Seasonal abundance of Allothrombium monochaetum and Allothrombium pulvinum in Navarra-Nafarroa (northern Spain), with notes on larval host preference and rate of parasitism

In this paper we report on the seasonal abundance of velvet mite larvae of Allothrombium pulvinum Ewing and Allothrombium monochaetum Goldarazena and Zhang in a meadow located in the mediterranean Navarra-Nafarroa region (northern Spain). Larvae of A. monochaetum peaked in numbers on Aphis fabae in mid-June while larvae of A. pulvinum peaked in the third week of July. Both species preferred the aphid thorax as attachment site. We also provide a list of aphid hosts of both mites and observed density of parasites on the hosts.     

Functional response ofAllothrombium pulvinum deutonymphs (Acari: Trombidiidae) on two-spotted spider mites (Acari: Tetranychidae)

Functional responses of deutonymphs of the predatory mite,Allothrombium pulvinum Ewing, on eggs and adult females of two-spotted spider mite,Tetranychus urticae Koch, were determined in the laboratory. Predation experiments were conducted on lima bean leaf discs over a 24-h period at 25±2°C, 30–50% RH and 24L: 0D photoperiod. Prey densities ranged from 10 to 120T. urticae eggs per disc or 2 to 32 adult females per disc.Allothrombium pulvinum deutonymphs were more effective againstT. urticae eggs than its adult females. The role ofA. pulvinum deutonymphs in integrated and biological pest control is discussed.     

Effects of gravity on positive phototaxis in fruit fly Drosophila melanogaster

Geotaxis and phototaxis are movements in response to gravity and light, respectively, and are commonly observed in nature. The interactions between these two types of movement have been shown to confer ecological advantages to many taxa. Although several studies have been conducted on phototaxis and geotaxis in various organisms, reports on the interactions between positive phototaxis and negative geotaxis are lacking. In the fruit fly, Drosophila melanogaster, any direct interactions that exist between positive phototaxis and negative geotaxis are yet to be determined and the ecological significance of such interactions remains unclear. In the present study, the effects of gravity on positive phototaxis in a Y‐maze were investigated using the Canton‐S wild type and gravity‐sensing‐deficient pyx³ mutant fruit flies. Gravity sensing was not necessary for horizontal positive phototaxis, but was required for vertical positive phototaxis. These results suggest that gravitoreception may selectively modulate positive phototaxis depending on the vertical and horizontal movements of the fruit flies.     

The role of two plant-derived volatiles in the foraging movement of 1st instar Helicoverpa armigera (Hübner): time to stop and smell the flowers

Positive phototaxis and negative geotaxis are behaviours that 1st instar Helicoverpa armigera use to direct their foraging movement upward towards nutritious new plant growth and reproductive structures. Odours emitted by fruits or seeds can attract larvae directly via chemotaxis. In this study we clarify the effect of leaf and flower odours on foraging 1st instar H. armigera. Using a Y-tube olfactometer we tested for chemotaxis towards two plant volatiles and found larvae were not attracted. Bioassays for phototaxis towards UV, blue, green and white light showed that a green leaf volatile ((Z)-3-hexenyl acetate) and a flower volatile (phenylacetaldehyde) reduced larval phototaxis towards blue light. Feeding on a host plant reduced phototaxis towards blue and green light. We concluded that the upward movement of 1st instars on plants is largely due to phototaxis towards the blue wavelengths of skylight. Plant attributes such as volatile chemicals affect the expression of phototaxis and therefore, indirectly influence larval movement to locate food resources. Handling Editor: Anna-Karin Borg-Karlson     

Host preference by Allothrombium pulvinum (Acari: Trombidiidae) larvae on aphids: Macrosiphum rosae,Aphis gossypii and Hyalopterus amygdali (Homoptera: Aphididae)

In this study aphid-plant association and its effect on host preference of parasitic Allothrombium pulvinum larvae was examined with multiple-choice tests. Host species selection, host size selection and superparasitism with mite larvae were studied with two-choice tests. Three aphid species were used: Macrosiphum rosae, Aphis gossypii and Hyalopterus amygdali. In multiple-choice tests, larvae of A. pulvinum showed no significant preference for any aphid-plant association when given M. rosae on rose, A. gossypii on cucumber and H. amygdali on apricot simultaneously. Two-choice tests showed that larval mites preferred H. amygdali to A. gossypii, but had no preference when offered a choice between A. gossypii and M. rosae or between H. amygdali and M. rosae. In host size selection and superparasitism tests, significantly more mites selected the larger host (M. rosae). Furthermore, parasitised H. amygdali were preferred to unparasitised ones. This revised version was published online in July 2006 with corrections to the Cover Date.     

