Digestion time and reemergence in the desert grassland scorpion Paruroctonus utahensis (Williams) (Scorpionida,Vaejovidae)

Summary The desert grassland scorpion Paruroctonus utahensis spends most of its life in its burrow. During the active season, only about 5% of the individuals in a population appear on the surface each night. Individuals do not appear on the surface for several nights following a meal. To determine if physiological digestion time could account for this delay in reemergence after eating, I measured changes in oxygen consumption immediately following a meal. Oxygen consumption exceeded 125 l O₂g^-1h^-1 just after completion of a meal, then dropped to normal levels (53 l O₂g^-1h^-1) within 6 h. I also measured the interval between completion of the meal and subsequent defecation. All individuals defecated by 72 h following ingestion (median 12 h). In field enclosures, scorpions returned to the surface after a mean of 20.3 days (median=16) following a successful predation event. Lack of correspondence between estimates of physiological digestion time and the reappearance interval lead me to reject the idea of a long digestive pause in Paruroctonus utahensis. This conclusion lends support to the hypothesis that scorpions remain in their burrows to minimize exposure to predation.     

Water detection in the desert sand scorpion,Paruroctonus mesaensis (Scorpionida,Vaejovidae)

Summary For the sand scorpion, Paruroctonus mesaensis, substrate moisture is a powerful and fast-acting stimulus of discrete behaviors related to localization and imbibitory uptake of water. These behaviors are readily observed in the field and quantified in the laboratory when free-roaming animals encounter sand substrates dampened by small amounts of water. Of 10 behaviors we monitored in laboratory tests, 5 (pedipalp-pull, rototiller-digging, prolonged stops, headstand, and backingup) occurred only after contact with a moistened substrate. These water-stimulated behaviors were selectively blocked when all 8 tarsal leg segments were coated with wax; coverings of the chemosensory pectine appendages had little to no effect. Electrophysiological recordings from chemoreceptor organs on the tarsi showed that neurons innervating the dorsal tarsal organ, were highly sensitive to humid air stimuli while the numerous, poretipped hairs on the ventral surface were responsive to aqueous solutions applied directly to their tips. Selective blocking of the 8 tarsal organs had no effect on water sensitive behavior indicating that the chemosensory hairs mediate detection of substrate moisture. Such localized, sensory triggering of a robust and directed behavior presents a useful model for further neuroethological studies.Abbreviations H headstand - PP pedipalp-pull - RD rototillerdigging - B backing-up - C wall-climbing - P pause - W walk-through - S stop - T turn - R rest - PST pacific standard time - SEM scanning electron microscopy     

Characteristics and environmental determinants of natality, growth and maturity in a natural population of the desert scorpion, Paruroctonus mesaensis (Scorpionida: Vaejovidae)

The scorpion, Paruroctonus mesaensis (Vaejovidae) has seven instar stages and moults six times. There is no post-reproductive moult. The average growth ratio between successive instars is 1–38. The average rate of weight increase is 000249/day. This rate fluctuates seasonally: it is highest in spring and lowest in winter and summer. Growth rate is a function of ambient burrow temperature and prey abundance. Body size is determinate. The growth rate of the pectines is significantly greater in males as compared with females. Analysis indicates that the pectines are a secondary sexual trait in males.
Animals mature from 19 to 24 months of age. Matings were observed from May through October. Gestation lasts 10–14 months. First surface appearance of newborn (instar II) is synchronous throughout the population and occurred in early August for five consecutive years. Some females were observed to be gravid for at least three consecutive years. Gravid females carry an average of 48–8 embryos. However, on the average only 19–9 young/female survive to appear on the surface. Factors which may be responsible for this observed decrease include resorption of embryos; death at birth, eclosion or first moult; and maternal and sibling cannibalism. The influence of various environmental factors on reproduction were analyzed. The number of newborn appearing on the surface and the population biomass of newborn showed a high positive correlation with precipitation and vegetation cover.     

