@article {2028, title = {An updated world checklist of velvet worms (Onychophora) with notes on nomenclature and status of names}, journal = {ZooKeys}, volume = {1184}, year = {2023}, month = {16/11/2023}, pages = {133-260}, type = {Checklist}, abstract = {

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.

Remark: Closing date of checklist data is 18 August 2023.

}, keywords = {catalogue, extant, fossil, Peripatidae, Peripatopsidae, species list, Taxonomy}, issn = {1313-2989, 1313-2970}, doi = {https://doi.org/10.3897/zookeys.1184.107286}, url = {https://zookeys.pensoft.net/article/107286/}, author = {Oliveira, Ivo de Sena} } @article {1910, title = {Sizing Up the Onychophoran Genome: Repeats, Introns, and Gene Family Expansion Contribute to Genome Gigantism in Epiperipatus broadwayi}, journal = {Genome Biology and Evolution}, volume = {15}, year = {2023}, month = {15/02/2023}, pages = {1-6}, abstract = {

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.

}, keywords = {hemocyanin, Panarthropoda, Peripatidae, velvet worm}, isbn = {1759-6653}, doi = {https://doi.org/10.1093/gbe/evad021}, url = {https://academic.oup.com/gbe/article/15/3/evad021/7039704}, author = {Sato, Shoyo and Cunha, Tauana J and de Medeiros, Bruno A S and Khost, Danielle E and Sackton, Timothy B and Giribet, Gonzalo} } @article {1508, title = {List of terrestrial invertebrates of St. John, U.S. Virgin Islands (exclusive of Acarina and Insecta), with some records of freshwater species}, journal = {Caribbean Journal of Science}, volume = {29}, year = {1993}, pages = {30-38}, abstract = {

Abstract: A list is given of the terrestrial invertebrates known to occur on St. John, with the exception of the Acarina and Insects. Included are: Platyhelminthes (land planarian) \– 1; Mollusca (snails and slugs) \– 32; Annelida (earthworms) \– 1; Onychophora (Peripatus) \– 1; Crustacea (woodlice, beachhoppers, crabs) \– 28; Arachnida (scorpions, pseudoscorpions, harvestmen, spiders, etc.) \– 120; Myriapoda (centipedes, millipedes, etc.) \– 19. In addition, a few freshwater species are noted. A bibliography of all works referring to these animals is provided.


Resumen: Se provee una lista de los invertebrados terrestres conocidos de St. John, con la excepci\ón de Acarina e Insecta. Se incluyen: Platyhelminthes (planaria terrestre) \– 1; Mollusca (caracoles y babosas) \– 32; Annelida (lombrices de tierra) \– 1; Onychophora (Peripatus) \– 1; Crustacea (carcomas, saltarines, cangrejos) - 28; Arachnida (escorpiones, pseudoescorpiones, opiliones, ara\ñas, etc.) \– 120; Myriapoda (ciempi\és, milpi\és) \– 19. Tambi\én son informadas varias especies dulceacu\ícolas. Se provee una bibliograf\ía que incluye todos los trabajos referentes a estos animales.

}, author = {Muchmore, William B.} } @article {1457, title = {Phylogenomic Analysis of Velvet Worms (Onychophora) Uncovers an Evolutionary Radiation in the Neotropics}, journal = {Molecular Biology and Evolution}, volume = {38}, year = {2021}, month = {24/08/2021}, pages = {5391 - 5404}, abstract = {

Onychophora (\“velvet worms\”) are charismatic soil invertebrates known for their status as a \“living fossil,\” their phylogenetic affiliation to arthropods, and their distinctive biogeographic patterns. However, several aspects of their internal phylogenetic relationships remain unresolved, limiting our understanding of the group\’s evolutionary history, particularly with regard to changes in reproductive mode and dispersal ability. To address these gaps, we used RNA sequencing and phylogenomic analysis of transcriptomes to reconstruct the evolutionary relationships and infer divergence times within the phylum. We recovered a fully resolved and well-supported phylogeny for the circum-Antarctic family Peripatopsidae, which retains signals of Gondwanan vicariance and showcases the evolutionary lability of reproductive mode in the family. Within the Neotropical clade of Peripatidae, though, we found that amino acid-translated sequence data masked nearly all phylogenetic signal, resulting in highly unstable and poorly supported relationships. Analyses using nucleotide sequence data were able to resolve many more relationships, though we still saw discordant phylogenetic signal between genes, probably indicative of a rapid, mid-Cretaceous radiation in the group. Finally, we hypothesize that the unique reproductive mode of placentotrophic viviparity found in all Neotropical peripatids may have facilitated the multiple inferred instances of over-water dispersal and establishment on oceanic islands.

