Temperature and altitude modulate feeding attributes of Mexican beetle,Zygogramma bicolorata Pallister on Parthenium hysterophorus

Zygogramma bicolorata Pallister (Coleoptera: Chrysomelidae) is an effective biocontrol agent of Parthenium hysterophorus L. which is an alien invasive herbaceous weed with a pan-tropical distribution. The present study aimed to assess the effects of temperature and altitude on feeding attributes (consumption rate, conversion efficiency and growth rate) of adults from the wild populations of Z. bicolorata inhabiting India and Nepal. Results revealed that adults inhabiting areas of low temperature (24°C ‒ 25°C) and high altitude (415 m ‒1400 m) were large and had higher food consumption rates. In contrast, those inhabiting areas of high temperature (34°C ‒ 36°C) and low altitude (81 m ‒ 229 m) were smaller and had higher food utilization efficiencies. In all the eco-climatic regions, females were larger than males and had higher feeding attributes than their counterparts. Temperature between 27°C and 30°C was found optimal for Z. bicolorata adults to convert and utilize the food biomass to body mass. Above the optimal temperature the feeding attributes decreased. Present results suggest that there exists a possibility for decrease in body size, and thereby weed biocontrol efficiency of Z. bicolorata adults with an increase in temperature due to global climate change.     

Food consumption,utilization and ecological efficiency of Parthenium beetle,Zygogramma bicolorata pallister (Coleoptera: Chrysomelidae)

Food consumption, utilization and ecological efficiency of Zygogramma bicolorata were studied in larvae and adults up to 45 days old. The study revealed that sex had a significant influence on dry food consumption in adults but not in larvae. The other parameters of ecological efficiencies, viz. efficiency of conversion of ingested food (E.C.I.), tissue growth and dry biomass, were highest in the fourth instar and in adult females. The Consumption Index (C.I.) was higher in males than females in both larval and adult stages. Developmental and growth rates were relatively higher in the first and second instars. The fourth instar and adult females were the most efficient consumers and converters of food. Food intake by females was an important determinant of the number of eggs laid.     

Host range and risk assessment of Zygogramma bicolorata,a defoliating agent released in South Africa for the biological control of Parthenium hysterophorus

Parthenium hysterophorus (parthenium) is a weed of international importance and is spreading rapidly in sub-Saharan Africa. Consequently, it has been targeted for biocontrol in South Africa since 2003. Based on precedents elsewhere in the world, the defoliating beetle Zygogramma bicolorata was prioritised as a candidate agent. Although no-choice tests, involving some 48 test plant species, indicated a significant preference for parthenium, significantly reduced feeding and oviposition was recorded on some species. Multiple-choice tests resolved many of these non-target results; however, Helianthus annuus (sunflower) was still selected for oviposition and feeding. Of the 12 sunflower cultivars tested, four were selected for oviposition, while two were selected for oviposition and feeding. These six cultivars were then subjected to larval development trials, together with three native and two weed species (in the Asteraceae). These trials showed high levels of complete development on parthenium, significantly reduced development on sunflower cultivars, and partial development on only one of the weed species. Finally, a risk assessment was conducted on the six sunflower cultivars to quantify Z. bicolorata feeding and reproductive performance. Feeding risk calculations revealed these cultivars to have an extremely low risk (<0.2%) of supporting Z. bicolorata feeding and development. Similarly, reproductive risk calculations showed a very low risk (<0.16%) of supporting viable Z. bicolorata populations. These data are supported by findings from both the native (Mexico) and introduced ranges (Australia, India) of Z. bicolorata, where it has never been recorded as a pest of sunflower. These considerations were accepted by the regulatory authorities and in August 2013, Z. bicolorata became the second insect agent to be released in South Africa for the biocontrol of parthenium.     

Semiochemical footprints of the predaceous coccinellid beetle,Menochilus sexmaculatus,deter Zygogramma bicolorata (Coleoptera: Chrysomelidae) from feeding on Parthenium

Present study assessed if semiochemical cues left by an aphidophagous beetle, M. sexmaculatus influenced foraging behaviour of a phytophagous beetle, Z. bicolorata on noxious weed, P. hysterophorus. The lower predation attributes of Z. bicolorata in presence of M. sexmaculatus semiochemical footprints confirms they interfere with weed biological control.     

