Morphology and Phylogenetic Position of Two New Gregarine Species (Apicomplexa: Eugregarinorida) Parasitizing the Lubber Grasshopper Taeniopoda centurio (Drury, 1770) (Insecta: Orthoptera: Romaleidae) in Mexico.

Eugregarines are understudied apicomplexan parasites of invertebrates inhabiting marine, freshwater, and terrestrial environments. Most currently known terrestrial eugregarines have been described parasitizing the gut from less than 1% of total insect diversity, with a high likelihood that the remaining insect species are infected. Eugregarine diversity in orthopterans (grasshoppers, locusts, katydids, and crickets) is still little known. We carried out a survey of the eugregarines parasitizing the Mexican lubber grasshopper, Taeniopoda centurio, an endemic species to the northwest of Mexico. We described two new eugregarine species from the gut of the host: Amoebogregarina taeniopoda n. sp. and Quadruspinospora mexicana n. sp. Both species are morphologically dissimilar in their life-cycle stages. Our SSU rDNA phylogenetic analysis showed that both species are phylogenetically distant to each other, even though they parasitize the same host. Amoebogregarina taeniopoda n. sp. clustered within the clade Gregarinoidea, being closely related to Amoebogregarina nigra from the grasshopper Melanoplus differentialis. Quadruspinospora mexicana n. sp. clustered within the clade Actinocephaloidea and grouped with Prismatospora evansi, a parasite from dragonfly naiads. Amoebogregarina taeniopoda n. sp. and Q. mexicana n. sp. represent the first record of eugregarines found to infect a species of the family Romaleidae.

A New Subspecies of Oxytricha granulifera (Hypotrichia: Oxytrichidae) from Mexico, with Notes on its Morphogenesis and Phylogenetic Position.

The genus Oxytricha Bory de Saint-Vincent in Lamouroux, Bory de Saint-Vincent and Deslongchamps, 1824 comprises about 38 species distributed worldwide and has been considered to be a nonmonophyletic group. Based on living observations, protargol preparations, and a small subunit ribosomal RNA (SSU rRNA) gene sequence, we describe a new subspecies Oxytricha granulifera chiapasensis n. subsp. This new taxon is morphologically characterized by undulating membranes basically in a Stylonychia-pattern, six dorsal kineties, size in vivo ca. 60-120 × 20-40 µm, 21-30 right and 21-31 left marginal cirri, 22-29 adoral membranelles, and spherical cortical granules arranged in longitudinal rows on the dorsal side. In terms of the SSU rRNA gene sequence, the new subspecies differs from populations of O. granulifera from GENBANK by 7-35 nucleotides. Phylogenetic analyses showed that Oxytricha granulifera gene sequences were nested into three groups, with the new subspecies included in one of them. Oxytricha granulifera chiapasensis n. subsp. is different from Oxytricha granulifera granulifera Foissner and Adam, 1983 and Oxytricha granulifera quadricirrata Blatterer and Foissner, 1988 based on: (i) undulating membranes in Stylonychia-pattern, (ii) formation of a sixth dorsal kinety during morphogenesis, (iii) the adoral membranelles number, and (iv) inhabiting freshwater habitats.

Epibiotic ciliate communities on the crayfish Procambarus (Austrocambarus) sp. cultivated in a rustic aquaculture in southern Mexico.

Rustic aquaculture systems represent a sustainable production alternative in the world, providing appropriate conditions for the establishment of epibiotic ciliates on crayfish. However, the impact of epibiotic ciliates on the aquaculture production has been insufficiently studied hitherto. The objective of this work was to identify the epibiotic ciliates on the surface of the crayfish Procambarus (Austrocambarus) sp. and analyze whether there is any correlation between seasonality and environmental conditions in an artificial pond in Chiapas, Mexico. The crayfish were collected during dry and rainy seasons to investigate their ciliate epibionts. Some physical and chemical parameters of the sampled water were measured and correlation matrix analyses were applied to detect possible relationships between the presence of ciliates and environmental parameters. Thirteen species belonging to the subclasses Peritrichia and Suctoria were identified. Significant differences were found for some environmental parameters and species recorded in both seasons. The present data could help to better understand the relationship bewteen the epibiotic ciliate community on crayfish and some environmental parameters in aquaculture systems.

