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Two new species of Manahunca, redescription of its type species, current conservation status of the genus and a survey of male glands in Stenostygninae (Opiliones: Laniatores: Biantidae)
Abstract
Manahunca bielawskii Šilhavý, 1973, the type species of the genus Manahunca Šilhavý, 1973, is redescribed based on abundant material from the type locality, including new data on its morphological variability and penis morphology. A neotype for M. bielawskii is herein designated due to the unknown whereabouts of the original holotype. Manahunca silhavyi Avram, 1977, is regarded as a new junior subjective synonym of M. bielawskii. In addition, two new species of Manahunca are described, M. turquino Alegre, Gainett & Giribet, n. sp. and M. matazon Alegre, Gainett & Giribet, n. sp. based on additional specimens from eastern Cuba, for which we provide new data on their geographical distribution, intraspecific variability and habitat. A new diagnosis and emended geographic distribution of the known species of Manahunca are provided, with hints on the current conservation status. The sexually dimorphic glandular structures found on the metatarsus III of males are explored for taxonomic significance in ten species of Stenostygninae. The existence of male dimorphism, most evident in the robustness of the chelicerae, is reported for two of the studied Manahunca species.
... The ongoing systematic review of the Cuban Stenostygninae, undertaken by one of us (AAB), rendered new information for some Cuban genera and for the entire subfamily. This revision has resulted in the redescription of Manahunca bielawskii Šilhavý, 1973 (the type species of the genus Manahunca Šilhavý, 1973), the recognition of Manahunca silhavyi as a junior subjective synonym of M. bielawskii, and the description of two new species from eastern Cuba (Manahunca turquino Alegre, Gainett &Giribet, 2019 andManahunca matazon Alegre, Gainett &Giribet, 2019). In addition, a new diagnosis for Manahunca and its included species were given, along with the update of the geographic distribution and a survey of the sexually dimorphic metatarsus III of several stenostygnines, which revealed the taxonomic potential of this structure in the group (Alegre et al. 2019). ...
... The ongoing systematic review of the Cuban Stenostygninae, undertaken by one of us (AAB), rendered new information for some Cuban genera and for the entire subfamily. This revision has resulted in the redescription of Manahunca bielawskii Šilhavý, 1973 (the type species of the genus Manahunca Šilhavý, 1973), the recognition of Manahunca silhavyi as a junior subjective synonym of M. bielawskii, and the description of two new species from eastern Cuba (Manahunca turquino Alegre, Gainett &Giribet, 2019 andManahunca matazon Alegre, Gainett &Giribet, 2019). In addition, a new diagnosis for Manahunca and its included species were given, along with the update of the geographic distribution and a survey of the sexually dimorphic metatarsus III of several stenostygnines, which revealed the taxonomic potential of this structure in the group (Alegre et al. 2019). ...
... This revision has resulted in the redescription of Manahunca bielawskii Šilhavý, 1973 (the type species of the genus Manahunca Šilhavý, 1973), the recognition of Manahunca silhavyi as a junior subjective synonym of M. bielawskii, and the description of two new species from eastern Cuba (Manahunca turquino Alegre, Gainett &Giribet, 2019 andManahunca matazon Alegre, Gainett &Giribet, 2019). In addition, a new diagnosis for Manahunca and its included species were given, along with the update of the geographic distribution and a survey of the sexually dimorphic metatarsus III of several stenostygnines, which revealed the taxonomic potential of this structure in the group (Alegre et al. 2019). However, the study of the key taxon Caribbiantes, although crucial for the systematic review of the subfamily, was still pending. ...
Caribbiantes cubanus Šilhavý, 1973, type species of genus Caribbiantes Šilhavý, 1973, is redescribed based on abundant material from the type locality, providing new data on its morphological variability and penis morphology. Two new species of Caribbiantes are described, C. obtusus sp. nov. and C. barbai sp. nov., based on additional specimens from central and eastern Cuba, for which we give information on their geographical distribution, intraspecific variability and habitat. A new diagnosis for Caribbiantes and updated geographic distributions of the included species are provided. The sexually dimorphic glandular structures on the metatarsus III of males for all Caribbiantes species are described. Male dimorphism, most evident in the robustness of the chelicerae, is reported for the studied species. The male genitalia patterns of Antillean stenostygnines species are described. The phylogenetic affinity of Antillean stenostygnines with Stenostygnus Simon, 1879, a genus from the north of South America, is discussed based on some morphological traits.
... In arachnids, certain lineages within the order Opiliones ('harvestmen' or 'daddy-long-legs') typify pronounced sexual dimorphism, which often affects the armature and elongation of appendages, and the presence of glands (Martens 1973;Willemart et al. 2010; Painting et al. 2015). The most speciose suborder, Laniatores, also known as armoured harvestmen, encompasses great part of this diversity; at least one-third of the~30 families display sexual dimorphism in robustness, size or armature of chelicerae, pedipalps or legs , and more than half of the families of Laniatores exhibit sexually dimorphic glands, particularly in the legs (reviewed in Willemart et al. 2007Willemart et al. , 2010Felgenhauer 2011, 2013;Pérez-González et al. 2016;Alegre et al. 2019). The dimorphic armature has been the focus of recent attention, specifically in the Neotropical laniatorean family Gonyleptidae, in terms of morphology, ecology and evolutionary history. ...
... These glands can usually be identified by correlation with swollen appendage segments in males, and they differ from the incrassate sexually dimorphic armature by the absence of spines and the presence of associated pores and sometimes setae (Willemart et al. 2010). Descriptions of species with glandlike swollen leg parts abound in the taxonomic literature of Laniatores (Willemart et al. 2010;Alegre et al. 2019) and similar glands are present in the tarsus of leg IV in males of all Cyphophthalmi. Ultrastructural and histological investigations also confirm the presence of glandular tissue that secrete chemicals through tegumental pore openings (Willemart et al. 2010;Felgenhauer 2011, 2013;Alegre et al. 2019). ...
