Anatomy and function of the ptychoid defensive mechanism in the mite Euphthiracarus cooki (Acari: Oribatida)

Ptychoidy is a defensive adaptation of several groups of oribatid mites in which legs and coxisternum can be fully retracted into the opisthosoma and protected by a ventrally deflected prodorsum, resulting in a seed-like appearance. Using Euphthiracarus cooki as a model, we examined details of exoskeletal and muscular anatomy in combination with studies of live individuals to provide the first functional analysis of ptychoidy. There are two main functional components: the first is a set of exoskeletal and muscular adaptations, mostly of the podosoma and prodorsum, that combine to effect leg withdrawal and prodorsal deflection; the second comprises adaptations of the opisthosoma that allow control of hydrostatic pressure during the large hemocoel volume adjustments associated with ptychoidy. Adaptations important in the closing process (enptychosis) are found in four body regions. Much of the podosomal exoskeleton (especially pleural) is unsclerotized, which facilitates leg retraction and prodorsal deflection during enptychosis. The coxisternum has several flexible furrows along which it folds in order to bring legs into a tightly parallel arrangement. The prodorsum has specialized attachment surfaces (manubrium and inferior retractor process) for retractor muscles and a paired bothridial scale that participates in prodorsal alignment during enptychosis. The subcapitulum has a prominent capitular apodeme on which important retractor muscles insert. The mineralized notogaster has an anterior "collar" that accommodates the retracted prodorsum; it includes paired notches and receptacles that accommodate the bothridial scales, thereby creating a temporary fixed axis for rotation of the prodorsum in a "lazy hinge" mechanism. Specialized muscles form the retractor system; most conspicuous are the large coxisternal retractors and prodorsal retractors, both of which originate on the notogaster. Other components have adjustor roles; among them are muscles of the endosternal system that control retraction of the subcapitulum and assist leg retraction, and the dorsoventral muscles which adjust the folded coxisternum. Hemolymph pressure control is a function of the opisthosoma, where the principle exoskeletal elements form a pleated venter, having a cross-sectional shape like an inverted "W." Paired holoventral plates (each representing fused genital, aggenital, anal, and anal plates) form the inner angle and are flanked by paired plicature plates. The holoventral plates are connected medially in two ways : 1) by three permanent bridges of sclerotized cuticle that include an anterior phragmatal bridge and two widely spaced, hollow apodemes (preanal, postanal); 2) by temporary zipper-like closures of two different types. Lateral compression of the pleats is effected by a series of transversely arranged, lateral compressor muscles that run from plicature to holoventral plate edges, and from holoventral plate edges to the medial apodemes. Compression increases hydrostatic pressure in the opisthosoma and stores energy in both the slightly deformed, mineralized notogaster and in the three holoventral bridges. During normal activity the compressor system is active and the inflated podosomal region provides support for the extended legs. When the mite is irritated, the prodorsum is hydraulically ejected from its active position in the notogastral collar, then relaxation of the compressors causes a fall in hemolymph pressure and return of the notogaster to an undeformed condition. Muscles of the retractor system then act in specific sequence to retract and adjust the coxisternum and prodorsum until they are precisely positioned at the completion of enptychosis. The process takes between 0.5 and 1 sec. When irritation ceases, partial opening allows sensory leg hairs to "test" the environment. Resumption of normal, active posture (ecptychosis) involves activation of the lateral compressor system and hydraulic inflation of the podosoma, through which legs are extended and the prodorsum is reflected.     

Morphology, microanatomy and sequence data of Sclerolinum contortum (Siboglindae, Annelida) of the Gulf of Mexico

Sclerolinum is a small genus of Siboglinidae (Annelida) living in an obligate mutualistic association with thiotrophic bacteria as adults. Its taxonomic position, based on morphology, has been controversial; however, molecular data point to a sister taxa relationship with vestimentiferans. 16S rRNA gene sequencing and comparative morphology revealed that the studied population from deep-sea hydrocarbon seeps of the Gulf of Mexico belongs to Sclerolinum contortum known from the Arctic Sea. Since no anatomical and microanatomical studies have been published yet, we conducted such a study on S. contortum using serial sectioning and light and transmission electron microscopy. We show that the Sclerolinum body, divided into a head, trunk, and opisthosoma, is very similar to that of the vestimentiferans, and therefore we propose that the body regions are homologous in both taxa.     

