Paper

1. Journal of South American Earth Sciences 37 (2012) 1e14
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Journal of South American Earth Sciences
journal homepage: www.elsevier.com/locate/jsames
The first recorded decline of the Malvinokaffric Devonian fauna in the Paraná
Basin (southern Brazil) and its cause; taphonomic and fossil evidences
O primeiro registro do declínio da fauna Malvinocáfrica do devoniano da bacia do
Paraná (sul do Brasil) e suas causas; evidências fósseis e tafonômicas
Elvio Pinto Bosetti a, Yngve Grahn b, *, Rodrigo Scalise Horodyski c, Paula Mendlowicz Mauller b
a
Universidade Estadual de Ponta Grossa, Departamento de Geociências, UEPG, Av. Carlos Cavalcanti 9500, Uvaranas 84010-919, Ponta Grossa, P.R., Brazil
b
Universidade do Estado do Rio de Janeiro, Faculdade de Geologia, Bloco A e Sala 4001, Rua São Francisco Xavier 524, 20550-013 Rio de Janeiro, R.J., Brazil
c
Universidade Federal do Rio Grande do Sul, Programa de Pós-graduação em Geociências, Av. Bento Gonçalves 9500, Bloco I, Prédio 43113, Campus do Vale,
91509-900 Porto Alegre, R.S., Brazil
a r t i c l e i n f o a b s t r a c t
Article history: The first recorded decline of the Malvinokaffric Devonian fauna in the Paraná Basin, south Brazil,
Received 7 December 2011 occurred between early and early late Emsian. The event is recognized on biostratigraphic implications,
Accepted 29 February 2012 and it coincides with a regional hiatus resulting from tectonic activity during a Precordilleran epeirogeny.
The Emsian beds, dated with palynomorphs, indicate a late Pragian - early Emsian (PoW Su spore Zone)
Keywords: age below, and an early late Emsian (upper FD e lower AP spore Zones) age above the hiatus. The
Malvinokaffric
extinctions that occurred between the late Pragian e early Emsian regression, and the initial trans-
Faunal decline
gression in the early late Emsian, were extensive and more dramatic than elsewhere on the globe. In the
Taphonomy
Palynology
Paraná Basin invertebrates 4 classes, 25 families, 41 genera, and 54 species become extinct. The Emsian
Invertebrates sedimentation in the Paraná Basin was realized under constant oscillation of sea level punctuated by
ParanáBasin storm events, probably due to Milankovitch orbital cycles. It is known that the eccentricity cycles are the
Palavraschave: most striking in regions at higher latitudes, which was the case of the ParanáBasin during the Emsian.
Malvinocáfrica This was, however, not a factor of major importance for the large decline of the fauna. The decline of the
declínio da fauna shelly fauna was an effect of the late Pragian e early Emsian regression, and the early late Emsian
tafonomia transgression introduced a reduced and less provincial shelly fauna.
palinologia Ó 2012 Elsevier Ltd. All rights reserved.
invertebrados
Bacia do Paraná r e s u m o
O primeiro registro do declínio da fauna Malvinocáfrica do Devoniano da Bacia do Paraná (sul do Brasil)
ocorreu entre o Eo e o início do Neo Emsiano. O evento é reconhecido por suas implicações bio-
estratigráficas e coincide com o hiato regional resultante de atividade tectônica durante a epirogenia da
Pré-Cordilheira andina. As camadas, datadas por meio de palinomorfos, indicam uma idade Neo Praguiana-
Eo Emsiana (Zona de esporo PoW Su) abaixo do hiato e uma idade do início ao fim do Neo- Emsiano (parte
superior das zonas de esporo FD e parte inferior AP) acima do hiato. As extinções que ocorreram entre
a regressão no Neo Praguiano - Eo Emsiano e o começo da transgressão no início do Neo Emsiano foram
amplas e mais dramáticas do que em qualquer outra parte do globo terrestre e entre os invertebrados da
Bacia do Paraná, 4 classes, 25 famílias, 41 gêneros e 54 espécies foram extintas. Durante o Emsiano, na Bacia
do Paraná, a sedimentação ocorreu sob constantes oscilações do nível do mar entremeadas por eventos de
tempestades, provavelmente devido à ciclos orbitais de Milankovitch. É de conhecimento geral, que os
ciclos de excentricidade são mais notáveis em regiões de latitudes mais altas, o que era o caso da Bacia do
Paraná durante o Emsiano. Contudo, este não foi o fator de maior importância para o grande declínio da
fauna. O declínio da malacofauna foi um efeito da regressão ocorrida no Neo Praguiana e Eo Emsiano e
a transgressão durante o início do Neo Emsiano introduziu uma fauna reduzida e menos provinciana.
Ó 2012 Elsevier Ltd. All rights reserved.
* Corresponding author. Fax: þ55 21 38657093.
E-mail address: yngvegrahn@gmail.com (Y. Grahn).
0895-9811/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jsames.2012.02.006

2. 2 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
Fig. 1. Location map of the localities from the ApucaranaSub-basin discussed in this study.