... Descriptions of species with glandlike swollen leg parts abound in the taxonomic literature of Laniatores (Willemart et al. 2010;Alegre et al. 2019) and similar glands are present in the tarsus of leg IV in males of all Cyphophthalmi. Ultrastructural and histological investigations also confirm the presence of glandular tissue that secrete chemicals through tegumental pore openings (Willemart et al. 2010;Felgenhauer 2011, 2013;Alegre et al. 2019). Accordingly, the only published behavioural evidence of their function are studies on two gonyleptid species, in which the males have been observed touching the substrate with these glands, probably to spread chemicals (Fernandes and Willemart 2014;Murayama and Willemart 2015). ...
Sexually dimorphic traits are widespread in animals, and include sex-specific weapons, ornamentation and, although less noticed, glands and associated structures. In arachnids, certain lineages of the order Opiliones exhibit diverse forms of dimorphism in the armature and length of appendages (common in Laniatores), as well as in the presence of sexually dimorphic glands (mostly investigated in Cyphophthalmi), positing harvestmen as promising models to study sexual dimorphism. Whereas the evolution and ecological significance of armature have been the focus of recent attention, sexually dimorphic glands remain understudied in groups other than Cyphophthalmi, despite being widespread in Opiliones. We therefore selected the amphi-Pacific family Zalmoxidae as an ideal taxon to investigate the evolutionary dynamics of this trait. We first describe four new species of Palaeotropical Zalmoxis, including a species with sexually dimorphic glands, and describe the morphology of zalmoxid species with sexually dimorphic glands using scanning electron microscopy. Using a previously assembled six-locus dataset supplemented with new terminals, and applying stochastic character mapping, we infer that sexually dimorphic glands evolved once in the Neotropics and at least four times in the Palaeotropic zalmoxids, revealing the evolutionary lability of this trait.
... The dimorphic condition is present in the Antillean stenostygnines and was first recorded by Šilhavý (1973) for Caribbiantes cubanus Šilhavý, 1973and Martibianta virginsulana Šilhavý, 1973(junior syn. of Heterolacurbs ovalis Roewer, 1912, by Pérez-González & Alegre 2009). Later, it was also noted in males of an undescribed species of Caribbiantes from Cuba (Pérez-González 2000), and finally in males of Manahunca bielawskii Šilhavý, 1973 and Manahunca matazon Alegre, Gainett & Giribet, 2019, by Alegre et al. (2019. Besides, in the Antillean Stenostygninae are seven other species in which the males have only hypertelic chelicerae and another seven species with males having only non-hypertelic chelicerae. ...
... The dimorphic condition is present in the Antillean stenostygnines and was first recorded by Šilhavý (1973) for Caribbiantes cubanus Šilhavý, 1973and Martibianta virginsulana Šilhavý, 1973(junior syn. of Heterolacurbs ovalis Roewer, 1912, by Pérez-González & Alegre 2009). Later, it was also noted in males of an undescribed species of Caribbiantes from Cuba (Pérez-González 2000), and finally in males of Manahunca bielawskii Šilhavý, 1973 and Manahunca matazon Alegre, Gainett & Giribet, 2019, by Alegre et al. (2019. Besides, in the Antillean Stenostygninae are seven other species in which the males have only hypertelic chelicerae and another seven species with males having only non-hypertelic chelicerae. ...
Stenostygnus martensi spec. nov. and Stenostygnus huberi spec. nov. are described from montane localities of the States of Mérida and Lara in Venezuela, and the diagnosis of the previously monotypic genus Stenostygnus Simon, 1879, is emended. The presence of these two new species as local endemics in Andean localities of Venezuela represents a huge range extension for the genus Stenostygnus, and the new localities are also ecologically different from previously known localities in the Amazon Basin. This could indicate the presence of a rich and uncharted diversity of stenostygnines in the Andean regions, particularly in the north of South America.
... This is the first time that a group of atypical setae has been described for the swollen calcaneus in Zalmoxoidea. Gainett et al. (2020) hypothesized a secretion paint-brush mechanism for the group of sulcated setae on the swollen parts of zalmoxid legs, aligning with the same hypothesis proposed to analogous setae on the metatarsus III of Biantidae (Alegre et al. 2019). We believe that this hypothesis is also plausible for the group of modified setae in the swollen metatarsus III of the described species of Icaleptidae. ...
The genus Icaleptes was proposed by Kury & Pérez-González in 2002 to accommodate a single Colombian species, Icaleptes malkini, collected in the southeastern slope of Sierra Nevada de Santa Marta, Cesar Department. In this study, we describe two new species for this previously monotypic genus collected on the northwestern slope of the Sierra Nevada de Santa Marta, in the department of Magdalena. The new species Icaleptes dimorphicus sp. nov. and Icaleptes armasi sp. nov. were included within Icaleptes based on external and genital similarities such as male scutum magnum continuously convex and lacking the sulci; pedipalps with highly reduced setae; chelicerae stout and unarmed with a low, wide bulla; capsula externa of the penis modified into a robust stragulum with two widely separated apical lobes and a wide capsula interna visible from the dorsal aspect with a well-developed parastylar collar. In contrast to Icaleptes malkini, the two new species have the ventral plate of the penis differentiated into two regions, which we interpreted here as a wide pergula basally and a short stout rutrum apically. The new species also have other remarkable sexual dimorphisms such as males with a strong protuberance on coxa IV, an arched movable finger in the chelicerae, and enlarged basitarsomeres on leg III. The modification of coxa IV in males is a strong diagnostic character that facilitates the identification of both new species. The description of two new species that are closely related to the type species Icaleptes malkini helps us to understand the morphological variation of the current concept of Icaleptidae.
... This rigid capsula externa also appears in other biantids such as Antillean Stenostygninae (e.g. Alegre et al. 2019;Alegre-Barroso and Pérez-González 2024) and the Mexican Stygnomma teapense Goodnight & Goodnight, 1951 (obs. pers.). ...
