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Anais da Academia Brasileira de Ciências

Print version ISSN 0001-3765On-line version ISSN 1678-2690

An. Acad. Bras. Ciênc. vol.80 no.3 Rio de Janeiro Sept. 2008 



Record of the genus Lycopodites in the Lower Permian of Paraná Basin, Brazil



Juliana SalviI; André JasperI; Fresia Ricardi-BrancoII; Mary Elizabeth C. Bernardes-de-OliveiraIII; Margot Guerra-SommerIV

ISetor de Botânica e Paleobotânica do Museu de Ciências Naturais da UNIVATES (UNIVATES), Programa de Pós-Graduação em Ambiente e Desenvolvimento, Rua Avelino Tallini, 171, Bairro Universitário, 95900-000 Lajeado, RS, Brasil
IIInstituto de Geociências, Universidade Estadual de Campinas (UNICAMP), Caixa Postal 6152, 13083-970 Campinas, SP, Brasil
IIIInstituto de Geociências da Universidade de São Paulo (USP), Universidade de Guarulhos (UnG), Centro de Ciências Exatas e Tecnológicas, Laboratório de Geociências, Praça Tereza Cristina, 1, Centro, 07023-070 Guarulhos, SP, Brasil
IVInstituto de Geociências da Universidade Federal do Rio Grande do Sul (UFRGS), Avenida Bento Gonçalves, 9500, Agronomia, 91509-900 Porto Alegre, RS, Brasil

Correspondence to




The genus Lycopodites, which encompasses the herbaceous forms of the lycopsids, presents broad time and spacial distribution during the Paleozoic in the Northern Hemisphere, with its initial records dating from the European Devonian. As to Gondwanan Paleozoic, to this moment, only Lycopodites amazonica Dolianiti had been reported for the Amazonian Middle Devonian (Curuá Group). Thus, the specimens reported in this study such as Lycopodites sp., coming from sedimentary rocks of the Itararé Subgroup, São Paulo State, and Lycopodites riograndensis sp. nov., collected in Rio Bonito Formation, Rio Grande do Sul, represent the oldest fertile forms recorded for Gondwana and the first ones to be described for the Paraná Basin. Its presence in layers, deposited after the end of the Neopaleozoic Glaciation, shows the appearance of new taxa in high latitudes, as well as the diversity of the lycopsids present in the Basin, previously indicated through the abundance of spores associated to the Class Lycopsida present in the palinomorphous assemblages.

Key words: Lycopodites sp., Lycopodites riograndensis sp. nov., Gondwana, Lower Permian, Paraná Basin.


O gênero Lycopodites, que engloba formas herbáceas de licópsidas, apresenta ampla distribuição temporal e espacial durante o Paleozóico no Hemisfério Norte, iniciando seu registro no Devoniano da Europa. Já no Paleozóico do Gondwana, até o presente momento, somente Lycopodites amazonica Dolianiti havia sido reportada para o Devoniano Médio da Amazônia (Grupo Curuá). Assim, os exemplares reportados no presente trabalho como Lycopodites sp., provenientes de rochas sedimentares do Subgrupo Itararé, SP, e Lycopodites riograndensis sp. nov., coletados na Formação Rio Bonito, RS, representam as formas férteis mais antigas registradas para o Gondwana e as primeiras a serem descritas para a Bacia do Paraná. A sua presença em camadas depositadas após o término da glaciação neopaleozóica, evidencia o ingresso de novos taxa em latitudes altas, bem como a diversidade das licópsidas presentes na Bacia, já esboçada através dos abundantes esporos associados à Classe Lycopsida presentes nas assembléias de palinomorfos.

Palavras-chave: Lycopodites sp., Lycopodites riograndensis sp. nov., Gondwana, Permiano Inferior, Bacia do Paraná.




The main purpose of this paper is to describe specimens of herbaceous lycopsids found in Permian sediments of the Brazilian portion of Paraná Basin, and to present an overview of this genus for the Paleozoic period in this basin. Although the lycopsids are little representative in the present floras, they are meaningful herbaceous and arborescent elements of Paleozoic associations (Jasper and Guerra-Sommer 1998). These plants arose in the Devonian, reaching their apex in the Upper Carboniferous, and started to decline from the Permo-Triassic limit on; from the end of the Meso-Cenozoic they are restrict registered, with only the herbaceous forms left. According to Mussa (2000), these plants are important elements in the Carboniferous Basins of the Northern Hemispheres and in the Permian coal systems distributed through several basins of the Gondwana.

