Bosetti et al. 2011 publicado em : Paläontologische Zeitschrift

RESEARCH PAPER

An earliest Givetian ‘‘Lilliput Effect’’ in the Parana Basin,and the collapse of the Malvinokaffric shelly fauna

Elvio Pinto Bosetti • Yngve Grahn • Rodrigo Scalise Horodyski •

Paula Mendlowicz Mauller • Pierre Breuer •

Carolina Zabini

Received: 22 December 2009 / Accepted: 16 June 2010 / Published online: 2 July 2010

� Springer-Verlag 2010

Abstract An earliest Givetian ‘‘Lilliput Effect’’ at Sıtio

Wolff and Sao Bento in the Parana Basin occurred after an

extinction event, possibly related to the latest Eifelian

KACAK Event. The Malvinokaffric fauna was reduced

from 65 genera before the extinction event to eight genera

after the event. However, the abundance of the individual

taxa is high. The size reduction of the surviving taxa was

up to 90%. The palynomorphs during the extinction and

post-extinction (‘‘Lilliput Effect’’) events are scarce. Nor-

mal palynomorph abundance and diversity is restored later

in the early Givetian immediately after the post-extinction

event. The relictual fauna in the beds with the ‘‘Lilliput

Effect’’ at Sıtio Wolff and Sao Bento constitute the last

survivors of the classical Malvinokaffric shelly fauna in the

Parana Basin, and are at these sites mixed with immigrants

and alien elements (e.g. orthoconic nautiloids).

Keywords ‘‘Lilliput Effect’’ � Earliest Givetian �KACAK Event � Malvinokaffric fauna � Parana Basin

Kurzfassung Im Gebiet von Sıtio Wolff und Sao Bento

des Parana-Beckens tritt im fruhesten Givetium ein ,,Lili-

put-ffekt’’ auf, der einem Aussterbeereignis folgt, das

moglicherweise Bezug zum KACAK-Event des spaten

Eifeliums hat. Die Malvinokaffrische Fauna wurde von 65

Gattungen vor dem Aussterbeereignis auf acht Gattungen

danach reduziert. Dennoch bleibt die Haufigkeit individu-

eller Taxa hoch. Die Großenreduktion uberlebender Taxa

erreicht 90%. Weiterhin sind Palynomorphen wahrend des

Aussterbeereignisses und des ,,Liliput-Effekts’’ selten.

Normale Haufigkeit und Diversitat erreichen die Palyno-

morphen erst spater im fruhen Givetium, unmittelbar

nach dem Auftreten des ,,Liliput-Effekts’’. Die Reliktfauna

aus den Lagen mit ,,Liliput-Effekt’’ im Gebiet von Sıtio

Wolff und Sao Bento beinhaltet die letzten Rudimente

der klassischen Malvinokaffrischen Schalen-Fauna im

Parana-Becken. Sie vermischt sich hier mit eingewanderten

Formen und exotischen Elementen (z. B. orthoconen

Nautiloiden).

Schlusselworter ,,Liliput-Effekt’’ � fruhestes Givetium �KACAK-Event �Malvinokaffrische Fauna � Parana-Becken

E. P. Bosetti

Laboratorio de Paleontologia, Universidade Estadual de Ponta

Grossa, Rua Otaviano Macedo Ribas 164, Ponta Grossa,

PR 84070-540, Brazil

e-mail: [email protected]

Y. Grahn (&) � P. Mendlowicz Mauller

Faculdade de Geologia, Universidade do Estado do Rio de

Janeiro, Bloco A, Sala 4001, Rua Sao Francisco Xavier 524,

Rio de Janeiro, RJ 20550-013, Brazil


P. Mendlowicz Mauller


R. S. Horodyski � C. Zabini

Programa de Pos-Graduacao em Geociencias,

Universidade Federal do Rio Grande do Sul, Av. Bento

Goncalves 9500, BL L Predio 43113, Campus do Vale,

Porto Alegre, RS 91509-900, Brazil


C. Zabini


P. Breuer

Saudi Aramco, Geological Technical Services Division,

Dhahran 31311, Saudi Arabia


123

Palaontol Z (2011) 85:49–65

DOI 10.1007/s12542-010-0075-8

Introduction

The term ‘‘Lilliput Effect’’ was introduced by Urbanek

(1993) to describe the size changes of faunas in extinction

events. In the aftermath of biotic crises, the organisms tend

to be much smaller than before the crises. According to

Twitchett (2006), this effect is one of the most widespread

evolutionary phenomena, but is virtually unstudied. Body

size is a key element in animal evolution, and many

paleontologists have observed that organisms that survived

mass extinctions often have a much smaller body size than

their predecessors. There are many reason why organisms

shrink, including drastic environmental changes (e.g.

effects of volcanic activity, increasing competition). By

definition, size reduction of individual taxa from pre-

extinction to immediate post-extinction is the ‘‘Lilliput

Effect’’ sensu stricto of Urbanek (personal communication

to E.P.B. 2009). A drastic reduction of body size was

observed in all megafossil taxa from the earliest Givetian

part of the Sao Domingos Formation. This interval gener-

ally contains few body fossils, and all of them are reduced

in size. The fossils are represented by conulariids, trilo-

bites, nautiloids, and brachiopods (rhynchonellids, disci-

nids, and lingulids). Additionally, ostracods, plant

fragments, and ichnofossils are preserved. Palynomorphs

are scarce, which is unusual for Middle Devonian strata in

Western Gondwana. The scarcity of acritarchs and chitin-

ozoans is probably because of the geochemical annexation

of carbon and nutrients such as nitrogen and phosphorus

during the black shale formation that characterized the

KACAK Event (cf. Riegel 2008), thus tending to deprive

the marine palynomorphs of essential food sources. Given

the normal progressive reduction in quantities of terrestrial

spores transported seawards from the shoreline, their rarity

in the investigated (distal shoreface) strata is not unex-

pected. In comparison with the dimensions of a typical

Malvinokaffric fauna, the sizes of the post-extinction fos-

sils are reduced by up to 90%. One taphonomic aspect that

may affect the size of the fossils in an assemblage is

hydrodynamic sorting. This is probably not the case

because of the low degree of fragmentation. Small ichno-

fossils are also present, corroborating the in situ size

reduction. The specimens here described are adult organ-

isms, and their preservation indicates autochthony or pa-

rautochthony conditions (Kidwell and Bosence 1991).