Centers of mass and weight distribution in spiders of the family uloboridae

In the Uloboridae, web reduction is accompanied by changes in opisthosomal shape, leg length, and web-monitoring tactics. These morphological changes make reduced-web spiders more cryptic and alter their leg leverage and centers of mass. When compared with the orb-weaver Uloborus glomosus, the irregular, reduced-web spider, Miagrammopes animotus, invests more mass in its prosoma and first legs. However, the latter species' elongate opisthosoma posteriorly shifts this region's center of mass, causing the relative position of its composite center of mass and the distribution of weight between its first and fourth legs to be similar to that of the orb-weaver. Like these species, the opisthosomal center of mass of the triangle-weaver, Hyptiotes cavatus, lies near its midpoint. However, the shorter first legs and rounder, heavier opisthosoma of Hyptiotes posteriorly shift its composite center of mass and distribute more of its weight onto its fourth legs. Consequently, the morphology of M. animotus can be adequately explained by its adaptiveness for web manipulation, balance, and weight distribution and the crypsis that these features confer as an ancillary advantage. In contrast, anatomical changes in H. cavatus are better explained as adaptations for web manipulation and crypsis.     

Mechanism of signal production in the vibratory communication of the wandering spider Cupiennius getazi (Arachnida,Araneae)

The communication with substrate vibrations produced by vibrations of the body or its appendages is widespread among arthropods, especially among spiders. Its biomechanics, however, is poorly understood. Males of the wandering spider Cupiennius getazi produce such substrate vibrations during courtship by means of dorsoventral movements of their opisthosoma without hitting their dwelling plant.Simultaneous recordings of the plant vibrations (accelerometry), of the opisthosoma movements (laser Doppler vibrometry) and of the electromyograms of the opisthosomal depressor muscle, revealed that the main frequency of the vibratory signal of about 80 Hz originates from the activity of the opisthosomal depressor muscle. The transfer functions of the spider's body show resonances which could amplify the main frequency before it is transmitted into the plant.A low frequency component of the opisthosomal movement (duration c. 0.3 s, displacement c. 6 mm (peak-peak) 30° deflection angle, frequency 10–20 Hz) can be distinguished from a main frequency component (duration c. 0.1 s, displacement c. 0.5 mm 2.5° deflection angle, frequency c. 80 Hz). The main frequency component is superimposed on an upward movement of the low frequency component.     

Respiratory system of arachnids I: morphology of the respiratory system of Salticus scenicus and Euophrys lanigera (Arachnida, Araneae, Salticidae)

The book-lungs and the tracheal systems of two species of jumping spider, Salticus scenicus and Euophrys lanigera, were investigated using gross anatomical, light and electron microscopic methods. Both species possess well-developed book-lungs of similar size and tracheal systems with a basically similar branching pattern. The tracheal spiracle opens into a single atrium, where it gives rise to four thick 'tube tracheae', from which small secondary tube tracheae originate in groups. The secondary tracheae (diameter 1-5 mum) run parallel, without further branching, into the prosoma. In the opisthosoma, they lie ventrolaterally, where they contact muscles and internal organs. In the prosoma, the secondary tracheae may penetrate the gut epithelium and central nervous tissue. The structure of the tracheal walls is very similar to that of insects, consisting of a striated inner cuticular layer with taenidial structures and a surrounding outer hypodermal layer. The wall thickness appears similar in all secondary tracheae, indicating that lateral gas diffusion may be possible through the walls of all small tube tracheae.     

