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Steven L Chown

Publications and source records attributed to Steven L Chown.

35 records · Page 2Linked to original sources
PubMed2005

Biological invasions in the Antarctic: extent, impacts and implications.

Alien microbes, fungi, plants and animals occur on most of the sub-Antarctic islands and some parts of the Antarctic continent. These have arrived over approximately the last two centuries, coincident with human activity in the region. Introduction routes have varied, but are largely associated with movement of people and cargo in connection with industrial, national scientific program and tourist operations. The large majority of aliens are European in origin. They have both direct and indirect impacts on the functioning of species-poor Antarctic ecosystems, in particular including substantial loss of local biodiversity and changes to ecosystem processes. With rapid climate change occurring in some parts of Antarctica, elevated numbers of introductions and enhanced success of colonization by aliens are likely, with consequent increases in impacts on ecosystems. Mitigation measures that will substantially reduce the risk of introductions to Antarctica and the sub-Antarctic must focus on reducing propagule loads on humans, and their food, cargo, and transport vessels.

Animals↗
PubMed2005

Deleterious effects of repeated cold exposure in a freeze-tolerant sub-Antarctic caterpillar.

Multiple freeze-thaw cycles are common in alpine, polar and temperate habitats. We investigated the effects of five consecutive cycles of approx. -5 degrees C on the freeze-tolerant larvae of Pringleophaga marioni Viette (Lepidoptera: Tineidae) on sub-Antarctic Marion Island. The likelihood of freezing was positively correlated with body mass, and decreased from 70% of caterpillars that froze on initial exposure to 55% of caterpillars that froze on subsequent exposures; however, caterpillars retained their freeze tolerance and did not appear to switch to a freeze-avoiding strategy. Apart from an increase in gut water, there was no difference in body composition of caterpillars frozen 0 to 5 times, suggesting that the observed effects were not due to freezing, but rather to exposure to cold per se. Repeated cold exposure did not result in mortality, but led to decreased mass, largely accounted for by a decreased gut mass caused by cessation of feeding by caterpillars. Treatment caterpillars had fragile guts with increased lipid content, suggesting damage to the gut epithelium. These effects persisted for 5 days after the final exposure to cold, and after 30 days, treatment caterpillars had regained their pre-exposure mass, whereas their control counterparts had significantly gained mass. We show that repeated cold exposure does occur in the field, and suggest that this may be responsible for the long life cycle in P. marioni. Although mean temperatures are increasing on Marion Island, several climate change scenarios predict an increase in exposures to sub-zero temperatures, which would result in an increased generation time for P. marioni. Coupled with increased predation from introduced house mice on Marion Island, this could have severe consequences for the P. marioni population.

Acclimatization↗
PubMed2005

Insect gas exchange patterns: a phylogenetic perspective.

Most investigations of insect gas exchange patterns and the hypotheses proposed to account for their evolution have been based either on small-scale, manipulative experiments, or comparisons of a few closely related species. Despite their potential utility, no explicit, phylogeny-based, broad-scale comparative studies of the evolution of gas exchange in insects have been undertaken. This may be due partly to the preponderance of information for the endopterygotes, and its scarcity for the apterygotes and exopterygotes. Here we undertake such a broad-scale study. Information on gas exchange patterns for the large majority of insects examined to date (eight orders, 99 species) is compiled, and new information on 19 exemplar species from a further ten orders, not previously represented in the literature (Archaeognatha, Zygentoma, Ephemeroptera, Odonata, Mantodea, Mantophasmatodea, Phasmatodea, Dermaptera, Neuroptera, Trichoptera), is provided. These data are then used in a formal, phylogeny-based parsimony analysis of the evolution of gas exchange patterns at the order level. Cyclic gas exchange is likely to be the ancestral gas exchange pattern at rest (recognizing that active individuals typically show continuous gas exchange), and discontinuous gas exchange probably originated independently a minimum of five times in the Insecta.

Animals↗
PubMed2004

Cold tolerance of Littorinidae from southern Africa: intertidal snails are not constrained to freeze tolerance.