Despite being one of the most conspicuous African opilionids, the members of Lacurbsinae remain one of the least known groups of harvestmen species. All eight previously-known species of Lacurbsinae are inadequately described and poorly illustrated, leaving the morphological characteristics of this subfamily obscure. After more than half a century, we describe a new species of Lacurbsinae. Metalacurbs foordisp. nov. is described, based on a male specimen collected in Ankasa National Park, Ghana, with a detailed description and illustration of its external and genital morphology. This marks the first modern taxonomic description of a species within Lacurbsinae, including an illustration and description of the male genital morphology, a crucial modern taxonomic characteristic for Opiliones and represents a starting point for the taxonomic revision of the subfamily.
The genus Neoscotolemon Roewer, 1912, is herein reviewed and re-diagnosed for the first time using modern taxonomic standards. Neoscotolemon is removed from Grassatores incertae sedis and transferred to the superfamily Samooidea incertae sedis, transl. nov. The genera Citranus Goodnight & Goodnight, 1942, Rula Goodnight & Goodnight, 1942, and Vlachiolus Šilhavý, 1979, are considered new subjective synonyms of Neoscotolemon Roewer, 1912. Neoscotolemon pictipes (Banks, 1908) is redescribed and fully illustrated, including, for the first time, the external and genital morphology of males. Neoscotolemon armasi spec. nov. is described from Isla de la Juventud, Cuba. Five additional species are transferred to Neoscotolemon, including some that were newly ranked from subspecies to species: Neoscotolemon bolivari (Goodnight & Goodnight, 1945) comb. nov., stat. rest. (transferred from Stygnomma), N. cotilla (Goodnight & Goodnight, 1945) comb. nov., nom. rest., stat. rest. (transferred from Stygnomma), N. spinifer (Packard, 1888) comb. rest. (transferred from Stygnomma), N. tancahensis (Goodnight & Goodnight, 1951) comb. nov., stat. prom. (transferred from Stygnomma), and N. vojtechi (Šilhavý, 1979) comb. nov. (transferred from Vlachiolus). Finally, upon reexamination of Neoscotolemon lutzi Goodnight & Goodnight, 1942, the male genital morphology, herein illustrated for the first time, indicates that this species is not related to Neoscotolemon and is therefore transferred to Metapellobunus and combined as Metapellobunus lutzi (Goodnight & Goodnight, 1942) comb. nov. A map is provided showing the known distribution of Neoscotolemon in the southeastern USA, Cuba, Mexico, and Belize with doubtful and unconfirmed records in Jamaica and Cayman Islands. Neoscotolemon is characterized by multiple somatic and genitalic traits. Atop the hourglass-shaped scutum magnum, there is a widely-separated pair of eyes with a large spiniform apophysis placed between them. The pedipalps are robust, and in major males they are elongated and thickened. In major and minor males, the metatarsus III is enlarged, ventrally covered with modified trichomes, and possesses aggregated pores distally. The penis has a ventral plate ending in a deep calyx, armed with two bilateral rows of macrosetae, and a short, pointed stylus that is basally fused to two laminar conductors. Although the family allocation remains uncertain, the re-diagnosis of Neoscotolemon, together with the re-description of the type species, makes this an easily recognizable genus, among Samooidea, that now contains seven species.
The troglobitic harvestman Jimeneziella decui Avram, 1970 is known from four neighboring caves (Cueva de Majana, Cueva de los Golondrinos, Cueva Perla del Agua, and Cueva de Mximo) located in eastern Cuba. We present the first ecological data on a population of this endangered species in Cueva de Mximo. The sex ratio of the population estimated in the main gallery of the cave was not different from 1:1. The spatial distribution observed was uniform, and the density of individuals was 0.48 individuals/m2 (FebruaryMarch) and 0.84 individuals/m2 (November). We describe morphological differences between the sexes and between males. Preliminary morphological and behavioral data suggests the possible existence of two male morphs in J. decui (robust and slender males). The slender males possessed less developed armature on leg IV; and the chelicerae, coxa IV and femur IV were less swollen than robust males. Our observations on male-male interactions suggest that robust males are more aggressive than slender males, which never initiated an attack on robust males, but in some occasions responded to attacks from robust males. The non-aggressive behavior exhibited by robust males towards slender males also suggests that the slender morph is not only a sneaker, but potentially a female mimic, which is also consistent with the morphology of slender males. Ecological and behavioral information also suggests the possible existence of territories defended by robust males with their well-developed weapons, and the presence of females inside them.
Opiliones are iconic arachnids with a Palaeozoic origin and a diversity that
reflects ancient biogeographic patterns dating back at least to the times of
Pangea. Owing to interest in harvestman diversity, evolution and biogeography,
their relationships have been thoroughly studied using morphology
and PCR-based Sanger approaches to infer their systematic relationships.
More recently, two studies utilized transcriptomics-based phylogenomics
to explore their basal relationships and diversification, but sampling was
limiting for understanding deep evolutionary patterns, as they lacked
good taxon representation at the family level. Here, we analysed a set of
the 14 existing transcriptomes with 40 additional ones generated for this
study, representing approximately 80% of the extant familial diversity in
Opiliones. Our phylogenetic analyses, including a set of data matrices
with different gene occupancy and evolutionary rates, and using a multitude
of methods correcting for a diversity of factors affecting phylogenomic data
matrices, provide a robust and stable Opiliones tree of life, where most families
and higher taxa are precisely placed. Our dating analyses using alternative
calibration points, methods and analytical parameters provide well-resolved
old divergences, consistent with ancient regionalization in Pangea in some
groups, and Pangean vicariance in others. The integration of state-of-the-art
molecular techniques and analyses, together with the broadest taxonomic
sampling to date presented in a phylogenomic study of harvestmen, provide
new insights into harvestmen interrelationships, as well as an overview of
the general biogeographic patterns of this ancient arthropod group.
