Undescribed Species Research Articles

Reticulate Evolution in Japanese Sceptridium (Ophioglossaceae), including diploid, tetraploid, and hexaploid species.

Homospory represents an intriguing reproductive strategy, including the potential for gametophytic selfing, considered the ultimate form of selfing. The plants of the fern genus Sceptridium (Ophioglossaceae) are generally considered to be predominantly selfing, making them suitable for analyzing speciation patterns and intraspecific variation characteristic of homosporous plants. The Sceptridium species in Japan not only exhibit variations in ploidy, including diploids, tetraploids, and hexaploids, but also pose taxonomic challenges due to morphologically distinctive forms that are difficult to be assigned to previously described species. We developed multiple single-copy nuclear markers and aimed to identify ancestral species of the polyploids and to elucidate the evolutionary entity of an undescribed species. Chromosome counts and analyses of individual gene trees confirmed that S. japonicum, S. atrovirens, and the undescribed species are allohexaploids originating through hybridization between a maternal tetraploid ancestor closely related to S. formosanum and a paternal diploid ancestor closely related to extant diploid species. By identifying homoeologs derived from the diploid ancestor, we successfully analyzed the phylogenetic relationship between the diploid ancestor of the hexaploid species and extant diploid species using the multispecies coalescent model. The examined undescribed species could not be distinguished from S. atrovirens by multiple nuclear markers or by SNP data obtained from multiplexed ISSR genotyping by sequencing. This species appears to represent one of the polymorphisms of S. atrovirens, which was fixed through gametophytic selfing.

Review of hymenolepidids associated with the tribe Sudarikovinini Spassky, 1991 (Eucestoda, Hymenolepididae), with descriptions of two unrecognized species from the Shaw's jird Meriones (Pallasiomys) shawi (Duvernoy) (Rodentia: Muridae: Gerbillinae) from Northern Africa.

The taxonomic position of the hymenolepidid cestode genera Paraoligorchis Wason et Johnson, 1977 and Sudarikovina Spassky, 1951 from gerbils and ground squirrels from Africa, Southwest Asia and South Asia is clarified based on re-examination of type and voucher materials. The generic diagnosis of Sudarikovina is amended. The main corrections for diagnostic characters at the generic level include: irregularly-spaced transverse anastomoses of ventral osmoregulatory canals; presence of both specimens with dextral genital pores and specimens with sinistral genital pores; vagina clearly covered externally by a dense layer of intensely-stained cells; young uterus reticulate and fully-developed uterus with a labyrinthine structure. The previously proposed generic character of a minimum number of testes per proglottis of species of Sudarikovina is reduced to 3. The genus Paraoligorchis is recognized as a junior synonym of Sudarikovina. For the type species of Paraoligorchis, P. taterae Wason et Johnson, 1977, the replacement name Sudarikovina wasojohni nom. nov. is proposed, thus resolving the secondary homonymy of the transferred species and its senior homonym S. taterae Hunkeler, 1972. The validity of the tribe Sudarikovinini sensu Spassky (1991), to which Paraoligorchis and Sudarikovina had been attributed, cannot be resolved due the lack of molecular data. The cestode specimens from Shaw's jird, Meriones (Pallasiomys) shawi (Duvernoy) from Algeria and Tunisia, initially designated as Hymenolepis sp. by Joyeux (1923) and Hymenolepis procera Janicki, 1904 by Joyeux & Foley (1930), represent undescribed species of Sudarikovina, which are described in this study as Sudarikovina pentatesticulata sp. n. and S. tetratesticulata sp. n., respectively. The main diagnostic characters of the two new species, which distinguish them from their congeners, are the number and the size of the testes.

Idiopyrgus Pilsbry, 1911 (Gastropoda, Tomichiidae): a relict genus radiating into subterranean environments

The freshwater gastropod fauna in Brazil presently consists of circa 180 known species, though this is deemed an underestimate. The little-studied cavern faunas have been considered a potential source of undescribed species, particularly regarding the Truncatelloidea. Here, based on new collection efforts in caves in Bahia state, Brazil, we describe two new troglobitic species of Idiopyrgus (Tomichiidae): Idiopyrgus eowynaesp. nov. and Idiopyrgus meriadocisp. nov. We improve upon the previous molecular phylogeny of this genus and take the opportunity to reclassify Spiripockia umbraticola Simone & Salvador, 2021, as Idiopyrgus umbraticola (Simone & Salvador, 2021), comb. nov. The genus Idiopyrgus now contains ten species, reinforcing the presence of Tomichiidae in Brazil as a relict lineage that largely radiated into cave environments. Furthermore, considering that so far only snapshots of their biology are known, we have collated all the information currently available from the literature and field observations on the cavernicolous species of Idiopyrgus. Notably, due to human action in the area where the two new species have been found, they can already be considered vulnerable.

