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Land Flatworms (Platyhelminthes: Tricladida) in Remnants of Deciduous Forest in the Northeast Region Of Southern Brazil

Planárias Terrestres (Platyhelminthes: Tricladida) Em Remanescentes De Floresta Decidual Do Nordeste Do Rio Grande Do Sul, Brasil

Abstracts

Land flatworms show high endemism due to their restricted mobility. In southern Brazil, land flatworm communities have been found mainly in areas of ombrophilous forests. Thus, this study documents land planarian species composition in remnants of deciduous seasonal forest in the northeast region of southern Brazil. Direct, diurnal samplings reveal the occurrence of 26 species of land flatworms, of which one belongs to the subfamily Rhynchodeminae and the others to the subfamily Geoplaninae. The Rhynchodeminae genus Rhynchodemus Leidy 1851 and the following Geoplaninae genera occurred: CephaloflexaCarbayo & Leal-Zanchet, 2003CARBAYO, F. & LEAL-ZANCHET, A.M. 2003. Two new genera of geoplanid land planarians (PLATYHELMINTHES: TRICLADIDA: TERRICOLA) of Brazil in the light of cephalic specialisations. Invert. Syst. 17(3):449-468. doi: 10.1071/IT01035.
https://doi.org/10.1071/IT01035...
, Choeradoplana Graff, 1896, Imbira Carbayo et al., 2013, IssocaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369., LuteostriataCarbayo, 2010CARBAYO, F. 2010. A new genus for seven Brazilian land planarian species, split off from Notogynaphallia (Platyhelminthes, Tricladida). Belg. J. Zool. 140 (Suppl.):91-101., Obama Carbayo et al., 2013, Paraba Carbayo et al., 2013, Pasipha Ogren & Kawakatsu, 1990 and XerapoaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369., besides the collective group Pseudogeoplana Ogren & Kawakatsu, 1990. The genus Obama had the highest species richness (S=6), followed by Paraba (S=4) and Pasipha (S=3). Eighteen species were recorded exclusively in one of the two study areas, and eight species occurred in both sites. The known distribution of Luteostriata abundans (Graff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.), Choeradoplana iheringiGraff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2., Obama ficki (Amaral & Leal-Zanchet, 2012), Imbira guaiana (Leal-Zanchet & Carbayo, 2001LEAL-ZANCHET, A.M. & CARBAYO, F. 2001. Two new species of Geoplanidae (Platyhelminthes, Tricladida, Terricola) from South Brazil. J. Zool. 253:433-446. doi: 10.1017/S0952836901000401.
https://doi.org/10.1017/S095283690100040...
) and Pasipha hauseri (Froehlich, 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265.) is increased. Results emphasize the relevance of expanding taxonomic studies on land flatworms and including more study areas in southern Brazil.

biodiversity inventory; planarians; community composition; Neotropical Region


Os tricladidos terrestres apresentam alto grau de endemismo, devido especialmente è sua capacidade de locomoção reduzida. No Rio Grande do Sul, dados sobre as comunidades de planárias terrestres são principalmente conhecidos de áreas de floresta ombrófila mista. O presente estudo teve como objetivo analisar a composição das espécies de planárias terrestres em remanescentes de floresta estacional decidual, situados na região nordeste do Rio Grande do Sul. Com base em coletas diurnas diretas foram registradas 26 espécies, pertencentes ès subfamílias Geoplaninae e Rhynchodeminae. Foram registrados os seguintes gêneros de Geoplaninae: CephaloflexaCarbayo & Leal-Zanchet, 2003CARBAYO, F. & LEAL-ZANCHET, A.M. 2003. Two new genera of geoplanid land planarians (PLATYHELMINTHES: TRICLADIDA: TERRICOLA) of Brazil in the light of cephalic specialisations. Invert. Syst. 17(3):449-468. doi: 10.1071/IT01035.
https://doi.org/10.1071/IT01035...
, Choeradoplana Graff, 1896, Imbira Carbayo et al., 2013, IssocaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369., LuteostriataCarbayo, 2010CARBAYO, F. 2010. A new genus for seven Brazilian land planarian species, split off from Notogynaphallia (Platyhelminthes, Tricladida). Belg. J. Zool. 140 (Suppl.):91-101., Obama Carbayo et al., 2013, Paraba Carbayo et al., 2013, Pasipha Ogren & Kawakatsu, 1990 e XerapoaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369., além do grupo coletivo Pseudogeoplana Ogren & Kawakatsu, 1990 e do gênero de Rhynchodeminae Rhynchodemus Leidy 1851. O gênero Obama apresentou a maior riqueza de espécies (S=6), seguido por Paraba (S=4) e Pasipha (S=3). Dezoito espécies foram registradas exclusivamente em uma das áreas de estudo, enquanto oito espécies ocorreram em ambas localidades. Amplia-se a distribuição conhecida de Luteostriata abundans (Graff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.), Choeradoplana iheringiGraff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2., Obama ficki (Amaral & Leal-Zanchet, 2012), Imbira guaiana (Leal-Zanchet & Carbayo, 2001LEAL-ZANCHET, A.M. & CARBAYO, F. 2001. Two new species of Geoplanidae (Platyhelminthes, Tricladida, Terricola) from South Brazil. J. Zool. 253:433-446. doi: 10.1017/S0952836901000401.
https://doi.org/10.1017/S095283690100040...
) e Pasipha hauseri (Froehlich, 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265.). Além disso, os resultados enfatizam a importância de ampliação dos estudos taxonômicos de planárias terrestres, bem como das áreas de estudo no sul do Brasil.