Biology of Allothrombium pulvinum Ewing (Acari: Trombidiidae) in West Mazandran, Iran

Allothrombium pulvinum Ewing is a common natural enemy of aphids and other arthropods in Iran. It is univoltine in Iran. The eggs hatch in spring, nymphs emerge in early summer and adults appear in autumn. Larvae are ectoparasites of aphids whereas deutonymphs and adults are free-living predators of aphids and spider mites. Adults hibernate in the soil and in cracks of tree trunks during winter. When spring comes, females lay eggs in the soil, on the soil surface and on weeds. Phytoseius plumifer (Phytoseiidae) was observed to be phoretic on deutonymphs of A. pulvinum on nettle trees (Celtis australis). In the laboratory, development from the egg to adult stage takes at least 107 days at 25±1°C. Soil is not essential for female oviposition. High humidity is the most important factor for oviposition and development in A. pulvinum.     

Predation by Allothrombium pulvinum on the spider mites Tetranychus urticae and Amphitetranychus viennensis: Predation Rate, Prey Preference and Functional Response

The deutonymphs of Allothrombium pulvinum Ewing (Acari: Trombidiidae) are among the most important natural enemies of spider mites in North, North East and West Iran. In this study, maximum predation rate and preference experiments were conducted with A. pulvinum deutonymphs on apple leaf discs, to determine their preference for either of two spider mite species: Amphitetranychus viennensis (Zacher) and Tetranychus urticae Koch (Acari: Tetranychidae). Maximum predation rate tests showed that the predatory mite consumed more eggs and females of T. urticae than of A. viennensis. Furthermore, the Manly’s preference index for eggs and females of T. urticae confirmed that T. urticae were the preferred prey. The functional response of A. pulvinum deutonymphs on females of T. urticae was examined over a 24-h period. Predation of A. pulvinum deutonymphs presented with females of T. urticae followed a type III functional response. Estimated handling time for the predatory mites was 4.51 h and attack coefficient b, which describes the changes in attack rate with prey densities in a type III functional response, was 0.021.     

Where to from here? The mechanisms enabling the movement of first instar caterpillars on whole plants using Helicoverpa armigera (Hübner)

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is an economically-important, polyphagous herbivore in Old World countries. The distribution of larvae within various host plants has been described, but few studies have tried to determine the behavioural mechanisms by which the given distributions arose. Our aim was to determine the mechanisms which enable larval movement on pea plants, starting with first instars. Observations and bioassays determined larval movement in response to light and angled surfaces, as well as the effect of feeding and plant volatiles on these responses. The majority (68–72%) of 1st instars were positively phototactic towards blue, green and white light and 42% towards UV light. In the dark, larvae showed negative geotaxis. The angle of their substrate also had a kinetic effect on larvae; the steeper the angle from horizontal the more larvae moved under all conditions. Phenylacetaldehyde (a flower volatile) suppressed larval movement except at 90°. (Z)-3-Hexenyl acetate (a green leaf volatile) reversed the direction of movement at the flattest angle. Feeding lessened the probability of moving. We suggest that phototaxis and geotaxis are behaviours common to larval lepidopterans (caterpillars), and that these basic behaviours are modulated by environmental, larval, and plant factors to give observed distributions. Using a multinomial model approach, we created a flow chart to qualitatively and quantitatively represent the decision-making process of first instar H. armigera in response to the factors influencing movement.     

Parasitism ofAcyrthosiphon pisum byAllothrombium pulvinum (Acariformes: Trombidiidae): Host attachment site,host size selection,superparasitism and effect on host

Several aspects of parasitism of the pea aphid,Acyrthosiphon pisum (Harris), by the mite parasiteAllothrombium pulvinum Ewing, were examined in the laboratory. Larvae ofA. pulvinum were fastmoving mites that find their host by contact of their foretarsi with the host. They can attach to all parts of the host body, but insert their chelicerae only into weakly sclerotized parts such as intersegmental membranes. Of the attached larval mites, most (63.5–74.1%) were on the thorax of their hosts, regardless of host size. In hosts of small and medium size, the ventral side receives most parasitism, whereas in large hosts the lateral sites are most often attacked. Larval mites prefer large hosts when allowed to select between pairs of large and small hosts, but show no significant preference when allowed to choose between pairs of large and medium hosts or pairs of medium and small hosts. In two-choice tests, larval mites prefer previously parasitized hosts to umparasitized hosts, which results in superparasitism of the hosts. When the mite load is fiveA. pulvinum kills all small hosts within three days, and all medium hosts and 50% of large hosts within four days, the reproduction of surviving adult aphids were significantly reduced. Host-finding behavior, attachment site preference, host size selection, superparasitism, and effect on hosts are briefly reviewed for larval parasites of Trombidiidae. The potential role of larvalAllothrombium in integrated and biological aphid control is also discussed.     

