Two new land planarian species (Platyhelminthes: Tricladida) from the Cerrado biome in southwestern Brazil.

Most Neotropical land flatworm species are known from areas of the Atlantic Forest. Herein, we describe two new land planarian species from areas of semideciduous forest in the Cerrado biome in southwestern Brazil. Paraba aurantia Marques & Leal-Zanchet, sp. nov. can be distinguished from its congeners by the wide black median band on the dorsum, eyes occupying almost the entire dorsal surface, prostatic vesicle twisted and asymmetrical with proximal portions forked, and female atrium with an ample lumen lined by a low epithelium with stratified appearance. Pasipha liviae Marques & Leal-Zanchet, sp. nov. shows a narrow longitudinal dorsal stripe, collar-shaped pharynx, and prostatic vesicle with a pear-shaped proximal portion and a globose distal portion separated by a constriction. In addition, P. liviae can be distinguished from its congeners by molecular analyses from the cytochrome c oxidase subunit I gene (COI). Phylogenetic analyses suggest that P. liviae is closely related to P. hauseri, which occurs in areas of semidseciduous forest in southern Brazil.

Mind the food: Survival, growth and fecundity of a Neotropical land planarian (Platyhelminthes, Geoplanidae) under different diets.

Although most land planarians are sensitive to environmental changes, some species are well adapted to human-disturbed areas and are easily transported to new places, having the potential to threaten native ecosystems. We investigated growth and survival in a land planarian common in human-disturbed areas in southern Brazil. Specimens of Obama anthropophila were divided into three groups that received different diets: (1) only the land planarian Luteostriata abundans (N=13), (2) only the slug Deroceras leave (N=12), and (3) alternating both prey types (N=13). We monitored the weight of the specimens for a month and counted and weighed egg capsules. Planarians receiving a mixed diet tended to survive less than the groups receiving a single prey type; there were significant differences between those feeding on D. laeve and the other groups. Planarians with the mixed diet ate L. abundans more often than D. laeve, and those feeding only on L. abundans tended to eat more than the other groups. Most egg capsules were laid by specimens with a diet based on D. laeve but the mixed-diet group laid heavier capsules. Both prey species are suitable as food for O. anthropophila, although it prefers planarians when both food items are available. The constant alternation between food items seems to have adverse effects, perhaps related to physiological changes to digest different food items. The heavier egg capsules of the mixed-diet group, considering its lower survival, suggest terminal investment, i.e., an increased reproductive effort when approaching death. The ability to feed on both snails and planarians, combined with its proximity to humans, make O. anthropophila a potentially invasive species.

Phylogeny and biogeography of the Cavernicola (Platyhelminthes: Tricladida): Relicts of an epigean group sheltering in caves?

The planarian suborder Cavernicola Sluys, 1990 was originally created to house five species of triclad flatworms with special morphological features and a surprisingly discontinuous and broad geographic distribution. These five species could not be accommodated with any degree of certainty in any of the three taxonomic groups existing at that moment, viz., Paludicola Hallez, 1892, Terricola Hallez, 1892, and Maricola Hallez, 1892. The scarce representation of the group and the peculiarities of the morphological features of the species, including several described more recently, have complicated new tests of the monophyly of the Cavernicola, the assessment of its taxonomic status, as well as the resolution of its internal relationships. Here we present the first molecular study including all genera currently known for the group, excepting one. We analysed newly generated 18S and 28S rDNA data for these species, together with a broad representation of other triclad flatworms. The resulting phylogenetic trees supported the monophyly of the Cavernicola, as well as its sister-group relationship to the Maricola. The sister-group relationship to the Maricola and affinities within the Cavernicola falsify the morphology-based phylogeny of the latter that was proposed previously. The relatively high diversity of some cavernicolan genera suggests that the presumed rarity of the group actually may in part be due to a collecting artefact. Ancestral state reconstruction analyses suggest that the ancestral habitat of the group concerned epigean freshwater conditions. Our results point to an evolutionary scenario in which the Cavernicola (a) originated in a freshwater habitat, (b) as the sister clade of the marine triclads, and (c) subsequently radiated and colonized both epigean and hypogean environments. Competition with other planarians, notably members of the Continenticola, or changes in epigean habitat conditions are two possible explanations -still to be tested- for the loss of most epigean diversity of the Cavernicola, which is currently reflected in their highly disjunct distributions.