Developmental response of the brown-winged green bug, Plautia stali (Heteroptera: Pentatomidae), to food shortage

To study developmental response of the brown-winged green bug, Plautia stali Scott, to food shortage we reared nymphs under restricted feeding conditions produced by shortening the feeding period after molt or withholding food from second to fifth instars. For second instars, molting rates were significantly reduced as the feeding period was shortened. Shortening the feeding periods for third to fifth instars also reduced molting rates, but less so; some nymphs were able to complete their developments even if no food was given to any of the instars. Compared with controls for which feeding was not restricted, nymphs that successfully reached the next instar had reduced postmolt body size as a result of restricted feeding for all instars except the fifth (2-day feeding for second instar, and 1-day feeding for third and fourth instars), whereas instar duration was unchanged or only slightly prolonged for all instars. These results suggest that only nymphs with nutritional accumulation over a specific threshold in each instar can progress to the next instar, and that, particularly for second to fourth instars, nymphs develop on schedule without prolonging instar durations to compensate for reduced growth under conditions of food shortage.     

Host range evaluation of the leaf-feeding beetle,Zygogramma bicolorata and the stem-boring weevil,Listronotus setosipennis demonstrates their suitability for biological control of the invasive weed,Parthenium hysterophorus in Ethiopia

The invasive weed, parthenium (Parthenium hysterophorus L.) (Asteraceae: tribe Heliantheae), damages agriculture, adversely impacts biodiversity and is hazardous to human and animal health in Ethiopia. The host range of two natural enemies, a leaf-feeding beetle, Zygogramma bicolorata (Coleoptera: Chrysomelidae) and a stem-boring weevil, Listronotus setosipennis (Coleoptera: Curculionidae) was evaluated for biological control of the weed in Ethiopia. The specificity of Z. bicolorata and L. setosipennis was assessed against 29 and 31 non-target plant species, respectively. The host range of Z. bicolorata and L. setosipennis was first assessed using no-choice tests to examine their oviposition and feeding response on non-target plants. Although oviposition by Z. bicolorata occurred on six non-target species in four Asteraceae species in no-choice tests, it was significantly lower than on parthenium and no larvae developed. Zygogramma bicolorata nibbled the leaves of one of the five niger seed (Guizotia abyssinica L. – an oil seed crop closely related to parthenium) cultivars tested, but feeding and oviposition were significantly less than on parthenium. Furthermore, choice tests indicated that Z. bicolorata did not oviposit nor feed on G. abyssinica when parthenium was present. In no-choice tests, L. setosipennis did not oviposit on any of the non-target species assessed. Mean oviposition on parthenium was 39.0?±?3.4 eggs per plant whereas no eggs were laid on any of the 31 species tested. Based on these and other host range tests, permission was obtained to field release Z. bicolorata and L. setosipennis in Ethiopia.     

Does Zygogramma bicolorata Pallister really affects the growth,density and reproductive performance of Parthenium hysterophorus L?

Zygogramma bicolorata Pallister (Coleoptera: Chrysomelidae) is a satisfactory potential candidate for controlling Parthenium hysterophorus L. The study was conducted in order to evaluate the bio-control potential of Z. bicolorata to satisfactory reduced the density of parthenium weed in natural field conditions. Therefore, beetles were released in the twelve study sites @ 12 adults and/or larvae/m² in their respective sites on 5th, 10th, 15th and 20th day after initial field releases. Observations were made on defoliation by the beetle, perthenium plant height, plant biomass, leaf injury rating and soil seed reservoir. Present study showed that field releases of Z. bicolorata considerably lowering down the parthenium density at all treated sites. At larval release sites by the 60th day of observation a complete defoliation was noted, however at adult release sites it took 80 days for complete defoliation. Moreover, significantly higher number of developmental stages i.e. eggs, larvae and adults were noted in the adults released sites than to larval released sites and the controls. It was also noted that plant height and its density, flowers & leaves production, biomass of parthenium plant and seed reservoir in soil was negatively affected in treated sites as compared to untreated sites. Thus, we have concluded that adults can be utilized in biocontrol program because population build-up was greater in those sites where adults were released.     