Developmental and functional ultrastructure of Ornithodiplostomum ptychocheilus diplostomula (Trematoda: Strigeoidea) during invasion of the brain of the fish intermediate host, Pimephales promelas

We examined tegumental development of the diplostomulum of Ornithodiplostomum ptychocheilus, with respect to structural transformations that have functional relevance to the invasion, migration, and site establishment processes in the brain of the fish second-intermediate host, Pimephales promelas. Using a combination of brightfield, scanning electron microscopy (SEM), transmission electron microscopy (TEM), and confocal microscopy (CM), we demonstrated that the diplostomula become established in the outer region of the optic lobes within 24-48 hr of penetration and continue to grow and transform over a period of 4-14 days. During this period, the J-shaped body consists of 2 distinct regions: (1) a highly motile prosoma with distinctive tegumental spines and (2) an opisthosoma, the tegument of which is elaborated into a dense uniform layer of long, thin microvilli. The prosoma is alternately invaginated into and everted from the opisthosoma, thus constituting a protrusible proboscis. By day 14 postinfection (PI), the body has lost this bipartite structure and has taken on the uniformly flattened form characteristic of metacercariae. The transitory complex structure of the diplostomula appears to be well suited to burrowing through host tissues (primarily by action of the prosoma), followed by rapid dissociation of host tissue and nutrient accumulation (primarily by action of the opisthosoma) in preparation for metacercaria encystment.     

The hydraulic interaction between prosoma and opisthosoma in Amaurobius ferox (Chelicerata,Araneae)

Summary Experiments are described which demonstrate that blood moves from the prosoma into the opisthosoma in Amaurobius ferox when the spider is held on a plasticine block and is stimulated with a small brush. This movement of blood is also seen during bouts of struggling when the spider is trying to free itself. The return flow of blood from the opisthosoma is to a large extent due to the pumping action of the heart.It is proposed that the locomotory exhaustion shown by artificially stimulated spiders is due to this loss of blood from the prosoma which leads not only to hydraulic insufficiency, but also to a lack of oxygen through interruption of the normal blood flow.     

Morphology of the prosomal endoskeleton of Scorpiones (Arachnida) and a new hypothesis for the evolution of cuticular cephalic endoskeletons in arthropods

Skeletomuscular anatomy of the scorpion prosoma is examined in an attempt to explain the evolution of two endoskeletal features, a muscular diaphragm dividing the prosoma and opisthosoma and cuticular epistomal entapophyses with a uniquely complex arrangement of muscles, tendons and ligaments. Both structures appear to be derived from modifications of the mesodermal intersegmental endoskeleton that is primitive for all major arthropod groups. The scorpion diaphragm is a compound structure comprising axial muscles and pericardial ligaments of segments VI to VIII and extrinsic muscles of leg 4 brought into contact by longitudinal reduction of segment VII and integrated into a continuous subvertical sheet. This finding reconciles a long-standing conflict between one interpretation of opisthosomal segmentation based on scorpion embryology and another derived from comparative skeletomuscular anatomy. A new evolutionary-developmental mechanism is proposed to account for the complex morphology of the epistomal entapophyses. Each entapophysis receives 14 muscles and tendons that in other taxa would attach to the anterior connective endoskeleton in the same relative positions. This observation suggests that the embryological precursor to the connective endoskeleton can initiate and guide ectodermal invagination and thereby serve as a spatial template for the development of cuticular apodemes. This mesoderm-template model of ectodermal invagination is potentially applicable to all arthropods and may explain structural diversity and convergence in cephalic apodemes throughout the group. The model is used to interpret the cephalic endoskeletons of two non-chelicerate arthropods, Archaeognatha (Hexapoda) and Symphyla (Myriapoda), to demonstrate the generality of the model.     

Rallinyssus sorae sp. n. (Acari: Dermanyssidae: rhinonyssinae) from the nasal passages of the sora, Porzana carolina (Aves: Rallidae)

Rallinyssus sorae sp. n. is described from the nasal turbinates of the sora, Porzana carolina, collected in Maryland and Ohio. The new species is most similar to Rallinyssus verheyeni Fain but differs in number of setae on the female ventral opisthosoma, structure of the fixed and movable digits of the male and female chelicerae, chaetotaxy of legs I to IV, and presence of a sternal plate in the male. The taxonomy and host-parasite relationships of the various species of Rallinyssus are discussed.     

