The effects of water depth and density on the growth of a unionid clam*

SUMMARY. 1. Unionid clams from Narrow Lake, Alberta, were collected to quantify the natural variation in growth, to assess the natural variation in abundance, age and size distribution, and growth with water depth in the lake, and to conduct in situ experiments to directly test the effects of water depth (temperature) and clam abundance on clam growth. 2. The unionid clam, Anodonta grandis simpsoniana, showed wide variation in length at a given age. There were no significant differences in growth between clams collected at 1,3, 5, and 7m depths in the lake despite marked differences in water temperature. The wide variation in clam biomass within each depth zone may have masked possible effects of water depth. 3. The effect of water depth and variation in clam density on clam growth was tested directly by stocking clams into small enclosures at densities equivalent to 50, 150, 250, 350 and 450g m-2 (live weight) at each of 1, 3, 5 and 7 m depths in Narrow Lake (each depth and abundance treatment in triplicate). A uniform sandy substrate was used in all enclosures to eliminate any possible effect of substrate type on growth. 4. Mortality was negligible (0.9%) during the experiment. Clam density had no significant effect on clam growth which suggests that clam growth was not food limited in the lake. 5. Clams reared at 7 m grew more slowly than clams reared at 1, 3 and 5 m. Clams reared at 5 m grew more slowly than clams reared at 1 and 3m. Growth of clams reared at 1 and 3m did not differ. These differences in growth were strongly correlated with the measured differences in water temperature between depths. 6. Migration between depths probably accounts for the lack of a depth effect on clams growing in the natural habitat.     

Distinguishing Energy Maximizers from Time Minimizers: A Comparative Study of Two Hummingbird Species

SYNOPSIS. The potential reproductive success of a food energymaximizer increases with foraging time, while that of a foragingtime minimizer increases with time spent in nonforaging activitiesgiven a set energy requirement has been met. How can these foraging"goals" be distinguished for nonbreeding animals in the field?If individuals of two species occupying the same habitat consumethe same foods, face similar foraging constraints, and havesimilar meal sizes (food intake per foraging bout), then relativeto a time minimizer, an energy maximizer should: (1) spend moretime foraging, with greater foragingbout frequency, but no differencein foraging-bout duration; (2) spend less time sitting, withlower sitting-bout duration yet greater sitting-bout frequency;(3) gain mass more rapidly, if net energy intake results inmass accumulation; and (4) exhibit no other differences in timebudgeting. These assumptions and predictions were verified bypopulation- and individual-level comparisons of immature malesof two species of nectar-feeding hummingbirds studied over threefield seasons. The results suggest that, relative to each other,migrant Rufous Hummingbirds are energy maximizers and nonmigrantCosta Hummingbirds are time minimizers. Despite significantdifferences in time budgeting, by far the most striking differencebetween the species was that the Rufous gained mass four toeight times as rapidly as the Costa. This was due to the Rufousentering torpor at night, resulting in relatively little overnightloss in body mass. These patterns underscore the importanceof measuring net energy intake as directly as possible (in thiscase by fat accumulation) in testing foraging theory. Indirectmeasures (such as time budgets) may not always provide the resolutionnecessary to detect important energetic differences betweendifferent foragers.     

The Cambrian origin of the circulatory system of crustaceans

The carapace of Recent crustaceans such as myodocope ostracodes and phyllocarids is pervaded with well-developed anastomosing sinuses conveying hemolymph from the metabolizing organs to the dorsal heart. The inner lamella cuticle, which separates the sinuses from seawater, is thin enough to allow gaseous diffusion (e.g., O₂ uptake) over its surface. Comparable radiating and/or anastomosing features, of possible vascular origin, are herein recognized in several possible Crustacea from the Cambrian: cambriid, svealutiid, hipponicharionid and beyrichonid Bradoriida and in Carnarvonia from the middle Cambrian Burgess Shale. The vascular network is basically the same in these groups, consisting of sinuses radiating from supposed adductorial areas or from inferred areas of dorsal attachment of the body. The integumental (carapace sinuses) and branchial (gills) systems of respiration in crustaceans and crustacean-like animals were probably already differentiated by the middle Cambrian. The oldest record of probable integumental circulation is in the bradoriid Petrianna from the early Cambrian of Greenland. Similar circulatory systems may be represented by radiating ridges on the cephalon of other Cambrian arthropod groups such as the arachnomorphs ( Burgessia ) and trilobites ( Naraioa ) and may also be manifest in the carapaces of Ordovician-Devonian leperditicope ostracodes. Organs on the thoracopods of Cambrian supposed crustaceans, such as Canadaspis , resemble the foliaceous thoracic gills of Recent nebaliid phyllocarids and therefore may have served the same (respiratory) function.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

The usefulness of destructive host feeding parasitoids in classical biological control: theory and observation conflict

Abstract.

