Tracking the diversity of the flatworm genus Imbira (Platyhelminthes)
in the Atlantic Forest
Silvana V. Amaral
Giovana G. Ribeiro
Mário J. Müller
Victor H. Valiati
Received: 23 August 2017 / Accepted: 10 November 2017 / Published online: 27 January 2018
Gesellschaft für Biologische Systematik 2018
The genus Imbira Carbayo et al., 2013 encompasses two species, Imbira guaiana (Leal-Zanchet & Carbayo, 2001) and Imbira
marcusi Carbayo et al., 2013, which occur in south Brazil, in areas originally covered by the Atlantic Forest. In the present study,
we examine the genetic diversity within the genus, investigate the occurrence of molecular autapomorphies for its species and
describe a new species for the genus based on an integrative approach. The Bayesian and maximum likelihood analyses based on
DNA barcoding recovered the monophyly of the genus Imbira, but indicate that specimens representing I. marcusi correspond to
five distinct lineages. These analyses, as well as sequence divergence data, revealed that the new species herein described is
closely related to I. guaiana and that the specific status of specimens of I. marcusi available in GenBank should be reviewed. In
addition, sequence analysis revealed 32 molecular autapomorphies for all independent evolutionary units within the genus. The
new species described herein seems to be endemic to its type locality, a private area without legal protection.
Land planarians (Tricladida: Continenticola), soil animals of
cryptic behaviour, depend on the humidity of their microhab-
itat because they do not have water-conserving adaptations
(Kawaguti 1932;Froehlich1955a;Winsoretal.1998). These
flatworms have restricted locomotion capacity over long dis-
tances, so that there are many endemic species (Sluys 1995).
The highest species richness of land planarians worldwide has
been documented in the southern hemisphere (Winsor et al.
1998), in areas which were originally covered by the
Brazilian Atlantic Rain Forest (Sluys 1998, 1999; Carbayo
et al. 2002;Ficketal.2006; Leal-Zanchet et al. 2011). Land
flatworms are predators of other soil invertebrates (Froehlich
Many land triclad species select relatively undisturbed areas,
such as native forests and plantations of a native tree species in
southern Brazil, rather than plantations of exotic trees
(Carbayo et al. 2002; Fonseca et al. 2009; Oliveira et al.
2014). The coexistence of several species in the same habitat
is possible due to differences in prey items, which reduces
interspecific competition (Boll and Leal-Zanchet 2016).
Land flatworms belong to the family Geoplanidae, which is
subdivided into four subfamilies, viz. Bipaliinae, Microplaninae,
Rhynchodeminae and Geoplaninae (Sluys et al. 2009). The sub-
family Geoplaninae has a Neotropical distribution and com-
prises about 270 species in 23 genera (Sluys et al. 2009;
Carbayo et al. 2013; Lemos et al. 2014; Negrete et al. 2014;
Rossi et al. 2014; Álvarez-Presas et al. 2015; Carbayo and
Almeida 2015; Rossi et al. 2015; Carbayo et al. 2016a, b).
Usually, taxonomic descriptions of land flatworms detail
external features, such as colour pattern and eye arrangement,
and anatomical features mainly related to the pharyngeal anat-
omy and anatomy of the reproductive apparatus. However, in
some genera, such features are very homogeneous among
species, making species delimitation a difficult task.
Therefore, more recently integrative approaches employing
molecular and morphological data have been used for species
identification in the subfamily Geoplaninae (Lemos et al.
2014;Álvarez-Presasetal.2015; Rossi et al. 2015;Carbayo
et al. 2016a, b
Electronic supplementary material The online version of this article
(https://doi.org/10.1007/s13127-018-0358-6) contains supplementary
material, which is available to authorized users.
* Ana Leal-Zanchet
Programa de Pós-Graduação em Biologia, Universidade do Vale do
Rio dos Sinos, São Leopoldo, Rio Grande do Sul 93022-000, Brazil
Organisms Diversity & Evolution (2018) 18:87–99