The type specimens of Fijicolana tuberculata Roewer, 1963 were re-examined and the male genital morphology is illustrated and described for the first time. Despite the presence of several morphological features that are typical of Samoidae, such as the presence of scopulae on legs III and IV, genital morphology unambiguously indicates that this species belongs to the Zalmoxidae rather than to the Samoidae. Fijicolana Roewer, 1963 is newly synonymized with Zalmoxis Sørensen, 1886. However, the newly implied combination is preoccupied by Z. tuberculatus Goodnight & Goodnight, 1948 thus the replacement name Zalmoxis roeweri nom. nov. is proposed to avoid secondary homonymy. The definition of Z. roeweri nom. nov. is amended, and the morphology of this species is compared with other representatives of Zalmoxidae and Samoidae. We conclude that the presence of scopulae alone is not a sufficiently diagnostic characteristic for Samoidae and, therefore, correctly placing taxa into families within Samooidea + Zalmoxoidea requires additional morphological evidence (e.g. genital morphology). In light of this result, we point out that the "scopulated" Australasian samoids Badessania metatarsalis Roewer, 1949, Sawaiellus berlandi Roewer, 1949 and Parasamoa gressitti Goodnight & Goodnight, 1957 require re-examination in order to detect potential errors in their family placement.
Alternative reproductive tactics in animals are commonly associated with distinct male phenotypes resulting in polymorphism of sexually selected weapons such as horns and spines. Typically, morphs are divided between small (unarmed) and large (armed) males according to one or more developmental thresholds in association with body size. Here, we describe remarkable weapon trimorphism within a single species, where two exaggerated weapon morphs and a third morph with reduced weaponry are present. Male Pantopsalis cheliferoides harvestmen display exaggerated chelicerae (jaws) which are highly variable in length among individuals. Across the same body size spectrum, however, some males belong to a distinct second exaggerated morph which possesses short, broad chelicerae. Multiple weapon morphs in a single species is a previously unknown phenomenon and our findings have significant implications for understanding weapon diversity and maintenance of polymorphism. Specifically, this species will be a valuable model for testing how weapons diverge by being able to test directly for the circumstances under which a certain weapon type is favoured and how weapon shape relates to performance.
Resumen: Se ofrecen dos nuevas sinonimias en Opiliones Laniatores neotropicales. La especie Stygnomma maya Goodnight & Goodnight, 1951 (Stygnommatidae) es considerada un sinónimo posterior de Stygnomma granulosa (Goodnight & Goodnight, 1947); de igual forma, Stygnoplus dominicanus (Roewer, 1943) (Stygnidae) es un sinónimo posterior de Stygnoplus tuberculatus (Goodnight & Goodnight, 1942). También se ofrecen algunos datos referentes a aspectos morfológicos y de distribución de las especies. Abstract: Two new synonymies of Neotropical laniatorid harvestmen are proposed. The species Stygnomma maya Goodnight & Goodnight, 1951 (Stygnommatidae) is considered as a junior synonym of Stygnomma granulosa (Goodnight & Goodnight, 1947); likewise, Stygnoplus dominicanus (Roewer, 1943) (Stygnidae) is seen as a junior synonym of Stygnoplus tuberculatus (Goodnight & Goodnight, 1942). Some information is also provided on the morphology and distribution of these species.
Males of Iporangaia pustulosa (Arachnida: Opiliones) have a sexually dimorphic metatarsus IV, which is thicker and with more glandular pores in males. Here we tested the hypothesis that this glandular area is used by males to leave chemicals in the environment, predicting that the animals would rub the metatarsus IV against the substrate. We have made recordings both in the field and in the laboratory, in several distinct contexts during the day and at night, comprising 67 hours of observations. We also experimentally tested the reaction of both sexes to a filter paper rubbed on the metatarsus gland, with adequate controls. We report and describe for the first time that the metatarsal gland of I. pustulosa is used to leave chemicals on the substrate by rubbing or touching it against the substrate. We also provide evidence that males can control the release of secretions of the metatarsal gland IV.
In Opiliones, one of the largest orders within the class Arachnida, with more than 6000 described species, sexual dimorphism can be widespread and exaggerated. This great variety of forms of gender-based dimorphism suggests that sexual selection may play an important role in the diversification of some lineages. It also impacts species identification, assignment of females to described species and biodiversity assessments. Here we use DNA-sequence-based species discovery methods (the Poisson Tree Processes model with Bayesian support values, bPTP, and the Generalized Mixed Yule–Coalescent approach, GMYC, accounting for phylogenetic uncertainty) to shed light on the morphological disparity displayed in several species of neopilionid harvestmen from New Zealand. Both species delimitation analyses recovered many clades that coincide with our prior assignment of morphospecies, based solely on males, and allowed us to assign females and juveniles to these species as well as to identify putative new species and to assign some unidentified species to genera. Several genetic species, particularly Forsteropsalis inconstans and Pantopsalis cheliceroides-listeri, showed complex morphological disparity in the size and shape of the male chelicerae, but also in the general size and coloration patterns of the males. The systematic implications of our results and a possible ecological explanation for the exaggerated traits are discussed. Following our findings, the following taxonomic action is taken: Forsteropsalis nigra is considered a junior synonym of F. inconstans (new synonymy).
Les glandes sexuellement dimorphiques ont évolué de manière convergente chez les animaux et sont souvent à l’origine de la production de phéromones. Dans le sous-ordre des Laniatores (Arachnida, Opiliones), ce type de glandes est très répandu, mais il n’existe aucune observation concernant la manière dont elles sont utilisées. À l’aide de la microscopie électronique à balayage et d’une approche comportementale, nous avons pu décrire l’existence d’ouvertures glandulaires et le mode d’utilisation de ces glandes chez deux opilions Cosmetidae : Gryne perlata et Gryne coccinelloides. Les mâles de ces deux espèces présentent des ouvertures glandulaires sur les métatarses des pattes I et IV. Différents types de comportements ont pu être observés selon l’espèce étudiée. Les glandes du métatarse I sont uniquement frottées contre les autres pattes, alors que celles situées sur le métatarse IV sont utilisées de différentes façons : elles peuvent être, soit frottées contre les autres pattes et contre le sol, soit mises uniquement en contact avec le sol.