What Predicts Gene Flow During Speciation? The Relative Roles of Time, Space, Morphology and Climate.

The processes that restrict gene flow between populations are fundamental to speciation. Here, we develop a simple framework for studying whether divergence in morphology, climatic niche, time and space contribute to reduced gene flow among populations and species. We apply this framework to a model system involving a clade of spiny lizards (Sceloporus) occurring mostly in northeastern Mexico, which show striking variation in morphology and habitat among closely related species and populations. We developed a new time-calibrated phylogeny for the group using RADseq data from 152 individuals. This phylogeny identified 12 putative species-level clades, including at least two undescribed species. We then estimated levels of gene flow among 21 geographically adjacent pairs of species and populations. We also estimated divergence in morphological and climatic niche variables among these same pairs, along with divergence times and geographic distances. Using Bayesian generalised linear models, we found that gene flow between pairs of lineages is negatively related to divergence time and morphological divergence among them (which are uncorrelated), and not to geographic distance or climatic divergence. The framework used here can be applied to study speciation in many other organisms having genomic data but lacking direct data on reproductive isolation. We also found several other intriguing patterns in this system, including the parallel evolution of a strikingly similar montane blue-red morph from more dull-coloured desert ancestors within two different, nonsister species.

A new species of Fissidentalium (Scaphopoda: Dentaliidae) in association with an actinostolid anemone from the abyssal Labrador Sea

The benthic biodiversity of the abyssal Labrador Sea was investigated using Agassiz trawl and in situ imagery. A megafaunal scaphopod associated with an epizoic anemone was recovered from soft sediments. Morphological and molecular investigations revealed the scaphopod to be an undescribed species in the dentaliid genus Fissidentalium P. Fischer, 1885. The new scaphopod species is characterised by a large size for the genus, is moderately curved, with numerous narrow, longitudinal ribs (60 ribs at 11 mm diameter ventral aperture), a dentaliid radula, and is described herein as Fissidentalium aurae sp. nov. The new species shows a close genetic relationship to congeners of Fissidentalium and separates from the sister genera Dentalium Linnaeus, 1758 and Antalis H. Adams & A. Adams, 1854. Genetic COI barcoding of the epizoic anemone suggests the species is a member of the family Actinostolidae Carlgren, 1932. The discovered association of a burrowing scaphopod with an epifaunal anemone at abyssal depth is a new record for the region and is indicative of how little is known about symbioses in the deep sea.

First records of moss-feeding aphids (Hemiptera: Sternorrhyncha: Aphididae) from Hungary

Four species of moss-feeding aphids were captured by Berlese samples collected in Heves county, NE Hungary. These data are establishing the first Hungarian records of moss-dwelling aphids, all the four species and three genera are reported from the country for the first time. Muscaphis (Aspidaphium) eschirichi (Börner, 1939) was yielded from a colony of Palustriella commutata (Hedw.) Ochyra collected from rock in a karst spring outlet, Muscaphis (Muscaphis) musci Börner, 1933 was yielded from colonies of Hypnum cupressiforme Hedw. collected from bark of Quercus pubescens Wild. in an open karst forest, Pseudacaudella rubida (Börner, 1939) was yielded from both of the above mosses, while a Myzodium sp. was yielded from a colony of Hypnum cupressiforme mixed with a single stem of Brachythecium salebrosum (Hoffm. ex F.Weber & D.Mohr) Schimp., collected from bark of dead trunk of Quercus cerris L. in a mixed deciduous forest. All moss species are new host records for the corresponding species. Myzodium sp. explain some distinctive characters different from the only European species of its genus and probably belong to an undescribed species, but we are avoiding of describing it on the basis of the single adult captured. Distinctive morphological and biological characteristics of the Hungarian specimens of the four aphid species are discussed and illustrated. A list of the accompanying taxa yielded from the Berlese samples are listed in a table.

