Interspecific variation in the microanatomy of cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

Scanning electron microscopy (SEM) is a useful tool for identifying interspecific variation in often overlooked structures that may represent useful sources for informative phylogenetic characters. In this study, we used SEM to compare the morphology of 12 cosmetid species from Central America, the Caribbean, and North America including multiple species for the genera Cynorta, Erginulus, and Paecilaema. To determine if microanatomical structures were unique to the cosmetid taxa under examination, we investigated the microanatomical structures of six additional species of gonyleptoidean harvestmen representing the families Agoristenidae, Cranaidae, Gonyleptidae, Manaosbiidae, and Stygnidae. Our results indicate that the shape of the ocularium (narrow, intermediate, or broad) did not vary within cosmetid genera, whereas the morphology of the rough pit glands on the eye mound varied considerably between species. Each cosmetid species had 10–20 rough pit glands on the ocularium whereas only the eye mounds of Avima intermedia (Agoristenidae) and Glysterus sp. (Gonyleptidae) had similar structures. With regards to the surface texture of the dorsal scutum, cosmetid harvestmen exhibited a rivulose‐microgranulate morphology (6 species), a microtuberculate‐rivulose‐microrgranulate morphology (4 species), or a microgranulate morphology (2 species). In contrast, each of the gonyleptoidean species exhibited a microgranulate pattern, with the exception of Stygnoplus clavotibialis, which had a rivulose‐microgranulate surface texture. For cosmetid harvestmen, we observed considerable interspecific variation in the shape and number of teeth on the fixed and moveable fingers of the male chelicerae. Similarly, we also observed interspecific variation in the distribution and shape of tubercles on the ventral and dorsal surfaces of the femur of the pedipalp. Overall, our results indicate that there are several microanatomical structures associated with the ocularium, dorsal scutum, male chelicera, and pedipalp that could represent informative phylogenetic characters in future taxonomic studies of cosmetid harvestmen. J. Morphol. 275:1386–1405, 2014. © 2014 Wiley Periodicals, Inc.     

Interspecific,ontogenetic, and sexual variation in ozopore morphology among cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

The ozopores of cosmetid harvestmen rest upon lateral projections of the carapace, have simple or highly reduced channels, and are partially obscured by enlarged dorsal processes associated with coxae I and II. Rather than use scent gland secretions to form a chemical shield on the dorsum, the cosmetid harvestman exhibits a unique defensive behavior known as “leg dabbing” in which the distal tip of tarsus I or II is dipped into fluid that accumulate at the base of coxa II and the droplet on the tarsus is pointed toward the predator. Relatively little is known about interspecific variation in ozopore morphology among cosmetid harvestmen. In this study, we used scanning electron microscopy to examine the ozopores of males and females of nine species as well as those of antepenultimate nymphs for two species. Among adults, we found differences between species in the shapes of the ozopores (round or subtriangular), the morphology of the dorsal and lateral channels (if present), and the relative size, shape and armature of the dorsal posterior process (dpp) of coxa I and the dorsal anterior process (dap) of coxa II. Our observations suggest that the morphology of dpp I and dap II could be sources for systematic characters in future phylogenetic studies of the Cosmetidae. We observed ontogenetic differences but relatively little intersexual variation in the morphology of the ozopore. The ozopores of nymphs are generally more oval than those of adults and the opening of the ozopore of the nymph is less obstructed, if at all, by the dorsal coxal processes of legs I–II. These morphological differences suggest that nymphs may use scent gland secretions in a manner different from that of adults.     