The Australian fauna is assessed for short-range endemism at the species level, i.e. the prevalence of species with naturally small ranges of less than 10,000 km². The phenomenon is found to be widespread and several groups are found to consist principally of short-range endemics: Gastropoda (snails and slugs, both freshwater and terrestrial), Oligochaeta (earthworms), Onychophora (velvet worms), Araneae (mygalomorph spiders), Schizomida (schizomids), Diplopoda (millipedes), Phreatoicidea (phreatoicidean crustaceans), and Decapoda (freshwater crayfish). The majority of taxa with high numbers of short-range endemics possess similar ecological and life-history characteristics, such as poor powers of dispersal and confinement to discontinuous habitats. The conservation of such groups is often hampered by poor taxonomic knowledge, but modern, comprehensive biotic surveys will be helpful in identifying short-range endemics.
All the systematic literature (774 references) of the suborder Laniatores of the Americas up
to year 2002 is tabulated to generate a thorough annotated classification. 2372 species in
746 genera of Laniatores of the New World are listed. 26 families of Laniatores are
recognized as valid, of which 21 occur in the New World. The most diverse family is
Gonyleptidae (823 species), followed by Cosmetidae (710 species), both endemic to the
New World. Synonymies, revalidations, replacement names and emended spellings are
done when necessary. The complete list of nomenclatural acts herein proposed is given. The
new family Escadabiidae Kury & Pérez is proposed, the new subfamily Ampycinae Kury is
proposed in Gonyleptidae. Countries and ultramarine departments included are 1) South
America: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, French Guyana, Guyana,
Paraguay, Peru, Suriname, Trinidad and Tobago, Uruguay and Venezuela; 2) Central
America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; 3)
Antilles: Bahamas, Bermuda Island, Caicos Islands, Cayman Islands, Cuba, Dominican
Republic, Haiti, Jamaica, Leeward Islands, Netherland Antilles, Puerto Rico, Tortuga Island,
UK Virgin Islands, U.S. Virgin Islands, Windward Islands; 4) North America: Canada,
Greenland, Mexico, U.S.A. First order administrative divisions (departments, provinces,
states) for all most diverse countries are interpolated in the locality names. A list of species
by first order administrative divisions is provided for all countries treated. The most diverse
country is Brazil, with 855 species of Laniatores, followed by Venezuela with 328 species.
An exhaustive list of the depository institutions of the type material with curators and
contacts addresses is given.
Amendments are made to inconsistencies, mistakes and omissions in the catalogue of American Laniatores by Kury (2003). Discrepancies between dates given in Kury (2003) and Neave's Nomenclator and the Zoological Record are dis-cussed. Accurate issue dates for relevant publications are used to define priorities. Etymologies are surveyed for generic names, establishing their grammatical gender, so specific names are inflected according to the provisions of and must be placed as an "incorrect subsequent spelling" of Hernandaria Sørensen, 1884. The valid genus name is Parahernandria Good-night & Goodnight, 1947 (stat. res.). Zaraxolia Strand, 1942 (stat. res.) is revalidated from the synonymy of Neocynorta Roewer, 1915 with Zarax devians Sørensen, 1932 as type species. Friburgoia Mello-Leitão, 1932 [December] is deemed a junior subjective synonym of Schenkelibunus Strand, 1932 [September] (stat. res.) (inverted precedence between syn-onyms). Liops Mello-Leitão, 1940 (non Fieber, 1870, nec Gidley, 1906) is a junior homonym and is replaced by its first available synonym, Corcovadesia Soares & Soares, 1954 (stat. res.). The following unavailable generic names are for-mally described as new: (1) Jimeneziella Kury & Alonso-Zarazaga, gen. nov. (type species: Jimeneziella decui Avram, 1973); (2) Euminua Kury & Alonso-Zarazaga, gen. nov. (type species: Euminua brevitarsa Sørensen, 1932). The follow-ing homonym generic names are replaced: (1) Cranellus Roewer 1932, with Narcellus Kury & Alonso-Zarazaga, nom. nov., (2) Metapachylus Pickard-Cambridge, 1905, with Pyropharynx Kury & Alonso-Zarazaga, nom. nov.; (3) Ovalia González-Sponga, 1987, with Oo Kury & Alonso-Zarazaga, nom. nov.; (4) Tiara González-Sponga, 1987 with Mitraia Kury & Alonso-Zarazaga, nom. nov.; (5) Limonia González-Sponga, 1998, with Manuelangelia Kury & Alonso-Zaraza-ga, nom. nov. Gonyleptes melloleitaoi Kury & Alonso-Zarazaga, nom. nov. is a replacement name for Gonyleptes curvi-cornis Mello-Leitão, 1932. Discocyrtus confusus Kury, 2003 is unavailable, so this species is here re-described as Discocyrtus confusus Kury n. sp. New combinations are: Parahernandria spinosa (Banks, 1909) (from Hernandaria), Schenkelibunus impar (Mello-Leitão, 1932) (from Friburgoia), Schenkelibunus perditus (Mello-Leitão, 1927) (from Friburgoia), Narcellus balthazar (Roewer, 1932) (from Cranellus), Narcellus montgomeryi (Goodnight & Goodnight, 1947) (from Cranellus), Pyropharynx gracilis (Pickard-Cambridge, 1905) (from Metapachylus), Oo spinosum (González-Sponga, 1999) (from Ovalia), Mitraia unispina (González-Sponga, 1987) (from Tiara), Manuelangelia tuberosa (González-Sponga, 1998) (from Limonia), Zaraxolia devians (Sørensen, 1932) (from Zarax), Corcovadesia hexabunus (Mello-Leitão, 1940) (from Liops) and Corcovadesia venefica (H. Soares, 1966) (from Liops). The following genera of Pachylinae, which appeared in conflicting subfamilies in Kury 2003, are formally transferred to the Ampycinae: Ampy-cella Roewer, 1929, Glysterus Roewer, 1931, Hernandarioides Pickard-Cambridge, 1905, Parahernandria Goodnight & Goodnight, 1947 and Hutamaia Soares & Soares, 1977. A list of taxa described in 2003 is interpolated. A complementary list of the 2004–2009 systematic literature on the subject is given.