Phylogenomic insights and geographic distribution of the New World genus Amphibolips Reinhard (Hymenoptera: Cynipidae, Cynipini) using ultraconserved elements

AbstractIn recent years, new wasp species and genera of Cynipidae have been described, and their species delimitation and evolutionary relationships have been supported using molecular markers. However, few studies have included comprehensive and extensive sampling of specimens across the complete distribution of a single genus. In this study, we analysed the phylogenetic relationships of the genus Amphibolips throughout its range using ultraconserved elements (UCEs). We collected 520 adult wasps from 401 sites in various geographic locations, predominantly in Mexico, and used 78 wasps for UCE analysis. Our results demonstrate that Amphibolips forms a monophyletic clade. Amphibolips can be further subdivided into four genetically well‐differentiated clades. Each clade had morphological traits that could be collectively characterised. Species delimitation using molecular sequencing highlights the taxonomic complexity involved in separating and assigning species using exclusively morphological criteria. This approach offers an opportunity to refine current morphological taxonomic criteria or propose new criteria to establish more accurate classifications. The rich diversity of host species and geographic regions in Mexico represents a biodiversity hotspot for Amphibolips. The number of undescribed species remains high. This study facilitates the determination of appropriate boundaries for putative new Amphibolips species.

A new species of Hypancistrus Isbrücker & Nijssen 1991 (Loricariidae, Siluriformes) from the rapids of the middle Rio Tocantins.

The Hypancistrus genus is recognized in the Río Orinoco basin and Rio Xingu in the Guiana and Brazilian Shields, respectively. Some of its species are important in ornamental fishing. Despite this significance, many other undescribed species are still awaiting to be named. Here we describe a new species of Hypancistrus found on bedrock in the Rio Tocantins, representing an extension of the distribution of the genus. Also, a multigene phylogeny is presented to evaluate the taxonomic position of this species concerning congeners. The new species differs from all congeners by (1) hypertrophied odontodes on cheeks reaching beyond the cleithrum, (2) a supraoccipital crest conspicuously elevated, (3) a supraorbital crest slightly convex, (4) oblique bars on the anterior part of the body, (5) a dark E-shaped mark on the snout, (6) three oblique dark bars on the anterior part of the body and horizontal vermicular bars from the pectoral girdle to the posterior insertion of the dorsal fin, (7) a thin light gray bar on the posterior of the head extending across the branchial opening, (8) a tan background color, (9) a developed suspensorium with a diminished appendix in the metapterygoid, and (10) a dentary plate robust significantly fused with the angulo-articular bone. The molecular phylogenetic results show the new species forming a group with Hypancistrus zebra (Brazilian Shield-Rio Xingu) as a clade, a sister group of a monophyletic group consisting of all congeners from the Río Orinoco.

Two new species of Thomasomys (Cricetidae: Sigmodontinae) from the western Andes of Ecuador and an updated phylogenetic hypothesis for the genus

Abstract The Andean cloud forests of Ecuador are home to numerous unique mammals. Rodents of the tribe Thomasomyini are particularly abundant in many Andean localities, with Thomasomys – the largest genus in the subfamily Sigmodontinae (51 species) – especially species-rich and diverse. Despite recent advances on the systematics of the genus, where seven species have been described in the last five years, there is tantalizing evidence that its true diversity remains completely understood. Over the course of approximately ten years of fieldwork in Ecuador, a significant number of Thomasomys specimens were collected from various localities in both, the eastern and western Andean ranges. Through an extensive genetic study of these specimens, augmented with what is available in public databases, we argue that there exist at least 20 undescribed species in the genus, with no less that twelve potential new species in Ecuador alone. In this paper, we describe two of these species belonging to the group cinereus, one recently collected and the other previously referred to as Thomasomys sp. 1; further, we present an updated cyt b gene tree of the genus. The gene tree includes at least 56 valid and putative species and supports the monophyly of the genus, while at the same time suggest a paraphyletic “aureus” group. Our findings suggest that the genus likely exhibits additional hidden diversity in significant portions of Colombia, Peru, and Bolivia which calls for the need for a comprehensive reassessment of the entire genus. The recognition of these two new species brings the total number of known Thomasomys to 53 species, 19 of which occur in Ecuador, including 17 that are endemic to this country.