}, keywords = {biogeography, dispersal, Gondwana, Onychophora, systematics, vicariance}, issn = {1537-1719}, doi = {https://doi.org/10.1093/molbev/msab251}, url = {https://academic.oup.com/mbe/article/38/12/5391/6357048}, author = {Baker, Caitlin M and Buckman-Young, Rebecca S and Costa, Cristiano Sampaio and Giribet, Gonzalo} } @article {1456, title = {Panamanian velvet worms in the genus Epiperipatus, with notes on their taxonomy and distribution and the description of a new species (Onychophora, Peripatidae)}, journal = {Invertebrate Biology}, volume = {140}, year = {2021}, month = {25/05/2021}, pages = {1-12}, abstract = {

Among the least studied clade of velvet worms (Onychophora) is a clade of Neotropical Peripatidae (=Neopatida). Neopatida has been traditionally divided into an Andicoles group and a Cara\ïbes group, which now correspond to Oroperipatus and the remaining genera, respectively. Panama is located in a unique position bridging the faunas of both clades and of two large continents, both of which host an important diversity of velvet worms, yet taxonomic study of the Panamanian onychophorans has been largely neglected. In this study we critically evaluate the onychophoran diversity of Panama, redescribe a species from the former Canal Zone and describe a new species from the Pacific side, close to Costa Rica, as Epiperipatus bernali sp. nov., with a unique combination of characters visible under the SEM. We further discuss the existence of multiple lineages of Onychophora in Panama, both from the Andicoles and Cara\ïbes groups, and within the latter, with connections to different lineages, both from South and Central America.

}, keywords = {Central America, morphology, Neotropics, Scanning electron microscopy, Taxonomy}, issn = {1077-8306, 1744-7410}, doi = {https://doi.org/10.1111/ivb.12336}, url = {https://onlinelibrary.wiley.com/doi/abs/10.1111/ivb.12336}, author = {Costa, Cristiano Sampaio and Giribet, Gonzalo} } @article {1452, title = {Fossil onychophorans from Dominican and Baltic amber: Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis balticus n.g., n.sp. (Succinipatopsidae n.fam.) with a proposed classification of the subphylum Onychophora}, journal = {Invertebrate Biology}, volume = {119}, year = {2000}, month = {03/2000}, pages = {104-109}, abstract = {

Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis balticus n.gen., n.sp. (Succinipatopsidae n.fam.) (Lobopodia: Onychophora), the first Tertiary fossils of the Lobopodia, are described from Dominican and Baltic amber, respectively. Both families are characterized by the presence of simple legs lacking foot portions with claws and pads. Tertiapatidae is further characterized by soluble body pigments and oral papillae shorter than the legs. Succinipatopsidae is characterized by non-soluble body pigments and oral papillae longer than the legs. Nomenclatural changes include the erection of the class Udeonychophora n. nom. for terrestrial onychophorans with a ventral mouth, the order Ontonychophora n.nom. for extant onychophorans possessing legs with a differentiated \“foot\” portion, and the family Helenodoridae n.nom. for the genus Helenodora from the Carboniferous. The biogeographical significance of these fossils and their phylogenetic relationship with previously described onychophorans are discussed.

}, keywords = {Euonychophora, fossil velvet worms}, issn = {1077-8306, 1744-7410}, doi = {https://doi.org/10.1111/j.1744-7410.2000.tb00178.x}, url = {https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7410.2000.tb00178.x}, author = {Poinar, George O.} } @article {1451, title = {Fossiliferous Cretaceous Amber from Myanmar (Burma): Its Rediscovery, Biotic Diversity, and Paleontological Significance}, journal = {American Museum Novitates}, year = {2002}, month = {26/03/2002}, pages = {1-72}, abstract = {

