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Relatively little is known about intraspecific variation in the use of chemical defenses by cosmetid harvestmen. In this study, we investigated sexual and ontogenetic variation in the emission of secretions from the scent glands and the morphology of the ozopores of Erginulus clavotibialis, a Neotropical species from Belize. Most individuals (> 94%) did not secrete when seized by leg IV, however, when the body was subsequently grasped firmly with forceps, 76% of the harvestmen (n = 96) responded by releasing enteric fluid (clear, no odor) or secretions from the scent glands (reddish-brown, distinct odor), most commonly in the form of globules near the ozopores. Adults and penultimate nymphs rarely (less than 5% of all trials) exhibited leg dabbing, an unusual behavior generally associated with cosmetid harvestmen. Males released enteric fluid (without scent gland secretions) significantly more than females or nymphs. Nymphs emitted secretions from scent glands at a significantly greater frequency than males or females. Using SEM, we examined the ozopores and the dorsal processes of coxae I and II and found that the morphology of these structures varies between adults and nymphs but not between males and females. The openings of the ozopores of nymphs are circular and are not obscured by the dorsal posterior process of coxa I and the dorsal anterior process of coxa II. In contrast, adults have subtriangular ozopores that are difficult to observe without dissection because they are partially blocked by the dorsal processes of coxae I and II.

The harvestmen family Cosmetidae contains more than 700 described species distributed from Argentina to the USA. The monophyly of the family remains undisputed; however, the majority of cosmetid genera are poorly defined and have not been revised using modern systematic approaches. This can be partially attributed to the paucity of information regarding genitalic morphology for most cosmetid species. A better understanding of the diversity of genitalic forms in this family will facilitate more in-depth systematic studies and taxonomic revisions within the family. In this study, we describe two distinct forms of genitalia exhibited by members of the genus Erginulus, compare these forms to the common genitalic morphology of other cosmetid harvestmen, and briefly discuss the systematic implications to be addressed in a subsequent revisionary study.

Morphological studies of harvestmen usually investigate the structures of adults, seldom including data regarding ontogenetic differences in the microanatomy of the appendages. In gonyleptoidean harvestmen, adults have clusters of pores on the distal-most tarsomeres of legs III and IV that are known as tarsal aggregate pores (TAPs). In gonyleptid harvestmen, these pores occur on the prolateral and retrolateral surfaces as groups adjacent to the tarsal claws. In this study, we used scanning electron microscopy (SEM) to compare the microanatomy of the TAPs occurring on leg IV of adults and antepenultimate nymphs for three common species of Neotropical cosmetid harvestmen, nymphs of an additional seven cosmetid morphospecies, and adults and antepenultimate nymphs of three species of gonyleptoidean harvestmen. The distal tarsomere of leg IV of adult cosmetid harvestmen features two pairs of aggregate pores including dorsal TAPs and ventrolateral vTAPs. The TAPs of cosmetid nymphs have denticulate bor...

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.

In many species of scorpions (Order Scorpiones) and spiders (Order Araneae), mating plugs (gel-like or sclerotized) are deposited in the female reproductive tract, minimizing the possibilities of further matings. Although harvestmen (Order Opiliones) have direct sperm transfer via copulatory organs (penis or spermatopositor in the male and ovipositor in the female), prior studies of the reproductive anatomy of these arachnids have not reported the occurrence of these structures. With the aid of scanning electron microscopy, we observed distinct masses of amorphous, gel-like material in the distal openings of the ovipositors of six different specimens including representatives of the families Cosmetidae (Cynorta marginalis and Flirtea picta), Gonyleptidae (Discocyrtulus bresslaui and Geraeocormobius sylvarum), Kimulidae (Kimula sp.) and Metasarcidae (Metasarcus clavifemur). Dissection of one of these structures from a specimen of the cosmetid harvestman C. marginalis revealed a detailed endocast of the internal surfaces of the female reproductive tract. Considerable interspecific variation in the relative sizes and shapes (smooth and rough) of the amorphous gel-like structures were observed, but evidence of sclerotized components (i.e., parts of the male genitalia) was not found. The location and morphology of the gel-like structures are consistent with observations of mating plugs in other arachnids. The composition, source (male, female or both) and function of these amorphous gel-like structures (i.e. effective matting plug) requires additional study. Fil: Townsend, Victor Robert. Virginia Wesleyan University; Estados Unidos Fil: Pérez González, Abel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Proud, Daniel Nathan. Moravian College; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

Previous studies of leg injuries in harvestmen have focused on the fitness consequences for individuals that use autospasy (voluntary detachment of the leg) as a secondary defense mechanism. Leg damage among non-autotomizing species of laniatorean harvestmen has not been investigated. Under laboratory conditions, we damaged femur IV of Cynorta marginalis and observed with scanning electron microscopy (SEM) the changes in these wounds over ten days. We also used SEM to examine leg damage from individuals of three species of cosmetid harvestmen that were collected in the field. On the basis of changes in the external surface of the hemolymph coagulum, we classified these wounds as fresh (coagulum forming), recent (coagulum with smooth surface), older (coagulum is scale-like with visible cell fragments), and fully healed (scale replaced by new cuticle growth on the terminal stump). Our observations indicate that wound healing in harvestmen occurs in a manner comparable to that of other chelicerates. Leg injuries exhibited interspecific variation with respect to the overall frequency of leg wounds and the specific legs that were most commonly damaged. In addition, we measured walking and climbing speeds of adult C. marginalis and found that individuals with fresh injuries (lab-induced) to femur IV walked at speeds significantly slower than uninjured adults or individuals collected from the field that had fully healed wounds to a single leg. J. Morphol. 278:73-88, 2017. ©© 2016 Wiley Periodicals,Inc.