Evolutionary insights and brief review of Loxodes Ehrenberg, 1830 (Ciliophora, Karyorelictea, Loxodidae) with description of a new species from Mexico.

Karyorelictids are a group of ciliates inhabiting marine and freshwater biotopes and possessing a non-dividing macronucleus. We describe a new freshwater species based on morphology and the 18S rRNA gene sequence data. Loxodes tziscaensis n. sp. can be easily distinguished from other Loxodes species by the arrangement of the nuclear apparatus and features of the buccal and somatic ciliature. The current proposed 18S rRNA phylogeny of Loxodes, including seven Loxodes species, shows two morphologically well-supported groups. Group A (L. rostrum, type species; L. vorax and L. tziscaensis n. sp.) includes species with a single nuclear group (two macronuclei and one micronucleus), in contrast to species of group B, which possess more than one nuclear group (L. striatus, L. magnus, L. kahli, L. penardi, and L. rex). We propose that the last common ancestor of Loxodes was a marine Remanella-like species possessing a single nuclear group. The division and differentiation of the micronucleus into a new macronucleus and the retention of the old macronuclei, independently of cell division, may have been two crucial processes during the evolution and diversification of Loxodes species with one nuclear group into species with multiple nuclear groups.

A checklist of species of Podophryidae (Ciliophora: Suctoria) as Epibionts of Crustaceans.

The epibiotic relationship between crustaceans and ciliates is frequent, for example, with species of suctorians. This work aims to present an updated list of species of the family Podophryidae worldwide distributed as epibionts of crustaceans and its geographic distribution. The available literature was checked, and all the species were verified. We found four species and four species indeterminata attached to 16 species of crustaceans. For the American continent, we obtained four species, and for Africa and Oceania, we did not find any species.

Cryptic Diversity in Paramecium multimicronucleatum Revealed with a Polyphasic Approach.

Paramecium (Ciliophora) systematics is well studied, and about twenty morphological species have been described. The morphological species may include several genetic species. However, molecular phylogenetic analyses revealed that the species diversity within Paramecium could be even higher and has raised a problem of cryptic species whose statuses remain uncertain. In the present study, we provide the morphological and molecular characterization of two novel Paramecium species. While Paramecium lynni n. sp., although morphologically similar to P. multimicronucleatum, is phylogenetically well separated from all other Paramecium species, Paramecium fokini n. sp. appears to be a cryptic sister species to P. multimicronucleatum. The latter two species can be distinguished only by molecular methods. The number and structure of micronuclei, traditionally utilized to discriminate species in Paramecium, vary not only between but also within each of the three studied species and, thus, cannot be considered a reliable feature for species identification. The geographic distribution of the P. multimicronucleatum and P. fokini n. sp. strains do not show defined patterns, still leaving space for a role of the geographic factor in initial speciation in Paramecium. Future findings of new Paramecium species can be predicted from the molecular data, while morphological characteristics appear to be unstable and overlapping at least in some species.

A checklist of species of the family Zoothamniidae (Ciliophora: Peritrichia), symbionts of crustaceans.

