Sexually Dimorphic Tegumental Gland Openings in Laniatores (Arachnida, Opiliones), With New Data on 23 Species

Sexually dimorphic glands often release sexual pheromones both in vertebrates and invertebrates. Species of Laniatores (Arachnida, Opiliones) seem to depend on chemical communication but few studies have addressed this topic. In this study, we review the literature for the Phalangida and present new data for 23 species of Laniatores. In 16 taxa, we found previously undescribed sexually dimorphic glandular openings on the femur, patella, metatarsus, and tarsus of legs I and metatarsus of legs III and IV For the other species, we provide scanning electron micrographs of previously undescribed sexually dimorphic setae and pegs located on swollen regions of the legs. We also list additional species in which males have swollen regions on the legs, including the tibia, metatarsus, and tarsus of legs I, trochanter and tibia of legs II, femur, metatarsus, and tarsus of legs III, and metatarsus and tarsus of legs IV. The function and biological role of the secretions released by these glands are discussed. J. Morphol. 271:641-653, 2010. (C) 2009 Wiley-Liss, Inc.

New familial assignments for three species of Neotropical harvestmen based on cladistic analysis (Arachnida: Opiliones: Laniatores)

We herein propose transfer of the two monotypic genera Globibunus (type-species G. rubrofemoratus Roewer, 1912) and Rivetinus (type-species R. minutus Roewer, 1919) to the subfamily Zamorinae (Agoristenidae) and Ramonus (type-species R. conifrons Roewer, 1956), previous placed in Agoristenidae, to the Prostygninae (Cranaidae) based on several characteristics of male genitalia. These transfers are corroborated by a cladistic analysis, which also recovered the three currently recognized agoristenid subfamilies.

First record of Stygnidae for the state of Espírito Santo and description of a new Protimesius (Arachnida: Opiliones: Laniatores)

Protimesius osvaldoi sp. nov. is described from the Reserva Biológica de Sooretama, state of Espírito Santo, southeastern Brazil, being the first record of Stygnidae from this State and the southernmost record of the family in the Brazilian Atlantic Forest (hitherto, the family was recorded down to Bahia only), extending in 210 km south of the previously known distribution. This is a large species, with armature of leg IV very reduced and penial morphology differing from the closest counterparts mainly in the ventral plate, which recedes deeply at the lateral borders and has the distal margin curved ventrally and by the presence of two small intermediate setae. Protimesius Roewer, 1913 consisted hitherto of 17 species, recorded from northern/northeastern Brazil and Amazonia of adjacent countries. A key is given for the 17 species of Protimesius for which males are known.

Systematic review and cladistic analysis of the genus Eusarcus Perty 1833 (Arachnida, Opiliones, Gonyleptidae)