Two new land planarian species (Platyhelminthes: Tricladida: Geoplanidae) from protected areas in southern Brazil.

We describe two new species of Neotropical land planarians found in protected areas of the Atlantic Forest in southern Brazil. Cratera obsidiana Amaral, Boll Leal-Zanchet, sp. nov., found in the State Park of Turvo, is the first species of the genus Cratera described from an area of deciduous seasonal forest. It can be distinguished from its congeners by the uniformly black dorsal color and the prostatic vesicle with proximal portion laterally expanded and directed upward. Luteostriata subtilis Boll, Amaral Leal-Zanchet, sp. nov., found in the National Park of Aparados da Serra, in an area covered by Araucaria moist forest, is distinguished from its congeners by only two conspicuous longitudinal stripes on the dorsal surface of the body and a large fold in the male atrium below the opening of the ejaculatory duct. We also provide an updated key for species of Cratera and the first key for species of Luteostriata.

Why be original? Two new species of Choeradoplana resembling the type species of the genus in their external aspects (Platyhelminthes, Continenticola).

The genus Choeradoplana Graff, 1896 encompasses 16 species, most of them found in Brazil. Herein two new species of this genus are described from remnants of Araucaria moist forests, located in the states of Paraná and Santa Catarina, south Brazil. Both species resemble the type-species of the genus, C.iheringi, showing brownish dorsal surface covered by dark-brown flecks. However, regarding their anatomy, the new species differ from C.iheringi and other congeners by a long and horizontal disposed permanent papilla. In such aspects, both species resemble C.benyai, but differ from this species, as well as from each other, in details of the prostatic vesicle, ejaculatory duct, and penis papilla.

Coming out in a harsh environment: a new genus and species for a land flatworm (Platyhelminthes: Tricladida) occurring in a ferruginous cave from the Brazilian savanna.

Faunal inventories in ferruginous caves from an area belonging to the Brazilian savanna (Cerrado phytophysiognomy), on the eastern margin of the Serra do Espinhaço Plateau, in southeastern Brazil, have revealed the occurrence of land flatworms. Herein, a flatworm sampled in such subterranean environment is described as a new genus and species of the Neotropical subfamily Geoplaninae, Difroehlichia elenae gen. nov., sp. nov. The new genus shows rare features within Geoplaninae, namely sub-cylindrical body, poorly developed sub-epidermal musculature and a narrow creeping sole. Some features, such as a small body and a broad sensory margin in the anterior region of the body, as well as the fact that the holotype showed signs of recent copula, may indicate an adaptation to the subterranean environment, probably representing a troglophile. Difroehlichia elenae is characterized by an almost homogeneous dark brown pigmentation over dorsal surface and body margins, a short cylindrical pharynx, and a tubular and unforked intrabulbar prostatic vesicle, among other features. The holotype shows a secondary male copulatory organ located immediately behind the primary one, both communicating with the female atrium and gonopore canal. Since the species seems to have low abundance and restricted distribution and its type-locality is affected by mining activities, major concern arises regarding its conservation.

Integrative approach reveals two new species of Obama (Platyhelminthes: Tricladida) from the South-Brazilian Atlantic Forest.

The genus Obama Carbayo et al., 2013 includes 38 species, being the most species-rich within Geoplaninae. Species of this genus show a similar anatomy regarding their copulatory apparatus, which may hinder species differentiation. In this study, we describe two new species, presenting a marbled colour pattern, found in two different phytophysionomies of the Atlantic Forest, namely Semi-deciduous Forest and Araucaria Forest. Both species can be distinguished from their congeners, as well from each other, by colour pattern and eye arrangement combined with characteristics of the pharynx, penis papilla and prostatic vesicle, confirmed by molecular analyses from cytochrome c oxidase subunit I gene (COI). Phylogenetic analyses suggest that both species herein studied are closely related to another species that occurs in areas of Araucaria Forest (O. maculipunctata). Results also indicate the need to use at least 600 bp of the gene COI in the definitions of interspecific divergences and for species delineation, at least for the genus Obama.