inventário de biodiversidade; planárias; composição de comunidades; Região Neotropical


Introduction

Land flatworms are susceptible to microclimatic changes since they live in humid, non-flooded habitats (Sluys 1998SLUYS, R. 1998. Land planarians (Platyhelminthes, Tricladida, Terrícola) in biodiversity and conservation studies. Pedobiologia 42:490-494., 1999SLUYS, R. 1999. Global diversity of land planarians (Platyhelminthes, Tricladida, Terricola): a new indicator- taxon in biodiversity and conservation studies. Biodiv. Conserv. 8:1663-1681. doi: 10.1023/A:1008994925673.
https://doi.org/10.1023/A:1008994925673...
). Their restricted motility leads to high endemism (Sluys 1995SLUYS, R. 1995. Platyhelminthes as paleogeographical indicators. Hydrobiologia 305:49-53. doi: 10.1007/BF00036362.
https://doi.org/10.1007/BF00036362...
, Ogren et al. 1997OGREN, R.E.; KAWAKATSU, M. & FROEHLICH, E.M. 1997. Additions and corrections of the previous lsnd planarian indices of the world (Turbellaria, Seriata, Tricladida, Terricola). Bull. Fuji Women's Coll. 35(2):55-61., Winsor et al. 1998WINSOR, L., JOHNS, P.M. & YEATES, G.W. 1998. Introduction, and ecological and systematic background, to the Terricola (Tricladida). Pedobiologia 42:389-404.). Land triclads are carnivorous and prey on other invertebrates such as snails, insect larvae, small arthropods and other land flatworms (Du Bois-Reymond Marcus 1951MARCUS, E. 1951. Turbellaria brasileiros. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 16:5-215., Froehlich 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369., Jones & Cumming 1998JONES, H.D. & CUMMING, M.S. 1998. Feeding behaviour of the termite-eating planarian Microplana termitophaga (Platyhelminthes: Turbellaria: Tricladida: Terricola) in Zimbabwe. J. Zool. 245:53-64. doi: 10.1111/j.1469-7998.1998.tb00071.x.
https://doi.org/10.1111/j.1469-7998.1998...
, Ogren 1995OGREN, R.E. 1995. Predation behaviour of land planarians. Hydrobiologia 305:105-111. doi: 10.1007/BF00036370.
https://doi.org/10.1007/BF00036370...
, Sluys 1999SLUYS, R. 1999. Global diversity of land planarians (Platyhelminthes, Tricladida, Terricola): a new indicator- taxon in biodiversity and conservation studies. Biodiv. Conserv. 8:1663-1681. doi: 10.1023/A:1008994925673.
https://doi.org/10.1023/A:1008994925673...
, Carbayo & Leal-Zanchet 2003CARBAYO, F. & LEAL-ZANCHET, A.M. 2003. Two new genera of geoplanid land planarians (PLATYHELMINTHES: TRICLADIDA: TERRICOLA) of Brazil in the light of cephalic specialisations. Invert. Syst. 17(3):449-468. doi: 10.1071/IT01035.
https://doi.org/10.1071/IT01035...
, Prasniski & Leal-Zanchet 2009PRASNISKI M.E.T. & LEAL-ZANCHET, A.M. 2009. Predatory behaviour of the land flatworm Notogynaphallia abundans (Platyhelminthes: Tricladida). Zoologia 26(4):606-612. doi: 10.1590/S1984-46702009005000011.
https://doi.org/10.1590/S1984-4670200900...
). Thus, flatworm diversity may reflect the diversity of other invertebrates that constitute their prey. There are records of about 170 species of land flatworms in Brazil, which has the highest species richness in the Neotropical Region (Carbayo & Froehlich 2008CARBAYO, F. & FROEHLICH, E.M. 2008. Estado do conhecimento dos macroturbelários (Platyhelminthes) do Brasil. Biota Neotrop. 8(4): http://www.biotaneotropica.org.br/v8n4/pt/abstract?thematic-review+bn01908042008 (last accessed on 30/08/2013). doi: 10.1590/S1676-06032008000400018
http://www.biotaneotropica.org.br/v8n4/p...
, 2012CARBAYO, F., & FROEHLICH E.M. 2012. Three new Brazilian species of the land planarian Choeradoplana (Platyhelminthes: Tricladida: Geoplaninae), and an emendation of the genus. J Nat. Hist. 46:1153-1177. doi: 10.1080/00222933.2012.657699.
