Stage-specific effects of temperature and dietary protein on growth and survival of Manduca sexta caterpillars

The effects of temperature and dietary protein concentration on growth and survival of Manduca sexta L. (Lepidoptera: Sphingidae) caterpillars during different larval stages were examined. Sets of caterpillars were raised from hatching at one of five constant temperatures (18, 22, 26, 30 or 34°C) and on one of two artificial diets (low or high protein concentration). Mass gain, duration (development time) and mean growth rate were measured for each caterpillar for the 1st to 3rd stadia, the 4th stadium, and the 5th stadium. Temperature significantly affected mass gain during each larval stage, resulting in smaller mass gains at higher temperatures at each stage. This effect was strongest at high temperatures during the 5th stadium. Temperature significantly affected durations of each larval stage, but the effect varied among stages: for example, the duration of stadia 1–3 decreased continuously with increasing temperature, whereas the duration of the 5th stadium was shortest at 26–30°C and increased at lower and higher temperatures. The effect of temperature on mean growth rate changed dramatically across larval stages: maximal growth rate occurred at 34°C during the 1st to 3rd stadia, at 30°C during the 4th stadium and at 26°C during the 5th stadium. Higher dietary protein concentration significantly decreased the duration of stadia 1–3 and of the 4th stadium, but had no significant effect on the duration of the 5th stadium. Temperature and dietary protein had little effect on mortality rates during any larval stadium, with one exception: mortality during the 5th stadium increased dramatically at temperatures of 30 and 34°C. These results demonstrate that the effects of temperature and dietary protein concentration on growth, development and survival in M. sexta vary markedly in different larval stadia during development; 5th instar caterpillars are particularly sensitive to higher temperatures.     

Anamorphosis and life-history of the millipede Nopoiulus kochii (Gervais, 1847), new for Egypt

Abstract

Monthly samples were taken to determine the anamorphosis and the life-history of the millipede Nopoiulus kochii, new to Egypt. Stadial determination was possible from counting the podous and apodous rings, rows of ocelli, and serial additions of defence glands. Sexual dimorphism occurred at stadium IV. Maturity is achieved at stadium VII. Few males, however, attained maturity at stadium VI. Further moultings of adult stadia occur so that additional adults are found from stadia VII to XIV. N. kochii breeds at the age of six months.     

Ecology of Chondromorpha kelaarti (Diplopoda, Polydesmida)

Stadia 11-VII and adults of Chondromorpha kelaarti (Humbert) can be separated by segment number, length, breadth and paired legs. Sexes are distinguished from stadium IV and the sex ratio vaned around SO%, though females were prevalent in May and June.
In females, a positive and significant correlation was found between number ofeggs present and body width at the 17th Segment.
Population density and biomass of C. kelaarti varied from 0.06 to 12.94 and 1.75 mg to 811.24 mg per 0.1 mz, respectively. From May to November, populations of C. kelaartiare aggregated in distribution.
Chondronrorphu keluurti passed the dry season in late stadia (VI, VII and adult) and may complete the life cycle in one or two years.     

The effect of temperature on feeding,growth, and metabolism during the last larval stadium of the female house cricket, Acheta domesticus

Eighth instar female house crickets at 35°C developed faster, gained slightly more wet weight, and consumed less food, water, and oxygen than at 25°C. The duration of the 8th stadium at 25°C was 13 days (undisturbed), but was 14 days when disturbed by daily weighing. The duration of the 8th stadium at 30°C was 8 days and at 35°C was 6 days. During the first half of the 8th stadium at 25, 30, and 35°C, there was a high rate of food and water consumption resulting in statistically equal maximum dry weight achievement (124 mg). Respiratory quotients greater than one during this time indicated the conversion of ingested carbohydrate to fat. During the latter half of the 8th stadium, food and water consumption declined and the crickets lost weight. The period of weight loss was proportionally much longer at 25°C than at 30 or 35°C. Respiratory quotients lower than 1.0 during the latter half of the 8th stadium at 30 and 35°C indicated the metabolism of stored lipids. The respiratory quotient at 25°C never fell below 1.0, possibly because some food remained in the gut. The absorption efficiency was not influenced by temperature (25–35°C). Though the caloric content of the faeces was lower at 25°C than at 30 or 35°C, which correlated to the much longer time for food passage at 25°C than at 35°C, the difference in total calories egested was insufficient to alter the absorption efficiency. A longer period of reduced feeding and greater dry weight loss during the latter half of the 8th stadium at 25°C resulted in a lower metabolic efficiency at 25°C than at 30 or 35°C. Eighth instar crickets in response to a step-function transfer from 30°C–25 or 35°C showed an immediate (<1 hr) and complete metabolic adjustment which was not affected by the temperature history during the 7th stadium. House crickets did not exhibit temperature acclimation in the range 20–40°C, the metabolic rate being determined by ambient temperature. The Q₁₀ for oxygen consumption in the range 20–40°C was about 2.     