In the Paraná Basin following lycopsids genera are registered: Brasilodendron, Bumbudendron, Leptophloeum and Lycopodiopsis (Chaloner et al.1979, Jasper and Guerra-Sommer 1998, Alarcon and Bernardes-de-Oliveira 1999, Ricardi-Branco and Torres 2003) all with arbustive habit; nevertheless, herbaceous lycopsids have not been previously mentioned for this area.

The material used in this study comes from two outcrops. The oldest one is associated to the Itararé Subgroup and is known as Sítio Itapema Outcrop, located in the Municipality of Cerquilho, State of São Paulo, Brazil (Fig. 1) (Rocha-Campos et al. 2000). The other one is Quitéria Outcrop, located in the Municipality of Encruzilhada do Sul, State of Rio Grande do Sul, Brazil (Fig. 1), which is related to the Rio Bonito Formation.




The Cerquilho area is stratigraphically located in the upper sedimentary sequence of the Itararé Subgroup or in the basal portion of the Tietê/Rio Bonito Formation, in the Tubarão Group. The lithostratigraphic position of the area is still a matter of dispute. Rohn (1987), Fúlfaro et al. (1991), among other authors, have considered this area as belonging to the postglacial Tietê Formation, correlated with the Rio Bonito Formation of Southern Brazil states. On the other hand, Nagali and Consoni (1987), Martini and Rocha-Campos (1991), Santos et al. (1996), Rocha-Campos et al. (2000) among others, have considered the Cerquilho coal measures as a result of an interglacial environment, included in the Itararé Subgroup. Some elements from this taphoflora were studied by Dolianiti and Millan (1973), Millan (1977, 1991, 1995), Millan and Dolianiti (1977, 1979, 1980a, b, 1981, 1982), Iannuzzi and Rohn (1995), Ricardi-Branco et al. (1999), Bernardes-de-Oliveira et al. (1999), Rohn and Lages (2000), Martins Neto et al. (1999, 2000) and Ricardi-Branco and Bernardes-de-Oliveira (2000). These elements are the following: sphenopsids, Cordaitales, protoglossopterids (?) and abundant seeds - Cornucarpuscerquilhensis, Samaropsis rigbyi, Samaropsis dolianitii and Samaropsis tietensis. The appearance of the first elements of the Glossopteris Flora in the Paraná Basin was recorded in this outcrop. It is considered Asselian-Sakmarian in age due to its palynological content, which is characterized by basal Protohaploxypinus goraiensis Subzone of the Vittatina constabilis Interval Zone (Souza and Marques-Toigo 2003, Souza 2006).

According to Jasper et al. (2006), Quitéria Outcrop is an isolated and fragmented exposition of the Rio Bonito Formation over the Sul Rio-Grandense Shield, preserved in a collapsed block following the North-West direction. The outcropping section is little thick (6.40 m) and does not present contact with the overlapped and eroded interval, neither with the one underlying, which is covered. The profile of the site can be divided into two clearly distinguished intervals: a basal one, with large incidence of organic matter, and an upper one, with incidence of plant megafossils, but with little, nearly unpreserved, organic matter. Thus, the lower level of the upper depositional interval corresponds to a roof level, with records of a unique megaflora, constituted by fragments (Jasper et al. 2006) of conifers, fronds of filicopsids, Pteridospermales and rare Glossopteridales. Based on palynoassemblages, Jasper et al. (2006) confirm that the interval of the Quitéria Outcrop is related to the upper part of the Vittatina costabilis Zone, described by Souza and Marques-Toigo (2003), conferring a Kungurian age for deposition of the analyzed sediments.



The fossil material collected at the Sítio Itapema Outcrop corresponds to impressions found in a light-brown siltite overlying carbonaceous bed. The whole sedimentary sequence lies within a coarse-grained sandstone. The studied specimens integrate a collection of more than 200 samples collected by several researchers during two decades, and now part of the Scientific Collection of the Laboratory of Systematic Paleontology of the Geosciences Institute of São Paulo University.

At the Quitéria Outcrop the specimens of herbaceous lycopsids impressions were collected from a yellowish 45 cm thick thin sandstone layer, in which it could also be seen the impressions of Coricladus quiteriensis, Botrychiopsis valida, contra-molds and molds of stem bases of Brasilodendron pedroanum, as well as impressions of Glossopteridales leaves. These specimens are stored in the Botanic and Paleobotanic Sector of the UNIVATES Natural Science Museum under the initials PbU.

