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Acanthogyrus (Acanthosentis) maroccanus (Dollfus, 1951), an insufficiently described quadrigyrid acanthocephalan of cyprinid fishes from Northwest Africa, is redescribed based on recently collected specimens from the Algerian barb Luciobarbus callensis (Valenciennes) in Algeria. Newly observed morphological features for A. (A.) maroccanus include the arrangement of proboscis hooks (not in regular circles), the male reproductive structures extending into the copulatory bursa and the presence of a para-receptacle structure and vaginal sleeve. The mechanism of copulation of this acanthocephalan is described based on several copulating pairs. The phylogenetic position of A. (A.) maroccanus within Eoacanthocephala was assessed based on partial 28S rDNA sequences. Maximum likelihood and Bayesian inference analyses placed A. (A.) maroccanus in a clade with Palliolisentis (Demidueterospinus) ophiocephalus (Thapar, 1931), both species included in the Quadrigyridae, the only family within the Gyracanthocephala.
Our study aimed at examining the phylogenetic position of the newly-found Setaria nematodes obtained from the red deer (Cervus elaphus) based on sequences of the mitochondrial cytochrome c oxidase subunit 1 (COX-1). Alignment and phylogenetic analyses, as well as SEM microscopic analysis, revealed the presence of two Setaria species: S. cervi and S. tundra. Setaria tundra was noted in only one individual, a calf of the red deer, while S. cervi was observed in three stages, two hinds and one calf of the red deer. According to our knowledge, it is the first case of S. cervi in the red deer in Poland confirmed in molecular studies and also the first case of S. tundra infection in the red deer.
Chapter 8 summarizes the ramifications of the Laryngeal Articular Model for the phonetic description of voice quality and investigates its place in phonetic theory. Predispositions for background voice qualities to underlie different vowel qualities are identified. It is argued that there is a parallel between the coarticulatory path that infants follow in their earliest speech development and elements in the process of phonetic sound change. The laryngeal articulator as an enabler of sound change is explored. Theories of ontogenetic speech development are reviewed (concepts of ‘speechlikeness,’ autogeneration, frame/content, cyclicity, reduplication, variegation) and reinterpreted to reflect the scope of laryngeal behaviour outlined in this volume. The discussion evaluates the implications of the revised view of laryngeal phonetic behaviour for the phylogeny of human speech. The physiology of the larynx is shown to permit a wider range of speech production than formerly assumed and to accommodate, on phonetic grounds, an earlier time period for the appearance of speech in human ancestors, as hypothesized in recent anthropological and genetic research.
Digenea Carus, 1863 represent a highly diverse group of parasitic platyhelminths that infect all major vertebrate groups as definitive hosts. Morphology is the cornerstone of digenean systematics, but molecular markers have been instrumental in searching for a stable classification system of the subclass and in establishing more accurate species limits. The first comprehensive molecular phylogenetic tree of Digenea published in 2003 used two nuclear rRNA genes (ssrDNA = 18S rDNA and lsrDNA = 28S rDNA) and was based on 163 taxa representing 77 nominal families, resulting in a widely accepted phylogenetic classification. The genetic library for the 28S rRNA gene has increased steadily over the last 15 years because this marker possesses a strong phylogenetic signal to resolve sister-group relationships among species and to infer phylogenetic relationships at higher levels of the taxonomic hierarchy. Here, we have updated the database of 18S and 28S rRNA genes until December 2017, we have added newly generated 28S rDNA sequences and we have reassessed phylogenetic relationships to test the current higher-level classification of digeneans (at the subordinal and subfamilial levels). The new dataset consisted of 1077 digenean taxa allocated to 106 nominal families for 28S and 419 taxa in 98 families for 18S. Overall, the results were consistent with previous higher-level classification schemes, and most superfamilies and suborders were recovered as monophyletic assemblages. With the advancement of next-generation sequencing (NGS) technologies, new phylogenetic hypotheses from complete mitochondrial genomes have been proposed, although their power to resolve deep levels of trees remains controversial. Since data from NGS methods are replacing other widely used markers for phylogenetic analyses, it is timely to reassess the phylogenetic relationships of digeneans with conventional nuclear rRNA genes, and to use the new analysis to test the performance of genomic information gathered from NGS, e.g. mitogenomes, to infer higher-level relationships of this group of parasitic platyhelminths.