The ultrastructure of hemocytopoietic organs in the desert scorpion, Paruroctonus

The light and electron microscopes were used to examine possible hemocytopoietic tissue in the desert scorpion, Paruroctonus mesaensis. Results agree with earlier light microscopic studies that cells are released into the blood from the two lateral lymphoid organs and the supraneural gland. The former are sacciform structures attached by their anterior ends to the diaphragm. The supraneural gland forms the thickened wall of the supraneural artery in the mesosoma from the first to the third abdominal ganglia. The lateral lymphoid glands have an acellular stroma in which are embedded granular and agranular cells. The stroma is apparently formed by specialized cells which release membranous cell fragments that become the matrix of the gland. Cells are released into the body cavity from the periphery of the two organs. The supraneural gland has a fibrous stroma in which are embedded a variety of cell types. The cells appear to be released in greatest abundance into the blood in the lumen of the gland. The gland has cells with opaque granules (0.9-1.4 micron diameter) and agranular cells of variable shape. The most abundant cell, possibly the stem-cell for the others, is about 10 micron diameter and often has processes of variable length. In addition, muscle cells at various stages of differentiation are found at the inner margin of the gland. These cells have thick and thin myofilaments (24-32 and 5-8 nm diameter) and dense bodies which sometimes become organized into sarcomeres with Z-bands before the cells are released into the gland lumen. The function of these muscle cells is unknown, but possibly they contribute to the maintenance of blood pressure and the release of cells into the blood from the inner margin of the gland.     

Friend or foe: behavioral responses to conspecifics in the northern scorpion, <Emphasis Type="Italic">Paruroctonus boreus</Emphasis> (Scorpionida: Vaejovidae)

We investigated the behavioral response of male Paruroctonus boreus to male and female pheromones of conspecifics. Adult male scorpions actively search for females during the mating season, but lack visual cues to direct their movement. Chemical cues (pheromones) are thought to be responsible for directing movement. In a series of laboratory experiments, in Y-mazes, males were exposed to areas that had contained males and females from the same population and areas that had contained conspecifics from different populations. Males spent significantly more time in areas that had contained females from the same population but did not show a preference for areas exposed to females from different populations. Males also significantly avoided areas exposed to males from the same population. Our results suggest that P. boreus has a sex-specific response to pheromones and that this response may be population-specific.     

Prey and feeding phenology of the desert sand scorpion Pamroctonus mesaensis (Scorpionidae: Vaejovidae)

The diet and feeding of the vaejovid scorpion Pamroctonus mesaensis Stahnke was investigated during a five-year study. Foraging and feeding behaviour are described. This scorpion is a "sit and wait" predator that eats a wide variety of cursorial, fossorial and aerial prey. Ninety-five prey species were recorded. Three year classes of scorpions exist. Each class captures a different proportion of major prey species and a significantly different average size of prey.
The proportion of each major taxon of prey in the diet is as follows: tenebrionid beetles (42%), Orthoptera (17%), other scorpions (16%) and Hymenoptera (12%). When analysed in relation to biomass, larger prey species assume more importance while smaller species are relatively less important. Although P. mesaensis ranks as the fourth most numerous prey species, they represent the most important diet item in terms of ingested biomass.
When observed, 3·75% of all scorpions were feeding. This percentage varied seasonally from a high in spring (7.0%) to a low in November and December (0.5 %). The proportions of major prey taxa in the diet also varied seasonally. Prey species were classified into three phenological categories: pulsed, seasonal and annual. Examples of each are given.     

Evidence of Chemical Signaling in the Sand Scorpion,Paruroctonus mesaensis (Scorpionida: Vaejovida)

This study presents evidence of intraspecific chemical communication in scorpions. The subject of our investigation was the nocturnal sand scorpion, Paruroctonus mesaensis. During the mating season, mature males show a sex-specific wandering behavior ostensibly directed at locating conspecific females that remain in the vicinity of their home burrows. Searching behavior was stimulated in the laboratory by releasing males onto substrates that had previously been occupied by females. Receptive males exhibited changes in locomotory behavior that favored occupation of the female-exposed area. Males occasionally displayed a precourtship behavior, called juddering, indicating the presence of a pheromone on the substrate. Juddering, and two newly described behavior patterns, tail-wagging and pedipalp-reaching, were also induced by solvent extracts of female cuticle. Most behavioral responses began vigorously within the first few s of stimulus contact and gradually adapted within 10 min. The potential importance of specialized sensory appendages, the pectines, for mediating chemosensitivity is discussed. These results are consistent with the hypothesis that mate identification and localization in sand scorpions are mediated in part by a contact sex pheromone.     

