Multispecies multicolor: resolving the century-old taxonomic trouble of Geoplana multicolor (Platyhelminthes: Geoplanidae)

ABSTRACT

Geoplana multicolor Graff, 1899 was described based on the external characters of a single specimen collected in São Paulo, southeastern Brazil. Posteriorly, its anatomy was described based on specimens considered conspecific collected in several nearby localities, despite differences in the color pattern. The internal anatomy of the holotype was unknown until now, making identifying G. multicolor problematic, a worrying situation considering its current status as the type species of Paraba. To perform a taxonomic revision, we analyzed the anatomy of the holotype of G. multicolor deposited at the Zoologisches Museum Hamburg and concluded that its morphology matches the current definition of Geoplana. Thus, we propose the revalidation of Geoplana multicolor Graff, 1899 stat. rev. Since the type-species of Paraba returned to Geoplana and Paraba becomes a junior synonym of Geoplana, we propose a new genus, Paraparaba gen. nov., for species previously in Paraba. Considering that the basis for the identification of G. multicolor has been the description of the anatomy of specimens sampled and studied by Marcus (1951), which correspond to a different species now included into Paraparaba, we propose a new nominal species for Geoplana multicolor sensu Marcus, Paraparaba impositrix gen. nov., sp. nov. We suggest a thorough revision of further material studied by subsequent authors, sampled in different Brazilian regions and Argentina, using Marcus’ description as the basis for identification.

KEYWORDS:
Geoplaninae; land flatworms; morphology; taxonomy; Paraba

INTRODUCTION

Land planarians constitute a family of free-living terrestrial flatworms, Geoplanidae, with a worldwide distribution. With about 960 species currently known, their peak in diversity occurs in tropical forests, such as those in the Neotropical, Indo-Malayan, and Australasian ecozones (Tyler et al. 2006-2022, Sluys 1999). Due to the few external features in land planarians, knowledge of their internal anatomy is essential for their identification and classification (Winsor 1998). Unfortunately, during the 19^th century and the first decades of the 20^th century, many land planarians were described based solely on external characters (Darwin 1844, Stimpson 1857, Humbert and Claparède 1862, Dendy 1890, Schirch 1929). This led to several problematic issues in their study in subsequent years, including many misidentifications, some of which have been, fortunately, recently resolved (e.g., Almeida et al. 2018).

In 1899, Ludwig von Graff published his monograph on land planarians (Graff 1899), describing dozens of new species, thus greatly increasing the number of known species. Although many of these new species had both external and internal characters reported, others were described solely on external aspects. One of these latter species was named Geoplana multicolor Graff, 1899.

Geoplana multicolor was described based on a single specimen collected in 1894 by J. Metz in the city of São Paulo, Brazil, and was deposited in the Zoologisches Museum Hamburg. Graff (1899) provided three drawings showing the dorsal and ventral aspects of the species (Fig. 1A-C). The entire description is presented below, translated from the original in German (Graff 1899: 326):

One of the most colorful land planarians, 45 mm long, up to 4.5 mm wide and 2.5 mm thick, appears posteriorly broadly blunt, anteriorly sharply pointed, with strongly arched back and sharply flat ventral surface. A broad, rust-red (fulvo-ferrugineus) band runs over the back, turns brown at the anterior end and does not fully reach the posterior end. This red band is bordered along its entire length by whitish spots, which stand out even more sharply because the black color of the parenchyma of the lateral portions is most densely accumulated in their surroundings. Similar lighter spots lacking parenchyma pigments cover the dark sides of the back; however, they do not appear as bright, as here a yellow (sulphureus) pigment runs over them. The ventral side is dull yellow (light-avellaneus) and bordered by a narrow black stripe. The overall pattern is reminiscent of the Brazilian G. splendida, although G. multicolor is even richer and more colorfully decorated. Regarding the eyes, nothing could be discovered. The mouth lies at 23 mm and the genital opening at 29 mm from the anterior end.

Figure 1
Drawings of the color pattern of Geoplana multicolor provided by Graff (1899) (A-C) and Marcus (1951) (D, E).

The second mention of G. multicolor in the literature occurred in 1901 in a list of animals that arrived in Hamburg on board ships (Kraepelin 1901). No description of the planarian was provided and the only information presented was that it came from the city of Santos, near São Paulo.