The new species Lenonchium zanjanense sp. n. is described from a natural habitat of Zanjan province, Iran, including line, light microscopy and scanning electron microscopy illustrations and a molecular (18S, 28S) study. It is characterized by its 3.50–4.51 mm long body, rounded lip region, continuous and 13.5–15.5 µm broad, odontostyle 21–24 µm long, neck 362–490 µm long, double guiding ring, pharyngeal expansion 190–285 µm long, female genital system didelphic–amphidelphic, uterus simple and 185–320 µm long or 3.4–5.9 times the corresponding body diameter, vulva nearly equatorial (V = 45–53), tail conical-elongated to filiform (90–165 µm, c = 23–43, c′ = 2.4–5.3) with three or four mucro-like projections at the tip, spicules 58–64 µm long and 16–21 contiguous ventromedian supplements ending at the level of the anterior end of the spicules. The taxonomy of the genus is updated, with an emended diagnosis, list of species, key to their identification and a compendium of their main morphometrics. Lenonchium asterocaudatum is regarded as identical and a junior synonym of L. denticaudatum. New insights into the phylogeny of the group are also provided, and the classification of Lenonchium within Nordiidae is seriously questioned.
Englerophytum and Synsepalum are two closely related genera of trees and shrubs from the African tropics. Previous molecular studies have shown that these genera collectively form a clade within the subfamily Chrysophylloideae (Sapotaceae). However, little is known about the inter-relationships of the taxa within the Englerophytum–Synsepalum clade. In this study, nuclear ribosomal DNA and plastid trnH–psbA sequences were used to estimate the phylogeny within the clade. Results indicate that the clade consists of six major lineages, two composed solely of taxa from the genus Englerophytum and four composed of taxa from the genus Synsepalum. Each lineage can be distinguished by suites of vegetative and floral characters. Leaf venation patterns, calyx fusion, style length and staminodal structure were among the most useful characters for distinguishing clades. Some of the subclades within the Englerophytum–Synsepalum clade were also found to closely fit descriptions of former genera, most of which were described by Aubréville, that have since been placed in synonymy with Englerophytum and Synsepalum. The clade with the type species of Englerophytum also contains the type species of the genera Wildemaniodoxa and Zeyherella, which are confirmed as synonyms.
To date, there are very few sequence data for Cyperaceae from mainland Southeast Asia. The aim of the present study was to contribute nuclear ribosomal internal transcribed spacer (ITS) sequences of selected species of Cambodian Cyperaceae to the overall phylogeny of the family. We generated ITS sequences of 38 accessions representing 26 species from Cambodia and used these sequences for phylogenetic analysis together with similar sequences from the National Center for Biotechnology Information GenBank. Our results corroborate recent phylogenetic work in the family and largely confirm established tribal relationships. The backbone of the phylogenetic tree of species-rich genera that have undergone rapid radiations is often weakly resolved (e.g. in Fimbristylis and in the C4 clade of Cyperus). Cryptic variation was revealed in the taxonomically difficult group of Fimbristylis dichotoma, with samples of this taxon appearing in two distinct clades within Fimbristylis. Further addition of geographically spread accessions of taxa will improve our understanding of the complex biogeographical history of the genera in the family. Eleocharis koyamae Tremetsb. & D.A.Simpson is proposed as a new name for E. macrorrhiza T. Koyama.