Temperature and laboratory feeding rates in Carcinusmaenas L. (Decapoda: Portunidae) larvae from hatching through metamorphosis

Larvae of Carcinus maenas L. were reared in the laboratory and temperature-dependent stage durations of successive instars were examined at 12, 15, 18, and 25°C. Feeding rates (FR), in numbers of newly hatched Artemia nauplii, biomass, and energy consumed by a single crab larva during 24 h, were measured daily through the moulting cycles of all larval stages at the four temperatures. Dry weight (DW) and elemental content of carbon (C), nitrogen (N) and hydrogen (H) were analysed in newly hatched (0 h) and 1-day-old (24 h) Artemia nauplii at six temperatures (6, 9, 12, 15, 18, 25°C). Due to a 24 h feeding regime, the temperature dependent “mean nutritive value” of newly hatched brine shrimp nauplii is defined, individual biomass (DW, C, N, H) and energy (Joule) 12 h after hatching. General figures in changing individual daily FR, and temperature-dependent peculiarities are discussed. The total amount of food ingested by successive larval instars increases exponentially, while the increasing cumulative quantity consumed by individual crab larvae on successive days of development is described by power functions. At higher temperatures significantly less absolute biomass and energy is consumed during the entire larval development. C. maenas megalops are the main energy consumers in larval life, contributing 41 to 67% (12 to 25°C) to the total larval energy intake between hatching and metamorphosis. Larval ability to adapt to increasing metabolic costs for maintenance in higher water temperatures is discussed with average daily feeding rates (AFR). Improved efficiencies are presented for the cumulative larval energy budget, 31% in assimilation, and 4.4% in gross growth (K₁).     

Impact of defoliation by the biocontrol agentZygogramma bicolorataon the weed Partheniumhysterophorus in Australia

The leaf-feeding beetle Zygogrammabicolorata Pallister was introduced from Mexico intoAustralia in 1980 as a biocontrol agent for the weedParthenium hysterophorus L. (Asteraceae). Z. bicolorata became abundant in 1990, and since 1992there has been regular outbreaks resulting in thedefoliation of the weed in central Queensland. In thisstudy we evaluated the impact of defoliation by Z. bicolorata on P. hysterophorus from 1996 to1998. Z. bicolorata caused 91–100% defoliationresulting in reductions in weed density by 32–93%,plant height by 18–65%, plant biomass by 55–89%,flower production by 75–100%, soil seed-bank by13–86% and seedling emergence in the following seasonby 73–90%. At sites with continued outbreaks ofZ. bicolorata, it is expected that the existing soilseed-bank will be minimised, resulting in reduceddensity of parthenium in 6 to 7 years.     

Slow and fast development in Parthenium beetle and its effect on reproductive attributes

The present study was designed to investigate the existence of slow and fast developing individuals and to evaluate their effect on reproductive attributes of Parthenium beetle, Zygogramma bicolorata Pallister (Coleoptera: Chrysomelidae); a biocontrol agent of Parthenium weed. Significantly different rates of development were found in a cohort of the beetle. Under constant rearing conditions, the emergence ratio of slow and fast developers was determined to be 0.47 and 0.53, respectively. Males developed faster than females, and significant differences in development were found among all immature stages and total developmental period. Slow developing females laid higher numbers of eggs with a higher percentage egg viability than fast developing females. This is the first attempt of such a study for the Parthenium beetle, and provides great insight into Parthenium beetle development for evolutionary biologists, improving our understanding of their ecological thresholds. The information could help in the selection of fast developing lines and thus in the mass multiplication of this bioagent.     

Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

Utilization of primary nutrients by the polyphagous xylophage,Homalodisca coagulata,reared on single host species

Feeding experiments were conducted to assess development of the polyphagous leafhopper, Homalodisca coagulata, on single host species. Insects were reared from eggs on two cultivars of Euonymus japonica, two cultivars of Lagerstroemia indica, and two species of Prunus. Only insects on nonvariegated E. japonica and L. indica cv. Osage survived to maturity, although a substantial proportion of those on P. salicinia developed to the fourth instar. Chemical profiles of food source (xylem fluid), insect excreta and body composition, and consumption rates were used to estimate assimilation efficiencies, daily assimilation rates, and efficiency of conversion of food into biomass for all primary nutrients identified in the xylem fluid. Rates of successful maturation were greatest on the highest nitrogen and carbon source (nonvariegated E. japonica) and were associated with low consumption rates, high assimilation efficiencies, and prolonged instar durations. Developmental period on L. indica was shorter and consumption rates were higher, yet assimilation efficiencies were reduced and young nymphs had lower survivorship. Low consumption rates, coupled with the low nutritional value of P. salicinia, precluded sufficient accumulation of nutrients to complete development. Insects fed on hosts with xylem fluid containing high ratios of amides to total organic compounds (indicative of low carbon to nitrogen ratios) had higher consumption rates but less efficient assimilation of primary nutrients. Carcass analyses of newly enclosed adults suggest that regulation of consumption rates and assimilation efficiencies to provide adequate accumulation of essential amino acids may be pivotal for successful development. © 1996 Wiley-Liss, Inc.     

Ecological energetics of Chaoborus in a tropical lake

Summary Ecological energetics of Chaoborus brasiliensis from Lake Valencia, Venezuela, were studied between February 1979 and February 1980. Direct measurements were made of the respiration rate, assimilation efficiency, and growth rate of all 4 larval instars and of the pupae. For the larval stages, respiration increased as the 0.67 power of body mass. Respiration rates of the larvae, when corrected for body size and temperature, were extraordinarily low by comparison with the rates for most aquatic insects. The respiration rates of pupae were 3 times as high as those of larvae the same size. Assimilation rates increased significantly with body size for the larvae and differed slightly but significantly among food types. Assimilation efficiencies fell within the expected range for carnivores. The growth efficiencies were exceptionally high for instars II-IV by comparison with other small aquatic organisms. High growth efficiency for Chaoborus brasiliensis, and possibly for Chaoborus generally, is explained by a very low maintenance cost and may be a significant explanation for the wide distribution and high degree of ecological success in this primary carnivore of plankton communities.     

Feeding and growth kinetics of the planktotrophic larvae of the spionid polychaete Polydora ciliata (Johnston)

We studied the effect of food concentration on the feeding and growth rates of different larval developmental stages of the spionid polychaete Polydora ciliata. We estimated larval feeding rates as a function of food abundance by incubation experiments with two different preys, presented separately, the cryptophyte Rhodomonas salina (ESD = 9.7 µm) and the diatom T.weissflogii (ESD = 12.9 µm). Additionally, we determined larval growth rates and gross growth efficiencies (GGE) as a function of R. salina concentration.P.ciliata larvae exhibited a type II functional response. Clearance rates decreased continuously with increasing food concentration, and ingestion rates increased up to a food saturation concentration above which ingestion remained fairly constant. The food concentration at which feeding became saturated varied depending on the food type, from ca. 2 µg C mL^− 1 when feeding on T. weissflogii to ca. 5 µg C mL^− 1 when feeding on R. salina. The maximum carbon specific ingestion rates were very similar for both prey types and decreased with increasing larval size/age, from 0.67 d^− 1 for early larvae to 0.45 d^− 1 for late stage larvae. Growth rates as a function of food concentration (R. salina) followed a saturation curve; the maximum specific growth rate decreased slightly during larval development from 0.22 to 0.17 d^− 1. Maximum growth rates were reached at food concentrations ranging from 2.5 to 1.4 µg C mL^− 1 depending on larval size. The GGE, estimated as the slope of the regression equations relating specific growth rates versus specific ingestion rates, were 0.29 and 0.16 for early and intermediate larvae, respectively. The GGE, calculated specifically for each food level, decreased as the food concentration increased, from 0.53 to 0.33 for early larvae and from 0.27 to 0.20 for intermediate larval stages.From an ecological perspective, we suggest that there is a trade-off between larval feeding/growth kinetics and larval dispersal. Natural selection may favor that some meroplanktonic larvae, such as P.ciliata, present low filtration efficiency and low growth rates despite inhabiting environments with high food availability. This larval performance allows a planktonic development sufficiently long to ensure efficient larval dispersion.     