Materials and methods

The paleontological material was collected from the Sao

Domingos Formation at Sıtio Ari, Sıtio Wolff, Sao Bento,

and Casa de Pedra (Fig. 1). The material (DEGEO/MPI-

3230 to DEGEO/MPI-3930) is deposited at Universidade

Estadual de Ponta Grossa (Paleontology Laboratory of the

Geosciences Department). Approximately 700 samples

were analyzed, each sample displaying one or more fossils.

The megafossils were processed by means of fine brushes

and needles. The microfossils were processed with Petro-

bras standard methods (Quadros and Melo 1987). The

palynomorph material is housed in the collections of the

Biostratigraphy and Paleoecology Management of Petro-

bras Research Center, Petrobras/Cenpes/Pdexp/Bpa, Ilha

do Fundao, Rio de Janeiro, RJ, Brazil (BPA).

Geologic setting

The Parana Basin (Fig. 1a) is classified as a polycyclic

intracratonic and intercontinental basin. In Brazil it is

represented by two sedimentary depocenters, i.e. the

Fig. 1 a Map showing the localities discussed or investigated in this

study. b Detail of the area studied. The dotted line shows the

administrative city limits of Tibagi. c Sample points in the Barreiro

section (localities 1–3) and at Sao Bento (locality 4)

50 E. P. Bosetti et al.

123

northern Alto Garcas and southern Apucarana Sub-basins.

The total extent of the Parana Basin is ca. 1,600,000 km2

and includes parts of southern Brazil, eastern Paraguay,

central Uruguay, and northeastern Argentina. Lange and

Petri (1967) formalized the Devonian lithostratigraphic

succession based on the geology of the Apucarana Sub-

basin. Their proposition was, in ascending order, Furnas

Formation and the tripartite division of the Ponta Grossa

Formation, viz., the Jaguariaıva, Tibagi, and Sao Do-

mingos Members. More recently, Grahn (1992), Gaugris

and Grahn (2006), and Grahn et al. (2000, 2002, 2010)

subdivided the Devonian of the Apucarana Sub-basin into

three formations, in ascending order, Furnas, Ponta

Grossa (including the Tibagi Member), and Sao Domin-

gos. This nomenclature will be adhered to in this paper.

The Sao Domingos Formation is dated as latest Emsian—

late early Frasnian with miospores (Loboziak et al. 1988;

Melo and Loboziak 2003; Mendlowicz Mauller 2008) and

chitinozoans (Grahn et al. 2000, 2002, 2010; Gaugris and

Grahn 2006; Mendlowicz Mauller et al. 2009). The

‘‘Lilliput Effect’’ occurs at Sıtio Wolff and Sao Bento

(Fig. 1b–c) close to the top of the outcropping part of the

Sao Domingos Formation, which is dated as early Give-

tian at the Casa de Pedra locality (Fig. 1b–c). It is pre-

ceded by the rocks at Sıtio Ari (Fig. 1b–c), dated as no

older than latest Eifelian and near the Eifelian/Givetian

boundary from acritarch evidence. The Barreiro region,

situated in the Sao Domingos district, Tibagi (Fig. 1b–c),

is the type–area of the Sao Domingos Formation (Bodziak

and Maack 1946). With the exception of Petri (1948),

Lange and Petri (1967), Melo (1985), Bosetti (2004), and

Bosetti et al. (2009), the references to the type-area are

rarely based on field data from the region. However, the

Sao Domingos shales reach their maximum surface

thickness, ca. 90 m, in the type-area itself (Bodziak and

Maack 1946; Lange and Petri 1967). According to Melo

(1985), these shales overlap the Tibagi Member sensu

stricto in virtually all its extent. The total thickness of the

Sao Domingos Formation is estimated at ca. 350 m (Melo

1988; Grahn et al. 2010).

Outcrops in the region vary in thickness between 1 and

25 m, and are commonly penetrated by diabase dikes.

This makes continuous stratigraphic sampling over geo-

graphic distances difficult. However, similar lithologies

are exposed along the Tibagi–Telemaco Borba highway

PR-340 (Bergamaschi 1999; Grahn et al. 2000, 2002), and

in a road-cut at km 424 on highway BR-376 in the Imbau

region (Bergamaschi and Pereira 2001); these enable

correlation of facies with the Barreiro section. Further-

more, a section with a ‘‘Lilliput’’ fauna followed by a

normal-sized fauna dated as earliest Givetian is known

from Sao Bento at km 280 along highway PR-340 (see

below).

Localities

Sıtio Ari (24�31021.7300S,50�28010.1700W)

Sıtio Ari is situated in the Barreiro region, Sao Domingos

district, Tibagi (Fig. 1b–c). This outcrop is located in the

basal part of the Barreiro section (Fig. 2) and in the Sao

Domingos Formation. The exposed thickness is ca. 10 m.

At the base occur cm-thick lenses of very fine to fine-

grained sandstone, and in the uppermost part siltstones with

lingulids. These siltstones also contain rare palynomorphs

(listed in Appendix 2). Of these, Chomotriletes vedugensis

indicates an age not older than latest Eifelian.

Sıtio Wolff (24�3304200S,50�3100000W)

Sıtio Wolff is situated in the Barreiro region near Salto Santa

Rosa waterfall, Sao Domingos district, Tibagi (Fig. 1b–c).

The locality corresponds to the middle part of the Barreiro

section (Fig. 2) and is within the Sao Domingos Formation.

The exposed thickness is 20.5 m (Figs. 2 and 3). At this site

bioclasts that represent the Malvinocaffric Realm with sub-

normal phenotypes are found (Bosetti et al. 2009). The Sıtio

Wolff outcrop is stratigraphically above the medium to

coarse-grained fossiliferous sandstones at Sıtio Ari referred

to interval D3 by Lange (1967), corresponding to latest

Emsian–Eifelian strata in the Apucarana Sub-basin. The silty

shales, ca. 5–7 m below the top of the section, contain (in

decreasing abundance order) ichnofossils (Phycosiphon),

Spongiophyton fragments and other unidentified plant

remains, conulariid fragments, discinid brachiopods,

calmoniid trilobites, rhynchonellid brachiopods, ostracods,

orthoconic nautiloids, lingulid brachiopods, and bivalves.