Ontogenetic variation in the sensory structures on the pedipalps of cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

In arachnids, pedipalps are highly variable appendages that may be used in feeding, courtship, defense, and agonistic encounters. In cosmetid harvestmen, adults have pedipalps that feature flattened femora, spoon‐shaped tibiae, and robust tarsal claws. In contrast, the pedipalps of nymphs are elongate with cylindrical podomeres and are adorned with delicate pretarsi. In this study, we used scanning electron microscopy to examine the distribution of cuticular structures (e.g., sensilla chaetica, pores) on the elements of the pedipalps of adults and nymphs of three species of cosmetid harvestmen. Our results indicate that there is considerable ontogenetic variation in the morphology of the trochanter, femur, patella, tibia, and tarsus. The pretarsus of the nymph has a ventral patch of setae that is absent from the adult tarsal claw. We observed this structure on all three cosmetid species as well as on the pedipalps of an additional seven morphospecies of nymphs collected in Belize and Costa Rica. This structure may represent a previously unrecognized autapomorphy for Cosmetidae. Examinations of the pedipalps of antepenultimate nymphs of additional gonyleptoidean harvestmen representing the families Ampycidae, Cranaidae, Manaosbiidae, and Stygnidae revealed the occurrence of unusual, plumose tarsal setae, but no setal patches on the tarsal claw.     

Interspecific variation in ovipositor morphology among Manaosbiid and Nomoclastid Harvestmen (Arachnida,Opiliones, Laniatores)

The external anatomy of the ovipositor has generally been overlooked as a source of informative characters in systematic studies of laniatorean harvestmen. In this study, we used scanning electron microscopy to examine the ovipositors of nine species representing the families Manaosbiidae (five species) and Nomoclastidae (four species). Similar to the ovipositor morphology of many gonyleptoidean families, the distal tips of the ovipositors of these harvestmen have four external lobes, with the margins most commonly adorned with 10 large peripheral setae. In manaosbiid and nomoclastid species, these peripheral setae have undivided bases, striated shafts and undivided distal tips. There are typically three setae on each anterior lobe and two setae on each posterior lobe. The medial setae on both anterior and posterior lobes insert into sockets that are slightly more dorsal. We observed small, surface denticles, usually associated with a pore, on the external surface of the lobes. There was interspecific and intraspecific variation in the number and shape of these surface denticles. The association of pores with denticles on the ovipositor appears to be a feature common to not only both families but is also a trait that has not been observed on ovipositors in other families of laniatorean harvestmen.     

Comparative study of the morphology of the gland opening area among Grassatores harvestmen (Arachnida,Opiliones, Laniatores)

Arachnids of the order Opiliones (harvestmen), which includes around 6000 specìes, 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.     

The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida,Opiliones)

Pedipalps are the most versatile appendages of arachnids. They can be equipped with spines (Amblypygi), chelae (Scorpiones), or adhesive pads (Solifugae), all of which are modifications to grasp and handle fast‐moving prey. Harvestmen (Opiliones) show a high diversity of pedipalpal morphologies. Some are obviously related to prey capture, like the enlargement and heavy spination of Laniatores pedipalps. Many Dyspnoi, by contrast, exhibit thin, thread‐like pedipalps that are covered with complex glandular setae (clavate setae). These extrude viscoelastic glue that is used to immobilize prey items. Comparable setae (plumose setae) have previously been found in representatives of both Eupnoi and Dyspnoi, yet comprehensive data on their distribution are lacking. This study examined the distribution and ultrastructure of glandular setae in harvestmen and related them to pedipalpal morphology. Pedipalpal and setal characters were analysed in a phylogenetic framework. We found that glandular setae are synapomorphic for and widespread in the Palpatores clade (Eupnoi plus Dyspnoi). Their occurrence correlates with pedipalp morphology and feeding habit. Remnants of arthropod cuticular structures or secretions, frequently found attached to glandular setae, and behavioural observations, underlined the importance of the setae for capturing and securing prey. We hypothesize that glandular setae evolved as an adaptation to capture small and agile prey, which are hard to catch with a capture basket. Details of ultrastructure indicate that the setae are derived sensilla chaetica, with both a secretory and sensory function. Derived ultrastructural characters of the glandular setae, such as slit‐like channel openings and a globular arrangement of the microtrichia, may increase their effectiveness. The functional role of further pedipalpal modifications, such as apophyses, stalked and hyperbendable joints, and curved segments, as well as sexual dimorphism and ontogenetic polymorphism, are discussed. Some implications of the results obtained for the taxonomic treatment of Phalangiidae are also discussed. These results shed new light on the biology and evolutionary history of this fascinating group of arthropods.     

