An updated world checklist of velvet worms (Onychophora) with notes on nomenclature and status of names.

More than a decade has passed since the publication of the only world checklist available for Onychophora. During this period, numerous nomenclatural acts and taxonomic changes have been suggested within the group and a wealth of novel data has been published on many taxa. Herein, the up-to-date taxonomic scenario within Onychophora is presented, with appraisal of name status. This checklist covers both extant (Peripatidae and Peripatopsidae) and fossil taxa, and each species is accompanied by information on synonyms, type designation, holotype location, type locality, and language of original description. Additional remarks include nomenclatural inconsistencies, synonymizations, name misspellings, conflicting collecting event data, availability of taxonomically informative molecular data, etc. According to the data, 237 species are currently assigned to Onychophora: 140 of Peripatopsidae, 92 of Peripatidae, and five fossil species with unclear relationship to extant taxa. Since the previous checklist, 37 species have been added to Onychophora, representing an increase of 18.5% in the diversity described for the group. Yet, taxonomic descriptions seem slow-paced, with an average of 3.6 onychophoran species being described annually. From the taxonomic standpoint, 216 species are valid, although many of them require morphological revision and molecular characterization; 21 species exhibit major taxonomic ambiguities and have been regarded as nomina dubia. Recurrent taxonomic issues identified in the literature include inaccurate collecting event data, doubtful taxonomic assignment of molecular sequences, and non-observance of nomenclatural rules. These and other taxonomic aspects are addressed herein in the light of the directives established by the International Code of Zoological Nomenclature.

Sizing Up the Onychophoran Genome: Repeats, Introns, and Gene Family Expansion Contribute to Genome Gigantism in Epiperipatus broadwayi.

Genome assemblies are growing at an exponential rate and have proved indispensable for studying evolution but the effort has been biased toward vertebrates and arthropods with a particular focus on insects. Onychophora or velvet worms are an ancient group of cryptic, soil dwelling worms noted for their unique mode of prey capture, biogeographic patterns, and diversity of reproductive strategies. They constitute a poorly understood phylum of exclusively terrestrial animals that is sister group to arthropods. Due to this phylogenetic position, they are crucial in understanding the origin of the largest phylum of animals. Despite their significance, there is a paucity of genomic resources for the phylum with only one highly fragmented and incomplete genome publicly available. Initial attempts at sequencing an onychophoran genome proved difficult due to its large genome size and high repeat content. However, leveraging recent advances in long-read sequencing technology, we present here the first annotated draft genome for the phylum. With a total size of 5.6Gb, the gigantism of the Epiperipatus broadwayi genome arises from having high repeat content, intron size inflation, and extensive gene family expansion. Additionally, we report a previously unknown diversity of onychophoran hemocyanins that suggests the diversification of copper-mediated oxygen carriers occurred independently in Onychophora after its split from Arthropoda, parallel to the independent diversification of hemocyanins in each of the main arthropod lineages.

Molecular analyses reveal the occurrence of three new sympatric lineages of velvet worms (Onychophora: Peripatidae) in the eastern Amazon basin

Abstract In the present study, we investigated the possible existence of new lineages of peripatids through comparisons between known Neotropical species and specimens obtained from two locations in Pará, a state in eastern Brazilian Amazonia using a molecular approach based on sequences of the mtDNA genes COI, 16Sr RNA, and 18S RNA. The analyses included also sequences of Asian and African taxa for a more systematic understanding of the phylogenetic relationships within the group. The analysis of the COI, 16S rRNA and 18S RNA sequences permitted the identification of three distinct lineages (A, B and C) based on two different phylogenetic approaches (Bayesian methods and ML). The three lineages presented here are completely distinct from all other peripatid taxa so far defined by molecular data. The presence of specimens of three independent onychophoran lineages occurring in sympatry in the Amazon basin was confirmed in all the analyses, providing consistent support for the phylogenies presented in this study. These findings reinforce the importance of the Amazon region in the diversification of Neotropical peripatids, and indicate that onychophoran diversity is much greater than previously thought, given that the number of taxa found at a single site was equivalent to the total number of allopatric species described for the entire region.