Then, Marcus (1951) presented a detailed description of a land planarian species collected at several localities in São Paulo that he considered conspecific with G. multicolor. According to him, some animals showed a thick body, similar to Graff’s specimen, but others were much flatter. His description of the color pattern is given below (translated from the original in Portuguese) (Marcus 1951: 67):

The species’ color is very variable. Typically, the dorsum has a ferruginous or brownish band bordered by thin pale stripes. White spots, not haloes, occur on the median band. The dorsolateral regions are black, darker in the zones that touch the pale stripes and less intense outwards. The venter is pale, often without the black margin of Graff’s worm. The dorso-median band can also be ochre, black, weak or absent. In the latter case, it results in a broad pale band as in G. metzi.

By analyzing this description and the drawings that Marcus provided (Fig. 1D-E), one can see that the color pattern does not match that of the original description. Marcus seems to have interpreted the white spots bordering the rust-red band of Graff’s specimen as the yellowish stripes (which Marcus describes as “pale”) that touch the dark sides of the body and that form a single yellowish band on the dorsum when a darker (reddish to black) median band is absent. Marcus also does not mention that the median band becomes darker toward the anterior end as in Graff’s specimen. These differences, together with the lack of black margins on the ventral side in most specimens, suggest that this material is not conspecific with G. multicolor Graff.

Subsequent works used Marcus’ description as the basis for identification (Froehlich 1955, 1956a, 1956b, 1957, Leal-Zanchet and Matos 2011), and the species even became the type-species of a new genus, Paraba Carbayo et al., 2013, based on its internal morphology as originally described by Marcus (Marcus 1951, Carbayo et al. 2013).

Considering that Geoplana multicolor Graff, 1899 currently is the type species of Paraba, the problematic taxonomic status of the species is particularly troublesome. Therefore, the presented study aimed at a taxonomic revision of Geoplana multicolor including an analysis of the holotype and comparison with Marcus’ description. This resulted in the formulation of several necessary taxonomic actions.

MATERIAL AND METHODS

We analyzed the specimen labeled as G. multicolor deposited by Ludwig von Graff at the Zoologisches Museum Hamburg (ZMH), Germany, to complement the descriptions presented in his monograph (Graff 1899) and, thus, clarify possible taxonomic misconceptions involving this species. Graff’s holotype (ZMH V2657) was loaned to AMLZ by the curator of Lower Invertebrates I of the Zoologisches Museum Hamburg. The specimen was preserved in ethanol and we received permission to perform histological processing of the material.

The specimen lent by the ZMH was analyzed regarding body shape, length and width, mouth and gonopore position, and distribution of eyes. The eyes were analyzed after immersion of the preserved specimen in clove oil for four days. Histological processing of the material was performed following the methods described by Rossi et al. (2015). The material was sectioned at intervals of 6 µm and stained with Masson’s trichrome method or hematoxylin and eosin (Romeis 1989).

The ratio of the height of the cutaneous musculature to the height of the body (mc:h index in Froehlich 1955) was determined in the median region of a transverse section of the pre-pharyngeal region. Mesenchymal muscle fibers were counted in transverse sections of the same region. The color descriptors used for classifying secretions with trichrome stain methods (erythrophil, xanthophil, and cyanophil) were based on the uptake of dyes of particular colors.

TAXONOMY

Redescription of Geoplana multicolor sensu Graff, 1899

Material examined: Holotype: ZMH V2657: J. Metz coll. 1894. São Paulo, state of São Paulo, Brazil: anterior tip: transverse sections on 17 slides; anterior region: horizontal sections on 37 slides; pre-pharyngeal region in two fragments: transverse sections on 11 slides and horizontal sections on 39 slides; pharyngeal region: sagittal sections on 41 slides; copulatory apparatus in two fragments: sagittal sections on 45 slides and 19 slides, respectively.

External features: Preserved, specimen approximately 40 mm in length and with a maximum width of 4 mm; body with parallel margins, dorsal surface strongly convex and ventral side flat; anterior tip rounded (Fig. 2A). Mouth and gonopore at the third quarter of the body from the anterior end (Table 1).

Figure 2
Geoplana multicolor, holotype (V2657): (A) anterior tip of the fixed specimen in dorsal view; (B) anterior tip of the specimen in clove oil. Anterior tip to the left. (ce) Conical eyes. Scale bars: A = 0.2 mm, B = 0.5 mm.

