2000), China (Geng el al.
1997), Welsh Borderland (Sutherland, 1994; Verniers,
1999; Mullins & Loydell , 2001) and mid -continental
U.S.A. (Grahn. 1985; Grahn & Bergstrom, 1985 ). A
glob al chitinozoan biozonation for the Siluri an was
proposed by Verniers et al. ( 1995).
6. Ordovician chitinozoan biozonation
of western Gondw ana
The Ordovicia n biozonation and its correlation within
western Go ndwana arc shown in Figure 2. A correl-
ation with the global zonation is demonstrated in Fig-
ure 3.
Y. G RA HN
6.a. Desmochitin a sp_2r. mitior Range Zon e
Zone base definiti on: FOB of Desmochitina sp. gr.
minor (Fig. 4a ).
Characterization o f zone: No other chitinozoan
species are present.
Reference section for zone base : In the Dique section
near Culpina in southern Bo livia, the zone base is
defined in the top of Cie neguillas Formation, ~ 5 m
below the base of the overlying Obispo Formation.
Representati ve occ urre nces of zone : Obispo Form -
ation, Sarna Anticline (Bolivia) and top of the
Cienegui llas Forma tion and the base of the Obispo
Formation, area ncar Culpina (Bo livia) .
Litho stratigraphic distribution: Top of Cieneg uillas
Format ion and Obispo Formation in Bolivia.
Age range : Late Tremadocian-earliest Arenigian
(Aorograptus victoriae--Tetragraptus phyllograpt oides
graptolite zones) (Figs 2, 3).
Rem arks : Desm och itina sp. gr. minor was first
described from the conglomerat ic series at M iedzygorz
in the Holy Cro ss Moun tain s (Poland) by Szani awski
in Chlebows ki & Szaniawski (1974). It has also been
described from the Leetse and lower Toila form-
ations in the Suhkrumag i section at Tallinn , Estonia
(Grahn, 1984 ). All these occurrences are from the
lower Arenigia n iCyath ochi tina primitive Zone) in
Balto scandia (Nolvak & Gra hn, 1993).
6. b. Conoc hitina decipiens Int erval Zone
Zon e base definiti on : FOB of Conoch itina decip iens
(Fig. 5a).
Charac teriza tion of zone: The co-occ urrence of
C. decipiens and Conochitina cf. C. hu vliceki.
Reference section for zone base: The west flank
of the Sarna Ant icline at Estancia Savinga Mayo in
southern Bolivia. The base is situated about 50 m above
the base of the Agua y Toro Formation.
Representative occ urrences of zone: Agua y Toro
Formation, Sarna Anticline, Bolivi a.
Lithos tratig raphic distribution: Ag ua y Toro Forma-
tion in Bolivia.
Age range: Early Arenig ian (Expansograptus
protobalticus-Baltograptus minutus graptolite zones)
(Figs 2, 3).
Remarks: Conochitina decip iens range from lower
to uppermost Areni gian . Only the lower part of its
stratigraphic rang e is repre sented in this zone.
6.c . Eremocbitina brevis Ran ge Zon e
Zone base definition: FO B of Eremo chitina brevis
(Fig. 4b) .
Characterization of zone: The co-occurre nce of
Eremochitina brevis and Conoc hiti na ordinaria.
Reference section for zone base: Section at Los
Co lorados Creek , abo ut 30 km west of Tilcara in
the Tumbaya Department, Jujuy Province, northwes t
Ordovician and Si lurian chitinozo an biozon es 513

Uthostratigraphy
Cluono· Conodonl Graptolite Chitinozoan
stratigraphy Sou1hem
Bolivia
Northern
Argentina
Braz ~
SolimOeo "'"".
Amazona'
zones
1
zone,
2
zone,
This paper
Chitinozoan ranges in the
Ordovician of western Gondwana

lJando,e')' Nhamunda
(Ahuddanian) Fm.
f----

Ashgill
. T.anticostiensis
~ § 1111
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uppe,

•
Benjamin
. naVIs
u. de ntatus
8 ustrodenla tu

I. victoriae
A. laoWOlfhi
,
L. ob6ligis

E. brevis
~
~~linnnt II
Arenig Fm. Constant Fm.
"".
AQua V t ore
Fm. . lower
E. hOlm;
E pIO/QbIJlItws
Conochitlna
decipians ~
E

Tphy/IogrlIptoides

ObispoFm.'" Proteus! H. copiosus Desmochitina

de/tatus A. murray; sp. gf. minor
A. victoriae
Tremadoc
Cieneguillas
• deltiler
Aoelograptvs
,
Fm.
Iscayachi Fm. . "'"'
R.
f1abelliformis

Figure 2. Correlation between Ordovician graptolite , co nodo nt. and chitinozoan biozones in we stern Gondwana. I - Albanesi &
Ortega. 2002. 2 - Albanesi & Onega, 200 2; Egcnhoff Maletz & Erdtmann, 2004 . Asterisk symbo lizes occ urrence o f chitinozoans.

Argentina . The base is situated about 500 m above the
base of the Acoite Formation.
Representative occurrences of zone: Sec reference
section for the base.
Lithostratigraphic distribution: Lower part of the
Acoite Formation in Argentina.
Age range: Middle Arenigian (Azygograp tus
lap worthi-Iower lsograp tus victor iae graptolite zones)
(Figs 2, 3).
Remarks: Eremochitina brevis is restricted to middle
Arenig strata.
The biozone was originally defined from the subsur-
face ofwestern Sahara by Paris ( 1990), where the index
species occurs together with L. esthonica, L. obeligis
and Velatachitina veligera in the upper part of the
E. brevis Zone (Appendix 3).
6.d. Lagenochitina obeligis Interv al Zon e
Zone base definition: FOB of Lagenochitina obeligis
(Fig. 5b).
Characterization of zone: The co-occurrence of
Lagenochitina obeligis and Conochitina brevis . Other
species present are Conochitina pe rvulgatal, C. micra-
cantha, C. sp. A sensu Grahn, 1992b and C. sp. sensu
Heuse, Grahn & Erdtmann , 1999.
Reference section for zone base: PETROBRAS 1-
JD- I-AM well at level 2594 m, which is 31 m above
the base of the Benjamin Constant Formation and the
pre-Cambrian basement of the Solimoes Basin. No
outcrop sections of the Benjamin Constant Formatio n
are known.
Representative occurrences of zone: Upper part
of the Acoite Formation, at Los Colorados Creek
in northwest Argentina (see Section 6.c) and the
Benjamin Constant Formation in the Solimoes
Basin (PET ROBRAS I-JD·I·AM and 2-BT- I-AM
wells).
Lithostratigraphic distribut ion: Benjamin Constant
Formation in Brazil and upper part of the Acoite
Forma tion in northern Argentina.
Age range: Late Arenigian (upper Isograp tus
victoriae-Undulograp tus dentatus graptolite zones)
(Figs 2, 3).
Remarks: Lagenochitina obeligis ranges from
middle Arenig to lower Llanvirn. Based on acritarchs
(e.g. Arkon ia virga ta), Quadros (1988) dated the
Benjamin Constant Formation as Arenigian/Llanvirn.
5 14 Y. GRA HN