Amber from Kachin, northern Burma, has been used in China for at least a millennium for carving decorative objects, but the only scientific collection of inclusion fossils, at the Natural History Museum, London (NHML), was made approximately 90 years ago. Age of the material was ambiguous, but probably Cretaceous. Numerous new records and taxa occur in this amber, based on newly excavated material in the American Museum of Natural History (AMNH) containing 3100 organisms. Without having all groups studied, significant new records and taxa thus far include the following (a \† refers to extinct taxa): For Plants: An angiosperm flower (only the third in Cretaceous amber), spores and apparent sporangia of an unusual but common fungus, hepatophyte thalli and an archegoniophore of Marchantiaceae, and leafy shoots of Metasequoia (Coniferae). Metasequoia is possibly the source of the amber. For Animals: Mermithidae and other Nematoda; the oldest ixodid tick (a larval Amblyomma); bird feathers; and the only Mesozoic record of the Onychophora (\&$\#$39;velvet\&$\#$39; worms), described as \†Cretoperipatus burmiticus, n. gen., n. sp. (Peripatidae). Poinar\&$\#$39;s classification of the Onychophora is substantially revised. Still largely unstudied, the fauna of mites (Acari) and spiders (Araneae) appears to be the most diverse ones known for the Mesozoic. For Insecta: Odonata indet. (wing fragment); Plecoptera indet.; new genera of Dermaptera, Embiidina, and Zoraptera (the latter two as the only definitive Mesozoic fossils of their orders). Within Hemiptera, there are primitive new genera in the Aradidae, Hydrometridae, Piesmatidae, Schizopteridae, and Cimicomorpha (Heteroptera), as well as in \†Tajmyraphididae (Aphidoidea), and \†Protopsyllidiidae. An adult snakefly (Raphidioptera: \†Mesoraphidiidae) is the smallest species in the order, and new genera occur in the Neuroptera: Coniopterygidae, Berothidae, and Psychopsidae, as well as larvae of apparent Nevrorthidae. Coleoptera are largely unstudied, but are probably the most diverse assemblage known from the Cretaceous, particularly for Staphylinidae. An adult lymexylid, the most primitive species of Atractocerus, is the first Mesozoic record of the family. In Hymenoptera there are primitive ants (Formicidae: Ponerinae n. gen., and \†Sphecomyrma n.sp (Sphecomyrminae)), the oldest record of the Pompilidae, and significant new records of \†Serphitidae and \†Stigmaphronidae, among others. Diptera are the most diverse and abundant, with the oldest definitive Blephariceridae and mosquito (Culicidae), as well as new genera in the Acroceridae, Bibionidae, Empidoidea; a new genus near the enigmatic genus Valeseguya, and an unusual new genus in the \†Archizelmiridae. \†Chimeromyia (Diptera: Eremoneura), known previously in ambers from the Lower Cretaceous, is also represented.

The stratigraphic distribution of exclusively Mesozoic arthropods in Burmese amber is reviewed, which indicates a probable Turonian-Cenomanian age of this material (90-100 Ma). Paleofaunal differences between the NHML and AMNH collections are discussed, as is the distinct tropical nature of the original biota. Burmese amber probably harbors the most diverse biota in amber from the Cretaceous, and one of the most diverse Mesozoic microbiotas now known.

}, issn = {0003-0082}, url = {http://hdl.handle.net/2246/2914}, author = {Grimaldi, David A. and Engel, Michael S. and Nascimbene, Paul C.} } @article {1394, title = {The highest chromosome number and first chromosome fluorescent in situ hybridization in the velvet worms of the family Peripatidae}, journal = {Zoological Studies}, volume = {59}, year = {2020}, month = {03/03/2020}, pages = {1-8}, abstract = {

The diversity of Onychophora is poorly studied, despite there being nearly 200 described species divided in two families: Peripatidae and Peripatopsidae. Peripatid velvet worms are found mainly in the Neotropical region. The low morphological diversity in Peripatidae is an obstacle to determining its taxonomy, and chromosomal analyses can help clarify this. The aim of this work was to chromosomally analyze one species of Epiperipatus from Mato Grosso do Sul, Brazil. Conventional staining and telomeric fluorescent in situ hybridization (FISH) were performed with the gonads of three males of Epiperipatus sp. The specimens showed 2n♂ = 73, the largest diploid number found in Onychophora to date, with the majority of chromosomes acro/telocentrics and the largest element submetacentric. The FISH marked the telomeric region of all elements and revealed one Interstitial Telomeric Site (ITS) on the proximal region of the long arm large submetacentric chromosome. The absence of male meiosis and female cell division in the analyzed specimens prevented us from determining whether the unpaired large submetacentric is a sex chromosome, which could lead to the description of a rare sex chromosome system (SCS) in Onychophora, or a case of fusion between autosomes. In either case, the presence of ITS is a clear indication of chromosomal fusion.