Table 1
Measurements, in millimeters, of G. multicolor holotype. (-) Not measured, (*) after fixation, (**) length, (DG) distance of gonopore from anterior end, (DM) distance of mouth from anterior end, (DMG) distance between mouth and gonopore, (DPVP) distance between prostatic vesicle and pharyngeal pouch. The numbers given in parentheses represent the position relative to body length.

Taxonomic characters	G. multicolor holotype V2657
Length*	45
Width*	4.5
DM*	23 (51)
DG*	29 (64)
DMG*	6
DPVP	-
Ovaries	-
Anteriormost testes	-
Posteriormost testes	-
Prostatic vesicle**	-
Penis papilla**	-
Male atrium**	-
Female atrium**	1.8
Common glandular ovovitelline duct**	0.06

Conical eyes (sugarloaf shaped), 110 µm in length and 25 µm in width, surround the anterior tip (Figs 2B, 3A-B). After the second millimeter of the body (approximately 4% of body length), the eyes become monolobate and dorsal, with pigment cups about 25-35 µm in diameter.

Internal morphology: Sensory pits (Fig. 3A, B), as simple invaginations (20-40 µm), contour the anterior tip and occur ventromarginally in an irregular, single row in the first 14 mm of the body (approximately 30% of body length). The creeping sole occupies 78% of the body width in the pre-pharyngeal region (Fig. 3C, Table 2). Two types of gland discharge through the epidermis of the pre-pharyngeal region: rhabditogen glands with xanthophil secretion and cyanophil glands with finely granular secretion. Cyanophil glands with fine granules are more abundant in the ventral epidermis, while rhabditogen glands are more abundant in the dorsal epidermis (Fig. 3C, D). Glandular margin absent (Fig. 3E).

Cutaneous musculature with the usual three layers as in Geoplaninae (circular, oblique, and longitudinal), with longitudinal muscle fibers organized in thick discrete bundles (Fig. 3C-E). Thickness of cutaneous musculature becoming progressively lower towards the body margins. Ventral musculature thicker than the dorsal musculature at the sagittal plane in the pre-pharyngeal region, 1.5 times thicker than the epidermis (Table 2). Mc:h 6%; however, the measurement of the body height is not fully accurate due to artifacts in the histological preparation.

Figure 3
Geoplana multicolor, holotype (V2657), in transverse (A-E) and horizontal sections (F; anterior tip to the left): (A) anterior region of the body; (B) detail of the anterior region of the body; (C) detail of the ventral surface of the pre-pharyngeal region; (D) detail of the dorsal surface of the pre-pharyngeal region; (E) detail of body margin of the pre-pharyngeal region; (F) detail of the longitudinal mesenchymal musculature around the intestine, in the anterior region of the body. (ce) Conical eyes, (cg) cyanophil glands, (cs) creeping sole, (de) dorsal epidermis, (dm) dorsal cutaneous musculature, (dsm) dorsal subcutaneous mesenchymatic musculature, (eg) erythrophil glands, (i) intestine, (lm) longitudinal mesenchymatic musculature, (n) nerve cord, (rg) rhabditogen glands, (sbm) sub-intestinal transverse mesenchymatic musculature, (sp) sensory pit, (spm) supra-intestinal transverse mesenchymatic musculature, (v) vitelline follicles, (vm) ventral cutaneous musculature. Scale bars: A = 0.5 mm, B-F = 0.2 mm.

Table 2
Body height, cutaneous musculature, and epidermis in the median region of a transverse section of the pre-pharyngeal region, in micrometers, and ratio of the height of cutaneous musculature to the height of the body (mc:h index) of G. multicolor holotype.

Taxonomic characters	G. multicolor holotype V2657
Dorsal musculature	45
Ventral musculature	85
Dorsal epidermis	37
Ventral epidermis	42
Body height	2200
Mc:h (%)	6
Creeping sole (%)	78

Mesenchymal musculature (Fig. 3A, C, D, F) consisting of three layers: dorsal subcutaneous, located close to the cutaneous longitudinal muscle bundles, with oblique decussate fibers; supra-intestinal transverse; and sub-intestinal transverse. In addition, there are longitudinal fibers around the intestine, forming a poorly developed muscular tube (Fig. 3F), as well as dorsoventral fibers.