The life history and ecology of the millipede Tymbodesmus falcatus (Polydesmida: Gomphoclesmidae) in northern Nigeria with notes on Sphenodesmus sheribongensis

The life history of the gomphodesmid millipede Tymbodesmus falcatus in Zaria, Nigeria is described. Larvae of stadia I to VI are soil-dwelling, stadium VII and adults are surface active.
Tymbodesmus probably completes its life cycle in two years; the adults are relatively shortlived and die before the onset of the dry season. The species survives the dry season as dia-pausing larvae in moulting chambers in the soil.
The biology of Sphenodesmus is, in general, similar to that of Tymbodesmus.
Body form in the species is described and compared with that found in other polydesmoids. A possible relationship between brood size and the degree of surface activity in polydesmoids is discussed.     

The effect of temperature on energy distribution during the last-larval stadium of the female house cricket, Acheta domesticus

The distribution of energy during the last stadium of the house cricket at two temperatures was the main theme of this study. Food consumption, growth, and oxygen consumption were greater in the first half of the stadium at both 25 and 35°C. An RQ > 1 indicated the conversion of carbohydrates to lipids during the first half of the instar at both temperatures. The duration of the stadium increased from 6 days at 35°C to 14 days at 25°C. The same maximal weight, protein content and lipid content were attained at both 25 and 35°C. A weight loss (mostly in stored lipids) after the midstadium peak weight was greater at the lower temperature. The absorption efficiency and the production of metabolic wastes were not affected by temperature, but the metabolic efficiency was much higher at 35 than at 25°C during the first half as well as the latter half of the stadium. Although during the first half of the stadium more energy was ingested, absorbed, and made available for growth at 25 than at 35°C, only slightly more growth occurred at 25°C. During the last half of the stadium less energy was ingested at 25 than at 35°C, and much more growth occurred at 35°C because of the even greater heat loss at 25 than at 35°C. Therefore at a lower temperature cricket larvae eat slightly more and reach the same maximal weight as at a higher temperature, but they end up smaller because they waste more energy during the extended duration of the stadium at the lower temperature.     

Artificial incubation of California condor Gymnogyps californianus eggs removed from the wild

The currrent California condor (Gymnogyps californianus) recovery plan entails increasing the reproductive rate via replacement-clutch manipulation of eggs. During the period from 1983 to 1985, 15 eggs were removed from wild nesting pairs for artificial incubation. The eggs were incubated at a dry bulb temperature of 36.4°C in modified forced-air Lyon Electric incubators. The incubation humidity was adjusted for individual eggs based on weight loss data (water = weight), 25.6–30.0°C wet bulb (41.0–63.0% Relative Humidity (RH)). The chicks were hatched initially under forced-air conditions of 36.1°C dry bulb, 31.1–01.7°C wet bulb (70.0–73.0% RH). In 1984, hatching parameters were changed to still-air conditions, 36.1°C dry bulb (top of the egg), 35.0°C dry bulb (bottom of the egg), 31.1–31.7°C wet bulb (70.0-73.0% RH). Tactile and auditory stimulation was utilized during the pip-to-hatch interval. From among 15 eggs collected, 13 hatched, and 12 condor chicks were raised successfully (hatchability: 86.7%; survivability: 92.3%).     