Earliest Onychophoran in Amber Reveals Gondwanan Migration Patterns.

The anomalous occurrence of supposedly Gondwanan taxa in Laurasian-derived regions remains an intriguing chapter of paleobiogeographical history. Representatives of Peripatidae, a major subgroup of velvet worms (Onychophora), show a disjointed distribution in the neotropics, tropical Africa, and Southeast Asia, the latter being the only landmass previously associated with Laurasia [1, 2]. The arrival of these animals in Southeast Asia is explained by two alternative, albeit not mutually exclusive, hypotheses: an early migration via Europe before continental drift (Eurogondwana hypothesis) or transportation via insular India during the Cretaceous and Paleogene ("out-of-India" hypothesis) [3-6]. The latter hypothesis is based on a single extant species of Peripatidae, Typhloperipatus williamsoni, in India. †Cretoperipatus burmiticus from Myanmar is the oldest fossil onychophoran found in amber [7], dating to sometime between the two proposed scenarios, and hence crucial for clarifying how Gondwanan lineages of these low-vagility animals reached Southeast Asia (see also Supplemental Information). Based on the anatomical reconstruction of †C. burmiticus using synchrotron radiation-based X-ray microtomography (SRµCT) and comparisons with extant taxa, we resolved this fossil species within Onychophora, particularly within Peripatidae, with T. williamsoni as its closest extant relative. This suggests that an early Eurogondwanan migration of peripatids was the most likely event, as Burmese amber is too old to be compatible with the out-of-India hypothesis. Moreover, peripatids probably colonized India only recently from Myanmar, refuting the putative Gondwanan relict status of Indian onychophorans. Finally, preservation artifacts identified in the novel amber material might have a major impact on studies of onychophoran stem and/or crown groups.

Macroperipatus ohausi: redescription and taxonomic notes on its status (Onychophora: Peripatidae) / Macroperipatus ohausi: redescripción y notas taxonómicas sobre su estatus (Onychophora: Peripatidae)

Onychophora have undergone a new wave of taxonomic revisions in the last three years. After a recent revision, only two species of Macroperipatus are registered from Brazil: Macroperipatus ohausi and Macroperipatus geayi. The former was only the second species of Onychophora to be described from Brazil, and is herein redescribed, illustrated, and its taxonomic status revised. Macroperipatus ohausi had only been collected at the type locality, Petrópolis, state of Rio de Janeiro, but has not been found there since its discovery. In 2009 we collected two specimens of this species in an Atlantic rain forest fragment in Nova Iguaçu in RPPN dos Petroleiros (also state of Rio de Jane iro). The color of the new specimens was interpreted and described based on the NBS/ISCC System of Color Designation, and corresponds only weakly to that described in earlier publications on M. ohausi. The identification of M. ohausi was based solely on its morphology and compared with its original description. In addition, a female syntype of M. ohausi deposited in the Zoologisches Museum (Hamburg) was examined. We reinvestigated its external morphology and concluded that the dorsal papillae have been misinterpreted in M. ohausi in previous studies of its integument. The new specimens examined are conspecific with M. ohausi mainly by having conical primary papillae, different sized accessory papillae, and by the number of legs. Our material resembles the original description of M. ohausi, for example, in spinous pad and nephridial tubercles, even though these features are not represented in the original description and rede- scription of M. ohausi; these features are further detailed here. We concluded that the dorsal papillae morphology of M. ohausi, shares diagnostic characters, for example, with representatives of the genera Peripatus and Epiperipatus. The dorsal papillae are rounded and not square as in the original description and redescription. Another argument that supports our conclusion is that the RPPN dos Petroleiros, the locality where the new samples were collected, belongs to the same mountain chain and Atlantic mosaic as the original material of M. ohausi. Furthermore, a male of M. ohausi is described here for the first time. Based on our morphological data, we transfered this species to the genus Epiperipatus, proposing a new combination, E. ohausi.