The pharynx is not visible in any of the analyzed body fragments.

Testes arranged in one irregular row on either side of the body, located between the dorsal transverse mesenchymal muscles and the intestinal branches. Vitellaria (Fig. 3E, F) scattered between intestinal branches, poorly developed. In the region of the copulatory apparatus, the only visible structures are the gonopore, the female atrium, and the female canal (Fig. 4A, B). Female atrium lined by pseudostratified epithelium (about 50 µm high), laterally filled by epithelium of stratified appearance (100-225 µm high) with large lacunae (Fig. 4C, D), receiving the openings of abundant erythrophil and cyanophil glands, both with finely granular secretion, and scarcer amorphous xanthophil secretion. Erythrophil and cyanophil secretions occur in the lumen of the female atrium. Muscularis of the female atrium (about 30-55 µm thick) composed of a thin subepithelial longitudinal layer, followed by a thicker circular layer, besides scarce external longitudinal fibers (Fig. 4C). Gonoduct vertical at the sagittal plane, lined with ciliated, columnar epithelium, receiving the openings of abundant xanthophil amorphous secretion and cyanophil glands producing a finely granular secretion, besides scarcer erythrophil glands with finely granular secretion. Muscularis of gonoduct consisting of a subepithelial layer of circular fibers, followed by a longitudinal layer.

Figure 4
Geoplana multicolor, holotype (V2657), photomicrographs of the copulatory apparatus in sagittal sections: (A) gonopore and female atrium; (B) detail of female canal; (C-D) details of the female atrium. Anterior tip to the left. The arrow indicates the lacunae. (cg) Cyanophil glands, (cov) common glandular ovovitelline duct, (eg) erythrophil glands, (fa) female atrium, (go) gonopore. Scale bars: A = 0.5 mm, B-D = 0.2 mm.

It is important to emphasize that the centenary specimen, borrowed from Zoologisches Museum Hamburg, was preserved in alcohol, presenting signals of previous damage at about the middle of the body as a possible result of accidental crushing. This is likely why we were unable to see the pharynx and most of the copulatory apparatus.

Comparative discussion

The analysis of the holotype of Geoplana multicolor Graff, 1899 indicates that this specimen does not present the diagnostic characteristics of Paraba Carbayo et al., 2013. Despite the impossibility of observing the anatomy of the pharynx and most of the copulatory apparatus of this specimen, characteristics such as the presence of conical eyes at the anterior end of the body and mesenchymal longitudinal musculature surrounding the intestine indicate that the specimen described by Graff (1899) belongs to Geoplana.

The diagnosis of Geoplana was recently emended, and, currently, the genus comprises 13 species (Carbayo et al. 2013, Almeida et al. 2018). One of these, Geoplana mogi Almeida & Carbayo, 2018, occurs in São Paulo and has a dorsal color pattern very similar to the one described by Graff for the holotype of G. multicolor. Geoplana mogi, has a “dorsal color pattern constituted by a median orange-ochre band, bounded on either side by a thin whitish stripe, and in turn this is bordered externally by a thin black line, externally to which is a grey band with dense white mottling, externally to which is a thin marginal black line” (Almeida et al. 2018: 20-30). Although Graff did not describe the white stripes, he stated that the wide rust-red band is bordered along its entire length by whitish spots, which in his illustration (Graff 1899: Pl. VII, fig. 12) are very similar to stripes. The thin black line that borders the external margin of these stripes in G. mogi can be compared to the black parenchyma color most densely accumulated in the surroundings of the whitish spots. Just as described by Graff, the dorsal band of G. mogi becomes darker toward the anterior end. Both descriptions indicate the presence of white spots on the dark lateral regions of the body and black margins on the ventral side (Graff 1899, Almeida et al. 2018). One of the few structures observed in the copulatory apparatus of Graff’s holotype is the female atrium, which is lined by epithelium of stratified appearance containing large lacunae, which possibly corresponds to what was described for G. mogi as “tall epithelium with stratified appearance and intraepithelial holes” (Almeida et al. 2018: 32), being the only species of Geoplana with this characteristic. Graff’s holotype corresponds to G. mogi concerning two diagnostic characteristics of this species: the dorsal color pattern and the presence of longitudinal mesenchymal muscle fibers. Only one characteristic present in the diagnosis of G. mogi, the diameter of the muscularis surrounding the ejaculatory duct, cannot be observed in G. multicolor.