In most taxonomic systems it is not uncommon tha t some autapomorphies are taken as "major features" whic h justify the creation of new monotypic genera . That is, suc h characters would denote a "higher grade", and therefor e should "deserve" a higher rank . It can be felt notably i n the opilioni' classification by ROEWER (see e . g . 1923 :10) , which is furthermore based too heavily on the armature o f the dorsal scute and tarsal segmentation . Consequently, al l possible combinations of spines and tubercles of the terga l areas become virtual genera, many of them defined only b y the absence of characters . The definition of genera of Gonyleptidae is critical , as can be seen by the discovery of a new mitobatine specie s from Southeastern Brazil, which would be undoubtedly regarde d as a new genus if the Roewerian approach was used . Here i s proposed instead its inclusion in Mitobates Sundevall,1833 . The characters involved were polarized by outgroup compariso n and will be explained in detail in a complete phylogeneti c analysis of the Mitobatinae (Kury, in prep .) .
In various animal species, male sexual dimorphic characters may be used during intrasexual contests as ornaments to attract females, or to hold them before, during, or after copulation. In the well-known harvestman, Phalangium opilio L., 1758, the behavioral functions of these male sexually dimorphic structures have never been studied in detail. Therefore, in addition to a morphometric study, 21 male contests and 43 sexual interactions were analyzed. Our observations revealed that during contests, the male cheliceral horns form a surface by which the contestants use to push each other face-to-face while rapidly tapping their long pedipalps against the pedipalps of the opponent, occasionally twisting the opponents pedipalp. Scanning electron micrographs revealed contact mechanoreceptors on the pedipalp that would detect the intensityfrequency of contact with the contenders pedipalp. Larger males won almost all contests, whereas the loser rapidly fled. During sexual interactions, the longer pedipalps of the male held legs IV of the female, whereas males with shorter pedipalps held the female by legs III. No contact with the male pedipalps and chelicerae by the females was visible before, during, or after copulation. Soon after copulating, males typically bent over the female, positioning their cheliceral horns against the femaless dorsum. Consequently, our data show that the cheliceral horns and the longer pedipalps of the male seem to play an important role, during both intersexual and intrasexual encountering.
We investigated the internal phylogeny of Laniatores, the most diverse suborder of Opiliones, using sequence data from 10 molecular loci: 12S rRNA, 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase subunit I (COI), cytochrome b, elongation factor-1α, histones H3 and H4, and U2 snRNA. Exemplars of all previously described families of Laniatores were included, in addition to two families – Petrobunidae, fam. nov. and Tithaeidae, fam. nov. – that we erect herein. Data analyses were based on maximum likelihood and Bayesian approaches on static alignments, and included phylogenetic tree estimation, molecular dating, and biogeographic analysis of ancestral area reconstruction. The results obtained include the monophyly of Laniatores and the infraorder Grassatores – the focus of this study – as well as support for numerous interfamilial relationships. The two new families described cluster with other South-east Asian families (Podoctidae and Epedanidae). Diversification of Laniatores is estimated at ~348 Mya, and origin of most Grassatores superfamilies occurs in a ~25 million year span of time immediately after the end-Permian mass extinction (254 Mya). Ancestral range reconstruction of the clade (Samooidea + Zalmoxoidea) suggests that the ancestral range of Samooidea comprises West Tropical Gondwana (West Africa + Neotropics), whereas that of Zalmoxoidea is exclusively Neotropical. The following additional taxonomic changes are proposed: (1) Remyus is transferred to Phalangodidae, and (2) Escadabiidae and Kimulidae are transferred to Zalmoxoidea.
Arachnids of the order Opiliones (harvestmen), which includes around 6000 species, have a pair of scent glands that open at the sides of the body, producing substances used as defence. Several types of behavioural, morphological and chemical defensive mechanisms have been identified in the order as a whole, although some of these tactics were restricted to particular groups. Only around 60 species have been studied from this perspective so far, more than half of which belong to the largest harvestman family within the order Laniatores, the Gonyleptidae, and have only recently been studied in an evolutionary perspective, showing the usefulness of defensive characters in taxonomy and evolutionary biology. Within Laniatores, the Grassatores clade includes the Gonyleptidae and 20 additional families, mostly poorly or not previously studied. We describe the morphology of the structures involved in fluid displacement during chemical defence in 15 of these families (data on two additional families are available from the literature) and discuss the evolution of such traits based on an available phylogenetic hypothesis of relationships within Grassatores, using the representatives of Triaenonychidae (a non-Grassatores family of Laniatores) for comparison. We conclude that most non-gonyleptoid Grassatores share (maybe plesiomorphically) a series of characteristics, mostly strongly different from what is observed within the gonyleptoids, and that smaller groups seem to share diagnostic features related to chemical defence, as is the case of stygnids, cosmetids and triaenonychines, and especially of manaosbiids and cranaids, whose defensive morphologies largely resemble those of derived gonyleptids. The following main synapomorphies were detected: (a) Grassatores: the presence of a deep and well-defined descending channel; (b) Samooidea + Zalmoxoidea + Assamioidea + Gonyleptoidea: lateral pegs along the lateral channel; (c) Samooidea + Zalmoxoidea: deep channels forming an H on the dorsal scute; (d) Gonyleptoidea: ozopore cutting dorsally (reversing in Agoristenidae and Stygnidae to a laterally placed oval ozopore), a wide and smooth lateral channel, reversing to a lateral channel whose bottom is covered with either small plates (Agoristenidae) or high tubercles (Stygnidae), and apophyses of coxa II close to or covering the ozopore.