All intertidal gastropods for which cold tolerance strategies have been assessed have been shown to be freeze tolerant. Thus, freeze tolerance is considered an adaptation to the intertidal environment. We investigated the cold tolerance strategies of three species of subtropical and temperate snails (Gastropoda: Littorinidae) to determine whether this group is phylogenetically constrained to freeze tolerance. We exposed 'dry' acclimated and 'wet' rehydrated snails to low temperatures to determine temperature of crystallisation (Tc), lower lethal temperature and LT(50) and to examine the relationship between ice formation and mortality. Tc was lowest in dry Afrolittorina knysnaensis (-13.6+/-0.4 degrees C), followed by dry Echinolittorina natalensis (-10.9+/-0.2 degrees C) and wet A. knysnaensis (-10.2+/-0.2 degrees C) . The Tc of both A. knysnaensis and E. natalensis increased with rehydration, whereas Tc of dry and wet Afrolittorina africana did not differ (-9.6+/-0.2 and -9.0+/-0.2 degrees C respectively). Wet snails of all species exhibited no or low survival of inoculative freezing, whereas dry individuals of A. knysnaensis could survive subzero temperatures above -8 degrees C when freezing was inoculated with ice . In the absence of external ice, Afrolittorina knysnaensis employs a freeze-avoidance strategy of cold tolerance, the first time this has been reported for an intertidal snail, indicating that there is no family-level phylogenetic constraint to freeze tolerance. Echinolittorina natalensis and A. africana both showed pre-freeze mortality and survival of some internal ice formation, but were not cold hardy in any strict sense.

Adaptation, Physiological↗
PubMed2004

Regulated bradycardia in the pulmonate limpet Siphonaria (Gastropoda: Mollusca) during pollutant exposure: implication for biomarker studies.

Although the cardiac activity of invertebrates is now widely used as a biomarker of marine pollution, few studies consider the intrinsic factors causing the variability of heart rate (HR), including the physiological states associated with metabolic depression and behavioural isolation. We examined the cardiac responses to copper exposure of a pulmonate limpet (Siphonaria capensis Quoy and Gaimard), known to adaptively depress heart rate (and metabolic rate) under naturally stressful conditions (hyposalinity). Analysis of variance was used to assess the effects of copper concentration (0-1000 microg/L), exposure time (0-2 h), and individual difference (eight limpets per concentration) on heart rate parameters. Minute by minute heart rate (HR) and heart rate variability (HRV), and bradycardia (< or =10 beats/min) were determined from continuous recording traces. Most of the HR variance was attributable to the interaction of concentration and individuals (35.8%). Considerably lower mean HRs for the higher range of copper concentrations (250 to 1000 microg/L), compared to the lower range (0 to 50 microg/L), were attributable to the virtually exclusive induction of bradycardia in the higher concentration treatments. With increasing concentration, bradycardia was induced sooner and became less interspersed with near normal heart rates (i.e., less episodic). This regulated bradycardial response is apparently associated with isolation (avoidance) behaviour rather than with copper uptake. These findings have implications for biomarker exercises which use heart rate, in cases where invertebrates depress metabolism.

Animals↗
PubMed2004

Metabolic rate in the whip-spider, Damon annulatipes (Arachnida: Amblypygi).

Metabolic rate estimates as well as a measure of their repeatability and response to laboratory acclimation are provided for the amblypygid Damon annulatipes (Wood). This species (mean +/- S.E. mass: 640+/-66 mg) shows continuous gas exchange, as might be expected from its possession of book lungs, and at 21 degrees C has a metabolic rate of 30.22+/-2.87 microl CO2 h(-1) (approximately 229.6+/-21.8 microW, R.Q. = 0.72). The intraclass correlation coefficient (r=0.74-0.89) indicated substantial repeatability in metabolic rate which did not change with laboratory acclimation over a period of 2 weeks. By contrast, absolute metabolic rate declined by c. 16-33%, although this was not a consequence of changes in mass (which were non-significant over the same period). Rather, it appears that a reduction in overall stress or activity in the laboratory might have been responsible for the decline in mass-independent metabolic rate. At the intraspecific level, metabolic rate scaled as microW = 342 M(0.857), where mass is in grams. Metabolic rates of this species are in keeping with its sedentary behaviour such that for a given body size they are lower than those of most arthropods (spiders and insects), higher than the very sedentary ticks, and equivalent to scorpions. These findings have implications for the understanding of the evolution of metabolic rates in arthropods.

Adaptation, Physiological↗
PubMed2004

Metabolism of the sub-Antarctic caterpillar Pringleophaga marioni during cooling, freezing and thawing.