Conservation and variation in Ubx expression among chelicerates

SUMMARY Chelicerates are an ancient arthropod group with a distinct body plan composed of an anterior (prosoma) and a posterior portion (opisthosoma). The expression of the Hox gene Ultrabithorax (Ubx) has been examined in a single representative of the chelicerates, the spider Cupiennius salei. In spiders, Ubx expression starts in the second opisthosomal segment (O2). Because the first opisthosomal segment (O1) in spiders is greatly reduced relative to other chelicerates, we hypothesized that the observed Ubx expression pattern might be secondarily modified. Shifts in the anterior boundary of the expression of Ubx have been correlated with functional shifts in morphology within malacostracan crustaceans. Thus, the boundary of Ubx expression between chelicerates with different morphologies in their anterior opisthosoma could also be variable. To test this prediction, we examined the expression patterns of Ubx and abdominal‐A (collectively referred to as UbdA) in two basal chelicerate lineages, scorpions and xiphosurans (horseshoe crabs), which exhibit variation in the morphology of their anterior opisthosoma. In the scorpion Paruroctonus mesaensis, the anterior border of early expression of UbdA is in a few cells in the medial, posterior region of the O2 segment, with a predominant expression in O3 and posterior. Expression later spreads to encompass the whole O2 segment and a ventral, posterior portion of the O1 segment. In the xiphosuran Limulus polyphemus, early expression of UbdA has an anterior boundary in the segment. Later in development, the anterior boundary moves forward one segment to the chilarial (O1) segment. Thus, the earliest expression boundary of UbdA lies within the second opisthosomal segment in all the chelicerates examined. These results suggest that rather than being derived, the spider UbdA expression in O2 likely reflects the ancestral expression boundary. Changes in the morphology of the first opisthosomal segment are either not associated with changes in UbdA expression or correlate with late developmental changes in UbdA expression.     

Heart and circulatory functions in a spider (Eurypelma californicum): the effects of hydraulic force generation

Summary In the tarantulaEurypelma californicum, the relationships between heart activity, circulation and the generation of hydraulic pressure for locomotion were studied. Several new techniques were employed.Mean resting heart rate was 21 beats min^–1 rising to 90 beats min^–1 after burst activity. Decay time to resting rates was related to the increase of heart rate. Post-recovery resting rates were usually elevated in comparison with rates after very long resting periods.A relative measure of heart amplitude was obtained. Four distinct patterns could be distinguished: (i) regular beats; (ii) short-term fluctuations of amplitude within a few heart beats; (iii) a slow rhythmic change of heart/pericardium filling, and (iv) non-periodic, stronger amplitude changes during periods of activity.During locomotion, heart rate rises with maximum rates often reached only minutes after the onset of activity. The rising phase is often characterized by irregularities and a reduction of heart amplitude.Prosomal hemolymph pressure in resting, restrained animals was 41±19 Torr, rising to ca. 90, and 217±48 Torr during walking and fast sprints, respectively. Values in unrestrained spiders were similar. Opisthosomal hemolymph pressures were ca. 20 Torr in resting animals, rising to 40–60 Torr during locomotion.Opisthosomal volume changes were measured. A small volume of hemolymph moved from the prosoma to the opisthosoma at the onset of locomotion, but following activity this volume quickly returned to the prosoma.The simultaneous measurement of carapace depression, opisthosomal volume changes and hemolymph pressures, and heart activity revealed the relationship between circulation and hydraulic force generation. The direction of hemolymph flow was also studied. In non-active animals, the heart occasionally changes its main pumping direction. During locomotion, hemolymph flow from the heart to the prosoma is often reduced or stopped. With a slight delay, hemolymph flow to the opisthosoma is increased. The critical pressure at which prosomal perfusion from the heart is halted is 50–70 Torr.It is concluded that anterior and posterior circulations are separate: hemolymph returning from the prosoma passes only through the anterior lungs, while hemolymph returning from the opisthosoma passes through the posterior lungs.Dedicated to Dr. Rosemarie John, in recognition of her unflagging enthusiasm and support for zoological researchProf. B. Linzen unexpectedly died on August 5, 1988     

