Some comments on the hydrostatic system of spiders (Chelicerata,Araneae)

An anatomical study of five spider types has shown that the musculi laterales of the prosoma, together with the subcuticular muscle sheet of the opisthosoma, may be jointly responsible for generating the internal hydrostatic pressures which control the leg extension mechanism.     

Segmentation and tagmosis in Chelicerata

Patterns of segmentation and tagmosis are reviewed for Chelicerata. Depending on the outgroup, chelicerate origins are either among taxa with an anterior tagma of six somites, or taxa in which the appendages of somite I became increasingly raptorial. All Chelicerata have appendage I as a chelate or clasp-knife chelicera. The basic trend has obviously been to consolidate food-gathering and walking limbs as a prosoma and respiratory appendages on the opisthosoma. However, the boundary of the prosoma is debatable in that some taxa have functionally incorporated somite VII and/or its appendages into the prosoma. Euchelicerata can be defined on having plate-like opisthosomal appendages, further modified within Arachnida. Total somite counts for Chelicerata range from a maximum of nineteen in groups like Scorpiones and the extinct Eurypterida down to seven in modern Pycnogonida. Mites may also show reduced somite counts, but reconstructing segmentation in these animals remains challenging. Several innovations relating to tagmosis or the appendages borne on particular somites are summarised here as putative apomorphies of individual higher taxa. We also present our observations within the concept of pseudotagma, whereby the true tagmata – the prosoma and opisthosoma – can be defined on a fundamental change in the limb series while pseudotagmata, such as the cephalosoma/proterosoma, are expressed as divisions in sclerites covering the body without an accompanying change in the appendages.     

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.     

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     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

Expression of pair-rule gene homologues in a chelicerate: early patterning of the two-spotted spider mite Tetranychus urticae

Embryo segmentation has been studied extensively in the fruit fly, DROSOPHILA: These studies have demonstrated that a mechanism acting with dual segment periodicity is required for correct patterning of the body plan in this insect, but the evolutionary origin of the mechanism, the pair-rule system, is unclear. We have examined the expression of the homologues of two Drosophila pair-rule genes, runt and paired (Pax Group III), in segmenting embryos of the two-spotted spider mite (Tetranychus urticae Koch). Spider mites are chelicerates, a group of arthropods that diverged from the lineage leading to Drosophila at least 520 million years ago. In T. urticae, the Pax Group III gene Tu-pax3/7 was expressed during patterning of the prosoma, but not the opisthosoma, in a series of stripes which appear first in even numbered segments, and then in odd numbered segments. The mite runt homologue (Tu-run) in contrast was expressed early in a circular domains that resolved into a segmental pattern. The expression patterns of both of these genes also indicated they are regulated very differently from their Drosophila homologues. The expression pattern of Tu-pax3/7 lends support to the possibility that a pair-rule patterning mechanism is active in the segmentation pathways of chelicerates.     

Patterning mechanisms and morphological diversity of spider appendages and their importance for spider evolution

The prosoma of spiders bears different gnathal (labrum, chelicerae, pedipalps) and locomotory appendages (legs). In most species these appendages are also used for additional functions, e.g. sensing, mating, and courtship. The opisthosoma is equipped with four pairs of highly specialized appendages. Two pairs of spinnerets are used for silk production and manipulation. The other two pairs of appendages are internalized during development and give rise to a complex respiratory system of book lungs and tracheae. Thus spiders have a number of different appendage types with radically different adult morphologies. Furthermore, all these appendage types display significant additional species specific diversity correlating with a large spectrum of functions of the appendages. Despite this importance of appendage diversity for the evolution of the spiders we know relatively little about the genetic patterning mechanisms producing this diversity of morphology. We review recent advances concerning the developmental genetics of spider appendage diversification, mainly concentrating on open questions and future directions of research. We conclude that the deeper understanding of appendage development and diversity in spiders can contribute significantly not only to evolutionary developmental biology, but also to behavioral biology, speciation research and population genetics, and the study of sexually dimorphic traits.     

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.     

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.     

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.     

