Skeletomuscular anatomy of the harvestman Leiobunum aldrichi (Weed, 1893) (Arachnida: Opiliones: Palpatores) and its evolutionary significance

Skeletal muscles of the North American harvestman Leiobunum aldrichi are exhaustively surveyed and compared with other chelicerates to clarify the evolutionary morphology and phylogenetic relationships of arachnids. Representatives of 104 muscle groups are described and illustrated, and their possible functions are proposed. Comparisons of the feeding apparatus of L. aldrichi with that of other opilions, especially Sim (Cyphophthalmi) and Acromares (Laniatores), and two scorpion genera ( Centruroides, Pandinus ) indicate that the pharyngeal apparatus in L. aldrichi is derived and that its ability to accommodate large food particles is a secondary rather than primitive condition. Comparisons reveal several possible synapomorphies between Opiliones and Scorpiones suggesting that these orders may be sister groups. Apparently unique synapomorphies include an extrinsic cheliceral muscle that arises from the carapace and inserts on the second cheliceral article (deutomerite); an epistome divided into distal and proximal parts by a transverse sulcus; pharyngeal dilator muscles supported by a peripharyngeal skeleton formed by one dorsomedial and two ventrolateral epistomal processes, the latter also with muscular attachments to the endosternite; a specialized preoral chamber (stomodieca) derived from extensions (coxapophyses) of the coxae of the pedipalp and first two leg pairs; internal processes associated with the coxapophyses that serve, in part, as an attachment for muscles operating the coxa-trochanter joints, and lateral endosternal suspensor muscles that insert on the arthrodial membrane between the leg coxae. These are the first observations providing explicit support for an Opiliones-Scorpiones clade.     

A divergent Cardinium found in daddy long-legs (Arachnida: Opiliones)

Recent studies indicate that a newly described bacterial endosymbiont, Cardinium, is widespread in arthropods and induces different reproductive manipulations in hosts. In this study, we used a portion of the 16S rRNA gene of the Cardinium to screen 16 Opilionid species from the suborder Palptores. We found the incidence of Cardinium in these Opiliones was significantly higher than in other pooled arthropods (31.2% versus 7.2%, P = 0.007). Phylogenetic analyses using maximum parsimony (MP) and Bayesian analysis revealed two distinct clades in Opiliones. One is a divergent monophyletic clade with strong support that has so far not been found in other arthropods, and a second one contains Cardinium both from Opiliones and other arthropods. There is not complete concordance of the Cardinium strains with host phylogeny, suggesting some horizontal movement of the bacteria among Opiliones. Although the divergence in the sequenced 16S rRNA region between the Cardinium infecting Opiliones and Cardinium from other arthropods is greater than among Cardinium found in other arthropods, all are monophyletic with respect to the outgroup bacteria (endosymbionts of Acanthamoeba). Based on high pairwise genetic distances, deep branch, and a distinct phylogenetic grouping, we conclude that some Opiliones harbor a newly discovered Cardinium clade.     

Sexual Differences in the Behavior of the Harvestman Leiobunum vittatum (Opiliones, Sclerosomatidae) Towards Conspecific Cues

Preliminary observations of the harvestman Leiobunum vittatum found that individuals rub their bodies against the substrate, presenting the possibility of chemical marking. To determine whether or not L. vittatum individuals can detect substrate-borne chemical cues, we compared responses of L. vittatum males and females to substrate-borne male and female cues. We found that individuals of L. vittatum do respond to conspecific cues and that their responses are sex-specific. In response to substrate-borne conspecific cues, male L. vittatum spent more time, engaged in more scraping with their sensory legs I, and engaged in pedipalpal tapping more often in the presence versus absence of conspecific cues (male and female equally). Furthermore, in the presence of conspecific cues, males engaged in two behaviors never observed in females—(a) “fast approach” and (b) “jerking”, the latter of which was never observed in the presence of cricket cues. In contrast to males, females did not spend more time on conspecific cues, but did spend more time tapping their pedipalps in the presence of male vs female cues, suggesting an ability to distinguish between them. A final experiment explored the possibility that females could discriminate among males of varying histories of agonistic interactions based upon their chemical cues. We found no support for this hypothesis. Our results demonstrate that L. vitattum do respond to conspecific cues, and introduce the possibility that intraspecific communication may be mediated in part by chemical cues.     

Hemocytes of Leiobunum limbatum and two other species of harvestmen (Arachnida,Opiliones): Morphological classification and functional aspects

The hemocytes of Leiobunum limbatum, Mitopus morio, and Opilio ravennae number from about 8,000 (juveniles) to 41,000 (pregnant females) per microliter of hemolymph. Five different types of hemocytes occur in all three species and both sexes. According to their ultrastructural appearance and their similarities to other arthropod hemocytes these five types are designated as prohemocyte, plasmatocyte, granulocyte, coagulocyte, and spherulocyte. From the ultrastructural point of view the prohemocytes are interpreted as stem cells for plasmatocytes which on their part differentiate into granulocytes. Transitional stages which would indicate the origin of coagulocytes and spherulocytes could not be found. Granulocytes and spherulocytes are interpreted as being storage cells; coagulocytes burst when hemolymph is transferred to a microscopic slide. Plasmatocytes are involved in the removal of dead cells or cell fragments. Plasmatocytes are demonstrated as being able to phagocytize and digest bacteria.     

