Exoskeletal cuticle differentiation during intramarsupial development of Porcellio scaber (Crustacea: Isopoda)

Exoskeletal crustacean cuticle is a calcified apical extracellular matrix of epidermal cells, illustrating the chitin-based organic scaffold for biomineralization. Studies of cuticle formation during molting reveal significant dynamics and complexity of the assembly processes, while cuticle formation during embryogenesis is poorly investigated. This study reveals in the terrestrial isopod Porcellio scaber, the ultrastructural organization of the differentiating precuticular matrices and exoskeletal cuticles during embryonic and larval intramarsupial development. The composition of the epidermal matrices was obtained by WGA lectin labelling and EDXS analysis. At least two precuticular matrices, consisting of loosely arranged material with overlying electron dense lamina, are secreted by the epidermis in the mid-stage embryo. The prehatching embryo is the earliest developmental stage with a cuticular matrix consisting of an epicuticle and a procuticle, displaying WGA binding and forming cuticular scales. In newly hatched marsupial larva manca, a new cuticle is formed and calcium sequestration in the cuticle is evident. Progression of larval development leads to the cuticle thickening, structural differentiation of cuticular layers and prominent cuticle calcification. Morphological characteristics of exoskeleton renewal in marsupial manca are described. Elaborated cuticle in marsupial larvae indicates the importance of the exoskeleton in protection and support of the larval body in the marsupium and during the release of larvae in the external environment.     

Aerial manoeuvrability in wingless gliding ants (Cephalotes atratus)

In contrast to the patagial membranes of gliding vertebrates, the aerodynamic surfaces used by falling wingless ants to direct their aerial descent are unknown. We conducted ablation experiments to assess the relative contributions of the hindlegs, midlegs and gaster to gliding success in workers of the Neotropical arboreal ant Cephalotes atratus (Hymenoptera: Formicidae). Removal of hindlegs significantly reduced the success rate of directed aerial descent as well as the glide index for successful flights. Removal of the gaster alone did not significantly alter performance relative to controls. Equilibrium glide angles during successful targeting to vertical columns were statistically equivalent between control ants and ants with either the gaster or the hindlegs removed. High-speed video recordings suggested possible use of bilaterally asymmetric motions of the hindlegs to effect body rotations about the vertical axis during targeting manoeuvre. Overall, the control of gliding flight was remarkably robust to dramatic anatomical perturbations, suggesting effective control mechanisms in the face of adverse initial conditions (e.g. falling upside down), variable targeting decisions and turbulent wind gusts during flight.     

Cuticular structure of the agnostine trilobite Homagnostus obesus

Study of the exoskeletal surface microstructure of specimens of Homagnostus obesus (Belt, 1867) from the Upper Cambrian of Sweden has given information about the structure of agnostine cuticle. It is likely that the very thin cuticle of agnostines (5-15 μm), unlike that of polymerid trilobites, was constructed only of a prismatic layer. The exoskeletons were strengthened by reticulation on the external surface, the ridges forming up to 15% of the total cuticle thickness. Pits on the visceral surface of the exoskeleton of H. obesus may have contained photoreceptors as their morphology is similar to that of the Nileus glabellar 'tubercle'. This would have allowed the animal to monitor changes in light intensity. Possible sensory receptors in other agnostine trilobites are reviewed. Most sense organs were positioned on the unmineralized ventral surface of the organism. □ Trilobita, Agnostina, exoskeleton. cuticle, micro-structure, sensory receptors.     

Temperature distribution and electrical properties along the Oriental hornet body

The hornet is an endothermic insect. Daily variations in hornet surface temperature were measured. Three peaks were found between 9:30 and 10: 30 a.m., 11 and 12 a.m. and between 2 and 3 p.m. Electrical current and voltage values were highest along the head. Electrical current along the gaster and the head flowed towards the thorax, i.e., from body parts with minimal temperature towards the body part with maximal temperature. Current and voltage values measured across the cuticle of the gaster were about 5nA and 100 mV, respectively, and these were of the same order of magnitude as the current and voltage values along the cuticle. It was found that: 1) temperature regulation most probably originates in the thorax and 2) there is a correlation between the temperature distribution along the hornet body surface and levels of the cuticular electrical signals.     