Onychophora experimentó una nueva serie de revisiones taxonómicas en los últimos tres años. Después de una reciente revisión, sólo dos especies de Macroperipatus fueron registradas para Brasil: Macroperipatus ohausi y Macroperipatus geayi. La primera fue solo la segunda especie de Onychophora en ser descrita en Brasil y aquí es redescrita, ilustrada, y su estatus taxonómico revisado. Macroperipatus ohausi sólo se había recolectado en la localidad tipo, Petropolis, Estado de Rio de Janeiro, pero no se ha encontrado allí desde su descubrimiento. En 2009, recolectamos dos ejemplares de esta especie en un fragmento de la ‘Mata Atlantica’ en Nova Iguaçu en el RPPN de los Petroleiros (también en el estado de Rio de Janeiro). El color de los ejemplares nuevos fue interpretado y descrito, basado en el sistema NBS System/ISCC de designación de colores, y correspondió débilmente al descrito en publicaciones anteriores sobre M. ohausi. La identificación de M. ohausi se basó únicamente en su morfología, en comparación con la descripción original. Además, se examinó un sintipo hembra de M. ohausi depositado en Zoologisches Museum (Hamburgo). Nosotros reinvestigamos su morfología externa y concluimos que las papilas dorsales fueron mal interpretadas en M. ohausi en los estudios previos de su integumento. Los nuevos especímenes examinados son coespecíficos con M. ohausi principalmente por tener las papilas primarias cónicas, diferente tamaño de las papilas accesorias, y por el número de patas. Nuestro material se asemeja a la descripción original de M. ohausi, por ejemplo, la almohadilla espinosa y tubérculos nefridiales, aunque estas características no están representadas en la descripción original y redescripcion de M. ohausi, estas características también se detallan aquí. Llegamos a la conclusión de que la morfología de las papilas dorsales de M. ohausi, comparte caracteres de diagnóstico, por ejemplo, con los representantes de los géneros Peripatus y Epiperipatus. Las papilas dorsales son redondeadas y no cuadradas como en la descripción original y redescripción. Otro argumento que apoya nuestra conclusión es que la RPPN dos Petroleiros, la ubicación en la que se recogieron las nuevas muestras, pertenece a la misma cadena de montañas y al mosaico Atlántico, tal como el material original de M. ohausi. Por otra parte, se describe aquí por primera vez un macho de M. ohausi. Basado en nuestros datos morfológicos, transferimos esta especie al género Epiperipatus, que propone una nueva combinación, E. ohausi.

Apodemes associated with limbs support serial homology of claws and jaws in Onychophora (velvet worms).

Although the onychophoran jaw blades are believed to be derivatives of foot claws, serial homology of these structures has not been demonstrated. To shed light on the evolutionary origin of the onychophoran jaws, we searched for morphological landmarks and compared the internal and external anatomy of jaws and distal leg portions in representatives of the two major onychophoran subgroups, the Peripatidae and Peripatopsidae. Our data revealed hitherto unknown structures associated with the onychophoran limbs, such as a soft diastemal membrane separating the anterior and posterior portions of the inner jaw blade (present only in Peripatidae), apodemes associated with feet, an eversible dorsal sac at the basis of each foot claw, and a specific arrangement of musculature associated with the sclerotised claws, jaws and their apodemes. Specific correspondences in structure and position of apodemes support serial homology of claws and jaws, suggesting that the onychophoran jaw evolved from the distal portion rather than the entire limb in the last common ancestor of Onychophora.

A world checklist of Onychophora (velvet worms), with notes on nomenclature and status of names.

Currently, the number of valid species of Onychophora is uncertain. To facilitate taxonomic work on this understudied animal group, we present an updated checklist for the two extant onychophoran subgroups, Peripatidae and Peripatopsidae, along with an assessment of the status of each species. According to our study, 82 species of Peripatidae and 115 species of Peripatopsidae have been described thus far. However, among these 197 species, 20 are nomina dubia due to major taxonomic inconsistencies. Apart from nomina dubia, many of the valid species also require revision, in particular representatives of Paraperipatus within the Peripatopsidae, and nearly all species of Peripatidae. In addition to extant representatives, the record of unambiguous fossils includes three species with uncertain relationship to the extant taxa. For all species, we provide a list of synonyms, information on types and type localities, as well as remarks on taxonomic and nomenclatural problems and misspellings. According to recent evidence of high endemism and cryptic speciation among the Peripatidae and Peripatopsidae, previous synonyms are revised. Putative mutations, subspecies and variations are either raised to the species status or synonymised with corresponding taxa. In our revised checklist, we follow the rules and recommendations of the International Code of Zoological Nomenclature to clarify previous inconsistencies.

