Notes on some tardigrades from Thailand,with descriptions of two new species

Abstract

Three species of tardigrades are reported from Thailand: Pseudechiniscus quadrilobatus Iharos, 1969, Calcarobiotus (Calcarobiotus) digeronimoi sp. n., and Mixibius sutirae sp. n. Calcarobiotus (C.) digeronimoi sp. n. has claws of Calcarobiotus type; those of the first three pairs of legs have a medial and a lateral spur, those of the hind legs lack basal spurs. Mixibius sutirae sp. n. has a densely sculptured dorsal cuticle and two macroplacoids; it lacks lunules and other thickenings on the legs. It is similar to Mixibius ornatus Pilato et al., 2001, from which it differs in characteristics of the cu‐ticular sculpture and by having the ventral surface unsculptured.     

Ground reaction forces in vertically ascending beetles and corresponding activity of the claw retractor muscle on smooth and rough substrates

We measured ground reaction forces in fore–aft and normal directions of single hind and front legs in vertically ascending Sagra femorata beetles (Coleoptera, Chrysomelidae) on a smooth and a rough substrate. Simultaneously, we performed electromyographic recordings (EMGs) of the hind leg claw retractor muscle that partly controls the attachment structures. On both substrates, hind legs produced upward- as well as downward-directed forces during one stance phase. Forces were equivalent in both directions. Front legs generated only upward-directed forces. The main function of hind legs in ascending beetles in the second half of the stance thus probably prevented the animals from tilting away from the substrate. The EMGs of hind legs showed an early spike during stance with large amplitude. It was mostly followed by few additional spikes with large amplitude and in some cases of spikes with smaller amplitude distributed throughout the stance phase. We found significantly more spikes on the rough substrate than on the smooth one. This is probably due to the more important role of pretarsal claws than tarsal hairy attachment pads on the rough substrate or to the reduced adhesive forces on the rough substrate that have to be compensated by additional muscle activity.     

Micropacteridae, a new family of Remipedia (Crustacea) from the Turks and Caicos Islands

Recent diving explorations of anchialine caves on the Turks and Caicos Islands yielded a rather small and slender new species of Remipedia. Micropacter yagerae n. gen., n. sp. is distinguished from all other species of nectiopod remipedes by a number of autapomorphic characters, including an oval body terminus with fused segments, unequal pairs of terminal claws on maxilla and maxilliped, an almost complete reduction of sternal bars and pleurotergites, molar processes with relatively few, but strong spines, and frontal filaments with bifurcate processes. Based on the unique combination of derived and primitive characters, we propose to erect a new family, Micropacteridae, for this new species and genus of Remipedia. Taxonomic diagnoses for the class Remipedia, order Nectiopoda (emended due to discovery that the maxilliped is 9-segmented), and for the families Speleonectidae and Godzilliidae are presented and discussed.     

Distal leg structures of the Aculeata (Hymenoptera): A comparative evolutionary study of Sceliphron (Sphecidae) and Formica (Formicidae)

The distal parts of the legs of Sceliphron caementarium (Sphecidae) and Formica rufa (Formicidae) are documented and discussed with respect to phylogenetic and functional aspects. The prolegs of Hymenoptera offer an array of evolutionary novelties, mainly linked with two functional syndromes, walking efficiently on different substrates and cleaning the body surface. The protibial-probasitarsomeral cleaning device is almost always well-developed. A complex evolutionary innovation is a triple set of tarsal and pretarsal attachment devices, including tarsal plantulae, probasitarsomeral spatulate setae, and an arolium with an internal spring-like arcus, a dorsal manubrium, and a ventral planta. The probasitarsal adhesive sole and a complex arolium are almost always preserved, whereas the plantulae are often missing. Sceliphron has retained most hymenopteran ground plan features of the legs, and also Formica, even though the adhesive apparatus of Formicidae shows some modifications, likely linked to ground-oriented habits of most ants. Plantulae are always absent in extant ants, and the arolium is often reduced in size, and sometimes vestigial. The arolium contains resilin in both examined species. Additionally, resilin enriched regions are also present in the antenna cleaners of both species, although they differ in which of the involved structures is more flexible, the calcar in Sceliphron and the basitarsal comb in Formica. Functionally, the hymenopteran distal leg combines (a) interlocking mechanisms (claws, spine-like setae) and (b) adhesion mechanisms (plantulae, arolium). On rough substrate, claws and spine-like setae interlock with asperities and secure a firm grip, whereas the unfolding arolium generates adhesive contact on smooth surfaces. Differences of the folded arolium of Sceliphron and Formica probably correlate with differences in the mechanism of folding/unfolding.     