Although general models of the processes involved in insect survival of freezing exist, there have been few studies directly investigating physiological processes during cooling, freezing and thawing, without which these models remain hypothetical. Here, we use open-flow respirometry to investigate the metabolism of the freeze-tolerant sub-Antarctic caterpillar Pringleophaga marioni Viette (Lepidoptera: Tineidae) during cooling, freezing and thawing and to compare animals exposed to non-lethal (-5.8 degrees C) and lethal (-6.0 degrees C, after which caterpillars are moribund for several days, and -18 degrees C, after which caterpillars are completely unresponsive) freezing stress. We found a large decrease in metabolic rate (that is not associated with freezing) at -0.6+/-0.1 degrees C and calculated a Q10 of 2.14 x 10(3) at this breakpoint. This breakpoint is coincident with the critical thermal minimum (CTmin) and is hypothesised to be a metabolic manifestation of the latter, possibly a failure of the Na+/K(+)-ATPase pump. This provides a plausible link between processes at the cellular level and observations of the action of the CTmin at tissue and whole-organism levels. Caterpillars froze at -4.6+/-0.1 degrees C and had detectable metabolism when frozen. Post-thaw, metabolic rates were lower than pre-freezing measurements. Post-thaw metabolic rates did not differ between temperatures that did and did not kill the caterpillars, which suggests that mortality may be a result of a breakdown in processes at the organismal, rather than cellular, level of organisation.

Animals↗
PubMed2004

Upper thermal tolerance and oxygen limitation in terrestrial arthropods.

The hypothesis of oxygen limitation of thermal tolerance proposes that critical temperatures are set by a transition to anaerobic metabolism, and that upper and lower tolerances are therefore coupled. Moreover, this hypothesis has been dubbed a unifying general principle and extended from marine to terrestrial ectotherms. By contrast, in insects the upper and lower limits are decoupled, suggesting that the oxygen limitation hypothesis might not be as general as proposed. However, no direct tests of this hypothesis or its predictions have been undertaken in terrestrial species. We use a terrestrial isopod (Armadillidium vulgare) and a tenebrionid beetle (Gonocephalum simplex) to test the prediction that thermal tolerance should vary with oxygen partial pressure. Whilst in the isopod critical thermal maximum declined with declining oxygen concentration, this was not the case in the beetle. Efficient oxygen delivery via a tracheal system makes oxygen limitation of thermal tolerance, at a whole organism level, unlikely in insects. By contrast, oxygen limitation of thermal tolerances is expected to apply to species, like the isopod, in which the circulatory system contributes significantly to oxygen delivery. Because insects dominate terrestrial systems, oxygen limitation of thermal tolerance cannot be considered pervasive in this habitat, although it is a characteristic of marine species.

Adaptation, Physiological↗
PubMed2003

Rates of species introduction to a remote oceanic island.

The introduction of species to areas beyond the limits of their natural distributions has a major homogenizing influence, making previously distinct biotas more similar. The scale of introductions has frequently been commented on, but their rate and spatial pervasiveness have been less well quantified. Here, we report the findings of a detailed study of pterygote insect introductions to Gough Island, one of the most remote and supposedly pristine temperate oceanic islands, and estimate the rate at which introduced species have successfully established. Out of 99 species recorded from Gough Island, 71 are established introductions, the highest proportion documented for any Southern Ocean island. Estimating a total of approximately 233 landings on Gough Island since first human landfall, this equates to one successful establishment for every three to four landings. Generalizations drawn from other areas suggest that this may be only one-tenth of the number of pterygote species that have arrived at the island, implying that most landings may lead to the arrival of at least one alien. These rates of introduction of new species are estimated to be two to three orders of magnitude greater than background levels for Gough Island, an increase comparable to that estimated for global species extinctions (many of which occur on islands) as a consequence of human activities.

Animals↗
PubMed2003

Diurnal variation in supercooling points of three species of Collembola from Cape Hallett, Antarctica.

Daily changes in microclimate temperature and supercooling point (SCP) of Collembola were measured during summer at Cape Hallett, North Victoria Land, Antarctica. Isotoma klovstadi and Cryptopygus cisantarcticus (Isotomidae) showed bimodal SCP distributions, predominantly in the high group during the day and in the low group during the night. There were no concurrent diurnal changes in water content or haemolymph osmolality. By contrast, Friesea grisea (Neanuridae) had a unimodal distribution of SCPs that was invariant between daytime and nighttime. Isotoma klovstadi collected foraging on moss had uniformly high SCPs, which shifted towards the low group when the animals were starved for 2-8 h. When I. klovstadi was acclimated for five days with lichen or algae, SCPs were higher than if they were supplied with moss, while those that were starved (with free water or 100% relative humidity) displayed a trimodal SCP distribution. A variety of pre-treatments, including cold, heat, desiccation and slow cooling were ineffective at inducing SCP shifts in C. cisantarcticus or I. klovstadi. It is postulated that behavioural avoidance of low temperatures by vertical migration may be key in I. klovstadi's short-term survival of nighttime temperatures. These data suggest that the full range of thermal responses of Antarctic Collembola is yet to be elucidated.