Preliminary assessment of metabolic costs of the nematode Myrmeconema neotropicum on its host, the tropical ant Cephalotes atratus

The parasitic nematode Myrmeconema neotropicum infects workers of the neotropical arboreal ant Cephalotes atratus. Infected ants exhibit altered behavior, e.g., reduced aggression and slower tempo, as well as physical traits, e.g., gaster changes from shiny black to bright red. These changes are thought to induce fruit mimicry and attract frugivorous birds, which are the presumed paratenic hosts for the nematodes. We used respirometry to measure the energetic costs of nematode infection, testing the prediction of higher metabolic rates for infected workers maintaining both ant and nematode biomass. Contrary to this prediction, infected workers had lower mass-specific metabolic rates than uninfected workers. Parasites are limited to the gasters (abdomens) of adult ants, and infected gasters had 57% more mass, but 37% lower metabolic rates, compared to uninfected gasters. These results use a metabolic currency to measure, in vivo, the energetic costs of parasitism, and they shed light on the complex co-evolutionary relationship between host and parasite.     

New Family,Genus, and Species of Microsporida (Protozoa: Microsporida) from the Tropical Fire Ant,Solenopsis geminata (Fabricius) (Insecta: Formicidae)*

SYNOPSIS. A new species of Microsporida, Burenella dimorpha sp. n., representing a new family, Burenellidae fam. n. and genus, is described on the basis of light- and electron-microscope observations. The family is characterized by 2 sequences of sporogony, each sequence having morphologically different sporonts and spores. The parasite infects the tropical fire ant, Solenopsis geminata (Fabricius), producing distinct pathologic manifestations (clearing of the cuticle and eye malformation) and death in the pupal stage of development. Transmission of the infection per os to healthy S. geminata, to the Southern fire ant, Solenopsis xyloni McCook, and to the red and black imported fire ants, Solenopsis invicta Buren and Solenopsis richteri Forel, is reported.     

A Diverse Ant Fauna from the Mid-Cretaceous of Myanmar (Hymenoptera: Formicidae)

A new collection of 24 wingless ant specimens from mid-Cretaceous Burmese amber (Albian-Cenomanian, 99 Ma) comprises nine new species belonging to the genus Sphecomyrmodes Engel and Grimaldi. Described taxa vary considerably with regard to total size, head and body proportion, cuticular sculpturing, and petiole structure while all species are unified by a distinct shared character. The assemblage represents the largest known diversification of closely related Cretaceous ants with respect to species number. These stem-group ants exhibit some characteristics previously known only from their extant counterparts along with presumed plesiomorphic morphology. Consequently, their morphology may inform hypotheses relating to basal relationships and general patterns of ant evolution. These and other uncovered Cretaceous species indicate that stem-group ants are not simply wasp-like, transitional formicids, but rather a group of considerable adaptive diversity, exhibiting innovations analogous to what crown-group ants would echo 100 million years later.     

Ant body posture: gaster curling increases ant speed

1. Body postures adopted by an animal can serve behavioural functions, homeostasis, or energy balance. 2. We investigated the function of holding the gaster curled forward under the thorax in acacia ants, Pseudomyrmex spinicola Emery, by testing whether ants adopted this posture for defence, thermoregulation, or for efficient locomotion. 3. For the defence hypothesis, we expected an increase in the proportion of ants with curled gasters after a visual threat, a vibrational disturbance of a branch, or the release of nestmate's alarm pheromones. Our data did not support these predictions. 4. For the thermoregulation hypothesis, we found a positive correlation between temperature and proportion of curled‐gaster ants. However, we did not find a reduction in the proportion of curled‐gaster ants after shading them, as predicted by this hypothesis. 5. Our data supported the locomotion hypothesis: curled‐gaster ants walked 1 cm s^?1 faster than ants with the gaster held straight. Straight‐gaster ants walked with the thorax closer to the surface, a posture that likely shifts the centre of gravity closer to the surface in a manner similar to gaster curling. 6. Studying the role of the body posture in acacia ants and other insects will provide a better understanding of the kinematics of walking in challenging angles with respect to gravity.     