Two new sympatric troglobitic freshwater flatworms (Platyhelminthes: Dugesiidae) from a hotspot of subterranean biodiversity in the Neotropics.

The Areias System, situated in the Atlantic Forest, is considered a hotspot of troglobitic diversity in Brazil. Herein we describe two new sympatric, troglobitic species of Girardia occurring in this system. Both species show minute eyes; one of them has a whitish body, and the other scattered, fine pigmentation over the dorsal surface. Regarding the copulatory apparatus, one of the species shows a bulbar cavity with multiple diverticula and a bursal canal of the angled type, whereas the other has a single, ovoid bulbar cavity and a smoothly curved bursal canal, among other distinguishing features. Thus, each species can be differentiated by a unique combination of features in its external morphology and copulatory apparatus. Both present a restricted distribution, occurring in a group of pools in travertine rock formed by water from the epikarst. The morphological features of both species and this sampling location indicate that both species are stygobionts.

Searching for shelter in a ferruginous cave? A new species of Pasipha from a plateau in the Brazilian savanna (Platyhelminthes, Tricladida).

In a fauna survey in the eastern margin of Serra do Espinhaço Plateau, in an area belonging to the Brazilian savanna (Cerrado phytophysiognomy), a land flatworm was sampled in a ferruginous cave. Anatomical and histological analyses indicated that it belongs to a new species of the genus Pasipha, which is herein described. The new species shows an almost homogenous dark brown dorsal pigmentation, eyes spreading over the dorsal surface, a collar-shaped pharynx, and a prostatic vesicle with two portions separated by a canal. It differs from similar species mainly by anatomical and histological details of the ejaculatory duct, as well as male and female atria. The flatworm shows no troglomorphic traits and was collected once in the entrance zone of the cave. Hence, despite representing the first land flatworm species described from a Neotropical cave, we consider that its occurrence in the cave is probably occasional, using it as a shelter.

Diversity out of simplicity: interaction behavior of land planarians with co-occurring invertebrates.

Land planarians have a simple anatomy and simple behavioral repertoire in relation to most bilaterian animals, which makes them adequate for the study of biological processes. In this study, we investigate the behavior of land planarians during interaction events with other invertebrates found in the same environment. We observed 16 different behavioral units, including seven different capture behaviors and three different prey ingestion behaviors. The capture behavior varied from very simple, such as simply covering the prey with the body, to more complex ones, including two forms of tube formation that are described for the first time. In general, the capture behaviors were similar among different predators but different for different prey. Similarly, prey ingestion type was more related to prey type than to predator species, with small soft prey being swallowed without fragmentation, large prey being crushed, and prey with a hard skeleton being perforated. Considering that land planarians face limitations due to their lack of efficient ways to retain water, thus being highly dependent on a moist environment, the set of behaviors shown by them in this study was considerably rich, especially concerning strategies to capture prey.

Three new species of Cratera Carbayo et al., 2013 from Araucaria forests with a key to species of the genus (Platyhelminthes, Continenticola).

Areas of Araucaria moist forest have been considered to constitute hotspots of land flatworm diversity, harbouring a high number of undescribed species. Herein we describe three new species of land flatworms of Cratera Carbayo et al., 2013 occurring in such type of forest in south Brazil. The three species are differentiated from their congeners mainly by their colour pattern, anatomy of the pharynx and prostatic vesicle, and details of the penis papilla and male atrium. An identification key to species of the genus in the Neotropical region is provided.

Two new species of Pasipha Ogren & Kawakatsu (Platyhelminthes: Continenticola) from areas of deciduous forest in southern Brazil.

Two new species of Geoplaninae from southern Brazil are described herein. The new species, belonging to the genus Pasipha Ogren & Kawakatsu, 1990, can be distinguished from each other and from their congeners by colour pattern and characteristics of the copulatory apparatus, especially regarding the female organs and prostatic vesicle. Both new species seem to be endemic to areas covered by deciduous forest.