A new giant species of placented worm and the mechanism by which onychophorans weave their nets (Onychophora: Peripatidae)

Onychophorans, or velvet worms, are poorly known and rare animals. Here we report the discovery of a new species that is also the largest onychophoran found so far, a 22cm long female from the Caribbean coastal forest of Costa Rica. Specimens were examined with Scanning Electron Microscopy; Peripatus solorzanoi sp. nov., is diagnosed as follows: primary papillae convex and conical with rounded bases, with more than 18 scale ranks. Apical section large, spherical, with a basal diameter of at least 20 ranks. Apical piece with 6-7 scale ranks. Outer blade 1 principal tooth, 1 accessory tooth, 1 vestigial accessory tooth (formula: 1/1/1); inner blade 1 principal tooth, 1 accessory tooth, 1 rudimentary accessory tooth, 9 to 10 denticles (formula: 1/1/1/9-10). Accessory tooth blunt in both blades. Four pads in the fourth and fifth oncopods; 4th. pad arched. The previously unknown mechanism by which onychophorans weave their adhesive is simple: muscular action produces a swinging movement of the adhesive-spelling organs; as a result, the streams cross in mid air, weaving the net. Like all onychophorans, P. solorzanoi is a rare species: active protection of the habitat of the largest onychophoran ever described, is considered urgent. Rev. Biol. Trop. 58 (4): 1127-1142. Epub 2010 December 01.

Los onicóforos o "peripatos" son animales escasos y poco conocidos. Aquí informamos el descubrimiento del onicóforo más grandes conocido, una hembra de 22cm de longitud del bosque costero caribeño de Costa Rica. Analizamos los especímenes con microscopia electrónica de barrido. La nueva especie, Peripatus solorzanoi, sp. nov. se caracteriza así: papilas primarias convexas y cónicas, con bases redondeadas y más de 18 filas de escamas de alto. Sección apical grande y esférica, con diámetro basal de al menos 20 filas. Parte apical con 6-7 filas de escamas de alto. Fórmula de lámina mandibular externa 1/1/1, lámina interna: 1/1/1/9-10. Diente accesorio romo en ambas láminas. Cuatro almohadillas en cuarto y quinto oncopodios; cuarta almohadilla arqueada. Describimos el mecanismo, previamente desconocido, mediante el cual tejen su red los onicóforos, usando fotografías para mostrar que es de naturaleza muscular. Como todos los demás onicóforos, P. solorzanoi es una especie escasa: consideramos urgente la protección del hábitat del onicóforo más grande del mundo.

Seminal receptacula in gravid and virgin female Peripatus (Macroperipatus) acacioi Marcus and Marcus (Onychophora, Peripatidae).

The ovoid seminal receptacula in Peripatus acacioi are located at the junctions of the short paired oviducts with the two horns of the uterus. Associated with each is a tubular funnel that opens into the haemocoel. In P. acacioi, spermatozoa may be stored in the seminal receptacula for several years (Campiglia and Walker '95, J. Morphol. 224:179-198). Observations of the structure of the seminal receptaculum using transmission electron microscopy (TEM) show that there are numerous tracheae within its wall indicating a good oxygen supply. The ultrastructure of the epithelium lining the seminal receptaculum indicates that these cells secrete the material that forms the luminal matrix that surrounds and provides nutrition for the stored spermatozoa. The ducts that interconnect the ovary, seminal receptaculum, funnel, and uterus are all packed with cilia. The structure of the seminal receptaculum and associated parts in the mature virgin female is identical to that of the gravid female, but the luminal matrix does not contain any spermatozoa. J. Morphol. 237:127-136, 1998. © 1998 Wiley-Liss, Inc.