1. Introduction Furthermore, the extinctions are known to be gradual and
spread over a longer time (House, 2002). In the type area (Bar-
The aim of this paper is to document the first recorded decline of randian), the Daleje Event is connected to a near-loss of loosely
the Malvinokaffric Devonian fauna in the Paraná Basin as suggested coiled mimosphinetids and Teicheroceratidae ammonites (House,
by taphonomic control and biostratigraphic implications. Paly- 2002). As pointed out by Boucot et al. (1969), there is a global
nomorph assemblages date the event to middle - early late Emsian, reduction of brachiopod generic diversity between early and late
which seemingly coincides with the global Daleje or gracilis-can- Emsian. Also the palynomorphs are affected (e.g. McGregor, 1979;
cellata Event (House, 1985, 2002; Chlupá and Kukal, 1986;
c Melo and Loboziak, 2003). The Daleje Event has also been recog-
Walliser, 1996). A global sea-rise (transgressive and gradual nized in other parts of Europe, Asia, North America, North Africa,
according to Chlupá and Kukal, 1986) occurred in the late Pragian -
c and Australia (Chlupá and Kukal, 1986:172e173; Talent et al.,
c
early Emsian (Sequence B in the Paraná Basin sensu Grahn et al., 1993:143; Becker and House, 1994:82e90). The absence of diag-
2011), followed by a rapid regression in the Paraná Basin nostic miospores such as Camarozonotriletes sextantii, Emphanis-
(Bergamaschi and Pereira, 2001), and a large diversity decline of the porites annulatus, Emphanisporites foveolatus, and Verruciretusispora
shelly taxa (this paper). The Daleje Event is in general not charac- dubia for the AB and lower FD spore zones in the Paraná Basin, and
terized by a mass-extinction, and it is considered as a low level the regression in late Pragian e early Emsian strata, below the AB
event. spore Zone, suggest the possibility of a tectonic cause to the hiatus

3. E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14 3
and the faunal extinction. The miospore species Rhabdosporites recently been published by Grahn et al. (2010, 2011). Unequivocal
minutus and Samarisporites praetervisus are known to have their middle Emsian strata have not been recorded in the Parana Basin (e.g.
first occurrences in the FD Zone (Streel et al., 1987), but in the Melo and Loboziak, 2003; Grahn et al., 2010, 2011), or any other
Paraná Basin they first occur in the late Emsian e early Eifelian GS intracratonic basin of Brazil or in the adjacent southern Bolivia
Zone (Melo and Loboziak, 2003; Grahn et al., 2010). Furthermore, (Racheboeuf et al., 1993; Grahn, 2002, 2005; Melo, 2000, 2005). This
E. annulatus also have a late Emsian first occurrence, and C. sextantii could be interpreted as caused by regional tectonic activity (e.g. Melo,
is known only from the late early e latest Eifelian Per Zone (Melo 2000; Pereira, 2000; Miranda and Della Favera, 2005) leading to
and Loboziak, 2003; Grahn et al., 2010). starved sedimentation or a very condensed accumulation. The pre-
hiatus interval, represented by Sequence B (sensu Grahn et al.,
2. Geologic setting 2011) shows, in general, a transgressive trend (retrogradation) with
distal facies, and increasing mud content toward the top and the
The Paraná Basin is one of the largest intracratonic basins of South Maximum Flooding Surface of the interval. Then followed a regres-
America, covering ca. 1,600,000 km2 across southern Brazil, eastern sion, possibly due to a basin sediment fill to depositional base-level
Paraguay, central Uruguay, and northeastern Argentina (Fig. 1). (cf. Tankard et al., 2009). The boundary to Sequence C is abrupt.
Updated lithostratigraphic and biostratigraphic information on the The AB and lower FD spore zones have not been identified in the
Devonian from the Paraná Basin in southern Brazil (Fig. 2) has Paraná Basin. The post-hiatus level is initiated by Sequence C (sensu
Chitinozoans
Paraná Basin Spores Grahn 2005
Age Sequen-
ces Mendlowicz SEA LEVEL CURVE
Apucarana Alto Garças Mauller et al.
Sub-basin Sub-basin * A B 2009
TP
U.bastosi fall rise
S. langei
IV
BMu
Frasnian
Hoegisphaera
BM BPi glabra
BJ
Chapada F
Group
TCo Fungochitina
unit 4 microspinata
Trg
Ancyrochitina
Givetian São taouratinensis
Domingos TA
Fm. Ramochitina
Lem
E Lli stiphrospinata
AD
Pre-Lem
Chapada Group unit 2
Alpenachitina
Per eisenacki
Cha-
pada D
Eifelian ?
Group
Vel
unit 3 Ancyrochitina
varispinosa**
AP
GS
C
Pre-Vel
Ancyrochitina
? parisi
Ti
Emsian ? ? FD Not
yet
defined Ancyrochitina
AB pachycerata ?
Ponta Chapada Su
B
Grossa Fm. Group unit 2
PoW
Pre-Su
Ramochitina
Pragian Ems magnifica
E
Urochitina
BZ
? ? E
Z loboi
Loch- Chapada
Furnas Fm. Group Angochitina
kovian A
unit 1 MN NsZ strigosa
***
Fig. 2. Diagram showing stratigraphy and sea level curves during late Pragian - Emsian to early Givetian of the Apucarana Sub-basin. The double-headed arrow next to the encircled
letter E (within the Ems miospore Zone) indicates the maximum stratigraphic span proposed by Rubinstein et al. (2005) for the D. emsiensis morphon Assemblage-zone. Other
conventions: A = Western European miospore zonation after Streel et al. (1987) and Steemans (1989). B = Western Gondwanan (North Brazil) miospore zonation after Melo
Loboziak (2003). Ti. ¼ Tibagi Member of the São Domingos Formation. * ¼ Alto Garças Sub-basin after Grahn et al. 2010. ** ¼ informal biozone. *** ¼ Angochitina praedensiba-
culata Zone.

4. 4 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
Grahn et al., 2011) approximately at the base of the serotinus con- marked MP-P have a permanent repository at the palynological
odont zone (upper FD spore zone sensu Melo and Loboziak, 2003). slide collection, Laboratory of Palynology Marleni Marques Toigo,
Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil.