The siltstones in the uppermost 5 m of the section contain

(also in order of decreasing abundance) ichnofossils (Phy-

cosiphon), Spongiophyton fragments and other unidentified

plant remains, orthoconic nautiloids, calmoniid trilobites,

discinid brachiopods, conulariid fragments, ostracods, and

lingulid brachiopods (Fig. 3).

A distinct feature of all bioclasts found in these beds is

their small dimensions when compared with the same taxa

in other facies of the Devonian sequences. Despite their

reduced size, all taxa collected are in an advanced onto-

genetic stage, and therefore represent adult forms. As they

are not randomly distributed in the section (Figs. 2 and 3),

the vertical distribution of the bioclasts is controlled by

lithologic variation. The identified fossils are listed in

Appendices 1 and 2.

Sao Bento (24�28011.2100S, 50�32008.4600W)

Sao Bento is situated at km 280 along highway PR-340

(Fig. 1b–c). The lower 30 m of this road-cut section

Earliest Givetian ‘‘Lilliput Effect’’ in the Parana Basin 51

123

contain a ‘‘lilliput fauna’’ (mainly conulariids) mixed with

normal-sized fossils in the upper part of the interval. The

uppermost 10 m of the section feature normal-sized

brachiopods and trilobites, and ‘‘lilliput’’ bioclasts are less

common. The total exposed thickness is ca. 40 m (Fig. 4).

A siltstone sample, ca. 21.5 m above the base of the

Fig. 2 Barreiro section with

taxonomic and taphonomic

distributions


123

section, yielded a characteristic earliest Givetian pal-

ynomorph assemblage. This indicates that the post-

extinction event with the ‘‘Lilliput Effect’’ was short-lived

and related to the initial Givetian transgression in the Pa-

rana Basin. The identified palynomorphs are listed in

Appendix 2.

Casa de Pedra (24�3400000S, 50�3005500W)

Casa de Pedra (Fig. 1b–c) is located in the Sao Domingos

Formation at the top of the Barreiro section (Fig. 2). The

exposed thickness is ca. 7.5 m. The base consists of ca.

1 m of dark gray, very fine-grained siltstone, with angular

multi–faceted pebbles. These are overlain by 4.5 m of

medium to coarse-grained siltstone with intercalated

sandstone lenses showing distinct hummocky cross strati-

fication. The sandstones seem to contain pebbles of the

same type as in the basal siltstones. Towards the top are

claystones and siltstones. The contact with the Pennsyl-

vanian beds is discordant. The palynomorphs (listed in

Appendix 2) are characteristic of the early Givetian and

display a normal abundance and diversity. Normal–sized

Lingulida spp., Pennaia pauliana and ichnofossils occur at

the locality.

The Malvinokaffric realm

The term ‘‘Malvinokaffric’’ was introduced by Richter

(1941) to replace the inappropriate term ‘‘austral’’ of

Clarke (1913). The term ‘‘austral’’, which defined the

Devonian forms in South America, became inadequate

because it implied that all Southern Hemisphere Devonian

faunas had an exclusively austral paleobiogeographic

character. This is not the case, and the morphological

characteristics of the euro-asiatic (boreal) faunas of New

Zealand and Australia were already known.

Clarke (1913) considered not only the trilobites but

also a faunistic set that characterized a vast Southern

Hemisphere region, and included different brachiopods

and other invertebrate groups. The derivation of the term

(Malvinocaffrische) came about with the reunion of two

regions of occurrence of Clarke’s (1913) austral fauna:

the Falkland (Malvinas) Islands and Cape Province (South

Africa). Richter and Richter (1942) concluded that the

Malvinokaffric Realm constituted one faunistic unit, as

Clarke (1913) had already mentioned. However, they

stressed through comparisons with the Northern Hemi-

sphere fauna that this paleobiogeographic realm had been

established during the Devonian, under constant migratory

exchanges between the Malvinokaffric austral and boreal

seas. Clarke (1913) conceived that the two isolated faunas

experienced a parallel development since the Silurian

(Bosetti 2004).

The Parana Basin is a center of the Malvinokaffric

Realm in South America. This realm developed essen-

tially in the Southern Hemisphere (South America, Ant-

arctica, South Africa, and Ghana) during the Early

Devonian and Eifelian. In contrast with the contempora-

neous zoogeographical entities that dominated the shallow

seas with warmer water in the Northern Hemisphere and

Oceania, the Malvinokaffric was characterized by a low

faunistic diversity, with few taxa represented by numerous

individuals having extensive regional distribution (Shirley

1965).

The Early Devonian–Eifelian strata in the Parana Basin

are characterized by the Malvinokaffric fauna. The entry of

Fig. 3 a Occurrence and distribution of bioclasts in the different

lithologies of the Sıtio Wolff outcrop. b Relative abundance of the

bioclasts in the Sıtio Wolff outcrop


123

the extra-Malvinokaffric articulate brachiopod Tropido-

leptus, in regions previously dominated by Malvinokaffric

forms, is considered by most geologists to represent the

irreversible decline of the faunistic realm defined by the

Malvinokaffric shelly fauna.

The Malvinokaffric extinction

The causes of the extinction of the Malvinokaffric shelly

fauna in Western Gondwana is a controversial matter

involving geochronology and physical environmental fac-

tors. Copper (1977) stated that the mass extinction of the

fauna occurred at the Frasnian–Famennian boundary. This

assertion was founded on the hypothesis that an extremely

cold climate would have caused recifal and peri-recifal

faunal extinction. Isaacson (1978) agreed with Copper that

the extinction resulted from a major marine regression at

the end of the Devonian. According to Melo (1985), there

were no records of Malvinokaffric forms in the upper

part of the Sao Domingos Formation, but he conceded a

temporal expansion of the fauna (particularly trilobites,

albeit uncommonly) into the Givetian. Assine and Petri

(1996) confirmed that the transgression at the Eifelian–

Givetian transition led to a drastic ecological change that

was responsible for the disappearance of the Malvinokaf-

fric Realm. This conclusion is corroborated by this study

and is discussed below.