New systematic assignments in Gonyleptoidea (Arachnida, Opiliones, Laniatores)

As part of an ongoing revision of the family Gonyleptidae, we have identified many species that are synonyms of previously described species or misplaced in this family. This article summarizes these findings, adding previously unavailable information or correcting imprecise observations to justify the presented taxonomic changes. The following new familial or subfamilial assignments are proposed: Nemastygnus Roewer, 1929 and Taulisa Roewer, 1956 are transferred to Agoristenidae, Agoristeninae; Napostygnus Roewer, 1929 to Cranaidae; Ceropachylinus peruvianus Roewer, 1956 and Pirunipygus Roewer, 1936 are transferred to Gonyleptidae, Ampycinae; Gyndesops Roewer, 1943, Haversia Roewer, 1913 and Oxapampeus Roewer, 1963 are transferred to Gonyleptidae, Pachylinae. The following generic synonymies are proposed for the family Gonyleptidae: Acanthogonyleptes Mello-Leitão, 1922 = Centroleptes Roewer, 1943; Acrographinotus Roewer, 1929 = Unduavius Roewer, 1929; Gonyleptes Kirby, 1819 = Collonychium Bertkau, 1880; Mischonyx Bertkau, 1880 = Eugonyleptes Roewer, 1913 and Gonazula Roewer, 1930; Parampheres Roewer, 1913 = Metapachyloides Roewer, 1917; Pseudopucrolia Roewer, 1912 = Meteusarcus Roewer, 1913; Haversia Roewer, 1913 = Hoggellula Roewer, 1930. The following specific synonymies are proposed for the family Gonyleptidae: Acanthogonyleptes singularis (Mello-Leitão, 1935) = Centroleptes flavus Roewer, 1943, syn. n.; Geraeocormobius sylvarum Holmberg, 1887 = Discocyrtus serrifemur Roewer, 1943, syn. n.; Gonyleptellus bimaculatus (Sørensen, 1884) = Gonyleptes cancellatus Roewer,1917, syn. n.; Gonyleptes atrus Mello-Leitão, 1923 = Weyhia brieni Giltay, 1928, syn. n.; Gonyleptes fragilis Mello-Leitão, 1923 = Gonyleptes banana Kury, 2003, syn. n.; Gonyleptes horridus Kirby, 1819 = Collonychium bicuspidatum Bertkau, 1880, syn. n., Gonyleptes borgmeyeri Mello-Leitão, 1932, syn. n., Gonyleptes curvicornis Mello-Leitão, 1932, syn. n., Metagonyleptes hamatus Roewer, 1913, syn. n. and Paragonyleptes simoni Roewer, 1930, syn. n.; Gonyleptes pustulatus Sørensen, 1884 = Gonyleptes guttatus Roewer, 1917, syn. n.; Haversia defensa (Butler, 1876) = Sadocus vallentini Hogg, 1913, syn. n.; Liogonyleptoides minensis (Piza, 1946) = Currala bahiensis Soares, 1972, syn. n.; Megapachylus grandis Roewer, 1913 = Metapachyloides almeidai Soares & Soares, 1946, syn. n.; Mischonyx cuspidatus (Roewer, 1913) = Gonazula gibbosa Roewer, 1930 syn. n.; Mischonyx scaber (Kirby, 1819) = Xundarava holacantha Mello-Leitão, 1927, syn. n.; Parampheres tibialis Roewer, 1917 = Metapachyloides rugosus Roewer, 1917, syn. n.; Parapachyloides uncinatus (Sørensen, 1879) = Goyazella armata Mello-Leitão, 1931, syn. n.; Pseudopucrolia mutica (Perty, 1833) = Meteusarcus armatus Roewer, 1913, syn. n. The following new combinations are proposed: Acrographinotus ornatus (Roewer, 1929), comb. n. (ex Unduavius); Gonyleptellus bimaculatus (Sørensen, 1884),comb. n. (ex Gonyleptes);Gonyleptes perlatus (Mello-Leitão, 1935), comb. n. (exMoojenia);Mischonyx scaber (Kirby, 1819), comb. n. (ex Gonyleptes); and Neopachyloides peruvianus (Roewer, 1956), comb. n. (ex Ceropachylus). The following species of Gonyleptidae, Gonyleptinae are revalidated: Gonyleptes atrus Mello-Leitão, 1923 and Gonyleptes curvicornis (Roewer, 1913).     