Sexual dimorphism in desiccation responses of the sand scorpion Smeringurus mesaensis (Vaejovidae)

The osmoregulatory and respiratory responses of male and female Smeringurus mesaensis (Vaejovidae) to prolonged desiccation were measured. No significant effect of sex on mass-loss rates (MLRs) was found. Still, females maintained their haemolymph osmolality when desiccated to 10% mass loss, whereas that of males increased significantly after loss of as little as 5% of initial mass. Females had a 3-fold larger hepatopancreas, significantly higher hepatopancreas water content and higher metabolic rates when adjusted to hepatopancreas-free dry mass. Thus, females not only store more water in the hepatopancreas but also mobilise it to the haemolymph at a higher rate during desiccation, thus maintaining haemolymph osmolality. Gas exchange rates of both males and females decrease as desiccation progresses. An initial respiratory exchange ratio (RER) of approximately 0.9 is followed by a significant increase at mass loss levels of 7.5% and higher. RER values greater than 1.0 may result from partial shift to anaerobic catabolism, which allows closure of the book lung spiracles for longer duration, thus minimising respiratory water loss. The effects of gas exchange rates on rates of water mobilisation between body compartments and water loss to the environment suggest a trade-off between maintaining osmotic stability and conserving body water stores under stressful conditions.     

A different oogenetic pattern in a US apoikogenous scorpion, Vaejovis spinigerus (Scorpiones, Vaejovidae)

The ovariuterus of the scorpion Vaejovis spinigerus (Scorpiones, Vaejovidae) differs from that of all other apoikogenic species studied so far in that the oocytes mature inside, rather than on the outer surface, of the ovariuterus tubes. Only the small, second and third oocyte generations are situated on the outer surface, sometimes even on top, of the mature oocytes. This oogenetic pattern has so far not been described in any other scorpion species. Since no males were found in the populations studied here, this species can be assumed to be parthenogenetic. However, this point needs more concrete proof.     

Wernerius inyoensis, an elusive new scorpion from the Inyo Mountains of California (Scorpiones, Vaejovidae)

A new scorpion species is described from the Inyo Mountains of California (USA). The presence of a strong subaculear spine, along with other characters, places the new species within Wernerius, an incredibly rare genus that until now consisted of only two species. Wernerius inyoensissp. n. can be most easily distinguished from the other members of the genus by smaller adult size, femur and pedipalp dimensions, and differences in hemispermatophore morphology. Previous studies have suggested that the elusive nature of this genus may be attributed to low densities and sporadic surface activity. Herein, we provide another hypothesis, that Wernerius are primarily subterranean. Mitochondrial sequence data are provided for the holotype.     

Negative influence of Gammarinema gammari (Nematoda) on the fecundity of Microphallus papillorobustus (Trematoda): field and experimental evidence

The gammarid amphipod Gammarus insensibilis frequently harbors adult individuals of the ectoparasitic nematode Gammarinema gammari as well as metacercariae of the trematode Microphallus papillorobostus. After the demonstration in a previous study of a negative relationship between the abundance of these 2 parasites, the nature of the relationship between these 2 parasites was explored in more detail by studying, in the field and in the laboratory, the influence of nematode abundance on trematode fecundity. In gammarids collected in the field, a negative relationship between metacercarial fecundity and the number of co-occurring nematodes was found. By manipulating the nematode abundance in the laboratory, it was confirmed that G. gammari has a negative effect on egg production in M. papillorobustus.     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Detection of vibrations in sand by tarsal sense organs of the nocturnal scorpion,Paruroctonus mesaensis

Summary The scorpionParuroctonus mesaensis locates prey by orienting to substrate vibrations produced by movements of the prey in sand. At the end of each walking leg of this scorpion there are two sense organs, the basitarsal compound slit sensillum and tarsal sensory hairs (Figs. 1, 3) that are excited by substrate vibrations conducted through sand. The slit sensilla appear to be most sensitive to surface (Rayleigh) waves while the tarsal sensory hairs respond best to compressional waves (Fig. 7). Both mechanoreceptors were activated by nearby disturbances of the substrate (Fig. 6) but only the slit sensilla responded to insects moving more than 15 cm away. Both receptors are highly sensitive to small amplitude (less than 10 Å) mechanical stimuli applied to the tarsus (Fig. 5).Behavioral studies of scorpions with ablated sense organs (Fig. 2) indicate that the basitarsal compound slit sensilla are necessary for determining vibration source direction.Abbreviation BCSS basitarsal compound slit sensillum (a) Supported by PHS Environmental Science and Regents Intern Fellowships (PB), and by intramural research funds from the University of California (RDF)     

The endogenous stages of Eimeria utahensis (Protozoa: Eimeriidae) in the kangaroo rat, Dipodomys ordii

