Embryonic Development of Ampheres leucopheus and lporangaia pustulosa (Arachnida: Opiliones: Gonyleptidae)

The first studies concerning the embryonic development of harvestmen started in the late 19th century, and focused mostly on holarctic species, and only three species of the suborder Laniatores (the largest, among the four suborders considered presently) were studied. Moreover, the last studies on embryology of harvestmen were made during the late 1970s. This study focused on the embryonic development of Ampheres leucopheus (Gonyleptidae, Caelopyginae) and Iporangaia pustulosa (Gonyleptidae, Progonyleptoidellinae). The embryonic development was followed in the field, by taking daily photographs of different eggs during about 2 months. When laid, eggs of A. leucopheus and I pustulosa have approximately 1.13 and 1.30 mm in diameter, respectively, and the second is embedded in a large amount of mucus. The eggs grow, mainly due to water absorption at the beginning of the process, and they reach a diameter of about 1.35 and 1.59 mm, respectively, close to hatching. It took, respectively, 29-56 days and 35-66 days from egg laying to hatching. For the description of the embryonic development, we use photographs from the field, SEM micrographs, and histological analysis. This allowed us, for instance, to document the progression of structures and pigmentation directly from live embryos in the field, and to record microstructures, such as the presence of perforations in the cuticle of the embryo in the place where eyes are developing. Yet, contrary to what was expected in the literature, we record an egg tooth in one of the studied laniatoreans. J. Exp. Zool. (Mol. Dev. Evol) 314B:489-502, 2010. (C) 2010 Wiley-Liss, Inc.

Review of the Brazilian harvestman genus Roeweria Mello-Leitao, 1923 (Opiliones: Gonyleptidae)

The genus Roeweria Mello-Leitao, 1923 is revised and a variation on external morphological characters and male genitalia is presented for Roeweria virescens (Mello-Leitao, 1923). The monotypic genus Harpachylus Roewer, 1943 is a junior synonym of Roeweria Mello-Leitao, 1923 because its type-species, H. tibialis Roewer, 1943 is a junior synonym of the type-species Roeweria bittencourti Mello-Leitao, 1923. Roeweria garrincha sp. n. from Cananeia, Sao Paulo, Brazil, is described and can be distinguished from other members of the genus by the presence of a ventral process on the penis and by very large ventral apophyses on the apex of the male femur and patella IV.

Conventional and ultrastructural analyses of the chromosomes of Discocyrtus pectinifemur (Opiliones, Laniatores, Gonyleptidae)

Among the Opiliones, species of the suborders Cyphophthalmi, Eupnoi, Dyspnoi and Laniatores have shown very diverse diploid chromosome numbers. However, only certain Eupnoi species exhibit XY/XX and ZZ/ZW sex chromosome systems. Considering the scarcity of karyotypical information and the absence of structurally identifiable sex chromosomes in the suborder Laniatores, we decided to analyse the chromosomes and bivalents of Discocyrtus pectinifemur (Gonyleptidae) to identify possible sex differences. Testicular cells examined under light microscopy showed it high diploid number, 2n = 88, meta/submetacentric chromosome morphology and a nucleolar organizer region on pair 35. Prophase I microspreading observed in transmission electron microscopy exhibited 44 synaptonemal complexes with similar electron density and thickness. The total and regular synapsis between the chromosomes of the bivalents was also noted in pachytene nuclei. Male mitotic and meiotic chromosomes revealed no distinct characteristic that could be related to the occurrence of heteromorphic sex chromosomes. Evolutionary trends of chromosome differentiation in the four suborders of Opiliones are discussed here.

A systematic revision of Goniosomatinae (Arachnida : Opiliones : Gonyleptidae), with a cladistic analysis and biogeographical notes