3. Materials and methods Slides marked CB and BPA have a permanent repository at the
Biostratigraphy and Paleoecology Management of Petrobras
The paleontological materials were investigated and collected Research Center, Petrobras/Cenpes/Pdexp/Bpa, Ilha do Fundão, Rio
from the lower São Domingos Formation at Desvio Ribas-Tibagi, de Janeiro, RJ, Brazil. The field work was performed in accordance
Metalúrgica Águia, Vendrami, Colônia Sutil, Caça e Pesca, Boa with the taphonomic protocol proposed by Simões and Ghilardi
Vista, and Rio Caniú (Figs. 1 and 3e5). The materials (DEGEO/MPI- (2000), and the collection procedures proposed by Bosetti (2004).
777 to DEGEO/MPI-996) are deposited at Universidade Estadual de For the characterization of the fossiliferous taphofacies seven
Ponta Grossa (Paleontology Laboratory of the Geosciences outcrops were standardized and investigated for the degree of
Department). The megafossils were processed with the help of fine packing, fragmentation, disarticulation of bioclastics, and position
brushes and needles, and the microfossils with Petrobras standard relative the bedding planes. Additionally, sedimentary structures
methods (Quadros and Melo, 1987). The palynological slides and textures were observed.
Fig. 3. Lithologic columns and palynomorph ranges in Metalúrgica Águia, Boa Vista, and Vendrami localities.

5. E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14 5
Fig. 4. Lithologic columns and palynomorph ranges in Desvio Ribas-Tibagi and Caça e Pesca localities. For legend, see Fig. 3.
4. Localities Orbiculoidea baini; partially fragmented valves of infaunal lin-
gulids, Nuculites sp., Palaeoneilo sp., Pleurodapis sp., Edmondia
Seven outcrops were examined, e.g. Desvio Ribas-Tibagi, Met- sp., and Modiolopsis sp., disarticulated valves on bedding planes,
alúrgica Águia, Vendrami, Colônia Sutil, Caça e Pesca, and Boa Vista, with infaunal lingulids, Australocoelia palmata, Nuculites sp.,
all situated in the southern part of Ponta Grossa city, and Rio Caniú Palaeoneilo sp., Pleurodapis sp., Edmondia sp., and Modiolopsis sp.
in Palmeira city (Figs. 1 and 3e5). Together they cover a geographic T1b: (moderate degree of fragmentation and lateral reworking).
distance of ca. 48 km. The outcrops vary in thickness, but the Composed of fragmented valves of infaunal lingulids, Austral-
lithofacies are the same. These include silty mudstones, siltstones, ocoelia palmata and Australospirifer spp., and disarticulated
claystones, locally sandy siltstones with cross-lamination, interca- valves of infaunal lingulids, Nuculites sp., Palaeoneilo sp., Pleu-
lated with a hard crust of silty, fine to medium-grained sandstone, rodapis sp., Edmondia sp., and Modiolopsis sp.
and sandstone with hummocky cross structures (HCS). Thickness Taphofacies T2: Composed of articulated bioclasts, whole and in
and length of these structures are decimeter to meter thick apparent life position. This concentration can be classified as
(uncommon in the Devonian of Paraná, which typically are milli- loosely packed, where the bioclasts may exhibit some physical
meters to centimeters). Most beds are overlain by plane-parallel, contact.
dark gray to black, silty shales. T2a: (moderate degree of articulation and most specimens in life
position). Composed of valves in apparent life position of
5. Taphofacies infaunal lingulids, Nuculites sp., A. palmata, disarticulated valves
parallel to bedding plane, consisting of Tentaculites crotalinus
Devonian outcrops of Ponta Grossa city were first analyzed and Styliolina sp., and homalonotid trilobites whole or with only
taphonomically by Bosetti (2004) and Ghilardi (2004). Through thorax or pygidium. Brachiopod valves in convex up position.
high resolution analysis, we have identified two taphofacies T2b: (shells mostly articulated and in life position with complete
sequences. Due to their particular characteristics it was possible to preservation of trilobtes exuviae, complete cephala and torax/
subdivide each taphofacies into two subtypes (Fig. 6) as follows: pygidium). Composed of valves in life position of infaunal lin-
gulids, Australospirifer spp., A. palmata, and Gigadiscina collis,
Taphofacies T1: Composed of disarticulated bioclasts of articulated complete pelmatozoan columns, perpendicular
brachiopods and molluscs, fragmented into tiny fragments. univalves parallel to the bedding plane of T. crotalinus and
These are randomly distributed in the matrix, and the degree of Styliolina sp., and homalonotid trilobites.
packing is dispersed.
T1a: (degree of fragmentation and extensive lateral reworking). The Desvio Ribas-Tibagi outcrop was chosen for further analysis,
Composed of tiny fragments of infaunal lingulids and since it is a well studied section, and has within a thickness of ca.

6. 6 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
Fig. 5. Lithologic columns and palynomorph ranges in Colônia Sutil and Rio Caniú localities. For legend, see Fig. 3.
10 m a considerable variation in lithology, sedimentary structures, indicate anoxic or dysoxic (oxygen-deprived) bottom-waters, occur
and a strikingly rich fossil content. at about the same time as the multiple extinction events in the
During the taphonomic analysis of the other six outcrops in Ponta Middle and Late Devonian (Algeo et al., 1995).
Grossa and Palmeira cities, we noted they share sedimentologic, Globally the sediments above those of the lower upper Emsian
stratigraphic, paleontologic and taphonomic characteristics. The show changes in faunal composition and extinction of certain
Emsian outcrops farther away, e.g. Lambedor, around Tibagi, and groups previously dominant. According to House (2002) none of
Jaguariaíva, in the intervals dated have the same depositional char- the events in the Lower Devonian can be considered as mass
acteristics, as well as the same taphofacies. These indicate similar extinctions and they are all small-scale events. Nor does it neces-
conditions prior to the late Pragian e early Emsian regression. sarily have the same association with the strong pulses of anoxia or
dysoxia that are characteristic of subsequent events (e.g. KǍCÁK
6. The first decline of the Malvinokaffric Realm in the Paraná Event).
Basin The decrease of provinciality in the late Emsian faunas may be
influenced by the contemporary transgressive event (House, 1985).
