Functional Response of Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) to Sugarcane Whitefly Aleurolobus barodensis (Maskell) in Laboratory Conditions

A functional response study of Chrysoperla carnea (Stephens) larvae to different densities of sugar cane whitefly Aleurolobus barodensis (Maskell) was conducted in test tubes at 26?±?2 °C, 65?±?5 % RH. Chrysoperla carnea showed two different types of functional response in larval instars. First instar exhibits type II. However, second and third larval instars revealed type III functional response. Based on modified Holling’s disk equation, the highest searching rates (a) of 0.82?±?0.0247 h^?1 was found for first instar larva. For second and third larval instars, the attack coefficient (b) were 0.002?±?0.030 and 0.0025?±?0.0424 respectively. The shortest handling time (T_h) per prey was observed at third instar stage (1.574?±?0.0568 h) followed by second and first instar with 1.72?±?0.0411 h and 1.919?±?0.0568 h respectively.     

Foraging behaviour of Scymnus coccivora Ayyar against cotton mealybug Phenacoccus solenopsis Tinsley

Predation is one of the significant biotic mortality factors reducing the insect pest population as functional response and the numerical response of the predator are the key factor regulating the population dynamics of predator prey species. This study is aimed to evaluate the functional response of all the developmental stages of Scymnus coccivora Ayyar (Coleoptera: Coccinellidae) against the different densities of cotton mealybug, Phenacoccus solenopsis (Tinsley) (Hemiptera: Pseudococcidae) and the numerical response of female predator. Experiments were carried out in controlled environment laboratory conditions at 25 ± 1 °C temperature, 60 ± 5% relative humidity and photoperiod of 16 h. Number of eggs consumed, number of eggs laid and the Efficiency of Conversion of Ingested food (ECI) were recorded daily. Results from the study revealed that all the developmental stages of S. coccivora exhibited a Type II response. Different parameters such as attack rate (a’), handling time (T_h) and the maximum rate of predation were estimated using Roger’s random attack equation and Holling Disc equation in which Rogers random attack equation was found best fit. Female has shown the highest attack rate (a’) followed by IV^th instar grub, male, III^rd, II^nd and I^st instar grub. With low handling time, IV^th instar grub has shown maximum predation rate of 76.40 per day followed by female (75.86), male (58.79), III^rd (22.84), II^nd (19.65) and I^st instar grub (15.39). The numerical response increase was curvilinearly related to different prey densities with the highest number of eggs (11.8 ± 3.44) produced at highest prey density (160). The Efficiency of Conversion of Ingested food (ECI) was highest (64.49 ± 8.03) at prey density of 10. Understanding the factors that lead to variation in functional response of predator in natural population will advance our understanding of the effects of predation on individual and the effectiveness of coccinellid predators as biocontrol agent against cotton mealybug.     

Interspecific variation in thermoregulation among three sympatric bats inhabiting a hot, semi-arid environment

Bats in hot roosts experience some of the most thermally challenging environments of any endotherms, but little is known about how heat tolerance and evaporative cooling capacity vary among species. We investigated thermoregulation in three sympatric species (Nycteris thebaica, Taphozous mauritianus and Sauromys petrophilus) in a hot, semi-arid environment by measuring body temperature (T _b), metabolic rate and evaporative water loss (EWL) at air temperatures (T _a) of 10?C42?°C. S. petrophilus was highly heterothermic with no clear thermoneutral zone, and exhibited rapid increases in EWL at high T _a to a maximum of 23.7?±?7.4?mg?g^?1?h^?1 at T _a????42?°C, with a concomitant maximum T _b of 43.7?±?1.0?°C. T. mauritianus remained largely normothermic at T _as below thermoneutrality and increased EWL to 14.7?±?1.3?mg?g^?1?h^?1 at T _a????42?°C, with a maximum T _b of 42.9?±?1.6?°C. In N. thebaica, EWL began increasing at lower T _a than in either of the other species and reached a maximum of 18.6?±?2.1?mg?g^?1?h^?1 at T _a?=?39.4?°C, with comparatively high maximum T _b values of 45.0?±?0.9?°C. Under the conditions of our study, N. thebaica was considerably less heat tolerant than the other two species. Among seven species of bats for which data on T _b as well as roost temperatures in comparison to outside T _a are available, we found limited evidence for a correlation between overall heat tolerance and the extent to which roosts are buffered from high T _a.     