Although the magnitude of Devonian extinctions is

widely accepted, the duration, number, and causes of these

events remain controversial: in particular, whether an event

should be considered as a prolonged extinction, or two

separate events, or a series of events.

Most geologists now regard the Frasnian–Famennian

extinction as being the most significant during the Devo-

nian. However, House (2002) stressed that some other

Devonian crises were at least comparable with the F/F

Event, and he pointed out that the Eifelian–Givetian

(KACAK) Event might have been the most striking

extinction.

Most of these events are diagnosed by lithofacies

and, according to House (2002), the 20 middle Paleozoic

Fig. 4 Sao Bento section with

taxonomic and taphonomic

distributions


123

extinction events share many common characteristics.

They are usually characterized by dark-colored sedimen-

tary facies (indicative of dysoxia or anoxia), hosting a

depauperate or no benthic fauna, and are frequently under

and overlain by regressive facies.

Many of these sediments are associated with faunal

extinction. The primary cause of dysoxia or anoxia is still

much debated, but this phenomenon has a clear short-term

effect on contemporary sea beds. According to House

(2002), many events show complex and progressive

changes in oxygenated conditions during the Devonian,

following a common pattern and probably also having a

common cause. These changes may have been induced by

climatic and consequential cycles of erosion or fluctuations

in sea level (eustatic or epeirogenic). Related cyclic climate

changes forced by orbital cycles (e.g. Milankovitch) may

also be implicated.

Most of the extinction events recorded in the Devonian

are associated with a transgression followed by a regressive

phase (House 2002 and references therein). These events

are usually manifested globally, but local or regional

variations of this pattern cannot be disregarded.

It is unclear if the event represented in the Sao Do-

mingos Formation was a consequence of one of these

global events. However, the studied sequence does show

sedimentary characteristics in accordance with the general

regression/transgression model whereby shallow water

ecosystems would be quickly replaced by anoxic deep

water ecosystems. The black shales characterizing parts of

the Sao Domingos Formation are accepted as maximum

flooding surfaces (Lange and Petri 1967; Melo 1985, 1988;

Assine and Petri 1996; Assine 1996; Bergamaschi 1999;

Bergamaschi and Pereira 2001; Bosetti 2004), thus indi-

cating dysoxic/anoxic conditions during the maximum

marine inundation of the basin.

Bosetti (2004) used a high resolution collecting

taphonomic method and described in some detail part of

the Sao Domingos Formation in the Barreiro region

(Tibagi), and new finds of calmoniid trilobites, conular-

iids, and rhynchonellid brachiopods (Schuchertella, Aus-

tralocelia, and Derbyina). All of these were supposedly

extinct at this time of deposition. Bosetti (2004) also

indicated that the Malvinokaffric fauna extended beyond

the Eifelian–Givetian boundary, reaching into the Give-

tian, and without apparent modifications in its paleobi-

odiversity. However, later finds by Bosetti et al. (2009)

in the same region demonstrated that only a few Malv-

inokaffric taxa transgressed this boundary and that these

taxa are represented by phenotypes of subnormal

dimensions compared with the typical representatives of

the Malvinokaffric fauna. Therefore, this Sao Domingos

shale fauna can be regarded as a relictual Malvinokaffric

assemblage.

It is generally believed that the Parana Basin exposures

of late Middle–Late Devonian strata are poorly fossilifer-

ous, and that the major proportion of the invertebrates

(especially cnidarians, trilobites, and articulate brachio-

pods) occurring in the Ponta Grossa Formation sensu

stricto had become extinct before the early Givetian.

However, this alone does not explain the absence of the

paleofauna, because even in the lower sequences there are

records of other trangressive peaks of equal intensity, and

with an abundance of invertebrates. In the early Givetian,

after the collapse of the classic Malvinokaffric shelly

fauna, the effect of warm water faunas with an Appala-

chian affinity is obvious in the Amazonas Basin (Melo

1988; previously noted by Rathbun 1874). The situation is

somewhat different in the Parana Basin, where a calmoniid

trilobite (i.e. Pennaia pauliana) ranges into the early Gi-

vetian after the collapse of the classic Malvinokaffric

shelly fauna.

Relictual assemblage in the Sao Domingos formation

The appearance of relictual fossil assemblages is common

in global paleontological records, especially during

immediate post-extinction events (e.g. latest Ordovician

Event; Ireviken Event, Silurian of Sweden, Erlfeldt 2006;

Trangrediens Event, Siluro—Devonian boundary in East-

ern Europe, Urbanek 1993; Late Devonian (Frasnian/

Famennian) Event and Permo—Triassic Event, Twitchett

2007; Late Triassic Event; K/T Event, etc.). Urbanek

(1970) defined relictual assemblages as sets of low-diver-

sity species or monospecific occurrences that survived

environmental disturbances in a given area. It is the

immediate effect of the extinction, associated with each

biotic crisis that leads to a drastic reduction of the number

of species as a result of ecological change. Such changes

open possibilities for new species to occupy the affected

area via speciation or immigration. Once the dispersion of

the species has been established, the local area is rapidly

recolonized (Krebs 1986).

Urbanek (1993) stated that in some cases the relictual

assemblages exhibit attributes of post-event syndrome, for

example extremely low diversity, high abundance of indi-

viduals, and subnormal phenotype like reduction in size.

They represent a brief delay in the evolutionary changes of

a specific fauna, before adaption to new environmental

conditions.