Modelling developmental changes in the carbon and nitrogen budgets of larval brachyuran crabs

The uptake and partitioning of nutritional carbon (C) and nitrogen (N) were studied during the complete larval development of a brachyuran crab,Hyas araneus, reared under constant conditions in the laboratory. Biochemical and physiological data were published in a foregoing paper, and complete budgets of C and N were now constructed from these data. Regression equations describing rates of feeding (F), growth (G), respiration (R), and ammonia excretion (U) as functions of time during individual larval moult cycles were inserted in a simulation model, in order to analyse time-dependent (i.e. developmental) patterns of variation in these parameters as well as in bioenergetic efficiencies. Absolute daily feeding rates (F; per individual) as well as carbon and nitrogen-specific rates (F/C, F/N) are in general maximum in early, and minimum in late stages of individual larval moult cycles (postmoult and premoult, respectively). Early crab zoeae may ingest equivalents of up to ca 40% body C and 30% body N per day, respectively, whereas megalopa larvae usually eat less than 10%. Also growth rates (G; G/C, G/N) reveal decreasing tendencies both during individual moult cycles and, on the average, in subsequent instars. Conversion of C and N data to lipid and protein, respectively, suggests that in all larval instars there is initially an increase in the lipid: protein ratio. Protein, however, remains clearly the predominant biochemical constituent in larval biomass. The absolute and specific values of respiration (R; R/C) and excretion (U; U/N) vary only little during the course of individual moult cycles. Thus, their significance in relation toG increases within the C and N budgets, and net growth efficiency (K ₂) decreases concurrently. Also gross growth and assimilation efficiency (K ₂; A/F) are, in general, maximum in early stages of the moult cycle (postmoult). Biochemical data suggest that lipid utilization efficiency is particularly high in early moult cycle stages, whereas protein utilization efficiency is higher in later stages. Only the zoea II appears to accumulate lipid from food constantly with a higher conversion efficiency than protein. The cumulative C and N budgets show in subsequent larval instars conspicuously increasing figures in all of their parameters.F andG increase to a particularly high extent from the first to the second zoeal instar, whereasR, U, exuvia production (G _E), and total assimilation (A) reveal a greater increase from the zoea II to the megalopa. Respiratory, excretory, and exuvial losses increase in subsequent larval instars at higher rates than tissue growth and, hence,K ₂ decreases in the same order. In the C budget,K ₂ values of 0.63 (zoea I). 0.56 (zoea II), and 0.29 (megalopa) were calculated (or: 0.56, 0.46, and 0.16 after subtraction of exuviae). In the N budget, corresponding values of 0.76, 0.66, and 0.45 (or: 0.72, 0.62, 0.38 without exuviae) were obtained. AlsoK ₁ decreases slightly in subsequent instars, whereasA/F reveals rather an increasing tendency, at least from the zoeal instars to the megalopa. Changes in the uptake and partitioning of matter in crab larvae are discussed in relation to developmental events and changes in life style before metamorphosis.     

In-vitro assessment of food consumption,utilization indices and losses promises of leafworm,Spodoptera litura (Fab.), on okra crop