The characteristics of Geoplana multicolor sensu Marcus do not match those seen in the holotype. Marcus (1951) describes his specimens as having trilobate dorsal eyes, whereas in the holotype the eyes are monolobate. In the holotype of G. multicolor, herein studied, the female atrium shows a pseudostratified epithelium with lacunae, whereas Marcus’ specimens have the female atrium almost entirely filled by an epithelium of stratified appearance, several cells high (Marcus 1951). In general, Marcus’ drawing of the copulatory apparatus of one of his specimens does not match the current definition of Geoplana and, therefore, this species was put in Paraba by Carbayo et al. (2013). However, since it was chosen as the type species mainly considering characteristics of Marcus’ description, while Geoplana multicolor in its original definition is a species of Geoplana, the genus Paraba becomes a junior synonym of Geoplana. Thus, a new genus must be erected to include the current species in Paraba and a new name must be chosen for Geoplana multicolor sensu Marcus.

Marcus (1951) describes his species as presenting a variety of different color patterns but provides a single drawing of a copulatory apparatus without specifying the color pattern of that specimen. All specimens analyzed by him were collected in the city of São Paulo. Later, land planarian specimens collected in the Brazilian states of Rio de Janeiro (Froehlich 1956), Paraná (Froehlich 1957) and Rio Grande do Sul (Leal-Zanchet and Matos 2011) and in the Argentinean province of Misiones (Negrete and Brusa 2017) were considered conspecific with Marcus’ material, even though some differences were often noticed in the external and internal anatomy. Since Marcus (1951), all works addressing Geoplana multicolor sensu Marcus in Brazil also noticed the similarity between this species and Geoplana metzi Graff, 1899, which, like G. multicolor, was also originally described based only on its external morphology. Riester (1938) described the internal morphology of specimens that he considered conspecific with Geoplana metzi. The most striking difference between Geoplanata metzi sensu Riester and Geoplana multicolor sensu Marcus seems to be the height of the epithelium in the female atrium. While Marcus’ specimen shows a tall epithelium typical of species of Paraba, in G. metzi the epithelium seems to be much lower. Due to this difference, Carbayo et al. (2013) decided to move G. metzi to Obama instead of Paraba, but, in fact, the species has not been examined in detail and was not among the ones used in the molecular phylogeny. Overall, the status of both Geoplana metzi and Geoplana multicolor sensu Marcus needs further investigation to confirm whether they belong to the same species or not. It is also likely that the specimens identified as Geoplana multicolor sensu Marcus across different states of Brazil and in Argentina represent a species complex rather than a single species. We suggest a thorough revision of such material that was identified based on Marcus’ description.

Taxonomic actions

Considering that the holotype of Paraba multicolor (Graff, 1899) presents characters exclusive to Geoplana, we propose revalidation of Geoplana multicolor Graff, 1899 stat. rev., which, thus, returns to its original genus. Despite the morphological similarities and the coincident distribution of G. multicolor and G. mogi, the impossibility of analyzing the complete morphology of the copulatory apparatus of the holotype of G. multicolor prevents the synonymization of these two species names. Therefore, Geoplana multicolor Graff, 1899 is herein regarded as a nomen dubium.

Since the currently recognized type-species of Paraba returned to Geoplana, Paraba becomes a junior synonym of Geoplana, and a new genus, Paraparaba gen. nov., is herein proposed for the species included in Paraba. We propose Geoplana rubidolineata Baptista & Leal-Zanchet, 2005 as the type species of the new genus and consider its diagnosis as the one established for Paraba by Rossi and Leal-Zanchet (2023).

Based on the description provided by Marcus (1951), we propose a new nominal species for Geoplana multicolor sensu Marcus, Paraparaba impositrix gen. nov., sp. nov. Since Marcus did not define a holotype in his original description, which seems to be based on an ensemble of non numbered specimens, we designate the specimen presented in figure 177 of the original description as the holotype. We did not ana lyze Marcus’ specimens to confirm or not that they belong to different species, so this task is yet to be accomplished.

The taxonomic actions are described below, viz. the synonymization of Paraba with Geoplana and the description of Paraparaba gen. nov. and Paraparaba impositrix sp. nov.