The global Devonian extinctions were particularly severe for Although the reasons for the extinctions are difficult to diag-
benthic marine organisms in shallow seas (notable exception: nose, the causes are mainly related to a rapid warming or global
Tropidoleptus carinatus, Isaacson and Perry, 1977). In fact, many of cooling (Twitchett, 2005). Still, the rates among low latitude
the taxa that thrived during and after the extinctions were typically inhabitants are most affected, because the dispersal ability is lower
deep-water or high-latitude relatives of the decimated forms. than those inhabiting tropical regions due to their sensitive toler-
Upper Devonian marine deposits are notable in part for the wide- ance of temperature gradients (or presence of paleogeographical
spread occurrence of black shales in the shallow inland seas of barriers). Another important aspect to be analyzed is the tapho-
North America and Eurasia. These organic-rich sediments, which nomic bias. It is known that certain taxa are dependent on

7. E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14 7
Fig. 6. Taphofacies in the Emsian of the ApucaranaSub-basin.
environmental factors such as salinity, temperature, oxygenation, misinterpretations due to possible taphonomic trends as outlined
and that these factors reflect the type of deposit in which they are by Twitchett (2005).
found (except for allochthonous fauna). The resolution of the It is known that marine transgressions are often associated with
vertical distribution is extremely important, because the fauna may a warmer climate, less intense thermohaline circulation, and less
not have been extinct, but influenced by stratigraphic control. oxygenation of deep waters. These phenomena obviously affect the
In the specific case of the Devonian of the Paraná Basin, the benthic fauna, particularly the sessile benthos. On the other hand,
Malvinokaffric realm, which is an assembly typical of these strata, the rapid regression accompanied by storms can disrupt the
has its greatest biodiversity in the basal sections of the sedimentary establishment of temporary biocenoses ecosystems.
succession (uppermost Pragian to lower Emsian), and this occurs Peck et al. (1999), reported that the dynamics of atmospheric
regardless of lithofacies and paleoenvironments (shoreface to circulation in regions with cold climates create conditions that are
offshore). The outcrops dated as late Emsian show a considerable extremely unstable, with strong shifts of air masses. This permits
decrease in biodiversity in relation to older strata. formation of storms. These have a severe destabilizing influence in
This fact is detected by the control of high resolution vertical the upper layers of unconsolidated marine substrate, contributing
distribution of fauna studied by stratigraphic surfaces. Many of the to an increased advection of organisms and subsequent restruc-
characters typically Malvinokafric disappear from the record in all turing of macrofaunal associations and communities.
facies (Tables 1and 2) in the upper Emsian. The Emsian outcrops are In this particular case, these properties occur in taphofacies T1
all indicators of abrupt changes in eustatic sea level, as well as beds, and T2 respectively. The distribution of most benthic macro-
disturbed by storms, as evidenced by the presence of multi-level invertebrates on unconsolidated sea-floor occurs in the form of
structure with HCS possession superior to most of these same patches, and as a result of complex interactions between physical
events in other layers in the same sequence (Figs. 3e5). However, and biological factors. On several unconsolidated sea-floors, for
the influence of storm deposits were less pronounced in the late instance, mosaics of patches can occur, each with a characteristic
Pragian e early Emsian (Sequence B). These phenomena are structure. This phenomenon was recently described for Devonian
probably related to Milankovitch orbital cycles. Zabini et al. (2010) discinids of the Paraná basin (Comniskey, 2011).
and Grahn and Bosetti (2010) characterized these phenomena in The large number of combinations between physical and bio-
the transitional offshore through lingulid fragments and lingulids logical factors can generate this mosaic, which will reflect the
reworked from shoreface, and pebbles of large size re-distributed diversity of benthic organisms and associations. In the present
by high-energy events. case, the patches are shown in taphofacies T2 i.e, where the
Barcellos Popp (1985) and Ciguel (1989) attempted to demon- material is in apparent life position or was not yet fragmented.
strate the biostratigraphic control of megafauna in selected locali- Interspersed with these taphocenoses, which apparently represent
ties, such as the classic outcrops of Jaguariaíva and the urban area of a moment of some environmental stability, is intense bioturbation.
Ponta Grossa. However the results were not conclusive. But in the These events show significant environmental changes, especially
broader analysis of the distribution of macroinvertebrates for the to the fauna with epibenthic and endobenthic suspension feeders
entire Devonian succession outcrops in the state of Paraná, distri- (taphofacies T1).
bution of fauna across strata becomes evident. For that they need to The species of coastal marine environments have two strategies
be considered not only the relative ages of outcrops, but also the for adapting to the regime of instability of the environment: resil-
changes of depositional environments (e.g. Zabini et al., 2010 and ience or resistance. Resilient biota have the capacity to quickly re-
Bosetti et.al., 2011). In the present study occurrences by age and colonize areas disturbed by events of high energy. It is the case of
general depositional environments are verified in order to avoid infaunal lingulids (Bosetti et al., 2010) occurring in our outcrops

8. 8 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
Table 1
Malvinokaffric shelly fossils (Brachiopoda, Cnidaria, Mollusca) in the Paraná Basin.