The fossil assemblages studied here can be regarded as

representatives of a relictual assemblage because, as typi-

cal Malvinokaffric fauna, they show low taxonomic

diversity and great abundance in the Sıtio Wolff and Sao

Bento outcrops. The distinctive factor of this new assem-

blage is that the diversity is even lower than that


123

characterizing the typical Malvinokaffric endemic fauna,

and the abundance levels of each taxon are high. The fact

that the fossil adult forms found in these localities are

much smaller than normal, and that alien elements (or-

thoconic nautiloids) are present in these beds (as likely

immigrants) suggest that the conditions inferred by Ur-

banek (1970, 1993) were established at these localities

during the Eifelian–Givetian transition. In the post-crisis

phase two lines of development in the relictual assem-

blages are envisaged:

1. A great numerical abundance creates favourable

conditions for generating a sufficient variation (‘‘raw

material’’ later used in adaptive radiation), and some

expansion of the niche is also involved, because of

both the abundance and increased variation of the

relictual species (Urbanek 1993). These factors

obstruct, at least in part, the invasion of empty habitats

by immigrants, thus facilitating rapid local speciation

of native elements. In a relatively short time, the main

niches available are inhabited again by new settlers,

hindering the possibility of entry of alien forms. These

latter are thus restricted but not necessarily absent

(Urbanek 1993).

2. When a delay of the evolutionary response of the

indigenous relictual species does not show distinct

post-event syndrome, and, especially, no sign of a

population explosion, the habitats remain sufficiently

empty to eventually be colonized by immigrants. The

delayed response to ecological change could create an

opportunity for rapid habitation by immigrants (Ur-

banek 1993).

In the Sao Domingos Formation, the faunal composi-

tion, distribution, and abundance indicate the first situation,

where the relictual assemblage itself occupies a substantial

part of the benthic niches; and the only alien element of the

assemblage, the immigrant orthoconic nautiloids, occupy

the newly created pelagic niches.

The lithologic variation observed in the study area

was produced by pronounced eustatic variations, con-

sidering the modest lithologic variation in major parts of

the Ponta Grossa and Sao Domingos formations. These

strata signify a succession of paleoenvironments; i.e. a

pattern of coarse sediments, at shoreface, to fine offshore

sediments, according to Walker and Plint (1992) and

Reading and Collinson (1996). The paleoenvironments

progress from proximal shoreface to distal offshore. The

base is represented by a coarse regressive phase with the

occurrence of conglomeratic (quartzite and quartz peb-

bles, maximum diameter ca. 1 cm), very coarse-grained

unfossiliferous sandstones (6 m thick at Sıtio Wolff)

suggestive of a proximal shoreface environment. Above

this facies is a succession of fine-grained sandstone to

medium-grained siltstone (2 m thick at Sıtio Wolff) with

ichnofossils (Phycosiphon) and fragmented plants (mm to

cm-sized, parallel to the bedding planes); this culminates

with an argillaceous dark shale (5 m thick at Sıtio Wolff)

lacking benthic fossils and indicating a distal offshore,

retrogradational environment, below storm wave base

(SWB). Towards the top a conglomeratic sandstone bed

(maximum thickness 0.5 m at Sıtio Wolff), similar to the

basal one represents a sudden recurrence of the shore

facies (proximal shoreface). Above this sequence, silty

shales (2 m at Sıtio Wolff) are covered by coarse to fine-

grained siltstones with wavy stratification (5 m at Sıtio

Wolff), similar to distal shoreface environments near fair

weather wave base (FWWB). These latter two facies are

abundantly fossiliferous, and contain subnormal-size

phenotypes. The distribution and abundance of the fossils

in the section are not random; they are linked to the

depositional tracts recognized by the sedimentary facies

analysis.

At Sıtio Wolff at least three main bathymetric chan-

ges are recognizable, reflecting changes in temperature

and oxygenation. The base of the section represents

the shore facies and the absence of bioclasts can be

explained taphonomically, whereby the preservation of

fossils in conglomeratic coarse-grained sandstones is

less probable. The middle part represents a brief interval

of maximum flooding; i.e. at the extreme peak of the

local transgression. The fact that no megafossils or

ichnofossils occur in this facies is evidently a conse-

quence of anoxia or extremely low oxygenation of the

sea floor.

The post-event syndrome would be expected in the

upper part of the section (distal shoreface environment)

where the bioclasts represent a phenotypic reaction to

conditions unfavourable for growth of individuals (conu-

lariids, brachiopods, and molluscs) and the natural selec-

tion of specimens and species of smaller size (ichnofossils

produced by small organisms, ostracods, and trilobites).

The conditions envisaged by Urbanek (1970, 1993) are

exemplified by these facies of the Sao Domingos

Formation.

To conclude, the stratigraphic intervals investigated here

are considered to have been deposited during the collapse

of the Malvinokaffric shelly biota, and the fauna can

therefore be considered as relictual with subnormal-size

phenotypes. This distinctive fauna clearly exemplifies the

post-event syndrome. Brayard et al. (2010) studied Early

Triassic gastropods in the aftermath of the Permian–Tri-

assic mass extinction. They found that large specimens had

already developed some 1–2 Ma following the mass

extinction, and concluded either that the lilliput effect was

an artifact (or not particularly significant), or that the post-

extinction recovery in the marine realm was rapid, in the


123

order of 1–2 Ma. Our results suggest that the recovery after

the KACAK Event was indeed rapid and probably less than

1 Ma.

Taphonomic considerations

The taphonomic analysis aimed primarily at determining

the following attributes:

1. the degree of valve fragmentation;

2. the degree of disarticulation of valves and component

parts;

3. the position of valves and parts in relation to bedding

planes;

4. the effects of bioerosion;

5. the effects of abrasion;

6. the degree of packing; and

7. distribution and abundance levels (Appendix 1).

The conulariids are very fragmented, whereas brachio-

pods and molluscs are intact, with the exception of one,

Fig. 5 Conulariids (a–g) and brachiopods (h–j) from Sıtio Wolff. aConularia quıchua. MPI 3697. b Paraconularia ulrichana. MPI 3654.

c Paraconularia ulrichana with plant fragments. MPI 3751. dParaconularia ulrichana. MPI 3772. e Paraconularia ulrichana. MPI

3401. f Paraconularia ulrichana with plant fragments. MPI 5678. gParaconularia ulrichana. MPI 3707. h Schuchetella agassizi. MPI

3664. i Australocoelia palmata. MPI 5679. j Australocoelia palmata.