A new species of Chaenotheca, C. biesboschii, has been found in the freshwater tidal area of the Biesbosch in the Netherlands, a national park well known for harbouring several rare and threatened mosses and lichens. A phylogenetic analysis of the ITS region revealed some strongly supported infrageneric clades in Chaenotheca which were given informal names, and some were assigned provisional names in anticipation of generic recognition. The analysis also showed that the new species differed in the sequenced region from other European Chaenotheca species. Chaenotheca biesboschii might be mistaken for C. gracillima but, in addition to a considerable difference in the ITS region, it also differs from this species in morphology. It is also similar to C. servitii but again differs in morphology. Chaenotheca biesboschii inhabits decorticated wood in the oldest stages of forest development of abandoned willow coppices. In 2016 and 2017 a fairly large population was found in an area comprising several square kilometres. In the Biesbosch area, extensive woodlands have developed only since the 1950s and therefore C. biesboschii might have been recently established in the area, possibly following climatic warming. The new species is characterized by having an immersed, glaucous green thallus; apothecia 0·9–1·4 mm high; capitulum on the lower side when young with a ring-like thickening covered by a yellow pruina; when mature with a rusty brown pruina on the capitulum and upper part of stalk; spherical spores, 3·5–5·5 µm diam., ornamented by irregular cracks, medium brown; photobiont Stichococcus. A key to the European species of Chaenotheca is provided.
This contribution provides morphological and molecular data for one new and one known species of the genus Pungentus. The first species, P. azarbaijanensis n. sp., was recovered from the rhizospheric soil of grasses, collected in West Azarbaijan province, Iran, and was characterized by 2082–2365 μm long females having an angular lip region separated from the rest of the body by a constriction, 33–35 μm long odontostyle, vulva at 43.5–51.0%, 27.0–29.5 μm long rounded-conoid tail, and males unknown. It was compared morphologically with five species: P. angulosus, P. crassus, P. marietani, P. parapungens and P. pungens, which have didelphic-amphidelphic female reproductive system, body longer than 1.5 mm, and odontostyle longer than 20 μm. The second species, P. engadinensis, was recovered in three different regions of Iran (Mazandaran, Semnan and East Azarbaijan provinces). It was compared morphologically with some other populations reported all over the world. Besides morphological studies, molecular phylogenetic studies using partial sequences of 28S rDNA D2-D3 fragments were performed, and the phylogenetic relations of the two Iranian populations with other species and genera were discussed.
Seed reserves play vital roles in seed germination and seedling growth and their variation may be related to various environment factors, plant traits and phylogenetic history. Here, the evolutionary correlation associated with seed mass and altitude and carbon (C), nitrogen (N) and phosphorus (P) allocation of seeds among 253 alpine herbaceous plants was tested. In this study, phylogeny had strong limitations on nutrient allocation of seeds across species, and species from younger phylogenetic groups tended to have higher N and P contents, which might be considered as the evolutionary selection of seed plants. Higher seed N and P content would help seedlings to gain more survival chance and stronger competitive capacity, and their progeny would be more likely to be preserved. When phylogeny was considered, altitude only had a significant positive effect on P content, but the negative effects on seed mass were all expressed. The independent effects of altitude and seed mass suggest that the nutrient allocation of seeds might be affected by both environment and plant traits. In addition, altitude and seed mass displayed partial overlapping effects on nutrient allocation of seeds. The negative effects of seed mass were affected slightly by altitude, whereas altitude only had a significant positive effect on P content when seed mass was controlled. Above all, seed P content showed obvious and general correlations with seed mass, altitude and age of clade, which indicated that higher seed P content might be an adaptive selection of species associated with growth and survival of progeny.