The effect of temperature on phototaxis and geotaxis by larvae of the crab Rhithropanopeus harrisi (Gould)

Investigations of the effect of sudden temperature change on the phototaxis of Stage I and IV zoeae upon stimulation from horizontal and vertical directions with 500-nm light indicate a temperature-induced geotactic response in larvae of the crab Rhithropanopeus harrisi (Gould). For the horizontal tests both zoea stages were reared at 20 °C. Stage I showed positive phototaxis at temperatures between 15 ° and 35 °C, while Stage IV responded over the range of 10–30 °C. For the vertical tests, larvae, reared at 25 °C, were stimulated with overhead lights. Stage I zoeae ascended at 15 °, 20 ° and 25 °C and descended at 5 °, 10 °, 30 ° and 35 °C. Stage IV zoeae ascended at 20 ° and 25 °C and descended at 5 °, 10 °, 15 °, 30 ° and 35 °C. Although the descent at high temperatures could result from a negative phototaxis, a reversal in phototactic sign at high temperatures was not found in the horizontal experiments and the same vertical movement pattern is observed in total darkness. Upon exposure to high temperatures near the water surface, larvae would descend by means of a positive geotaxis rather than a negative phototaxis. This response involves active swimming by Stage IV larvae and passive sinking by Stage I.     

Biology ofAllothrombium pulvinum (Acariformes: Trombidiidae), a potential biological control agent of aphids in China

Allothrombium pulvinum Ewing is a common natural enemy of aphids and other small arthropods in China. It is univoltine species. The eggs hatch in spring, nymphs emerge in early summer, and adults emerge in autumn. Larvae are ectoparasites of aphids, whereas deutonymphs and adults are free-living predators of aphids, spider mites and some lepidopterous eggs. Adults hibernate in soil during winter and females lay their eggs in spring. In the laboratory the duration of immature stages is 74 days at 20–30°C. Larvae kill their host within 1–3 days when the mite load of the host is 2 or more. They decrease the reproductive rate of adult aphids and arrest the development of early-instar aphid nymphs when the mite load is 1. Larvae are capable of controlling the host population growth when the level of parasitism is high. Owing to its wide occurrence,A. pulvinum could be a potential candidate in the biological control of aphids. It is suggested that its impact on the host pest be evaluated in IPM programs in the future.     

Geotaxis in the springtail Folsomia candida

Laboratory assays demonstrated the presence of a small positive geotaxis response to a 15° incline by Folsomia candida Willem (Collembola: Isotomidae). Negative phototaxis played an additive role to positive geotaxis when the experimental apparatus were exposed to light. The geotactic response was negatively affected by cold acclimation and decreasing surrounding temperature, but unaffected by food deprivation. The reduced mobility of springtails at low temperature did not seem to play a role in the corresponding decreased geotaxis. The low level of geotaxis and its further decrease with exposure to low temperature support an earlier suggestion that F. candida do not respond to cooling temperatures of fall by relocation to warmer deeper soil layers, but remain in the upper soil layers and increase their cold tolerance to continue foraging in the food‐rich upper soil layers.     

Host plant choice and location by larvae of the wheat bulb fly (Delia coarctata)