Ant-mimicry in some Brazilian salticid and clubionid spiders (Araneae: Salticidae,Glubionidae)*

This paper describes the morphological and behavioural adaptations responsible for ant-like appearance in eight species (genera Zuniga, Synemosyna, Sphecotypus, and Myrmecium) of salticid and clubionid spiders studied in Amazonian and SE Brazil. All ant-mimicking spiders have body and legs thin, and the shiny integument typical of their models. Light horizontal hair bands and constrictions of the cephalothorax and abdomen simulate, respectively, the head-thorax joint and segmented gaster of ants. The petiole and postpetiole of the ants are usually mimicked by a lengthened pedicel, together with a narrowing of the posterior cephalothorax and/or anterior abdomen. The prominent pedipalps of the spiders often simulate ant mandibles, but they may also be strikingly similar to an ant's head. All ant-mimicking spiders walked in a zig-zag ant-like pattern, and frequently raised and moved about the first pair of legs as ‘antennae’. The mimics were found in the same microhabitats (foliage or ground) as their models, and displayed strong avoidance reactions toward the latter both in the field and in captivity. The inoffensive characteristics of the mimics and the noxious traits of their models (strong mandibles, potent sting, hard integument, venomous secretions) strongly suggest that the spiders are Batesian ant-mimics. The detailed structural and behavioural adaptations enhancing ant-mimicry provide strong circumstantial evidence that the selective agents involved must have good visual acuity, and are probably small insectivorous vertebrates (e.g. birds, lizards and toads) or arthropods (e.g. wasps and spiders) which avoid ants.     

Cataglyphis desert ants improve their mobility by raising the gaster

We analyze theoretically the moment of inertia of the desert ant Cataglyphis (C. bicolor and C. fortis) around a vertical axis through its own center of mass when the animal raises its gaster to a vertical position. Compared to the value when the gaster is horizontal, the moment of inertia is reduced to one half; this implies that when increasing its angular acceleration the ant need apply only half the level of torque when the gaster is raised, compared to when the gaster is lowered. As an example, we analyze the cases of an ant running on circular and sinusoidal paths. In both cases, the ant must apply a sideways thrust, anti-roll and anti-pitch torques to avoid toppling, and, on the circular path when accelerating and throughout the sinusoidal trajectory, a torque to enable turning as the path curves. When the ant is accelerating in a very tight circle or running on a very narrow sinusoidal path, in which the amplitude of the sinusoid is less than the length of the ant's body, the forces required for the turning torque can equal and exceed those required for the sideways thrust, and can be reduced significantly by the ant raising the gaster, whereas the foot-thrust for the anti-roll and anti-pitch torques rises only modestly when the gaster is up. This suggests that there may be an evolutionary advantage for employing the gaster-raising mode of locomotion, since this habit will allow desert ants to use lower forces and less energy, and perhaps run faster on more tortuous paths.     

Mechanisms affecting load size determination inAtta cephalotes L. (Hymenoptera,Formicidae)

Summary The size of leaf-cutting ant foragers correlates more closely with the weight than the area of the leaf fragments they cut. This implies that the mechanism of load size determination is not a simple function of body geometry. Ants were found not to adjust the radius of cut to compensate for experimental changes in leaf weight during the process of cutting. However, ants changed their cutting behaviour according to the thickness of the region of the fragment being cut. Ants decreased their cut radius when cutting through leaf veins. Similarly, when cutting artificial laminae of varying thicknesses, ants reduced the cut arc radius when traversing thicker regions.Cut radius was not principally controlled by the position of the rear legs nor the overall body length but by mechanisms associated with the head and thorax of the ant.     

Behavioral mechanisms of spatial competition between red wood ants (Formica aquilonia) and ground beetles (Carabidae)