Northern Western
Laurentia Avalonia Gondwana Baltica Saudi-Arabia Gondwana
Chrono- Quebec and Oklahoma Brabant N. Africa Baltoscandia Paris,
stratigraphy western New- ~en kins, 1969 , Shropshire Massif SW Europe. Niil vak & Verniers
foundland 1970 Grahn Jenkins,1967 Samuelsson l:! Paris,1990, Grahn, 1993 This paper
& AI-Hajri,
Achab,1989 & Miller,1986 Verniers,200C • 2004 Nii lvak,1999 2000
'au u a U' lB. ct. ga1~achian A. oulebsiri lscsbt'atfatJgO(JrrJe8U BeJonechitirvJsp. 1
B amaChlsna T el a ra amach,ana
$ . lepta S. taugourd8au;
T. anticostiensis 10
Ancyrochitina Conochitina Ancyroctlitina r anticostienSiS
Ashgill ""ga ruga'a merga
C. vaureafensis P. sytvanica
A meroa T. bergstroemi A. nigenca A. nigerica
C. ssnta T. bergstroemi

H. nyalOphrys
A . oornlcU/ans
B. seriespinosa
9
Aca~:;:C;;:lfJa ,
C. ·~ 2 B. micracantha A. reticulifera r lis tuiosa 1:frstulosa
Ancyrochitina
C. pygmaeatH.
B. robus ta onniensis
f-- B Df:1()f1e:vlltma F. fungiformis

~ I-- L baltica
H. crickmayi Ancyrochitina robusta 3
F= 7
Caradoc
i
grscquilmu!tlspmst
liCatvS!(JrimitiV8
,nach,tma S .A
S. comoectitis alatlcornis
L. dalbyensis 6
51 bulmanlS
nvi/I n i
S. cervicornis 4

e.hirsutaILageno H. bromidensis
Lagenochitina
~~
- L. deunffi
L. stentor 9
chitina sp.A m ras mosa
L. plssotensis 1 L. pissotansis
Rhabdochitina L. cJavata 11 L clavata
usiata L striata
A. armoric<!Jna
1
Llanvirn C. jenkinsi C. cafix
4 C. jenkinsi

c.wrpna Siphonochitina Cyathochitina

$ . formosa
C Iom>osa ragnalll
S. tenuicollis - C. calix -

B. srum C rotoca/;x
D. ornensis 3 L. obe/igis
C. d . dispar 2 = g ~~ C.cucumis
C. tangel D, ornensis
Arenig C. brevis E. brevis 1 . brevis E. brevis
L.esmomca CyathOChitina
C. fa mondi E. baculata Conochitme
primitive
deciplans
C. symmetrica C. symmetr/ca
Lagenochitina
L. ron/fundus esthonica
L destombesi Desmochitina
Lagenochitina sp.gr. mmor
destombesi
Tremadoc

Figure 3. Correlation between Ordovician chitinozoan biozones in western Gondwana and other areas on a global scale. · 2004 -
Webby et al. (2004 ).

6.e. Tanuchitino antieostiensis In terval Zone In this interval, many different Herc ochitina species
occurtogether with Con ochitina COllc a va . Conochitina
Zone base definition: FOB of Tanuchitina anticostien -
armifera and Ancyro chitina longispina (Appendix 3)
sis (Fig. 5c).
(Acha b, 1989).
Characte rization of zone: The co-occurrence of the
index species, Armo ricochitina nigerica, Lagenochit-
ina prussica and Rhabdochitin a aff gracilis. 7. Silurian chitinozoan biozonation
Reference section for zone base: PETROBRAS of western Gondwana
l -AM-l -AM well level 2091 m, Amazonas Basin,
northern Brazil. No outcrop sections of the Auras- The Silurian biozonation and its correlation within
Mirim Formation are known. western Gondwana are shown in Figures 6-8 and 11.
Representative occurrences of zone: Upper part of A correlation with the global zonation is demonstrated
the Autas-Mirirn Formatio n. in Figures 9 and 10.
Lithostratigraphic distribution: Upper part of the
Autas-Mirim Formation in northern Brazil.
7.a. Belonechitina postrobus ta Range Zone
Age range: Upper Ashgill (Figs 2. 3).
Remarks: Tanuchitina anticostiensis is restricted to Zone base definition: FOB of Belon echitina p ostro-
upper Ashgillian (Rawtheyan) strata. busta (Fig. 5d) has not been defined in west-
In Quebec , Tanuchitina anticostiensis and Her- ern Gondwana. The last occurre nce is in the top
cochitina crickmayi zones roughly correspond to Rhuddanian strata in eastern Paraguay (Grahn, Pereira
the T. anticostiensis Zone in western Gondwana. & Bergamaschi, 2000).
Ordovician and Silurian chitinozoan biozones
b Age range: M iddle Lland overy (Aeronia n) (F igs 6.
Figure a . Ordovic ian chitinozoan index species from western
Gondwana. The scale bars represent 100 J.1m. ca) Desmocbitinu
sp. gr. minor [Szaniawski ill Chlebowski & Szaniawski.
19 7~) . Dique Sect ion ncar Culpina. base of the Obispo
Formation. By courtesy of OrThomas Hcuse (lena. Germany).
(b) Ervmocbi tina bsvvis Benoit & Taugourdeau. 1961. Los
Colorado Creek. lower partof the Acoite Formation. By courtesy
of Or Eduardo Guil lermo Ottonc (Buenos Aires. Argentina).
Characterizat ion o f zone: In the uppermost part ,
Spinachitina wolfarti. Sphaerochitina palestinaense
and Ancyrochiti na llncyn'u appear.
Reference section for zone base: No section known,
Representative occurrences of zone: Anschutz Min-
erals RD-1 16 well. lowermost part of the Vargas Pefia
Frn", Parana Basin. eas tern Paraguay, The upper part
o f the B. !H,strobustu Zone is also present in the
ELET RONO RTE shallow bo reho le SM 10 17, lower-
most part of the Pitinga Form at ion . Amazonas Basin.
northern Brazil (Gra hn. 2005u ).
Lith ostrat igraph ic distribution : Sec repre sent ative
occurrences o f zon e above.
Age range: Early Llandovery (Rhuddanian) (Figs 6,
8- 11).
Remarks: Belonech itina postro bnsta is restricted to
lower Lland overy (Rhuddanian) strata.
In the g loba l zonalion (Vernie rs et al. 1995 )
the Rhuddan ian includes the Spinachitina frugili...
Belonechitina postrobusta. Conochitina electa and
lowermost Spin achitina 11I11e1l1rili zones. Other im-
portant species in this interval include Ancyrochit-
ina laevaensis, Belonechitina asperu, Plectochitina
nodi/era. Angochitina seuratt, Enconochitina iklaensis,
Plectochitina paraguayensis and Pterochitina deichaii
(A ppendix 3) .
7.h. Sp inQchitinQ hQrrinx toni RanJ,:t' Zone
Zo ne base de finition : FO B of Spinachit ina harringt oni
(Fig. 5e).
Characterization o f zo ne: In the up pe r part is a
subzone with Sphuerochitina solutidina defined (see
5 15
Section 7.c ). Besides the index spec ies are other
diagnostic species Plectochitina sp. A. Spha erochltina
silurica, Spina chitina sp. A. Cono chitinu sp. A !ie/l.m
Gra hn i ll Gra hn. Pereira & Bergam asch i. 2000,
Plectochitina sp. C se' '''', Grahn ill Gra hn. Pereira &
Bergarnaschi, 2000 and Sphaerochuina sp. C .'Ot' /I .\"II
Grahn ill Gra hn, Pereira & Bcrgam asch i, 2000 .
Reference section for zone base: Outcrop Minas
Cue (G rahn . 2005h ). lower part of the Vargas Pena
Formation. Parana Basin. eastern Paraguay,
Rep resentative occurrences of zone: Lower part of
the Vargas Pefia Formation in the Parana Basin. eastern
Paraguay.
Lithostrati graph ic d istribut ion : See rep resent at ive
occurrcncies of zone.
8- 11 ).
Remarks: Spina chitina harringtoni is restricted to
middle Lland overy (Aeronian) strata.
In the globa l zonatio n (Verniers et al. 1995) the
Aeronian includes the Spinachitina maennili and
Conochitina ulargadu zones. Other important spe-
cies in this interval include Ancyrochitina com'exa.
Conochitina vitrea and in the uppermost part (of
the C. alurgada Zone). Plectochitina saharicu and
Plectochitina p.\'c!UdOllgg/W;JltllJs (Appendix 3 ).
7.f:. Spl,"~rtIch itin" ,w /Iltidill" Ran.:,e Su hze ne
Zonebase defi nition: FOB of Spha erochitina solu tidinu
(Fig. 5f).
Charac terization of zone : Besides the index species
are ocherdiagnostic spec ies such as Conochitina sp. A
sellSIl Gra hn ill Gra hn, Pereira & Berg amaschi. 2000.
Plectochitina sp. C sensu Grah n ill Grahn. Pereira &
Bergamasch i, 2000 and Sphuerochitina sp , C ..emil
Gra hn i ll Grahn . Pereira & Bcrgamasch i, 2000.
Refer ence section for zone base : An schutz Minerals
RD-11 6 well. level 300 m, lower part o f the Varg as
Pefia Formation. Parana Basin. eastern Paraguay.
Representative occurrences of zone: Lower part of
the Vargas Pena Formation in the Parana Basin. castern
Paraguay.
Lithostra tigraphic distribution : See represent at ive
occurrences of zone above.
Age ra nge : M iddle Lland ove ry (Aero nia n ) (F igs 6,
8- 11 ).
Remarks: Sphaerochitina solutidina is restricted to
upper midd le Lland ove ry (upper Aeron ian) strata,
In the global zonation (Verniers et ul. 1995 )
this subzone corresponds to the upper Conochitina
alargada zone. Ot her importa nt species in this inte rval
arc Plectochitina saharica and Plectochitina pseudo-
agglutinans.
? d. PfJ/:on fJc!litin a dj a/m ai Int erv al Zon e
Zo ne base definition: FO B o f Pogolloclrililla djulmai
(Fig. 5g ).
5 16 Y. G RA H N