Functional response of different stages of Hippodamia variegata (Col.: Coccinellidae) to Diuraphis noxia (Hemiptera: Aphididae) on two wheat cultivars

As one of the most important natural enemies of aphids, lady beetles have been widely used for the control of pest aphids. Important to the efficiency of natural enemies is their functional response to varying prey densities. In this study, the functional response of Hippodamia variegata Goeze (Col.: Coccienllidae) to different densities of Diuraphis noxia Mordvilko was examined on two wheat cultivars with different resistance levels under laboratory conditions (25 ± 1°C, 65 ± 5% RH and 16:8 L:D photoperiod). Our results revealed that all stages of H. variegata exhibited a type II functional response, in that predation decreased with increasing prey density. Searching efficiency (a) and handling time (T_h) of third instar larvae on the Sardari cultivar were 0.0285/h and 0.0079 h, while for fourth instar larvae these values were 0.0561/h and 1 × 10^?8 h, for males these values were 0.0412/h and 0.0173 h and for females were 0.059/h and 0.0168 h. The corresponding values for the back cross cultivar for third instar larvae were 0.0475/h and 0.0782 h, for fourth instar larvae were 0.0753/h and 0.0529 h, for males were 0.045/h and 0.0060 h and for females were 0.0516/h and 0.0077 h. This study demonstrated that these wheat cultivars do not have a significant effect on the functional response parameters of H. variegata life stages, except for the searching efficiency of third instar larvae and the handling time of male adults. The possible application of this species against Russian wheat aphids in an integrated pest management programme is also discussed.     

A new method to study the functional response of Scymnus syriacus (Coleoptera: Coccinellidae) to different densities of Aphis gossypii

Functional response is basic to any investigation of predator–prey relationships. In this study, the functional response of female Scymnus syriacus Marseul (Col.: Coccinellidae) to different densities (10, 20, 40, 60, 80, 100) of third instar nymphs of Aphis gossypii Glover as prey was studied in an open patch experiment in a growth chamber (25 °C, 65 ± 5% RH and a photoperiod of 16L:8D h ). Using logistic regression, a type II functional response for female Scymnus syriacus was determined. The searching efficiency (a') and handling time (T_h) of the female predator using non linear least-square regression were estimated as 0.0769 ± 0.0136 h^? 1 and 0.3103 ± 0.0438 h., respectively. Mean times required for the female predator to settle in a patch were 10.20 ± 4.28, 6.58 ± 2.58, 12.58 ± 4.50, 4.53 ± 1.48, 5.14 ± 2.59, 3.87 ± 3.52 min at different prey densities, respectively. Maximum theoretical predation rate (T/T_h) estimated by Rogers' model for the female predator was 77.34. The proportion of female predators remaining in open patches at the end of the experiment was dependent on prey density (R² = 0.876). The type of functional response obtained here agrees with studies on this predator in closed patches.     

Responses of Episyrphus balteatus DeGeer (Diptera: Syrphidae) in relation to prey density and predator size

Laboratory experiments were conducted to determine the functional and numerical responses of the aphidophagous hover fly Episyrphus balteatus DeGeer (Diptera: Syrphidae) to different densities of 4th instar black bean aphids, Aphis fabae Scopoli (Homoptera: Aphididae), on broad bean, Vicia faba L. (Fabaceae). Two different-sized larvae of predators were tested in different densities of similar-sized prey to determine whether functional response parameters depended on the body sizes of predator and prey. In numerical response experiments, gravid E. balteatus females were exposed individually to different densities of 4th instars of A. fabae on cut sections of the broad bean plant, V. faba L. Logistic regression suggested a type II functional response for both larval sizes of E. balteatus. The searching efficiency (a) of the larger larvae was higher than that of the smaller ones. Prey consumption was higher, and handling time (T_h) was lower for larger larvae than smaller ones. The theoretical maximum number of A. fabae nymphs eaten by the different-sized larvae was 125 and 269 nymphs per day. Larger E. balteatus larvae are more efficient predators for aphid management strategies. The reproductive numerical response, in terms of the number of eggs laid, increased curvilinearly with increasing prey density, but the proportion of eggs laid (egg number/prey density) decreased as the initial density of prey increased.     