https://doi.org/10.1080/00222933.2012.65...
, Amaral et al. 2012AMARAL, S.V., OLIVEIRA, S.M., & LEAL-ZANCHET, A.M. 2012. Three new species of land flatworms and comments on a complex of species in the genus Geoplana Stimpson Platyhelminthes: Continenticola). Zootaxa 3338:1-32., Leal-Zanchet et al. 2012LEAL-ZANCHET, A.M., ROSSI, I., SEITENFUS, A.L.R. & ALVARENGA, J. 2012. Two new species of land flatworms and comments on the genus Pasipha Ogren & Kawakatsu, 1990 (Platyhelminthes: Continenticola). Zootaxa 3583:1-21.). Recent inventories and studies on flatworm communities reveal the occurrence of at least 90 morphospecies of land flatworms, most of which are unknown to science, in forests of southern Brazil (Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38., Baptista et al. 2010BAPTISTA, V.A., OLIVEIRA, S.M. & LEAL-ZANCHET, A.M 2010. Inventário de planárias terrestres (Platyhelminthes, Tricladida) em remanescente de Floresta Estacional Decidual do sul do Brasil. Biota Neotrop. 10(2): http://www.biotaneotropica.org.br/v10n2/en/abstract?inventory+bn02110022010 (last accessed on 13/09/2013). doi: 10.1590/S1676-06032010000200027
http://www.biotaneotropica.org.br/v10n2/...
). Most of these studies occurred in ombrophilous forests (Leal-Zanchet & Carbayo 2000LEAL-ZANCHET, A.M. & CARBAYO, F. 2000. Fauna de planárias terrestres da Floresta Nacional de São Francisco de Paula, Rio Grande do Sul: uma análise preliminar. Acta Biol. Leopold. 22:19-25., Carbayo et al. 2002CARBAYO, F., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2002. Flatworms (Platyhelminthes, Tricladida, Terricola) diversity versus man-induced disturbance in ombrophilous rainforest from Southern Brazil. Biodiv. Conserv. 11:1091-1104. doi: 10.1023/A:1015865005604.
https://doi.org/10.1023/A:1015865005604...
, Fick et al. 2003FICK, I.A.; LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2003. Comparação da estrutura de comunidades de tricladidos terrestres (PLATYHELMINTHES: TERRICOLA) em duas formações florestais do sul do Brasil. In VI Congresso de Ecologia do Brasil. Anais de trabalhos completos. Universidade Federal do Ceará, Fortaleza, p.152-153., 2006FICK, I.A., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2006. Community structure of land flatworms (Platyhelminthes, Terricola): comparisons between Araucaria and Atlantic forest in Southern Brazil. Inv. Biol. 125(4):306-313. doi: 10.1111/j.1744-7410.2006.00062.x.
https://doi.org/10.1111/j.1744-7410.2006...
, Leal-Zanchet et al. 2011LEAL-ZANCHET, A.M., BAPTISTA, V.A., CAMPOS, L.M., RAFFO, J.F. 2011 Spatial and temporal patterns of land flatworm assemblages in Brazilian Araucaria forests. Invert. Biol. 130:25-33. doi: 10.1111/j.1744-7410.2010.00215.x.
https://doi.org/10.1111/j.1744-7410.2010...
, Antunes et al. 2012ANTUNES, M.B., LEAL-ZANCHET, A.M. & FONSECA, C.R. 2012. Habitat structure, soil properties, and food availability do not predict terrestrial flatworms occurrence in Araucaria Forest sites. Pedobiologia 55:25-31. doi: 10.1016/j.pedobi.2011.09.010.
https://doi.org/10.1016/j.pedobi.2011.09...
). Two inventories have been performed in deciduous forest, one in the central region and the other in the northwest region of southern Brazil (Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Baptista et al. 2010BAPTISTA, V.A., OLIVEIRA, S.M. & LEAL-ZANCHET, A.M 2010. Inventário de planárias terrestres (Platyhelminthes, Tricladida) em remanescente de Floresta Estacional Decidual do sul do Brasil. Biota Neotrop. 10(2): http://www.biotaneotropica.org.br/v10n2/en/abstract?inventory+bn02110022010 (last accessed on 13/09/2013). doi: 10.1590/S1676-06032010000200027
http://www.biotaneotropica.org.br/v10n2/...
).