Two new species of freshwater flatworms (Platyhelminthes: Tricladida: Continenticola) from South American caves.

The diversity of freshwater triclads in the Neotropical region is considered to be low, but extensive areas of South America remain almost unexplored. Herein we describe two cave-dwelling, new species of Girardia, one from a transition zone of the Cerrado and Caatinga phytophysiognomies and the other from the Cerrado phytophysiognomy. The species from the Cerrado-Caatinga transition is a troglobite, eyeless and whitish; the species from the Cerrado area is possibly a troglophile, since it shows heavily pigmented body and eyes. Each species is easily recognized by a unique combination of features in its external morphology and copulatory apparatus. The two new species of Girardia show a restricted distribution, even the troglophile, and occur in caves without legal protection. Therefore, they must be considered as vulnerable organisms in a conservation context.

Preference for different prey allows the coexistence of several land planarians in areas of the Atlantic Forest.

Land planarians are recognized as important predators, yet studies on their feeding habits are usually restricted to invasive species. Thus, it is difficult to determine the real ecological role of this group in ecosystems and how their communities are structured. In the present study, we analyzed the diet of six co-occurring Neotropical land planarians and their success in capturing prey, based on experiments in the laboratory, in order to determine how they share resources in the same environment. We also calculated indices of food niche breadth and food niche overlap for land planarians for the first time. The diet of Luteostriata abundans comprises only woodlice and the diets of Obama ficki and Obama ladislavii are composed only of gastropods, while Paraba multicolor and Obama anthropophila feed on both gastropods and other land planarians. An invasive species recently found in Western Europe, Obama nungara, showed the highest food niche breadth, feeding on gastropods, earthworms and planarians. We found the highest niche overlap between O. anthropophila and P. multicolor. The results suggest that land planarians are frequent predators of woodlice and land gastropods in the Neotropical ecozone and thus are important for the maintenance of native ecosystems and for the control of invasive species. The coexistence of several species in the same habitat is possible due to the use of different species as main prey, which reduces interspecific competition.

A taste for exotic food: Neotropical land planarians feeding on an invasive flatworm.

Invasive species establish successfully in new habitats especially due to their ability to include new species in their diet and due to the freedom from natural enemies. However, native species may also adapt to the use of new elements in their ecosystem. The planarian Endeavouria septemlineata, first recorded in Hawaii, was later found in Brazil. Recently, we found it in human-disturbed areas in southern Brazil and here we investigate its interactions with other invertebrates both in the field and in the laboratory. We observed the species in the field during collecting activities and hence maintained some specimens alive in small terraria in the laboratory, where we offered different invertebrate species as potential prey and also put them in contact with native land planarians in order to examine their interaction. Both in the field and in the laboratory, E. septemlineata showed a gregarious behavior and was found feeding on woodlice, millipedes, earwigs and gastropods. In the laboratory, specimens often did not attack live prey, but immediately approached dead specimens, indicating a scavenging behavior. In an experiment using the slug Deroceras laeve and the woodlouse Atlantoscia floridana, there was a higher consumption of dead specimens of woodlice and slugs compared to live specimens, as well as a higher consumption of dead woodlice over dead slugs. Four native land planarians of the genus Obama and one of the genus Paraba attacked and consumed E. septemlineata, which, after the beginning of the attack, tried to escape by tumbling or using autotomy. As a scavenger, E. septemlineata would have no impact on the populations of species used as food, but could possibly exclude native scavengers by competition. On the other hand, its consumption by native land planarians may control its spread and thus reduce its impact on the ecosystem.

The first troglobitic species of freshwater flatworm of the suborder Continenticola (Platyhelminthes) from South America.