Figure 5. Ordovic ian and Silurian ch itinozoa n index species from western Gon dwana . The scale bars re present I DO urn. (a l COIlOC!litiJW
decipiens Taugo urdcau & Jek hows ky, 19M), PET ROB RAS I-JD- l · Al\t well level 2590 m. Benjamin Constant Fill. (b) t.agenochitina
ohdigis Par is. \9 XI. PET RO BRA S I·J D- I· A~1 well. core 18 (l096 m). Benja min Co nstant Fm . (e ) Tanucttitina allticostiellsi.'i(A chab.
1( 77) . PET ROB RAS l -AM- I-At'\'l well. level 209 1 m. Auta s-Miri rn Fm . (d) Belonechitina postrobusta (Nestor. 1( 80 ). Ansch utz
Mineral s RD· 116 wel l. level 400.95 111. Vargas Pcf m Fm. (e ) Spinachitina harringtoni Gra hn ill Gra hn. Pereira & Bcrgam aschi. 2000.
An sch utz Minera ls RD-115 well. level 14 1.77 m. Varga s Pena Fm. (f) Sphaerochitinu soluti dina Paris. 1988. Anschutz Minerals
RD-116 well. level 280 .70 m. Vargas Pcna Fill. (g) Pogonochitina dj ahnoi (Sommer & van Boekcl . 1( 65 ). Outcro p lgarape da Rainha
4-65-68. Pitinga Frn. (lower par t). (h) Dt·smochitina cf D. densu Eisc nnck. 1962 . Ansch utz Minerals RD·11 6 well. level 222 .60 ru,
Vargas Pefta Fill. (i ) Marg achitina margaritana tEiscnack. 1( 37). PET ROBRAS I-AM -I -AM wel l. cuttings 1746 -1 749 In. Pitinga
Ordovician and Silurian chitinozoan biozones 517

Chitinozoan ranges in the Silurian of western Gondwana

f----------------------------------1 oi j ~
~ ~
.s
.~
0
N
s:
o
f--

o
"0
.;::
ll..

5 .'
2'·3
1

Figure 6. Chronostratigraphy and chitinozoan ranges in the Early Silurian of western Gondwana. Chitinozoan Zone 1 - Belonechitina
postrabusta , 2 - Spinachitina harringtoni, 3 - Sphaerochitina solutidina, 4 - Desmochitina cf. D. densa, 5 - Pogonochitina djalma i,
and 6 - Margachitina margaritana-Saiopochitina monterrosae.

Characterization of zone: In the upper part is a Lithostratigraphic distribution: See representative

subzone with Desmochitina cf. D. densa defined
(see Section 7.e). Besides the index species are
other diagnostic species such as Angochitina gurupi-
ense , Cing ulochit ina bouinensis, Eisena ckitina bejui,
Plecto chitina pa raguayensis, Ancyrochitina erichseni,
Allgochiti lla longicollis, Pogonochitina inornata and
Pogonochitina tiunguaense.
Reference section for zone base: PETROBRAS
l- AM-I -AM well, level 1755 m, Amazonas Basin,
northern Brazil. No outcrop section with the base of
the P. dj almai Zone is known.
Representative occurrences of zone: Several local-
ities within the lower part of the Pitinga Formation in
the Amazonas Basin and the Tiangua Formation in the
Parnaiba Basin, north ern Brazil.
occurrences of zone above.
Age range: Late middle Llandovery (Acronian) to
late Llandovery (Telychian) (Monograptus turricu-
latus grapto lite Zone in the lower part) (Figs 6,
8- 11).
Remarks: Pogonochitina dja lmai range from upper
middle Llandovery (uppermost Acronia n) to lower
Wenlock (lower Sheinwoodian ) strata.
Pogonochitina dja lmai occur together with Tclychian
graptolites in the Bove Basin (Guinea) (Paris in
Villeneuve et af. 1989). In the globa l zonation (Verniers
er al. 1995) this zone can be correlated with the
Zone of Eisena ckitina dolioliformis. Other important
species in this interval include Pterochitina deichoii.
Conochitina vitrea, Plectochit ino pseudoagglutinans,