Acclimatization↗
PubMed2003

Rapid responses to high temperature and desiccation but not to low temperature in the freeze tolerant sub-Antarctic caterpillar Pringleophaga marioni (Lepidoptera, Tineidae).

A broad definition of rapid cold hardening (RCH) is that it is the process whereby insects increase their survival of a sub-zero temperature after a brief (h) pre-exposure to a less severe low temperature. The effects of various pre-treatments on survival of two h at -7.9 degrees C were investigated in the freeze tolerant sub-Antarctic caterpillar Pringleophaga marioni (Lepidoptera: Tineidae), the first time RCH has been investigated in a freeze tolerant arthropod. All caterpillars froze when exposed to -7.9 degrees C, and none of the low temperature pre-treatments (-5, 0, 5 and 15 degrees C, as well as -5 degrees C and 0 degrees C with a delay before freezing) nor slow cooling (0.1 degrees C/min) elicited any improvement in survival of -7.9 degrees C as compared to controls. However, high temperature treatments (25, 30 and 35 degrees C), desiccation and acclimation for 5 days at 0 degrees C did result in significant increases in survival of the test temperature, possibly as a result of heat shock protein production. Haemolymph osmolality was elevated only by the 35 degrees C pre-treatment. It is suggested that the unpredictable environment of Marion Island means that P. marioni must always be physiologically prepared to survive cold snaps, and that this year-round cold hardiness therefore supersedes a rapid cold hardening response.

Acclimatization↗
PubMed2003

Climatic variability and the evolution of insect freeze tolerance.

Insects may survive subzero temperatures by two general strategies: Freeze-tolerant insects withstand the formation of internal ice, while freeze-avoiding insects die upon freezing. While it is widely recognized that these represent alternative strategies to survive low temperatures, and mechanistic understanding of the physical and molecular process of cold tolerance are becoming well elucidated, the reasons why one strategy or the other is adopted remain unclear. Freeze avoidance is clearly basal within the arthropod lineages, and it seems that freeze tolerance has evolved convergently at least six times among the insects (in the Blattaria, Orthoptera, Coleoptera, Hymenoptera, Diptera and Lepidoptera). Of the pterygote insect species whose cold-tolerance strategy has been reported in the literature, 29% (69 of 241 species studied) of those in the Northern Hemisphere, whereas 85 % (11 of 13 species) in the Southern Hemisphere exhibit freeze tolerance. A randomization test indicates that this predominance of freeze tolerance in the Southern Hemisphere is too great to be due to chance, and there is no evidence of a recent publication bias in favour of new reports of freeze-tolerant species. We conclude from this that the specific nature of cold insect habitats in the Southern Hemisphere, which are characterized by oceanic influence and climate variability must lead to strong selection in favour of freeze tolerance in this hemisphere. We envisage two main scenarios where it would prove advantageous for insects to be freeze tolerant. In the first, characteristic of cold continental habitats of the Northern Hemisphere, freeze tolerance allows insects to survive very low temperatures for long periods of time, and to avoid desiccation. These responses tend to be strongly seasonal, and insects in these habitats are only freeze tolerant for the overwintering period. By contrast, in mild and unpredictable environments, characteristic of habitats influenced by the Southern Ocean, freeze tolerance allows insects which habitually have ice nucleators in their guts to survive summer cold snaps, and to take advantage of mild winter periods without the need for extensive seasonal cold hardening. Thus, we conclude that the climates of the two hemispheres have led to the parallel evolution of freeze tolerance for very different reasons, and that this hemispheric difference is symptomatic of many wide-scale disparities in Northern and Southern ecological processes.

Acclimatization↗
PubMed2003

Discontinuous gas exchange and the significance of respiratory water loss in Scarabaeine beetles.