Relatively little is known about the natural history of the cosmetid harvestmen that inhabit the forests of Central America. The primary objective of this study was to investigate habitat use among adult Cynorta marginalis. In the field, we established 15 transects (40 m in length) and sampled them repeatedly in the morning (0830–1100 h) and evening (1800–2300 h). During 45 morning and 35 evening surveys, we captured and marked 146 males and 112 females. Only three individuals (all females) were recaptured over the course of the study, indicating that the population size at this field site was relatively large. Heavy rains significantly reduced the surface activity of adults (1.1 individuals per transect during heavy rain vs. 3.7 individuals per transect during light or no rain). Harvestmen most commonly used leaves (65% of captures) or tree trunks (31%) as perches, and rarely occupied branches or the leaf litter. Only 18 adults were observed with leg injuries, with two individuals having damage to multiple legs. Perch height was significantly lower during the evening (110.9 cm at night vs. 155.1 cm during the day). Adults were most commonly found alone (85%), however, we observed several harvestmen in male-female pairs (8%), same sex pairs (4%) and heterosexual groups (3%). Although we did not observe reproductive or aggressive behaviors in the field, we observed intraspecific interactions under laboratory conditions and found that some individuals (15%) engaged in courtship suggesting that the sharing of perches by male-female pairs may be related to reproductive activity.

The cuticular scales of spiders are flattened setae that may occur in a diverse array of colors and shapes on the dorsal and lateral surfaces of the prosoma, opisthosoma, and walking legs. In this study, we used transmission electron microscopy (of both sections and wholemounts) and scanning electron microscopy (in concert with paraffin carving) to examine the internal anatomy and ultrastructure of the cuticular scales of several species of lynx spiders (Oxyopidae) and jumping spiders (Salticidae). We also examined iridescent and noniridescent pigmented scales for species in both families. In addition to discovering intra- and interspecific and sexual differences in scale ultrastructure, the results of our research also indicate that the ultrastructure of the scales of these spiders varies directly with coloration. For iridescent scales, we found a general absence of trabeculae, a lack of pigment granules, and an almost complete fusion of the the upper and lower laminae. For noniridescent scales, we observed granules, well-formed trabeculae, and a complex internal structure consisting of internal elements within the lumen of the scale. Our examination of the scales of these spiders represents the first complete description of the ultrastructure of the cuticular scales of any species of spider. J. Morphol. 240:77-92, 1999. © 1999 Wiley-Liss, Inc.

An examination of the cuticular scales of the lynx spiders Oxyopes aglossus, O. salticus, and Peucetia viridans using scanning electron microscopy revealed that scales in these spiders are morphologically distinct, yet similar to the scales of the jumping spiders Eris militaris and Hentzia mitrata. Like the cuticular scales of jumping spiders, the cuticular scales of lynx spiders exhibit morphological differentiation in regard to location of occurrence on the body, with scales near the eyes tending to have more numerous and larger spines on the superior surface than scales on other regions of the prosoma and opisthosoma. The functional significance of this differentiation in scale morphology is unknown. Sexual dimorphism and ontogenetic variation in scale morphology and color were observed in the genus Oxyopes,but not in Peucetia. In addition, the scales of P. viridans were distinguishable from the scales of Oxyopes spp. on the basis of the number of apical spines (1 in P. viridans instead of 3-7 in Oxyopesspp.) and on the presence of spines on the inferior surface (many in P. viridans and none in Oxyopesspp.). J. Morphol. 236:223-231, 1998. © 1998 Wiley-Liss, Inc.

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.

The legs of laniatorean harvestmen feature a considerable diversity of setae, glandular openings, sensory structures, and tubercles. In this study, we compared the morphology of cuticular structures occurring on leg IV of 12 species of cosmetid harvestmen and six species of other gonyleptoidean harvestmen representing the families Agoristenidae, Gonyleptidae (three subfamilies), and Stygnidae. Our results indicate that there is considerable interspecific variation in rough pit glands and the tarsal perforated organ among cosmetid and gonyleptoidean harvestmen. We also found interspecific variation in the relative size, number, distribution, and morphology of the tubercles on the femur, patella and tibia. We observed relatively little interspecific variation in the morphology of sensilla chaetica, trichomes, spines, single pores, tarsal aggregated pores, and slit sensilla. Additional studies of the ultrastructure of cuticular structures, especially rough pit glands, are needed to assess their functional significance. Future systematic revisions of cosmetid taxa may also benefit from the inclusion of characters based upon interspecific variation in the microanatomical structures of the legs.