Behavioral aspects of spatial competition between red wood ants (Formica aquilonia) and six mass species of Carabidae were studied in field and laboratory experiments. We showed that red wood ants essentially influence spatial distribution of ground beetles on their common territories. Transplantation experiments suggest that in newly established ants' settlements stronger forms of interrelations arise than in old stable colony. To examine the ability of beetles to avoid collisions with ants we used two experimental techniques. In laboratory, we tested carabids ability to avoid a clash in a Y-shaped labyrinth containing an active tethered ant in one section. In field experiments we compared quantitative characteristics of movements (such as crookedness of individual trajectories, speed of movement, the time spent on stops) for beetles placed close to ants foraging routes and on ant-free plots. All beetles studied displayed a clear tendency to learn, that is, to modity their behavior in order to avoid collisions with ants. Species that exhibited best parameters of learning were closer to ants by their size and characteristic movement, namely, Pterostichus oblogopunctatus and P. magus. Beetles' stereotyped behavioral tactics can be considered universal for avoiding collisions with any subject (for instance, with an ant) of a certain size and speed of movements. A set of tactics in the labyrinth included: (1) attempts to round the ant; (2) turns away after touching the ant with antennae; (3) turns away without a contact; (4) avoidances of a dangerous section; (5) stops near the ant with the antennae hidden. Comparing pairwise difference between four species shows that beetles use species-specific preference for definite combinations of tactics. Effective learning allows carabids to penetrate into ant foraging territory and partly avoide interference competition. It seems that red wood ants are not inclined to learn to avoid collisions with competing carabid species. Instead, they recognize an "enemy's image" and selectively attack relatively small predatory carabids rather than herbivorous species. Experiments with dummy beetles suggest that ants react for several hierarchically organized key characteristics of competitors such as speed of movement, dark color, bilateral symmetry, protuberances (legs, antennae), and smell. Among "professional" groups in ant family, guards and hunters react to beetles aggressively, whereas aphid tenders ignore them. Sophisticated combination of flexible and innate behavioral patterns enables insects to share territories and interact in the mode of relatively mild, spatial, competition instead of predation. Eliciting sets of hierarchically organized features of competitors that govern adjustment of spatial distribution in insects' community enable us to integrate ideas of classic and cognitive ethology.     

The Effect of Diet and Opponent Size on Aggressive Interactions Involving Caribbean Crazy Ants (Nylanderia fulva)

Biotic interactions are often important in the establishment and spread of invasive species. In particular, competition between introduced and native species can strongly influence the distribution and spread of exotic species and in some cases competition among introduced species can be important. The Caribbean crazy ant, Nylanderia fulva, was recently introduced to the Gulf Coast of Texas, and appears to be spreading inland. It has been hypothesized that competition with the red imported fire ant, Solenopsis invicta, may be an important factor in the spread of crazy ants. We investigated the potential of interspecific competition among these two introduced ants by measuring interspecific aggression between Caribbean crazy ant workers and workers of Solenopsis invicta. Specifically, we examined the effect of body size and diet on individual-level aggressive interactions among crazy ant workers and fire ants. We found that differences in diet did not alter interactions between crazy ant workers from different nests, but carbohydrate level did play an important role in antagonistic interactions with fire ants: crazy ants on low sugar diets were more aggressive and less likely to be killed in aggressive encounters with fire ants. We found that large fire ants engaged in fewer fights with crazy ants than small fire ants, but fire ant size affected neither fire ant nor crazy ant mortality. Overall, crazy ants experienced higher mortality than fire ants after aggressive encounters. Our findings suggest that fire ant workers might outcompete crazy ant workers on an individual level, providing some biotic resistance to crazy ant range expansion. However, this resistance may be overcome by crazy ants that have a restricted sugar intake, which may occur when crazy ants are excluded from resources by fire ants.     

Shear adhesive performance of leaf‐cutting ant workers (Atta cephalotes)

Wingless arboreal ants must resist the force of gravity while traversing substrates in their environment. For leaf‐cutting ants like Atta cephalotes, foraging may also include a ca. 30 m vertical descent while carrying a load 1–6 times their body mass. We hypothesized that heavier and larger ants would carry heavier and larger loads and that adhesive performance would positively correlate with load mass. We found no relationship between ant mass, body length, head width, or adhesive performance, and the load size an ant carried. In addition to workers carrying vegetative loads (most often leaves), workers in an active foraging trail also include smaller workers riding on the leaves carried by larger workers, and large major workers, providing protection from aerial and ground attacks (Soldiers), respectively. Despite varying functional roles, all foraging ants require secure attachment to the substrate. We measured shear adhesive performance of each foraging role and found that Soldiers produced the highest shear adhesive forces. However, when controlling for tarsal pad area, we found that ants carrying loads have higher shear adhesive performance per unit area than those riding on leaves, and that Soldiers have the lowest shear adhesive performance per unit area. This suggests that while leaf choice does not appear to be dictated by size, mass, or shear adhesive performance of individual ants, overall, ants who carry leaves adhere more strongly given their pad size than those who do not. Abstract in Spanish is available with online material.     