Order Tricladida Lang, 1884

Suborder Continenticola Carranza, Littlewood, Clough, Ruiz-Trillo, Baguña & Riutort, 1998

Geoplanidae Stimpson, 1857

Geoplaninae Stimpson, 1857

Geoplana Stimpson, 1857

Junior synonym: Paraba Carbayo, Álvarez-Presas, Olivares, Marques, Froehlich & Riutort, 2013.

Geoplana multicolor Graff, 1899 stat. rev.

Paraparaba Rossi, Boll & Leal-Zanchet, gen. nov.

https://zoobank.org/383031C9-9920-4A30-A778-1C7E8BF30099

Type species: Geoplana rubidolineata Baptista & Leal- Zanchet, 2005.

Diagnosis: Geoplaninae with small-to-medium-sized body, 6-80 mm in length; body slender, with margins nearly parallel; dorsum and ventral side slightly convex; eyes mono-, bi-, and trilobate; pharynx cylindrical; prostatic vesicle extrabulbar, horizontal, or with ventrally oriented proximal portion, with or without forked portion; penis papilla protrusible, conical or cylindrical; ascending portion of ovovitelline ducts lateral to the gonopore canal or the female atrium, joining each other above the female atrium; female canal dorsoanteriorly flexed, arising from the posterodorsal region of the female atrium; female atrium rounded to ovoid, lined with epithelium of multilayered aspect (Rossi & Leal-Zanchet 2023).

Etymology: From Tupi paraparaba, multicolored, with many colors, a word formed by reduplication of paraba, variegated, with two colors (Tibiriçá 1984). Additionally, the name could be interpreted as coming from the Greek para, beside, next to, plus the name of the genus Paraba.

Paraparaba impositrix Rossi, Boll & Leal-Zanchet, sp. nov.

https://zoobank.org/B4735B35-3076-45E7-A86C-971619A85398

Geoplana multicolor: Marcus (1951)

Geoplana multicolor: Froehlich (1955, 1956a, 1956b, 1958)

Geoplana multicolor: Leal-Zanchet & Matos (2011)

Paraba multicolor: Negrete & Brusa (2017)

Holotype: Specimen represented by Marcus (1951: fig. 177). The current location of this specimen is unknown.

Diagnosis: Species of Paraparaba with ferruginous or brownish dorsal band bordered by narrow pale stripes and black body sides; eyes mono- and trilobated; pharynx cylindrical with dorsal insertion displaced posteriorly; prostatic vesicle with globose forked proximal portion; penis papilla conical; female atrium ovoid, with narrow lumen; constriction between male and female atria.

Etymology: A feminine derivation of Late Latin imposi tor, one that applies or imposes something, an impostor, a deceiver; therefore, a female impostor, as this species “deceived” taxonomists by assuming the identity of Geoplana multicolor. The word impositor was also used in classical times by polymath Marcus Terentius Varro in his work ‘De Lingua Latina’ to refer to someone that applies a name to a thing (Lewis and Short 1891); thus, impositrix could also be interpreted as the species that applied a name, in this case Paraba, to a group of land planarian species. The word is used as a noun in apposition.

Species assigned to Paraparaba gen. nov.

Paraparaba aurantia (Marques & Leal-Zanchet, 2022), comb. nov.

Paraparaba bresslaui (Schirch, 1929), comb. nov.

Paraparaba caapora (Froehlich, 1958), comb. nov.

Paraparaba cassula (E. M. Froehlich, 1955), comb. nov.

Paraparaba franciscana (Leal-Zanchet & Carbayo, 2001), comb. nov.

Paraparaba gaucha (Froehlich, 1959), comb. nov.

Paraparaba goettei (Schirch, 1929), comb. nov.

Paraparaba hortulana (Rossi & Leal-Zanchet, 2023), comb. nov.

Paraparaba iguassuensis (Peres, Rossi & Leal-Zanchet, 2020), comb. nov.

Paraparaba impositrix Rossi, Boll & Leal-Zanchet, gen. nov., sp. nov

Paraparaba incognita (Riester, 1938), comb. nov.

Paraparaba pankaru (Amaral & Leal-Zanchet, 2019), comb. nov.

Paraparaba phocaica (Marcus, 1951), comb. nov.