Phylum Class Family Genus Species
a a a
Brachiopoda Rhynchonellata Cryptonellidae Cryptonella Cryptonella ? baini
a a a
Meristellidae Meristella Meristella septata
a a a
Anoplothecidae Coelospira? Coelospira ? colona
a
Hysterolitidae Australospirifer Australospirifer parana
a
Australospirifer contrarius
Australospirifer antarticus
Australospirifer kayserianus
Australospirifer iheringi
Meganteididae Derbyina Derbyina whitiorum
Leptocoeliidae Australocoelia Australocoelia palmata
Strophomenata Strophochonetidae Australostrophia Australostrophiamesembria
a a a
Chonetidae Notiochonetes Notiochonetes falklandicus
Schuchertellidae Schuchertella Schuchertella agassizi
Schuchertella sulivani
a a
Lingulata Obolidae Lingulepis Lingulepis wagoneri
a a
Discinidae Gigadiscina Gigadiscina collis
Orbiculoidea Orbiculoidea bondenbenderi
Orbiculoidea excentrica
Orbiculoidea baini
Lingulidae Infaunal lingulids Infaunal lingulids
Cnidaria Scyphozoa Conulariidae Conularia Conularia quíchua
Paraconularia Paraconularia africana
a a
Mollusca Tentaculita Tentaculitidae Seretites Seretites jaculus
Uniconus Uniconus crotalinus
a a
Styliolina Styliolina sp.
Homoctenus Homoctenus sp.
a a a
Cephalopoda Orthoceratidae Orthoceras Orthoceras sp.
a a a
Pseudorthoceratidae Spyroceras Spyroceras zoilus
a a a a
Hyolitha Hyolithidae Hyolithes Hyolithes subaequalis
a a a a
Gastropoda Sinuitidae Bucanella Bucanella laticarinata
a a
Ptomatis Ptomatis moreirai
a a a
Pleurotomariidae Pleurotomaria Pleurotomaria (?) kayseri
a a a
Platyceratidae Platyceras Platyceras allardycei
a a a
Bellerophontidae Bellerophon Bellerophon sp
a a a
Plectenotidae Plectonotus Plectonotus hapsideus
a a a
Bivalvia Pterineidae Actinopteria Actinopteria langei
a a
Leptodesma Leptodesma austronotica
a a a
Cetoconchidae Pleurodapis Pleurodapis multicincta
a a a
Grammysiidae Cardiomorpha Cardiomorpha colossea
Prothyris Prothyris knodi
a a
Mytilidae Phthonia Phthonia ? epops
Solemyidae Janeia Janeia Bokkeveldensis
Janeia Braziliensis
Pholamyoidea Edmondia Edmondia sp.
Malletiidae Palaeoneilo Palaeoneilo rhysa
Palaeoneilo magnífica
Paleoneilo sancticrucis
Modiomorphidae Modiolopsis Modiolopsis abbreviata
Modiomorphidae Modiomorpha Modiomorpha scaphula
Malletiidae Nuculites Nuculites reedi
Nuculites sharpie
Nuculites pacatus
Nuculanidae Nuculana Nuculana viator
Crassatelidae Cypricardella Cypricardella oliveira
a a a
Sanguinolitidae Sanguinolites Sanguinolites lagoensis
a
Taxon extinct before the late Emsian.
(undetermined infaunal lingulids and Lingulepis wagoneri), that and taphonomic modes of the fauna. The sandy layers are always
after the storm events recolonized the substrate via re-excavation associated with movement of bioclasts and their fragmentation.
processes (T1). Resistance biota demonstrate a good capacity of Among these storm cycles (Milankovitch orbital cycles), there is
resistance to disturbance, as the case of trilobites (vagrant benthos) a distinct fauna, which probably represents the re-colonization of
and spiriferid brachiopods (T2). the sea-floor. This occurs in mudstones containing endobiont fauna
Resilient and resistent communities alternate in the sedimen- and in siltstones with epibiont fauna. Substrate re-colonization
tary strata studied according to lithologic changes caused by the between the storm events is evident and has three intervals of
constant oscillation of sea level. These oscillations are punctuated faunal succession. However, not always the same taxa are observed
by storm events and were diagnosed in all investigated outcrops (see description of taphofacies T1 and T2 above). The degree of
that are stratigraphically below the black shales from offshore fragmentation of the valves increases sharply near the highest-
facies indicative of the transgressions that characterized the energy events, demonstrating a reworking by waves, and rework-
earliest and late Emsian. ing of previously buried material.
The sections vary in their lithologies, showing changes in sea The observation of these attributes suggests that after these
level. The associated fossil content allows control of the distribution catastrophic events, periods of relative calm occurred long enough

9. E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14 9
Table 2
Malvinokaffric shelly fossils (Echinodermata, Arthropoda) in the Paraná Basin
Phylum Class Family Genus Species
a a a a
Echinodermata Ophiuroidea Encrinasteridae Encrinaster Encrinaster pontis
a a a a
Asteroidea Helianthasteridae Echinasterella Echinasterella darwini
a a a a
Crinoidea Flucticharadae Crenatames Crenatames amicabilis
a
Crenatames sp A
a
Crenatames sp B
a a
Laudonomphalus Laudonomphalus multituberculatus
a a
Marettocrinus Marettocrinus sp. D
a a a
Opsiocrinidae Ophiocrinus Ophiocrinus stangeri
a
Dimerocrinitidae Indet. Indet.
a
Pisocrinidae Indet. Indet.
a a a
Botryocrinidae Costalocrinus? Costalocrinus? sp
a a
Blastoidea Pentremetidae Indet. Indet.