The lepidopteran insect pests have significant importance in vegetable production. The present study was performed to investigate the baseline studies about the assessment of feeding and consumption potential, utilization indices and losses promises of leafworm, Spodoptera litura (Fab.) on Okra. The data regarding feeding potential, food utilization and consumption indices as well as losses of different larval instars were recorded and subjected to appropriate statistical analysis. The results showed that, in the beginning, the approximate digestibility of various instars was increase, e.g. third instar (51.36%–64.03%), fourth instar (63.42%–69.45%) and fifth instar (70.25%–76.10%). However, after a certain period, the digestibility was decreased and efficiency to convert the ingested food into biomass varied significantly. The consumption index values increased with an increase in time but the consumption and growth rate was declined of fourth instar larvae. The ingestion and digestion increased of third (10.01–13.06, 8.32–11.91 mg), fourth (11.27–17.28, 10.96–14.03 mg) and fifth (12.60–19.40, 11.93–15.28 mg) larval instars. The corrected weight of consumed leaves increased with a gain in body weight. However, in the third instar, a decline was observed on the last day of feeding. Maximum leaf area was consumed by fifth instar larvae (44.66 cm²) followed by fourth (35.41 cm²) and third (27.98 cm²) instars. In conclusion, all the dependent parameters, including food utilization potential, consumption indices and losses were higher for fifth instar larvae than others. These results emphasized the re-establishment of fundamental (economic threshold level: ETL, economic injury level: EIL) integrated pest management concepts.     

Dry-matter budgets for Diacrisia casignetum larvae fed on sunflower leaves

Larvae of Diacrisia casignetum Kollar were reared in groups of 100, 50 and 10 on fresh sunflower leaves at 27 ± 1°C temperature, 75 ± 5% r.h. and 12 h light each day. Rates of feeding, excretion and weight gain increased with age of the larva and were greatest in the sixth instar. Differences in these rates attributable to density were statistically heterogeneous (P < 0.001) whereas, of those attributable to instar, the weight gain was significant (P < 0.10). The second-instar larvae had highest levels of protein and carbohydrates in the body tissues; and the sixth-instar larvae had, the least. The pattern of protein assimilation in various instars was similar to that of glycogen except that proportional differences in the latter were pronounced in fifth and sixth instars. Consumption index, growth rate and approximate digestibility were high in the early instars whereas efficiencies of conversion of ingested and digested food were high in advanced instars. Food utilization, as reflected by efficiency of conversion of ingested and digested values in advanced instars, indicated the amount needed for maintenance decreased and more energy was left for growth and reproduction.     

Food deprivation-dependent development and fecundity in Ophraella communa

Ophraella communa LeSage is native to North America and a biological control agent of the invasive weed Ambrosia artemisiifolia L. Since A. artemisiifolia plants grow old and die after September annually, O. communa suffers from food shortage. To understand the effect of food shortage or deprivation on population fitness of O. communa, the development and fecundity and hatchability of its progeny eggs were observed when larvae were offered A. artemisiifolia plants for either 3, 6, 12 or 24 hours daily. The results showed that larval food deprivation significantly influenced survival and developmental durations of larvae and pupae. Survival rates and developmental durations of larvae and pupae decreased and were prolonged significantly with decreasing time of larval daily food intake. Longevity and fecundity of adults shortened and decreased significantly with decreasing time of larval daily food intake. In addition, the hatch rates of progeny eggs decreased significantly with decreasing time of larval daily food intake. The present study suggests that food shortage is one of the most critical factors that suppresses O. communa populations. This can explain why field populations of the beetle decrease significantly with ageing and death of A. artemisiifolia plants after late September.     

Biology and host range of Heterapoderopsis bicallosicollis: a potential biological control agent for Chinese tallow Triadica sebifera

Chinese tallow, Triadica sebifera, is an invasive weed that infests natural and agricultural areas of the south-eastern USA. A candidate insect for biological control of Chinese tallow has been studied under quarantine conditions. The biology and host range of a primitive leaf-feeding beetle, Heterapoderopsis bicallosicollis, were examined between July 2008 and February 2010. H. bicallosicollis adults fed on 16 of 29 non-target species and survived >10 days on all 29 species, more than the number of days that adults survived when deprived of food. Additionally, adult feeding, oviposition and partial larval development occurred on a Florida endangered species, Heterosavia bahamensis. These data indicated that H. bicallosicollis adult feeding and oviposition may not be limited to the target weed and included several valued North American natives. Therefore, as its release could pose unacceptable risks, testing of this species was discontinued and the quarantine colony destroyed.