Goniosomatine harvestmen have strongly armed pedipalps, generally large bodies and, commonly, very long legs (sometimes more than 20 cm), and are distributed in the Brazilian Atlantic forest, from southern Bahia to Santa Catarina. Since they are conspicuous animals and individuals of some species tend to concentrate in caves (and also under rock boulders), they have been (and still are) the target of several studies, especially those focusing on reproductive and defensive behavior, population ecology, physiology, chromosomes, etc. In spite of their importance for biological studies (some species constitute important and frequently used models for these studies), the taxonomy of Goniosomatinae has faced some problems, including misidentification, a large number of undescribed species and the lack of a phylogenetic hypothesis for the relationships among its species (which would allow evolutionary studies to be made). The last taxonomic changes in the subfamily were made 60 years ago. Considering a taxonomic revision and cladistic analysis of the subfamily to be of paramount importance, the main scope of the present paper is to provide a cladistic analysis and taxonomic revision of the species of Goniosomatinae and a new arrangement of genera (and species). The main taxonomic changes are given as follows. Six genera are recognised within the subfamily: Goniosoma; the newly described genus Pyatan; the reestablished genera Serracutisoma, Heteromitobates and Mitogoniella; and Acutisoma. New generic synonyms include: Glyptogoniosoma = Goniosomella = Lyogoniosoma = Metalyogoniosoma = Xulapona = Goniosoma, Acutisomelloides = Pygosomoides = Spelaeosoma = Serracutisoma; and Acutisomella = Heteromitobates. Newly described species include: Goniosoma capixaba; G. apoain; Pyatan insperatum DaSilva, Stefanini-Jim & Gnaspini; Serracutisoma pseudovarium; S. fritzmuelleri; S. guaricana; Heteromitobates anarchus; H. harlequin; H. alienus; Mitogoniella taquara; M. unicornis; and Acutisoma coriaceum. New combinations include: Goniosoma macracanthum (Mello-Leitao, 1922); G. unicolor (Mello-Leitao, 1932); G. carum (Mello-Leitao, 1936); Serracutisoma proximum (Mello-Leitao, 1922); S. banhadoae (Soares & Soares, 1947); S. molle (Mello-Leitao, 1933); S. thalassinum (Simon, 1879); S. catarina (Machado, Pinto-da-Rocha & Ramires, 2002); S. inerme (Mello-Leitao, 1927); S. spelaeum (MelloLeitao, 1933); Heteromitobates inscriptus (Mello-Leitao, 1922); H. albiscriptus (Mello-Leitao, 1932); Mitogoniella modesta (Perty, 1833); and M. badia (Koch, 1839). Reestablished combinations include: Mitogoniella indistincta MelloLeitao, 1936 and Acutisoma longipes Roewer, 1913. New speci. c synonyms include: Acutisomella cryptoleuca = Acutisomella intermedia = Goniosoma junceum = Goniosoma patruele = Goniosoma xanthophthalmum = Metalyogoniosoma unum = Goniosoma varium, Goniosoma geniculatum = Goniosoma venustum; Goniosomella perlata = Progoniosoma minense = Goniosoma vatrax, Glyptogoniosoma perditum = Progoniosoma cruciferum = Progoniosoma tijuca = Goniosoma dentipes; Leitaoius iguapensis = Leitaoius viridifrons = Serracutisoma proximum; Acutisoma marumbicola = Acutisoma patens = Serracutisoma thalassinum; Progoniosoma tetrasetae = Serracutisoma inerme; and Acutisoma monticola = Leitaoius nitidissimus = Leitaoius xanthomus = Mitogoniella mutila = Acutisoma longipes. The following species are considered species inquirenda: Goniosoma lepidum Gervais, 1844; G. monacanthum Gervais, 1844; G. obscurum Perty, 1833; G. versicolor Perty, 1833; and Mitogoniella badia (Koch, 1839). The monotpic genus Goniosomoides Mello-Leitao, 1932 (and its species, G. viridans Mello-Leitao, 1932) is removed from Goniosomatinae and considered incertae sedis.

Systematic revision and cladistic analysis of the Brazilian subfamily Sodreaninae (Opiliones: Gonyleptidae)

Sodreaninae is reviewed and all ten species are combined under its type genus, Sodreana Mello-Leitao, 1922, according to a cladistic analysis of morphological characters, which revealed a pectinate pattern of clades. The subfamily is endemic to the Brazilian Atlantic rainforest from Santa Catarina state to Rio de Janeiro state. Sodreana is herein considered a senior synonym of Stygnobates Mello-Leitao, 1927, Zortalia Mello-Leitao, 1936, Gertia B. Soares & H. Soares, 1946 and Annampheres H. Soares, 1979. The following new combinations are proposed: Sodreana barbiellinii (Mello-Leitao, 1927), Sodreana hatschbachi (B. Soares & H. Soares, 1946), Sodreana inscripta (Mello-Leitao, 1939), Sodreana leprevosti (B. Soares & H. Soares, 1947b), Sodreana bicalcarata (Mello-Leitao, 1936). Sodreana granulata (Mello-Leitao, 1937) is revalidated from the synonymy of Sodreana sodreana Mello-Leitao, 1922. Three new species are described: Sodreana glaucoi from Ilhabela and Boraceia, Sao Paulo state; S. curupira from Parque Nacional da Serra dos Orgaos, Rio de Janeiro state, and S. caipora from Ubatuba, Sao Paulo state. Sodreaninae species are restricted to forested areas and most occur in the southern part of the coastal Atlantic rainforest, one species occurs in interior Atlantic rainforest. The biogeographical analysis (Brooks Parsimony Analysis) resulted in a single and fully resolved most parsimonious tree with three main: components: northern (Bahia and Serra do Espinhaco), southern (Santa Catarina, Parana, Serra do Mar of Sao Paulo), and central (Espirito Santo, Serra da Bocaina, southern state of Rio de Janeiro, Serra dos Orgaos, Serra da Mantiqueira, Serra do Mar of Sao Paulo).