There are relatively few published illustrations of ovipositors for harvestmen belonging to the suborder Laniatores. As a result, the ovipositor of these harvestmen has largely been ignored as a source of informative characters. We used scanning electron microscopy to examine the ovipositor of eight species representing several major lineages (Gonyleptoidea, Phalangodoidea, Samooidea, and Zalmoxoidea). We observed interspecific variation with respect to the number of external lobes on the distal tip, the surface texture of the distal tip, and the morphology of the peripheral setae. The ovipositors of Bishopella (Phalangodoidea) and Stygnomma (Samooidea) were similar in appearance and differed with respect to the number and position of peripheral setae. We observed significant interspecific variation among the gonyleptoidean species, especially with respect to the peripheral setae. The ovipositors of Zalmoxoidea harvestmen had smooth, spatulate peripheral setae but differed with respect to the surface texture of the distal tip. The unusual morphology of these setae has not been observed previously and may represent a new synapomorphy for the family. Characters based upon ovipositor morphology have the potential to illuminate phylogenetic relationships within and between families and genera in these harvestmen.

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.

Many species roost communally but the proximate causes and ultimate functions of this widespread behavior remain poorly understood. We studied the communal roosts of two undescribed species of harvestmen in the genus Prionostemma Pocock 1903 at a Caribbean rainforest site in southeastern Nicaragua. The species are quite similar in gross morphology but differ in body coloration, male genitalia, and roosting behavior. One species roosts primarily on spiny palms while the other species, which is darker in coloration, roosts inside buttress root cavities. In a mark-recapture study, the cavity-roosting species had higher levels of individual site fidelity than found previously in the spiny palm-roosting species, perhaps because suitable cavities are scarcer than spiny palms. The tree cavity aggregations were strongly male-biased, which our review of the literature suggests is unusual for harvestman roosts. The overall sex ratio of the spiny palm aggregations was 1∶1, but some roost sites were strongly...

The external morphology of the penis is an important source of systematic characters in phylogenetic studies of harvestmen. Modern taxonomic studies generally include micrographs generated by scanning electron microscopy (SEM) to illustrate penis morphology. In contrast, the external morphology of the ovipositor has largely been ignored for harvestmen belonging to the suborder Laniatores. Comparative studies of ovipositor microanatomy using SEM are especially lacking for species belonging to the superfamily Gonyleptoidea. In an effort to determine if the ovipositor could be a useful source of informative characters for these harvestmen, we investigated interspecific variation in the external morphology of the ovipositor for 14 species from the family Cosmetidae. Our SEM-based study revealed that the external surface of the distal tips of the ovipositors of most species was generally divided into four symmetrical lobes, although we observed a bilobed condition in Erginulus clavotibialis and Erginulus subserialis. The distal surfaces were also generally smooth, with the exception of the ovipositor of Erginulus weyerensis, which featured small surface setae. In addition, we observed considerable interspecific variation in the morphology of the peripheral setae on the distal tip, especially with respect to relative size, morphology of the shaft, and number, symmetry, and shapes of the distal tips. The functional significance, if any, of variation in the structure of the peripheral setae is unclear. Additional behavioral studies of copulation and oviposition are needed to determine the functional relationships between reproductive morphology and behavior. The morphological variation that we observed suggests that future taxonomic studies of cosmetid harvestmen, and potentially other gonyleptoidean taxa, would benefit from the inclusion of descriptions of ovipositor morphology.

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.

Most species descriptions of harvestmen from Central America that belong to the suborder Eupnoi are based on characters such as total body length, relative size of leg segments, coloration of the body and legs, and armature of the eye mound and dorsal scutum. Characters based on reproductive morphology and microanatomical structures are generally absent from most taxonomic works. Not surprisingly, the taxonomy of these harvestmen from this region requires extensive revision. In an effort to identify novel characters, we used scanning electron microscopy to compare the morphology of somatic and reproductive structures of Prionostemma vittatum (Roewer, 1910), Metopilio niger (Goodnight and Goodnight, 1942), and Metopilio ornatipes (Banks, 1909). Specifically, we investigated the morphology of the ocularium, ozopore, dorsal scutum, genital plate, pedipalp, spiracular opening on tibia II, ovipositor, and penis. We compared the morphology of these Neotropical taxa with that of a temperate sclerosomatid species, Leiobunum formosum (Wood, 1870). In general, we observed interspecific variation in the surface texture of the cuticle and the distribution of setae on the body and appendages. Potentially informative characters include cuticular structures associated with the ocularium, anterior propeltidium, meso- and metapeltidium, dorsal scutum, genital plate, pedipalp, and penis. We also observed intersexual variation with respect to morphology of Spicer's tarsal organ and the claw of the pedipalp.

Most harvestmen emit secretions from scent glands as a secondary defence mechanism. Prior studies of Phareicranaus calcariferus described the general morphology of the ozopore, noted jet emission as the mechanism for the release of secretions, but did not investigate intraspecific variation in emissions or the morphology of the ozopore. In this study, we compared the modes of emission by adult females and alpha males and found that when seized by forceps, approximately 50% of adults emitted enteric fluids (colourless, odourless), globules of secretions near the ozopores, or jets of fluid that were directed upwards and backwards or laterally. Individuals that used jet emissions usually released secretions from one scent gland and in a single burst. We observed considerable intersexual variation with respect to the type of emissions. Females emitted jets 32% of the time (n = 28), whereas males produced jets in only 4% of our observations (1 out of 26). Males more frequently produced enteric fluid (31%) in comparison to females (14%). Our SEM-based investigation of ozopore morphology did not reveal intersexual differences in microanatomy. However, we observed intraspecific variation in the morphology of the dorsal openings and dorsal channels of the ozopores.