• 1 We examine the conventional wisdom among biological control practitioners that destructive host feeding is a desirable attribute in parasitoids employed for classical biological control, using both the predictions of population dynamics theory and historical data on biological control introductions of Hymenoptera against Homoptera.
• 2 Population dynamics theory predicts that destructive host feeders, compared with other parasitoids, are (a) either just as likely or more likely to become established, and (b) unable to depress host equilibria as strongly.
• 3 Analyses of the B IOC AT database suggest that among parasitoids of Homoptera destructive host feeders are superior to other parasitoids with respect to both establishment rate and success rate.
• 4 We present likely explanations for the disparity between the predictions of population dynamics theory and the results of database analysis. A partial explanation for the mismatch between theory and observation with respect to the degree of pest suppression may be that females of destructive host feeding parasitoids rely less upon hosts as a food source when alternative foods such as honeydew and nectar are plentiful.
• 5 We conclude that, despite the predictions of population dynamics theory, destructive host feeders are probably better biological control agents than other parasitoids, and certainly no worse, but that it would imprudent to use destructive host feeding as the sole, or even primary, selection criterion when seeking agents for classical biological control.

     

Factors affecting the co-existence of the serpentine endemic Mimulus nudatus Curran and its presumed progenitor, Mimulus guttatus Fischer ex DC

Population genetics theory suggests that two species which are reproductively isolated only by postzygotic barriers cannot co-exist in sympatry since the rarer population will become extinct. Mimulus nudatus and Mimulus guttatus are two closely related species that are isolated by a postzygotic barrier operating at the seed provisioning stage. On several sites on die serpentine soils of Lake and Napa counties, California, M. nudatus and M. guttatus live in sympatry and flowering times of the species overlap, so the species are sometimes flowering next to each other. We investigated whether there was any reduction in fertility of M. nudatus and M. guttatus caused by interspecific crosses when growing in sympatry. The pollinators of M. nudatus and M. guttatus were identified. Small sweat bees, Dialictus sp., preferentially visited the smaller flowered M. nudatus species and honey bees, Apis mellifera , preferred the larger flowered M. guttatus . In spite of most pollinator visits being intra-specinc, individuals of bodi bee species made transitions between the Mimulus species. This will result in greater pollen transfer from M. guttatus to M. nudatus than vice versa because firstly, M. guttatus produces more pollen and secondly, the sweat bees were too small to touch the stigma of a M. guttatus flower. This asymmetry in gene flow was detected by a greater reduction in viable seed produced by M. nudatus plants when surrounded by M. guttatus plants dian vice versa. Only when M. nudatus was the maternal parent could any hybrids be detected in field produced seed. To enable M. nudatus and M. guttatus to co-exist in sympatry, the two species may thus need to be sufficiendy ecologically different. Such ecological differentiation was attributed to the greater drought tolerance of M. nudatus . In addition, greenhouse experiments suggested that M. nudatus may have evolved greater tolerance to calcium deficient soils.     

Comparative body size relationships in pocket gophers and their chewing lice

In this paper, we use the method of independent contrasts to study body size relationships between pocket gophers and their chewing lice, a host-parasite system in which both host and parasite phylogcnies are well studied. The evolution of body size of chewing lice appears to be dependent only on the body size of their hosts, which confirms the 1991 findings of Harvey and Keymer. We show that there is a positive relationship between body size and hair-shaft diameter in pocket gophers, and that there is also a positive relationship between body size and head-groove width in chewing lice. Finally, we show a positive relationship between gopher hair-shaft diameter and louse head-groove width. We postulate that changes in body size of chewing lice are driven by a mechanical relationship between the parasite's head-groove dimension and the diameter of the hairs of its host. Louse species living"on larger host species may be larger simply because their hosts have thicker hairs, which requires that the lice have a wider head groove. Our study of gopher hair-shaft diameter and louse head-groove dimensions suggest that there is a 'lock-and-key' relationship between these two anatomical features.     

Salt tolerance in Eucalyptus spp.: identity and response of putative osmolytes

In four species of salt-tolerant eucalypts (Eucalyptus raveretiana, E. spathulata, E. sargentii and E. loxophleba), we found substantial concentrations of quercitol – a cyclitol known for its accumulation in seeds of Quercus. Quercitol was absent in old foliage of E. globulus, a species noted for greater susceptibility to salinity, and also absent in the moderately tolerant E. camaldulensis, but, relative to other species, both had higher foliar concentrations of inositol. Simple sugars and cyclitols accumulated to osmotically significant concentrations in all species. The osmotic potential of expressed sap was always less than that of the external ‘soil’ solution and increasing salinity produced predictable reductions in growth and increases in ion concentrations in foliage of saplings of four eucalypt species. The more salt-tolerant species, E. spathulata, E. loxophleba and E. sargentii, were able to maintain well-regulated leaf Na⁺ concentrations even at 300 mol m⁻³ NaCl. These more salt-tolerant species also showed an apparent increase in net selectivity for K⁺ over Na⁺ as salinity increased, irrespective of the Na⁺ : Ca²⁺ ratio of the external medium (range 25 : 1 to 75 : 1; Ca²⁺ always ≥ 4.0 mol m⁻³). By contrast, E. globulus was unable to exclude Na⁺ when exposed to higher NaCl concentrations (e.g. 200 and 300 mol m⁻³). Carbon isotope signatures of foliage reflected imposed salinity but were not strongly enough correlated with growth to support previous suggestions that isotope discrimination be a means of evaluating salt tolerance. On the other hand, patterns of sugar and cyclitol accumulation should be further explored in eucalypts as traits contributing to salt tolerance, and with potential use as markers in breeding programmes.