Paraparaba piriana (Almeida & Carbayo, 2012), comb. nov.

Paraparaba preta (Riester, 1938), comb. nov.

Paraparaba pseudogaucha (Rossi & Leal-Zanchet, 2023), comb. nov.

Paraparaba rubidolineata (Baptista & Leal-Zanchet, 2005), comb. nov.

Paraparaba smaragdina (Rossi, Negrete & Leal-Zanchet, 2020), comb. nov.

Paraparaba suva (Froehlich, 1959), comb. nov.

Paraparaba tapira (Froehlich, 1958), comb. nov.

Paraparaba tata (Bolonhezi, Lago-Barcia & Carbayo, 2020), comb. nov.

Paraparaba tingauna (Kishimoto & Carbayo, 2012), comb. nov.

Paraparaba viricornuta (Rossi & Leal-Zanchet, 2023), comb. nov.

ACKNOWLEDGMENTS

The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq,167467/2017-4 and 313691/2018-5), the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior and the Fundação de Amparo à Pesquisa do Rio Grande do Sul are thanked for research grants and fellowships in support of this study. We acknowledge Andreas Schmidt-Rhaesa, curator of Lower Invertebrates I of the Zoologisches Museum Hamburg, for his kindness and understanding, lending us the holotype of G. multicolor and granting permission to perform its histological processing. We acknowledge the laboratory technicians C. Kremer and L. Guterres for their help in section preparation. We thank two anonymous reviewers for their valuable suggestions in an earlier draft of the manuscript.