Eight genera are included in the family Zoothamniidae, and at the present the data of the epibiotic species on crustaceans is not updated. Thus, the main goal of the present work is to provide the checklist of zoothamniids worldwide reported as associated to several groups of crustaceans, including some data of their geographic distribution, and also to provide the complete list of species of this peritrich group. We obtained all available data of family Zoothamniidae and checked their taxonomic status. Then we provided the complete list of species including those reported as epibionts on crustacean hosts. We obtained 156 species included in eight genera of Zoothamniidae, being 85 species recorded as symbionts on 86 species of hosts. Genus Zoothamnium contained the higher number (56) of epibiotic species on crustaceans, and genus Pseudohaplocaulus has not been reported for any crustacean. The reviewed data highlights the concentration of records in Europe, and stands out the need to amplify sampling and studies in the Southern Hemisphere. The main groups of crustaceans infested by zoothamniids are the amphipods, isopods and decapods.

Periacineta mexicana n. sp. (Ciliophora, Suctoria, Discophryida), epizoic on Mexican backswimmers of the genus Buenoa (Insecta, Hemiptera, Notonectidae).

A new species of suctorian in the genus Periacineta, epibiotic on aquatic bugs (Hemiptera, Heteroptera, Nepomorpha), is described on the basis of morphological characteristics of the cell body, lorica, tentacular placement, and stalk, and its 18S rRNA partial sequence gene. Periacineta mexicana n. sp. is a loricate suctorian with elongate body and rounded apical region; tentacles are distributed randomly over apical region and not grouped into fascicles. Macronucleus in adult is elongate and located centrally. The suctorian usually forms closely aggregated pseudocolonies. We provide morphological data based on optical and scanning electron microscopy. A comparison with similar congeners, and emended diagnosis of the genus Periacineta also are provided. The ciliates were found attached to the first two pairs of legs of Mexican notonectids Buenoa pallens and Buenoa spp. (backswimmers).

Euplotes octocarinatus Carter, 1972 (Ciliophora, Spirotrichea, Euplotidae): Considerations on its morphology, phylogeny, and biogeography.

The cosmopolitan genus Euplotes Ehrenberg, 1830 comprises a highly distinguishable group of ciliates. However, details of the cell surface, the ciliature, and molecular data are still scarce for some species. We studied Euplotes octocarinatus Carter, 1972 from two Mexican freshwater bodies, providing data on its morphology, SSU rRNA gene sequence, and phylogeny. In addition, we obtained all data of previous records to show its geographic distribution and biogeographical pattern. The current populations showed some differences as compared with the original description and we provide an improved diagnosis. Morphologically, the species is very similar to Euplotes patella and E. daidaleos but differs by invariably having eight dorsolateral kineties (vs. nine in Euplotes patella and E. daidaleos), and lacking endosymbiotic green algae (vs. present in E. daidaleos). Phylogenetically, the Mexican population of E. octocarinatus nested with four isolates of the species lacking morphological characterization. The Euplotes octocarinatus described herein grouped into a fully-supported clade, which includes E. patella, E. amieti, E. daidaleos, E. eurystomus, E. woodruffi and E. aediculatus. Biogeographically, E. octocarinatus seems to have a wide distribution.

A checklist of ciliates (Ciliophora) inhabiting on ostracods (Crustacea, Ostracoda).

A compilation of the ciliated species found on freshwater and marine ostracods as epibiont or parasite (endobiont) has been carried out based on published records. The checklist includes the taxonomic position of each species of epibiontic and endobiontic ciliate, the species of basibiont ostracodes, the geographic zones and the bibliographic references where they were recorded. Altogether 7 suctorian, 29 peritrich, one apostome and one scuticociliatid species were listed. Two of recorded suctorian species are possible specific to marine ostracodes, whereas only one, Tokophrya sibirica to freshwater hosts. Fourteen species of peritrichs are likely specific to freshwater ostracodes, while three possible specific to marine ostracode hosts. Other suctorian and peritrich ciliate species were found on a variety of host taxa. One species of scuticociliatid was recorded as endobiont in ostracod.

Checklist of ciliates (Alveolata: Ciliophora) that inhabit in bromeliads from the Neotropical Region.

