The UK risk assessment scheme for all non-native species

1. A pest risk assessment scheme, adapted from the EPPO (European and Mediterranean Plant Protection Organisation) scheme, was developed to assess the risks posed to UK species, habitats and ecosystems by non-native taxa. 2. The scheme provides a structured framework for evaluating the potential for non-native organisms, whether intentional or unintentional introductions, to enter, establish, spread and cause significant impacts in all or part of the UK. Specialist modules permit the relative importance of entry pathways, the vulnerability of receptors and the consequences of policies to be assessed and appropriate risk management options to be selected. Spreadsheets for summarising the level of risk and uncertainty, invasive attributes and economic impact were created. In addition, new methods for quantifying economic impact and summarising risk and uncertainty were explored. 3. Although designed for the UK, the scheme can readily be applied elsewhere

Cold tolerance of microarthropods

1. Microarthropods (Acari and Collembola) are dominant components of the terrestrial fauna in the Antarctic. Their cold tolerance, which forms the mainspring of their adaptational strategy, is reviewed against a background of their structure and function, and by comparison with other arthropods. 2. Two species, the isotomid collembolan Cryptopygus antarcticus Willem and the oribatid mite Alaskozetes antarcticus (Michael), are examined in detail, and afford a comparative approach to the mechanisms underlying cold tolerance in insect and arachnid types. 3. All microarthropods appear to be freezing‐susceptible (unable to tolerate tissue ice), and they utilize varying levels of supercooling to avoid freezing. Gut contents are considered to be the prime nucleation site in most arthropods when supercooled, particularly for Antarctic species. Moulting also increases individual supercooling ability especially in Collembola, and the activity of ice‐nucleating bacteria in cold‐hardy arthropods may be important. 4. Sources of ice nucleators are many and varied, originating externally (motes) or internally (ice‐nucleating agents). They act either extracellularly (mainly in the haemolymph) to promote freezing in ice‐tolerant life stages, or intracellularly in freezing‐susceptible forms. Thermal hysteresis proteins, acting colligatively, occur in many arthropods including Collembola; they depress both the freezing point of body fluids and the whole‐body supercooling point of freezing‐ susceptible and freezing‐tolerant species. 5. Bimodal supercooling point distributions are a feature of microarthropods and water droplets. Samples of field populations of Antarctic mites and springtails show significant seasonal changes in these distributions, which in some respects are analogous to purely physical systems of water droplets. Supercooling points are confirmed as accurate measures of cold‐hardiness and survival for Antarctic species, but not necessarily for other arthropods. The effects of constant sub‐zero temperatures approaching the limit of the supercooling ability of arthropods require study. 6. Desiccation and dehydration influence microarthropod physiology in several ways; in Alaskozetes it triggers glycerol synthesis. Glycerol may aid binding of water in severely dehydrated insects, but the relationship of such ‘bound’ water to cold‐hardiness is unclear. 7. Sugar alcohols (polyols) and sugars are accumulated as potential cryoprotectants in many arthropods at low temperatures, and antifreeze systems may be single or multi‐component in structure. Cryoprotectant synthesis and regulation have been studied principally in insects, and fresh weight concentrations of 0–3‐5 M of polyols have been found. Trehalose accumulation may also influence cold‐hardiness. 8. Microarthropods fall within the spectrum of cold tolerance observed for arthropods and other invertebrates. No special adaptations are found in Antarctic species, and similar strategies and mechanisms are present in both insects and arachnids. The colonization and maintenance of microarthropod populations of polar land habitats seem not to have required the evolution of any novel features with respect to cold tolerance

Temporal resolution of cold acclimation and de-acclimation in the Antarctic collembolan, Cryptopygus antarcticus

The Antarctic collembolan, Cryptopygus antarcticus (Willem), can switch its supercooling point (SCP) between 'winter' and 'summer' modes of cold hardiness over a matter of hours. High resolution temporal scaling of the acquisition and loss of cold hardiness is undertaken by assaying changes in the proportion of animals freezing below −15 °C in response to cooling rate, acclimation temperature, and access to food and moisture. Rapid de-acclimation to the 'summer' modal state is readily achieved after 1–6 h in response to warming and access to food; however, rapid acclimation to the 'winter' modal state is only evident in response to slow cooling and narrow ranges of temperature (0–5 °C). The rapid loss of cold tolerance at higher temperatures with access to food, in particular, emphasizes this species' opportunistic responses to resource availability in the short polar summers. Cold hardiness is apparently more readily traded off against nutrient acquisition than vice versa in this maritime Antarctic species