Illustrations of penis morphology are essential components of species descriptions for harvestmen belonging to the suborder Laniatores. Male genitalia are important sources of taxonomic characters and are generally assumed to exhibit relatively little intraspecific variation. In contrast, descriptions of female reproductive morphology are rarely included in taxonomic descriptions of laniatorean harvestmen. As a result, relatively little is known about variation in the external features of the ovipositor. In this study, we used scanning electron microscopy to examine variation in male and female reproductive morphology among three species of harvestmen that are members of the superfamily Gonyleptoidea. Specifically, we examined the microanatomy of penises and ovipositors of Cynortula granulata (Cosmetidae), Phareicranaus calcariferus (Cranaidae), and Rhopalocranaus albilineatus (Manaosbiidae). Our results support the general observation that male reproductive morphology is conservative and displays little intraspecific variation. We observed considerable intraspecific variation in the number and shape of marginal setae on the ventral plate of the penis, but little or no variation in the morphology of the distal border of the ventral plate or the shape of the glans penis or stylus. With respect to female genitalia, we observed considerable intraspecific variation in the number of peripheral setae on the distal tip of the ovipositor. We also observed interspecific variation in the morphology of the peripheral setae (surface of the shaft and at shape of the distal tip), the distribution and morphology of smaller setae on the external surface of the ovipositor, and the surface texture of the external lobes. Our results indicate that there are several features associated with ovipositor morphology among laniatorean harvestmen that may represent potentially informative taxonomic characters. J. Morphol. 274:1415–1424, 2013. © 2013 Wiley Periodicals, Inc.

Descriptions of defensive behaviors used by Neotropical harvestmen are rarely based upon direct observations of encounters between individuals and syntopic predators. In this study, we investigated the defensive responses exhibited by five harvestman species from Costa Rica in interactions with ctenid spiders (Cupiennius spp.), whip spiders (Phrynus pseudoparvulus), onychophorans (Epiperipatus sp.), and centipedes (Scolopendra sp.). We also observed interactions between harvestmen and a non-predatory invertebrate, the millipede Nyssodesmus python. In 64 encounters, ctenid spiders consumed 11 harvestmen, including eight Prionostemma sp., two Cynorta marginalis, and one Eupoecilaema magnum. The most frequent defensive behavior exhibited by harvestmen was fleeing. Other behaviors included aggression (striking with leg II), bobbing, leg autotomy, lying flat, and thanatosis. Prionostemma exhibited the most variable responses, especially in the presence of ctenid spiders. In addition to the lab-based e...

Previous studies of leg injuries in harvestmen have focused on the fitness consequences for individuals that use autospasy (voluntary detachment of the leg) as a secondary defense mechanism. Leg damage among non-autotomizing species of laniatorean harvestmen has not been investigated. Under laboratory conditions, we damaged femur IV of Cynorta marginalis and observed with scanning electron microscopy (SEM) the changes in these wounds over ten days. We also used SEM to examine leg damage from individuals of three species of cosmetid harvestmen that were collected in the field. On the basis of changes in the external surface of the hemolymph coagulum, we classified these wounds as fresh (coagulum forming), recent (coagulum with smooth surface), older (coagulum is scale-like with visible cell fragments), and fully healed (scale replaced by new cuticle growth on the terminal stump). Our observations indicate that wound healing in harvestmen occurs in a manner comparable to that of other chelicerates. Leg injuries exhibited interspecific variation with respect to the overall frequency of leg wounds and the specific legs that were most commonly damaged. In addition, we measured walking and climbing speeds of adult C. marginalis and found that individuals with fresh injuries (lab-induced) to femur IV walked at speeds significantly slower than uninjured adults or individuals collected from the field that had fully healed wounds to a single leg. J. Morphol. 278:73-88, 2017. ©© 2016 Wiley Periodicals,Inc.

Relatively little is known about factors that contribute to microhabitat selection among Central American harvestmen. In this study, we compared walking and climbing speeds for five common species from Costa Rica representing the families Cosmetidae (3 species), Gonyleptidae (1 species) and Sclerosomatidae (1 species). Our sample included two arboreal species, two species that infrequently occupy perches in the vegetation, and one species that rarely climbs. Our analyses revealed no significant interspecific differences in climbing speed, although species with relatively long legs walked significantly faster than species with shorter legs. An arboreal habit did not correlate with increased climbing speed, and all species walked significantly faster than they climbed.

In tropical rain forests, harvestmen assemblages are extremely diverse, with richness often exceeding 25 species. In the neotropics, there are published accounts of harvestmen faunas in South America rainforests (especially Amazonia), but relatively little is known about the community ecology of harvestmen in tropical forests of Central America. In this paper, we provide the first insights into the diverse assemblage of harvestmen inhabiting a wet forest at La Selva Biological Station, Costa Rica. Over five field seasons, we recorded 38 species. During our 2009 field season, we examined variation in species abundance, richness, and composition between adjacent successional forests (young secondary, mature secondary, and primary forests) as well as between distinct habitats (ground/litter layer and shrub/tree layer). Based on night samples (but not day), our results indicate that there are only minor differences in species composition and relative abundance between the forest ages, but no differences in richness. The ground/litter layer and shrub/tree layer habitats differed markedly in species composition, species richness, and relative abundance of several species. Our analysis of covariance supports the hypothesis that leg length is related to climbing behavior for several species belonging to Eupnoi and Laniatores.

