Microhabitat use by a post-settlement stage estuarine fish: evidence from relative abundance and predation among habitats

Seagrass beds provide food and shelter for many fish species. However, the manner in which fishes use seagrass bed habitats often varies with life stage. Juvenile fishes can be especially dependent on seagrass beds because seagrass and associated habitats (drift macroalgae) may provide an effective tradeoff between shelter from predation and availability of prey. This study addressed aspects of habitat use by post-settlement pinfish, Lagodon rhomboides (Linneaus), an abundant and trophically important species in seagrass beds in the western North Atlantic and Gulf of Mexico. Abundance of post-settlement fish in seagrass beds was positively related to volume of drift macroalgae, but not to percent cover of seagrass, indicating a possible shelter advantage of the spatially complex algae. Tethering experiments indicated higher rates of predation in seagrass without drift macroalgae than in seagrass with drift macroalgae. Aquarium experiments showed lower predation with higher habitat complexity, but differences were only significant for the most extreme cases (unvegetated bottom, highest macrophyte cover). Levels of dissolved oxygen did not differ between vegetated and unvegetated habitats, indicating no physiological advantage for any habitat. Seagrass beds with drift macroalgae provide the most advantageous tradeoff between foraging and protection from predation for post-settlement L. rhomboides. The complex three-dimensional shelter of drift macroalgae provides an effective shelter that is embedded in the foraging habitat provided by seagrass. Drift macroalgae in seagrass beds is a beneficial habitat for post-settlement L. rhomboides by reducing the risk of predation, and by providing post-settlement habitat within the mosaic (seagrass beds) of adult habitat, thus reducing risks associated with ontogenetic habitat shifts.     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Effect of the removal of cleaner fish on the abundance and species composition of reef fish

The ecological significance of cleaner fish on coral reefs was investigated. I removed all cleaner fish, Labroides dimidiatus, from eight small reefs, measured the subsequent effect on the abundance and species composition of all reef fish after 3 and 6 months, and compared it with eight control reefs with cleaner fish. The removal of cleaner fish had no detectable effect on the total abundance of fish on reefs and the total number of fish species at both times. Multivariate analysis by non-metric multidimensional scaling and ANOSIM pairwise tests based on 191 fish species revealed no effect of cleaners on the community structure of fish. Similar results were obtained using principal components analysis on subsets of the data using the 33 most common fish species and the 15 most abundant species (≥5 individuals per reef ) with both log10 (x + 1) transformed data and with fish numbers standardized for abundance. This study demonstrates that the removal of cleaner fish for 6 months did not result in fish suffering increased mortality nor in fish leaving reefs to seek cleaning elsewhere. Received: 28 October 1996 / Accepted: 7 February 1997     

Life on the edge: conspecific attraction and recruitment of populations to disturbed habitats

Variation in the recruitment of benthic marine invertebrates is often attributed to the interaction of the supply of new individuals to a habitat and the availability of space for colonisation when they arrive. Also important in determining variation in recruitment is the response of the larvae to the characteristics of the habitat. Larvae of many benthic marine invertebrates have shown great specificity of requirements in setting their limits of distribution at the time of selection of a habitat. The tubeworm Galeolaria caespitosa shows great variation in recruitment from place to place on rocky intertidal seashores and is a gregarious animal with larvae showing directed responses to conspecific adults on the substratum. I hypothesised that, if variation in recruitment of G. caespitosa were independent of conditions on the substratum, the magnitude of recruitment in patches of the same shape but different sizes cleared within continuous mats of conspecific adults would be directly related to the area available for colonisation in the patch. Alternatively, if variation in recruitment were due to the response of larvae to conspecific adults on the substratum, the magnitude of recruitment would be a function of the perimeter of the patch, which, given patches of the same shape, is a measure of the influence of conspecific adults in that patch. To distinguish between these alternatives, small (area = 25 cm²; perimeter = 20 cm) and large (area = 225 cm²; perimeter = 60 cm) square patches were cleared within continuous mats of conspecific adults at four sites and recruitment of G. caespitosa was monitored over two seasons of recruitment. The density of recruits per unit area was, on average, almost three times greater in small than in large patches, indicating that recruitment of G. caespitosa is not directly related to the area of the patch. In contrast, the density of recruits per unit perimeter was not significantly different between small and large patches, indicating that recruitment of G. caespitosa is related to the proximity of conspecific adults in the patch. Therefore, at a given site, the perimeter of patches within mats of G. caespitosa is a better predictor of the relative magnitude of recruitment among patches than that provided by their areas. These results are contrary to many models of invertebrate recruitment that assume close linkage between available space on the substratum and settlement. Moreover, they highlight the importance of behavioural responses of animals at the time of selection of habitat in accounting for variation in recruitment. For populations of organisms that display gregarious behaviour at settlement, or conspecific attraction, this direct relationship between the perimeter of patches and recruitment could be used as a tool in restoring populations to disturbed habitats. The added benefit of such facilitative interactions in restoring populations is that they provide increasing returns to the population for a given supply of potential colonists to a habitat. Received: 1 November 1996 / Accepted: 20 January 1997     

Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

The effect of light environment, leaf area, and stored carbohydrates on inflorescence production by a rain forest understory palm

Variation in flowering by long-lived plants may be correlated with current resource availability. If, however, there are trade-offs between current and future reproduction, or between reproduction and storage or growth, then understanding variation requires a whole-plant, longer-term perspective. Inflorescence production by Calyptrogyne ghiesbreghtiana Linden ex. H. Wendl., an understory palm, was studied over 3 years. Annual inflorescence production varied greatly and was correlated with variation in plant size and light environment. There was no trade-off between past inflorescence production and the frequency of future inflorescence production. On the contrary, individuals that produced more inflorescences than predicted from their size and light environment tended to continue to do so in subsequent years also. I manipulated the resource environment of a subset of plants by removal of leaves and/or reproductive spikes. Leaf removal suppressed inflorescence production for the following 2 years, but spike removal had no effect. One year after leaf removal stored reserves were, on average, back to pre-treatment levels. There was, however, a negative effect of recent inflorescence production on storage. Plants with higher levels of storage had higher inflorescence production in the next 75 days. In C. ghiesbreghtiana the resource cost of reproduction is apparent in short-term variation in stored reserves. In contrast, annual inflorescence production does not follow a trade-off pattern between successive years, but consistently reflects both plant size and the light environment. Received: 20 October 1996 / Accepted: 25 January 1997     

The birch tube-maker Acrobasis betulella in a fragmented habitat: the importance of patch isolation and edges

We examined the response of the birch tube-maker Acrobasis betulella Hulst (Lepidoptera: Pyralidae) to habitat patch isolation and edges. Density of A.␣betulella larva was higher on distant islands than on islands close to the shore. Following experimental removal of all larvae from the islands, adults were able to recolonize even the most distant islands, and larval density was again positively correlated with the degree of isolation. Larval density was not correlated with island size or the amount of birch present on the islands. Larvae on more distant islands did not have lower mortality than those in less isolated sites. Larvae were found more often on edges than in the interior of birch stands and developed faster on edges. This positive edge effect, coupled with the ability of the adults to disperse to even our most distant islands, suggests that A. betulella would be favored in an environment fragmented at the same scale as our island system. Received: 15 April 1996 / Accepted: 30 September 1996     

The effect of exercise and restraint on pectoral muscle metabolism in pigeons

The effect of various activity regimes on metabolism of pigeon pectoralis was examined by measurement of blood lactate following exercise, total lactate dehydrogenase activity of pectoral muscle, and proportions of specific isoenzymes of pectoral muscle lactate dehydrogenase. Sprint-trained birds had the highest pectoral muscle lactate dehydrogenase activity (1409 IU · g⁻¹ wet tissue), while endurance-trained birds had the highest peak lactate levels (287 mg · dl⁻¹, extrapolated from decay curves) and fastest half-time of the lactate response (4.8 min) following exercise, but the lowest lactate dehydrogenase activity (115 IU · g⁻¹ wet tissue). Immobilization of one wing for 3 weeks following endurance training produced a marked increase in lactate dehydrogenase activity of the immobilized muscle, compared to that in the contralateral pectoralis and endurance-trained muscle. Aerobic forms of the lactate dehydrogenase enzyme (that favor conversion of lactate to pyruvate) predominated in pectoral muscle of endurance-trained birds, while cage-confined birds exhibited primarily the anaerobic isoenzymes. These results demonstrate that conversion of pectoral muscle lactate dehydrogenase isoenzymes, total lactate dehydrogenase activity, and half-time of lactate response after exercise is dependent on activity regime in pigeons. In this respect, pigeon pectoral muscle responds to training and disuse in a manner similar to that of mammalian skeletal muscle. Accepted: 10 September 1996     