Eusarcus Perty 1833 is one of the oldest described genera of Pachylinae, comprising 36 species distributed from northeastern to southern Brazil (including the central west region), northeastern Argentina, eastern Paraguay and Uruguay. The genus is reviewed and a new classification is proposed based on a cladistic analysis. A cladistic analysis was performed with the 34 valid species of Eusarcus and 11 species belonging to certain Gonyleptidae subfamilies. The data matrix has 67 characters: 14 from dorsal scutum and pedipalp, 38 from male legs and 15 from male genitalia. Two equally parsimonious trees were found (L=319; C. I.=0.26, R. I.=0.61). Pygophalangodus gemignanii uruguayensis Ringuelet 1955a and Pygophalangodus gemignanii gemignanii Mello-Leitao 1931b are here elevated to the category of species, and the following new combinations are proposed: E. catharinensis (Mello-Leitao 1927); E. berlae (Mello-Leitao 1932); E. gemignanii (Mello-Leitao 1931b); E. signatus(Roewer 1949); E. sooretamae (Soares & Soares 1946a); E. uruguayensis (Ringuelet 1955a). The following generic synonymies are proposed: Eusarcus Perty 1833 (type species E. armatus Perty 1833) = Metagraphinotus Mello-Leitao 1927 (type species M. catharinensis Mello-Leitao 1927), Pareusarcus Roewer 1929 (type species P. corniculatus Roewer 1929), Pygophalangodus Mello-Leitao 1931b (type species P. gemignanii-gemignanii Mello-Leitao 1931b) and Antetriceras Roewer 1949 (type species A. signatus Roewer 1949). The following specific synonymies are proposed: Eusarcus hastatus Sorensen 1884 = Pucrolioides argentina Roewer 1913, E. guimaraensi H. Soares 1945, Jacarepaguana pectinifemur Piza 1943, Canestrinia canalsi Mello-Leitao 1931a, and E. maquinensis H. Soares 1966b; E. armatus Perty 1833 = E. curvispinosus Mello-Leitao 1923b, and Enantiocentron montis Mello-Leitao 1936; Eusarcus catharinensis (Mello-Leitao 1927) = E. antoninae Mello-Leitao 1936, E. perpusillus Mello-Leitao 1945, E. tripos Mello-Leitao 1940, and Metagraphinotus trochanterspinosus Soares & Soares 1947b; E. nigrimaculatus Mello-Leitao 1924 = Pareusarcus centromelos Mello-Leitao 1935a, E. furcatus Roewer 1929, Orguesia armata Roewer 1913, and Pareusarcus corniculatus Roewer 1929; E. oxyacanthus Kollar in Koch 1839a = Enantiocentron doriphorus Mello-Leitao 1932, and E. spinimanu Mello-Leitao 1932; E. pusillus Sorensen 1884 = E. vervloeti B. Soares 1944c; E. berlae Mello-Leitao 1932 = Metagraphinotus arlei Mello-Leitao 1935a. Metapucrolia armata (Sorensen 1895) is revalidated, transferred to Eusarcus and considered as a species inquirenda. A new name, Eusarcus metapucrolia is proposed for this species to avoid homonymy with the type species of Eusarcus, E. armatus Perty 1833. Eusarcus aberrans Mello-Leitao 1939a is considered as a species inquirenda. The male of E. teresincola Soares & Soares 1946a is described. Female of the following species are described: E. bifidus Roewer 1929; E. dubius B. Soares 1943b; E. insperatus B. Soares 1944a; E. schubarti Soares & Soares 1946a; E. sooretamae (Soares & Soares 1946a). The following new species are described from Brazil: E. acrophthalmus (type locality: Bahia, Ilheus, Parataquice); E. alpinus (Rio de Janeiro, Santa Maria Madalena, Parque Estadual do Desengano); E. caparaoensis (Minas Gerais, Alto Caparao, Parque Nacional do Caparao); E. cavernicola (Goias, Sao Domingos, Parque Estadual de Terra Ronca, Lapa da Angelica); E. didactylus (Rio de Janeiro, Teresopolis, Parque Nacional Serra dos Orgaos); E. garibaldiae (Santa Catarina, Itajai); E. geometricus (Rio de Janeiro, Teresopolis, Parque Nacional Serra dos Orgaos); E. manero (Rio de Janeiro, Marica, Itaipuacu); E. matogrossensis (Mato Grosso, Chapada dos Guimaraes); E. mirabilis (Minas Gerais, Marlieria, Parque Estadual Rio Doce); E. sergipanus (Sergipe, Itabaiana, Parque Nacional de Itabaiana) and E. tripectinatus (Minas Gerais, Rio Preto). The holotype of E. curvispinosus is proposed as the neotype of E. armatus Perty 1833, the type material of which has been lost. Lectotypes for the following species were designated: E. aduncus; E. hastatus; E. oxyacanthus.

6-Alkyl-3,4-dihydro-2H-pyrans: Chemical Secretion Compounds in Neotropical Harvestmen

The defensive, secretions, of five neotropical) species of harvestmen, (Opiliones: Gonyleptidae) from the Brazilian Atlantic Forest were analyzed and chemically characterized by GC-MS and NMR Methods. Three of the species, Cobania picea, Roweria virescens, and Serracutisoma proximum, secrete a mixture of 2,3-dimethyl-1,4-benzoquinone and 2-ethyl-3methyl-1,4,4-benzoquinone. The secretions produced,by the Other two species Iporangaia pustulosa and Neosadocus maximus, contain 1-hepten-3-one, 5-methyl-1-hexen-3-one, and 1-(6-butyl-3,4-dihydro-2H-pyran-2-yl)pentanone. (1)as major components, as well as,2,3-dimethyl-1.,4-benzoquinone and 2-ethyl-3 methyl-1,4-benzoquinone as minor,constituents. The. dihydropyran 1-(6-butyl-3,4-dihydro-2H-pyran-2-yl)pentanone (1) is a new natural product, composed of two 1-hepten-3-one, subunits formally linked in a hetero-Diels-Alder reaction. The natural product was proven to be racemic, and its biogenetic origin is discussed.