Fm. (low er part). (j) Salopochltina monterrosae (Cramer, 1969). PETROB RAS t· AM- l -AM we ll, cuttings 1746--1749 m, Pitinga
Fm. (lower part). (k) Ango chitin a echinate Eiscnack, 193 1. PETROBRAS I·AM· I-AM well , core 46 (1607.00-16 13 m) , Pitinga Fm.
(upper part). (I) Eisenackitina g ranulata (Cramer, 1964 ). PETROBRAS I-AM - I-AM well , core 46 (1607.00- 1613 m) , Pitinga Fm .
(upper part). (m) Fungochitina kosovensis Paris, 1981. PETROBRAS I-A M-l ~ A M well, core 46 ( 1607.00--16 13 m). Pitinga
Fm. (upp er part) .
5 18 Y. G RA HN

Chitinozoan ranges in the Silurian of western Gondwana

"'"No>
0 ",

50 "0
~ N
s:
o

... .... ....'. '.'.'.'. JJIJJIJJr.IJJIJJIJ.r.l. 9

Figure 7. Chronostrat igraphy and chitin ozoan ranges in the Late Silurian of western Gondwana. Chitinozoan Zone 6 - Angochitina
ecbinata , 7 - Eisenuckitin a grunulutu, and 8 - Fungochit ina kosovens is. T & J 1960- Taugou rdeau & Jekhowsky (1960).

Plectochitina saharica, Conochitina edje lensis and
Conochitina emmastensis (Appendix 3).
7.e. Desmochitina d. D. densa Range Subzo ne
Zone base de finition: FOB of Desmochitina cf. D.
densa (Fig. 5h).
Characteriza tion o f zone: Besides the index species,
other diagnostic spec ies are Pog onochitina inorna ta.
Ancyrochitina paranaensis and Linochit ina sp. A sensu
Grahn in Grahn, Pereira & Bergamaschi, 2000, have
their first appearance in this subzone, and Conoeltitina
elongata disap pears in its topmost pan .
Reference section for zonc base: Anschu tz Minerals
RD-116 well, level 230 m, upper pan of thc Vargas
Pena Format ion, Parana Basin , eastern Paraguay, No
outcrop section with the base of the Desmochitina ef.
D. dens a Subzo ne is known.
Repre sentative occurrences of zone: Upper Vargas
Pcna Formation in the Parana Basin, eastern Paraguay,
Lithostratigraph ic distribution: See repr esentative
occ urrences of zone.
Age range: Early late Llandovery (early Telychian )
(Monogmp tus turriculatus graptolite Zone in the lower
pan ) (Figs 6, 8- 11).
Remarks: Desmochitina cf. D. densa is restricted to
lower upper Llandovery (lower Telychian) strata.
In the global zonation (Verniers et al. (995) the
Desmochitina cf. D. densa subzone is coeva l with
the upper Eisena ckitina dolioliformis Zone. See above
for characteristic chitinozoan species in this zone (sec
Section 7.d).
7.r. .\ /arxachir;na margari/ana-Salopochitina mon terrose e
Ran ge Zone
Zone base definition: FO B o f Margachitina margar-
itana (Fig. 5i) or Salopoelritina monterrosae (Fig . 5j ).
Characterization of zone: In addition to the index
species, other diagnostic species are Angochit ina
sp. A sensu Gra hn & Pari s, 1992, Belonechitino'l
Ordovicion and Silurian chiti nozoan biozones 5 19

Lithostratigraphy
Chrono- Brazil Paraguay Arge ntina Boliv ia Peru Ch itinozoan
strat igra phy zones
Chaco-
SolimOes Amazonas Parnaiba Paran a Parana Tarija
Parana Peru-Bolivia Basin
Basin Basin Basin Basin Basin Basin
Basin

?
(5 Cabure Anoyo Tarabuco

.
"0
;:: Furnas Fm .
Fm.
•
Colorado Fm. Fm.
•
C- . . .. . .
J utal Fm. *
Lower
....Mana cap uru
... ?~
- - - -\ --- Fungochitina
kosovensis
c
?- ?
\ . . .. ..
;;• EisenadUtina
\
0
;: ',"
.=.
-e Upper Pitinga
Fm .
CopoFm.
U_ \
gnJnulata

. . . .. .
"0 c caenipunco
• • •
::l
..J i
e
Fm. KiruSillas
Fm.
* Angochitina
flChina ta

e
,.....-- ---
"•
-"" iii
r ?
o E
0 0

c: ~ Lower caehi·
Ql I oonco Fm.*
~ -~ f---
3: ~8
v. Mb<.1
"'i ·.!1
? Sacta Marpachitina
4'e6n fm . margaritana -
salopochitina
JaicOs f m. CanyFm. *
>-
ii c
Lowe r Pitinga • --? - - - - - '--
zapla *
-lapla
- -* - -- -- -*
Cancafliri * Sa n Gabtn
motI lerrosae
~;'i~
~
~
Fm. .e,
Ql ':- • • Vargas Pena Fm.
- - -
Fm .
- - - - - - - - . . .. . . a
Fm. Fm dja /rnai
>
0
"0 -.
.-
§
-,
c • jpu Fm. Vila Maria
• Fm. * -
Spinachifiria •
c:
t1l ' c
'g .s f- - -
Fm.
--- .na.rri. ~i.
Beloned litina
:J ~c
Euse bio Ayala postrobusts
ao1l Fm .

Figure 8. Correl ation between Silurian lithostratigraphic unit s in western Gondwan a. a - Chitinozo an Subzone of Sphaerochitina
solutidina and b - Subzone of Desmochilina cf. D. densa. Asteri sk symbolizes occurrence of chitinozoan s.