Brazilian cave diversity, especially of invertebrates, is poorly known. The Bodoquena Plateau, which is located in the Cerrado Biome in central Brazil, has approximately 200 recorded caves with a rich system of subterranean water resources and high troglobitic diversity. Herein we describe a new troglobitic species of Girardia that represents the first obligate cave-dwelling species of the suborder Continenticola in South America. Specimens of the new species, which occur in a limestone cave in the Bodoquena Plateau, in the Cerrado biome, are unpigmented and eyeless. Species recognition in the genus Girardia is difficult, due to their great morphological resemblance. However, the new species can be easily recognized by a unique feature in its copulatory apparatus, namely a large, branched bulbar cavity with multiple diverticula.

A new species of Notogynaphallia (Platyhelminthes, Geoplanidae) extends the known distribution of land planarians in Chacoan province (Chacoan subregion), South America.

BACKGROUND: The subfamily Geoplaninae (Geoplanidae) includes land planarian species of the Neotropical Region. In Argentina, the knowledge about land planarian diversity is still incipient, although this has recently increased mainly in the Atlantic Forest ecosystem. However, other regions like Chacoan forests remain virtually unexplored. RESULTS: In this paper, we describe a new species of the genus Notogynaphallia of the Chacoan subregion. This species is characterized by a black pigmentation on the dorsum and a dark grey ventral surface. The eyes with clear halos extend to the dorsal surface. The pharynx is cylindrical. The main features of the reproductive system involve testes anterior to the ovaries, prostatic vesicle intrabulbar (with a tubular proximal portion and a globose distal portion) opening broadly in a richly folded male atrium, common glandular ovovitelline duct and female genital canal dorso-anteriorly flexed constituting a "C",female atrium tubular proximally and widening distally. CONCLUSIONS: This is the first report of the genus Notogynaphallia in Argentina (Chacoan subregion, Neotropical Region) which increases its geographic distribution in South America. Also, as a consequence of features observed in species of the genus, we propose an emendation of the generic diagnosis.

A new genus and species for the first recorded cave-dwelling Cavernicola (Platyhelminthes) from South America.

Species diversity of Brazilian cave fauna has been seriously underestimated. A karst area located in Felipe Guerra, northeastern Brazil, which is a hotspot of subterranean diversity in Brazil, has revealed more than 20 troglobitic species, most of them still undescribed. Based on recent samplings in this karst area, we document the occurrence of the suborder Cavernicola (Platyhelminthes) in South American hypogean environments for the first time and describe a new genus and species for this suborder. Hausera Leal-Zanchet & Souza, gen. n. has features concordant with those defined for the family Dimarcusidae. The new genus is characterized by two unique features, viz. an intestine extending dorsally to the brain and ovovitelline ducts located dorsally to the nerve cords, which is complemented by a combination of other characters. The type-specimens of Hauserahauseri Leal-Zanchet & Souza, sp. n. are typical stygobionts, unpigmented and eyeless, and they may constitute an oceanic relict as is the case of other stygobiotic invertebrates found in this karst area in northeastern Brazil.

A new species of land flatworm (Platyhelminthes: Continenticola) from areas of Araucaria Forest in southern Brazil.

The genus Cratera Carbayo et al. was proposed to encompass five species of Geoplaninae from southeastern Brazil that were mainly recorded in the state of São Paulo. Here we describe a new species of the genus, C. steffeni sp. nov., that occurs in areas of Araucaria Forest in southern Brazil, which augments the known distribution of Cratera. The new species is distinguished from others of the genus by its characteristic colour pattern and a combination of internal morphological characters.

A new species of Supramontana Carbayo & Leal-Zanchet (Platyhelminthes, Continenticola, Geoplanidae) from the Interior Atlantic Forest.

Supramontana argentina sp. nov. (Platyhelminthes, Continenticola, Geoplanidae) from north-eastern Argentina is herein described. The new species differs from Supramontana irritata Carbayo & Leal-Zanchet, 2003 from Brazil, the only species of this genus so far described, by external and internal morphological characters. Supramontana argentina sp. nov. is characterized by having a colour pattern with a yellowish median band, thin para-median black stripes, and two dark grey lateral bands on the dorsal surface. The most outstanding features of the internal morphology are a ventral cephalic retractor muscle almost circular in cross section, prostatic vesicle extrabulbar, tubular and very long, and penis papilla conical and blunt with a sinuous ejaculatory duct.