a a a
Nimphaeoblastidae Pachyblastus? Pachyblastus ? sp
a a a
Stylophora Paranacystidae Paranacystis Paranacystis petrii
a a
Anomalocystitidae Australocystis Australocystis langei
a a
Arthropoda Trilobita Calmoniidae Calmonia Calmonia signifer
a
Calmonia subseciva
a
Calmonia michrischia
a a
Paracalmonia Paracalmonia cuspidata
a
Paracalmonia pessulus
a
Paracalmonia mendesi
a
Paracalmonia paranaenses
a
Paracalmonia salamunii
Metacryphaeus Metacryphae
a
Metacryphaeus rotundatus
Pennaia Pennaia pauliana
a a
Dalmanites Dalmanites gonzaganus
a a
?Gamoedaspis ?Gamonedaspis accola
a a
Phacopina Phacopina braziliensis
a a
Acaste Acaste lombardi
Tibagya Tibagya parana
a a a
Homalonotidae Burmeisteria Burmeisteria notica
a
Burmeisteria hershelii
a
Taxon extinct before the late Emsian.
for the fauna to re-colonize the substrate. However, the overlying a sharp decline in the diversity and fauna abundance, with a low
fauna is never the same, which supports this interpretation. degree of disarticulation.
Mudstones and sandstones overlap from top to bottom sections, The intense eustatic sea level oscillations, and the reccurring
where differentiation is observed in fauna and taphonomic mode of storms interspersed with periods of apparent stability of the
fossil concentrations. marine substrate, were probably caused by orbital cycles
The sections also have HCS midsize structures in the sandy responsible for these abrupt changes. Movements of tectonic
strata, where no bioclasts are found. Below and above these, the subsidence due to mantle extension (see Tankard et al., 2009),
bioclasts are totally fragmented and disarticulated, although recorded stratigraphically in the early late Emsian, resulted in
sometimes they can be complete and in apparent life position. This relatively rapid subsidence and sea level rise. During the Early
supports the hypothesis that variations in the relative level radi- Devonian the ParanáBasin was located close to the South Pole
cally interfered in the distribution of fauna and taphonomic mode (Eriksson et al., 2011; Scotese, 2011). The Emsian hiatus in the
on a local scale (Figs. 3e5). basin may have had a duration of up to ca. 7 Ma (Fig. 2; Weddige,
Sections with the presence of tiny fragments of shells, associ- 1996, Deutsche Stratigraphische Kommission, 2002). The reccur-
ated with fragmented valves of infaunal lingulids and bivalves rence of the six recognized taphofacies cycles during the early late
(T1a), record strong bidirectional flows. Immediately above these Emsian transgression would then be over a period of ca. 2.5 Ma,
layers, the lithology is finer and fossils in apparent life position with an estimated variation of the Milankovitch eccentricity
occur in this facies (T2a). Layers of fine to medium sandstone with orbital cycles of ca. 400.000 years. It is known that the cycles of
HCS structure coincide with a high degree of fragmentation and eccentricity are the most striking in regions at higher latitudes,
disarticulation of valves and other invertebrate parts. Tiny frag- which is the case of the ParanáBasin during the late Emsian (De
ments are not found (T1b) in these strata. Thin layers of mudstone Boer and Smith, 1994).
are present in the mid section and present again fossil in apparent The extinctions that occurred between the late Pragian e early
life position (belonging to the same taxonomic groups occurring Emsian regression, and the initial transgression in the early late
beneath). Sandstone layers override this feature and associated Emsian, were extensive and more dramatic than elsewhere on the
reworked, fragmented, and disarticulated bioclasts (T1b). globe. With the exception of the phylum Cnidaria, other phyla of
In all profiles changes of taphonomic attributes parallel the the Malvinokaffric Realm experienced significant casualties.
lithologic changes. Over the upper sandstone layers occur thick, These included the disappearance of 4 classes, 25 families, 41
hard, siltstones, with a unique fauna, and a large number of fossils genera and 54 species. This is the most critical moment in the
in apparent life position (e.g. Australospirifer spp, and A. palmata Malvinokaffric shelly fauna, where 55% of families, 65% of the
and pelmatozoans). This represents a new set of pedunculate genera, and 62% of species went extinct (see Tables 1and 2). These
epifauna and homalonotid trilobites. At the top of the section, dark data, when compared to data presented by Bosetti et al. (2011), are
shale is superimposed by layers of thick siltstone. As is common in more devastating than those that led to the collapse of the clas-
this type of feature, a maximum flooding surface is marked by sical Malvinokaffric shelly fauna. Apparently, when the KǍCÁK

10. 10 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
Fig. 7. Selected late Emsian spores from ApucaranaSub-basin. 1. Acinosporites apiculatus. Rio Caniú, CB 2011-02-10862-2, D61/2. 2. A. apiculatus. Desvio Ribas-Tibagi, CB 2011-02-
10860-2, H63. 3. Acinosporites lindlarensis. Colônia Sutil, BPA 20094542, H52/1. 4. A. lindlarensis? Desvio Ribas-Tibagi, CB 2011-02-10860, M50/1-2. 5. Brochotriletes hudsonii? Rio
Caniú afloramento A, BPA 200402225, N21/3. 6. Apiculiretusispora plicata. Caça e Pesca, MP-P 12A, G44/3. 7. A. plicata. Caça e Pesca, CB 2011-02-10859, L55/1. 8. Archaeozonotriletes
chulus. Desvio Ribas-Tibagi, CB 2011-02-10860, R51/2. 9. Dibolisporites echinaceus. Desvio Ribas-Tibagi, CB 2011-02-10860, P62/1. 10. Dictyotriletes sp. cf. D. richardsonii. Rio Caniú,
MP-P 15A, S46/2. 11. Dictyotriletes sp. cf. D. richardsonii. Colônia Sutil, BPA 20094542, Z51/1. 12. Emphanisporites annulatus. Colônia Sutil, BPA 20094540, H50/1. 13. E. annulatus. Rio
Caniú, CB 2011-02-10862, D39. 14. Emphanisporites mcgregori. Rio Caniú afloramento A. BPA 200402225, N21/3. 15. Emphanisporites rotatus. Colônia Sutil, BPA 20094540, F61/1. 16.