Effects of pupal treatment with Proteus® and Metarhizium anisopliae sensu lato on functional response of Habrobracon hebetor parasitising Helicoverpa armigera in an enclosed experiment system

The ectoparasitoid wasp, Habrobracon hebetor Say (Hymenoptera: Braconidae) is a valuable biocontrol agent attacking larval stages of many lepidopteran pests including Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae). The possibility of combined application of this parasitoid wasp with other biological control agents and chemical insecticides is necessary for the success of an Integrated Pest Management (IPM) programme. One of the most important behaviours of a parasitoid refers to the number of hosts attacked by per parasitoid as a function of host density. In this research, the single and joint effects of the insecticide, Proteus® and the entomopathogenic fungus, Metarhizium anisopliae sensu lato (Metsch.) Sorokin (Hypocreales: Clavicipitaceae) (isolate M14) were studied on the functional response of H. hebetor. Newly mated females (<24?h) from treated pupae of H. hebetor were used to study the functional response of this parasitoid wasp to different densities of H. armigera larvae. A type II functional response was observed in all treatments. The results revealed that control and Proteus® treatments developed the lowest (0.5599?±?0.0373?h) and the highest (0.5709?±?0.0443?h) handling time, respectively. The most and the least values of attack rate were observed in control (0.0996?±?0.0164?h^?1) and Proteus® (0.0776?±?0.0119?h^?1) treatments, respectively. The maximum theoretical parasitism rate (T/T_h) was estimated to be 42.86 for female control wasps. After conducting field experiments on functional response along with life table parameters, M. anisopliae sensu lato (isolate M14) may be used as a compatible biocontrol agent with H. hebetor in IPM programmes of H. armigera.     

Predation and functional response of the multi-coloured Asian ladybeetle Harmonia axyridis on the adult Asian citrus psyllid Diaphorina citri

Diaphorina citri (Hemiptera: Liviidae) is a vector for the citrus Huanglongbing greening disease, which can reduce crop yields. Harmonia axyridis (Coleoptera: Coccinellidae) is known to prey on D. citri nymphs; we investigated the predation capacity and functional response of H. axyridis on adult D. citri. H. axyridis larvae exhibited Holling’s type II functional response to different densities of adult D. citri. For second, third and fourth instar H. axyridis larvae, the successful attack rates were 0.75?±?0.22, 1.06?±?0.11, and 1.04?±?0.26, respectively; the handling times were 18.08?±?5.37, 0.45?±?0.07, and 0.24?±?0.07?h, respectively; and the estimated maximum predation rates were 4, 54, and 102, respectively. Fourth instar larvae had the lowest handling time. Adult H. axyridis consumed more D. citri adults maintained at 15–20°C than at 30–35°C, with an optimal temperature of 17.28°C. More adults were consumed during the day than at night, with peak consumption between 12:00 and 14:00?h, not differing significantly differ between predator genders. Adult H. axyridis reared on the aphid Acyrthosiphon pisum were tested for D. citri predation; these H. axyridis had successful attack rates, handling times, and estimated maximal predation rates of 0.66?±?0.18, 0.82?±?0.26?h, and 29, respectively. Conversely, those fed and tested on D. citri had corresponding rates of 0.99?±?0.17, 0.38?±?0.12?h, and 63, respectively. Significant H. axyridis predation on adult D. citri indicated its enhanced role in suppressing both nymph and adult D. citri.     

Age-dependent functional response of Aphidius matricariae (Hymenoptera: Braconidae) on tobacco aphid,Myzus persicae nicotianae (Hemiptera: Aphididae)

This study was performed to investigate the age-dependent functional response of Aphidius matricariae Haliday (Hymenoptera: Braconidae) at varying densities of tobacco aphid, Myzus persicae nicotianae Blackman (Hemiptera: Aphididae). Six densities (2, 4, 8, 16, 32, and 64 numbers) of third instar nymphs of M. persicae nicotianae were daily exposed to one pair of 1-day-old A. matricariae until the parasitoid female died. Each host aphid density was replicated 10 times. The experiments were conducted at constant environmental conditions (25 ± 1 °C, 70 ± 5% RH and 16:8h L:D photoperiod). The logistic regression model demonstrated that the functional response of A. matricariae was type III in the second day and type II in the first day and from third to the end of adult parasitoid lifetime. The handling time (T_h) was lowest in the first day (0.2305 h) and highest in the sixth day (2.9296 h) of the parasitoid’s life. The searching efficiency (a) values were 0.0573, 0.0649, 0.0503, 0.0229, and 0.0077 h^?1 in the first, third, fourth, fifth, and sixth days of the parasitoid’s life, respectively. Also, the b value for type III functional response was 0.0147 in the second day. The values of maximum attack rate (T/T_h) ranged from 104.12 nymphs/24 h in the first day to 19.8 nymphs/24 h in the sixth day of parasitoid’s life. Based on the results, A. matricariae showed a high potential to control the population of tobacco aphid and the highest efficiency gained in the first half of the female parasitoid life.     