Variation in the geometry of foreleg claws in sympatric giant water bug species: an adaptive trait for catching prey?

When giant water bugs (Heteroptera: Belostomatidae) encounter prey animals that are larger than they are themselves, they first hook the claw of their raptorial legs onto the animal, and then use all their legs to pin it. The claws of the raptorial legs in giant water bugs play an important role in catching larger prey, but the relationship between the claws, body lengths of predators, and prey size has not been fully investigated. To elucidate the functioning of claws in catching prey, we investigated prey body size relative to predator size in nymphs of two sympatric belostomatid giant water bug species, the vertebrate eater Kirkaldyia (=Lethocerus) deyrolli Vuillefroy and the invertebrate eater Appasus japonicus Vuillefroy, captured in rice fields. The younger nymphs of K. deyrolli caught preys that were larger than themselves, whereas those of A. japonicus caught preys that were smaller. Younger nymphs of K. deyrolli had claws that were curved more sharply than those of A. japonicus. The more curved claws of younger nymphs of K. deyrolli probably hook more easily onto larger vertebrates and thus this shape represents an adaptation for acquiring such prey.     

The D1 family dopamine receptor,DopR, potentiates hind leg grooming behavior in Drosophila

Drosophila groom away debris and pathogens from the body using their legs in a stereotyped sequence of innate motor behaviors. Here, we investigated one aspect of the grooming repertoire by characterizing the D1 family dopamine receptor, DopR. Removal of DopR results in decreased hind leg grooming, as substantiated by quantitation of dye remaining on mutant and RNAi animals vs. controls and direct scoring of behavioral events. These data are also supported by pharmacological results that D1 receptor agonists fail to potentiate grooming behaviors in headless DopR flies. DopR protein is broadly expressed in the neuropil of the thoracic ganglion and overlaps with TH‐positive dopaminergic neurons. Broad neuronal expression of dopamine receptor in mutant animals restored normal grooming behaviors. These data provide evidence for the role of DopR in potentiating hind leg grooming behaviors in the thoracic ganglion of adult Drosophila. This is a remarkable juxtaposition to the considerable role of D1 family dopamine receptors in rodent grooming, and future investigations of evolutionary relationships of circuitry may be warranted.     

Postcranial morphology,functional adaptations and palaeobiology of Callistoe vincei,a predaceous metatherian from the Eocene of Salta,north‐western Argentina

Abstract: Callistoe vincei Babot et al., 2002 is a Paleogene borhyaenoid known from exceptionally complete postcranial elements, which provides rare information about the anatomy and evolutionary history of metatherian predators during the South American Cenozoic. The axial skeleton of Callistoe is characterized by the peculiar transverse processes of the cervical vertebrae emphasizing lateral instead of sagittal traction. There is no clavicle and eighteen thoracolumbar vertebrae, of which only five are lumbars. The shoulder and elbow joints suggest movements restricted to parasagittal flexion/extension that are consistent with primarily terrestrial locomotion, as is also emphasized in Borhyaena tuberata and Lycopsis longirostrus. On the manus, the pollex is not reduced and the ungual phalanges indicate very long claws, similar to those observed in some extant digging taxa. This feature is unique to C. vincei among borhyaenoids. The knee joint is characterized by the presence of ossified patellae but shallow femoral trochleae. This joint suggests that the leg was nearly parasagittal, a position also inferred for Borhyaena. The astragalus shape is consistent with parasagittal flexion/extension, as in all Miocene–Pliocene borhyaenoids. The hind foot is characterized by reduced claws in comparison with the manus as well as the slenderness of the first and fifth digits, another peculiarity of C. vincei. The habitat of Callistoe was a temperate humid forest and according to the known fossil record, Callistoe was the largest mammalian predator of its time, sharing the predator ecological niche with crocodiles.     