Temperature regulation and metabolism of an Australian bat, Chalinolobus gouldii (Chiroptera: Vespertilionidae) when euthermic and torpid

The thermal and metabolic physiology of Chalinolobus gouldii, an Australian vespertilionid bat, was studied in the laboratory using flow-through respirometry. Chalinolobus gouldii exhibits a clear pattern of euthermic thermoregulation, typical of endotherms with respect to body temperature and rate of oxygen consumption. The basal metabolic rate of euthermic Chalinolobus gouldii is approximately 86% of that predicted for a 17.5-g mammal and falls into the range of mass-specific basal metabolic rates ascribed to vespertilionid bats. However, like most vespertilionid bats, Chalinolobus gouldii displays extreme thermolability. It is able to enter into torpor and spontaneously arouse at ambient temperatures as low as 5 °C. Torpid bats thermoconform at moderate ambient temperature, with body temperature ≈ ambient temperature, and have a low rate of oxygen consumption determined primarily by Q ₁₀ effects. At low ambient temperature (< 10 °C), torpid C. gouldii begin to regulate their body temperature by increased metabolic heat production; they tend to maintain a higher body temperature at low ambient temperature than do many northern hemisphere hibernating bats. Use of torpor leads to significant energy savings. The evaporative water loss of euthermic bats is relatively high, which seems unusual for a bat whose range includes extremely arid areas of Australia, and is reduced during torpor. The thermal conductance of euthermic C. gouldii is less than that predicted for a mammal of its size. The thermal conductance is considerably lower for torpid bats at intermediate body temperature and ambient temperature, but increases to euthermic values for torpid bats when thermoregulating at low ambient temperature. Accepted: 22 August 1996     

The effect of gender on left ventricular function immediately after the wingate test

The effect of gender on left ventricular systolic function and exercise haemodynamics in healthy young subjects was studied during 30-s all-out sudden strenuous dynamic exercise. A group of 22 men [19.3 (SD 1) years] 20 women [19.1 (SD 1) years] volunteered to participate in this study. Two-dimensional direct M-mode and Doppler echocardiograph studies were performed with the subject in the sitting position. The Doppler examination of flow was located with continuous-wave, interrogating ascending aorta measurements. The subjects completed the study without showing any electrocardiograph abnormalities. An interaction effect with stroke volume (P < 0.05) was characterized by a decrease in the men and an increase of stroke volume in the women. Cardiac output rose significantly (P < 0.05) up to 14.5 (SD 6) l · min⁻¹) for the men and 12.1 (SD 4) l · min⁻¹ for the women compared to the rest values [5.8 (SD 0.4) and 4.7 (SD 0.5) l · min⁻¹, respectively]. Flow velocity integral and acceleration time differed significantly between the two groups at rest (P < 0.05). During exercise these differences showed an interaction effect (P < 0.05). These results would indicate that normal men and women respond to sudden strenuous exercise by reducing their left ventricular systolic function, with a significantly greater decrease in women (P < 0.05). The gender differences in the haemodynamic responses during the present study, may, as suggested by others, be attributable to differences in energy metabolism. In addition, changes in Doppler parameters of aortic flow, haemodynamics and blood pressure responses during sudden strenuous exercise differed markedly from those seen before with endurance exercise. Accepted: 8 January 1997     

Responses to object approach by a wide field visual neurone, the LGMD2 of the locust: Characterization and image cues