Wheat bulb fly (Delia coarctata Fallen, Diptera: Anthomyiidae) is an important pest of winter wheat in the eastern half of the UK, and in northern and eastern Europe. The larvae must find a host plant and invade a tiller soon after hatching in late January. Chemical controls are costly and weather conditions may reduce their efficacy or prevent their application. Post‐emergence control relies on organophosphate insecticides, which may soon be withdrawn due to concerns about their negative health and environmental effects. Winter wheat (Triticum aestivum L.) is the preferred cereal host, but other winter cereals and related grasses may also be attacked, while oats (Avena spp.) are shunned. In choice test bioassays, neonate larvae chose couch grass (Elytrigia repens (L.) Nevski syn. Elymus repens (L.) Gould, Agropyearon repens (L.) Beauv.) seedlings and exudates over wheat seedlings and exudates, and exhibited geotaxis and negative phototaxis. Analysis of larval trails in choice test bioassays of seedling exudates showed that couch exudates are more attractive than wheat exudates, and that wheat exudates are more arrestant than couch exudates. This suggests that infochemicals isolated from couch, wheat, and oats could be used in wheat bulb fly control; possible delivery mechanisms are discussed. These findings, previous research, and a comparison of the phenologies and geographical distributions of D. coarctata and its hosts suggest that E. repens is the natural host of D. coarctata.     

Photoresponses of Chaoborus larvae

The diel vertical migration of Chaoborus larvae is a well known phenomenon. In order to quantify the ability of larvae to utilize underwater light cues in their migration, we measured photoresponses of fourth-instar Chaoborus punctipennis larvae in the laboratory. The action spectrum for these larvae was characterized by a maximum in sensitivity at 400 nm, a plateau at a lower sensitivity from 480 to 560 nm, and a region of much lower sensitivity at wavelengths longer than 620 nm. Dark-adapted larvae exhibited a positive phototaxis at low light intensity which shifted to a negative phototaxix as light intensity increased. At 540 nm the threshold intensity was 1.5 × 10^?9 W/m² for positive phototaxis and about 10^?6 W/m² for negative phototaxis. Light adaptation decreased sensitivity and altered the phototactic pattern. Larvae have a clear circadian rhythm in negative phototaxis, in which greatest responsiveness occurs early in the day. We suggest that the rhythm in photoresponsiveness primarily controls the timing of the downward migration at dawn.     

Overdispersion of Allothrombium pulvinum larvae (Acari: Trombidiidae) parasitic on Aphis gossypii (Homoptera: Aphididae) in cotton fields

Abstract.

• 1 Dispersion patterns of the protelean parasite, Allothrombium pulvinum Ewing, among individuals of an aphid host, Aphis gossypii Glover, were examined during spring 1991 in several cotton fields in Jiangsu Province, China.
• 2 The variance-to-mean ratios (i.e. dispersion index) of larval mites per host were greater than 1, indicating that the mite parasites were overdispersed among aphid hosts. The variance increased with the mean according to the power law, variance = 1.51 mean¹⁰⁶, which explained 99.7% of the variation in the data.
• 3 The negative binomial distribution adequately describes the patterns of larval mite dispersion among aphid hosts in eight out of ten populations. The degree of clumping (1/k) decreased curvilinearly with parasite density (mites per host).
• 4 Mites were more clumped among adult aphids than among immature ones.
• 5 Ecological and evolutionary consequences of mite overdispersion within host populations are discussed. The role of Allothrombium in pest control is also discussed.

     

A laboratory study on the short-term zonal oscillations of the trochid Monodonta turbinata (Born) (Mollusca: Gastropoda)

Zonal oscillations of Monodonta turbinata (Born) were monitored with an actographic device which precluded clustering behaviour. Unfed snails maintained their cyclic pattern of behaviour for up to 8 days under a light-dark cycle which simulated the natural one. Under constant conditions of light or dark, however, the snails ceased migration and occupied a zonal position typical of day and night, respectively. Experiments in diffuse light and with light from below the floor of the experimental tank showed that the downward migration of M. turbinata in the daytime depends on positive geotaxis combined with negative phototaxis whilst the upward migration at night depends on a negative geotaxis. This mechanism is similar to that described in other littoral molluscs. There was no evidence of an endogenous control of rhythmic zonal activity.     

Ontogenetic shift in geotaxis for walleye pollock,Theragra chalcogramma free embryos and larvae: potential role in controlling vertical distribution

Synopsis The behavioral capability of walleye pollock,Theragra chalcogramma free embryos and larvae to control vertical distribution was assessed by examining buoyancy during resting and swimming orientation and activity as they developed in complete darkness from hatching to first feeding readiness (1 to 7 d post hatching at 6° C). Free embryos exhibited positive geotaxis 1 d post hatching, actively swimming through a density gradient to remain in the lower water column. Activity increased with free embryo development and by 7 d post hatching, feeding-ready larvae reversed their vertical orientation, now exhibiting negative geotaxis as they migrated to the upper water column. The results indicate that even at the earliest developmental stages, walleye pollock possess the capability to control vertical distribution. Laboratory results are compared with patterns of vertical distribution observed in the sea.     