Efficacy of pesticides on the functional response on larval ectoparasitoid,Habrobracon hebetor Say (Hymenoptera: Braconidae)

Exposing to sub-lethal and low lethal doses of pesticides may cause changes in natural enemy behavioural, such as functional, response. In this study, the effects of chlorpyrifos, carbaryl, abamectin and spinosad were evaluated on the functional response of the Habrobracon hebetor to different densities of last instar larvae of Anagasta kuehniella Zeller. Young adult females of the parasitoid were exposed to LC₃₀ of chlorpyrifos, carbaryl, abamectin and spinosad that were 0.32, 4.03, 3.05 and 17.51?mg a.i./l for 24?h, respectively. Host densities of 2, 4, 8, 16, 32 and 64 were offered to treated young females for 2?h in 10-cm Petri dishes and then the parasitism data were recorded. Experiments were conducted in eight replications. Functional response type was determined using logistic regression and the parameters were appraised by non-linear regression using statistical analysis software. Functional response was type Ш in control and insecticide treatments. Searching efficiency in control, chlorpyrifos-, carbaryl-, abamectin- and spinosad-treated wasps were 0.008?±?0.002, 0.002?±?0.0009, 0.0034?±?0.0013, 0.0076?±?0.002 and 0.0073?±?0.002?h^?1and handling times were 1.38?±?0.1, 7.64?±?1.01, 3.3?±?0.315, 1.55?±?0.1 and 1.46?±?0.11?h, respectively. Chlorpyrifos and carbaryl had the highest effect on searching efficiency of H. hebetor. Spinosad and abamectin showed less adverse effect on the functional response parameters. Finally, after conducting the advanced field studies, spinosad and abamectin may be used as a compatible chemical material with biological control agent in integrated pest management programmes.     

Parasitism capacity and searching efficiency of Diaeretiella rapae parasitizing Brevicoryne brassicae on susceptible and resistant canola cultivars

Different cultivars of aplant species can affect the foraging and efficiency of natural enemies, both directly through physical and biochemical properties or indirectly through the herbivore's diet. In this study, the parasitism capacity and functional response of Diaeretiella rapae McIntosh were determined on the cabbage aphid, Brevicoryne brassicae (L.) reared on susceptible (Opera) and resistant (Okapi) canola cultivars under laboratory conditions at 25?±?1?°C, 60?±?5% RH and a16:8?h L:D photoperiod. The parasitoid exhibited Type II and Type III functional responses on the resistant and susceptible cultivars, respectively. The estimated value of searching efficiency (a) was 0.1637?±?0.1095?h^?1 on the resistant cultivar whereas its value was dependent on host density on the susceptible cultivar. The handling times (T_h) on the susceptible and resistant canola cultivars were 0.108?±?0.040 and 0.320?±?0.048?h, respectively. The net parasitism rate (C₀) of the parasitoid wasp varied from 128.09 hosts per parasitoid lifetime on the susceptible to 71.01hosts on the resistant canola cultivar. The transformation rate from host population to parasitoid offspring (Qp) was equal to 1 on both cultivars (C₀?=?R₀). The finite parasitism rate (ω) on the susceptible cultivar (0.819 hosts per parasitoid per day) was significantly higher than that on the resistant one (0.578 hosts per parasitoid per day). In conclusion, canola cultivars affected the performance of D. rapae in controlled small-scale laboratory experiments and compared with the susceptible cultivar, the resistant one had anadverseeffect on the efficiency of the parasitoid.     