Conventional and ultrastructural analyses of the chromosomes of Discocyrtus pectinifemur (Opiliones, Laniatores, Gonyleptidae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Cytogenetics of three Brazilian Goniosoma species: A new record for diploid number in Laniatores (Opiliones, Gonyleptidae, Goniosomatinae)

Currently, 60 species of harvestmen have been karyotyped and all of these are from the Nearctic and Palearctic regions. This work is the first cytogenetic report of three gonyleptid species of the suborder Laniatores: Goniosoina aff. badiuln, G. proxiinuni and G. spelaeum of the Neotropical region, from the southeastern region of Brazil. Conventional Giemsa stain chromosome preparations were obtained from embryonic cells and adult male testes. Embryo mitotic plates of G. aff. badium and G. proximum indicated 88 chromosomes, and mitotic spermatogonial plates of G. spelaeum males revealed intra- and interindividual variation of chromosome number, ranging from 92-109 chromosomes. In the three analyzed species, the mitotic chromosomes were meta- or subinetacentric with no obvious sex chromosomes being identified during mitosis. Prophase I spermatocytes of G. spelaeum also revealed intra- and interindividual bivalent number variation and furthermore indicated the presence of multivalence. The karyotypes of these three Goniosoina species exhibited the largest chromosome pair with a negative heteropycnosis in the distal region of the shortest arrn chromosomes of G. spelaeum submitted to silver impregnation evidenced this negative heteropycnotic region as nucleolus organizer region (NOR). These results, when compared with cytogenetic data of other Laniatores species from the Palearctic region, indicated that a new record for diploid chromosome number probably characterize the genus Goniosoma in the Neotropical region.

A comparative analysis of the chemical nature of defensive secretions of Gonyleptidae (Arachnida : Opiliones : Laniatores)

The arachnids of the order Opiliones (harvestmen) produce substances used in defense. In the present paper, we analyzed 22 species of Gonyleptidae to explore the use of defensive substances in taxonomy and evolutionary biology. Thirty-seven different compounds were detected, 18 of which were preliminarily identified. These compounds were mapped onto a phylogenetic tree showing the relationships within the Gonyleptidae. Data from Cosmetidae were used as an outgroup. Five ketones and six alkyl phenols were reported for the first time in harvestmen. (c) 2005 Elsevier Ltd. All rights reserved.

Maternal care in the Neotropical harvestman Liogonyleptoides tetracanthus (Opiliones: Gonyleptidae)

We describe post-ovipositional maternal care in Liogonyleptoides tetracanthus Mello-Leitao 1932 (Gonyleptinae) and experimentally evaluate the protective role of this behavior against egg predation under laboratory conditions. Females laid 138.8 eggs on average and remained close to the clutch during the entire day. Eggs hatched after 11-15 days and nymphs dispersed from maternal protection after one to two days. Most of the experimental clutches left unattended were entirely consumed by conspecifics in 2-3 days. There was no reduction in egg number in the clutches protected by females. Although biological data are scarce, there are cases of egg hiding, paternal and maternal care within the subfamily Gonyleptinae. This diversity of forms of parental care is unusual when compared to other gonyleptid subfamilies, and future systematic revisions of the polyphyletic Gonyleptinae should include parental care as a potential source of phylogenetic information.