Species from almost all classes of ciliates are prone to be found inhabiting bromeliads in the Neotropics, from Mexico to Brazil, and the Antilles. Studies of ciliates recorded from bromeliads have been carried out from few bromeliad species, mainly in tropical forest. We compiled all available data of free living and sessile ciliates from bromeliads, including their geographic distribution and bromeliad identity. We provide a list of 170 ciliate species that have been recorded in 52 epiphytic and terrestrial bromeliad species, distributed in ten Neotropical countries . Most of the species belong to the Classes Oligohymenophorea, Colpodea and Spirotrichea. The largest number of ciliate species has been recorded in Brazil and Mexico. Bromeliothrix metopoides and Glaucomides bromelicola were the two species with the widest geographical distribution, 19 species have been recorded only in Mexico, 11 in the Antillean islands, and 89 only in Southamerica. Free living species prevailed over sessile species, and both represent 2% of the total ciliate species number. Sixteen ciliate species have been recorded only inhabiting in bromeliads. Although bromeliads show a high endemicity, their specificity is low in relation to ciliates.

Suctorians (Ciliophora: Suctoria) as epibionts of decapods of families Cambaridae and Pseudothelphusidae.

Epibiosis is common among ciliates and crustaceans because the calcified surface of the crustacean body offers a suitable site for colonization by the epibiont. Suctoria are among the most common epibiont ciliate groups of freshwater decapod crustaceans. The aim of this study is to increase knowledge of suctorians as epibionts of freshwater decapod crustaceans of the families Cambaridae and Pseudothelphusidae. Crustaceans were collected from three rivers and one pond in Chiapas, Mexico. Six species of suctorian epibionts were recorded: Acineta tuberosa, Podophrya maupasi, P. sandi, Tokophrya cyclopum, T. quadripartita and Trichophrya epistylidis. This is the first record of ciliate epibionts on members of family Pseudothelphusidae (represented by Phrygiopilus montebelloensis and Raddaus bocourti), and the first record of suctorians epibionts on Procambarus (Austrocambarus) sp. (Cambaridae).

Ciliate species from tank-less bromeliads in a dry tropical forest and their geographical distribution in the Neotropics.

The study of ciliate diversity in tropical environments remains scarce. In Neotropical forests, bromeliads are a common component of the vascular flora; bromeliads with tank morphology intercept rain water and detritus, resulting in the formation of a phytotelm, where heterotrophic protist communities like ciliates can establish. However, it is not known if ciliates inhabit tank-less bromeliads. For this reason, the goal of the present study was to investigate if ciliates can establish between the leaf axils of five terrestrial and epiphytic tank-less bromeliad species in a dry tropical forest in west Mexico. We collected samples of rain water and detritus from the leaf axils during the humid season of years 2015 and 2016. For ciliate taxonomical identification, we used optical microscopy, in vivo observation, and silver impregnation techniques. To summarize information about geographical distribution of ciliates identified at species level in the Neotropics, we provided their records from previous works, at country level with locality and georeferenciation. We recorded 27 taxa of ciliates, where the class Oligohymenophorea contained the largest richness of taxa. Drepanomonas revoluta, Leptopharynx bromeliophilus, and Tetrahymena sp. were recorded from all the species of bromeliads. Bromelia karatas was the species that hosted the largest number of ciliate taxa (22). Our results indicated that Glaucomides bromelicola, Gonostomum bromelicola, Leptopharynx bromelicola and L. bromeliophilus, species which are considered endemic to tank bromeliads, can also inhabit tank-less bromeliads. We provided previous records of 19 ciliate species from eleven countries within the Neotropical region, and Bromeliothrix metopoides was the species most frequently recorded in Neotropical countries (9). Therefore, tank-less bromeliads can constitute a temporal habitat for ciliates, and function as cysts reservoirs in environments with a pronounced seasonality like dry tropical forests.

A checklist of species of Vorticellidae (Ciliophora: Peritrichia) epibionts of crustaceans.

