Desiccation as a factor in the intertidal zonation of barnacles

Four species of balanomorph barnacles, Balanus crenatus Brugière, B. balanoides (L.), Elminius modestus Darwin and Chthamalus stellatus (Poli), were studied to assess the susceptibility of intertidal barnacle species to desiccation. Known sized samples of barnacles were exposed to controlled desiccating conditions and subsequent survival and water loss were determined. It is clear that the ability to live high on the shore is dependent on a reduction of the overall permeability to water loss. Because of greater surface area to volume ratios, small stages are particularly prone to desiccation. In normal intertidal emersion periods, small stages of B. crenatus particularly, and also of B. balanoides and E. modestus which are similar in their desiccation resistance, would be susceptible to desiccation at normal temperatures and low humidities. Large barnacles would be more prone to death from high temperatures when the tide is out. The spat of C. stellatus, although surviving much longer than spat of larger dimensions of the other species, must also be prone to prolonged emersion conditions at high shore levels.     

The ecophysiological complex of Bathyporeia pilosa and B. pelagica (Crustacea: Amphipoda). I. Respiration rates

The metabolic rates of high and low shore estuarine populations of Bathyporeia pilosa Lindström and an open coast population of B. pelagica (Bate) have been determined over a range of temperatures during January and February, and June and July, 1968. Changes in oxygen uptake have also been measured monthly at 15°C. During the winter, oxygen uptake was in the order: high shore B. pilosa>low shore B. pilosa>B. pelagica. During the summer, high shore B. pilosa and B. pelagica had similar metabolic rates, but both were significantly higher than low shore B. pilosa. Both populations of B. pilosa had lower metabolic rates in summer than in winter, whereas the metabolic rate of B. pelagica remained much the same. Seasonal changes in metabolic rate are closely correlated with reproductive cycles. The possible influences of environmental parameters are discussed.     

Heat tolerance,growth and regeneration in three North Sea bryozoans exposed to different constant temperatures

Three species of bryozoans—Membranipora membranacea (L.), Electra pilosa (L.) and Conopeum reticulum (L.) — are capable of acclimating to elevated temperatures, above the normal range experienced in nature, when exposed to a gradual increase in ambient temperature. Conspicuous differences in LD 50 values, as a consequence of acclimation, occur between representatives of the same species acclimated and grown at constant temperatures in the laboratory. The tolerance range of these species is influenced by their thermal history in the laboratory. While increased ambient temperatures accelerate growth rate, final colony size attained after prolonged exposure declines at higher temperatures. The size of zooecia attained is inversely proportional to the test temperature. Colonies of E. pilosa maintained at 22°C develop erect branches. Hence, it is probable that E. pilosa forma erecta is only a growth form of normally encrusting colonies of E. pilosa. Temperature affects rate of regeneration.     

The life cycle ofClytia attenuata (calyptoblastea: Campanulariidae)

The life cycle of the hydroidClytia attenuata (Calkins) (Calyptoblastea: Campanulariidae) has been completed in the laboratory including development of the medusa, previously described asPhialidium lomae Torrey (Leptomedusae: Campanulariidae). Under laboratory conditions, the hydroid exhibits some morphological variation. Characteristic branching of the hydroid occurs at temperatures between 17° to 19°C. At 13° to 15°C the colonies are unbranched and cannot be distinguished fromClytia cylindrica L. Agassiz. Young medusae are similar to other young species ofPhialidium. Development to the adult form requires 25 to 30 days at 17° to 24°C. The adult medusae are 6 to 10 mm in diameter, watch-glass shaped, and have 20 to 28 tentacles. Based on the adult medusa,Clytia attenuata is maintained as a valid species.     

Temperature tolerance of the polychaete worms Diopatra cuprea and Clymenella torquata

The resistance to high temperatures of two species of intertidal tube-dwelling polychaete worms has been tested to show seasonal and geographic variations. The summer 50% survival temperature at Beaufort, North Carolina, was 42.5 °C for Diopatra cuprea (Bosc) and 40.5 °C for Clymenella torquata (Leidy). Winter 50% survival temperatures for both species at Beaufort were approximately 4.0 C° lower. Both species showed a geographic difference in 50% survival temperature of more than 4.0 C° between North Carolina and Massachusetts in summer. D. cuprea from the Mississippi coast showed a lower survival temperature, probably due to combined temperature and salinity effects. Laboratory acclimation of C. torquata from Beaufort at low temperatures, during summer months, produced experimental results similar to those from winter animals. The seasonal differences in temperature tolerance are phenotypic expressions of a physiological response which can be related to environmental temperature patterns.     