Deciduous forest was the greatest forest cover in southern Brazil before colonization. About 24% of the original area now remains in the southern Brazilian state Rio Grande do Sul. The northeastern region has been highly impacted by human colonization (Teixeira & Coura-Neto 1986TEIXEIRA, M.B. & COURA-NETO, A.B. 1986. Vegetação. As regiões fitoecológicas, sua natureza, seus recursos econômicos. Estudo fitogeográfico. In Levantamento de Recursos Naturais. IBGE, Rio de Janeiro, v. 33, p. 541-632.). This study offers the first inventory of land flatworms in the northeastern deciduous forest of Rio Grande do Sul and expands our knowledge of flatworm diversity and distribution in southern Brazil.

Material and methods

The study areas are located in two cities (Salvador do Sul and São Pedro da Serra) of the northeast hillside of Rio Grande do Sul (Figure 1), between 29°26′S - 51°30′W and 29°25′S - 51°28′W, respectively. The climate is warm temperate (subtropical) and humid, without any marked dry periods (Nimer 1989NIMER, E. 1989. Climatologia do Brasil. Instituto Brasileiro de Geografia e Estatística, Rio de Janeiro.). The region experiences a long cold period with a mean temperature of 15°C and a warm period with mean temperature of 20°C (Quadros & Pillar 2002QUADROS, F.L.F. & PILLAR, V.D.P. 2002. Transições floresta-campo no Rio Grande do Sul. Ciênc. Amb. 24:109-118.). The annual rainfall is 1,200-1,750 mm year−1 with the heaviest rains occurring in the summer (Oliveira & Ribeiro 1986OLIVEIRA, A.A.B. & RIBEIRO, A.G. 1986. Climatologia. In Levantamento de Recursos Naturais. IBGE, Rio de Janeiro, v. 33, p. 757-791.). The rough relief has a maximum altitude of 556 m a.s.l. (Leite 2002LEITE, P.F. 2002. Contribuição ao conhecimento fitoecológico do sul do Brasil. Ciênc. Amb. 24:51-73.). The coverage consists of seasonal deciduous forest defined by a canopy dominated by caducifolious species and a dominance of Pachystroma longifolium (Nees) I.M.Johnst. and Eugenia rostrifolia D.Legrand (Teixeira & Coura-Neto 1986TEIXEIRA, M.B. & COURA-NETO, A.B. 1986. Vegetação. As regiões fitoecológicas, sua natureza, seus recursos econômicos. Estudo fitogeográfico. In Levantamento de Recursos Naturais. IBGE, Rio de Janeiro, v. 33, p. 541-632.). Both study areas have suffered intense anthropogenic influence, with high level of deforestation to prepare sites for agriculture and cattle farming (Teixeira & Coura-Neto 1986TEIXEIRA, M.B. & COURA-NETO, A.B. 1986. Vegetação. As regiões fitoecológicas, sua natureza, seus recursos econômicos. Estudo fitogeográfico. In Levantamento de Recursos Naturais. IBGE, Rio de Janeiro, v. 33, p. 541-632.), and have undergone consequent structural and physiognomic changes (A. Backes, personal communication).

Figure 1
Study areas of the northeastern deciduous forest of Southern Brazil. SP: São Pedro da Serra, SS: Salvador do Sul.

Sixteen diurnal samplings were performed with random and non-random methods from June 1996 to March 2007. Land flatworms were directly sampled under fallen logs and branches, rocks and leaf litter, which, after inspection, were returned to their original positions in order to avoid altering soil microhabitats (Ball & Reynoldson 1981BALL, I.R. & REYNOLDSON, T.B. 1981. British Planarians. Cambridge University Press, Cambridge., Winsor 1997WINSOR, L. 1997. The biodiversity of terrestrial flatworms (Tricladida: Terricola) in Queensland: a preliminary report. Mem. Mus. Vict. 56:575-579.).

In the field, specimens were identified as morphospecies. Their external morphology was studied under a stereomicroscope. Body fragments of the pre-pharyngeal region, pharynx, and copulatory apparatus were histologically processed to be studied under optical microscopy according to techniques described by Baptista & Leal-Zanchet (2005). Adult specimens without all the necessary morphological characteristics required for identification at the genus level, which could be differentiated from specimens of other species occurring in the study areas, were placed in the collective group Pseudogeoplana Ogren & Kawakatsu, 1990.

The sampled specimens were placed in the scientific collection of the Planarian Research Institute, Universidade do Vale do Rio dos Sinos (UNISINOS).

Results

In the two study areas, we found 26 species of land flatworms (Table 1). Twenty-five of these species belong to the subfamily Geoplaninae and one to Rhynchodeminae, of which at least 13 are unknown to science. Species of the genus Obama Carbayo et al., 2013 represented 23% of the species found (S=6), followed by Paraba Carbayo et al., 2013 (S=4) (Figures 2-4) and Pasipha Ogren & Kawakatsu, 1990 (S=3). Another seven Geoplaninae genera were also documented: CephaloflexaCarbayo & Leal-Zanchet, 2003CARBAYO, F. & LEAL-ZANCHET, A.M. 2003. Two new genera of geoplanid land planarians (PLATYHELMINTHES: TRICLADIDA: TERRICOLA) of Brazil in the light of cephalic specialisations. Invert. Syst. 17(3):449-468. doi: 10.1071/IT01035.
https://doi.org/10.1071/IT01035...
, Choeradoplana Graff, 1896, Cratera Carbayo et al., 2013, LuteostriataCarbayo, 2010CARBAYO, F. 2010. A new genus for seven Brazilian land planarian species, split off from Notogynaphallia (Platyhelminthes, Tricladida). Belg. J. Zool. 140 (Suppl.):91-101., Imbira Carbayo et al., 2013, IssocaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369. and XerapoaFroehlich, 1955FROEHLICH, E.M. 1955. Sôbre espécies brasileiras do gênero Geoplana. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 19:289-369.. Five morphospecies were included into the collective group Pseudogeoplana Ogren & Kawakatsu, 1990. The genus Rhynchodemus Leidy, 1851 (Rhynchodeminae) was also documented.