The structure and potential loasaceous affinities of Schismocarpus

The affinity of Schismocarpus with Loasaceae has been questioned. Characteristic trichomic features are most often used for definition of Loasaceae, and Schismocarpus possesses a distinctly loasaceous syndrome of trichomes. Its inflorescence morphology is similar to that of Eucnide and Mentzelia , and it shares floral features with Eucnide. Although Schismocarpus possesses unique features (e.g., perennating "rootstock" and striate pollen) relative to Loasaceae, it is not aberrant considering the broad range of diversity that has been long accepted within the family. The apparently divergent floral morphology may be explained by its evolution of a "pollen flower" syndrome.     

山羊蠕形螨扫描电镜观察

本文通过扫描电镜对寄生虫山羊体的山羊蠕形螨生活史各期颚体,足体和末体的超微结构观察,发现以下超微结构;一个圆锥状须乳突位于雌螨触须第２节背面;叶状背阳茎侧突和腹阳茎侧突竖立于阳茎同一基部,未受精雌螨船形阴门的一对瓣汇成纵线,雌螨交配后阴门半开,产卵后全部敞开,同时描述该螨的背基刺,锥状突,口下板,螯肢,基内叶,口孔,须爪和肛道等的超微结构,文中还讨论了锥状突,口下板,螯肢,基内叶,须爪和须乳突的功     

A histochemical study of aldehyde fuchsin-positive material and "high-esterase cells" in the pineal gland of the Mongolian gerbil.

The pineal gland of the Mongolian gerbil consists of a superficial gland, stalk and deep pineal. The deep pineal differentiates postnatally. Histochemical studies of the superficial pineal gland indicate that it may be involved in the secretion of protein. Presumptive secretory material visualized by aldehyde fuchsin (AF) and chrome hematoxylin was observed along the course of blood vessels and among the pinealocytes. The distribution and texture of the AF-positive material was distinctive. It did not correspond to the pattern and texture of material stained with PAS, Sudan Black or acid orcein. Staining with AF was markedly reduced after incubation with trypsin, indicating that the AF-positive material is at least partially protein. The amount of stainable material increased with age. The AF-positive material was observed in what appeared to be interstitial or glial cells and processes, and in the processes of perivascular cells. Cells and fibrous processes with high non-specific esterase activity ("high-esterase cells") were observed among the pinealocytes and along the course of blood vessels. The distribution of the "high-esterase cells" and the morphology and texture of their esterase-containing processes were remarkably similar to the morphology and distribution of the material that stained with AF. It may be that the "high-esterase cells" contain AF-positive material. The "high-esterase cells" hydrolyzed both alpha-naphthyl acetate and alpha-naphthyl butyrate. The pinealocytes hydrolyzed only alpha-naphthyl acetate. The "high-esterase cells" appear to form a distinct class of cells within the superficial pineal gland. They are tentatively identified as a type of glial cell.     

Allopsoroptoides galli n. g., n. sp., a new genus and species of feather mites (Acari: Analgoidea: Psoroptoididae) causing mange in commercially raised domestic chicken in Brazil

A new feather mite, Allopsoroptoides galli n. g., n. sp. (Psoroptoididae: Pandalurinae), is described from the domestic chicken, Gallus gallus (Linnaeus) (Galliformes: Phasianidae), from Brazil. This is the first record of a representative of the feather mite family Psoroptoididae from an avian host of the order Galliformes. The new genus is closely related to the genus Cygnocoptes Fain & Bochkov, 2003 but clearly differs from the latter and all other genera of the family by the loss of four median pairs of hysteronotal setae (c1, d1, e1, and h1) in both sexes and by the unique shape of the male opisthosoma. Instead of the bilobate opisthosoma, the male opisthosoma in this genus has a narrow and long median projection, ending with a pair of semi-ovate terminal lamellae. These mites were first detected during a mange outbreak in several commercial poultry facilities in the state of São Paulo, Brazil.     