Two new harvestmen species (Arachnida: Opiliones) from the Caucasus

Two new harvestmen species of the family Phalangiidae, Rilaena caucasica sp. n. and Rilaena silhavyi sp. n. are diagnosed, illustrated, and described from the Caucasus region. Comparative illustration of the related Rilaena anatolica (Roewer, 1956), R. atrolutea (Roewer, 1915) and R. kelbajarica Snegovaya &; Pkhakadze, 2014 Snegovaya, N. Y., &; Pkhakadze, V. D. (2014): New species of the genus Rilaena (Opiliones, Phalangiidae) from the mount Gyamish, Azerbaijan. Vestnik zoologii, 48, 313–318. doi: 10.2478/vzoo-2014-0037[Crossref] , [Google Scholar] are given.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:7B29FD94-45A2-4E32-A41E-3276E016410B     

Four new Laniatorean harvestmen (Arachnida: Opiliones) from mid-Cretaceous Burmese amber

Four new laniatorean harvestmen specimens (Arachnida: Opiliones: Laniatores) are described from the mid-Cretaceous (upper Albian–lower Cenomanian) Burmese amber of Northern Myanmar. One is placed as Insidiatores indet., but is not formally named as it is probably immature. Burmalomanius circularis n. gen. n. sp. and Petroburma tarsomeria n. gen. n. sp. represent the first fossil records of the extant families Podoctidae and Petrobunidae respectively. Finally, Mesodibunus tourinhoae n. gen. n. sp. belongs to Epedanidae, a family previously recorded from Burmese amber. These new records bring the total number of Burmese amber laniatorean species to ten, and the total number of fossil laniatoreans to fifteen. The new finds offer additional calibration points for the Laniatores tree of life and are consistent with the hypothesis that the modern Laniatores fauna of Southeast Asia may have had Gondwanan, as opposed to a Laurasian, origins.     

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.     

Metabolic rates in harvestmen (Arachnida, Opiliones): the influence of running activity

Abstract. Metabolic rates of adult Lophopilio palpinalis (Herbst, 1799) (Arachnida, Opiliones, Phalangioidea) and Paranemastoma quadripunctatum (Perty, 1833) (Arachnida, Opiliones, Troguloidea) are measured during rest and activity. Carbon dioxide release during rest is continuous in both species. Mean values at 20 °C are 4.2 µL min⁻¹ g⁻¹ for the males of P. quadripunctatum, 4.1 µL min⁻¹ g⁻¹ for the males of L. palpinalis and 4.7 µL min⁻¹ g⁻¹ for the females of L. palpinalis, thus being significantly higher in the egg-producing females. In L. palpinalis, respiratory quotient at rest is 0.84. Spontaneous walking activity with speeds of 15–30 cm min⁻¹ raises the metabolic rate by up to three-fold in both species. Lophopilio palpinalis is made to undertake constant running on a treadmill with speeds of 60, 72 and 96 cm min⁻¹. Enforced activity causes the animals to raise their metabolic rates by up to five-fold above resting rates. Animals reach a steady state of CO₂ release on the treadmill and show a fast t_1/2 on-response, indicating aerobic exercise. The minimum cost of locomotion is determined to be 2.5 × 10⁻³ J cm⁻¹ g⁻¹, thus fitting the predicted values for terrestrial locomotion.     