Quercus purhepecha (Fagaceae), a new species of shrub oak endemic to the state of Michoacán, Mexico

Within the genus Quercus there are species with a shrubby habit that have been little studied worldwide. In Mexico, several shrub species within section Quercus are taxonomically highly problematic. These include eight named taxa that share traits such as abaxially tomentose leaves and growth of stems from one or multiple rhizomes, and there is evidence of additional undescribed species. Here, Quercus purhepecha is described and illustrated as a new tomentose shrubby white oak species, with a very restricted distribution within the Cuitzeo basin in the Trans-Mexican Volcanic Belt (TMVB), Michoacán state, Mexico. This new species is morphologically compared with Q. frutex, Q. microphylla and Q. repanda, which are similar taxa also distributed in the TMVB. In addition, a potential distribution model of Q. purhepecha for the Cuitzeo basin is presented.

Phylogeny, Diversification, and Biogeography of Garra (Cypriniformes: Cyprinidae) Reveals Multiple Cross‐Drainage Dispersals in Southeast Asia

ABSTRACTGarra, a kind of small‐ to medium‐sized fish, is widely distributed from southern Eurasia to central Africa. As one of the most diverse genera of Cyprinidae, investigating the phylogeny and biogeography of Garra remains challenging. In this study, we combined sequences of Garra samples collected from Myanmar with sequences downloaded from GenBank to investigate the phylogeny, diversification, and biogeography of Garra on a global scale, with an emphasis on Southeast Asia. Species delimitation results indicated that there are at least 22 potential species, including eight undescribed species in Myanmar, suggesting that the diversity of Garra in this region have been largely underestimated. Diversification analysis suggested a relatively high diversification rate in the early branches of Garra. Ancestral distribution reconstruction results revealed that Garra originated from the Irrawaddy River basin in the late Eocene, approximately 34 million years ago, with subsequent dispersals across different drainages influenced by the uplift of the Qinghai‐Tibet Plateau. Our study provided a new insight into the evolutionary history of Garra and the basis for further research on this genus.

Peptide Toxin Diversity and a Novel Antimicrobial Peptide from the Spider Oxyopes forcipiformis

Spider venoms are emerging as a rich source of bioactive peptide toxins with therapeutic potential. Lynx spiders of the genus Oxyopes are small, cursorial hunters that employ complex venom to subdue arthropod prey. However, extracting crude venom from these diminutive arachnids poses significant challenges. This study presents a transcriptome analysis of venom glands from an undescribed Oxyopes forcipiformis species, revealing 339 putative protein and peptide toxin sequences categorized into seven functional groups. The venom composition was dominated by membrane-active peptides (40.71%), venom auxiliary proteins (22.71%), neurotoxins (15.63%), channel active peptides (7.08%) and uncharacterized components (13.87%). Additionally, phylogenetic analysis of 65 disulfide-bond-rich peptides yielded six distinct families based on sequence homology and cysteine framework. Finally, a novel antimicrobial peptide, GK37, was identified using in silico and homology analyses. Our data suggested that GK37 presented significant antibacterial activity against Gram-positive bacteria Staphylococcus aureus with a minimum inhibitory concentration (MIC) of 1.552 µM by disrupting bacterial membranes. At 4× MICs, GK37 almost showed no hemolytic activity on blood cells or toxicity against Hek293T cells. Our findings provided a basis for targeted studies of the diversity and pharmacological effects of lynx spider peptide. We elucidated a valuable high-throughput approach for obtaining proteins and peptides from small-group spiders.

New species of braconid parasitoid wasps from the subfamilies Doryctinae and Brachistinae from late Eocene Baltic amber

Three new fossil species of braconid wasps are described and illustrated from late Eocene Baltic amber: Palaeorhoptrocentrus tenuicornis sp. nov. (Doryctinae), Taphaeus obscurus sp. nov. (Brachistinae: Diospilini), and Eubazus electrus sp. nov. (Brachistinae: Brachistini). Keys to the known species of Palaeorhoptrocentrus Belokobylskij, 2023 and to species of Eubazus Nees, 1812 described from Baltic amber are provided. An illustrated redescription of Blacus (Electroblacus) facialis Brues, 1933 (Brachistinae: Blacini) is also provided, along with digital photographs for the first time of the female of B. (Electroblacus) facialis, whose type material was lost during the World War II. We also provide morphological notes and digital photographs of a male belonging to an undescribed species of Meteorus (Euphorinae).