MPI 3599


123

partly fragmented specimen of Schuchertella agassizi Hartt

1874. Trilobites are mostly articulate (entire specimens and

articulated thorax/pygidium), isolated pygidia and cephala

being incommon.

Apart from Spongiophyton, the plant fragments are

difficult to classify. There are no bioclasts with evidence of

bioerosion and abrasion, and the bioclasts are poorly

packed and matrix-supported.

Fig. 6 Bivalves (a) and trilobites (b–e) from Sıtio Wolff. a Nuculana? viator. MPI 3535. b Thorax of Pennaia pauliana. MPI 3818 c Pygidium

of Pennaia pauliana. MPI 3662. d Cephalon of Pennaia pauliana. MPI 3748–B. e Pennaia pauliana. MPI 3367–A


123

Fig. 7 Brachiopods (a–d), nautiloids (e–f), and ichnofossils (g–h)

from Sıtio Wolff. a Orbiculoidea baini, Brachial valve uncompressed.

MPI 3669. b Orbiculoidea excentrica (1– ventral valve; 2– brachial

valve). MPI 3663. c Lingulid indet in ‘‘scissor’’–position. MPI 3582.

d Lingulid indet. MPI 3769–B. e ?Ctenoceras sp. MPI 5680. f?Ctenoceras sp. MPI 5681. g Phycosiphon isp. MPI 3666. hPhycosiphon isp. MPI 5682


123

Morphological variations and apparently subnormal size

phenotypes must be carefully analyzed, because the fos-

silization processes, especially diagenetic factors, can

affect the original morphology of the bioclasts. Lucas

(2001) introduced the term ‘‘taphotaxa’’, referring to taxa

based on morphological characters produced by the fos-

silization processes. The identification of invalid taxa in the

Ponta Grossa Formation (conulariids and trilobites) by

Simoes et al. (2003) and Soares et al. (2008), demonstrates

that diagenetic and weathering alterations can modify the

original structure of fossils, thus leading to erroneous tax-

onomic assignments.

The subnormal-sized phenotypes have been described

above and compared with those of normal size that occur in

other facies; and taphotaxa have not been found in the

analyzed concentration, because morphological alterations

linked to taphonomic characteristics have not been

observed. Moreover, all the fossils display growth lines,

striae, and morphological features of the carapaces and

exuviae indicative of advanced ontogenetic growth.

Twitchett (2007) stressed that, in relation to body size

of specimens, some preservational aspects can affect the

totality of the fossils in an assemblage, and the hydrody-

namic selection of bioclasts (by some kind of water flux) is

one of the factors to be considered in search of taphonomic

bias.

In this study, entire fossils and fragments of variable

size occur in the same sample and on the same bedding

plane. Fragile plant fragments are intimately associated

with entire trilobite carapaces and minute orbiculoid

valves, all of them of different sizes and densities, but

occurring side by side. This suggests that, if transportation

of bioclasts had occurred, it was insufficient to cause

selection by size or density.

Taphonomic classes linked to paleobathymetry and to

autochthony/parautochthony/allochthony factors were pro-

posed by Rodrigues et al. (2003) based on conulariids, and

by Simoes et al. (2009) based on homalonotid trilobites.

The conulariids (Fig. 5) are represented at Sıtio Wolff by

isolated, horizontally oriented and incomplete specimens

without apertural parts and are attributable to taphonomic

class 3-IV of Rodrigues et al. (2003). They occur in bio-

turbated siltstones (bioturbation index 3 of Miller and

Smail 1997) at the FWWB. The taphonomy of the conu-

lariids indicates that the taphocoenosis is parautochthonous

to allochthonous.

In relation to the degrees of disarticulation and frag-

mentation, the basic, distinctive morphologies of the

diverse taxonomic groups was considered. All rhyncho-

nellids and bivalves (Figs. 5, 6) are disarticulated and

concordant with the bedding planes, thus suggesting a

time interval between death and ultimate burial of the

bioclasts, but without hydrodynamic selection. Regarding

the trilobites (Fig. 6), three situations are evident (in

descending frequency):

1. the predominant occurrence of complete exuviae

(extended skeleton);

2. the occurrence of articulated thorax/pygidium; and

3. subordinate occurrence of isolated pygidia, and more

rarely isolated cephala.

Overall, this points to autochthony/parautochthony

(Fig. 6).

Discinid brachiopods (Fig. 7) are abundant and repre-

sented exclusively by the genus Orbiculoidea. The typical

dorso-ventral flattening of the brachial valve was not

observed.

Dorsal and ventral valves normally occur disarticulated,

but very close to each other and, in some cases, they belong

to the same individual. Lingulids (Fig. 7), are normally

concordant with the bedding planes, complete or with the

valves in scissor position, demonstrating that there was no

significant transport of these specimens; hence they are

regarded as autochthonous to parautochthonous.