Genomic and gene regulatory signatures of cryptozoic adaptation: Loss of blue sensitive photoreceptors through expansion of long wavelength-opsin expression in the red flour beetle Tribolium castaneum

Background

Hox genes are expressed in specific domains along the anterior posterior body axis and define the regional identity. In most animals these genes are organized in a single cluster in the genome and the order of the genes in the cluster is correlated with the anterior to posterior expression of the genes in the embryo. The conserved order of the various Hox gene orthologs in the cluster among most bilaterians implies that such a Hox cluster was present in their last common ancestor. Vertebrates are the only metazoans so far that have been shown to contain duplicated Hox clusters, while all other bilaterians seem to possess only a single cluster.

Results

We here show that at least three Hox genes of the spider Cupiennius salei are present as two copies in this spider. In addition to the previously described duplicated Ultrabithorax gene, we here present sequence and expression data of a second Deformed gene, and of two Sex comb reduced genes. In addition, we describe the sequence and expression of the Cupiennius proboscipedia gene. The spider Cupiennius salei is the first chelicerate for which orthologs of all ten classes of arthropod Hox genes have been described. The posterior expression boundary of all anterior Hox genes is at the tagma border of the prosoma and opisthosoma, while the posterior boundary of the posterior Hox genes is at the posterior end of the embryo.

Conclusion

The presence of at least three duplicated Hox genes points to a major duplication event in the lineage to this spider, perhaps even of the complete Hox cluster as has taken place in the lineage to the vertebrates. The combined data of all Cupiennius Hox genes reveal the existence of two distinct posterior expression boundaries that correspond to morphological tagmata boundaries.     

Teratological deformities of pedipalps in the Tegenaria atrica spider,induced by low and high temperatures applied alternately

The study was aimed at demonstrating the relationship between temperature (as a factor which disturbs morphogenesis) and deformities in the front part of the prosoma in Tegenaria atrica. By exposing spider embryos to alternating temperatures of 14 and 32 °C for the first 10 days of embryonic development, i.e. until the first metameres of the prosoma appeared on the germ band, we obtained individuals with a range of anomalies including those which affected the prosomal morphology. We selected five spiders with deformities of pedipalps or of pedipalps and walking legs for comprehensive analysis. The results indicate a relationship between temperature applied as a teratogenic factor and anomalies in the front part of the prosoma.     

The delineation of the fourth walking leg segment is temporally linked to posterior segmentation in the mite Archegozetes longisetosus (Acari: Oribatida,Trhypochthoniidae)

Acari (mites and ticks) lack external segmentation, with the only indication of segmentation being the appendages of the prosoma (chelicerae, pedipalps, and four pairs of walking legs). Acari also have a mode of development in which the formation of the fourth walking leg is suppressed until the nymphal stages, following a hexapodal larva. To determine the number of segments in the posterior body region (opisthosoma) of mites, and to also determine when the fourth walking leg segment is delineated during embryogenesis, we followed the development of segmentation in the oribatid mite Archegozetes longisetosus using time‐lapse and scanning electron microscopy, as well as in situ hybridizations of the A. longisetosus orthologues of the segmentation genes engrailed and hedgehog. Our data show that A. longisetosus patterns only two opisthosomal segments, indicating a large degree of segmental fusion or loss. Also, we show that the formation of the fourth walking leg segment is temporally tied to opisthosomal segmentation, the first such observation in any arachnid.     

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.     

Gross morphology,histology, and ultrastructure of the alimentary system of Ricinulei (Arachnida) with emphasis on functional and phylogenetic implications