Efficiency of uniparental male and female care against egg predators in two closely related syntopic harvestmen

Although the benefits of maternal care have been investigated in many species, the caring role of males in species with exclusive paternal care has received less attention. We experimentally quantified the protective role of paternal care in the harvestman Iporangaia pustulosa. Additionally, we compared the effectiveness of paternal care against predation in this species with a syntopic harvestman with maternal care, Acutisoma proximum. We demonstrated that nearly one-third of the unprotected Iporangaia clutches disappeared entirely in 12 days, while the other two-thirds suffered a mean reduction of 55% in egg number. Conversely, 50% of the control clutches did not suffer any reduction, and only one was entirely consumed by predators. We also demonstrated that the mucus coat that covers Iporangaia clutches has an important deterrent role against predation by conspecifics: 58.3% of the clutches without mucus were attacked and three of them were entirely consumed, whereas only three clutches with mucus were attacked, suffering a reduction of up to three eggs. Iporangaia males were as efficient as Acutisoma females in protecting eggs. However, unattended Acutisoma eggs were attacked 20% more frequently than unattended Iporangaia eggs. Unattended Iporangaia eggs are protected by a mucus coat that prevents or decreases predation rate, whereas Acutisoma eggs are more susceptible to predation, probably because they lack this mucus coat. Thus, besides the fact that Iporangaia males efficiently protect the offspring against egg predators, females also contribute to egg protection by providing a mucus coat that deters egg predators. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Sexually dimorphic legs in a neotropical harvestman (Arachnida, Opiliones): Ornament or weapon?

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 mates 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. (C) 2008 Elsevier B.V. All rights reserved.

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.

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.

First record of Stygnidae for the state of Espirito Santo and description of a new Protimesius (Arachnida : Opiliones : Laniatores)

Protimesius osvaldoi sp. nov. is described from the Reserva Biologica de Sooretama, state of Espirito Santo, Southeastern Brazil, being the First record of Stygnidae from this State and the southernmost record of the family in the Brazilian Atlantic Forest (hitherto, the family was recorded down to Bahia only), extending in 210 km south of the previously known distribution. This is a large species, with armature of leg IV very reduced and penial morphology differing from the closest counterparts mainly in the ventral plate, which recedes deeply at the lateral borders and has the distal margin curved ventrally and by the presence of two small intermediate setae. Protimesius Roewer, 1913 consisted hitherto of 17 species, recorded from northern/northeastern Brazil and Amazonia of adjacent Countries. A key is given for the 17 species of Protimesius for which males are known.

Litter size, effects of maternal body size, and date of birth in South American scorpions (Arachnida: Scorpiones)

We present new data on litter size and date of birth (month) for 21 South American scorpions species. We provide data for one katoikogenic species, the liochelid Opisthacanthus cayaporum Vellard, 1932 (offspring = 3; birth month: Jan); and for several apoikogenic species, such as the bothriurids Bothriurus araguayae Vellard, 1934 (53; Sep), B. rochensis San Martín, 1965 (22-28; Jan, Aug); the buthids Ananteris balzanii Thorell, 1891 (10-34; Jan-Mar), Physoctonus debilis (Koch, 1840) (2; Sep), Rhopalurus amazonicus Lourenço, 1986 (19; Nov), R. lacrau Lourenço & Pinto-da-Rocha, 1997 (30; Dec), R. laticauda Thorell, 1876 (41; Nov), R. rochai Borelli, 1910 (11-47; Dec-Jan, Mar-Apr), Tityus bahiensis (Perty, 1833) (4-23; Oct-Mar), T. clathratus Koch, 1844 (8-18; Nov-Jan), T. costatus (Karsch, 1879) (21-25; Jan, Apr), T. kuryi Lourenço, 1997 (4-16; Mar), T. mattogrossensis Borelli, 1901(8-9; May), T. obscurus (Gervais, 1843) (16-31; Jan-Feb, May, Jul), T. serrulatus Lutz & Mello, 1922 (8-36; Dec, Feb-Apr), T. silvestris Pocock, 1897 (5-14; Dec-Jan, Apr), T. stigmurus (Thorell, 1876) (10-18; Nov, Jan, Mar), Tityus sp. 1 (T. clathratus group - 7-12; Feb-Apr), Tityus sp. 2 (T. bahiensis group - 2; Mar); and the chactid Brotheas sp. (8-21; Jan, Apr). We observed multiple broods: R. lacrau (offspring in the 2nd brood = 27), T. kuryi (6-16), T. obscurus (2-32), T. silvestris (8), T. stigmurus (4-9), T. bahiensis (offspring in the 2nd brood = 2-18; 3rd = 1), and T. costatus (2nd brood = 18; 3rd = 4). We found statistically significant positive correlation between female size and litter size for T. bahiensis and T. silvestris, and nonsignificant correlation for T. serrulatus.