Nautiloids (Fig. 7) are relatively rare in the Parana Basin,

and only two genera are known (Orthoceras and Spyroc-

eras). Orthoconic nautiloids, here referred questionably to

?Ctenoceras Noetling 1884, are abundant in the studied area

and are recorded for the first time in the Sao Domingos

Formation. These bioclasts are found complete or without

the apical extremity, concordant with the bedding planes,

and without preferred orientation, thus indicating the

absence of paleocurrents or occurrence in the nearshore

swash zone (Grahn 1986). The cyrtoconic shell of the

orthoceratids is strongly curved, and when fragmented

can be confused with the shells of tentaculitds of the genus

Fig. 8 Selected acritarchs (a–b), miospores (c–o), and chitinozoans

(p–t) from the Barreiro section and Sao Bento. The scale bar

represents 20 lm. a Chomotriletes vedugensis. Sıtio Ari, sample 2,

BPA 200912041, G67/3. b Lunulidia micropunctata, Sıtio Wolff,

sample 7, BPA 200912046, O45/3. c Grandispora mammillata, Casa

de Pedra, BPA 200904539, D43. d Craspedispora paranaensis, Casa

de Pedra, BPA 200904539, D54. e Grandispora pseudoreticulata,

Casa de Pedra, BPA 200904539, J61. f Chelinospora ligurata, Casa

de Pedra, BPA 200904539, G51/2. g Zonotriletes armillatus, Casa de

Pedra, BPA 200904539, K41/4. h Leiotriletes balapucensis, Casa de

Pedra, BPA 200904539, O54. i Cristatisporites sp.1, Casa de Pedra,

BPA 200904539, R50. j Archaeozonotriletes variabilis, Casa de

Pedra, BPA 200904539, S40. k Dibolisporites turriculatus, Casa de

Pedra, BPA 200904539, U40/3. l Grandispora douglastownense, Sao

Bento, BPA 200913384, L39/1. m Verrucosisporites premnus, Sao

Bento, BPA 200913384, V58/4. n Grandispora permulta, Sao Bento,

BPA 200913384, W61/4. o Chelinospora timanica, Sao Bento, BPA

200913384, M52. p Alpenachitina matogrossensis?, Sao Bento, BPA

200913384, D65c. q Alpenachitina petrovinensis, Sao Bento, BPA

200913384, E42c. r Ancyrochitina sp. cf. A. cornigera, Sao Bento,

BPA 200913384, L38/1. s Ramochitina aff. R. boliviensis, Casa de

Pedra, BPA 200904539, Q38/4. t Ramochitina ramosi. Casa de Pedra,

BPA 200904539, Q55

c


123


123

Homoctenus Ljaschenko 1955, which was recorded by

Ciguel (1989) in strata dated as Givetian. Despite the simi-

larities, the specimens studied are clearly chambered, longer

than tentaculitids and the curvature is located near the mid-

line of the shell, rather than in the apical region. Because of

the characteristic morphology of the shell it was decided to

use the genus ?Ctenoceras to define this group; however,

taxonomic studies are clearly necessary.

Ichnofossils (Fig. 7) belonging to the ichnogenus Phy-

cosiphon (ichnofacies Zoophycos) occur as U–shaped

loops, frequently ramified, and when in great numbers they

form large systems parallel or oblique to the bedding

planes. These structures are recorded for the first time in

the Parana Basin, and the animal that originated them was

probably a small (possibly mm-sized) vermiform organism.

This ichnofossil has been recorded in the Devonian of the

Parnaıba Basin, and is likely to represent another migratory

element in the assemblages here analyzed.

Palynomorphs (Fig. 8) are scarce immediately before

the extinction event (Sıtio Ari) and during the post–

extinction ‘‘Lilliput Effect’’ (Sıtio Wolff and Sao Bento),

but become common subsequently (Casa de Pedra). The

siltstones above the basal sandstones at Sıtio Ari contain,

inter alia, Chomotriletes vedugensis, an acritarch that first

occurs in latest Eifelian strata near the Eifelian–Givetian

boundary (Le Herisse, personal communication 2009) in

Bolivia (base of Los Monos Formation). Lunulidia micro-

punctata, an ecological phenotype of Navifusa bacilla, that

occurs at Sıtio Wolff and has been recorded in stressed

environments of latest Emsian–Frasnian age in the Parana

Basin. Thus, its presence at Sıtio Wolff likewise signifies a

stressed environment. Miospores and chitinozoans from

Sao Bento and Casa de Pedra are diversified and abundant

(Appendix 2), and indicative of earliest and early Givetian

age, respectively (the miospore index species Chelinospora

ligurata is present at Casa de Pedra). The section at Sao

Bento also contains chitinozoans restricted to the earliest

Givetian (i.e. Alpenachitina matogrossensis? and Alpen-

achitina petrovinensis).

On the basis of all the data obtained from this tapho-

nomic analysis, it can be concluded that the taphocoenose

is parautochthonous being preserved essentially in a near

life position with little or no evidence of transportation.

Concluding remarks

Three sections (Sıtio Ari, Sıtio Wolff, and Casa de Pe-

dra) in the lower part of the Sao Domingos Formation in

its type area near Tibagi and at a locality at Sao Bento

(Figs. 1 and 2) have been investigated. They demonstrate

a pre-extinction Malvinokaffric sequence at Sıtio Ari;

followed at Sıtio Wolff and Sao Bento, by an extinction

event possibly related to the latest Eifelian KACAK

Event, and an earliest Givetian post-extinction event

characterized by the ‘‘Lilliput Effect’’. The extinction

event reduced the Malvinokaffric genera from 65 to

eight in the post-extinction event. The individual taxa

from the surviving genera are notably abundant and, as a

manifestation of the ‘‘Lilliput Effect’’, they display a size

reduction of up to 90%. Palynomorphs are scarce during

the extinction and post-extinction events. The Malvino-

kaffric relictual fauna at Sıtio Wolff and Sao Bento

represents the last survivors of the distinctive Malvino-

kaffric fauna in the Parana Basin. They are here mixed

with immigrants and alien elements (e.g. orthoconic

nautiloids). At Casa de Pedra, the Sao Domingos For-

mation is stratigraphically immediately above the post-

extinction beds and yields an early Givetian warmer

water assemblage with Malvinokaffric survivors (e.g.

Pennaia pauliana).

Acknowledgments Elvio Pinto Bosetti acknowledges Conselho

Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq, PQ

480427/2007-0), Rafael Costa da Silva CPRM-RJ, for improving

ichnofossil taxonomy, Juliana de Morais Leme (Universidade de

Sao Paulo—USP), for improving conulariid taxonomy, and Dmitry

A. Ruban (Rostov-na-Donu, Russia), for valuable suggestions.

Yngve Grahn thanks the Faculty of Geology at Universidade do

Estado do Rio de Janeiro (UERJ) and Dr C. S. Valladares, head of

the post-graduate program at the Faculty of Geology, for access to

the facilities, and the Conselho Nacional de Desenvolvimento Ci-

entifico e Tecnologico (CNPq, PQ 309751/2007-1) which made his

work possible through grants. Rodrigo Scalise Horodyski thanks

CAPES (Coordenacao de Aperfeicoamento de Pessoal de Nıvel

Superior), Paula Mendlowicz Mauller CAPES (Coordenacao de

Aperfeicoamento de Pessoal de Nıvel Superior, BEX 4515/05-6),

and Carolina Zabini the Conselho Nacional de Desenvolvimento

Cientifico e Tecnologico (CNPq, 140659/2007-2) for grants. Pierre

Breuer acknowledges the Saudi Arabian Oil company (Saudi

Aramco) for granting permission to work on this published mate-

rial. Prof. emer. Art Boucot (Corvallis, Oregon) and Prof. emer.