Swimming propagules (embryos and larvae) are a critical component of the life histories of benthic marine animals. Larvae that feed (planktotrophic) have been assumed to swim faster, disperse farther and have more complex behavioural patterns than non-feeding (lecithotrophic) larvae. However, a number of recent studies challenge these early assumptions, suggesting a need to revisit them more formally. The current review presents a quantitative analysis of swimming speed and body size in planktotrophic and lecithotrophic propagules across five major marine phyla (Porifera, Cnidaria, Annelida, Mollusca and Echinodermata). Results of the comparative study showed that swimming speed differences among ciliated propagules can be driven by taxonomy, adult mobility (motile vs sessile) and/or larval nutritional mode. On a phylogenetic level, distinct patterns emerge across phyla and life stages, whereby planktotrophic propagules swim faster in some of them, and lecithotrophic propagules swim faster in others. Interestingly, adults with sessile and sedentary lifestyles produce propagules that swam faster than the propagules produced by motile adults. Understanding similarities and differences among marine propagules associated with different reproductive strategies and adult lifestyles are significant from ecological, evolutionary and applied perspectives. Patterns of swimming can directly impact the dispersal/recruitment potential with incidence on the design of larval rearing methods and marine protected areas.
Neoechinorhynchus is one of the most speciose genera of acanthocephalans, with approximately 116 described species. A recent study, aimed at establishing the genetic diversity of Neoechinorhynchus in Middle American freshwater fishes, validated nine species molecularly and morphologically and revealed the existence of 10 putative candidate species. Neoechinorhynchus golvani, a parasite commonly found in cichlids throughout Middle America with an allegedly large intraspecific morphological variability, was found to represent a species complex; species delimitation methods uncovered three additional genetic lineages. Here, we re-analyse the morphological and molecular data for N. golvani species complex infecting cichlids in that geographical area. A multivariate analysis of variance (MANOVA) was conducted particularly for the length of apical, middle and posterior hooks of the species/lineages of Neoechinorhynchus in cichlids, revealing morphological variation in the length of apical hooks for Lineage 8, although no morphological distinction was observed for Lineages 9 and 10. A new concatenated phylogenetic analysis of one mitochondrial and two ribosomal DNA genes was used to further corroborate the species delimitation among lineages; Neoechinorhynchus Lineage 8 was found to be morphologically and genetically distinct from its sister taxa, N. golvani and other two undescribed genetic lineages, and is formally described as a new species. Neoechinorhynchus costarricense n. sp. is described from the intestines of eight species of cichlids in Costa Rica. The new species is distinguished from the other species/lineages of Neoechinorhynchus in cichlids mainly by the size of the apical hooks of the proboscis.
Species identification based on the morphometry of opisthaptoral hard parts, in combination with internal transcribed spacer ribosomal DNA (ITS rDNA) region sequences, confirmed the presence of four viviparous Gyrodactylus von Nordman, 1832 (Plathyhelminthes, Monogenea) species on Nototheniid fish from the Prince Gustav Channel (Weddell Sea, Antarctica). Gyrodactylus antarcticus Gusev, 1967 was found mostly on Trematomus newnesi Boulenger (93 specimens) but also on T. bernacchii Boulenger (one specimen), the latter representing a new host record for this species. Gyrodactylus byrdi Hargis & Dillon, 1968 and G. coriicepsi Rokicka, Lumme & Ziętara, 2009 were recorded on their type hosts, T. newnesi and Notothenia coriiceps Richardson, respectively. Gyrodactylus wilkesi Hargis & Dillon, 1968 was found mostly on the fins of T. bernacchii (29 specimens), but also on T. hansoni Boulenger (one specimen) and T. newnesi (three specimens). The finding of G. wilkesi on T. newnesi represents a new host record. The low number of Gyrodactylus specimens may indicate an accidental infection. The occurence of all four Gyrodactylus species in the Prince Gustav Channel represents a new locality record. According to phylogentic methods, the newly redescribed monogeneans belong to the Antarctic lineage, forming a sister group to North American and European marine Gyrodactylus species, and consist of two species groups, one comprising G. coriicepsi and G. nudifronsi Rokicka, Lumme & Ziętara, 2009, and the other G. anarcticus and G. wilkesi.