Variability in adult group compositions of a prosimian primate

Many species fall into specific mating-system categories, and that category is usually associated with a suite of behavioral and morphological characteristics. Several lemur species, including Propithecus diadema edwardsi, have been labeled "idiosyncratic" because variation in socionomic sex ratios among groups is consistent with wide variation in social structure. We used several hypotheses founded in behavioral ecology to assess variability in P. d. edwardsi. First we examined 46 group-mating seasons to quantify variability. We then tested predictions that the number of males per group would increase as the number of adult females increased, and the number of males would increase as female mating synchrony increased. Examining variation in offspring survival relative to the number of adult males in a group may tell us which composition is likely to persist into the future, so we also hypothesized that as the number of males in a group increased, fertility and offspring survival would increase. We found an equal distribution of polygynous, polygynandrous, pairs, and polyandrous groups. Furthermore, female distribution and mating synchrony did not predict the number of males, and offspring survival was not correlated with the number of males. Since infants survived equally well in groups of all compositions, sifakas experienced no pressure to maintain a particular number of adult males per adult female. The small number of adults per group (mean=3.2) may result from balancing feeding competition against predator detection. Augmenting the mate pool available from the group with mates from neighboring groups may promote the notable variability seen in the adult group compositions of sifakas.     

Environmental control of the breeding of three boreo-arctic cirripedes

Three species of barnacles with boreo-aretic distribution were shown to require maintenance for several weeks below a critical temperature before the breeding condition could be attained. The temperatures critical for Balanus balanoides (L.) Balanus balanus (L.) and Balanus crenatus (Bruguière) were found to be between 10° and 12°C, 10° and 14°C, and at about 17°C, respectively. Although the strong influence of continuous light and the weak influence of continued feeding in delaying the onset of breeding in B. balanoides were confirmed, there remained some outstanding anomalies between the breeding behaviour of this species under laboratory conditions and between the tide marks. It was found impossible to initiate breeding by the application of conditioning procedures significantly in advance of the time of the normal Autumn breeding season in B. balanoides. Breeding appears to be inhibited, independently of external conditions, for a set period after the preceding brood cycle. Evidence points to a similar, largely endogenous, control of breeding in B. balanus which also breeds once annually, but not in B. crenatus which breeds continuously so long as food and temperature levels permit.     

Symbiosis in sacoglossan opisthobranchs: functional capacity of symbiotic chloroplasts

It has been previously reported that many species of the order Sacoglossa (Mollusca: Opisthobranchia) contain algal chloroplasts within the cells of their digestive gland and maintain them in a symbiotic condition. In the present study, two species, Elysia hedgpethi Marcus and Placobranchus ianthobapsus Gould, were compared as to their abilities to retain functional chloroplasts in their tissues. Animals were starved for varying lengths of time, and the functional capacity of the plastids was ascertained at intervals. The chlorophyll content of whole animals, and the ability to incorporate ¹⁴CO₂ were used as the assay for functional capacity. E. hedgpethi decreased in chlorophyll content during starvation until the tenth day, when no chlorophyll was detectable spectrophoto-metrically. Incorporation of ¹⁴CO₂ paralleled the decline in chlorophyll, and was at control levels by the tenth day. P. ianthobapsus showed no decline in chlorophyll content over 27 days starvation, although the ability to incorporate ¹⁴CO₂ showed a decrease.     

Respiration and nitrogen excretion of zooplankton. II. Studies of the metabolic characteristics of starved animals

Respiration and nitrogen-excretion studies were carried out on several species of zooplankton (Meganyctiphanes norvegica, Phronima sedentaria, Acartia clausi and Sagitta setosa) under starvation. Although all the species were mainly ammonotelic, apparently a significant amount of organic nitrogen was excreted; the validity of the measurements and their significance are discussed. The effect of duration of starvation showed for M. norvegica and A. clausi two different patterns of behaviour, which were chiefly a function of the rate of biomass turnover of the species studied. The rates of metabolism, chemical composition, and reaction to starvation varied with season in M. norvegica. The physiological balance of the experimental animals was examined by calculating the protein carbon equivalent to respiratory and excretory catabolism, and by use of atomic O:N ratios. Starved individuals catabolized more protein carbon than can be accounted for by the amount of respiratory oxygen utilized. A hypothesis, which suggests that there are three levels of resistance to starvation, is proposed to explain this paradox, and its metabolic basis is discussed.     