plumula , Conochitina acuminata, Desm ochitina densa, densa, Conochitina acumin ata , Conochitina probos-
Euconochiti na cruzi, Linochitina penequadrata, Po- cifera, Sa/opochitina monterrosae and Ramochi tina
gono chitina cf. P. djalmai, Pterochitina deichaii and corniculata (Appendix 3).
othe rs (Fig. 3).
Reference section for zone base: PETROBRAS l-
7.g. A ngochitina echinate Interval Zone
AM - I -AM well.Ievel 1752 m, lower part ofthe Pitinga
Forma tion, Amazonas Basin, northern Brazil. No Zone base definition: FOB of Angochitina echinata
outcrop section with the base of the M. margaritana-S. (Fig. 5k).
monterrosae Zone is known. Characterization of zone: An abundant and diverse
Representative occ urre ncies of zone: Several local- chitinozoan fauna appears at the base of this zone
ities within the lower part of the Pitinga Formation in (Fig. 6).
the Amazonas Basin, northern Brazil. Reference section for zone base : Outcrop at
Lithostratigraphic distribution: See representative Coc habamba (Cramer, Diez & Cue rda, 1974), Peru-
occurrences of zone. Bolivia Basin, southern Bol ivia.
Age range: Latest Llandovery (late Telychian) to Repre sentative occurrences of zone: Several localit-
early Wenlock (Sheinwoodian) (Figs 6, 8-11). ies within the upper part of the Pitinga Formation in
Remarks: Margachitina margaritana and Sa/opo- the Amazonas Basin, north ern Brazil. Localities with
chitina monterrosae range from upperm ost Llandovery the upper part of the Kirus illas Formation in southern
(upper Telychian) to lower Wenlock (lower Sheinwoo- Bolivia.
dian) strata . Lithostratigraphic distribution: Upper part of the
In the global zonation (Verniers et a/. 1995) this Pitinga Formation in the Amazonas Basin, north ern
zone can be corre lated with the Angochitina /ongicollis Brazil, upper part of the Kiru sillas Form ation, Peru-
and lower Margachitina margaritana zones. Other Bolivia Basin, southern Bolivia and upper part of
important species in this inter val include Eisenackitina the Cachipunco Form ation, Tar ij a Basin , nort hern
dolioliform is, Conochitina emmastensis, Desmochitina Argentina.
Ord ovician and Silurian chitinozoan biozo nes 521

Western Global
Northern Gondwa na Ba ltica China Gondwa na zonat ion
Chrono- Ukraine Yangtze
Southwestern Saudi Arabia Baltoscandia La ufe ld,1971, Region
strat igraphy Europe Pa ris et al., La ufeld,1974, Nestor,1994, Ge ng et aI., This paper Verniers et al.
Paris,1981 1995, AI-Hajri Nestor,1994 Pa ris & 1997, Gra hn, 1995
& Pa ris,1998 Grah n,1996 unpub. data
UrnOChitina Cafpichitina AnthOChitina
24 velata superba
E. tiJifers
F.fungiformis
0 Margachitina
'C elegans
?
&: Ancyrochitina
Urnochitina
urna
fragifis FungOChitina Fungochitina
23
kosovensis kOSOvensis
S sCJluletlX:Pt'JhllJfa , rr.2UP'~ eo
c
•
E
E. ranosa
Eisenackitina
""
0 E. philiP; ? Eisenackitina
;: '5~ granuJata philo'"
E lagenomorplla
0
'0 -=-c B. latJfrons
A. elongata -- - - A. elongata

"
...J ~
G. militaris ? Ang<x:hitina
~
o
C. sp.2 echinata
C so. 1
22 Int rzone V
c 1 - - 21 - S.lycoperdoides
.~
•E 1--- 20 _ AnthOChitina A. indecora
?

~
jawfensis Conochitina
""s 0
I pachycephala
c: - c~
Q)
;;: s
c
.~
Fungochitina
fongicoflis
C. cinaulata
Margachitina
s: margaritana
(/) Bursachitina
M.m n Mm rit n M. margaritana
"

*
'SCI era ro ISCI era A. longicolfis S. monterrosae A. Jongicoflis
"g. 1 - - ' 9 ' ,
m
. "'-
r
A. loncJicollls
e ~
-'-,- Interzone III Eisenackitina
doliofiformis
P. djalmai ~
Eisenackitina
dOlioliformis
~ c
c.alargada
. emmas ensl
n erzooe
0 e C. edJelensis C. afargada
'C
c •
.s, P oaraauavensis
x C. 'Ossica S. harrington;
S. maennili
'" ~nI n,,
.~
...J -e

~
-c
~
n s sa B. postrooosta B. postrobusta
sa . sa
s:
a: t. , ,
" "
Figure 10. Correlation betweenSilurian chitinozoanbiozones in western Gondwana and otherareas on a global scale. I - Chitinozoan
Subzone of Sphaerochitina solutidina and 2 - Subzone of Desmochltina cf. D. densa ,

7.i. Fungochitina IW.·ww!tl.'l;S Ran ge Zo ne
Zone base definition: FOB of Fungochitina kosovensis
(Fig.5m).
Characterization of zone: The co-occurrence of
the index species, Ancyrochitina afT. A. libyensis,
Angochitina n.sp. afT. A. cyrenaicensis, Ango chitina?
sp. sens u Grahn & Paris. 1992, Armigutta urubuense,
Clat hroc hitina sp. A sel/Sll Grahn. 2003, Ramochitina
bjomsundquisti, Rhabdoch itina conocephala , Sahar-
och itina gomphos, Urnochitina llrlla. Urochitina n.sp.
A sensu Grahn. 2005a , Vinnalochitina corinnae and
others.
Referenc e section for zone base: PETROBRAS 1-
AM-I -AM well, core 46 (1613 m), uppermost part
of the Pitinga Form ation , Amazona s Basin, northern
Brazil. No outcrop section with the base of F. kosovensis
Zone is known.
Representative occ urrences of zone: Seve ral local-
ities within the uppermost part of the Pitinga and/or
lowermo st part of the Manacapuru format ions in the
Amazonas Basin, northern Brazil.
Lithostratigraphic distribution : Upper most part of
the Pitinga and lowermost part of the Manacap uru
formation s in the Amazona s Basin , nort hern Brazil and
Kirusillas Formation, Peru -Bolivia Basin, southern
Bolivia.
Age range : Early Pridoli (Figs 7-11 ).
Remarks: Fu ngochitina kosovensis is restricted to
lower Pridoli strata.
Paris & Kriz (198 4) de fined the F. kosovensis
Zone in Bohem ia. and the accompanying species
include Calpichitina gregoria, Pterochit ina perivelata,
Eisenackitina cf. inte rmedia, Urnochitin a llnw and
others, In the global zonation (Vern iers et al. 1995 )
the Fungochitina kosovensis Zone is coeval with
this zone in western Go ndwana. Other important
species in this inter val include Eisenackitina il/-
term edia and Eisenackitina lageno mo rpha (Appe n-
dix 3).
522 Y. G RA H N

S ilurian biostrat igraphic correlat ion of western Gondwana

>.
, .c Grapo tolite
00. Acritarch Spore zon es
c ~ zones Cono dont Spore zones Chitinozoan
e o>
~ -;:; Suarez-Soru co, zones
zones
Le Herisse, Gray,198 5,
Le Herisse
et al. 2001,
zones
O~
CJ) -
1975, Mal etz ,
Suarez-Soruco
& Egenh off 200
Me rino , 1991
Haytham & van
der Eem ,1995,
Le H et al. 200 1
1989, Gray
et al. 1992
Steemans &
Pereira,2002
Th is paper