Figure 2-7
Land flatworms occurring in the northeastern deciduous forest of Southern Brazil. (2) Paraba sp.1; (3) Luteostriata abundans; (4) Obama carrierei; (5) Obama ficki; (6) Paraba gaucha; (7) Obama ladislavii. Scale bar: 5 mm.

Table 1
Land planarians recorded in areas of the northeastern deciduous forest of southern Brazil.

Four species had a high number of records, with 16 or more sampled specimens (Table 1). Two of these species, Paraba sp.1 (Figure 2) and Obama sp.1, are still undescribed (Table 1). Luteostriata abundans (Graff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.) (Figure 3) and Obama carrierei (Graff, 1897) (Figure 4) had the third and fourth highest records, respectively. Two species, Obama sp.2 and Choeradoplana iheringiGraff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2., had a moderate number of occurrences at the study sites (Table 1). The other 12 species had few occurrences (less than five specimens), and five of these are formally described taxa. Eighteen species were recorded exclusively in one of the two study areas, and eight species occurred in both sites (Table 1).

Discussion

The flatworm species richness recorded in the areas of deciduous forest of this study was similar to the richness of other areas of deciduous forest in southern Brazil where land flatworms inventories have been performed (Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Baptista et al. 2010BAPTISTA, V.A., OLIVEIRA, S.M. & LEAL-ZANCHET, A.M 2010. Inventário de planárias terrestres (Platyhelminthes, Tricladida) em remanescente de Floresta Estacional Decidual do sul do Brasil. Biota Neotrop. 10(2): http://www.biotaneotropica.org.br/v10n2/en/abstract?inventory+bn02110022010 (last accessed on 13/09/2013). doi: 10.1590/S1676-06032010000200027
http://www.biotaneotropica.org.br/v10n2/...
). Two of these locations are Turvo State Park and Santa Maria, in northwest and central Rio Grande do Sul, respectively. The species composition, however, differs among the three areas. A single species recorded in Turvo State Park (Geoplana sp.2 = Obama sp.2) and two species in Santa Maria, Obama sp.2 and O. ladislavii (Graff, 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.), were also found in our study areas. In addition, both dominant species in the study areas, Paraba sp.1 and Obama sp.1, were not yet recorded in other sites of southern Brazil where land flatworms have been studied (Leal-Zanchet & Carbayo 2000LEAL-ZANCHET, A.M. & CARBAYO, F. 2000. Fauna de planárias terrestres da Floresta Nacional de São Francisco de Paula, Rio Grande do Sul: uma análise preliminar. Acta Biol. Leopold. 22:19-25., Carbayo et al. 2002CARBAYO, F., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2002. Flatworms (Platyhelminthes, Tricladida, Terricola) diversity versus man-induced disturbance in ombrophilous rainforest from Southern Brazil. Biodiv. Conserv. 11:1091-1104. doi: 10.1023/A:1015865005604.
https://doi.org/10.1023/A:1015865005604...
, Fick et al. 2003FICK, I.A.; LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2003. Comparação da estrutura de comunidades de tricladidos terrestres (PLATYHELMINTHES: TERRICOLA) em duas formações florestais do sul do Brasil. In VI Congresso de Ecologia do Brasil. Anais de trabalhos completos. Universidade Federal do Ceará, Fortaleza, p.152-153., 2006FICK, I.A., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2006. Community structure of land flatworms (Platyhelminthes, Terricola): comparisons between Araucaria and Atlantic forest in Southern Brazil. Inv. Biol. 125(4):306-313. doi: 10.1111/j.1744-7410.2006.00062.x.
https://doi.org/10.1111/j.1744-7410.2006...
, Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Leal-Zanchet et al. 2012LEAL-ZANCHET, A.M., ROSSI, I., SEITENFUS, A.L.R. & ALVARENGA, J. 2012. Two new species of land flatworms and comments on the genus Pasipha Ogren & Kawakatsu, 1990 (Platyhelminthes: Continenticola). Zootaxa 3583:1-21., Baptista et al. 2010BAPTISTA, V.A., OLIVEIRA, S.M. & LEAL-ZANCHET, A.M 2010. Inventário de planárias terrestres (Platyhelminthes, Tricladida) em remanescente de Floresta Estacional Decidual do sul do Brasil. Biota Neotrop. 10(2): http://www.biotaneotropica.org.br/v10n2/en/abstract?inventory+bn02110022010 (last accessed on 13/09/2013). doi: 10.1590/S1676-06032010000200027
http://www.biotaneotropica.org.br/v10n2/...
).