Adam Urbanek (Warsaw, Poland) read the manuscript, and their

comments were most useful. The comments and linguistic correc-

tion by the two reviewers, Prof. emer. Art Boucot (Corvallis,

Oregon) and Prof. emer. Geoffrey Playford (Brisbane, Australia)

greatly improved the content of the manuscript. Dr Thomas Heuse

(Jena, Germany) made the German translations. Willian Mikio Kurita

Matsumura (UEPG, Ponta Grossa) is acknowledged for help in field and

improvement of the illustrations. Our sincere thanks to all.

Appendix 1

See Table 1.


123

Appendix 2: Palynomorphs from Sıtio Ari, Sıtio Wolf,

Sao Bento, and Casa de Pedra

Sıtio Ari

Sample 1

Acritarchs: Leiosphaeridia spp.

Sample 2

Acritarchs: Chomotriletes vedugensis Naumova 1953,

?Navifusa sp, and Tasmanites spp.

Sıtio Wolf

Sample 3

Acritarchs: cf. Dactylofusa sp., Acritarcha gen. sp. indet.

Chitinozoans: Ancyrochitina? spp.

Spores: Spore gen. sp. indet.

Sample 4

Spores: Spore gen. sp. indet

Sample 5

Barren

Sample 6

Barren

Sample 7

Acritarchs: Lunulidia micropunctata Pothe de Baldis 1979

Sample 8

Barren

Sample 9

Barren

Sao Bento

Sample 1

Barren

Sample 2

Acritarchs: Navifusa bacilla (Deunff) Playford 1977 and

other unidentified acritarchs.

Chitinozoans: Alpenachitina matogrossensis? Burjack &

Paris 1989, Alpenachitina petrovinensis Burjack & Paris

1989, Ancyrochitina langei Sommer & Boekel 1964, An-

cyrochitina sp. cf. A. cornigera Collinson & Scott 1958,

Ancyrochitina spp., Fungochitina pilosa Collinson & Scott

1958, and Ramochitina sp.

Table 1 Fossils and taphonomic data from Sıtio Wolf

Taxon Fragmentation Articulation Position

Conulatae

Conularia quıchuaUlrich 1890

VF X OB

Paraconulariaulrichana Clarke 1913

VF X OB

Lingulida

Orbiculoidea bainiSharpe 1856

NF D OB

OrbicoloideaexcentricaLange 1943

NF D OB

Lingulids indet NF D OB

Rhynchonellida

Derbyina whitiorumClarke 1913

NF D OB

Australocoeliapalmata

NF D OB

(Morris and Sharpe 1846)

Schuchertella agassiziHartt 1874

F D OB

Bivalvia

Nuculana? viatorReed 1925

NF D OB

Nautiloida

?Ctenoceras sp.

Noetling 1884

LF X OB

Trilobita

Pennaia paulianaClarke 1913

X D–A OB

Crustacea

Ostracoda indet. NF A OB

Zoophycos

Phycosiphon isp X X OB

Spongiophytaceae

Spongiophyton spp.

Krausel 1954

F X OB

Algae

Algae indet. VF X OB

F fragmented, VF very fragmented, LF little fragmented, NF not

fragmented, D disarticulated, A articulated, OB on the bedding plane,

X not considered


123

Spores: cf. Acinosporites lindlarensis Riegel 1968, Cheli-

nospora timanica (Naumova) Loboziak & Streel 1989,

Grandispora douglastownense McGregor 1973, cf. Grandis-

pora douglastownense McGregor 1973, Grandispora liby-

ensis Moreau–Benoit 1980, Grandispora permulta (Daemon)

Loboziak, Streel & Melo 1999, cf. Grandispora sp., Murati-

cavea sp., Verrucosisporites premnus Richardson 1965.

Casa de Pedra

Acritarchs: Gorgonisphaeridium spp., Leiosphaeridia spp.,

Navifusa bacilla (Deunff) Playford 1977, and other

unidentified acritarchs.

Chitinozoans: Ancyrochitina spp., Ancyrochitina langei

Sommer & Boekel 1964, Ramochitina aff. R. boliviensis

Grahn 2002, R. ramosi Sommer & Boekel 1964, Sphae-

rochitina? spp., and Chitinozoa gen. et sp. indet.

Spores: Acinosporites acanthomammillatus Richardson

1965, A. lindlarensis Riegel 1968, A. macrospinosus

Richardson 1965, Apiculiretusispora spp., Archaeozono-

triletes variabilis (Naumova) Allen 1965, Auroraspora

minuta Richardson 1965, Camarozonotriletes? concavus

Loboziak & Streel 1989, Chelinospora ligurata Allen

1965, C. timanica (Naumova) Loboziak & Streel 1989,

Craspedispora paranaensis Loboziak, Streel & Burjack

1988, Cristatisporites sp.1, Diatomozonotriletes franklinii

McGregor & Camfield 1982, Dibolisporites turriculatus

Balme 1988, Emphanisporites mcgregorii Cramer 1966, E.

rotatus McGregor 1961, Geminospora svalbardiae (Vi-

gran) Allen 1965, Grandispora incognita (Kedo) McGre-

gor & Camfield 1976, G. libyensis Moreau–Benoit 1980,

G. mammillata Owens 1971, G. permulta (Daemon) Lob-

oziak, Streel & Melo 1999, G. pseudoreticulata (Menendez

& Pothe de Baldis) Ottone 1996, Leiotriletes balapucensis

di Pasquo 2007, Retusotriletes paraguayensis Menendez &

Pothe de Baldis 1967, Retusotriletes spp., Samarisporites

spp., Verrucosisporites scurrus (Naumova) McGregor and

Camfield 1982, Verrucosisporites spp., and Zonotriletes

armillatus Breuer et al. 2007.

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Bosetti et al. 2011 publicado em : Paläontologische Zeitschrift