The LGMD2 belongs to a group of giant movement-detecting neurones which have fan-shaped arbors in the lobula of the locust optic lobe and respond to movements of objects. One of these neurones, the LGMD1, has been shown to respond directionally to movements of objects in depth, generating vigorous, maintained spike discharges during object approach. Here we compare the responses of the LGMD2 neurone with those of the LGMD1 to simulated movements of objects in depth and examine different image cues which could allow the LGMD2 to distinguish approaching from receding objects. In the absence of stimulation, the LGMD2 has a resting discharge of 10–40 spikes s⁻¹ compared with <1 spike s⁻¹ for the LGMD1. The most powerful excitatory stimulus for the LGMD2 is a dark object approaching the eye. Responses to approaching objects are suppressed by wide field movements of the background. Unlike the LGMD1, the LGMD2 is not excited by the approach of light objects; it specifically responds to movement of edges in the light to dark direction. Both neurones rely on the same monocular image cues to distinguish approaching from receding objects: an increase in the velocity with which edges of images travel over the eye; and an increase in the extent of edges in the image during approach. Accepted: 23 October 1996     

Cold hardiness in Entomobrya nivalis (Collembola, Entomobryidae): annual cycle of polyols and antifreeze proteins, and antifreeze triggering by temperature and photoperiod

In the Swiss Prealps Entomobrya nivalis hibernates in an inactive state, hidden under bark flakes on spruce. For freeze avoidance it relies on thermal hysteresis proteins (THPs) and polyols (mainly ribitol, with small amounts arabitol and threitol). Polyols are present only during the inactive state, THPs additionally protect during the transition phase in spring and autumn, when animals are still active but frosts may occur. Peak values were recorded in February/March for THPs (3.5 °C hysteresis between melting and freezing point) and for polyols (26 μg mg⁻¹ FW; hemolymph osmolality 680 mosmol l⁻¹). E. nivalis is able to control its hemolymph osmolality independently of body water content. Mean osmolality in summer was 350– 440 mosmol l⁻¹, in winter it was elevated to 650 mosmol l⁻¹, due to a synthesis mainly of ribitol. Body water content varied between 1.8 and 3.3 mg H₂O mg⁻¹ DW, depending on humidity conditions. Experiments on triggering of antifreeze synthesis showed the action of temperature and photoperiod as cues, but there was also evidence for an endogenous rhythm. No clear correlation between antifreeze concentration and supercooling ability could be established, suggesting that gut content or other parameters also play an inportant role. Accepted: 18 November 1995     

Effects of ultraviolet radiation on early larval stages of the Alpine newt, Triturus alpestris, under natural and laboratory conditions

Although Alpine newts (Triturus alpestris) are found at altitudes up to 2500 m, their larvae proved to be extremely sensitive to UV radiation when exposed in clear tapwater to natural sunlight or to comparable artificial UV-B radiation in the laboratory. The experiments revealed severe skin damages (lysis of epithelial cells) and mortality after a few days of exposure. In their natural habitats above the timberline, however, the larvae are protected by the high concentration of dissolved organic carbon in the water leading to an almost complete absorption of UV radiation within the first few centimeters of the water layer. Furthermore, when exposed to UV radiation larvae show erratic swimming activities that may carry them into protected areas. Only in very shallow natural habitats did we detect sublethal UV-like histological effects. Shallow high mountain ponds with clear water normally lack newt populations, probably as a consequence of both low temperature and the effects of UV radiation. Received: 4 October 1996 / Accepted: 7 February 1997     

The heat increment of feeding in house wren chicks: magnitude, duration, and substitution for thermostatic costs

The heat increment of feeding (HIF), a transient postprandial increase in metabolic rate, is the energy cost of processing a meal. We measured HIF in house wren chicks (Troglodytes aedon) ranging in mass from 1.6 to 10.3 g. This mass range (age 2–10 days) spanned a transition from blind, naked, ectothermic chicks through alert, endothermic birds with nearly complete feathering. We fed chicks crickets (2.7–10% of chick body mass) and determined HIF from continuous measurements of oxygen consumption rate (O₂) before and after meals. At warm ambient temperatures (T _a) of 33–36 °C, the magnitude of HIF (in ml O₂ or joules) was linearly related to meal mass and was not affected by chick mass. HIF accounted for 6.3% of ingested energy, which is within the range of results for other carnivorous vertebrates. The duration of HIF was inversely related to chick mass; 10-g chicks processed a standard meal approximately twice as fast as 2-g chicks. HIF duration increased with increasing meal mass. The peak O₂ during HIF, expressed as the factorial increase above resting metabolism, was independent of body mass and meal mass. In large, endothermic chicks ( > 8 g), HIF substituted for thermoregulatory heat production at low T _a. Accepted: 11 December 1996     