"6 .
'0
.;;:
e,
Fungochitina
kosovensis
m
:e
0 . ... Eisenacki tina
;: '5 S. leintwardi- g ranula ta
o ~ 9
,...d. nen sis ~----
'0 C
3 .~ S. scanicus . Angochitina
~ 8
0 echinata
C)
N. nils soni
C
'C
Q) '" P. pra edeubeli . .
~ E
o 0
o I ~-
c-;-
OJ 0
? -- 7 ?
~ c;: 0. s ag itta
'iii rhenana Micro fossil
s: ?
Ul I· Assemblage
6 Archaeozono- M. margaritana
.t:::
oc-, Interval . .. " triletes cnutas- S. monterrosae
nanus
~~ w ith P. dj almai ~
0>7 * .-* Climaco-
>
5 L. divellomedia
o 0~ graptus Microfossil II ...J.
'0 Q)
S. harringtoni
Assem blage
lij ~ 4 Laevolancis
-
...J -o
-0
_ 1-3 I divellomedia
~
Belon echitina
No zonation I postrobus ta
ff *
Figure II . Correlation between Silurian graptolite. conodont, acritarch, spore and chitinozoan biozones in western Gondwana. I -
Chitinozoan Subzoneof Sphaerocbitina solutidina and 2 - Subzoneof Desmochitina cr. D . densa. " Basedon graptolites. Rickards et at.
(2002), considered the lowermost part o f the Lipe6n Formation as early Rhuddanian (acuminatus- atavus zones). Acritarchs (Bultynck
& Martin. 1982; Rubinstein, 1997) and chitinozoans (Grahn & Gutierrez, 200 1). however, suggest a Telychian-Sheinwoodian age
for that part. Interval with Climacograptus indicate finds of Monograptus cf. gregarius and Climacograptus innotatus brasi liensis in
the Amazonas Basin (Jaeger, 1976) and Climacograp tus scalaris sca laris in the Parnaiba Basin (H. Jaeger, pers. comm. 1992)... -
Monogra ptus turriculatus Zone is present at the base of the Kirusillas Formation (Ahlfeld & Branisa, 1960; Suarez-Soru co, 1975).