Geminospora svalbardiae. Rio Caniú. CB 2011-02-10862, S54/1. 17. Gneudnospora divellomedia var. minor? Colônia Sutil, BPA 20094540, J46/1.
event occurred (at the EifelianeGivetian transition), the shelly 7. Biostratigraphy of the late Emsian beds in the investigated
fauna had already been strongly affected during the Emsian. It is localities
true that many families still persisted through the Eifelian, but at
present it is not known if another extinction occurred during the Palynomorph dating of samples from the investigated locali-
Eifelian. The fact is that only 7 families, 9 genera and 11 species are
ties in the lower Sao Domingos Formation (Grahn et al., 2011)
recorded after the KǍCÁK Event, and all show subnormal size shows a late Emsian age (upper FD e lower AP spore zones sensu
phenotype (Lilliput Effect; Bosetti et al., 2011). It also discounts Streel et al., 1987, equivalent to the lower GS spore Zone sensu
taphonomic bias, since the same depositional environments Melo and Loboziak, 2003). The absence of spores from the
identified for the interval late Pragian - Emsian are also identified Grandispora/Samarisporites complex in most of the localities
in Middle Devonian layers. To sum up, the first decline of the
indicate a somewhat older age than the AP Zone for the basal Sao
Malvinokaffric shelly fauna in the Emsian, as a result of the events Domingos Formation, which is in agreement with Melo
described above, was lead to the genetic and ecological unsus- and Loboziak (2003) and Grahn et al. (2011). The section at Boa
tainability of the fauna later in the Devonian. The Emsian decline Vista (Figs. 1 and 3) is barren of palynomorphs, but all other
had dramatically increased the vulnerability of the shelly fauna in investigated sections yielded age-diagnostic palynomorphs
relation to environmental changes and the inevitable final (Figs. 1, 3e5). The occurrence of Acinosporites lindlarensis
collapse. and E. annulatus in the sections suggests an age not older than

11. E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14 11
Fig. 8. Selected late Emsian spores and acritarchs from ApucaranaSub-basin. 1. Grandispora sp. Rio Caniú. CB 2011-02-10862, R46. 2. Granulatisporites concavus. Rio Caniú aflor-
amento A. BPA 200402225, W28/1. 3. Retusotriletes paraguayensis. Colônia Sutil, BPA 20094542, O36/2. 4. Synorisporites cf. S. lobatus. Colônia Sutil, BPA 20094542, V35/2. 5. Syn-
orisporites cf. S. lobatus. Colônia Sutil, BPA 20094540, P49/1. 6. Zonotriletes brevivelatus. Desvio Ribas-Tibagi. MP-P 13A, L40/1. 7. Cordobesia oriental. Vendrami, MP-P 11A, P50/1. 8.
C. oriental. Vendrami. CB 2011-02-10858, J53. 9. Duvernaysphaera tenuicingulata. Rio Caniú afloramento A. BPA 200402225, E19/2. 10. Zonotriletes rotundus. Colônia Sutil, BPA
20094542, O41/1. 11. Z. rotundus. Desvio Ribas-Tibagi, CB 2011-02-10860-2, Q61. 12. Navifusa bacilla. Rio Caniú, CB 2011-02-10862-2, C56/2. 13. Palacanthus ledanoisii. Vendrami, MP-
P 11A, G52/3-4. 14. D. tenuicingulata. Colônia Sutil, BPA 20094542, R64/2. 15. N. bacilla. Rio Caniú afloramento B. BPA 200402225, B21/1. 16. P. ledanoisii. Rio Caniú afloramento B. BPA
200402225, Q18/2-4. 17. Pterospermopsis circumstriata. Vendrami, CB 2011-02-10858, N48/3. 18. Triangulina alargada. Rio Caniú, CB 2011-02-10862, Z56. 19. Muraticavea munificus.
Vendrami, MP-P 11A, S34/3.
the base of the serotinus conodont zone (¼upper FD spore Zone). characteristic for the late Emsian in the ParanáBasin. The first
In the Colônia Sutil and Rio Caniú sections (Fig. 5) occur e.g., occurrence of Acinosporites apiculatus is a good indicator for the
Geminospora svalbardiae, Synorisporites cf. S. lobatus, and Zono- base of the AP spore Zone (Melo and Loboziak, 2003), and it is
triletes rotundus, and these species are known from late Emsian - known to range from near the base of Sequence C (¼base of the
early Eifelian assemblages in North Africa (Breuer and Steemans,
Sao Domingos Formation sensu Grahn et al., 2011) in the Paraná
in press). Other diagnostic spore species with a stratigraphic Basin. This species is often associated with A. lindlarensis, Dic-
range from late Emsian are Granulatisporites concavus tyotriletes emsiensis, Emphanisporites rotatus, G. svalbardiae,
and Zonotriletes brevivelatus. Acritarch species as Duvernay- Retusotriletes paraguayensis, and Synorisporites specimens (Grahn
sphaera tenuicingulata and Navifusa bacilla are present and et al., 2010). These spore species are also present in the

12. 12 E.P. Bosetti et al. / Journal of South American Earth Sciences 37 (2012) 1e14
(CNPq no 401796/2010-8). Eduardo Premaor and Paulo Alves de Souza
(Universidade Federal do Rio Grande do Sul, Porto Alegre) are warmly
acknowledged for laboratory work and the making of slides, Peter
Isaacson (Moscow, Idaho) for checking the English, and Pierre Breuer
(Dhahran, Saudi Arabia) and Philippe Steemans (Liege, Belgium) for
comments on the palynomorphs, Our sincere thanks to all.