In encounters with predators, sclerosomatid harvestmen may employ a variety of defensive tactics including the voluntary detachment of legs (autotomy). The long-term costs of this evasive defense are not fully understood, but prior studies have documented negative consequences for terrestrial locomotion and foraging. In this study, we investigated the impact of leg loss upon locomotion in adult harvestmen (Leiobunum spp.). In southeastern Virginia, these harvestmen regularly climb vegetation and occupy perches on tree trunks, branches, and leaves that are often 1–2 m or more above the ground. In our study, we measured walking and climbing speeds for individuals with 5, 6, 7, and 8 legs. The results of our field surveys conducted over three seasons revealed relatively high frequencies (36–63%) of leg loss. We also found that individuals with six legs occupied perches that were significantly lower in the understory than those with eight legs. In the lab, we observed significantly slower walking spe...

Natural history studies serve as important foundations for future investigations of behavioral ecology and provide biologists with critical information needed for the development of empirical research programs. Relatively little is known about the biology of most species of Neotropical harvestmen in Central America. In this field study, we investigated the activity patterns of two common species Cynorta marginalis (Cosmetidae) and Prionostemma sp. (Sclerosomatidae) in a wet tropical forest in Costa Rica. We used visual encounter surveys along randomly selected transects to compare the relative abundance of these species during morning, afternoon, evening, and late night sampling periods. To generate ethograms, we observed up to three individuals of each species for 10 min along each transect, generating 69 behavioral observations for C. marginalis and 51 for Prionostemma. Both species occupy perches on vegetation and are generally nocturnal with respect to activity. After dusk, individuals of C. marginalis actively climb, interact with conspecifics, and forage on the surfaces of small and large trees. During the day, adult Prionostemma form relatively inactive clusters of up to 40 individuals on arboreal perches and descend to the leaf litter after dusk to forage.

In Central America, the family Manaosbiidae is recorded only from Panama and Costa Rica. Four species occur in this region: Barrona williamsi Goodnight & Goodnight 1942, Bugabitia triacantha Roewer 1915, Poassa limbata Roewer 1943, and Zygopachylus albomarginis Chamberlin 1925. In this paper, we describe Barrona felgenhaueri new species (Cocle Province, Panama) and Bugabitia akini new species (Cocle Province, Panama) and report a new record for B. williamsi (Cocle Province, Panama). We used SEM to examine the penis morphology of Barrona Goodnight & Goodnight 1942 and the Caribbean species Cranellus montgomeryi Goodnight & Goodnight 1947 and Rhopalocranaus albilineatus Roewer 1932. We compared genital morphology of these species with published descriptions for Manaosbiidae from South America. With respect to genital morphology, we found that the most variable characters were the number and relative sizes of the setae that occur on the lateral margins of the ventral plate. Other features that exhib...

In July and August 2005, we investigated the use of two distinct microhabitats by harvestmen (Arachnida: Opiliones) in crappo-cocorite seasonal forest, lower montane rain forest, and upper montane rain forest of the Northern Range of Trinidad, West Indies. In addition to new locality records, geographic distributions, and microhabitat use, this article provides a preliminary understanding of the species diversity, compositional similarity, and relative harvestmen abundance among forest stands located within three types of rain forest of different altitudes found in Trinidad. We collected harvestmen inhabiting the space within and underneath decaying logs as well as the leaf litter associated with the logs. In upper montane rain forest, in addition to logs, we sampled the sheaths of palm (Euterpe broadwayi Becc. ex Broadway) fronds, another shelter commonly used by harvestmen that was especially abundant at the higher elevations. We collected 173 individuals representing eight families and 14 species. The number of harvestmen per shelter varied from 3.2 individuals per log in crappo-cocorite forest to 0.8 individual per log or palm frond sheath in upper montane rain forest. In the crappo-cocorite seasonal forest, cosmetids, sclerosomatids and stygnids were most abundant. In upper montane rain forest, cranaids and manaosbiids were the most abundant taxa, whereas cosmetids and sclerosomatids were relatively rare. Crappo-cocorite and lower montane forests were very similar in species composition but differed greatly from upper montane forest. The cosmetids Paecilaema ingleiGoodnight and Goodnight 1947, Cynortula granulata Roewer 1912, and the stygnid Stygnoplus clavotibialis (Goodnight and Goodnight 1947) were found to be strongly correlated with the crappo-cocorite and lower montane forests, whereas in upper montane forests Santinezia serratotibialisRoewer 1932 and Cranellus montgomeryiGoodnight and Goodnight 1947 were closely correlated.

A new species, Santinezia noctiscansor, most similar to members of the S. festae group, is described from central Panama; this species is the only member of the genus that occurs in Central America. Our examinations of adults and nymphs collected from Coclea nd Panama Provinces revealed that males are sexually mature as penultimate nymphs. The subadult strongly resembles the adult with respect to coloration, scutal outline, armature of femora III-IV, and penis morphology, but differs in having fewer tarsomeres on each leg, a smaller body size (scutal length and width), and a much smaller tubercle on the ventral surface of coxa IV. Natural history observations are provided for specimens collected from the field site in CocleP rovince.