8. Conclusions informatio n from the investigated areas wi ll al ter and

Thi s erection of the chi ti nozoan bio zonation for
the Ordovician and Silurian of western Gondwana
(South A merica) is hampered by the scattered Ordovi-
cian in formation avail able. Future information from
Ecuador, Colombia and Venezuela and complementary
refine this biozonatio n, especiall y for the Ordov ician.
It is clear that the Ordovician chit inozoan assemblages
have a mi xed provincia li ty with elements from both
Laurentia, Baltica and Gondwana. The Silurian chitino-
zoan assemblages di splay a more distin ct Gondwana
affinity. South America was in a polar to circumpolar
Ordovician and S ilurian ch itinozoa n biozones
positi on during the Ordovician an d Si lu rian, and
the c hitinozoan assemblages there fo re should refl ect
a cold wa te r fa una . As po inte d out by G rahn &
Bergstrom ( 198 4), chiti nozoan co ld wa ter assem blages
seem to be more di verse and abundant th an tro p ical
assemblages . T his is a lso true fo r western Gondwana
(cf contempora ry tro pical fau nas in Ba lt ica an d
La ur en tia). The nature of th e sam p ling (mostly oil
co m pany borings and iso lated outcrops) have preve nted
a more dense sam pling, w hich undoubtedly could
reveal a m ore deta ile d chi tinozoan b iozo nat ion for
we ste rn Gondwana. A ltogether five chi tino zoan zo nes
arc defi ned in th e Ordovician (zones of Desmochitina
sp . g r. minor, Conoc hitina de cipiens, Eremoch itina
brevis, Lagenochitina obeligis and Tanuchitina an -
ticostiensis), an d nine in the Silu ri an (zo ne s of
Belonechitina postrobusta, Spina chitina harringtoni,
Pogonochitina djalm ai , Margachitina margaritana-
Salopochitina monterrosae, Angochitina ech ina ta, Eis-
enacki tina gra nulate, Fu ngo chitina kos ovensis and th e
s ubzones of Sp haerochitina so lutidina and Desmochit-
ina cf. D. densa). These b iozo ne s ca n be correlated
w ith o ther areas o n a g lo bal scale .
Acknowledge me nts. The author thanks the Faculty of
Geo logy at Universidade do Estado do Rio de Janeiro
(UERJ ), and Dr Egberto Pereira, head of the post-graduate
program at the Faculty of Geo logy at UERJ for access to the
facilities, and the Conselho Nacio nal de Dcsenvolvimen to
Cientifico e Tccnologico (CNPq, PQ 303777/02-8), which
made this work possib le throug h grants. Through the co urtesy
of Dr Sylvia Maria Couto dos Anjo s, former head of BPA
at CENPES (PETROBRAS, Rio de Jane iro), and Prof.
Florent in Paris (Rennes, France), I had access to Scanning
Electron Micro scope s. Prof. Art Boucot (Corva llis, Oregon)
is acknow ledged for checking the English. I am very grateful
to Profe ssors Stig M. Bergstrom (Co lumb us, Ohio) and
Jacque s Verniers (Ghe nt, Belgium) for their carefu l revisions
of the final version of the ms., whic h ted to a much improved
work. All technical help from Jorge Louiz dos Santos and
Maria Rosalva Campos Coelho at UERJ and Rogerio da
Silva Mart ins da Costa at CENPES is grea tly appreciated .
My sincere thanks to all.
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Ordovician and Silurian ch itin ozo a n biozones
Appendix 1. Revised chit inozoan species
So uthern Bolivia
Cramer, Diez & Cuerda (1974)
Ancy rochitina ancyrea Bolivian variant = Ancyrochitina
n.sp . A sensu Grahn. 20050
Ango chitina sp. 2 = Ango chitina echina ta Eisenack, 1931
Ango chitin a sp. 1 = Ango chitina sp.
Linochitina erratica . . .. L. cingulata = Cingulochitina con-
vexa (Laufeld, 1974) and Cingulochitina wmnui Paris,
1984
Conochitina proboscifera = Conochitina tuba Eisenack,
1932
Ptero chltina perivetata (Eisenack, 1937)
Desmochitina urna = Urnochitina uma (Eisenack, 1934)
Liachenko ( 1994)
Ancyrochiti na cf. ancyrea . Fungochitina sp. A, Gotlan-
dochitina sp. I , and Ancyrochitina sp. = Ancyrochitina
spp.
Angochitina cf. echin ata and Angochitina sp. A-E = Ango-
chitina echinata Eisenack, 1931
Sp haeroc hitina sp. I =Angochitina sp.
Sp haerochitina sp. I (PI. 2, fig. 9) = Fungochitina kosov en-
sis? Paris, 1981
Cing ulochitina aff. sp. se rrata and Cingulochitina se rrata =
Cingulo chit ina se rrata (Taugourdeau & Jekhowsky, 1960)
Conochitina probos ciftra and Conochitin a sp. A = Cono-
chitina go rdonensis (Cramer, 1964)
Co nochitina sp. C = Conochitina probiscifera Eisenack,
1937
Co noc hitina sp. B = Co nochitina tuba'! Eiscnack, 1932
Eise nackitina? sp. = Eisena ckitina granulata (Cramer, 1964)
Marga chitina sp. I and Calpichitina (D ensichitina )
sp. I = Margachitina catenaria Obut, 1973
Pte rochitina perive lata (Eisenack, 1937)
Northern Argentina
Otton e, Toro & Waisfeld ( 1992)
Conochltina sp. A = Eremochitina brevis Benoit &
Taugourdeau, 1961
Spathachitina sp. A = Con ochitina ordinaria Achab, 1980
Sp athachitina sp. Band Lageno chitina sp. A = Conochitina
brevis Taugourdeau & Jekhow sky, 1960
Eastern Paraguay
Wood & Miller ( 1997)
Ancyrochitina sp. 5 = Ancyrochitina p aran aensis Grahn in
Grahn, Pereira & Bergamaschi, 2000
Desmochitin a cf. D. densa Eisenac k, 1962
Conochitina sp. (PI. 16, fig. 5) = Conochitina p robisciferal
Eisenack, 1937
Canochitina sp. (PI. 16, fig. 6) = Conochitina sp. A sensu
Grahn, Pereira & Bergama schi, 2000
Eisenackitina cf. E. bejui Wood & Miller, 1991
Linochitina sp . = Lino chitina sp. A sensu Grahn, Pereira &
Bergamaschi, 2000
Brazil
Sommer & van Boeke l (196 3, 1965)
Ancyrochitina erichse ni Sommer & van Boekel, 1963
Conochitina dj almai = Pogono chitina djalmai Sommer &
van Boeke l, 1965
527
Spha erochitina lucian oi = Sphaero chitina palestinaense
Grahn , Melo & Steemans, 2005
Costa (1970, 1971 a)
Pallachitina wilh elmi and Palla chitina dep ressa = Bursa -
chitina wilhelmi (Costa , J 970)
Spathachitina cruzi, Spatha chitina clarindot, Spathachit-
ina reticulata, and Spathachit ina tenuis = Euconochitina
cruzt (Costa, 1970)
Spath achitina maga /hues i and Sputhu chitina raih ensi s in
Costa, 1970, and in Costa, 1971a Conochitina acuminate
(PI. I, fig. I), Conochitina dec ipiens (PI. 1, fig. 5),
Conochitina ftavioi (PI. 2, figs 4-5 ), Conochitina ped-
unculosa (PI. 3, figs 1- 6, PI. 4, figs 1-2), Con ochitina
simplex (PI. 4, fig. 3), Conoc hitina turris (PI. 4, fig. 5),
Spathachitina rathensis (PI. 7, fig. 4), Sp athachitina
maga/haesi (PI. 7, figs 5--6), Tanuchitina ontariensis
(PI. 8, figs 1-4 ), Tam/chitina proemtnentts (PI. 8, figs 5-6),
Tanuchitina paraensis (PI. 9, figs 1- 7), I/lichitina multiplex
(PI. 12, fig. 5), Pogonochitina grandis (PI. 13, figs 3, 5),
Cyathochitina brasili ca (PI. 15, figs 1-3), Cyathochitina
contca (PI. 16, fig. I ), Cyathochitina dispar verrucata
(PI. 16, figs 2-3 ), Cyathoc hitina campanulaeformis (PI. 16,
fig. 5), Cy athochitina cap utoi (PI. 16, fig. 6), C yathochitina
elinitae (PI. 17, figs 1-4), aod Cyathochitina cf. gra nulata
(PI. 17, fig. 7) = Pogonochiti na djalllla i Sommer & van
Boekel, 1965
Cy athoc hitina hym enophora nigerica (PI. 17, figs 5-6) =
Sa/opochitina monterrosae (Cramer, 1969)
Conoc hitina latifrons (PI. 2, fig. 3), Pogono chitina gra ndi
(PI. 12, fig. 7, PI. 13, figs 1-2, 4,6), and Pogonochitina
sp inifera (PI. 14, fig. I ) = Pogonochitin a aff. P. inomata
(Costa, 197 Ia)