Morphological changes during postembryonic development in two species of neotropical harvestmen (Opiliones,Laniatores, Cranaidae)

Morphological changes during postembryonic development in the Cranaidae are described on the basis of the examination of an incomplete series of larvae, nymphs, and adults of Phareicranaus calcariferus and Santinezia serratotibialis. The life histories of these species are hypothesized to consist of six nymphal stages, featuring the appearance of secondary male sexual characteristics in the antepenultimate nymph (N5). Color and body shape change dramatically during development. Growth rates for nymphs based upon leg measurements were similar for both species. In S. serratotibialis, the greatest increase in leg size occurred from larva to 1st nymph. The tarsomeres of legs I–IV varied by 1–2 segments per leg for each nymph stage, with the number of tarsal segments increased by 1–2 segments at each stage. Adults had nearly twice as many tarsomeres on leg II than other legs. Ontogenetic changes were observed in the armature of the proximal cheliceral segment, ocularium, pedipalp, opisthosoma, distitarsus III and IV, and leg IV. Morphological changes in postembryonic development in cranaid harvestmen are similar to those reported for other Laniatores. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Ultrastructure of the sexually dimorphic tarsal glands and tegumental glands in gonyleptoid harvestmen (Opiliones,Laniatores)

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. J. Morphol. 274:1203–1215, 2013. © 2013 Wiley Periodicals, Inc.     

Maternal care, defensive behavior, and sociality in neotropical Goniosoma harvestmen (Arachnida, Opiliones)

Summary: Goniosoma includes large and conspicuous species of harvestmen that occur exclusively in the Brazilian Atlantic forest. This paper reports on the parental activities, defensive behavior, and gregariousness of five species of Goniosoma, and also summarizes the published biological data for the genus. The behavioral patterns within the genus are discussed and a hypothesis for the emergence of gregariousness in the group is presented. Several Goniosoma live inside caves, mainly as trogloxenes. Although species of the genus comprise only 5% of the harvestmen fauna in Brazil, they account for 20.6% of all harvestmen species recorded in Brazilian caves. In addition to several morphological and physiological features that may favor the occupation of caves, species of Goniosoma also present subsocial behavior, which may confer a special advantage in this particular environment. Female protection is crucial for egg survival since predators may consume entire batches in a single night. Most species of Goniosoma form dense diurnal aggregations ranging from three to 200 individuals (mainly subadults and adults of both sexes). In a hypothetical scenario for the evolution of sociality in the Goniosoma, physiological constraints acting on individual harvestmen would lead to a behavioral response for the selection of sites with appropriate microclimatic conditions. The lack of cannibalism among adults and subadults may favor tolerance towards conspecifics, and may represent a pre-adaptation to gregariousness. The derived functions of gregarious behavior may include strengthening of the defensive signal through the collective release of a repugnant secretion, the dilution effect, and promptness in fleeing a predator attack as a consequence of the additional alarm role of scent gland secretions. These defensive functions may overcome the costs of group living and may be responsible for the maintenance of gregariousness in Goniosoma.     

Evolution of hyperflexible joints in sticky prey capture appendages of harvestmen (Arachnida,Opiliones)

The rigid leg segments of arthropods are flexibly connected by joints, which usually consist of two ball-and-bowl hinges, permitting a uniaxial pivoting up to 140°. Here, we report the occurrence of hyperflexible joints (range of movements?=?160–200°) in the pedipalps (second pair of appendages) of some harvestmen (Sabaconidae and Nemastomatidae), representing some of the most flexible leg joints among arthropods. Hyperflexion is achieved by a reduction of hinges and a strong constriction of the joint region. We demonstrate that hyperflexion occurs during prey capture and is used to clamp appendages of the prey, in addition to attachment by glue secreted by specialized setae. By means of high-speed video recordings, we found that in the Sabaconidae the tibiotarsal joint of the pedipalp can flex extremely rapidly (<5 ms), limiting prey escape. This is the fastest reported predatory strike in arachnids and caused both by leverage and a click mechanism. By comparative analysis of different related taxa, we retraced joint evolution and found that hyperflexion has independently evolved in Sabaconidae and Nemastomatidae, with totally different joint kinematics. We hypothesize that (rapid) hyperflexion evolved to enhance the efficiency of the pedipalp as a means of prey capture, because in springtails detachable scales limit the action of the sticky secretion of pedipalpal setae.     

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.