The family Vorticellidae includes 21 genera, mainly with species worldwide distributed, as free-living, and attached to diverse organisms as epibionts. The goal of this paper is to provide an updated checklist of the worldwide vorticellids reported attached to the crustaceans. We revised and checked all available records of species of vorticellids including their crustacean hosts. We found 96 vorticellid species belonging to 14 genera with at least a report on the body of 81 species of crustaceans. Four genera of ciliates (Vorticella, Pseudovorticella, Carchesium and Intranstylum) had the major number of species living as epibionts. We provide also the complete list of the species of vorticellids, and conclude the need of a taxonomic revision of the family.

New record of Epistylishentscheli (Ciliophora, Peritrichia) as an epibiont of Procambarus (Austrocambarus) sp. (Crustacea, Decapoda) in Chiapas, Mexico.

Epibiosis is very common between crustaceans and ciliates where the calcified surface of the crustacean body provides a suitable substrate for ciliate colonization. The aim of this contribution is to provide data about a new record between the epistylid ciliate Epistylishentscheli Kahl, 1935, and the crayfish Procambarus (Austrocambarus) sp. The distribution of the epistylid on the basibiont body and its cellular/colonial characteristics were analyzed. Procambarus (Austrocambarus) sp. harbored colonies of E.hentscheli only on the pereiopods. This is the first record of this peritrich ciliate as an epibiont on Crustacea, having been previously found on algae and fish.

An annotated checklist of species in the family Lagenophryidae (Ciliophora, Oligohymenophorea, Peritrichia), With a brief review of their taxonomy, morphology, and biogeography.

The genera Lagenophrys Stein, 1852, Paralagenophrys Clamp, 1987, Clistolagenophrys Clamp, 1991, and Operculigera Kane, 1969 make up the family Lagenophryidae and, together, contain more than 80 species worldwide that live mainly as ectosymbionts of crustaceans. Lagenophryids are characterized by possession of a lorica, but the lorica aperture and the associated parts of the body differ between genera. Despite their widespread distribution, relatively few papers have been published on lagenophryids in recent years, and the present paper is intended to promote research on lagenophryids by providing an annotated checklist of all known species and a brief review of the family, including data of their geographic distribution and their hosts.

A taxonomic revision of order Pilisuctorida (Ciliophora, Apostomatia) with keys to the subordinate taxa.

The article is to present a review of the taxonomic literature of pilisuctorid ciliates and new data from samples taken in Ukraine and Mexico. In order to contribute to the knowledge of this specialised ciliate group, the present article contains the general characteristics of order Pilisuctorida members, information on morphology and life cycles of all its representatives and some own data on distribution of three species of genus Conidophrys in Ukraine and Mexico. The taxonomic composition of this order, and diagnoses of all taxa are also included, as well the keys to the taxa.

Free-living ciliates from epiphytic tank bromeliads in Mexico.

The ciliate diversity of Mexican bromeliads is poorly known. We studied the ciliate community of two species of epiphytic tank bromeliads from 48 individuals of Tillandsia heterophylla and four of T. prodigiosa. The bromeliads occurred on over 22 tree host species. Samples were collected during 2009 and 2010 in a mountain cloud forest and in two coffee plantations and in a pine-oak forest. The ciliates were identified in live and protargol preparations. We recorded 61 ciliate species distributed in 39 genera grouped in eight classes. Ten species were frequent in the 52 samples (20 ± 3.2) and Leptopharynx bromeliophilus was the most frequent recorded in 25 samples. Thirty-three species are new for the fauna of Mexico, 24 species have been recorded for the first time in tank bromeliads. The classes Spirotrichea, Oligohymenophorea and Colpodea presented the highest number of species, 16, 14, and 12, respectively. Colpoda was the most species-rich genus being present with six species. A low similarity between areas and seasons was obtained with Jaccard's index. We conclude that the two bromeliads species host a rich ciliate diversity whose knowledge contributes to the question of ciliate distribution and specifically, in tank bromeliads.