Regulation of sex-specific feeding behavior in fiddler crabs: physiological properties of chemoreceptor neurons in claws and legs of males and females

This study examined properties of chemoreceptor neurons in the claws and legs of the fiddler crabs Uca pugilator and U. pugnax. The primary goal was to establish the neural basis of previously observed greater female sensitivity to feeding stimulants, and secondarily to compare physiological properties of chemoreceptor neurons in these semi-terrestrial crustaceans with those of fully aquatic forms. Sensitivity of chemoreceptor neurons in claws and legs is sex-specific; individual neurons of females respond to lower stimulus concentrations than male chemoreceptor neurons, and equivalent concentrations elicit greater spiking in female vs male chemoreceptor neurons. Thus, the population of chemoreceptor neurons in females expresses lower thresholds and greater average sensitivity than in males. Greater sensitivity of claw neurons explains observations indicating that females continue to feed at food levels too low to stimulate males. Sensitivity differences in leg neurons of males vs females have no clear behavioral correlate, but suggest that females can orient to more dilute stimuli than males. Chemoreceptor neurons of fiddler crabs have low sensitivities and slow rates of adaptation compared to other crustaceans. Also, neurons in claws adapt less slowly than neurons in legs, which may reflect subtle differences in the chemical stimulus environment experienced by claws vs legs.     

Osteochondrosis / Osteoarthrosis and Claw Disorders in Sows,Associated with Leg Weakness

The objective of this study was to investigate the associations between different leg weakness symptoms and osteochondrosis/osteoarthrosis and claw disorders in sows together with the influence of age on these findings. One hundred and seventeen sows in one herd were followed from 6 months of age until culling and judged for leg weakness once in every gestation using a scale from 1 (normal) to 4 (severe changes). At slaughter changes in joints, growth plates and claws were scored on a scale from 1 (normal) to 5 (very severe changes). Osteoarthrotic changes were strongly associated with osteochondral changes in humeral and femoral condyles. The clinical signs of osteochondrosis and osteoarthrosis were found to be: buck-kneed forelegs, turn out of fore and hind legs, upright pasterns on hind legs, stiff locomotion, lameness and tendency to slip. The clinical signs of claw lesions were found to be: buck-kneed forelegs, upright pasterns, steep hock joints, turn out of hind legs, standing under position on hind legs, stiff movements, swaying hindquarters, goose-stepping hind legs, tendency to slip and lameness. Overgrown claws were strongly associated with leg weakness indicating the need for claw trimming in sow populations.     

Tardigrades from Ecuador,with the Description of Two New Species: Mixibius ornatus n. sp. and Diphascon (Adropion) onorei n. sp. (Eutardigrada,Hypsibiidae)

Seven species are reported from Ecuador, South America, two of which, Mixibius ornatus and Diphascon (Adropion) onorei, are new to science. Mixibius ornatus n. sp. differs from Mixibius saracenus (Pilato 1973) and from M. fueginus Pilato & Binda 1997 by having a sculptured cuticle, broader claws, lunulae present, stylet supports inserted on the buccal tube in a more caudal position, and shorter placoids. Diphascon (A.) onorei n. sp. has an elongate pharyngeal bulb with small apophyses, three rod-shaped macroplacoids, a granular microplacoid, and septulum; claw bases enlarged, smooth (first three pairs of legs) or crenate (outer claws on the hind legs); a cuticular bar is present near the inner claws on the first three pairs of legs. Other species reported are Echiniscus aliquantillus Grigarick, Schuster & Nelson 1983; Minibiotus intermedius (Plate 1888); Macrobiotus coronatus De Barros 1942; Macrobiotus danielae Pilato, Binda, Napolitano & Moncada 2001, and Astatumen bartosi Weglarska 1959.     