Morphological changes during postem- bryonic development in the Cranaidae are described on the basis of the examination of an incomplete series of larvae, nymphs, and adults of Phareicranaus calcarife- rus and Santinezia serratotibialis. The life histories of these species are hypothesized to consist of six nymphal stages, featuring the appearance of secondary male sex- ual characteristics in the antepenultimate nymph (N5). Color and body shape change dramatically during devel- opment. Growth rates for nymphs based upon leg mea- surements were similar for both species. In S. serratoti- bialis, 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 seg- ment, ocularium, pedipalp, opisthosoma, distitarsus III and IV, and leg IV. Morphological changes in postem- bryonic development in cranaid harvestmen are similar to those reported for other Laniatores. J. Morphol. 270:1055-1068, 2009. 2009 Wiley-Liss, Inc.

Larval erythraeid mites are common ectoparasites of harvestmen (Opiliones). However, interactions between these mites and hosts in tropical environments have received little attention. In this study, we examined the prevalence and intensity of mite infestations for Neotropical harvestmen from the Caribbean island of Trinidad. Overall, the prevalence of infestation was 9.9%, with significant interspecific variation in the intensity of mite infestation and significant differences in parasitism of hosts between habitats. Harvestmen collected from lowland, crappo-cocorite forest exhibited the highest infestation rates. In our samples, the intensity of infestation varied from 1 to 17 mites per host. Examination by scanning electron microscopy revealed that harvestmen were infested by two taxa of mites, a species of Leptus and another unidentified erythraeid species. Our observations represent the first host records for erythraeid mites parasitizing harvestmen from the families Cranaidae and Manaosbiidae. Mites were found to attach preferentially to the tibia and femur of the leg (175 of 178 observations). In general, the harvestmen of Trinidad exhibited mite parasitism in a fashion similar to temperate species with respect to site of attachment and prevalence and intensity of mite infestation. In the field, we also collected an adult Cynortula sp. (Cosmetidae) that was transporting 30 gravid adults of an oribatid mite (Acariformes, Oribatida, Mixonomata). Cases of mite phoresy upon harvestmen are rare, with our observation representing the first report of phoresy by an oribatid mite upon a harvestman and the first documentation of mite phoresy upon harvestmen in the Caribbean.

Relatively little is known about the microhabitat preferences of most species of Neotropical harvestmen. We investigated the use of trunks, buttresses, and the leaf litter in the immediate vicinity of trees by multiple species of harvestmen in a Trinidad, W. I. rainforest. A total of 238 individuals were collected. This included adults of species from the families Cosmetidae, Manaosbiidae, Sclerosomatidae, and Stygnidae. Our results indicate that the cosmetids (2 species), especially Cynortula sp., were the most abundant species to occur on the trees. We also found significant, positive correlations for several species with regard to tree size and the number of individuals present. We hypothesize that harvestmen may use the surfaces of trees as well as the leaf litter in the immediate vicinity of the buttresses as either shelters from potential predators or as areas that provide favorable microclimates (e.g., relatively high humidity).

Relatively little is known about the food habits of neotropical harvestmen. We used Erginulus clavotibialis (Pickard-Cambridge 1905), a locally abundant species of cosmetid harvestman in Belize, in a food choice experiment. Individuals were presented with fresh fruit (pineapple) and live invertebrate prey (termites) in an experimental chamber. This species showed a strong preference for fruit, as 72% of individuals ate it first and 67% spent the most time in the fruit-containing portion of the experimental chamber. Five E. clavotibialis (13%) consumed termites, confirming this species' ability to capture and consume live invertebrate prey. Adult males located food more quickly than nymphs. Harvestmen feeding on fruit were also significantly more active than non-feeding individuals or those preying upon termites first. Opportunistic frugivory may be important to E. clavotibialis during times when fruit is available (e.g., wet season). We hypothesize that this species exhibits a generalist diet in ...

We describe the sequence of cranial ossification in the Tailed Frog, Ascaphus truei, and examine intraspecific variation in the timing and sequence of cranial development. Because tadpoles of A. truei lack the subarticular tubercles that normally delineate Gosner stages 39 and 40, we provide a modified Gosner staging system more appropriate for this species. Ossification sequence was highly variable among the populations and greater than that reported for other anuran species. In general, early ossification events in A. truei are similar to that reported for other anuran species. However, compared with other taxa, both onset of initial ossification events and appearance of specific elements appear relatively later in the development of Ascaphus.

Adults and larvae of the loberine erotylid beetle Loberus impressus (LeConte) were found associated with fungi growing on corolla and seed capsule tissue of the blue flag iris, Iris hexagona. We examined adult beetle specimens using light and scanning electron microscopy to determine if specialized structures (mycangia) may function in transporting fungi. Two pairs of deep pits on the ventral aspect of the gena between the eyes and the maxillae of both sexes contained fungal spores and hyphae, suggesting a possible role as mycangia in addition to their role as glandular outlets. Inoculation from the surface of cleaned beetle specimens produced colonies of Cladosporium and Fusarium. These genera are widespread, usually air-dispersed conidial fungi that sometimes are associated with insects.