Effect of treating eggs of cotton bollworm with Bacillus thuringiensis Berliner on functional response of Trichogramma brassicae Bezdenko

Cotton bollworm, Helicoverpa armigera (Hübner) is a cosmopolite insect pest of a wide spectrum of crops such as cotton, maize, tomato, soybean, etc. Egg parasitoids mainly Trichogramma brassicae Bezdenko and Bacillus thuringiensis Berliner (Bt) are biological control agents, that are used as components of sustainable and environmentally compatible IPM systems. Although Bt does not come in direct contact with egg parasitoids, it may persist within the host’s body and affect the quality of the host’s eggs via biochemical changes in their mother and possibly behaviour and potency of the parasitoids. In this study, the functional response of T. brassicae to different densities of H. armigera eggs was investigated in two sets of experiments at 26?±?1?°C, 65?±?5% RH, and 16: 8?h photoperiod. The first group was a control and the second one were eggs laid by hosts treated as 3rd instar larvae with LC₂₀ of Bt (determined as 9.8?×?10^5?IU/l of artificial diet based on a preliminary bioassay). A type III functional response was observed in both treatments with a direct density dependent mortality up to eight host eggs and an inverse one upward. Both handling time and searching efficiency were affected by Bt treatment as the handling time was increased by a factor of 1.5 and the searching efficiency was decreased by a factor of 0.6. The searching efficiencies were 0.0310?±?0.003 and 0.0182?±?0.005?h^?1, and handling times were 1.134?±?0.042 and 1.672?±?0.082?h in control and Bt treatment respectively.     

Life table of Gaeolaelaps aculeifer (Acari: Laelapidae) feeding on larvae of Lycoriella auripila (Diptera: Sciaridae) with stage-specific estimates of consumption

Gaeolaelaps aculeifer (Canestrini, 1883) is a soil-dwelling predatory mite with potential for use as a biological control agent of fungus gnats (Diptera: Sciaridae) in mushroom production. The life table, predation rate and population growth rate of G. aculeifer on a diet of larvae of the sciarid fly, Lycoriella auripila, at 23?±?1°C, 60?±?5% RH and a photoperiod of 0:24 (L:D)?h was investigated. The results revealed that the duration of egg, larva, protonymph, deutonymph, females and males of G. aculeifer were 3.8?±?0.1, 1.4?±?0.1, 3.9?±?0.1, 4.1?±?0.1, 67.7?±?2.8 and 60.3?±?3.1 days, respectively. Net reproductive rate (R₀) was 54.8?±?7.1 offspring, intrinsic rate of increase (r) was 0.12?±?0.01 offspring day^?1, finite rate of increase (λ) was 1.13?±?0.01 day^?1and mean generation time (T) was 32.3?±?0.6 days. The predator consumed a mean of 0.08?±?0.05, 1.73?±?0.18, 3.16?±?0.28 and 75.9?±?7.1 third instar L. auripila larvae during the larval (1.3?±?0.1 days), protonymph (3.9?±?0.1 days), deutonymph (4.1?±?0.1 days) and adult (52.6?±?2.2 days) stages. Population parameters and consumption rates suggest that G. aculeifer has good potential as a biological control agent of L. auripila in mushroom production.     

Consumption rate and functional response of the predaceous mite<Emphasis Type="Italic"> Kampimodromus aberrans</Emphasis> to two-spotted spider mite <Emphasis Type="Italic">Tetranychus urticae</Emphasis> in the laboratory

Prey stage preference of female Kampimodromus aberrans (Oudemans) (Phytoseiidae) at constant densities of different stages of Tetranychus urticae Koch (Tetranychidae), functional response types and parameters of the predator females to the varying densities of eggs, larvae, protonymphs and deutonymps of T. urticae were determined in order to establish its potential for the mite biological control. Experiments were conducted at 25 ± 1°C, 65 ± 10% RH and 16:8 (L:D) photoperiod. Our results indicated that the predator consumed significantly more prey larvae than other prey stages. Functional response type of predator was determined by a logistic regression model. The predator exhibited a Type II response on all prey stages. The attack rate (α) and handling time (T _h ) coefficients of a Type II response were estimated by fitting a “random-predator” equation to the data. The lowest estimated value α and the highest value of T _h (including digestion) were obtanined for the predator feeding on deutonmph. The lowest value of T _h were obtained for the predator feeding on prey larvae, but the attack rate value obtained on larva wasn’t different than that obtained on egg and protonymph. According to our results, K. aberrans could be an efficient biological control agent of T. urticae at least at low prey densities. However, further field based studies are needed to draw firm conclusions.     