A scanning electron microscopic survey of the cuticle in Cyphophthalmi (Arachnida, Opiliones) with the description of novel sensory and glandular structures

The cuticular surfaces of Cyphophthalmi (Opiliones) were studied in detail, covering a wide range of their taxonomic diversity. Previously unknown structures are described, including a sexually dimorphic row of spines and glandular openings on leg I of Fangensis cavernarum. Scanning electron micrographs of the prosomal paired hairs and the subapical process are provided for the first time. Evidence for the multi-pored nature of the shaft of solenidia as well as the hollowed nature and absence of wall pores of sensilla chaetica are also shown for the first time using scanning electron microscopy. The prosomal paired hairs may constitute a novel autapomorphy for Cyphophthalmi, as they are absent in all studied members of the other species of Opiliones. Finally, the variation in shape of some of the structures examined may be of great taxonomic value.

Harvest-ironman: heavy armature, and not its defensive secretions, protects a harvestman against a spider

Natural selection has caused prey species to evolve distinct defensive mechanisms. One of such mechanisms was the evolution of noxious or distasteful chemicals, which have appeared independently in a number of vertebrates and invertebrates. In detailed analyses of arthropod behaviour, scent gland secretions have consistently been shown to be responsible for repelling specific predators. Because using such chemicals is costly, animals with alternative cheaper defences are expected not to release such secretions when alternative options exist. In this study, we sought to determine the defensive mechanisms of the harvestman Discocyrtus invalidus, a heavy bodied species that bears a pair of repugnatorial glands. The spider Enoploctenus cyclothorax was used as the predator, and the cricket Gryllus sp. was used as a control. In a first set of experiments, the harvestmen were preyed upon significantly less than the crickets. In two other experiments, we found that harvestmen did not use their scent gland secretions to deter the predator. Moreover, results of a fourth experiment revealed that these spiders are not repelled by defensive secretions. Discocyrtus invalidus has a thick cuticle on the entire body: scanning electron micrographs revealed that only the mouth, the articulations of appendages and the tips of the legs are not covered by a hard integument. In a fifth experiment, we found that these spiders had difficulty piercing the harvestmen body. This is the first experimental evidence that a chemically defended arachnid does not use its scent gland secretions to repel a much larger predator but instead relies on its heavily built body. (c) 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Alternative mating tactics in dimorphic males of the harvestman Longiperna concolor (Arachnida: Opiliones)

Intense male-male competition for females may drive the evolution of male morphological dimorphism, which is frequently associated with alternative mating tactics. Using modern techniques for the detection of discontinuous allometries, we describe male dimorphism in the Neotropical harvestman Longiperna concolor, the males of which use their elongated, sexually dimorphic legs IV in fights for the possession of territories where females lay eggs. We also tested three predictions related to the existence of alternative mating tactics: (1) if individuals with relatively longer legs IV (majors) are more likely to monopolize access to reproductive resources, they are expected to remain close to stable groups of females more than individuals with relatively shorter legs IV (minors) do; (2) if minors achieve fertilization by moving between territories, they are expected to be less faithful to specific sites; and (3) majors should be observed in aggressive interactions more often. We individually marked all the individuals from a population of Longiperna during the reproductive season and recorded the location of each sighting for males and females as well as the identity of males involved in fights. Majors were more likely to have harems, and large majors were even more likely to do so. Majors were more philopatric and all males involved in fights belonged to this morph. These results strongly suggest that the mating tactic of the majors is based on resource defense whereas that of the minors probably relies on sneaking into the territories of the majors and furtively copulating with females.

Mating system and exclusive postzygotic paternal care in a Neotropical harvestman (Arachnida: Opiliones)

This study tests predictions of the hypothesis of evolution of paternal care via sexual selection by using the Neotropical harvestman Pseudopucrolia sp. as the model organism. Females use natural cavities in roadside banks as nesting sites, which are defended by males against other males. Females leave the nests after oviposition, and all postzygotic parental care is accomplished by males, which protect the eggs and nymphs from predators. We provided artificial mud nests to individuals in the laboratory and conducted observations on the reproduction of the species. Male reproductive success was directly related to nest ownership time: the longer a male held a nest, the higher his chances of obtaining copulations. All males that succeeded in mating and obtaining one clutch eventually mated with additional females that added eggs to the clutch. Thus, desirable males were not limited to monogamy by paternal care. Experimental manipulations demonstrated that guarding males were more attractive to females than were nonguarding males and also that males guarded unrelated eggs. Finally, we found that females and nonguarding males spent more time foraging than guarding males. We use our data to contrast hypotheses on the origin and maintenance of paternal care and to provide a critical assessment of the hypothesis of the evolution of paternal care via sexual selection. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.