Desiccation,higher temperatures and upper intertidal limits of three species of sea mussels (Mollusca: Bivalvia) in New Zealand

Distribution, abundance, and resistance adaptations to higher temperature and desiccation of three species of intertidal mussels (Mytilus edulis aoteanus, Perna canaliculus and Aulacomya maoriana) were studied in New Zealand. M. edulis aoteanus generally was more abundant upshore, with P. canaliculus dominating downshore. M. edulis aoteanus was more common than P. canaliculus on the outside of mixed-species clumps. Abundance of A. maoriana was variable, with individuals favouring damp habitats such as inside mussel clumps. In moving air at 75% relative humidity and at 20°C or 30°C, median lethal levels of water loss were similar for P. canaliculus and M. edulis aoteanus but lower for A. maoriana. Rates of desiccation varied inversely with size and were higher for P. canaliculus, due mainly to valve gaping with resultant loss of water from the mantle cavity. M. edulis aoteanus was more tolerant of higher water temperatures than were the other species. Success in colonizing upshore or more aerially exposed habitats seems to be related to ability of small mussels to tolerate desiccation, especially during hot, windy weather.     

Goniadella gracilis,a polychaete new to British seas

Polychaetes of the genus Goniadella Hartman were found in Liverpool Bay, England, in sand and sandy gravel below depths of 15 m. The worms had most of the characteristics of G. gracilis (Verrill), a species previously recorded only from eastern North America and South Africa.     

Interactions between an invasive and a native bryozoan (<Emphasis Type="Italic">Membranipora membranacea</Emphasis> and <Emphasis Type="Italic">Electra pilosa</Emphasis>) species on kelp and <Emphasis Type="Italic">Fucus</Emphasis> substrates in Nova Scotia,Canada

Most research on biological invasions to date has focused on the population dynamics of very successful and disruptive introduced species; however, additional knowledge of the biology of the native species is essential for understanding interactions between the two and may reveal factors that limit invasion success. The invasive bryozoan Membranipora membranacea interacts with native Electra pilosa on two substrates in northwest Atlantic subtidal habitats: highly dynamic and fast-growing kelps; and smaller, more stable, and slow-growing fucoid algae. We quantified the relative abundance and evaluated encounter outcomes in different seasons of these two bryozoans on kelp and Fucus at four sites in Nova Scotia. We also examined the effects of substrate (kelp, Fucus), temperature (7, 10, 13°C), and food (limited, unlimited) on growth rates of E. pilosa in laboratory experiments and using field manipulations. We compared our findings on factors affecting the growth of E. pilosa to those on M. membranacea obtained in similar and thus directly comparable experiments from a previous study. The proportional abundance of M. membranacea was greater than that of E. pilosa on kelps, but the opposite was observed on Fucus. Competitive standoffs between the two bryozoans were more frequent than expected, with no differences recorded between substrates; most encounters were won by M. membranacea. Growth of E. pilosa was faster on Fucus than kelp, decreased with increasing temperature only on Fucus, and was not affected by food. Growth rate of E. pilosa in all treatments was slower than that previously measured for M. membranacea. Faster growth and strong overgrowth abilities likely interact on kelps to ensure success of the invasive bryozoan. Success can be limited by low space availability, which in turn restricts growth rate, and consequently, colony size, such as on fucoid substrates. The incorporation of alternative contexts into invasion research can reveal factors involved in the resilience of native communities.     