Estimating reaction norms for predictive population parameters,age specific mortality,and mean longevity in temperature‐dependent cohorts of Culex quinquefasciatus Say (Diptera: Culicidae)

Culex quinquefasciatus plays a major role in the transmission of important parasites and viruses throughout the world. Because temperature is an important limiting factor on growth and longevity of all mosquito species, estimating the reaction norms provides very important basic information for understanding both plasticity and individual variations of the population. In the present study, Cx. quinquefasciatus were maintained at five different constant temperatures (15°, 20°, 23°, 27°, and 30°C) for two subsequent generations. Reproductive population parameters in blood‐fed mated females and longevities of virgin and blood‐fed mated adults reared at different temperatures were compared for the two generations. Longevity increased as temperature decreased within a range of 15° to 30°C for the unmated adults, and 15° to 27°C for the mated and blood‐fed adults. Generation times were as long as 124.07 and 106.76 days for two subsequent generations reared at 15°C, and the highest intrinsic rate of increase (r_m) values were estimated at 0.22 and 0.18, respectively, from the cohorts reared at 27°C. For survival rates, reproductive rates (R₀), and r_m values, 30°C was found to be a critical temperature for this species. These cohorts produced the smallest amount of eggs (R₀= 5.06), r_m values decreasing across generations (from 0.11 to 0.06), and the survival rates from egg to adult were found to be insufficient (16.1 and 10.8%). Additionally, the rate of exponential increase with age and age specific mortalities (b) were calculated for the virgin cohorts. Age specific mortality rates increased as temperature decreased. The increase in mortality rates started to accelerate at 27°C and was more pronounced at 30°C, for both females and males. We estimated the coefficients of variation for the b values in which females have smaller coefficients than those of the males at all temperatures.     

Interactions of Adult Stoneflies (Plecoptera) with Riparian Zones I. Effects of Air Temperature and Humidity on Longevity

The effects of constant air temperature and relative humidity on the longevity of three species of gripopterygid stonefly adults from New Zealand were investigated in laboratory experiments, and the results were compared to field measurements of air temperature and humidity obtained during summer. Greatest longevity for Zelandobius furcillatus, Zelandoperla decorata and Acroperla trivacuata was recorded in cool humid conditions (10°C, 100% humidity) for adults fed water and a 5% sucrose solution. Absence of feeding reduced survival by 37–73% at 17°C and 100% humidity. Survival decreased significantly with increasing constant air temperature (10, 17 and 25°C) and decreasing mean relative humidity (100, 81 and 15%). Males survived significantly longer than females in all temperature treatments for Z. furcillatus, but longevity was not influenced by gender in other species or in the humidity experiments. Interpolated LT 50 values over 96 h for female stoneflies in the temperature treatments averaged of 22–23°C. Field measurements at near-ground level and 1.5 m above the streambank during summer indicated that these air temperatures were exceeded for 25% of the time in a pasture catchment compared to &lt;0.1% of the time in a native forest catchment. These findings implicate air temperature as a factor potentially influencing the longevity of adult stoneflies, and suggest that maintenance of appropriate microclimate conditions should be a consideration in riparian management.     

The effect of temperature on embryo survival and development in pallid sturgeon Scaphirhynchus albus (Forbes & Richardson 1905) and shovelnose sturgeon S. platorynchus (Rafinesque, 1820)

The thermal response of pallid sturgeon Scaphirhynchus albus and shovelnose sturgeon S. platorynchus embryos was determined at incubation temperatures from 8 to 26°C and 8 to 28°C, respectively. The upper and lower temperatures with 100% (LT₁₀₀) embryo mortality were 8 and 26°C for pallid sturgeon and 8 and 28°C for shovelnose sturgeon. It was concluded that 12–24°C is the approximate thermal niche for embryos of both species. Generalized additive and additive‐mixed models were used to analyze survival, developmental rate and dry weight data, and predict an optimal temperature for embryo incubation. Pallid sturgeon and shovelnose sturgeon embryo survival rates were different in intermediate and extreme temperatures. The estimated optimal temperature for embryo survival was 17–18°C for both species. A significant interaction between rate of development and temperature was found in each species. No evidence was found for a difference in timing of blastopore, neural tube closure, or formation of an S‐shaped heart between species at similar temperatures. The estimated effects of temperature on developmental rate ranged from linear to exponential shapes. The relationship for rate of development to temperature was relatively linear from 12°C to 20°C and increasingly curvilinear at temperatures exceeding 20°C, suggesting an optimal temperature near 20°C. Though significant differences in mean dry weights between species were observed, both predicted maximum weights occurred at approximately 18°C, suggesting a temperature optimum near 18°C for metabolic processes. Using thermal optimums and tolerances of embryos as a proxy to estimate spawning distributions of adults in a river with a naturally vernalized thermal regime, it is predicted that pallid sturgeon and shovelnose sturgeon spawn in the wild from 12°C to 24°C, with mass spawning likely occurring from 16°C to 20°C and with fewer individuals spawning from 12 to 15°C and 21 to 24°C. Hypolimnetic releases from Missouri River dams were examined; it was concluded that the cooler water has the potential to inhibit and delay sturgeon spawning and impede embryo incubation in areas downstream of the dams. Further investigations into this area, including potential mitigative solutions, are warranted.     