Luteostriata abundans that is dominant in areas of semideciduous forest (Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38.), was recorded for the first time in a deciduous forest. The present record may be explained by the proximity between the northeastern part of the deciduous forest and semideciduous forest range. Furthermore, anthropogenic impacts in the prospected area may favour occupation by L. abundans, which is tolerant to such impacts (Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38., Prasniski & Leal-Zanchet 2009PRASNISKI M.E.T. & LEAL-ZANCHET, A.M. 2009. Predatory behaviour of the land flatworm Notogynaphallia abundans (Platyhelminthes: Tricladida). Zoologia 26(4):606-612. doi: 10.1590/S1984-46702009005000011.
https://doi.org/10.1590/S1984-4670200900...
). The distribution of this species is restricted to northeastern Rio Grande do Sul (Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38., Carbayo 2010CARBAYO, F. 2010. A new genus for seven Brazilian land planarian species, split off from Notogynaphallia (Platyhelminthes, Tricladida). Belg. J. Zool. 140 (Suppl.):91-101.). Obama carrierei was previously recorded in Argentina and Bolivia (Graff 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.) and in areas of semideciduous forest (Hauser & Maurmann 1959HAUSER, J. & MAURMANN, E. 1959. Studien über die Bewegungen des genus Geoplana. Pesquisas 3:631-646.).

Obama sp.2 has frequently been observed on the border of different types of forests and around man-made buildings (Leal-Zanchet & Carbayo 2000LEAL-ZANCHET, A.M. & CARBAYO, F. 2000. Fauna de planárias terrestres da Floresta Nacional de São Francisco de Paula, Rio Grande do Sul: uma análise preliminar. Acta Biol. Leopold. 22:19-25., Carbayo et al. 2002CARBAYO, F., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2002. Flatworms (Platyhelminthes, Tricladida, Terricola) diversity versus man-induced disturbance in ombrophilous rainforest from Southern Brazil. Biodiv. Conserv. 11:1091-1104. doi: 10.1023/A:1015865005604.
https://doi.org/10.1023/A:1015865005604...
, Fick et al. 2003FICK, I.A.; LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2003. Comparação da estrutura de comunidades de tricladidos terrestres (PLATYHELMINTHES: TERRICOLA) em duas formações florestais do sul do Brasil. In VI Congresso de Ecologia do Brasil. Anais de trabalhos completos. Universidade Federal do Ceará, Fortaleza, p.152-153., 2006FICK, I.A., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2006. Community structure of land flatworms (Platyhelminthes, Terricola): comparisons between Araucaria and Atlantic forest in Southern Brazil. Inv. Biol. 125(4):306-313. doi: 10.1111/j.1744-7410.2006.00062.x.
https://doi.org/10.1111/j.1744-7410.2006...
, Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38., Leal-Zanchet et al. 2012LEAL-ZANCHET, A.M., ROSSI, I., SEITENFUS, A.L.R. & ALVARENGA, J. 2012. Two new species of land flatworms and comments on the genus Pasipha Ogren & Kawakatsu, 1990 (Platyhelminthes: Continenticola). Zootaxa 3583:1-21.). It has been considered a generalist species regarding habitat use and tolerance (Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38.). Choeradoplana iheringi has not yet been recorded in areas of seasonal forests from southern Brazil (Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Baptista et al. 2010BAPTISTA, V.A., OLIVEIRA, S.M. & LEAL-ZANCHET, A.M 2010. Inventário de planárias terrestres (Platyhelminthes, Tricladida) em remanescente de Floresta Estacional Decidual do sul do Brasil. Biota Neotrop. 10(2): http://www.biotaneotropica.org.br/v10n2/en/abstract?inventory+bn02110022010 (last accessed on 13/09/2013). doi: 10.1590/S1676-06032010000200027
http://www.biotaneotropica.org.br/v10n2/...
, Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38.) despite its ample distribution in Minas Gerais, Rio de Janeiro, São Paulo, Santa Catarina and Rio Grande do Sul (Graff 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2., Riester 1938RIESTER, A. 1938. Beiträge zur Geoplaniden-Fauna Brasiliens. Abh. senckenberg. naturforsch. Ges. 441:75-78., Marcus 1951MARCUS, E. 1951. Turbellaria brasileiros. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 16:5-215., Froehlich 1956FROEHLICH, C.G. 1956. Tricladida Terricola das regiões de Teresópolis e Ubatuba. Pap. Av. Dep. Zool. 12(16):313-344., 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265., Leal-Zanchet & Souza 2003LEAL-ZANCHET, A.M. & SOUZA, S.A. 2003. Redescrição de Choeradoplana iheringi Graff (Platyhelminthes, Tricladida, Terricola). Rev. Bras. Zool. 20(3):523-530. doi: 10.1590/S0101-81752003000300026.
https://doi.org/10.1590/S0101-8175200300...
, Baptista et al. 2006BAPTISTA, V.A., MATOS, L.B., FICK, I.A. & LEAL-ZANCHET, A.M. 2006. Composição de comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) do Parque Nacional dos Aparados da Serra, Brasil. Iheringia 96:293-297. doi: 10.1590/S0073-47212006000300004.
https://doi.org/10.1590/S0073-4721200600...
). Until now, its distribution in southern Brazil included areas of ombrophilous forest (Leal-Zanchet & Souza 2003LEAL-ZANCHET, A.M. & SOUZA, S.A. 2003. Redescrição de Choeradoplana iheringi Graff (Platyhelminthes, Tricladida, Terricola). Rev. Bras. Zool. 20(3):523-530. doi: 10.1590/S0101-81752003000300026.
https://doi.org/10.1590/S0101-8175200300...
, Baptista et al. 2006BAPTISTA, V.A., MATOS, L.B., FICK, I.A. & LEAL-ZANCHET, A.M. 2006. Composição de comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) do Parque Nacional dos Aparados da Serra, Brasil. Iheringia 96:293-297. doi: 10.1590/S0073-47212006000300004.
https://doi.org/10.1590/S0073-4721200600...
, Antunes et al 2012ANTUNES, M.B., LEAL-ZANCHET, A.M. & FONSECA, C.R. 2012. Habitat structure, soil properties, and food availability do not predict terrestrial flatworms occurrence in Araucaria Forest sites. Pedobiologia 55:25-31. doi: 10.1016/j.pedobi.2011.09.010.
https://doi.org/10.1016/j.pedobi.2011.09...
).