The endogenous life cycle of Eimeria utahensis is described from experimentally infected kangaroo rats, Dipodomys ordii. The endogenous asexual cycle consisted of 4 generations of meronts. First-generation meronts were concentrated in the anterior third of the small intestine. The succeeding generations of meronts and the sexual stages were concentrated in the middle third of the small intestine. First-generation meronts had a mean diameter of 9.7 micrometer and contained 12 to 16 merozoites. Second-generation meronts had a mean diameter of 8.0 micrometer and contained 12 to 16 merozoites and a residual body. Third-generation meronts had a mean diameter of 12.4 micrometer and contained 4 to 8 merozoites. Fourth-generation meronts had a mean diameter of 8.6 micrometer and contained 16 to 24 merozoites. Young gamonts were located in epithelial cells of the crypts of the small intestine. Shortly after the parasites entered the epithelial cells, the infected cells became displaced into the lamina propria, and most of the mature gamonts were in this location. The nuclei of host cells containing young sexual stages became greatly elongated and flattened. A few young gamonts were seen in cells in which the host cell nuclei were dividing. During development, nuclei of microgamonts became arranged on the periphery of numerous compartments. Only one type of wall-forming body could be distinguished in the macrogamonts.     

The influence of the food size,distance and food site on food carrying behavior in rats (Rattus norvegicus)

The behavior to foods in rats (Rattus norvegicus) was investigated in a seminatural enclosure and in a straight alley in the laboratory. In both situations, the size of food (0.045-, 1-, and 3-g pellets were used in the enclosure and 0.045-, 0.2-, 1-, and 3-g pellets in the alley) and distance between food sites and nest site (1.8, 3.6, 5.4, and 7.2 m in the enclosure and 2, 4, and 6 m in the alley) were varied. In both situations, when food was presented in only 1 place, the food carrying behavior increased as the size of food increased from 0.045 to 1-g. One and three g pellets were always carried, regardless of the distance to nest. On the other hand, when multiple food sites were presented in the alley, rats were more likely to carry small pellet from the further food site, whereas large pellet was always carried and were not influenced by the distance. The behavior to foods in the enclosure, however, was not influenced in either food site condition. This might be due to the difference in familiarity to the environment and anxiety from perceived predation risk.     

Partitioning of transpiratory water loss of the desert scorpion, Hadrurus arizonensis (Iuridae)

Terrestrial arthropods lose body water to the environment mainly through transpiration. The aim of this study was to determine the fraction of respiratory losses from total transpiratory water loss in scorpions, as relatively high respiratory losses would indicate a fitness benefit from regulation of gas-exchange rate under stressful desiccating conditions. We measured metabolic rates and water-loss rates of Hadrurus arizonensis (Iuridae) at a range of ecologically-relevant temperatures. Calculation of respiratory water losses was based on increased metabolic and water-loss rates during nocturnal activity (assuming no change in cuticular resistance at a given constant experimental temperature). Respiratory losses accounted for 9.0 ± 1.7% of total transpiratory losses at 25 °C, doubled to 17.9 ± 1.8% at 30 °C and increased to 31.0 ± 2.0% at 35 °C (n = 5, 15 and 15, respectively). Furthermore, the relative importance of respiratory transpiration is likely to be higher at temperatures above 35 °C, which have been recorded even within the burrows of H. arizonensis. Measurements of cuticular lipid melting points do not provide evidence for increased cuticular resistance to water loss at higher temperatures. However, the relatively high fraction of respiratory water losses reported here for H. arizonensis supports the notion of respiratory regulation as an evolved mechanism for conserving scorpion body water stores under stressful conditions.     

An experimental test of frequency-dependent food selection: winter browsing by moose

Penned moose Alces alces (L.) offered birch and rowan trees in various proportions in experimental tests showed a clear preference for birch both in terms of selection and use of trees. Both tree selection and tree use was frequency-independent, i.e. the preference for birches did not vary with either the number of birches, or total birch biomass available. The preference for birch was not absolute, i.e. rowans were also present in the diet. Birch were more profitable than rowans in terms of biomass intake rate, whereas rowans were more nutritious than birches. The results are consistent with both the optimal diet theory and the nutrient mixture hypothesis, although neither of the hypotheses are unambiguous regarding frequency-dependent food selection. It is argued, however, that tree selection and use by moose rather should be considered as a patch use problem and that trees should not be regarded as particulate food items as in most foraging models.