Respiratory system of arachnids II: morphology of the tracheal system of Leiobunum rotundum and Nemastoma lugubre (Arachnida, Opiliones)

The tracheal system of two species of harvestmen with different life styles was investigated: the long-legged, Leiobunum rotundum (Phalangioidea, Phalangiidae) and the short-legged Nemastoma lugubre (Troguloidea, Nemastomatidae). The morphology of the tracheae is very similar in both species: The branching pattern is basically asymmetric and dichotomous, and the tracheae taper between branching points. The tracheal diameters range from 160 to 0.3 mum in L. rotundum (mean body mass 25.3 mg), and from 70 to 0.5 mum in N. lugubre (mean body mass 3.8 mg). Ultrastructurally, the tracheal walls are similar to those of insects, consisting of an outer hypodermal layer and an inner cuticular layer with taenidial structures. Tracheae of all diameters have close contact with organs and muscles, indicating that diffusive gas exchange may take place through the walls of all tracheae. The finest tracheae end freely in the hemolymph or at the surface of organs and muscles. Exceptions are tracheal penetration of the central nerve mass in the prosoma in both species, and of the muscles in L. rotundum. The latter may correlate with the active perambulatory life style of L. rotundum.     

Neotropical harvestmen (Arachnida,Opiliones) use sexually dimorphic glands to spread chemicals in the environment

Sexually dimorphic glands have convergently appeared in animals and are often responsible for the production of pheromones. In the suborder Laniatores of the order Opiliones (Arachnida), glands of such type are widespread, but there is not a single paper on how they are used. Using Scanning Electron Microscopy and a behavioral approach, we describe glandular openings and how these glands are used, in the harvestmen Gryne perlata and Gryne coccinelloides (Cosmetidae). Males of these two species have glandular openings on the metatarsi of legs I and on the metatarsi IV. Males were shown rubbing the glands of the metatarsi I against their other legs, whereas glands on the metatarsi IV are gently touched on the substrate or rubbed either against other legs, or against the substrate. Not all behaviors were seen in both species.     

Screening pigment granule formation in Eumesosoma roeweri (Arachnida: Opiliones)

A unique pathway that utilizes endoplasmic reticulum (ER) networks is proposed for screening pigment granule formation in the retina of adult Eumesosoma roeweri. Smooth endoplasmic reticulum (SER) functions to transport pigment particles to the formative site. Each site is composed of concentric, interconnected rings of SER that are filled with dense-cored, spherical pigment particles. Formation of the screening pigment granule begins by the release of particles from the innermost rings of carrier SER. Continued release followed by fusion and condensation of the pigment particles results in the formation of a mature pigment granule.     

Biogeographical variation in the harvestman Mitopus morio (Opiliones, Arachnida)

The taxonomic status of three forms of the harvestman Mitopus morio in Britain have been examined. These have been identified by previous workers as a small, upland M, morio cinerascens , a larger, lowland M. morio morio , and a northern M. morio borealis. In this paper, the delimitation of trinomials has been found to be inappropriate for M. morio. since all three forms have been shown to lie at different points on a continuum of phenotypic variation within the species M. morio. It is suggested that this phenotypic variation is mediated by the variation in temperature regimes between different localities.     

Appearance of ocular electrical responses during development of cavernicolous and epigaeic opilionids (Arachnida)

The appearance of the electroretinogram was established during the development of two species of opilionids, an epigaeic species, Pachylus chilensis, and a cavernicolous one, Ischyropsalis luteipes. In nymphs and adults of the two species, electroretinograms of about the same amplitude can be recorded. During embryonic and larval development they are quite different. In Ischyropsalis luteipes, it is impossible to record an electrical response before hatching. In Pachylus chilensis on the other hand an initial inconstant and very small response occurs from the 50th hour of development of the retina and can always be recorded from the 72nd hour. In older embryos, the response has a progressively greater amplitude, but remains a simple negative wave. On the tenth day, when the first synaptic relays appear, the electroretinogram takes the shape of that of the adult. Given that transmission electronic microscopy cannot detect differences between the microvilli of these two species, it is tempting to compare Ischyropsalis luteipes lack of response with that of the mutant norp A of Drosophila melanogaster in which transmission electron microscopy shows no marked difference, whereas freeze-etching shows a very low number of intramembranous particles.     

Ultrastructure and function of the circulatory organs of Leiobunum limbatum and two other species of harvestmen (arachnida: Opiliones)

The ultrastructure of the circulatory organs of Leiobunum limbatum, Mitopus morio, and Rilaena triangularis (Arachnida: Opiliones) has been investigated. We studied the organization of the heart, the myocardial contractile elements, its tubular system, organelles, and cell-junctions. The epicardium exhibits a large number of longitudinally arranged microtubules and is attached to the myocardium by special membrane complexes. Because of these structures, the epicardium is interpreted as the elastic antagonist to the heart muscle. At both ends of the heart the epicardium continues into a nonbranching aorta. Consequently the ultrastructure of the aortal wall, containing numerous microtubules, is similar to the ultrastructure of the epicardium. At the posterior end of the heart is a muscular slit-valve, while at the anterior end is a lobe-valve that lacks contractile elements. The heart is innervated by a dorsal ganglion. It covers the entire length of the heart and contains neural and glial cells. Two different types of neurosecretory granules and both electron-dense and electron-lucent synaptic vesicles are described.     