Developmental changes in the patterns of feeding in fourth- and fifth-instar Helicoverpa armigera caterpillars

Data are presented for developmental changes in feeding behaviour within and across the fourth and fifth stadium of Helicoverpa armigera (Lepidoptera, Noctuidae) caterpillars fed nutritionally homogeneous semi‐synthetic foods. We recorded the microstructure of feeding over continuous 12‐h periods on consecutive days throughout the two stadia, and in one experiment recorded continuously for 21 h. Larvae in the two stadia showed the same general pattern of macro‐events in feeding, including a similar duration of post‐ecdysis fast, which was usually broken by consumption of the exuviae, and then a sustained period in which discrete meals on the experimental food were taken regularly. There were, however, some distinct differences in the patterns of meal‐taking both between stadia and across different one‐third time segments within stadia. Considering between‐stadium differences, the proportion of time spent feeding differed significantly only in the last segment of the feeding period of the two stadia, with the value for the fourth‐instar larvae being substantially greater than for fifth‐instar larvae. As regards within stadium changes, the proportion of time feeding increased from the first to the second segment of both stadia. However, whereas the proportion of time feeding increased from the second to the final segment of the fourth stadium, it decreased across the same period in the fifth stadium. These patterns of changes in the proportion of time feeding within and between stadia, and their behavioural mechanisms (combination of meal durations and meal frequencies), can be explained only partially with reference to increasing food requirements with development. Three areas are identified where further study might help elucidate the reasons for the observed developmental changes in the microstructure of feeding: allometric constraint, the dynamic links between ingestion and post‐ingestive processing, and ecological factors such as predation.     

Effect of temperature on yield and night biomass loss in Spirulina platensis grown outdoors in tubular photobioreactors

Outdoor experiments carried out in Florence, Italy (latitude 43.8° N, longitude 11.3° E), using tubular photobioreactors have shown that in summer the average net productivity of a Spirulina platensis culture grown at the optimal temperature of 35 °C was superior by 23% to that observed in a culture grown at 25 °C. The rates of night biomass loss were higher in the culture grown at 25 °C (average 7.6% of total dry weight) than in the one grown at 35 °C (average 5%). Night biomass loss depended on the temperature and light irradiance at which the cultures were grown, since these factors influenced the biomass composition. A net increase in carbohydrate synthesis occurred when the culture was grown at a low biomass concentration under high light irradiance or at the suboptimal temperature of 25 °C. Excess carbohydrate synthesized during the day was only partially utilized for night protein synthesis.     

Calorimetry of rat tail tendon collagen before and after denaturation: the heat of fusion of its absorbed water

The specific heat, of rat tail tendon at various water contents was measured as a function of temperature. The resulting graphs showed peaks arising from the melting, near 50°C, of helical material in the collagen, and from the melting of absorbed water in the range -40°C to 0°C. The heat of melting of helical material was 11.7 cal per gram of dry tendon. Determination of the heat and temperature of fusion of the absorbed water allowed resolution of the water into four states in the case of tendon before denaturation, and three states after denaturation. The four states are (1) water not freezable on cooling to - 70°C, (2) freezable water with-both heat and temperature of fusion different from the values for ordinary water, (3) freezable water with the heat of fusion of ordinary water, but a different temperature of fusion, and (4) water not distinguished from ordinary water. The fourth state was absent in denatured tendon. The results are discussed in terms of increasing size of clusters of absorbed water molecules.     