A series of laboratory experiments were conducted to determine if adult wolf spiders Gladicosa pulchra were capable of acting as predators of juvenile spotted salamanders Ambystoma maculatum. Spiders preyed upon salamanders in two of 20 trials and attacked salamanders in five trials. As attacks did not always kill salamanders, multiple attacks per trial were possible. A total of 11 attacks was observed: two complete attacks, and nine incomplete attacks. Attacks were considered complete if the spider inserted its chelicerae into the salamander and incomplete if the spider did not. In response to complete attacks, the salamander would flip onto its back. Salamanders occasionally responded to incomplete attacks by continuously raising and lowering their heads. To our knowledge, this behaviour is undescribed in this group of salamanders. An additional experiment investigated the influence of increased levels of salamander movement on this interaction. In this experiment, spiders preyed upon salamanders in one of eight trials and attacked salamanders in six trials (28 total attacks: four complete attacks, 24 incomplete attacks). The results of this study indicate that wolf spiders can prey upon juvenile spotted salamanders, and that the ecological relationship between wolf spiders and amphibians may be quite complex.

We describe multiple observations of epizoic cyanobacteria occurring on external surfaces of a species of sclerosomatid harvestman (Prionostemma sp.) in Costa Rica. In the field we collected four adults (three males, one female) that had green films growing upon the dorsal surfaces of the carapace and abdominal scutum. Examination by SEM revealed dense clusters of what appeared to be small prokaryotic cells (1–5 µm in diameter) covering the external surfaces of the carapace, abdominal scutum and coxae. We extracted DNA from the films of two specimens. The DNA was used as a template to amplify the intergenic spacer region (IGS) between the beta and alpha phycocyanin subunits (a signature DNA sequence, unique to cyanobacteria) by PCR. We successfully amplified an approximately 700 base pair product using DNA extracted from the film and did not obtain any product from the harvestman lacking the film. Our observation represents the second confirmed occurrence of epizoic cyanobacteria on Neotropical h...

In Acheta domesticus, the Malpighian tubules (Mt) are composed of three morphologically distinct regions (proximal, mid and distal), each consisting of a single cell type. The bulk of the Mt is composed of the midtubule, which shows the greatest response to corticotropin releasing factor-related diuretic peptides (CRF-DP). We know from previous laboratory studies that the second messenger cAMP and its analog dibutyryl cAMP (db-cAMP) cause an approximate doubling in the secretion rate and that this is accompanied by notable ultrastructural changes in the midtubule, especially membrane reorganization in the basal area and extensive vesiculation of the cytoplasm. In this study, we examined the morphological changes in membranes both at the cell surface and internally. By enzymatically removing the basal lamina, we examined the increase in spacing between infolded membranes initiated by db-cAMP stimulation. To examine the intracellular membranes, we used a technique developed for use in invertebrate tissues. This allowed the removal of the cytoplasm for high resolution scanning electron microscopy (HR-SEM) while maintaining the integrity of the lipid constituents of the cell. By using HR-SEM and confocal laser scanning microscopy (CLSM), we gained a unique three-dimensional perspective of the complexity of the internal membrane system of the A. domesticus Mt in both the unstimulated and db-cAMP-stimulated states.

Although relatively rare among harvestmen in the superfamily Gonyleptoidea, paternal care has been observed in the families Manaosbiidae and Gonyleptidae, but not previously in the Cosmetidae. In this study, we describe multiple observations of egg guarding by adult males of an undescribed species of cosmetid harvestman from Volcan Cacao, Guanacaste Province, Costa Rica. Observations were made from 26–28 July 2010, during the wet season. In this species, males only guard eggs after dusk, leaving eggs unattended during the day. Based upon differences in color and size, males guarded eggs through several stages of development. When guarding, males contacted the first two pairs of legs with the eggs. Oviposition sites consisted of the undersides of leaves of small plants, with eggs closely packed together in a single layer covered by abundant, transparent mucus. The largest, darkest eggs were located near the distal tip of the leaf.

The external morphology of the penis is an important source of systematic characters in phylogenetic studies of harvestmen. Modern taxonomic studies generally include micrographs generated by scanning electron microscopy (SEM) to illustrate penis morphology. In contrast, the external morphology of the ovipositor has largely been ignored for harvestmen belonging to the suborder Laniatores. Comparative studies of ovipositor microanatomy using SEM are especially lacking for species belonging to the superfamily Gonyleptoidea. In an effort to determine if the ovipositor could be a useful source of informative characters for these harvestmen, we investigated interspecific variation in the external morphology of the ovipositor for 14 species from the family Cosmetidae. Our SEM-based study revealed that the external surface of the distal tips of the ovipositors of most species was generally divided into four symmetrical lobes, although we observed a bilobed condition in Erginulus clavotibialis and Erginulus subserialis. The distal surfaces were also generally smooth, with the exception of the ovipositor of Erginulus weyerensis, which featured small surface setae. In addition, we observed considerable interspecific variation in the morphology of the peripheral setae on the distal tip, especially with respect to relative size, morphology of the shaft, and number, symmetry, and shapes of the distal tips. The functional significance, if any, of variation in the structure of the peripheral setae is unclear. Additional behavioral studies of copulation and oviposition are needed to determine the functional relationships between reproductive morphology and behavior. The morphological variation that we observed suggests that future taxonomic studies of cosmetid harvestmen, and potentially other gonyleptoidean taxa, would benefit from the inclusion of descriptions of ovipositor morphology.

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.