Photoresponses of second-instar Chaoborus larvae

The diel vertical migration of Chaoborus larvae varies with larval instar. Although light is involved in the control of vertical migration the contribution of larval photoresponses is unknown. In order to describe ontogenetic changes in larval photoresponses we measured photoresponses of second-instar Chaoborus punctipennis larvae in the laboratory. The response spectrum of these larvae had peaks in sensitivity at 420 and 620 nm with a wide plateau of lower sensitivity from 460 to 600 and 640 to 680 nm. Dark adapted larvae were positively phototactic at intensities from 10^?7 to 10¹ Wm^?2 at 420 nm. The level of response decreased somewhat above 10^?4 Wm^?2, and above 10^?2 Wm^?2 a small proportion of larvae shifted to a negative phototaxis. At 420 nm the threshold intensity was about 10^?7 Wm^?2 for positive phototaxis and 10^?2 Wm^?2 for negative phototaxis. Light adaptation increased the threshold intensity for positive phototaxis. The differences in larval photoresponses between second- and fourth-instar larvae suggests that the young instars are adapted to the photoenvironment of the water column and older larvae are adapted to avoid the water column except at very low light intensities. These predictions match the diel distribution of these larvae.     

Photoresponses of late instar <Emphasis Type="Italic">Chaoborus punctipennis</Emphasis> larvae to UVR

With a few clear exceptions (e.g., Daphnia) it is uncertain if most aquatic invertebrates can detect and respond to ultraviolet radiation (UVR). It is known that many aquatic invertebrates are vulnerable to UVR and that anthropogenically-induced increases in surface UVR have occurred in recent decades. We examined the photoresponses of late larval instars of Chaoborus punctipennis to different combinations of UVA (320–400 nm), UVB (300–320 nm) and visible light (400–700 nm) to determine whether the larvae can detect and/or avoid UVR. To accomplish this, we exposed late instar C. punctipennis larvae to a directional light source of UVR only (peak wavelength at 360 nm), visible light only or visible plus various wavebands of UVR. We examined negative phototaxis for 10 min at a quantum flux of 2.62 x 10¹³ quanta s^–1 cm^–2 (S.D. = 3.63 x 10¹² quanta s^–1 cm^–2). In the dark, larvae stayed close to the surface of the experimental vessels. Under all treatments containing visible light the larvae exhibited negative phototaxis and occupied the bottom of the vessels. Under UVR only, the larvae occupied the middle of the water column. Our results suggest that late instar C. punctipennis larvae are unable to detect and avoid UVB and short UVA wavelengths but they can detect long UVA wavelengths.     

GEOTAXIS/PHOTOTAXIS AND BIOCHEMICAL PATTERNS IN HETEROCAPSA (=CACHONINA) ILLDEFINA (DINOPHYCEAE) DURING DIEL VERTICAL MIGRATIONS

Two separate experiments with Heterocapsa (= Cachonina ) illdefina Herman et Sweeney, one with and the other without water volume replacement, were performed in a 250-L laboratory mesocosm (45-cm diameter × 150-cm height) to examine how diel vertical migration (DVM) relates to taxis sign and strength and to cellular biochemical state. Although only the cell population grown with water volume replacement maintained a division per day over the course of the experiment, periodic measurements during both experiments demonstrated that cells aggregating at the surface during the light period generally were deficient in all measured biochemical constituents compared to cells obtained from a midcolumn depth. More specifically, H. illdefina cells that aggregated at the surface during the light period in both experiments exhibited weakened positive geotaxis but strengthened positive phototaxis and were very deficient in lipid and free amino acid compared to midcolumn cells. Cells sampled at midcolumn during the light period exhibited similar but weaker taxes changes compared to surface samples, and geotaxis strength was inversely correlated with cell diameter, cellular DNA and protein content, and RNA/DNA ratio. In comparison, published data on Gymnodinium breve Davis, a harmful algal bloom species, showed that cells aggregating at the surface during the light period generally exhibited weakened negative geotaxis and strengthened positive phototaxis and were very deficient in lipid and chl a compared to midcolumn cells. Although the persistent tendency toward negative geotaxis was weaker in midcolumn subpopulations throughout the day, its strength was inversely correlated with cell diameter and cellular lipid content. The combined results for both species support a revised conceptual model of optimized DVM in autotrophic marine dinoflagellates incorporating generalized expressions of taxis and biochemical state of individual cells.