The spinnerets and epiandrous glands of spiders

The spinnerets of spiders are carried on the fourth and fifth segments of the abdomen (opisthosoma). Primitively there are two pairs, anterior lateral ( al ) and anterior median ( am ) on the fourth segment, and two pairs, posterior lateral ( pl ) and posterior median ( pm ) on the fifth, am are present in Liphistius but are never functional. In mygalomorphs am are invariably absent, and usually al also. In araneomorphs am are either reduced to a function-less colulus, perhaps absent altogether, or represented by the cribellum, which is a specialized spinning organ.
It seems unlikely that the lateral and median spinnerets correspond to the exopodites and endopodites of a biramous limb, which limbs are characteristic of the Crustacea, a group having no close relationship to the Arachnida. From embryology it seems clear that the lateral spinnerets are the segmental appendages. Glands, here described as the epiandrous glands, very similar to spinning glands, occur on the second abdominal segment of most male spiders. It is suggested that these may be serially homologous with the median spinnerets, which would then not be appendicular in origin but would be modifications of ventral glandular structures.     

Morphology and ultrastructure of the Malpighian tubules of the Chilean common tarantula (Araneae: Theraphosidae).

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.     

Morphology, phylogeny and biology of Gliocephalis hyalina, a biotrophic contact mycoparasite of Fusarium species

Gliocephalis hyalina, a rarely seen microfungus with a morphology similar to the hyphomycete genus Aspergillus but with slimy conidia was found in a mixed microbial culture from soybean roots. This species has been reported sporadically since 1899, each time in association with other fungi or bacteria. Gliocephalis hyalina has not been maintained in monoxenic culture and requires other fungi to grow. Light and scanning electron microcope studies indicate that it is a biotrophic contact parasite of Fusarium species. The fungus may penetrate the cells but has no apparent deleterious effect on the growth or plant pathogenicity of its host. Phylogenetic analyses of partial nuclear small subunit rDNA sequences place G. hyalina near the Laboulbeniales, an order of obligate insect parasitic microfungi, and the related mycelial genus Pyxidiophora. Gliocephalis hyalina is mycoparasitic along with many Pyxidiophora species. These discoveries suggest that some "unculturable" microorganisms or "cryptic DNA" recovered from environmental DNA samples might represent obligate biotrophs that could be cultured and studied with simple techniques.     

Shared human-chimpanzee pattern of perinatal femoral shaft morphology and its implications for the evolution of hominin locomotor adaptations

Background

Acquisition of bipedality is a hallmark of human evolution. How bipedality evolved from great ape-like locomotor behaviors, however, is still highly debated. This is mainly because it is difficult to infer locomotor function, and even more so locomotor kinematics, from fossil hominin long bones. Structure-function relationships are complex, as long bone morphology reflects phyletic history, developmental programs, and loading history during an individual’s lifetime. Here we discriminate between these factors by investigating the morphology of long bones in fetal and neonate great apes and humans, before the onset of locomotion.

Methodology/Principal Findings

Comparative morphometric analysis of the femoral diaphysis indicates that its morphology reflects phyletic relationships between hominoid taxa to a greater extent than taxon-specific locomotor adaptations. Diaphyseal morphology in humans and chimpanzees exhibits several shared-derived features, despite substantial differences in locomotor adaptations. Orangutan and gorilla morphologies are largely similar, and likely represent the primitive hominoid state.

Conclusions/Significance

These findings are compatible with two possible evolutionary scenarios. Diaphyseal morphology may reflect retained adaptive traits of ancestral taxa, hence human-chimpanzee shared-derived features may be indicative of the locomotor behavior of our last common ancestor. Alternatively, diaphyseal morphology might reflect evolution by genetic drift (neutral evolution) rather than selection, and might thus be more informative about phyletic relationships between taxa than about locomotor adaptations. Both scenarios are consistent with the hypothesis that knuckle-walking in chimpanzees and gorillas resulted from convergent evolution, and that the evolution of human bipedality is unrelated to extant great ape locomotor specializations.