Killer Whale Predation on a Giant Manta Ray (Mobula birostris), a Sicklefin Devil Ray (Mobula tarapacana) and a Tiger Shark (Galeocerdo cuvier) in the Southwest Indian Ocean

ABSTRACTKnowledge of killer whale (Orcinus orca) feeding ecology in tropical waters is scarce. In the southwest Indian Ocean, opportunistic sightings provide a valuable source of information to better understand their behaviour, diet and distribution. Here, we compile existing records of killer whale predation on elasmobranchs in the southwest Indian Ocean, including sightings of three undescribed prey species regionally: a giant manta ray (Mobula birostris), a sicklefin devil ray (Mobula tarapacana) and a tiger shark (Galeocerdo cuvier). Documenting such observations is important to increase knowledge on killer whale ecology in the tropics and the cascading impact they might have on prey populations.

Characterization and phylogenetic analysis of the chloroplast genome of Diospyros aff. oleifera

The Diospyros genus (Ebenaceae) has significant economic value. During field surveys, we discovered a Diospyros specimen showing morphological overlap with both D. oleifera and D. kaki var. silvestris, provisionally named Diospyros aff. oleifera. To resolve its taxonomy, we sequenced and analyzed its chloroplast genome. The complete chloroplast genome is 157,732 bp with a quadripartite structure. mVISTA analysis revealed unique sequence variations compared to related species. Phylogenetic analysis using 75 protein-coding genes grouped it with D. oleifera, indicating their close relationship. Our findings suggest this specimen likely represents a novel, undescribed species. This study provides insights into Diospyros diversity and a foundation for future research.

Integrative taxonomy reveals two new species of Maritrema Nicoll, 1907 (Digenea: Microphallidae Ward, 1901), parasitizing birds in the Gulf of Mexico.

Members of the genus Maritrema Nicoll, 1907 include endoparasites of aquatic birds and mammals, distributed worldwide. Adult specimens were collected from the intestines of three bird species (the great black hawks, Buteogallus urubitinga Gmelin; laughing gull, Leucophaeus atricilla Linnaeus; and the willet, Tringa semipalmata Gmelin) in three localities along the Gulf of Mexico. Photogenophores were sequenced for the large subunit (LSU) from nuclear rDNA, and the new sequences were aligned with other microphalloid sequences available in GenBank. The maximum likelihood and Bayesian inference analyses revealed three independent lineages, one corresponding to a previously described species, Maritrema corai Hernández-Orts, Pinacho-Pinacho, García-Varela & Kostadinova, 2016, and two representing two undescribed species. These two new species are described in the current study. Maritrema itzamnai n. sp. can be morphologically differentiated from its four congeneric species occurring in coastal areas of Mexico by having smaller oral (20-29 × 20-38 μm) and ventral (20-39 × 19-33 μm) suckers. In addition, Maritrema itzamnai n. sp. possess annular vitellarium instead of a horseshoe-shaped vitellarium. Maritrema kukulkanni n. sp. can be distinguished from its congeneric species reported from Mexico by possessing a larger oesophagus (44-117μm) and by having a vitellarium distributed in both the hindbody and forebody. Maritrema corai is the third species recorded in this study parasitizing the great black hawk (Buteogallus urubitinga); this represents a new host and locality record, expanding its distribution range from the Mexican Pacific Ocean to the Gulf of Mexico.

Vishniacozyma floricola sp. nov., a flower-related tremellomycetous yeast species from Europe.

During the course of two independent studies conducted in Hungary and Spain, four conspecific yeast strains were isolated from flowers of different plant species. DNA sequences of two barcoding regions, the D1/D2 domain of the LSU rRNA gene and the internal transcribed spacer (ITS) region (ITS1-5.8S rRNA gene-ITS2), revealed that the four strains represent an undescribed Vishniacozyma (family Bulleribasidiaceae, Basidiomycota) species. In terms of pairwise sequence similarities and according to our phylogenetic analyses of the concatenated DNA sequences of the ITS region and the D1/D2 domain of the LSU rRNA gene, the undescribed species is most closely related to Vishniacozyma melezitolytica, a yeast species of phylloplane origin. The novel species differs from the type strain of V. melezitolytica by 8 substitutions and 3 insertion/deletion (indels) and 11 substitutions and 5 indels along the D1/D2 domain of the LSU rRNA gene and the ITS region, respectively. In addition to the DNA sequence divergences, the two species differ in some physiological characters as well. We propose the species Vishniacozyma floricola sp. nov. to accommodate the above-noted strains (holotype, NCAIM Y.02320; isotype, CBS 18939; MycoBank number, 856028).