The Carboniferous tetrapod Crassigyrinus scoticus is an enigmatic animal in terms of its morphology and its phylogenetic position. Crassigyrinus had extremely reduced forelimbs, and was aquatic, perhaps secondarily. Recent phylogenetic analyses tentatively place Crassigyrinus close to the whatcheeriids. Many Carboniferous tetrapods exhibit several characteristics associated with terrestrial locomotion, and much research has focused on how this novel locomotor mode evolved. However, to estimate the selective pressures and constraints during this important time in vertebrate evolution, it is also important to study early tetrapods like Crassigyrinus that either remained aquatic or secondarily became aquatic. We used computed tomographic scanning to search for more data about the skeletal morphology of Crassigyrinus and discovered several elements previously hidden by the matrix. These elements include more ribs, another neural arch, potential evidence of an ossified pubis and maybe of pleurocentra. We also discovered several additional metatarsals with interesting asymmetrical morphology that may have functional implications. Finally, we reclassify what was previously thought to be a left sacral rib as a left fibula and show previously unknown aspects of the morphology of the radius. These discoveries are examined in functional and phylogenetic contexts.
The genus Megalobatrachonema is a rare group of nematode parasites within Ascaridida. The systematic status of Megalobatrachonema in the superfamily Cosmocercoidea (Ascaridida) has long been controversial. The relationship of Megalobatrachonema and Chabaudgolvania remains unsolved. In the present study, a new species of Megalobatrachonema, M. hainanensis sp. nov., was described based on specimens collected in Amolops hainanensis (Boulenger) and Hylarana spinulosa (Smith) (Amphibia: Anura) from Hainan Island, China. The large ribosomal DNA (28S) and internal transcribed spacer (ITS1-5.8S-ITS2) were also sequenced for molecular identification and phylogenetic study. Phylogenetic analyses using maximum likelihood (ML) inference and Bayesian inference (BI) based on 28S and ITS1 sequence data, respectively, supported that Megalobatrachonema is a member of the family Kathlaniidae. In addition, the genetic comparison and phylogenetic results based on ITS1 sequence data also supported that the genus Chabaudgolvania should be considered as a synonym of Megalobatrachonema.
The pestiviruses bovine viral diarrhea virus 1 (BVDV-1), 2 (BVDV-2), and HoBi-like (HoBiPeV) are endemic among Brazilian cattle, the world's largest commercial bovine herd. In the last two decades (1998–2018) over 300 bovine pestiviruses have been partially or fully sequenced in Brazil, including viruses from different regions, different epidemiological backgrounds, and associated with diverse clinical presentations. Phylogenetic analysis of these viruses demonstrated a predominance of BVDV-1 (54.4%), with subgenotypes −1a (33.9% of total) and −1b (16.3%) being more frequent and subgenotypes −1d, −1e, and −1i at very low frequencies. The overall BVDV-2 frequency was 25.7% but it varied largely by region, reaching up to 48% in Southern states. BVDV-2b was the predominant subgenotype (84.8% of BVDV-2), followed by BVDV-2a (8.86%). HoBiPeV accounted for 19.9% (61/307) of the genotyped viruses and were detected at high frequency in cattle from Northeastern states. These findings demonstrate a unique mix of pestivirus species and subgenotypes, unlike that seen in Europe or North America. The design of effective diagnostic tools, vaccines, and control programs for limiting bovine pestivirus infections in Brazil must take into consideration this unique mix of viruses. This article provides a critical review of two decades of genetic identification of pestiviruses in Brazil.
Despite the reduction in the prevalence of soil-transmitted helminthiases in many regions of the world, morbidity rates remain high in some rural regions. The Kato–Katz technique is a simple, inexpensive and field-applicable tool commonly used for the diagnosis and worm-burden characterization of these infections. Molecular studies have revolutionized our understanding of the epidemiology and evolutionary genetics of parasites. In this study we recovered helminthic DNA from Kato–Katz slides (n = 93) prepared in 2011 in the Brazilian Amazon. We achieved DNA recovery by polymerase chain reaction (PCR) in 84% of cases for Ascaris sp. and 75% of cases for hookworms. The sequencing confirmed the specific species of the amplicons. The slides stored for a few years could be analysed using this methodology, allowing access to DNA from a large collection of samples. We must consider the Kato–Katz thick smears as a source of helminth DNA. This can significantly reduce logistical difficulties in the field in terms of obtaining, preserving, transporting and initial processing of samples.