Latitudinal variation and coevolutionary diversification of sexually dimorphic traits in the false blister beetle Oedemera sexualis

Sexual traits are subject to evolutionary forces that maximize reproductive benefits and minimize survival costs, both of which can depend on environmental conditions. Latitude explains substantial variation in environmental conditions. However, little is known about the relationship between sexual trait variation and latitude, although body size often correlates with latitude. We examined latitudinal variation in male and female sexual traits in 22 populations of the false blister beetle Oedemera sexualis in the Japanese Archipelago. Males possess massive hind legs that function as a female‐grasping apparatus, while females possess slender hind legs that are used to dislodge mounting males. Morphometric analyses revealed that male and female body size (elytron length), length and width of the hind femur and tibia, and allometric slopes of these four hind leg dimensions differed significantly among populations. Of these, three traits showed latitudinal variation, namely, male hind femur was stouter; female hind tibia was slenderer, and female body was smaller at lower latitudes than at higher latitudes. Hind leg sizes and shapes, as measured by principal component analysis of these four hind leg dimensions in each sex, covaried significantly between sexes, suggesting coevolutionary diversification in sexual traits. Covariation between sexes was weaker when variation in these traits with latitude was removed. These results suggest that coevolutionary diversification between male and female sexual traits is mediated by environmental conditions that vary with latitude.     

Use of forest canopy by European red squirrelsSciurus vulgaris in Northern Greece: claws and the small branch niche

Moving and standing in trees impose multiple problems to arboreal mammals. Among them, the major ones are the negotiation of slender terminal branches and of large vertical supports. Both microhabitats are important as they have been linked alternatively to the evolutionary loss of claws in early primates. Therefore, rates of use of these different supports by claw-bearing arboreal mammals may offer insights to their actual significance in the adaptive evolution of early primates. In this context, canopy, tree crown, branch size, inclination, and texture use were recorded on four adult free ranging European red squirrelsSciurus vulgaris Linnaeus, 1758 in a mixed coniferous forest in northern Greece.S. vulgaris was mainly arboreal, exploiting the terminal branch zone, using frequently oblique and intermediately textured supports<5 cm and moderately large vertical branches. Furthermore, comparative data from other sciurid species and clawed primates showed positive correlations of small and horizontal support use, and negative ones of vertical support use to body mass. These findings show that keeled functional claws do not impede habitual use of slender branches and may not facilitate efficient climbing on large vertical trunks. These observations partly question the association between habitual use of the small branch niche and primate adaptations and lend support to alternative hypotheses, underscoring the importance of inquiring for more complex mechanisms that lead to the evolution of the unique set of primate morphological adaptations.     

Achilaophonte maxima gen.n., spec.n., a new taxon of the Laophontidae (Copepoda,harpacticoida) from the high Antartic (Weddell Sea)

Male and female of a new genus and species of the family Laophontidae, Archilaophonte maxima, are described. The specimen was found in the high Antartic (Weddell Sea) and apprears to be the most primitive genus up to now within the superfamily Laophontoidea as defined by Huys (1990). Based on its setation of legs and mouth parts, however, it can be placed unequivocally into the family Laophontidae. Archilaophonte maxima gen. n. shows close affinities to the laophontid genus Esola Edwards 1891. Both genera form a monophyletic group which is interpreted here as the first and most primitive offshot in the evolution of the Laophontidae. The synapomorphies of the former lineage are the shape of the protopodite of the P1 and shape and setation of the female P5.     

The Anatomy of the Tarsi of Schistocerca gregaria Forskål

Summary The tarsus of S. gregaria is divided into three units (here called segments) and an arolium set between a pair of claws. The first segment bears three pairs of pulvilli in the fore and middle legs, and one pair and two single pulvilli in the hind legs. Segment two bears a pair of pulvilli, segment three one long pulvillus and the arolium a similar pad on the undersurface. The outer layers of the arolium pad differ from those of the pulvilli in possibly lacking an epicuticle and in having a layer of cuticle which, unlike the corresponding layer in the pulvilli, does not stain with protein stains. The claws and dorsal surfaces bear trichoid sensilla, basiconic sensilla and campaniform sensilla. Smaller basiconic sensilla and canal sensilla occur on the proximal part of the pulvilli, and basiconic sensilla on the arolium undersurface. Internally the cuticle is modified in the arolium and pulvilli so that rods of probably chitin and resilin are formed. This would impart flexibility to the undersurfaces whilst retaining some degree of rigidity which might prevent damage to the small and delicate sense organs on the pulvilli. The tip of the arolium is specialised for adhesion, and there are two large neurones internally which could conceivably monitor attachment or detachment of the tip. There are chordotonal organs in segment three, and several other large neurones throughout the tarsus, some of which are associated with the slings of tissue holding the apodeme in a ventral position. Gland cells occurring in the dorsal epidermis of the adult mature male are also briefly described.     