LITERATURE CITED

• Almeida AL, Marques FPL, Carbayo F (2018) ‘Endless forms most beautiful’: taxonomic revision of the planarian Geoplana vaginuloides (Darwin, 1844) and discovery of numerous congeners (Platyhelminthes: Tricladida). Zoological Journal of the Linnean Society 185: 1-65. https://doi.org/10.1093/zoolinnean/zly022
  » https://doi.org/10.1093/zoolinnean/zly022
• Baptista VA, Leal-Zanchet AM (2005) Nova espécie de Geoplana Stimpson (Platyhelminthes, Tricladida, Terricola) do sul do Brasil. Revista Brasileira de Zoologia 22(4): 875-882. https://doi.org/10.1590/S0101-81752005000400011
  » https://doi.org/10.1590/S0101-81752005000400011
• Carbayo F, Álvarez-Presas M, Olivares CT, Marques FPL, Froehlich EM, Riutort M (2013) Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: proposal of taxonomic actions. Zoologica Scripta 42(5): 508-528. https://doi.org/10.1111/zsc.12019
  » https://doi.org/10.1111/zsc.12019
• Darwin C (1844) Brief Description of several Terrestrial Planariae, and of some remarkable Marine Species, with an Account of their Habits. Annals and Magazine of Natural History 14: 241-251.
• Dendy A (1890) On the Victorian Land Planarians. Transactions of the Royal Society of Victoria 2: 65-80.
• Froehlich CG (1955) Sobre Morfologia e Taxonomia das Geoplanidae. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Série Zoologia 19: 195-279.
• Froehlich CG (1956a) Tricladida Terricola das regiões de Teresópolis e Ubatuba. Papéis Avulsos do Departamento de Zoologia Secretaria de Agricultura São Paulo 12(16): 313-344.
• Froehlich CG (1956b) Planárias Terrestres do Paraná. Dusenia 7(4): 173-191.
• Froehlich CG (1957) On a collection of Brazilian land planarians. Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Série Zoologia 21: 93-121.
• Graff L von (1899) Monographie der Turbellarien. II. Tricladida Terricola (Landplanarien). Wilhelm Engelmann, Leipzig, 574 pp.
• Humbert A, Claparède E (1862) Description de quelques espèces nouvelles de planaires terrestres de Ceylan suivie d’observations anatomiques sur le genre Bipalium. Memoires de la Société de Physique et d’Histoire Naturelle de Genève 16: 293-306.
• Kraepelin K (1901) Ueber die durch den Schiffsverkehr in Hamburg eingschleppten Tiere. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 18(2): 183-209.
• Leal-Zanchet AM, Matos LB (2011) Geoplana multicolor Graff (Platyhelminthes: Tricladida): new records and taxonomic re-interpretation of morphological characters. Zootaxa 2804: 41-55.
• Lewis CT, Short C (1891) A new Latin Dictionary. Clarendon Press, Oxford.
• Marcus E (1951) Turbellaria Brasileiros (9). Boletim da Faculdade de Filosofia, Ciências e Letras da Universidade de São Paulo, Série Zoologia 16: 5-215.
• Negrete L, Brusa F (2017) Increasing diversity of land planarians (Platyhelminthes: Geoplanidae) in the Interior Atlantic Forest with the description of two new species and new records from Argentina. Zootaxa 4362: 99-127. https://doi.org/10.11646/zootaxa.4362.1.5
  » https://doi.org/10.11646/zootaxa.4362.1.5
• Riester A (1938) Beiträge zur Geoplaniden-Fauna Brasiliens. Abhandlungen der senkenbergischen naturforschenden Gesellschaft 44: 1-88.
• Romeis B (1989) Mikroskopische Technik. Urban und Schwarzenberg, München, 17^th ed.
• Rossi I, Amaral SV, Ribeiro GG, Cauduro GP, Fick I, Valiati VH, Leal-Zanchet AM (2015) Two new Geoplaninae species (Platyhelminthes: Continenticola) from Southern Brazil based on an integrative taxonomic approach. Journal of Natural History 50(13-14): 787-815. https://doi.org/10.1080/00222933.2015.1084057
  » https://doi.org/10.1080/00222933.2015.1084057
• Rossi I, Leal-Zanchet AM (2023) Hidden diversity under stripes: three new species of land flatworms of the genus Paraba (Platyhelminthes: Geoplanidae) from the southern Atlantic Forest. Studies on Neotropical Fauna and Environment 59(2): 509-536. https://doi.org/10.1080/01650521.2023.2278220
  » https://doi.org/10.1080/01650521.2023.2278220
• Schirch PF (1929) Sobre as planarias terrestres do Brasil. Boletim do Museu Nacional 5: 27-38.
• Sluys R (1999) Global diversity of land planarians (Platyhelmin thes, Tricladida, Terricola): a new indicator-taxon in biodiversity and conservation studies. Biodiversity & Conservation 8(12): 1663-1681. https://doi.org/10.1023/A:1008994925673
  » https://doi.org/10.1023/A:1008994925673
• Stimpson W (1857) Prodromus descriptionis animalium evertebratorum quæ in Expeditione ad Oceanum, Pacificum Septentrionalem a Republica Federata missa, Johanne Rodgers Duce, observavit et descripsit. Pars I. Turbellaria Dendrocœla. Proceedings of the Academy of Natural Sciences of Philadelphia 9: 19-31.
• Tibiriçá LC (1984) Dicionário Tupi-Português. Traço Editora, São Paulo.
• Tyler S, Schilling S, Hooge M, Bush LF (2006-2022) Turbellarian taxonomic database. Version 2.07 http://turbellaria.umaine.edu
  » http://turbellaria.umaine.edu
• Winsor L (1998) Aspects of taxonomy and functional histology in terrestrial flatworms (Tricladida: Terricola). Pedobiologia 42: 412-432.

ADDITIONAL NOTES

• ZooBank register
  https://zoobank.org/78C8A589-860A-4A5B-976D-50FA796A881A
• How to cite this article
  Rossi I, Boll PK, Leal-Zanchet AM (2024) Multispecies multicolor: resolving the century-old taxonomic trouble of Geoplana multicolor (Platyhelminthes: Geoplanidae). Zoologia 41: e23093. https://doi.org/10.1590/S1984-4689.v41.e23093
• Published by
  Sociedade Brasileira de Zoologia at Scientific Electronic Library Online - https://www.scielo.br/zool

• Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq,167467/2017-4 and 313691/2018-5) The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq,167467/2017-4 and 313691/2018-5), the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior and the Fundação de Amparo à Pesquisa do Rio Grande do Sul are thanked for research grants and fellowships in support of this study.

Edited by

• Editorial responsibility
  Fernando Carbayo

Data availability

Data citations

Tyler S, Schilling S, Hooge M, Bush LF (2006-2022) Turbellarian taxonomic database. Version 2.07 http://turbellaria.umaine.edu

Publication Dates

• Publication in this collection
  29 Nov 2024
• Date of issue
  2024

History

• Received
  30 Nov 2023
• Accepted
  08 Aug 2024