All the specimens corresponding to herbaceous lycopsids were analyzed with the use of a stereoscopic microscope, photographed and drawn with the lightcamera resource, and measures were recorded, such as the width and length of branches, microphylls, sporangia and stems. Observations made with the stereoscopic microscope indicate that there has not been preservation of cuticles and spores.



The systematic classification of fossil plants, used in this study is the one used by Stewart and Roth well (1993), who used cladistic methods for including fossil forms in the taxonomic groups of fossil and modern plants.

Realm Plantae
Division Trachaeophyta
Class Lycopsida
Order Lycopodiales
Family Lycopodiaceae

The family Lycopodiaceae includes land herbaceous plants and epiphytes approximately 5-20 cm high presenting dichotomous branching and simple leaves (microphylls) 0.2-2 cm long (Judd et al. 2002). According to Raven et al. (2003) in this family are included the modern genera Lycopodium, Phylloglossum, Huperzia, Phlegmariurus, among others, adding up to approximately 15 tropical and 7 temperate zone genera (United States and Canada).

According to Chaloner (1967), the fossil record of this family is scarce, represented by the herbaceousgenus Lycopodites Lindley and Hutton (1833). This genus resembles the modern Lycopodium (Doubinger et al. 1995). The generic name Lycopodites was used for the first time to describe some third order stems densely covered by small foliar cushions which, later on were described as conifer fragments (Taylor and Taylor 1993). In more recent descriptions this genus includes stems or axes organized in a helicoid arrangement and, if presenting sporangia in the adaxial side of the leaves, they were isolated or organized in strobiles. Specimens of Lycopodites have been described as important elements from Devonian deposits (Boureau 1967), remaining actual systems, including isophyllous and an isophyllous forms (Taylor and Taylor 1993).

Genus Lycopodites Lindley and Hutton 1833
Lycopodites sp.
(Fig. 2)



Specimen: GP/3T 2276

Provenance: Sítio Itapema, Cerquilho, São Paulo State, Brazil.

Stratigraphical level: Upper portion of Itararé Subgroup or lower portion of Tiete Formation.

Age: Early Permian/Asselian-Sakmarian.


Impression of delicate stem fragment, presented in two discontinuous portions caused by fragmentation upon specimen collection, belonging to the same axes. Its total length is 45.0 mm and it is 2.5 mm wide. It presents lepidodendroid phyllotaxy, scars in the foliar basis not clearly outlined, fusiform, with 2.5 mm of length by 1.0 mm of width, with acute inferior and superior ends. The scar of the circular vascular bundle is little clear. No evidences of ligule or parichnos. Falcate uninerved microphylls, arranged helicoidally, connected to the stem through a wide swollen base with 2.0 mm of width; their laminar portion is aciculated and do not present preserved apex, and it is 2.0 mm long and 0.5 mm wide. They emerge from the stem at 45°. It presents sparse sporophylls between the sterile microphylls, bearing ovate pedicellate sporangia of axillary position. Rounded sporangia with approximately 0.6 mm diameter, supported by 0.4 mm long pedicels.


Although it has not been possible to verify the homospory due to the poor preservation of the specimen, the supposed absence of ligule, the presence of uninerved microphylls and most importantly, the presence of sporangia on the axils of sporophylls similar to vegetative microphylls and scattered among them enable a reasonably safe identification of this specimen as belonging to the genus Lycopodites Lindley and Hutton 1833.

The specimen GP/3T - 2276 resembles the specimen Lycopodites sahnii Lele apud Surange (1966:15, Fig. 6), originated in the Triassic of South Rewa Gondwana Basin, India, concerning the following characteristics: dimensions, shape of the scars in the foliar base, and helicoidal arrangement of the scars. It differs in the following: emergence angles of 45° in the Brazilian specimen, whereas the Indian presents 90° angles; the width and the swollen shape of the microphyll base and the presence of pedicellate axillary sporangia in the Brazilian specimen.

The specimen studied differentiates itself from all the fertile species of the genus Lycopodites recorded by Chaloner (1967). As for the species based solely on vegetative bodies presenting isophyllia, Lycopodites carbonaceous Feistmantel 1875, from the Upper Carboniferous of Czech Republic and other European sites, would be the closest representative. Nonetheless, its microphylls of contracted base are quite distinct from those, with the swollen base of the specimen from São Paulo; furthermore, the lack of description of reproductive parts and dimensions prevents us from making a more accurate comparison. The specimen studied isdistinct from Lycopodites riograndensis sp. nov. once the sporangia it presents are pedicellate and smaller.