A new genus and species of demodecid mites from the tongue of a house mouse Mus musculus: description of adult and immature stages with data on parasitism

The study of the parasitofauna of the house mouse Mus musculus (Rodentia: Muridae) Linnaeus is particularly important owing to its multiple relationships with humans – as a cosmopolitan, synanthropic rodent, bred for pets, food for other animals or laboratory animal. This article proposes and describes a new genus and species of the parasitic mite based on adult and immature stages from the house mouse. Glossicodex musculi gen. n., sp. n. is a medium‐sized demodecid mite (adult stages on average 199 µm in length) found in mouse tissue of the tongue. It is characterized by two large, hooked claws on each tarsus of the legs; the legs are relatively massive, consisting of large, non‐overlapping segments. The palps consist of three slender, clearly separated, relatively narrow segments, wherein their coxal segments are also quite narrow and spaced. Also, segments of the palps of larva and nymphs are clearly isolated, and on the terminal segment, trident claws that resemble legs' claws can be found. On the ventral side, in immature stages, triangular scuta, topped with sclerotized spur, can be also observed. Glossicodex musculi was noted in 10.8% of mice with a mean infection intensity of 2.2 parasites per host.     

Pincer-like claws in centipedes (Chilopoda): multiple evolutionary origin of similar form and serial pattern

In most Chilopoda, the walking legs end in a single-tip claw usually accompanied by short accessory spines. Instead, in all species of three small and only distantly related geophilomorph taxa (Diphyonyx, Neogeophilidae, Eucratonyx), the claws of an anterior set of leg pairs are unusually pincer-like. By integrating different microscopic techniques, including confocal laser scanning microscopy, we found that these modified claws are very similar in form, internal structure, and pattern of variation in shape along the trunk in all three taxa: the claws are distinctly swollen and bent, provided with peculiar bulges, and flanked by a conspicuous additional branch, either cylindrical or flattened, which overreaches the tip of the claw; instead, the internal cuticular features are not modified with respect to the condition in the other centipedes, claiming against the possibility of controlled abduction/adduction between claw and branch. Irrespective of the total number of leg pairs (63–129), the claws change gradually from pincer-like to usual shape invariantly in the range spanning between the 34 and the 45% of the total number of leg pairs. Despite these similarities, pincer-like claws originated independently in the three taxa, and by way of fundamentally different changes, either by the dramatic modification of the already existent anterior accessory spine (Diphyonyx, Neogeophilidae) or by the production of a novel cuticular projection (Eucratonyx). Moreover, their shared pattern of variation along the body was most probably constrained by already operating developmental processes controlling the longitudinal patterning of the trunk.     

A new genus and species of Renaudarctidae (Heterotardigrada: Arthrotardigrada) from Ryukyu Archipelago,Japan

A new armoured marine tardigrade, Nodarctus hallucis gen. et sp. nov., is described from specimens collected from sandy beaches of Haterumajima, Ishigakijima and Okinawajima Islands, Ryukyu Archipelago, Japan. It is a member of the rare family Renaudarctidae, which is characterized by a stocky body with dorsal and ventral plates and stumpy legs with digits, claws and accessory filaments. However, the collected animals have prominent characters that differ from the two known species of the family: subdivided dorsal segmental plates I–III lacking lateral processes, an acute caudal spike, indented sensory plates, short spikes and peculiar papillae on legs I–III, basal spikes of legs IV and only two digits on legs IV. These differences justify the erection of a new genus and species for the animal. Morphological affinities with other marine taxa, the function of the leg papillae and the distribution pattern of the new species are discussed.http://zoobank.org/urn:lsid:zoobank.org:pub:041878AA-B437-45BF-80E3-32591D21D965     