Comparison of Acyrthosiphon pisum (Homoptera: Aphididae) and Frankliniella occidentalis (Thysanoptera: Thripidae) as prey for the development and survival of Chrysopa pallens (Neuroptera: Chrysopidae)

The effects of two different prey species on the pre-imaginal development and survival of the green lacewing, Chrysopa pallens were determined under laboratory conditions. The developmental duration from first instar to adult emergence was shorter when larvae were fed on Acyrthosiphon pisum (22.56 ± 0.39 d) than on Frankliniella occidentalis (29 ± 0.45 d). The survival rate of lacewings from first instar to adult emergence was 22.6% higher when they were fed on A. pisum than when fed on F. occidentalis. The possible implications of these findings are applicable to mass-rearing of C. pallens and for understanding predator's biology and survival capacity in the presence of prey species evaluated and could be used in the development of an effective biological control program.     

Host-stage preference and functional response of aphid parasitoid Diaeretiella rapae (McIntosh) (Hym.: Braconidae) on greenbug,Schizaphis graminum (Rondani) (Hem: Aphididae)

Searching capacity is of great importance in the efficiency of natural enemies. In this research, the host-stage preference and functional response of Diaeretiella rapae (McIntosh) to different densities of greenbug, Schizaphis graminum (Rondani) were determined on Pishtaz wheat leaves at 25 ± 1°C, 60–70% relative humidity, and a photoperiod of 16:8 h (L:D). The host-stage preference was investigated by exposing 60 number of all nymphal instars and adults of greenbug to individual parasitoid females at eight replicates. Results indicated significant preference of D. rapae to second instar nymphs of S. graminum to perform functional response experiment. Densities of 2, 4, 8, 16, 32, 50, 64, 90 and 130 second instar nymphs of greenbugs were exposed to individual mated female wasps at 10 replicates for 24 h. Type of functional response was determined by logistic regression and its parameters, searching efficiency (a) and handling time (T _h), were estimated by non linear regression. The results indicated that the functional response of D. rapae was type III and values of searching efficiency, handling time and maximum rate of parasitism were calculated by two models as 0.1364 ± 0.084, 0.5761 ± 0.0179, 41.6594; 0.06489 ± 0.00652 (h^?1), 0.5628 ± 0.0173 (h) and 42.6439 (aphids/parasitoid/day), respectively.     

Interspecific variation in the escape responses of aphids: effect on risk of predation from foliar-foraging and ground-foraging predators

A series of laboratory experiments was conducted to determine the effect of interspecific differences on prey defensive behavior on the susceptibility of two aphid species (Acyrthosiphon pisum and A. kondoi) to a ground-foraging predator, Harpalus pennsylvanicus, and a foliar-foraging predator, Coccinella septempunctata. These organisms are representative of a biologically and economically important predator/prey system in alfalfa. The primary defensive behavior of both aphid species toward C. septempunctata was to “drop” from the plant. Both aphid species were significantly more likely to drop from the plant in the presence of C. septempunctata. However, when C. septempunctata was present, a significantly lower proportion of A. kondoi individuals dropped (0.42 ± 0.07) compared to A. pisum (0.73 ± 0.08). As a result of their lower propensity to drop from the plant A. kondoi individuals are significantly more likely to be consumed by C. septempunctata. Conversely, the higher propensity of A. pisum individuals to drop increased their susceptibility to ground-foraging predators. When A. pisum was the prey species, ground-foraging predators made a significant contribution to overall aphid suppression and there was a significant synergistic interaction between ground and foliar-foraging predators. When A. kondoi was the prey there was no interaction between the predator species. As either a cause or consequence of its higher propensity to drop, A. pisum seems to be more adapted for survival and dispersal off the plant. In comparison to A. kondoi individuals, A. pisum individuals relocate plants more quickly (63 ± 41 s vs. 164 ± 39 s), disperse farther (18 ± 1.7 cm vs. 13 ± 0.66 cm), and survive longer (37 ± 2.0 h vs. 25 ± 2.0) off the plant. This study demonstrates the importance of prey defensive behavior in determining the susceptibility of a prey species to a multiple-predator complex. Received: 24 February 1997 / Accepted:17 December 1997     

Ontogeny of hypoxic modulation of cardiac performance and its allometry in the African clawed frog Xenopus laevis