A comparison of traditional,cluster and Zürich-Montpellier analyses of infaunal pelecypod associations from two adjacent sediment beds

A Zürich-Montpellier analysis provided more detailed information about pelecypod associations in two sediment beds than either a traditional subjective approach or a cluster analysis based on Jaccard's coefficient. The Zürich-Montpellier test separated till and sand pelecypod associations, and indicated distinguishing species and atypical samples. Species distinguishing the till association were Cardita ventricosa, Yoldia myalis, Semele rubropicta, Venerupis kennerlyi, Clinocardium nuttallii, Nuculana minuta and Macoma incongrua. Species distinguishing the sand were Macoma elimata, Compsomyax subdiaphana, Yoldia ensifera, Macoma calcarea, M. lipara, M. brota, Yoldia limatula and Macoma alaskana. Two samples contained species from both associations plus some species rare elsewhere. Combined R-and Q-type grouping procedures, of which the Zürich-Montpellier approach is an example, are useful in evaluating benthic faunal similarities, but the suitability of various forms of the procedure needs further study.Supported by the Defense Research Board of Canada, Grant No. 9520-23. Based on a thesis submitted by J. D. Popham to the University of Victoria in partial fulfillment of the requirements for the degree of B.Sc. (Honours).     

Respiration and nitrogen excretion of zooplankton. IV. The influence of starvation on the metabolism and the biochemical composition of some species

Changes in the respiration, ammonia excretion and biochemical composition were studied for three species of starving zooplankton (Calanus finmarchicus, Sagitta elegans, and Acartia clausi). Over the period of starvation, the respiration rate of all three species followed the same pattern of an initial decrease followed by a more or less constant level. A similar pattern was observed for the ammonia excretion rate of S. elegans and A. clausi, whereas C. finmarchicus excretion appeared to oscillate between high and low levels of protein catabolism. Study of the biochemical changes showed that C. finmarchicus consumed primarily lipids, and at times proteins, to meet its energy requirement whereas S. elegans and A. clausi primarily used protein. Variations in the elemental composition as well as the O:N ratio confirmed that C. finmarchicus alternated between periods of protein-dominant catabolism and lipid-dominant catabolism during starvation. No similar change in catabolism was observed in the two other species. The results are discussed in terms of physiological mechanisms of resistance to starvation and were used to calculate the energy budget of S. elegans and C. finmarchicus during the period of total starvation. The significance of such budgets is discussed and some of the sources of error examined.Bedford Institute of Oceanography Contribution.     

Demographic and temporal structure of an allele frequency cline in the mussel Mytilus edulis

Feeding rates, patterns of prey selection, and starvation tolerance were investigated for adult males and females of the cyclopoid copepod Corycaeus anglicus collected from the waters of Friday Harbor, Washington, USA. Selection by C. anglicus was determined largely by prey body-size, but was also affected by species and developmental stage. Small developmental stages of all prey species were fed upon at relatively low rates. The small calanoid species Acartia clausii was increasingly vulnerable to predation by C. anglicus as it progressed through successive developmental stages. Larger prey species, Pseudocalanus sp. and Calanus pacificus, were more vulnerable in intermediate stages, the C3 and N6 stages, respectively. Larger and smaller prey were characteristically attacked at different sites on their bodies; however, attack sites fell within a similar range of body widths, 130 to 170 m. Males of Corycaeus anglicus killed a maximum of 1.4 prey d^-1 when feeding on the optimally-sized adult females of Acartia clausii, which are approximately equivalent to its own body length. Males fed at approximately double the rates of females. Despite its small size and apparent lack of metabolic stores, this cyclopoid is highly tolerant of starvation conditions. Median survival time without food is at least 2 wk for both males and females. In its predatory behavior, C. anglicus employs an ambush-type strategy and seems to be adapted for infrequent encounters with relatively large prey.Contribution No. 1412 from the School of Oceanography, University of Washington, Seattle     

The chromosomes of Trypetesa lampas (Cirripedia,Acrothoracica)

Observations were made of the chromosomes of the burrowing barnacle Trypetesa lampas (Hancock). A method of squash preparation was used, which incorporated staining the material in a solution of 2% orcein in 45% acetic acid. The diploid number of chromosomes of T. lampas was found to be 12 and the haploid number 6, in both males and females. No obvious chromosomal mechanism of sex determination was found. There was, therefore, no cytogenetic confirmation of Kühnert's view (1934) that the sex of the larva is predetermined genetically. During mitosis in the females and embryos of Trypetesa lateralis Tomlinson and Kochlorine floridana Wells and Tomlinson, the diploid number of chromosomes was found to be about 14. The numbers of chromosomes in the 3 species of Acrothoracica studied were approximately half those observed in all 12 species of Thoracica and all 5 species of Rhizocephala previously investigated.     