Appendix A
Miospore species mentioned in text and figures
A. apiculatus (Streel) Streel, 1967 (Fig. 7: 1e2)
A. lindlarensis Riegel, 1968 (Fig. 7: 3e4)
Apiculiretusispora plicata (Allen) Streel, 1967 (Fig. 7: 6e7)
Apiculiretusispora spp.
Brochotriletes hudsonii McGregor and Camfield, 1976 (Fig. 7: 5)
Archaeozonotriletes chulus (Cramer) Richardson and Lister, 1969
(Fig. 7: 8)
C. sextantii McGregor and Camfield, 1976
Dibolisporites echinaceus (Eisenack) Richardson, 1965 (Fig. 7: 9)
Dibolisporites spp.
D. emsiensis (Allen) McGregor, 1973
Fig. 9. Selected late Emsian acritarchs and chitinozoans from ApucaranaSub-basin. 1. Dictyotriletes sp. cf.
Ancyrochitina varispinosa. Caça e Pesca, MP-P 12A, W57/1. 2. A. varispinosa. Caça e D. richardsonii Steemans, 1989 (Fig. 7: 10e11)
Pesca, CB 2011-02-10859, G61/3. 3. Ancyrochitina sp.B? sensu Grahn et al., 2000. E. annulatus McGregor, 1961 (Fig. 7: 12e13)
Metalúrgica Águia, CB 2011-02-10861-2, Q41.
Emphanisporites foveolatus Schultz, 1968
Emphanisporites mcgregorii Cramer, 1966 (Fig. 7: 14)
investigated sections. The only chitinozoan species encountered E. rotatus McGregor, 1973 (Fig. 7: 15)
in this study are Ancyrochitina varispinosa and Ancyrochitina sp. G. svalbardiae (Vigran) Allen, 1965 (Fig. 7: 16)
B? sensu Grahn et al., 2000 (Fig. 9). The former species Geminospora sp.1
range from late Emsian to earliest Givetian (Grahn et al., 2010), Gneudnaspora divellomedia (Chibrikova) Balme, 1988 var. divel-
and the latter is a characteristic late Emsian species (Grahn et al., lomedia Breuer et al., 2007
2000). Gneudnaspora divellomedia (Chibrikova) Balme, 1988 var. minor?
Breuer et al., 2007 (Fig. 7: 17)
Grandispora spp.(Fig. 8: 1)
8. Concluding remarks
G. concavus Breuer and Steemans, in press (Fig. 8: 2)
R. paraguayensis Menéndez and Pöthe de Baldis, 1967 (Fig. 8: 3)
In the Paraná Basin the first decline of the Malvinokaffric Realm
Retusotriletes spp.
is connected to a hiatus (or strongly condensed sedimentation)
R. minutus Tiwari and Schaarschmidt, 1975
from the late Pragian e early Emsian (PoW Su spore Zone) to early
S. praetervisus (Naumova) Allen, 1965
late Emsian (upper FD spore Zone). The initial late Emsian trans-
Synorisporites cf. S. lobatus (Rodriguez, 1978) Breuer and
gression is characterized by six recognized Milankovitch eccen-
Steemans, in press (Fig. 8: 4e5)
tricity orbital cycles with a periodicity of ca. 400.000 years. These
Synorisporites spp.
cycles reflects intense eustatic sea level oscillations, and recurring
V. dubia (Eisenack) Richardson and Rasul, 1978
storms interspersed with periods of apparent stability of the
Z. brevivelatus Breuer and Steemans, in press (Fig. 8: 6)
marine substrate. However, these phenomena were not the cause
Z. rotundus Breuer and Steemans, in press (Fig. 8: 10e11)
of the faunal diversity decline. The number of taxa that disappeared
after the late Pragian e early Emsian regression are more extensive
than commonly documented in other regions of the world Acritarch species mentioned in text and figures
including 4 classes, 25 families, 41 genera, and 54 species. The taxa
that occur during the early late Emsian transgression are less Cordobesia oriental Pöthe de Baldis, 1977 (Fig. 8: 7e8)
provincial than the late Pragian e early Emsian fauna. Diexallophasis remota (Deunff) emend. Playford, 1977
D. tenuicingulata Staplin, 1961 (Fig. 8: 9, 14)
Acknowledgements Hemiruptia spp.
Leiosphaeridia spp.
Elvio P. Bosetti thanks the Conselho Nacional de Desenvolvimento Muraticavea munificus Wicander and Wood, 1981 (Fig. 8: 19)
Cientifico e Tecnológico (CNPq, PQ 401796/2010-8) for financial Muraticavea? spp.
support, Sandro Scheffler (UIFESP, São Paulo) for echinoderm N. bacilla (Deunff) Playford, 1977 (Fig. 8: 12, 15)
taxonomy, Renato Pirani Ghilardi (UNESP, São Paulo) for trilobite Palacanthus ledanoisii (Deunff) Playford, 1977 (Fig. 8: 13, 16)
taxonomy, and William Mikio Kurita Matsumura (UFRGS, Ponta Pterospermopsis circumstriata Jardiné et al., 1972 (Fig. 8:17)
Grossa) for field assistance and discussions. Yngve Grahn acknowl- Pterospermopsis spp.
edges economical support for field-work (CNPq no 401796/2010-8), Tasmanites spp.
Rodrigo Scalise Horodyski a grant from CNPq (141256/2010-9), and Triangulina alargada (Cramer) Playford, 1977 (Fig. 8: 18)
Paula Mendlowicz Mauller from CAPES (Coordenação de Aperfei- Triangulina spp.
çoamento de Pessoal de Nível Superior, BEX 4515/05-6) and CNPq Veryhachium spp.

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