Temperature preferences of the mite, Alaskozetes antarcticus, and the collembolan, Cryptopygus antarcticus from the maritime Antarctic

The thermal preferences of Alaskozetes antarcticus (Acari, Cryptostigmata) and Cryptopygus antarcticus (Collembola, Isotomidae) were investigated over 6 h within a temperature gradient (−3 to +13 °C), under 100% relative humidity (RH) conditions. After 10 days of acclimation at −2 or +11 °C, individual supercooling points (SCP) and thermopreferences were assessed, and compared with animals maintained for 10 days under fluctuating field conditions (−6 to +7 °C). Acclimation at −2 °C lowered the mean SCP of both A. antarcticus (−24.2 ± 9.1) and C. antarcticus (−14.7 ± 7.7) compared to field samples (−19.0 ± 9.0 and −10.7 ± 5.2, respectively). Acclimation at +11 °C increased A. antarcticus mean SCP values (−13.0 ± 8.5) relative to field samples, whereas those of C. antarcticus again decreased (−16.7 ± 9.1). Mites acclimated under field conditions or at +11 °C selected temperatures between −3 and +1 °C. After acclimation at −2 °C, both species preferred +1 to +5 °C. Cryptopygus antarcticus maintained under field conditions preferred +5 to +9 °C, whereas individuals acclimated at +11 °C selected +9 to +13 °C. For A. antarcticus, thermopreference was not influenced by its cold hardened state. The distribution of field specimens was further assessed within two combined temperature and humidity gradient systems: (i) 0–3 °C/12% RH, 3–6 °C/33% RH, 6–9 °C/75% RH and 9–12 °C/100% RH and (ii) 0–3 °C/100% RH, 3–6 °C/75% RH, 6–9 °C/33% RH and 9–12 °C/12% RH. In gradient (i), C. antarcticus distributed homogeneously, but, in gradient (ii), C. antarcticus preferred 0–3 °C/100% RH. Alaskozetes antarcticus selected temperatures between 0 and +6 °C regardless of RH conditions. Cryptopygus antarcticus appears better able than A. antarcticus to opportunistically utilize developmentally favourable thermal microclimates, when moisture availability is not restricted. The distribution of A. antarcticus appears more influenced by temperature, especially during regular freeze-thaw transitions, when this species may select low temperature microhabitats to maintain a cold-hardened state

Rafting in Antarctic Collembola

Darwin was an early exponent of the importance of ‘occasional means of dispersal’ in accounting for the present-day distribution of plants and animals. This study examined the implications of capture on the water surface of meltwater and seawater for the local and long-range dispersal of Antarctic springtails. Individuals of the maritime Antarctic collembolan Cryptopygus antarcticus, were floated on tap water and seawater at 0, 5 and 10°C. LT50s on seawater were 34 (10°C), 65 (5°C) and 75 (0°C) days. On tap water, LT50s were 69 (10°C), 126 (5°C) and 239 (0°C) days. Less than 20% escaped from the water surface. A significantly greater proportion of springtails moulted on tap water and viable offspring were produced on both tap water and seawater. Comparison across treatments of survival of moulting and non-moulting individuals found significantly greater survival in moulting animals for three of the treatment combinations. It is suggested that moult exuviae facilitate survival on the water film through the simultaneous provision of a flotation aid and a source of nourishment – that is, an ‘edible raft’. A separate experiment measuring changes in haemolymph osmolality over time on tap water and seawater at 2 and 5°C found significant differences in all treatments. Causes of mortality are discussed in relation to osmoregulatory failure and starvation

Supercooling point frequency distributions in Collembola are affected by moulting

1. Many arthropods depress the freezing point of their body fluids (supercool) to avoid freezing at subzero temperatures. This is normally a seasonal response and is achieved by the production of specific biomolecules including cryoprotectants, a cessation in feeding, and the removal or masking of ice-nucleating material from their bodies. 2. In springtails, the mid-gut is shed during moulting which results in the complete evacuation of the gut and a concomitant reduction in the supercooling point (SCP). We determined whether this non-adaptive explanation could account for the variability observed in the SCP of summer-acclimatized springtails. 3. Moult preparation resulted in a highly significant reduction in the SCP. Feeding after moulting restored the SCP to previous high levels. 4. Significant differences in SCP between springtails sampled from vegetation and the soil surface, on different days, and at different sites on the same day were also documented, demonstrating that not all variation in SCP is environmentally induced. 5. Investigations of the responses of the SCP to environmental variation in springtails and other arthropods should take into account the effects of moulting before solely adaptive conclusions are drawn