Pogon ochitina spinife ra inorn ata (PI. 14, figs 2-4)=PoglI-
nochitina inom ata (Costa, 1971 a)
Conochitina intermedia (PI. 2. figs 1- 2) = Pogono chitina cf.
P. djalllloi Sommer & van Boekel, \ 965
Desm ochitina cing ulata (PI. 18, fig. 3), Desmochitina
cing ulata serrata (PI. 18, figs 4-7), and Hoegisp haera
complanata (PL 18, fig. 8) = Cingulochitina ex. gr. serrata
{Taugourdeau & Jekhowsky, 1960)
D esmochitin a erratica (PI. 18, figs 1- 2) = Linochitin a ex. gr.
erratica Eiscnack, 193 1
Cono chuina vascu liformis (PI. 4, fig. 6), Sp atha chitina
cn a t (PI. 6, figs 1- 2), Spathachitina clarindoi (PI. 6,
figs 3-5 ), Spath achirina reticulate (PI. 7, figs 1- 3), and
Spathachitina tenuis (PI. 7, fig. 7) = Eucon ochitina cn u i
(Costa, 1970)
An cyrochitina spinosa (PI. I I, figs 1- 2) = Ango chitina sp. A
sensu Grahn & Paris, 1992
Palla chitina wilhelm i (PI. 5, fig. I). Pa llachitina rotunda
(PI. 5, fig. 2), aod Pallachitina depressa (PI. 5, fig. 3) =
Bursa chit ina wilhelmi (Co sta, 1970)
Angochitina amazontca (PI. II , fig. 6) , Angoc hitina crumena
(PI. 11, figs 7-8), Lagenochitina so mmeri (PI. 14,
figs 5- 8), and Lagen ochitina ovoidea (PI. 14, figs 9-
10) = Lagenochitin a sp. aff. L. navicula Taugourdeau &
Jekhowsky, 1960
Cyathochitina caputo! (PI. 15, figs 6-7 ) = Cyathochitina sp.
B se nsu Paris, 1981
An cy rochitina ancy rea (P I. 10, figs 2-4), Ancyrochitina
megastyla (PI. 10, fig. 5?), and Ancyrochitina primittva (P I.
10, figs 6-8) = Ancyrochitina primitiva? Eisenack, 1964
PleclOchitina sahariea (PI. 1\ , figs 3- 5), Sph aerochitina
collinsoni (PI. 12, fig. 2), and Cyathochitina campanu -
laef ormis (PI. 15, figs 4-5 ) =Ancyrochirina sp. A sens u
Grahn & Paris, 1992.
 528
Quadros ( 1982)
Desmochitina sp. = Cingulochitina bouniensis Verniers,
1999
Cyathochitina sp. = Pogonochitina tianguaense Grahn ,
Me lo & Steemans, 2005
Conochitina dolosa = Conochitina elongata Taugourdcau &
Jekhowsky, 1960
A p pen d ix 2. Li st of chitin oz o an species repo rted fr om
western Go ndwana and cited in text an d figures
An cy rochitina ancyrea (Eiscnack, 1931)
Ancyrochitina brevis Taugourdeau & Jekhowsky, 1960
An cyrochitina erichseni Sommer & van Boekel, 1963
Ancyrochit ina fragiiis Eise nack , 19550
Ancyrochitina para naensis Grahn in Grahn. Pe reira &
Bcrgamasch i. 2000
An cyro chitina p itinga ense Grahn, 20050
An cyro chitina polentinensis Schweincbcrg, 1987
An cyro chitina primitiva Eisenack, 1964
An cy rochitina regularis Taugourdeau & Jekh owsky, 1960
Ancyrochitina aff. A . Iiby ensis Jaglin, 1986
Ancyrochitina aff A. regu laris Tau gourdcau & Jekh owsky,
1960
Ancy rochitina aff. A . tom elllo.m Taugo urdcau & Jekhowsky,
1960
Ancyrochitina cf. A. brevis Taugourdcau & Jekhowsky, 1960
An cyrochitina cr. A. desm ea Eise nack, 1964
An cyrochitina sp. A se nsu Grahn & Paris. 1992
Ancyrochitina sp. A. sens u Gra hn & Melo. 2003
An cy rochitina sp. B sens u Grahn & Me lo, 2003
An cy rochitin a sp. B. sens u Grahn, Melo & Steemans , 2005
Ancyrochitina sp. C sens u Grahn & Mel o, 2003
Ancyrochitina n. sp. A. se ns u Grahn. 2005 a
An cy rochitina n. sp. B. se ns u Grahn, 2005 a
Ancy rochitina ex . gr. ancyrea (Ei senack, 193 1)
Ancyrochitina ex . gr.jloris Jagli n, 1986
A ncyrochitina ex. gr. gutnica Laufeld, 1974
A ngochitina echina ta Eisenac k, 1931
A ngochitina elongate Eisenac k, 1931
Angochitinafi/osa Eisenack, 1955a
Angochitina gurupiens e Grahn. Melo & Steemans, 2005
An gochitina longicollis Eisenaek, 1959
Angochitina strigosa Boumcndjel, 2002
Angochitina n. sp. afT. A. cyre natcensts sensu Grahn & Pari s,
1992
Ango chitina aff.jiloSQ Eisena ck , 1955a
Angochitina sp. aff. A . mourai non Lange , 1952 se nsu
Schweineberg, 1987
Angochit ina cf. A. echinate Eisc nack. L'B t sensu Gra hn &
Paris. 1992
Ango chitina cf. A . e/on gata Eisenac k, 1931
Angochitina sp. A sensu Grahn & Paris, 1992
Angochitina sp. A sensu Grahn & Gutie rre z, 2001
Angochitina sp. 1 sensu Grahn & Gutierrez. 2001
Angochitina sp. I . sensu Grahn . 2005 b
Angochitina sp. 2 se nsu Grahn & Guti errez. 2001
Angochitina sp. B. sensll Grahn & Melo , 2003
Angochitina? thodeui Paris, 1981
An gochitina? sp. sensu Grahn & Paris. 1992
Armigutta urubuense Grahn & Me lo, 2003
Armoricochitina nig erica (Bo uche, 1965)
Belonechi tina? p lumu lu n. sp. Grahn , 2005a
Belonechitinapostrobusta (Nestor, 1980)
Belonechitina sp. A. sens u Gra hn. 2005a
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Y. G R A HN
Be/onechitina sp. B. sens u Grahn. 2005a
Bursachitina wilhelmi (Co sta. 1970)
Bursachit ina sp. A. sensu Gra hn & Me lo, 2003
Cingu lochitina bo uinensis Verniers. 1999
Cing ulochitina canvex a (Laufel d, 1974)
Cing uloch itina cylindrica (Taugourdeau & Jekh owsky, 1960)
Cingulochitina serrata (Taugourdeau & Jekhowsky, 1960)
Cingulochltina wronai Pari s. 1984
Cingu lachitina aff. C. conwxa (Laufeld, 1974)
Cingu/oehi tina aff. C. ervensts (Paris in Babin et al., 1979 )
Cingu/ochitina aff. C. serrata (Taugo urdea u & Jekhowsky.
1960 )
Cingulochitina sp . A. Paris. 19 81
Ctathrochitina sp . A. sens u Grahn. 2003
Conochitina acuminata Eisenac k, 1959
Conochltina brevis (Taugo urdea u & Jek howsky, 1960 )
Conochltlna deciptens (Taugo urdea u & Jekh owsky, 1960)
Conochitina edjelensis (Taugo urdea u & Jekhowsky, 1960)
Conochitina elongata (Taugo urdeau & Jekhowsky, 1960)
Conochiti na gordonensis Cramer. 1964
Con ochitina micracantha (Eisenack, 1959)
Canochitina ordinaria Achab, 1980
Conochitina pachycephala Eisenack, 1964
Conochitina p ervulgata ? Umnova, 196 9
Conochitina prob oscifera Eisena ck , 1937
Conochitina tuba Eise nack . 1932
Conochitina cf. C. acuminate Eiscnack, 1959
Conochulna cf C. armillata Taugourdeau & Jckhowsky,
1960
Conochitina ef. C. havliceki Paris & Mc rgl. 1984
Conochitina cf. C. tuba Eisenack, 1932
Con oehitina sp . A sensu Gra hn. 1992
Con ochitina sp. A sensu Grahn in Grahn. Pereira &
Bergamaschi, 2000
Conochitina sp. sens u Heuse. Grahn & Erdtmann, 1999
Cyathochitina campanulaefo rmis (Eisc nack, 1931)
Cyathodntina eap uto i Co sta. 1971a
Cyathoc hitina sp_B sens u Paris, 198 1
Cyathochilina cf Ci kuckerstana (Eisenack. 1934 )
Desmochitina cortesiana Schweinebe rg, 1987
Des mocJritina densa Eise nack , 1962
Desmochitina sp. gr. min or (Sza niawski in Chlebowski &
Szaniawski, 1974 )
Desm ochitina ef. D. densa Eisenack, 1962
Eisena ckitina bej ui Wood & Miller, 1991
Eisenackit ina ef. E. bej ui Wood & Miller, 199 1
Eisenackitina g ranulata (Cramer. 1964 )
Erem ochitina brevis Benoit & Taugourdea u, 1961
Eucon ochitina cruzi (Co sta, 1970)
Eucon ochitina ik/aensis (Nestor, 1984)
Euconochitina patu/a (Co sta, 1971 a )
Euconochitina sulc ata (Co sta, 1971 a )
Euconochitina sp. A. sensu Gra hn, 2005a
Fungoehitina kosovensis Paris, 1981
Fungochitina sp. A. se ns u Grahn & Melo, 2003
Lagenochitina obeligis Paris. 1981
Lagenochitina prussi ca Eiscnack, 1931
Lagenochitina aff. L. na vicula Taugourdcau & Jekh owsky,
1960
Linochitina p enequadrata n. sp. Grahn. 2005a
Linochitina aff. L. klonk ensis Paris, Lau feld & Chlupae ,
198 1
Linochitina ex . gr. erratica (Eisenac k, 193 1)
Linochltina sp. A sensu Grahn in Grahn. Pereira &
Bergam asehi, 2000
Lino chitina sp. A sens u Gra hn & Me lo, 2003