Opiliones are one of the largest arachnid orders, with more than 6,500 species in 50 families. Many of these families have been erected or reorganized in the last few years since the publication of The Biology of Opiliones. Recent years have also seen an explosion in phylogenetic work on Opiliones, as well as in studies using Opiliones as test cases to address biogeographic and evolutionary questions more broadly. Accelerated activity in the study of Opiliones evolution has been facilitated by the discovery of several key fossils, including the oldest known Opiliones fossil, which represents a new, extinct suborder. Study of the group's biology has also benefited from rapid accrual of genomic resources, particularly with respect to transcriptomes and functional genetic tools. The rapid emergence and utility of Phalangium opilio as a model for evolutionary developmental biology of arthropods serve as demonstrative evidence of a new area of study in Opiliones biology, made possible through transcriptomic data. Expected final online publication date for the Annual Review of Entomology Volume 60 is January 07, 2014. Please see http://www.annualreviews.org/catalog/pubdates.aspx for revised estimates.
In at least four closely related families of the diverse harvestmen lineage Gonyleptoidea, males may possess sexually dimorphic tarsal glands in the swollen tarsomeres of the basitarsus and/or metatarsus of leg I. The first histological and ultrastructural examination of the sexually dimorphic tarsal glands in leg I focused only on Manaosbiidae. In this study, we examine the morphology and ultrastructure of the sexually dimorphic glands, and their associated glandular openings, found in the basitarsus and/or metatarsus of leg I of males representing Cosmetidae, Gonyleptidae, and Cranaidae (glandular openings only). In cosmetids and gonyleptids, the tarsal glands are made up of 20–60 glandular units that form distinct groups within the prolateral and retrolateral half of the tarsomere. Each glandular unit consists of a pair of terminal secretory cells, an intercalary cell wrapped around the receiving canal, and a canal cell tightly wrapped around the length of the conducting canal. Cosmetidae, Gonyleptidae, and Cranaidae exhibit remarkably similar tarsal glands and gland openings although the location of the glands in the leg differs slightly among them. Males of these three families exhibit markedly different glands and glandular openings compared to males of the family Manaosbiidae. The sexually dimorphic tarsal glands may provide an important morphological character for determining phylogenetic relationships among gonyleptoid families. Finally, we provide morphological and ultrastructural data for the common tegumental glands. These data indicate that the sexually dimorphic tarsal glands are strikingly similar to, and may possibly be derived from, the tegumental glands.
Studies, especially with transmission electron microscopy of developmental stages, show that the many superficially different spines, bristles, hairs, etc. can be classified into 4 major types: (1) multicellular with cells similar to those of the general epidermis (spines); (2) multicellular with differentiation of a segregated trichoid complex (setae); (3) unicellular (acanthae); and (4) subcellular in the sense of several or many per cell (microtrichia). Except for eversible pouches, there are no flaccid projections because the cuticle of setae and other projections is always sclerotized.Spines, as the term is used above, occur primarily on external surfaces, except for comparable structures in the proventriculi of some insects. Setae are on external surfaces only, and most of them retain a sensory function. Acanthae are best known on internal surfaces of foregut and reproductive ducts but can be found on external surfaces also. Microtrichia occur mostly on external surfaces but occasionally on the taenidia of tracheae. Only the socketed setae include one or more sense cells, and hence only setae are sensory (but spines can bear setae). Several or numerous subtypes can be recognized for each of these types. All of the types can have the cells withdraw; they then usually become filled with cuticle and even commonly become underlaid by cuticle; the producing cells may or may not persist. Such solid hairs commonly cannot be identified as to type without knowledge of their development.
Morphological characters are essential for establishing phylogenetic relationships, delimiting higher-level taxa, and testing phylogenetic relationships inferred from molecular sequence data. In cases where relationships between large clades remain unresolved, it becomes imperative to establish which character systems are sound predictors of phylogenetic signal. In the case of Laniatores, the largest suborder of Opiliones, some superfamilial relationships remain unresolved or unsupported, and traditionally employed phenotypic characters are typically of utility only at the family level. Here we investigated a promising set of morphological characters that can be discretized and scored in all Opiliones: cuticular structures of the distal podomeres (metatarsi and tarsi). We intensively sampled members of all known families of Laniatores, and define here three new, discrete appendicular characters toward refinement of Laniatores superfamilial systematics: metatarsal paired slits (MPS; occurring in all Laniatores except Sandokanidae), proximal tarsomeric gland (PTG; in Icaleptidae, Fissiphalliidae, and Zalmoxidae), and tarsal aggregate pores (TAP; found in Gonyleptoidea, Epedanoidea, and Pyramidopidae). We conducted statistical tests on each character to characterize the strength of phylogenetic signal and assess character independence, based on alternative tree topologies of Laniatores. All three characters had high retention indices and bore significantly strong phylogenetic signal. Excepting one pairwise comparison, morphological characters did not evolve in a correlated manner, indicating that appendicular morphology does not constitute a single character system. Our results demonstrate the predictive power and utility of appendicular characters in Opiliones phylogeny, and proffer a promising source of diagnostic synapomorphies for delimiting superfamilies.
The limits of zalmoxid distribution in Southeast Asia are poorly understood, but a focus of recent research. Here we describe six new species of litter-inhabiting harvestmen in the genus Zalmoxis Sørensen, 1886 (Opiliones: Laniatores: Zalmoxidae) using light microscopy and SEM. Three of these species are from the Philippine Islands (Zalmoxis gebeleizis sp. nov., Zalmoxis derzelas sp. nov., and Zalmoxis sabazios sp. nov.) and the other three from Borneo (Zalmoxis zibelthiurdos sp. nov., Zalmoxis bendis sp. nov., and Zalmoxis kotys sp. nov.). The collecting localities of these species add to the known range of Zalmoxidae, which have not previously been reported from Borneo. The new species add to known morphological variation of Zalmoxis, specifically with respect to sexually dimorphic tarsomeres, body size, and armature of the anal plate.