Metabolic characteristics of midwater zooplankton: Ammonia excretion,O:N rations,and the effect of starvation

The nature of protein catabolism in a wide range of species of midwater zooplankton was investigated. The weight-specific ammonia excretion rates (g NH₃–N g^–1 dry wt h^–1, y) decline exponentially with minimum depth of occurreece (MDO, x), y=163.4 x^{–0.479±0.212} (95%ci) (CI=confidence interval), when temperature is held constant. The change in ammonia excretion can be partially explained by the decrease in percent protein (%P) with MDO, %P=80.17 MDO^{–0.148±0.122 (95%ci)} The atomic O:N ratio of freshly caught zooplankters ranged from 9.1 to 91, with most measurements between 9 and 25. Detailed studies were carried out on the response of one of the species studied (Gnathophausia ingens) to starvation (28 d). After 14 d of starvation the average ammonia excretion rate declined by more than 75% to less than 1 g NH₃–N g^–1 wet wt h^–1, although the average oxygen consumption declined by only 13% within the first 7 d of starvation and then remained stable. This differential response of oxygen consumption and ammonia excretion to starvation resulted in an increase in the average O:N ratio of starved animals from an initial 33 to 165 after 21 d. The average O:N ratios of fed mysids remained below 38 during the experiment. G. ingens maintains a relatively uniform metabolic rate during starvation by relying more heavily on its large lipid stores than when being fed.     

Activity budgets in two species of bee flies (Lordotus: Bombyliidae,Diptera): a comparison of species and sexes

Summary This study compares time budgets of males and females of two sympatric species of bee flies, Lordotus pulchrissimus and L. miscellus. These species occur synchronously in sand dune habitat on the north shore of Mono Lake, Mono Co., California, and feed almost exclusively on the flowers of a composite shrub, Chrysothamnus nauseosus. The two closely related species, with similar ecological requirements and in an identical environment, allocate time and energy in very different ways, even when accomplishing the same ecological goals. L. pulchrissimus males and females engage respectively in aggressive interactions and feeding primarily in the morning. In contrast, L. miscellus males and females intersperse brief periods of resting with flying when defending territories and feeding, and they keep up these activities until later in the day. Similarly within species, males are involved in aggressive interactions for a shorter period each day, and females feed over a longer period. Flies in all sex-species classes but male pulchrissimus allocate their time energetically in similar ways. Male pulchrissimus spend more time each day in energetically costly activities; they engage in continuous hovering flight and intense interactions in aerial aggregations. While male miscellus feed little, male pulchrissimus spend a large portion of their time feeding, approximately as much as females, contrary to the expectation that males should be foraging-time minimizers. This study corroborates the conclusions of previous studies on bee-fly communities by showing that nectar and pollen are important resources for adult bee flies, at least for some species: Individuals of these two species spend a large proportion of their adult lifetimes feeding.     

Distribution of Trypetesa lampas (Cirripedia,Acrothoracica) in various gastropod shells

The distribution of Trypetesa lampas (Hanoock) and its rates of infection of different species of gastropod shells in the waters of Anglesey were studied. T. lampas was found in depths from 3 to at least 25 fathoms (5 to 46 m). It infected dead shells, mainly of Buccinum undatum L., Colus gracilis (da Costa), Neptunea antiqua (L.) and Natica alderi (Forbes); it was absent from shells occupied by hermit crabs in the intertidal zone. Significantly more B. undatum shells were infected than shells of N. antiqua and Natica alderi, and more of both C. gracilis and N. antiqua than of Natica alderi. Larger shells of B. undatum were more likely to be infected than were the smaller ones. The number of females per shell varied with the shell length raised to the power 2.39 instead of as expected with the square of the length.     

Upper temperature tolerances of some European molluscs II. Donax vittatus,D. semistriatus and D. trunculus

The upper temperature tolerances of 3 European species of Donax [D. vittatus (da Costa), D. semistriatus Poli, D. trunculus L.] are compared by means of median lethal temperatures (LT₅₀) and median burial temperature (BT₅₀) determinations for periods of exposure up to 96 h. The influence of previous acclimation temperatures on these determinations is also considered. D. trunculus shows the greatest thermal tolerance, reflecting its distribution in shallower water; D. vittatus and D. semistriatus display much closer thermal tolerances, although D. semistriatus which has a more southerly geographical distribution has a slightly higher thermal tolerance. All 3 species exhibit a significant effect of previous acclimation temperature, the effect on the burrowing responses being greater than that on LT₅₀.