PHYSIOLOGICAL COMPARISON OF THE ISOMORPHIC LIFE HISTORY PHASES OF THE HIGH INTERTIDAL ALGA ENDOCLADIA MURICATA (RHODOPHYTA)1

The isomorphic phases of Endocladia muricata (Post. & Rupr.) J. Ag. Were compared for photosynthetic and respiratory difference in response to a variety of environmental manipulations. Photosynthetic light response during submergence at 15° C and the pattern of respiratory recovery following prolonged emergence (3 h) at either 15° or 30° C were similar between gametophytes and tetrasporophytes. The phases showed the same ability to photosynthesize and respire during emergence at each temperature tested (15°, 25°, and 35° C, fully hydrated thalli) and at various desiccation state (measured at 25° C only). Submerged rates of photosynthesis following prolonged emergence at 15° and 30° C were, however slightly greater (17%) for tetrasporophytes as compared to gametophytes. Regardless of the life history phase, plants incubated at 15° C during emergence recovered more completely than plants incubated at 30° C. Photosynthetic recovery after 1 h in plants incubated at 15° C often “spiked” and yielded rates as great as 185% of pretreatment rates. Increased photosynthetic rates during recovery were absent for the 30° C incubations. The initial photosynthetic recovery of plants collected from the upper limits of distribution was greater than that of plants collected from the lower limits. Recovered rates of respiration were highly variable over time. Respiration often exceeded pretreatment values more then threefold, and the elevated rates were sustained for 12 h. Photosynthesis and respiration in air were comparable to rates in seawater and varied slightly with increasing temperature. Photosynthetic and respiratory rates also decreased with increasing tissue water loss. Thus, only slight differences in physiological performance were observed between phases and individuals collected from different vertical positions. Metabolic differences were transient and apparent only under experimental conditions that modeled extreme environmental conditions.     

Variation in growth attributes of contrasting populations of Trifolium repens

A range of growth attributes was measured in seedlings of 10 Trifolium repens populations, differing in leaf size and origin, grown in three temperature and two glasshouse environments. Growth rates of large leaf types of Mediterranean origin were higher than those of smaller leaf types at 10°C. However, the greater temperature response of the smaller leaf types resulted in higher growth rates for S.100 and S.184 than for a large leaf type from Israel at 20°C. The increase of growth rate with temperature was associated with changes in leaf area ratio and net assimilation rate between 10° and 15°C but only with changes in net assimilation rate between 15° and 20°C. Within each temperature environment, population differences in growth rate were related to differences in net assimilation rate rather than leaf expansion. At low temperature a greater proportion of dry matter was distributed to leaf tissue in large leaf types particularly those of Mediterranean origin but they showed a proportionately smaller increase in allocation to leaves with increasing temperature compared with small leaf types. In the glasshouse environments growth rates in spring were more than double those in the autumn. This difference was associated with net assimilation rates which were about five times greater in the spring environment. However, leaf area ratios in the spring were only half those in the autumn. These differences in leaf area ratio between the glasshouse environments were closely related to differences in specific leaf area but not to differences in distribution of dry matter to leaf tissue which was greater in the spring environment.     

Effects of temperature on growth and water relations of cacao (Theobroma cacao var.Comum) seedlings

Growth of 55-day-oldTheobroma cacao var.Comum seedlings varied with temperature regimes, various plant parts, growth parameters, and time of harvesting. Over a 60-day period the optimal day-temperature regimes were near 33.3°C for dry weight increase and relative growth rates of seedlings and leaves; 30.5°C for increase in leaf area, height growth, and leaf abscission; 22.2°C for dry weight increase of stems or roots, stem diameter growth, and root-shoot ratio. The rates of increase in dry weights of stems or roots as well as root-shoot ratios declined progressively at temperatures above 22.2°C Partitioning of dry matter was affected by temperature regime, with proportionally more photosynthate retained by shoots and less translocated to roots at high temperatures. The progressive decrease in the root-shoot ratio at temperatures above 22.2°C may decrease drought tolerance of seedlings because roots will be less capable of absorbing endugh water to replace transpirational losses. This was shown by more negative shoot water potentials at high temperatures. Research supported by the College of Agricultural and Life Science. University of Wisconsin, Madison, Wi, USA and the Centro de Pesquisas do Cacau (CEPLAC/CEPEC), 45600 Itabuna, Ba, Brazil.     