The studied specimen also differs from Lycopodites amazonica described by Dolianti (1967) for the Amazonian Basin (Devonian - Curuá Group) once the latter presents an extremely thin (approximately 1.00 mm) and stiff stem with erect linear leaves; these are apparently stiff, forming a 55° angle, arranged in a pseudo-spiral way that does not resemble at all the slightly flexous stems with falcate and decurrent leaves typical of the specimens of Lycopodites Lindley and Hutton, 1833 (non Brongniart). Due to the above it was decided to classify the studied specimen as "sp.", and wait for more specimens, which could then allow a safer identification.

Genus Lycopodites Lindley and Hutton
Lycopodites riograndensis sp. nov.
(Fig. 3)



Holotype: PbU 190 (displayed in the Paleobotanic Collection of the Botanic and Paleobotanic Sector of the UNIVATES Natural Science Museum).

Paratypes: PbU 033, 038, 294, 262, 427.

Name derivation: specific epithet derived from the name of the State of Rio Grande do Sul, location of this species occurrence.

Provenance: Quitéria Outcrop, Encruzilhada do Sul, State of Rio Grande do Sul, Brazil.

Statigraphical level: Rio Bonito Formation, Lower Permian, Paraná Basin.

Age: Early Permian/Kungurian


The studied material consists of vascular plants branch fragment of lepidodendroid phyllotaxy, preserved as impressions (Fig. 3). Their total length varies from 6.82 to 46.22 mm and their width, from 4.81 to 12.48 mm; their stems are between 0.73 to 2.28 mm wide.

Its stem is densely covered by uninerved microphylls, rounded at their base, occasionally tapering gradually towards the apex (Fig. 3) or keeping their rounded shape to the end (Fig. 3). These microphylls vary between 0.63-5.84 mm in length, and 0.25-1.37 mm in width, and are arranged helicoidally along the stem,emerging from the thallus at 45° (Fig. 3). Neither foliar scars nor ligules are preserved.

Some specimens present sporangia with ovate-elliptical shape, with measures varying from 0.78-3.21 mm in length and 0.09-1.47 mm in width, distributed among the microphylls, developing in the adaxial faces and emerging from the axils.


A difficulty found when studying fossils of the Lycopodites type is distinguishing these remains from distal branches of the species of the order Lepidodendrales. However, when there is the presence of sporangia distributed longitudinally along the stem, in association with leaves, as is the case with the material here studied, this distinction is clear. The absence of the ligule is another aspect to be used to distinguish the two groups. Furthermore, there are some species of Lycopodites with strobiles, that are also found in some species of microsporangiate Lepidostrobus which can, in fact, represent an herbaceous coned Lycopodium plant (Taylor and Taylor 1993).

Nevertheless, the specimens found in Quitéria Outcrop can be clearly included in the genus Lycopodites for presenting stems densely covered with microphylls with helicoidal distribution, having ovate-helicoidal sporangia clearly preserved in the basal portions of the adaxial faces of the microphylls (Taylor and Taylor 1993).

Another comparison that could be made is with Oxroadia gracilis, that, according to Taylor and Taylor (1993) is a small lycopsid dichotomically branched without clear preservation of the foliar cushion, but that presents the bases of the decurrent leaves. Nevertheless, the description of this genus made by Lindley and Hutton (1833) is based on specimens of the Lower Carboniferous (Calciferous Sandstone Series, Scotland). The microphylls do not present ligule and are vascularized by a single bundle. The sporangia are elongated and formed on sporophylls whose apex is turned downward. Nothing is known about the spores. This genus is considered as an herbaceous lycopsid, unlike the distal branches of the arborescent form, due to the absence of secondary tissues in the vascular system and cortex. However, the specimens analyzed here do not present evidence of dichotomy nor decurrence of the foliar bases, but do present ovate-elliptical sporangia in the adaxial faces of the non-curved sporophylls.