Ricinuleid functional mouthparts are the cucullus, the chelicerae, the pedipalps, and the labrum. These structures are movably jointed to the rest of the prosoma, most likely protruded upon hydrostatic hemolymph pressure and retracted by prosomal muscles. Seta‐like protrusions from the labrum and the pedipalpal coxae form a sieve‐like filter inside the preoral cavity and the mouth. Although the tip of the labrum can be elevated upon muscle constriction, ingestion of large, solid food particles is unlikely. The mouth has a crescent‐shaped cross section. The cuticle‐lined, also crescent‐shaped pharynx is equipped with a large dilator muscle but lacks antagonistic constrictor muscles. It represents a precerebral sucking pump. The triangular to Y‐shaped, cuticle‐lined esophagus is equipped with constrictor and dilator muscles. Its posterior part represents a postcerebral sucking pump. Four blind ending diverticula ramify from the anterior prosomal part of the entodermal midgut tube. Two of these diverticula remain inside the prosoma and form few short branches. The other two extend through the pedicel into the opisthosoma and ramify and coil there. A stercoral pocket protrudes ventrally out of the midgut tube. The most distal part of the midgut tube is modified into a contractile rectal gland. Its secretions may have defensive or physiological functions. A short anal atrium is formed by the cuticle‐lined ectodermal hindgut which opens at the end of the three‐segmented metasoma. The telescoping segments of the metasoma are protruded by hemolymph pressure and retracted by muscles. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Bicephality,a seldom occurring developmental deformity in Tegenaria atrica caused by alternating temperatures

The experiment was aimed at demonstrating the relationship between deformities of the front part of the prosoma accompanied by changes in the brain structure in bicephalous Tegenaria atrica and exposure of their embryos to temperature fluctuations. By exposing spider embryos to alternating temperatures of 14 and 32 °C for the first 10 days of embryonic development, we obtained eight two-headed individuals, subsequently divided into three groups according to morphological differences. We described in detail morphological abnormalities of the prosoma identified in members of each group. Histological examination confirmed a close relationship between morphological deformities and the brain structure of teratogenically changed spiders. The fusion of appendages (pedipalps and chalicerae) was accompanied by the fusion of corresponding ganglia. The absence of appendages (pedipalps) was accompanied by the absence of corresponding ganglia. This correlation may have resulted from previously impaired neuromere development which led to changes in the morphological structure of the prosoma. Since no deformities were identified in control animals, it can be concluded that bicephaly was caused by exposing embryos to alternating temperatures.     

Morphological Characterization of First Instar Larvae of Asian Horseshoe Crabs and their Hybrids*

The morphological characters of first instar larvae of three Asian horseshoe crabs and their hybrids were examined. For this purpose, the length of six parts on the larvae, the pattern of pigmentation, the structure of the margin of the opisthosoma, and the distribution and shape of spines on the extracardiac region of the prosoma were recorded. In general, the phenotypes of the hybrids were intermediate between those of the parents, suggesting that both male and female genomes contribute to development of the hybrids. However, there were some exceptional characters and phenotypic variations. Some characters were inherited from only one parent. This was suggested to imply that if the maternal species did not have the gene regulating expression of a given paternal character, the paternal gene did not express this character.     

A New Three-Dimensionally Preserved Xiphosuran Chelicerate from the Montceau-Les-Mines Lagerstätte (Carboniferous, France)

Excellent three dimensional preservation of 142 specimens of Alanops magnificus gen. et sp. nov. (Chelicerata: Xiphosura) from the Stephanian Konservat-Lagerstätte of Montceau-les-Mines (Saône-et-Loire, France), exposes the carapace design and hitherto unrecorded details of fossil xiphosuran ventral anatomy, and makes possible an interpretation of appendicular functional morphology. All legs are long, slender and chelate. The chelate condition of the fifth leg and the shape of the prosoma (highly vaulted with a subvertical frontal area) indicate that the animal was neither a burrower like Recent xiphosurans (e.g. Limulus ) nor an active swimmer as suggested for other extinct forms, but was more likely to have been a benthic crawler. The ability to fold up is attested by both partly enrolled specimens showing appendages withdrawn into the prosomal cavity, and by coaptive devices on the external and internal margins of both prosoma and opisthosoma. The low-level articulation system at the boundary between the two shields allowed both the carapace closure and the complete outstretching of the animal. The unusually small adult size of Alanops magnificus gen. et sp. nov. combined with the loss of ophthalmic ridges and spines is interpreted as indicating a paedomorphic derivation from a bellinurine stock. The depositional environment (limnic intramontane basin) and faunal association (dominated by syncarids and bivalves) of Alanops indicate that the animal probably lived in fresh water.