We investigate the phylogeny, biogeography, time of origin and diversification, ancestral area reconstruction and large-scale distributional patterns of an ancient group of arachnids, the harvestman suborder Cyphophthalmi. Analysis of molecular and morphological data allow us to propose a new classification system for the group; Pettalidae constitutes the infraorder Scopulophthalmi new clade, sister group to all other families, which are divided into the infraorders Sternophthalmi new clade and Boreophthalmi new clade. Sternophthalmi includes the families Troglosironidae, Ogoveidae, and Neogoveidae; Boreophthalmi includes Stylocellidae and Sironidae, the latter family of questionable monophyly. The internal resolution of each family is discussed and traced back to its geological time origin, as well as to its original landmass, using methods for estimating divergence times and ancestral area reconstruction. The origin of Cyphophthalmi can be traced back to the Carboniferous, whereas the diversification time of most families ranges between the Carboniferous and the Jurassic, with the exception of Troglosironidae, whose current diversity originates in the Cretaceous/Tertiary. Ancestral area reconstruction is ambiguous in most cases. Sternophthalmi is traced back to an ancestral land mass that contained New Caledonia and West Africa in the Permian, whereas the ancestral landmass for Neogoveidae included the south-eastern USA and West Africa, dating back to the Triassic. For Pettalidae, most results include South Africa, or a combination of South Africa with the Australian plate of New Zealand or Sri Lanka, as the most likely ancestral landmass, back in the Jurassic. Stylocellidae is reconstructed to the Thai-Malay Penisula during the Jurassic. Combination of the molecular and morphological data results in a hypothesis for all the cyphophthalmid genera, although the limited data available for some taxa represented only in the morphological partition negatively affects the phylogenetic reconstruction by decreasing nodal support in most clades. However, it resolves the position of many monotypic genera not available for molecular analysis, such as Iberosiro, Odontosiro, Speleosiro, Managotria or Marwe, although it does not place Shearogovea or Ankaratra within any existing family. The biogeographical data show a strong correlation between relatedness and formerly adjacent landmasses, and oceanic dispersal does not need to be postulated to explain disjunct distributions, especially when considering the time of divergence. The data also allow testing of the hypotheses of the supposed total submersion of New Zealand and New Caledonia, clearly falsifying submersion of the former, although the data cannot reject the latter. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 92–130.
Chemical communication is an important aspect of arthropod biology especially for those arthropods with limited abilities to detect visual and acoustic signals. Sexually dimorphic glands are often associated with the production of pheromones, which play a role in reproductive processes. In the family Manaosbiidae (Opiliones: Laniatores), males exhibit an enlarged, swollen, often fused, and spindle-like basitarsus on leg I. In this study, we provide a novel description of the morphology and ultrastructure of the glandular structures found in the proximal swollen tarsomeres of the male manaosbiid Rhopalocranaus albilineatus Roewer, 1932 and compare the external leg I morphology with that of two other manaosbiid harvestmen (Barrona williamsi Goodnight and Goodnight, 1942 and Cranellus montgomeryi Goodnight and Goodnight, 1947). The two proximal tarsomeres of the male R. albilineatus leg I contain two large, paired, acinar glands consisting of many glandular cells. Cells empty their secretory products into a large, branched epicuticular duct, which exits the leg via a pore on the ventral region. Thus, a total of four glandular structures are present within the two swollen tarsomeres and each possesses a conducting canal and pore. Finally, we discuss possible roles of these basitarsal glands in manaosbiid reproductive biology based on the present understanding of sexually dimorphic glands in other terrestrial arthropods (i.e., insects and spiders).
The evolution of sexually dimorphic traits has been the focus of much theoretical work, but empirical approaches to this topic have not been equally prolific. Males of the neotropical family Gonyleptidae usually present a strong fourth pair of legs armed with spines, but their functional significance is unknown. We investigated the putative functions of the leg armature in the harvestman Neosadocus maximus. Being a non-visual species, the spines on male legs can only be perceived by females through physical contact. Thus, we could expect females to touch the armature on the legs of their mates if they were to evaluate it. However, we found no support for this hypothesis. We did show that (1) leg armature is used as a weapon in contests between males and (2) spines and associated sensilla are sexually dimorphic structures involved in "nipping behavior", during which a winner emerged in most fights. Finally, we demonstrate that five body structures directly involved in male-male fights show positive allometry in males, presenting slopes higher than 1, whereas the same structures show either no or negative allometry in the case of females. In conclusion, leg armature in male harvestmen is clearly used as a device in intrasexual contests.
The arachnids of the order Opiliones (harvestmen) have a pair of scent glands opening at the sides of the body, the substances of which are used in defense. Several types of behavioral, morphological and chemical defensive mechanisms have been assigned to the order as a whole, although some of these tactics were restricted to particular groups. Only around 25 species have been studied from this perspective so far. In the present paper, we analyzed 33 species (mostly from the largest harvestmen family, the Gonyleptidae) aiming at recognizing the usefulness of the defensive characters in taxonomy and evolutionary biology. We observed the morphology of the gland opening (ozopore) area and the defensive behavior, and their relationship, and mapped these traits on an available phylogenetic hypothesis of relationship within Gonyleptidae. As outgroups, we analyzed Cosmetidae and Stygnidae. Combining the observed behavioral characters of the emission of defensive secretion (near the ozopore, with liquid displacement through an integumentary groove, or in form of a jet) with the morphological types of the gland opening (direction of the integumentary dome that surrounds the gland opening, presence of two openings and the relationship between their sizes, and presence of a V-shaped cut at the anterior opening), we recognized eight patterns. In addition, we could examine the evolution of such traits within Gonyleptidae.
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