Little is known about the genetic and morphological characters of Taenia ovis. The purpose of the present study was to characterize sheep isolates of T. ovis using rostellar hook morphometry as well as mitochondrial genes sequence analysis. Ninety sheep specimens of Cysticercus ovis were collected from 18 slaughterhouses in Iran. The mean ± s.d. for total length of large and small hooks were 174.1 ± 6.4 and 116.7 ± 5.4 µm, respectively. CO1 and 12S rRNA sequence analysis showed 11 and nine haplotypes, respectively. The level of pairwise nucleotide variations between individual haplotypes of CO1 and 12S rRNA genes were 0.3–1.1 and 0.2–1.0%, respectively. Level of nucleotide variation in CO1 and 12S rRNA between T. ovis haplotypes from present study and eight other Taenia species was found to be 11.3–17.8 and 5.3–16.3%, respectively. Phylogenetic analysis clustered all T. ovis isolates into a single clade comprised of the all CO1 and 12S rRNA haplotypes. CO1 nucleotide difference between T. ovis ovis and T. asiatica was 13.6% that is lesser than the corresponding difference between T. ovis ovis and T. ovis krabbei, warranting the designation of two separate species as T. ovis and T. krabbei. Interclass correlation coefficients showed that there was no significant association between rostellar hook length variation and the variability of the mitochondrial genes.
The early actinopterygian Mesopoma planti is reassigned to a new genus on the basis of data obtained from high-resolution computed tomography (CT) scans of an unusually well-preserved specimen from the Early Pennsylvanian of Lancashire, UK. The former M. planti is joined by two further Mesopoma species from the Late Mississippian of Scotland. CT scans of the key planti specimen bring to light new details of the dermal skull, pectoral girdle and fin. Among the cranial features, CT data reveal a specialised, anteriorly projecting preopercular bone, the location of the spiracular duct opening, presence of a so-called coronoid process on the lower jaw and the full three-dimensional shape of the snout. Of the pectoral girdle and fin, for the first time in a Palaeozoic actinopterygian it has been possible to complete a three-dimensional reconstruction of the entire endoskeleton in articulation. The fin presents new diversity within a conservative general pattern, revealing for the first time a double propterygium. Girdle shape shows that the fin orientation is derived: rotated with the leading edge dorsalmost. These details are used to identify unexploited character states for use in phylogenetic analyses, while functional implications of the fin and girdle suggest advanced locomotory control emerging among different groups of post-Devonian ray-finned fishes.
In the present paper, the phylogenetic relationships between genera, subfamilies and families of the Hemiuroidea are explored. Twelve new sequences of 28 rDNA and data taken from GenBank (NSBI) on 43 species affiliated to 34 genera were included in the analysis. Most of the hemiuroidean trematodes form two highly supported clades (A and B), which are sister groups to each other. Hemipera manteri joined with Gonocerca spp. with moderate statistical support. This clade is basal relative to the clades A and B. Сlade A is polytomic and contains representatives of the families Accacoeliidae, Syncoeliidae, Didymozoidae, Hirudinellidae and Sclerodistomidae, and derogenid subfamilies Derogeninae and Halipeginae. At the same time, the Syncoeliidae, Hirudinellidae and Accacoeliidae form a well-supported monophyletic group. The phylogenetic relationship between Derogeninae and Halipeginae is poorly resolved. Сlade B unites the isoparorchiid, bunocotylid, lecithasterid and hemiurid trematodes. Our data re-establishes the family Bunocotylidae, which consists of two subfamilies, Opisthadeninae and Bunocotylinae, and the Machidatrema chilostoma + Hysterolecithoides frontilatus group. The Bunocotylidae is the sister group to the Hemiuridae + Lecithasteridae group and the Isoparorchiidae is a basal relative to the representatives of these three hemiuroid families.