Social influences on nymphal development in the cockroach, Diploptera punctata

Solitary male nymphs of the cockroach Diploptera punctata (Eschscholtz) (Blattaria: Blaberidae) took significantly longer to reach adulthood than males paired with either a male or female nymph or grouped with four other male nymphs since birth. When isolated throughout nymphal development, 15.8% of males passed through 3 stadia before adult eclosion, and the remainder went through 4 stadia. In contrast, 61.3% of paired males became adults in 3 stadia. Males need not, however, be isolated or paired for the entire nymphal period to express isolated or paired patterns of development. About 60% of males paired in just the first stadium or its initial 9 days became adults in 3 stadia, and only 20.4% of males isolated in the first stadium and the first 3 days of the second reached adulthood within 3 stadia. Although the first stadium was a critical period in which social condition determined the course of future development, analyses of covariance showed that isolated males gained less weight than paired ones, not only in the first stadium, but in the second as well. Moreover, the degree of growth of a male in the second stadium, measured as either weight gain or relative growth rate, did not depend on the male's social condition in the first stadium, because isolated second-instar males grew less than paired ones, even when both sets of insects had been paired in the first stadium. Female nymphal development, unlike that of males, was not greatly affected by social factors.     

Effect of temperature on development, overwintering and establishment potential of Franklinothrips vespiformis in the UK

This study investigated the effect of temperature on the development and overwintering potential of the predatory thrips Franklinothrips vespiformis (Crawford) (Thysanoptera: Aeolothripidae), a biological control agent used against glasshouse pests in continental Europe and Israel. Developmental rates increased linearly with rearing temperatures. It was estimated that 304.9 degree days, above a lower threshold temperature of 11.9 °C, were required for F. vespiformis to complete development from egg to adult eclosion. The effect of low temperatures (–5, 0, and 5 °C) was examined on adult female and larval survival. Subsequent reproductive and developmental attributes of survivors were also investigated. Lethal time experiments indicated that larval stages are more cold tolerant than adult F. vespiformis females. Surviving larvae increased their developmental times to adults with decreasing temperature and increasing exposure periods and second instars were significantly more successful than first instars in reaching adulthood. Surviving adult females decreased their oviposition rate with decreasing temperature and increasing exposure periods, and exposures to low temperatures affected the number of viable eggs produced. The results are discussed in the context of overwintering and establishment potential of F. vespiformis in the UK in the event of introducing the predatory thrips as a biological control agent against glasshouse pests.     

Water and energy fluxes during summer in an arid‐zone passerine bird

Endothermic animals resident in hot, arid terrestrial environments are likely to face a trade‐off between their ability to obtain water and elevated thermoregulatory water requirements. We assessed whether daily water flux (DWF) is higher on hot days, reflecting increases in evaporative cooling demands, in an arid‐zone bird that obtains its water through food intake. We obtained measurements of DWF (partitioned into water influx and efflux rates) in 71 White‐browed Sparrow‐Weavers Plocepasser mahali at a desert site and a semi‐desert site, during summer in the Kalahari Desert of southern Africa. We found no evidence that DWF varied with maximum daily air temperature (T_air, range = 27.6–39.2 °C). Instead, DWF was lower during dry periods than in the wet season at the semi‐desert site. Furthermore, birds showed deficits in water balance (water influx/water efflux) during the dry periods at both sites. Our data show that DWF is low in a non‐drinking bird that obtains its water through food, and that demands for evaporative water loss on very hot days (maximum T_air of 40–44 °C) may exceed water intake rates during hot and dry periods. Species that do not have opportunities to drink will experience strong trade‐offs between thermoregulation, hydration state and activity levels as temperatures increase.     

Increasing temperature decreases the predatory effect of the intertidal shanny Lipophrys pholis on an amphipod prey

Interactions between Lipophrys pholis and its amphipod prey Echinogammarus marinus were used to investigate the effect of changing water temperatures, comparing current and predicted mean summer temperatures. Contrary to expectations, predator attack rates significantly decreased with increasing temperature. Handling times were significantly longer at 19° C than at 17 and 15° C and the maximum feeding estimate was significantly lower at 19° C than at 17° C. Functional‐response type changed from a destabilizing type II to the more stabilizing type III with a temperature increase to 19° C. This suggests that a temperature increase can mediate refuge for prey at low densities. Predatory pressure by teleosts may be dampened by a large increase in temperature (here from 15 to 19° C), but a short‐term and smaller temperature increase (to 17° C) may increase destabilizing resource consumption due to high maximum feeding rates; this has implications for the stability of important intertidal ecosystems during warming events.