The ontogeny of cardiac hypoxic responses, and how such responses may be modified by rearing environment, are poorly understood in amphibians. In this study, cardiac performance was investigated in Xenopus laevis from 2 to 25 days post-fertilization (dpf). Larvae were reared under either normoxia or moderate hypoxia (PO₂ = 110 mmHg), and each population was assessed in both normoxia and acute hypoxia. Heart rate (f _h ) of normoxic-reared larvae exhibited an early increase from 77 ± 1 beats min^?1 at 2 dpf to 153 ± 1 beats min^?1 at 4 dpf, followed by gradual decreases to 123 ± 3 beats min^?1 at 25 dpf. Stroke volume (SV), 6 ± 1 nl, and cardiac output (CO), 0.8 ± 0.1 μl min^?1, at 5 dpf both increased by more than 40-fold to 25 dpf with rapid larval growth (~30-fold increase in body mass). When exposed to acute hypoxia, normoxic-reared larvae increased f _h and CO between 5 and 25 dpf. Increased SV in acute hypoxia, produced by increased end-diastolic volume (EDV), only occurred before 10 dpf. Hypoxic-reared larvae showed decreased acute hypoxic responses of EDV, SV and CO at 7 and 10 dpf. Over the period of 2–25 dpf, cardiac scaling with mass showed scaling coefficients of ?0.04 (f _h ), 1.23 (SV) and 1.19 (CO), contrary to the cardiac scaling relationships described in birds and mammals. In addition, f _h scaling in hypoxic-reared larvae was altered to a shallower slope of ?0.01. Collectively, these results indicate that acute cardiac hypoxic responses develop before 5 dpf. Chronic hypoxia at a moderate level can not only modulate this cardiac reflex, but also changes cardiac scaling relationship with mass.     

Temperature-dependent functional response of Euborellia annulipes (Dermaptera: Anisolabididae) preying on Plutella xylostella (Lepidoptera: Plutellidae) larvae

Temperature mediates trophic interactions, including relationships between insect pests and predators, and functional response studies are often used to determine the suitability of predators as biocontrol agents. We investigated the effects of temperature on the functional response of Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae) preying on Plutella xylostella (L.) (Lepidoptera: Plutellidae) larvae. Predation rate, type of functional response, attack rate (a'), handling time (T_h), and maximum predation rate (T/T_h) of the predator were estimated using seven prey densities and three thermal conditions. The functional response of E. annulipes to P. xylostella was temperature-dependent, type III under the lower temperatur (18°C and 25 °C) , and type II at 32 °C. We observed increasing values of a’ in 25 °C and 32 °C, decreasing values of Th and highest T/T_h as the thermal condition increased. Our findings suggest that E. annulipes could be effective to control P. xylostella under different thermal conditions, however its predation behavior changes according to temperature variation.     

Effects of Bacillus thuringiensis Cry1C toxin on the metabolic rate of Cry1C resistant and susceptible Spodoptera exigua (Lepidoptera: Noctuidae)

Abstract. The effects of Bacillus thuringiensis (Bt) Cry1C toxin on the metabolic rate of Cry1C resistant and susceptible Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) are investigated using closed‐system respirometry. Mechanisms of resistance to the Bt toxin may be associated with an energetic cost that can be measured as an increase in metabolic rate compared with Bt‐susceptible insects. This hypothesis is tested using third‐ and fifth‐instar larvae and 1–7‐day‐old pupae. Metabolic rate is measured as the amount of O₂ consumed and CO₂ produced. V?_O2 and V?_CO2 (mL g^?1 h^?1) of third‐instar Cry1C resistant larvae reared continuously on a diet containing 320 µg Cry1C toxin per g diet (CryonT) are significantly greater than third‐instar Cry1C resistant larvae reared on toxin for 5 days and reared thereafter on untreated diet (Cry5dT), Cry1C resistant larvae reared on untreated diet (CryReg) and the susceptible parental strain (SeA) reared on untreated diet. There are no differences in V?_O2 and V?_CO2 (mL g^?1 h^?1) among treatment groups for fifth‐instar larvae. CryonT larvae and pupae weigh significantly less than larvae and pupae receiving other treatments. Smaller body mass may be an important biological cost to individuals exposed continuously to Bt toxin. One‐day‐old pupae of all treatment groups exhibit a high V?_O2 (mean approximately 0.174 mL g^?1 h^?1) with CryonT having a significantly greater value than all other treatments; there are no differences among the other treatments. Pupal metabolic rates of all treatment groups decline to a minimum between days 2 and 4 then increase linearly between days 4 and 7 until adult emergence. These results demonstrate no difference in metabolic rates, and possibly fitness costs, between resistant (CryReg and Cry5dT) and susceptible (SeA) S. exigua except when larvae were reared continuously on toxin (CryonT).