Wandering the taxonomic mine-field: the Podocotyle species complex (Digenea: Opecoelidae).

This study focuses on the opecoelid genus Podocotyle, particularly the species in the northern European seas (White, Barents and Pechora), with additional data from few isolates from the Far East. The research combines molecular analyses (28S, 5.8S+ITS2 rDNA, and cox1 mtDNA) and morphological examinations. Molecular analyses suggested that there are most likely four species of Podocotyle in the northern European seas and three more in the Far East. Morphological redescriptions are provided for three northern European species: P. atomon, P. reflexa, and P. odhneri, with the latter recognized as a valid species. The study also verified the life cycle of P. atomon and identified Lacuna vincta (Littorinidae) as the first intermediate host for P. reflexa. In the Sea of Okhotsk, L. turrita is the first intermediate host for two apparently undescribed Podocotyle species. The results also highlight the morphological variability of P. atomon, emphasizing the need for an integrative approach in the taxonomy and biodiversity studies.

Hymenochaete ametzii sp. nov. (Hymenochaetales, Basidiomycota), an endangered bark-dwelling species inhabiting old Quercus pyrenaica trees from the Iberian Peninsula

Bark-dwelling fungi represent a group of ecologically highly specialized organisms. This study deals with an undescribed species of Hymenochaete characterized by specifically inhabiting the bark of Pyrenean oak (Quercus pyrenaica), and producing effuse-reflexed basidiomata and mainly globose to subglobose basidiospores. Maximum Likelihood and Bayesian analyses of the nuclear ITS-LSU regions revealed that the sequences of H. ametzii form a monophyletic group with a low intraspecific variation and substantially different from closest taxa, further supporting its recognition as a species. The seven localities H. ametzii is known from are old forests with a long ecological continuity, containing large old Q. pyrenaica trees, and are located in the supramediterranean belt of the Mediterranean biogeographical region. Based on habitat availability and field counts of colonized trees, the global population of H. ametzii is estimated at 8,670 mature individuals. Considering that the traditional use of old Q. pyrenaica stands ceased several decades ago in the Iberian Peninsula, with a consequent decline in habitat quality and availability, it is concluded that H. ametzii is Endangered (EN) according to the IUCN criteria.

To be, or not tu-be? Population structure and connectivity in three Arctic kinorhynch species: Echinoderes aquilonius, Echinoderes eximus and Echinoderes remanei

The Arctic, despite being profoundly affected by global climate change, remains largely understudied. Its local ecosystems and fauna are undergoing significant changes, including marine meiobenthic invertebrates, such as kinorhynchs. However, although they are an important component of the benthic meiofauna, our understanding of their communities remains relatively limited. The present study focuses on the structure and connectivity of the populations of three Echinoderes species: Echinoderes aquilonius, Echinoderes eximus and Echinoderes remanei, inhabiting the coastal area of Disko Island (Western Greenland). A hypothesis of low population diversity was tested through analyses based on both morphology and the molecular cox1 marker. For the first time the distribution and relationships between and within species were described. Phylogenetic analysis and haplotype networks reflected the results of population statistics (nucleotide and haplotype diversity, Tajima's D and Fu's Fs), revealing expected homogeneity and low diversity, without any noticeable patterns of geographical distribution. The results indicate that Arctic kinorhynchs form progressively developing and expanding communities. Additionally, the analyses revealed a taxonomically important example of intraspecific polymorphism. Presence or absence of tubes are usually considered to be species diagnostic characters, and the finding an isolated population of E. eximus-like specimens with sublateral tubes on segment 9, suggested that this could be a new, yet undescribed species. However, the genetic divergence between these specimens and other populations of E. eximus without tubes on segment 9 was so low that they all have to be considered conspecific. This is the first time molecular sequence data has provided evidence for intraspecific tube polymorphism among kinorhynchs.