Relatively little is known about the morphology and ultrastructure of the Malpighian tubules of spiders (Arachnida: Araneae). Our study represents the first investigation of the Malpighian tubules of a theraphosid spider and is the only study to examine the living Malpighian tubules using confocal laser scanning microscopy. In theraphosid spiders, the Malpighian tubules originate from the stercoral pocket in the posterior portion of the opisthosoma and extend forward toward the prosoma in a dendritic pattern. There are three distinct segments (initial, main, and terminal), all dark brown in appearance. Each segment has distinctive ultrastructural features. Both the terminal and the main segment appear to be composed of at least two cell types with finger-like cytoplasmic protrusions associated with one of these types. The terminal segment, which is most proximal to the stercoral pocket, is the largest in diameter. It is composed of large, cuboidal cells containing many mitochondria and lipid inclusions. The main segment is intermediate in diameter with many mitochondria and secretory vesicles present. The initial segment is relatively thin in comparison to the other segments and is intimately associated with the digestive gland. The cells of the initial segment contain very little cytoplasm, fewer mitochondria, secretory vesicles, and prominent inclusions. J. Morphol. 251:73–82, 2002. © 2002 Wiley-Liss, Inc.

Intraguild predation is described as predation among organisms that exploit similar resources. As wolf spiders (Araneae, Lycosidae) are generalist predators that share habitat and food resources with the ground skink Scincella lateralis, we conducted a series of laboratory experiments to determine if wolf spiders are capable of preying upon ground skinks. Wolf spiders (Gladicosa pulchra) successfully preyed on skinks during 3 of 20 encounters, but did not overtly respond to chemical stimuli from the skinks. Skinks employed antipredatory behaviors (i.e., immobility) when exposed to visual and chemical stimuli from spiders, providing evidence that they recognize spiders as predators prior to an encounter. In an additional experiment, the hypothesis that increased skink movement would lead to an increase in spider attacks was tested. Skinks exhibited high levels of movement in this experiment, with spider predation occurring during 4 of 10 encounters. The presence of structural refugia played a significant role in this predator–prey interaction by increasing the amount of time required for a spider to prey upon a skink. Our results indicate that G. pulchra can prey upon S. lateralis, and that skinks may recognize cues deposited on the substrate by spiders. These data indicate that these two species may interact in an ecologically meaningful manner.

In spiders, cuticular scales are flattened, hollow setae that occur in a variety of shapes and colours. Using transmission electron microscopy and scanning electron microscopy, we examine and provide descriptions of the external and internal anatomy of prosomal and opisthosomal scales of 70 species of lynx spiders (Araneae, Oxyopidae), representing the genera Hamataliwa, Hostus, Oxyopes, Peucetia, Schaenicoscelis, Tapinillus, and Tapponia. The cuticular scales of these taxa exhibit regional (with respect to location on the body), sexual, inter- and intrageneric variation in external and internal morphology. In the genera, Peucetia, Schaenicoscelis, and Tapinillus, cuticular scales are plumose and only occur on the prosoma. Species of the genera Hamataliwa and Tapponia, have lanceolate scales that occur on the prosoma, legs, and opisthosoma. Hostus has spatulate scales in these areas. In the genus Oxyopes, most species have lanceolate scales on the prosoma and spatulate scales on the legs and opisthosoma. Examinations of the genitalia of 82 species of lynx spiders, revealed a strong correlation between genitalic morphology and scale shape and distribution.

The larva of Loberus impressus LeConte is described based on adult-associated and reared specimens collected from corollas and seed pods of Iris hexagona on the gulf coast of Louisiana. The larva of L. impressus is similar to larvae of Zavaljus brunneus (Gyllenhal) Hapalips prolixus Sharp, Pharaxonotha spp., and Bolerus angulosus (Arrow), the other described larvae within the Xenoscelinae. Larval characters typical for the Loberini that are found in L. impressus include arrangement of integumental granules into rows, frayed and aciculate setae borne on spiny tubercles, and urogomphi spiny or tuberculate and recurved. The tarsungular setae of L. impressus and B. angulosus are single, in contrast to the dual setae present in other described larvae of Languriidae. Larva and adults of L. impressus were common in dried corollas and seed pods of I. hexagona that had begun to split open or had been invaded by Crematogaster ants. Fusarium and Cladosporium fungi were potential sources of food for L. impressus and other fungivorous beetles in the iris seed pods and on corollas.

This study presents the first complete description of the external and internal morphology of cuticular scales of spiders of the genus Hamataliwa and is also the first examination of intrageneric variation in the morphology of the scales of the lynx spiders (Oxyopidae). The cuticular scales of nine species, including taxa from Australia and Central and North America, were examined using scanning electron microscopy, paraffin carving, and transmission electron microscopy (whole mounts only). The surface morphology and internal anatomy of the scales exhibit considerable intra- and inter-specific variation. The structures that exhibit the most variation are (i) the plicae, small lateral ridges on the dorsal surface, which show variation in both morphology and pattern, and (ii) the rod-shaped, branched cuticular elements that occur within the lumina, which vary most dramatically in pattern. Scale morphology also varies regionally (i.e., with respect to location on the body) in most species. A comparison of the morphology of scales of Hamataliwa spp. with those of the striped lynx spider, Oxyopes salticus, and the green lynx spider, Peucetia viridans, suggests that scales may represent a useful phylogenetic character for understanding relationships both within and between genera in the Oxyopidae.