Synlycostrobus tyrmensis is also an interesting lycopsid of the end of the Jurassic and beginning of the Cretaceous of Siberia that has an uncommon cone arrangement (Taylor and Taylor 1993). It is thought to have been a crawling plant, probably not very different from the modern Lycopodium. The ligulate leaves areisophyllous. The cones emerge from what have been called fertile shoots, each of them located at the axis of the leave or bract. The cones are small (5.0 mm long) and consist of approximately 20 sporophylls helicoidally arranged. Each sporophyll has a conspicuous distal blade and an end leaning downward. Superficially, the fertile branches of the Synlycostrobus resemble primary axes and smaller shoots (cones) which characterize the Cordaitales. Although the morphological characteristics suggest proximity with the taxon, the presence of the ligule in the Synlycostrobus dismisses this correlation.

Thus, the specimens here studied, originating from the Quitéria Outcrop, can be clearly classified as belonging to genus Lycopodites. However, all its unique characteristics prevent us from relating it with species of it or of other genera of herbaceous lycopsids of the Permian, which justifies the creation of a specific name. Therefore, it is proposed the creation of the specific epithet "riograndensis" to this new species of the genus Lycopodites, resulting in the name Lycopodites riograndensis.



The apparent absence of the ligule, the presence of uninerved microphylls and mainly, the presence of sporophylls with axillary sporangia similar to vegetative mycrophylls scattered among them allow a relatively safe identification both from the Itapema Outcrop (Itararé Subgroup) and the Quitéria Outcrop (Rio Bonito Formation) within the genus Lycopodites.

Regarding the distribution of the genus Lycopodites (Fig. 4) and its presence in the Paraná Basin, some comments can be made. Records of this genus begin in Europe with Boureau (1967), mentioning Lycopoditesin the Devonian of Czech Republic and Germany, Carboniferous of Scotland, France and England, as well as Upper Carboniferous in Northern America. The other occurrences in the Northern Hemisphere are related to the Mesozoic. In Gondwana, all occurrences of Lycopodites already recorded, with the exception of Lycopodites amazonica, described by Dolianiti (1967), for Devonian, are associated, according to Chaloner(1967), to the Mesozoic (Lycopodites gracilis and Lycopodites sahnii in India's Triassic; Lycopodites aberi in New Zealand's Jurassic and Lycopodites victoriae in Australia's Jurassic). Thus, the forms here described for Paraná Basin are the oldest fertile structures of this genus recorded for the Southern Hemisphere.



The records of microspore and megaspore species is abundant for the Neopaleozoic of Paraná Basin; many of them are associated to lycopsids (eg. Sublagenicula, Punctatisporistes, Foveosporites, Lundbladispora, Densosporites, Cirratriradites, Kraeuselisporites, Vallatisporites and Cristatisporites). Thus, in biostratigraphic zonings proposed for this basin in paleopalynologic studies (Ricardi-Branco et al. 2002, Amaral and Ricardi-Branco 2004, Souza and Marques-Toigo 2003, Souza 2006), the presence of lycopsids spores becomes a relevant aspect, although it is not known its kinship with the genera of the stems of lycopsids described for the basin. The occurrence of the lycopsids, mainly the ones with arbustive habit (Brasilodendron, Bumbudendron, Leptophloeum and Lycopodiopsis, Chaloner et al. 1979, Jasper and Guerra-Sommer 1998, 1999, Alarcon et al. 1999, Ricardi-Branco et al. 2002), and now herbaceous, is frequent in swampy sites associated to the generation of coal levels and/or strata. This continuous presence of lycopsids, from the Carboniferous to the end of the Permian in vegetable fossil assemblages, supports the hypothesis that the Paraná Basin was inhabited by this group since the oldest records of Carboniferous (Pennsylvanian; Souza et al. 2006) to the newest in the Permian (Rohn 1997). This distribution and their reaching the western southern Gondwana might have happened firstly with the Northern Hemisphere lycophytes; this possibility is also suggested by Piérart (1962, 1975, 1981, 1984) for some megaspore species such as Sublagenicula brasiliensis, that can be considered transgressive megaspores that migrated during the Carboniferous from regions near the limits of glaciation to regions that subsequently would be taken by the Glossopteris Flora.



The authors are grateful to Biol. MSc. Cátia Viviane Gonçalves for her collaboration in the final edition of the figures. The authors also wish to acknowledge the financial support granted to this research by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), through Project N. 97/03639-8: "Survey of the Upper Carboniferous - Lower Permian (Tubarão Group) Paleofloristic Composition and Succession in the State of São Paulo"; Fundação de Amparo à Pesquisa do Estado do Rio Grande do Sul (FAPERGS) and Conselho Nacional de Desenvolvimento e Tecnologia (CNPq).



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