Obama ficki (Amaral & Leal-Zanchet, 2012) (Figure 5) has been documented in ombrophilous forest areas of Rio Grande do Sul and Santa Catarina (Leal-Zanchet & Carbayo 2000LEAL-ZANCHET, A.M. & CARBAYO, F. 2000. Fauna de planárias terrestres da Floresta Nacional de São Francisco de Paula, Rio Grande do Sul: uma análise preliminar. Acta Biol. Leopold. 22:19-25., Fick et al. 2003FICK, I.A.; LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2003. Comparação da estrutura de comunidades de tricladidos terrestres (PLATYHELMINTHES: TERRICOLA) em duas formações florestais do sul do Brasil. In VI Congresso de Ecologia do Brasil. Anais de trabalhos completos. Universidade Federal do Ceará, Fortaleza, p.152-153., 2006FICK, I.A., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2006. Community structure of land flatworms (Platyhelminthes, Terricola): comparisons between Araucaria and Atlantic forest in Southern Brazil. Inv. Biol. 125(4):306-313. doi: 10.1111/j.1744-7410.2006.00062.x.
https://doi.org/10.1111/j.1744-7410.2006...
, Baptista et al. 2006BAPTISTA, V.A., MATOS, L.B., FICK, I.A. & LEAL-ZANCHET, A.M. 2006. Composição de comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) do Parque Nacional dos Aparados da Serra, Brasil. Iheringia 96:293-297. doi: 10.1590/S0073-47212006000300004.
https://doi.org/10.1590/S0073-4721200600...
, Amaral et al. 2012AMARAL, S.V., OLIVEIRA, S.M., & LEAL-ZANCHET, A.M. 2012. Three new species of land flatworms and comments on a complex of species in the genus Geoplana Stimpson Platyhelminthes: Continenticola). Zootaxa 3338:1-32.). The type-locality of Paraba gaucha (Froehlich, 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265.) (Figure 6) is Salvador do Sul. It has also been found in areas of semideciduous forest (Froehlich 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265., Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38.). Obama ladislavii (Graff 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2.) (Figure 7) was recorded in all four main types of forest formations in southern Brazil (Graff 1899GRAFF, L.V. 1899. Monographie der Turbellarien. Tricladida, Terricola (Landplanarien). Engelmann, Leipzig, v. 2., Carbayo et al. 2002CARBAYO, F., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2002. Flatworms (Platyhelminthes, Tricladida, Terricola) diversity versus man-induced disturbance in ombrophilous rainforest from Southern Brazil. Biodiv. Conserv. 11:1091-1104. doi: 10.1023/A:1015865005604.
https://doi.org/10.1023/A:1015865005604...
, Fick et al. 2003FICK, I.A.; LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2003. Comparação da estrutura de comunidades de tricladidos terrestres (PLATYHELMINTHES: TERRICOLA) em duas formações florestais do sul do Brasil. In VI Congresso de Ecologia do Brasil. Anais de trabalhos completos. Universidade Federal do Ceará, Fortaleza, p.152-153., 2006FICK, I.A., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2006. Community structure of land flatworms (Platyhelminthes, Terricola): comparisons between Araucaria and Atlantic forest in Southern Brazil. Inv. Biol. 125(4):306-313. doi: 10.1111/j.1744-7410.2006.00062.x.
https://doi.org/10.1111/j.1744-7410.2006...
, Castro & Leal-Zanchet 2005CASTRO, R.A & LEAL-ZANCHET, A.M. 2005. Composição de comunidades de planárias terrestres (Platyhelminthes) em área de floresta estacional e de campo na região central do Rio Grande do Sul, Brasil. Acta Biol. Leopold. 27:147-150., Baptista et al. 2006BAPTISTA, V.A., MATOS, L.B., FICK, I.A. & LEAL-ZANCHET, A.M. 2006. Composição de comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) do Parque Nacional dos Aparados da Serra, Brasil. Iheringia 96:293-297. doi: 10.1590/S0073-47212006000300004.
https://doi.org/10.1590/S0073-4721200600...
, Leal-Zanchet & Baptista 2009LEAL-ZANCHET, A.M. & BAPTISTA, V.A. 2009. Planárias terrestres (Platyhelminthes: Tricladida) em áreas de floresta com araucária no Rio Grande do Sul. In Floresta com Araucária: Ecologia, Conservação e Desenvolvimento Sustentável (C.R.S. Fonseca, A.F. Souza, T.L. Dutra, A.M. Leal-Zanchet, A. Backes & G.M.S. Ganade, eds.), Holos, Ribeirão Preto, p.199-207., Antunes et al. 2008ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38., 2012ANTUNES, M.B., LEAL-ZANCHET, A.M. & FONSECA, C.R. 2012. Habitat structure, soil properties, and food availability do not predict terrestrial flatworms occurrence in Araucaria Forest sites. Pedobiologia 55:25-31. doi: 10.1016/j.pedobi.2011.09.010.
https://doi.org/10.1016/j.pedobi.2011.09...
). Its known distribution includes locations in Rio Grande do Sul and Santa Catarina. Imbira guaiana (Leal-Zanchet & Carbayo, 2001LEAL-ZANCHET, A.M. & CARBAYO, F. 2001. Two new species of Geoplanidae (Platyhelminthes, Tricladida, Terricola) from South Brazil. J. Zool. 253:433-446. doi: 10.1017/S0952836901000401.
https://doi.org/10.1017/S095283690100040...
) was only recorded in São Francisco de Paula, Rio Grande do Sul (type-locality), in areas of ombrophilous forest and plantations of Araucaria angustifolia (Carbayo et al. 2001CARBAYO, F. & LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2001. Land planarians (PLATYHELMINTHES, TRICLADIDA, TERRICOLA) as indicators of man-induced disturbance in South Brazilian rainforest. Belg. J. Zool. 131(1):223-224., 2002CARBAYO, F., LEAL-ZANCHET, A.M. & VIEIRA, E.M. 2002. Flatworms (Platyhelminthes, Tricladida, Terricola) diversity versus man-induced disturbance in ombrophilous rainforest from Southern Brazil. Biodiv. Conserv. 11:1091-1104. doi: 10.1023/A:1015865005604.
https://doi.org/10.1023/A:1015865005604...
). Pasipha hauseri (Froehlich, 1959FROEHLICH, C.G. 1959. On geoplanids from Brazil. Bol. Fac. Fil. Ciênc. Letr. Univ. São Paulo, série Zoologia, 22:201-265.) was previously recorded in the distribution range of the semideciduous forest, but also in the Amazon Forest (Froehlich & Froehlich 1972FROEHLICH E.M. & FROEHLICH, C.G. 1972. Land planarians from the Amazonian region. Pap. Av. Zool. 26(2):29-45.).