Proteins of the Crustacean Exoskeleton

We describe here some of the components of the exoskeleton ofthe decapod crustacean with emphasis on the constituent proteins,including both structural and enzymatic. All four layers, butparticularly the inner three, of the exoskeletons of four brachyuranscontain high concentrations of proteins 31 kDa; the innermostmembranous layer is especially rich in such proteins. A numberof crab exoskeletal proteins resemble insect cuticle proteinsin size (M_r) and isoelectric point (pI). A further similarityis the cross reactivity of crab exoskeletal proteins with fourdifferent antibodies against cuticular proteins of two speciesof insects. One of the small M_r exoskeletal proteins in theBermuda land crab Gecarcinus lateralis has a similar distributionas a protein of similar size in the cuticle of the tobacco hornworm Manduca sexta. The partial dissolution of an old exoskeletonand formation of the two outer layers of a new exoskeleton aremajor events in readying a crustacean for the increase in sizethat occurs at each molt. Expressing both parallel and sequentialactivation of a number of genes, a single layer of epidermalcells that bounds a crustacean such as G. lateralis synthesizesspecific proteins at different stages of the intermolt cycleas the outermost epicuticle and exocuticle are formed duringproecdysis and as the endocuticle and membranous layer are formedduring metecdysis. Finally, two sets of proteinases isolatedfrom integumentary tissues of land crabs degrade the same exoskeletalproteins in vitro as are degraded in vivo during proecdysis.     

Cuticle collagen genes. Expression in Caenorhabditis elegans

Collagen is a structural protein used in the generation of a wide variety of animal extracellular matrices. The exoskeleton of the free-living nematode, Caenorhabditis elegans, is a complex collagen matrix that is tractable to genetic research. Mutations in individual cuticle collagen genes can cause exoskeletal defects that alter the shape of the animal. The complete sequence of the C. elegans genome indicates upwards of 150 distinct collagen genes that probably contribute to this structure. During the synthesis of this matrix, individual collagen genes are expressed in distinct temporal periods, which might facilitate the formation of specific interactions between distinct collagens.     

Huffmanela cf. carcharhini (Nematoda: Trichosomoididae: Huffmanelinae) from skin of a sandbar shark, Carcharhinus plumbeus, in the Pacific Ocean

Eggs of Huffmanela cf. carcharhini from the skin of an aquarium-held, juvenile sandbar shark, Carcharhinus plumbeus , from the Pacific Ocean were studied using light and scanning electron microscopy. Grossly, eggs imparted a scribble-like skin marking approximately 130 × 60 mm on the right side of the shark's snout adjacent to its eye and nostril. Fresh (unfixed) eggs were elliptical, 75-95 μm long (xˉ = 85 μm, SD = ±4.5; n = 75), 48-63 μm wide (53 ± 3.4; 75), 8-10 μm in shell thickness (9 ± 1.3; 27), 45-68 μm in vitelline mass length (52 ± 6.9; 8); had a smooth shell surface and nonprotruding polar plugs 8-13 μm wide (10 ± 1.5; 73); lacked thin filaments, superficial envelope, and shell spines; sank in 35 ppt artificial seawater; and did not spontaneously hatch after 12 hr in 35 ppt artificial seawater. Formalin-fixed eggs measured 193 days postfixation were 75-95 μm long (84 ± 3.9; 150), 45-60 μm wide (50 ± 2.2; 150), 5-10 μm in shell thickness (8 ± 1.2; 87), 45-60 μm in vitelline mass length (51 ± 3.0; 92), and 30-40 μm in vitelline mass width (33 ± 2.0; 84), and had nonprotruding polar plugs that were 10-15 μm long (11 ± 1.4; 93) and 8-10 μm wide (9 ± 1.1; 108). Forcibly hatched first-stage larvae (unfixed) were filiform, 188-273 μm long (212 ± 25.5; 13), 8-13 μm wide (10 ± 1.2; 13), and had fine transverse striations. Eggs infected the epidermis only. Histology revealed intra-epithelial inflammation with eosinophilic granulocytes and hyperplasia, plus dermal lymphofollicular hyperplasia associated with the infection. The eggs of H. cf. carcharhini likely undergo considerable ex utero development before being sloughed (unhatched) from the host, along with epidermal cells.     