Revision of Nanophareus, a mysterious harvestman genus from Chile, with descriptions of three new species (Opiliones: Laniatores: Gonyleptidae)

The Chilean genus Nanophareus Roewer, 1929 is revised and three new species are described: N. araucanus sp. nov. (type locality: Parque Nacional La Campana, Valparaiso, Chile); N. bipartitus sp. nov. (type locality: Parque Nacional La Campana, Valparaiso, Chile); N. bosqenublado sp. nov. (type locality: Parque Nacional Fray Jorge, Coquimbo, Chile). The type species, N. palpalis Roewer, 1929, is redescribed and a lectotype is designated. A cladistic analysis was performed using these three new species plus N. palpalis and 14 more laniatorid species, and a data matrix of 72 characters: Seven from the ocularium, 22 from the dorsal scutum, one from the venter, one from the chelicera, eight from the pedipalp, 24 from male legs, and nine from male genitalia. Two equally most parsimonious trees were found (L = 210; C.I. = 0.41; R.I. = 0.51). Nanophareus was recovered as nested within a paraphyletic subfamily Pachylinae. The genus Nanophareus was found to be monophyletic based on the following exclusive synapomorphies: An external row of enlarged tubercles inserted among small ones on lateral margin of the dorsal scutum (innapplicable in N. bosqenublado); the ventro-basal margin of pedipalpal tibia curved 90 degrees in lateral view; and retrolateral seta of the pedipalpal tibia with a socket apically bifid (socket and seta longer than pedipalpal tibia length).

Gregarious behavior of two species of Neotropical harvestmen (Arachnida: Opiliones: Gonyleptidae)

We present the first record and description of the gregarious behavior of the Neotropical harvestmen Serracutisoma proximum (Mello-Leitao 1922) and Serracutisoma spelaeum (Mello-Leitao 1933) (Opiliones: Gonyleptidae: Goniosomatinae) (DaSilva & Gnaspini 2010). We followed and described the pattern of these aggregations over a period of 17 months in a cave in southeastern Brazil. Individuals of the two species aggregated with both conspecifics and heterospecifics during the non-reproductive season (i.e., from October to March, the cool and dry season). Aggregations contained up to 81 individuals, usually with a female-biased adult sex ratio. Multispecific aggregations were usually composed mainly of representatives of one of the two species, suggesting that although these species also aggregate with heterospecifics, there is a preference for aggregating with conspecifics. This study provides novel information on the social behavior of harvestmen, specifically regarding the composition of multispecific aggregations.

Descriptions of Thereza murutinga sp. nov. and Pristocnemis caipira sp. nov., and new records of Caelopyginae (Opiliones: Laniatores: Gonyleptidae)

Two new species of Caelopyginae are described: Pristocnemis caipira sp. nov. from Parque Nacional da Serra da Canastra, Minas Gerais, Brazil, and Thereza murutinga sp. nov., from Parati, Rio de Janeiro, Brazil. Pristocnemis caipira sp. nov. is diagnosed by shallow grooves on the dorsal scutum, a lower number of tarsal segments, presence of a spine on the anal operculum, large, pointed tubercles on the lateral margins of the dorsal scutum, and an absence of white patches on the dorsal scutum. Thereza murutinga sp. nov. is diagnosed by the combination of the following characters: incomplete median longitudinal groove on area III and absent on area II, prosoma much smaller than opisthosoma, and a large white patch covering the prosoma and whole areas of the opisthosomal scutum. A cladistic analysis including the two new species was performed to investigate their relationship within the subfamily. We obtained 927 trees of 185 steps (CI=0.4, RI=0.74), which corroborated the generic assignments of the new species, but infrageneric relationships remained unsolved. In addition, male genitalia of Ampheres luteus (Giltay) and genera Pristocnemis and Thereza are redescribed, new distribution records for Caelopyginae species are presented, and biogeographical implications are discussed.

Behavioral analysis of the interaction between the spitting spider Scytodes globula (Araneae: Scytodidae) and the harvestman Discocyrtus invalidus (Opiliones: Gonyleptidae)

Spitting spiders (Scytodidae) have a distinct predatory strategy in which they eject a sticky secretion from their cheliceral fangs to immobilize prey. This behavior could potentially allow the spider not only to avoid defensive secretions but also to bite specific vulnerable spots of a potential prey such as a harvestman. We used an ethogram, a fluxogram and an experiment to analyze the interaction between the harvestman Discocyrtus invalidus Piza 1938 (Arachnida: Opiliones) and the syntopic spider Scytodes globula (Nicolet 1849) (Arachnida: Araneae). These spiders, while readily taking crickets as prey, seldom spat at and never bit the harvestmen, which apparently did not exude repugnatorial secretions. We therefore tested, by clogging the glands and using appropriate controls, whether non-visible amounts of secretions could cause the rejection, but the harvestmen were still refused. This is the first detailed and quantified description of an interaction between a spitting spider and a harvestman. The general conclusions are that S. globula avoids preying on D. invalidus, S. globula behaves differently when attacking harvestmen and crickets and the scent gland secretions of D. invalidus do not play a direct role in this predator-prey interaction.