Respiratory water loss in insects is a controversial topic. Whilst earlier studies considered respiratory transpiration a significant component of overall water loss, to the extent that it was thought to be responsible not only for the evolution of discontinuous gas exchange cycles (DGCs) but also for variation in DGC patterns, later work repeatedly questioned its importance. In particular, investigations of the proportional contribution of respiratory transpiration to total water loss in species showing DGCs suggested that respiratory transpiration was unlikely to be important in these species. In turn, these studies have been criticized on analytical grounds. In this study we investigated variation in cuticular and respiratory water loss rates in five Scarabaeus dung beetle species, all of which show discontinuous gas exchange cycles, to ascertain the significance of respiratory water loss using modern analytical techniques. In particular, we determined whether there is variation in water loss rates amongst these beetles, whether both respiratory and cuticular water loss rates contribute significantly to variation in the former, and whether metabolic rate variation and variation in the duration of the DGC periods contribute significantly to variation in respiratory water loss rate. Total water loss rate varied such that species from arid areas had the lowest rates of water loss, and both cuticular and spiracular transpiration contributed significantly to variation in overall water loss rate. Moreover, variation in metabolic rate and in the duration of the DGC periods contributed significantly to variation in respiratory water loss rate. By contrast, examination of proportional water loss revealed little other than that it varies between 6.5% and 21%, depending on the species and the temperature at which it was examined. Cuticular water loss scaled as mass(0.721), but did not differ from that expected from geometric considerations alone. By contrast, respiratory water loss scaled as mass(0.531), suggesting that gas exchange takes place by diffusion and convection. Our results provide direct evidence that respiratory water loss forms a significant component of water balance, and that changes in both metabolic rate and DGC characteristics contribute to modulation of respiratory water loss.

Animals↗
PubMed2003

Repeatability of standard metabolic rate and gas exchange characteristics in a highly variable cockroach, Perisphaeria sp.

For natural selection to take place several conditions must be met, including consistent variation among individuals. Although this assumption is increasingly being explored in vertebrates, it has rarely been investigated for insect physiological traits, although variation in these traits is usually assumed to be adaptive. We investigated repeatability (r) of metabolic rate and gas exchange characteristics in a highly variable Perisphaeria cockroach species. Although this species shows four distinct gas exchange patterns at rest, metabolic rate (r=0.51) and the bulk of the gas exchange characteristics (r=0.08-0.91, median=0.42) showed high and significant repeatabilities. Repeatabilities were generally lower in those cases where the effects of body size were removed prior to estimation of r. However, we argue that because selection is likely to act on the trait of an animal of a given size, rather than on the residual variation of that trait once size has been accounted for, size correction is inappropriate. Our results provide support for consistency of variation among individuals, which is one of the prerequisites of natural selection that is infrequently tested in insects.

Animals↗
PubMed2002

Discontinuous gas-exchange in centipedes and its convergent evolution in tracheated arthropods.

We have examined the gas-exchange characteristics of five southern African centipede species from three orders. Two scolopendromorph species exhibit discontinuous gas-exchange cycles (DGCs) identical to those recorded for several insect and chelicerate species. Another scolopendromorph and a lithobiomorph species exhibit weak periodic patterns, and a scutigermorph species shows continuous gas exchange. A crucial component for DGCs in tracheated arthropods is the presence of occludible spiracles. However, on the basis of studies of temperate centipedes, most recent invertebrate biology texts hold the view that centipedes, as a group, cannot close their spiracles. Using flow-through normoxic and normoxic-anoxic-normoxic respirometry and electron microscopy, we conclusively demonstrate that at least one of the scolopendromorph species, Cormocephalus morsitans L., can close its spiracles fully, thus accounting for its DGCs. Homologies in spiracular structure and DGCs suggest that several other tracheated arthropod taxa probably have this ability too and that DGCs have evolved convergently at least four times in the Arthropoda. Spiracular closure and discontinuous gas-exchange cycles are probably more widespread in arthropods than has previously been suspected.

Animals↗
PubMedDate not supplied

Water-balance characteristics respond to changes in body size in subantarctic weevils.

Several environmental factors leading to size-dependent mortality influence insect body size. Few investigations have been concerned with the ways in which the mechanisms underlying variation in water-balance characteristics evolve in response to changes in body size that occur independently of water-balance requirements. Using an explicitly phylogenetic analysis, we show how body size has changed over time in the Ectemnorhinus group of weevils and how water-balance characteristics have evolved in response to this change and changes in habitat use. The basal species in the group are all large bodied and from moist environments. In response to a change in resource availability, there was a marked decline in size within the group. Despite the reduction in water content and dehydration tolerance that this meant, evolution of low whole-animal water-loss rates and high tolerance of dehydration resulted in conservation of desiccation resistance. The return to moist habitats in the group resulted in a reduction in dehydration tolerance and an increase in water-loss rate. Thus, dehydration tolerance and water-loss rate respond rapidly both when there is selection for water conservation and when this requirement is relaxed. Future laboratory selection experiments might usefully explore both directions of water-balance evolution.

Animals↗