The tale of the shrinking weapon: seasonal changes in nutrition affect weapon size and sexual dimorphism,but not contemporary evolution

Sexually selected traits are often highly variable in size within populations due to their close link with the physical condition of individuals. Nutrition has a large impact on physical condition, and thus, any seasonal changes in nutritional quality are predicted to alter the average size of sexually selected traits as well as the degree of sexual dimorphism in populations. However, although traits affected by mate choice are well studied, we have a surprising lack of knowledge of how natural variation in nutrition affects the expression of sexually selected weapons and sexual dimorphism. Further, few studies explicitly test for differences in the heritability and mean‐scaled evolvability of sexually selected traits across conditions. We studied Narnia femorata (Hemiptera: Coreidae), an insect where males use their hind legs as weapons and the femurs are enlarged, to understand the extent to which weapon expression, sexual dimorphism and evolvability change across the actual range of nutrition available in the wild. We found that insects raised on a poor diet (cactus without fruit) are nearly monomorphic, whereas those raised on a high‐quality diet (cactus with ripe fruit) are distinctly sexually dimorphic via the expression of large hind leg weapons in males. Contrary to our expectations, we found little evidence of a potential for evolutionary change for any trait measured. Thus, although we show weapons are highly condition dependent, and changes in weapon expression and dimorphism could alter evolutionary dynamics, our populations are unlikely to experience further evolutionary changes under current conditions.     

The mechanics of elevation control in locust jumping

How do animals control the trajectory of ballistic motions like jumping? Targeted jumps by a locust, which are powered by a rapid extension of the tibiae of both hind legs, require control of the take-off angle and speed. To determine how the locust controls these parameters, we used high speed images of jumps and mechanical analysis to reach three conclusions: (1) the extensor tibiae muscle applies equal and opposite torques to the femur and tibia, which ensures that tibial extension accelerates the centre of mass of the body along a straight line; (2) this line is parallel to a line drawn from the distal end of the tibia through the proximal end of the femur; (3) the slope of this line (the angle of elevation) is not affected if the two hind legs extend asynchronously. The mechanics thus uncouple the control of elevation and speed, allowing simplified and independent control mechanisms. Jump elevation is controlled mechanically by the initial positions of the hind legs and jump speed is determined by the energy stored within their elastic processes, which allows us to then propose which proprioceptors are involved in controlling these quantities.     

A remarkable new genus of Atalophlebiinae (Ephemeroptera: Leptophlebiidae) from the Neotropics

An unusual species of Leptophlebiidae is described based on males, females, and nymphs from Rio de Janeiro State, Brazil. As a consequence of its distinct characteristics on all stages, a new genus is established. The new genus can be distinguished from other South American Leptophlebiid genera mostly by: Adults: vein MA of fore wings asymmetrical; hind wings with costal projection well developed, Sc ending at cross vein near costal projection; tarsal claws dissimilar, one blunt other acute; projections of styliger plate forming two well developed lobes with rounded apex, ventrally obstructing view of the penes; penes fused on basal half, each lobe with a ventral furrow and a long and slender spine directed anteriorly. Nymphs: Head prognathous, wider than labrum; labrum with prominent median emargination, with three subtle crenulations; body flattened; hind wings pads present; tarsal claws with over 20 denticles, subapical denticle much larger than remaining denticles; gills long and narrow, present on abdominal segments I-VII; posterolateral projections present on abdominal segments VIII-IX. Phylogenetic analyses conducted based on a previously published data matrix that included other South America leptophlebiid genera placed Poranga nessimiani gen. nov. et sp. nov. as sister to Bessierus + Perissophlebioides. In most analyses this clade was recovered within the Farrodes complex. Nymphs of the new taxa are particularly similar to Bessierus, whereas adults share the very acute costal projection on hind wings seen in non-dipterous members of the Farrodes complex.