}, keywords = {Cryptic species, Epiperipatus, Heteromorphic chromosome, Interstitial Telomeric Site, karyotype}, issn = {1810-522X}, doi = {https://doi.org/10.6620/ZS.2020.59-05}, url = {http://zoolstud.sinica.edu.tw/Journals/59/59-05.html}, author = {Duarte Dutra, D{\'e}bora and Bonfim Souza, Lucas Henrique and Medeiros Cordeiro, L{\'\i}via and Araujo, Douglas} } @article {1389, title = {Burmese (Myanmar) amber checklist and bibliography 2018}, journal = {Palaeoentomology}, volume = {2}, year = {2019}, month = {28/02/2019}, pages = {22{\textendash}84}, abstract = {

A list of all known taxa described or recorded from Burmese amber from the published literature up to the end of 2018 is given, along with a comprehensive bibliography. The history of the study of inclusions is summarised, and demonstrates that the number of species has risen exponentially over the past two decades. The first three species were named in 1916 and by the end of 1920 a total of 42 species had been named by T.D.A. Cockerell. Only three more species were named by 1999 though by the end of 2018 the total had risen to an incredible 1,192 species, of which over half were named in the past three years. Some 320 species were named in 2018, the highest number described from one type of amber in any one year in the entire history of amber studies.

}, keywords = {Arachnids, Burmese amber, Cretaceous, fungi, insects, invertebrates, Myanmar, plants, protists, vertebrates}, issn = {2624-2826}, doi = {https://doi.org/10.11646/palaeoentomology.2.1.5}, url = {https://www.mapress.com/pe/article/view/palaeoentomology.2.1.5}, author = {Ross, Andrew J.} } @article {426, title = {Morphological and molecular phylogeny of Epiperipatus (Onychophora: Peripatidae): a combined approach}, journal = {Zoological Journal of the Linnean Society}, year = {2020}, month = {07/11/2020}, pages = {1-31}, abstract = {

Onychophora, or velvet worms, are a key group for understanding ecdysozoan evolution. It comprises two families: Peripatopsidae, largely of Austral distribution, and Peripatidae, which is circumtropical. The interrelationships between the members of Peripatidae present many taxonomic issues exacerbated in the radiation of the Neotropical species or Neopatida. To understand the phylogeny of Neopatida, and to test the information of such morphological characters, we gathered novel molecular and morphological datasets focusing on Neotropical specimens. Our data were analysed using a combination of parsimony and maximum likelihood for the individual and combined molecular and morphological datasets. An analysis of morphology alone was inconclusive, supporting the notion that morphological characters used in peripatid taxonomy have little power to resolve phylogenetic relationships among higher taxa in Neopatida. However, the analyses of molecular or combined data show a split of the Neotropical species into two clades, which we use to reassign genera. Epiperipatus, as currently understood, is non-monophyletic, because it includes species of monotypic genera. To avoid paraphyly of Epiperipatus, the following new combinations are proposed: Epiperipatus bouvieri (Fuhrmann, 1913), Epiperipatus hitoyensis (Oliveira et al., 2012a), Epiperipatus solorzanoi (Morera-Brenes \& Monge-N\ájera, 2010) and Epiperipatus sucuriuensis (Oliveira et al., 2015).

}, keywords = {biogeography, cara{\"\i}bes onychophorans, conserved morphology, Epiperipatus, morphology, Neopatida, New World, Peripatidae, South America, systematics}, issn = {0024-4082}, doi = {10.1093/zoolinnean/zlaa100}, url = {https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zlaa100/5959943}, author = {Costa, Cristiano Sampaio and Giribet, Gonzalo and Pinto-Da-Rocha, Ricardo} } @article {285, title = {Conservation of terrestrial invertebrates: a review of IUCN and regional Red Lists for Myriapoda}, journal = {ZooKeys}, volume = {930}, year = {2020}, month = {28/04/2020}, pages = {221-229}, abstract = {

Red Listing of Threatened species is recognized as the most objective approach for evaluating extinction risk of living organisms which can be applied at global or national scales. Invertebrates account for nearly 97\% of all animals on the planet but are insufficiently represented in the IUCN Red Lists at both scales. To analyze the occurrence of species present in regional Red Lists, accounts of 48 different countries and regions all over the world were consulted and all data about myriapods (Myriapoda) ever assessed in Red Lists at any level assembled. Myriapod species assessments were found in eleven regional Red Lists; however, no overlap between the species included in the global IUCN Red List and the regional ones was established. This means that myriapod species considered threatened at regional level may not be eligible for international funding specific for protection of native threatened species (more than US$ 25 million were available in the last decade) as most financial instruments tend to support only threatened species included in the IUCN Red List. As the lack of financial resources may limit protection for species in risk of extinction, it is urgent to increase the possibilities of getting financial support for implementation of measures for their protection. A Red List of all Myriapoda species recorded in Red Lists at national or local (596) and global (210) scales totaling 806 species is presented. This list shows for the first time an overview of the current conservation status of Myriapoda species. Here, the urgent need of establishing a Myriapoda Specialist Group in the Species Survival Commission of IUCN is also stressed.