Surface structures of the antennular flagella of the hermit crab Pagurus alaskensis (Benedict): A light and scanning electron microscopy study

The surface structures of the antennular flagella of Pagurus alaskensis are described in detail. Attention is directed towards the surface morphology of two types of possible sensilla: (1) exoskeletal pores (1.0–3.0 μm in diameter); (2) setae of various kinds. In addition, small (0.1–0.2 μm) pits occur in the exoskeleton which are not considered to be sensory in function. The exoskeletal pores are found at fairly specific locations on both the inner and outer flagella, particularly on the short segments of the outer flagella. Neither the inner nor the outer flagella are bilaterally symmetrical with respect to their setal armature. On the outer flagellum six groups of setae may be distinguished: lateralmesial; dorsal; ventral; accessory; aesthetasc; setae of the distal segment. On the inner flagellum setae of the mesial and lateral rows form distinctive groups. The morphology, orientation and locations of all the flagellar setae are defined and where possible the numbers of the various morphological types within the specific setal groups are given. It is noteworthy that many setal types have obvious apical pores and yet no pores could be found in the chemoreceptive aesthetasc setae. The functions of the various setae are discussed in relation to their topographical position and to existing electrophysiological and behavioral data. Some suggestions are made about future experiments to demonstrate the central connections of specific sensilla or groups of sensilla and to show their significance in the whole animal.     

The cuticle of the nematode Caenorhabditis elegans: A complex collagen structure

The cuticle of the nematode Caenorhabditis elegans forms the barrier between the animal and its environment. In addition to being a protective layer, it is an exoskeleton which is important in maintaining and defining the normal shape of the nematode. The cuticle is an extracellular matrix consisting predominantly of small collagen-like proteins that are extensively crosslinked. Although it also contains other protein and non-protein compounds that undoubtedly play a significant part in its function, the specific role of collagen in cuticle structure and morphology is considered here. The C. elegans genome contains between 50 and 150 collagen genes, most of which are believed to encode cuticular collagens. Mutations that result in cuticular defects and grossly altered body form have been identified in more than 40 genes. Six of these genes are now known to encode cuticular collagens, a finding that confirms the importance of this group of structural proteins to the formation of the cuticle and the role of the cuticle as an exoskeleton in shaping the worm. It is likely that many more of the genes identified by mutations giving altered body form, will be collagen genes. Mutations in the cuticular collagen genes provide a powerful tool for investigating the mechanisms by which this group of proteins interact to form the nematode cuticle.     

Blastodermic cuticles of a springtail,Tomocerus ishibashii Yosii (Collembola : Tomoceridae)

The formation and structure of the blastodermic cuticles of a springtail, Tomocerus ishibashii Yosii (Collembola : Tomoceridae) are described together with the change of egg membrane. The blastodermic cuticles of the Collembola are 2-layered, and formed in the early stages of the embryonic development, preceding the differentiation of germ band. The first blastodermic cuticle is thicker (about 0.8-1.5 μm in thickness) and its surface is provided with complex structures, whereas the second one is thinner (about 0.2-0.4 μm in thickness) and smooth. About 3 days after oviposition, the chorion (about 2 μm in thickness) splits into 2 and the first blastodermic cuticle, provided with many projections and 4 large spines appear on the surface of the egg. Three types of projections are distinguished: button-, cone- and seta-like structures. The halves of the ruptured chorion are attached to the first blastodermic cuticle on both sides below the spines, and no projections are found in the regions concealed by the ruptured chorion. The projections of the first blastodermic cuticle are formed by cellular protrusions of the blastoderm. The conspicuous large spines on the first blastodermic cuticle are formed by the evaginations of the blastoderm. Tendrils of the primary dorsal organ run between the first and second blastodermic cuticles.