Chemical and behavioral defenses of the neotropical harvestman Camarana flavipalpi (Arachnida: Opiliones)

The defensive secretion of the harvestman Camarana flavipalpi (Gonyleptidae) was chemically characterized, revealing the presence of a single, highly volatile component (2-methyl-5-ethyl-phenol). Individuals do not frequently use chemical defenses and rely mainly on thanatosis as defense. Two types of emission of secretions were recorded: a globule at the gland opening without liquid displacement and liquid displacement along the lateral margins of the dorsal scute with accumulation of fluid at the lateroposterior area of the body. The anterior opening of the ozopore is covered by an integumentary dome that faces the laterodorsal area of the scute. The lateral channel is well defined near the opening of the ozopore and along the lateral margin of the dorsal scute as rows of small granules. The ozopore morphology of C. flavipalpi is similar to closely related species and simplified when compared with other species that promptly release chemical secretions.     

Fissiphalliidae, a new family of South American laniatorean harvestmen (Arachnida: Opiliones)

Fissiphalliidae, a new family of the Opiliones suborder Laniatores, is described in the superfamily Gonyleptoidea, based on a new genus (Fissiphallius n. gen.) und 3 new species (F. sturmi n. sp., F. spinulatus n. sp., F. sympatricus n. sp.) from Colombia. Most emphasized are the male genitalic characters, unique in Opiliones: A honzontally split truncus which results in a long movable (glans) and an immovable finger (distal part of the truncus). Both hide the extremely long stylus with the seminal opening at its end. In an expanded state, the movable finger is bent dorsally by an erectile vesicle. It is shown that male genitalic characters in Opiliones display clear functional and constructional traits on the family level. Those characters, if used appropriately, will demonstrate that several families in gonyleptoid Laniatores are polyphyletic and should be divided into several taxa of the family level.     

Comparative Density of Hair Sensilla on the Legs of Cavernicolous and Epigean Harvestmen (Arachnida: Opiliones)

To allow an animal to behave appropriately, the location of sensorial structures is expected to be related to their function. As the different leg pairs of arachnids may have different functions (probing x supporting the body), one could expect them to have a different density of sensilla. Moreover, different regions of the same leg (dorsal, lateral, and ventral) would also be expected to have different densities of sensilla, according to the use of each region (e.g., the ventral part is often in contact with the substrate while the dorsal part is not). As cavernicolous animals are expected to be more sensitive than their epigean relatives, one could also expect a different density of sensilla when comparing cavernicolous and epigean animals. Using three epigean and three cavernicolous species of harvestmen (Arachnida, Opiliones), this study aimed at describing the morphology of hair sensilla on the legs and answering three questions: (1) Are there differences in the density of hair sensilla between the dorsal, lateral and ventral regions of each leg pair of the same individual? (2) Are there differences in the density of hair sensilla between the leg pairs of the same individual? (3) Are there differences in the density of hair sensilla when comparing the leg pairs of individuals of cavernicolous and non-cavernicolous species? The tarsi and metatarsi of all right legs of the six studied species were analyzed under a scanning electron microscope. The results (P < 0.05) showed that, in general: the ventral region of the tarsus was denser in sensilla trichodea than the lateral and dorsal regions, particularly on legs I and II; the density of sensilla chaetica did not differ on legs III and IV, but was greater on the dorsal region of legs I and II; the ventral part of legs I had the higher density of sensilla trichodea of the four pairs, whereas the second pair had the lower density; Holcobunus citrinus (Eupnoi) was the species with higher density of sensilla trichodea, on all legs; the cavernicolous species had a lower density of sensilla than the epigean species. The results are tentatively related to harvestmen behavior.     

Electron microscope study on the meiotic cycle of Acanthopachylus aculeatus (Arachnida; Opiliones)

Summary The primary spermatocytes of an Opilionid, Acanthopachylus aculeatus show (at least as a rule) two bodies which in longitudinal sections appear as integrated of up to 12 dark bands each about 300 Å wide, interlaced with one another by a curtain of thin filaments. The same bodies appear in transversal sections as formed by an hexagonal lattice the nodal points of which are the cross sections of the dark bands. The study of spermatocytes at early prophase demonstrated that at the inception these bodies are formed by association of several structures comparable to those observed in the axis of paired autosomes and called synaptinemal complexes.The findings in this species are compared with their similar in other species, particularly with species of Gryllidae.This investigation was supported by United States Public Health Service Research Grant GM 08337 from the Research Grants Branch, Division of Medical Sciences, and partly by Grant RF 61034 from The Rockefeller Foundation.