Axillary bud banks of two semiarid perennial grasses: occurrence, longevity, and contribution to population persistence

The occurrence, longevity, and contribution of axillary bud banks to population maintenance were investigated in a late-seral perennial grass, Bouteloua curtipendula, and a mid-seral perennial grass, Hilaria belangeri, in a semiarid oak-juniper savanna. Axillary buds of both species were evaluated over a 2-year period in communities with contrasting histories of grazing by domestic herbivores. A double staining procedure utilizing triphenyl tetrazolium chloride and Evan's blue indicated that both viable and dormant axillary buds remained attached to the base of reproductive parental tillers for 18–24 months which exceeded parental tiller longevity by approximately 12 months. Bud longevity of the late-seral species, B. curtipendula, exceeded bud longevity of the mid-seral species, H. belangeri, by approximately 6 months. Younger buds located on the distal portion of the tiller base were 3.2 and 1.4 times more likely to grow out than older proximal buds of B. curtipendula and H. belangeri, respectively. The percentage of older proximal buds, which included comparable portions of viable and dormant buds, that grew out to produce tillers following mortality of parental tillers was 6.0% for B. curtipendula and 8.4% for H. belangeri. In spite of the occurrence of relative large axillary bud banks for both species, the magnitude of proximal bud growth did not appear sufficient to maintain viable tiller populations. We found no evidence to support the hypothesis of compensatory bud growth on an individual tiller basis for either species. Grazing history of the communities from which the buds were collected did not substantially affect the number, status, longevity, or outgrowth of axillary buds on an individual tiller basis for either species. However, long-term grazing by domestic herbivores influenced axillary bud availability by modifying population structure of these two species. Bud number per square meter for B. curtipendula was 25% lower in the long-term grazed compared to the long-term ungrazed community based on a reduction in both tiller number per plant and plant number per square meter. In contrast, bud number per square meter for H. belangeri was 190% greater in the long-term grazed than in the long-term ungrazed community based on a large increase in plant density per square meter. Minimal contributions of axillary bud banks to annual maintenance of tiller populations in this mid- and late-seral species underscores the ecological importance of consistent tiller recruitment from recently developed axillary buds. Consistent tiller recruitment in grasslands and savannas characterized by intensive grazing and periodic drought implies that (1) bud differentiation and maturation must be remarkably tolerant of adverse environmental conditions and/or (2) tiller recruitment may resume from buds that mature following the cessation of severe drought and/or grazing, rather than from mature buds that survive these disturbances. These scenarios warrant additional research emphasis given the critical importance of this demographic process to tiller replacement in species populations and the maintenance of relative species abundance in grasslands and savannas. Received: 12 August 1996 / Accepted: 30 December 1996     

An Aspergillus nidulans nuclear protein, AnCP, involved in enhancement of Taka-amylase A gene expression, binds to the CCAAT-containing taaG2 , amdS , and gatA promoters

Using AnCP (Aspergillus nidulans CCAAT-binding protein) as a CCAAT-specific binding factor model, the possibility that one factor is able to recognize CCAAT sequences in several different genes in A.␣nidulans was examined. DNase I protection analysis showed that AnCP specifically bound to CCAAT sequence-containing regions comprising 21 to 36 bp of the taa, amdS and gatA genes. Furthermore, replacement of the CCAAT sequence with CGTAA was found to abolish the binding of AnCP and to have an inhibitory effect on taa promoter activity. This clearly demonstrates a positive function of the CCAAT element. However, amylase was induced by starch and repressed by glucose in a CCAAT-box disruptant, as in wild-type cells. Received: 28 June 1996 / Accepted: 7 October 1996