}, keywords = {Arthropoda, Chilopoda, Diplopoda, extinction, national red lists, Pauropoda, risk assessment, Symphyla, threatened species}, issn = {1313-2989}, doi = {10.3897/zookeys.930.48943}, url = {https://zookeys.pensoft.net/article/48943/}, author = {Karam-Gemael, Manoela and Decker, Peter and Stoev, Pavel and Marques, Marinez I. and Chagas Jr, Amazonas} } @article {264, title = {{\'E}tudes pour servir {\`a} l{\textquoteright}histoire naturelle des Myriapodes}, journal = {Annales des sciences naturelles, s{\'e}rie 2, Zoologie}, volume = {7}, year = {1837}, month = {01/1837}, pages = {35-60}, abstract = {

Recommended citation: Gervais, P. (1837): \Études pour servir \à l\’histoire naturelle des Myriapodes. Annales des sciences naturelles, s\érie 2, Zoologie, 7: 35-60. https://www.biodiversitylibrary.org/part/28836

}, url = {https://www.biodiversitylibrary.org/part/28836}, author = {Gervais, P.} } @article {229, title = {Notes on the genus Capromys of Desmarest}, journal = {The Zoological Journal}, volume = {4}, year = {1829}, month = {01/1829}, pages = {269-278}, abstract = {
}, keywords = {Cuba}, url = {https://www.biodiversitylibrary.org/page/27485185}, author = {MacLeay, William Sharp} } @article {228, title = {A Monograph on the Species and Distribution of the Genus Peripatus (Guilding)}, journal = {Quarterly Journal of Microscopical Science, new series}, volume = {28}, year = {1888}, month = {04/1888}, pages = {431-493}, url = {https://www.biodiversitylibrary.org/page/13827081}, author = {Sedgwick, Adam} } @article {118, title = {A new giant velvet worm from Costa Rica suggests absence of the genus Peripatus (Onychophora: Peripatidae) in Central America}, journal = {Revista de Biolog{\'\i}a Tropical}, volume = {68}, year = {2020}, month = {03/02/2020}, pages = {300-320}, abstract = {

Introduction: Neotropical onychophoran taxonomy and diversity has been poorly investigated. Recent studies have discovered problems in species classification: they have questioned the accepted genera and the actual number of species. This is true in Costa Rica, where several unidentified species have been reported. Objective: The objective of this investigation was to evaluate the occurrence of the accepted genera in this country, and to describe a new genus and species from Central America. Methods: In 2017, we collected one onychophoran in the Kek\öldi Indigenous Reserve in Talamanca, Lim\ón, Costa Rica. The specimen gave birth to several offspring. Therefore, seven organisms were analyzed. Light microscopy was used to observe the gross morphology in all samples. The detailed morphology was studied in the biggest specimen with scanning electron microscopy; after that, we performed a phylogenetic analysis with the corresponding sequence of COI. Results: According to our results, a new genus and species of giant onychophoran was found. The genus was identified by its giant size, apical piece of seven scale ranks, large conical primary papillae, dorso-median furrow flanked by two-three accessory papillae, the absence of hyaline organs and a marked sexual dimorphism with respect to the number of legs. The new species presents a particular head pattern, as well as novel structures like cephalic papillae, accessory papillae with rudimentary apical pieces, and a lack of antennal chemoreceptors. Phylogenetic analysis rendered our genus as monophyletic and includes Peripatus solorzanoi, which is grouped within the Central American clade. As our species is clustered inside the Costa Rica-Panamanian group, it is not related to the Caribbean Island nor Guyanan Shield samples, home of Epiperipatus and Peripatus respectively. Therefore, we suggest that those genera do not occur in Central America, and a new genus exists: Mongeperipatus, gen. nov. Conclusion: We concluded that Costa Rica is home to a diversity of undescribed onychophorans that requires specific studies to help clarify the taxonomy and evolutionary relationships of the group to justify their protection.

}, keywords = {Costa Rican biodiversity, endemism, onychophoran speciation, Phylogenetics, saproxilic taxa}, issn = {0034-7744}, doi = {https://doi.org/10.15517/rbt.v68i1.37675}, url = {https://www.revistas.ucr.ac.cr/index.php/rbt/article/view/37675}, author = {Barquero-Gonz{\'a}lez, Jos{\'e} Pablo and S{\'a}nchez-Vargas, Steven and Morera-Brenes, Bernal} }