Deciduous forest remnants in northeastern southern Brazil suffer a greater amount of anthropogenic impacts than remnants in the northwest and central regions. Nevertheless, Geoplaninae species richness is high in this region. The occurrence of at least 13 species of land planarians that are probably new to science emphasises the importance of increasing taxonomic studies of the group, as was suggested by Antunes et al. (2008)ANTUNES, M.B., MARQUES, D.I.L. & LEAL-ZANCHET, A.M. 2008. Composição das comunidades de planárias terrestres (Platyhelminthes, Tricladida, Terricola) em duas áreas de floresta estacional decidual do sul do Brasil. Neotrop. Biol. Conserv. 31:34-38.. These results, as well as differences in species composition among areas, also indicate the need to further develop studies of land planarian communities in other areas of southern Brazil.

We are grateful to the Brazilian Research Council (CNPq), the Fundação de Amparo è Pesquisa do Rio Grande do Sul (FAPERGS) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for grants and fellowships in support of this study. We thank Prof. Dr. A. Backes (in memoriam) for his generosity and for sharing his deep knowledge of the forests of southern Brazil over many years. We acknowledge A.L. Seitenfus, C. Palacios, D. Rocha, E. Benya, I. Fick, L. Teixeira, M. Fontoura, M. Cardoso, M. Antunes, M. Gallon, N. Rodrigues, R. Murowaniecki, R. Castro, S. Souza, S.M. Oliveira, S.T. Souza, S. Schneider, V. Baptista and W. Santos for their help in sampling flatworms. Dr. L. Negrete is thanked for photographs of live specimens of flatworms in the figures 2, 3 and 7. We also thank MSc. E. Toriani for the English review of the manuscript. Three anonymous reviewers are acknowledged for their suggestions.

References

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Publication Dates

  • Publication in this collection
    Mar 2014

History

  • Received
    08 Oct 2013
  • Reviewed
    24 Jan 2014
  • Accepted
    14 Jan 2014
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