Significance of burrow-opening diameter as a flood-prevention mechanism for air-filled burrows of small intertidal arthropods

Small air-breathing terrestrial arthropods (e.g. beetles, ants, crabs) living within the intertidal zone of mud and sand flats construct burrows with a narrow opening to the substrate surface. Despite frequently remaining open, these burrows are not flooded by the incoming tide. We examine, from a theoretical standpoint, the rôle of burrow-opening diameter in preventing burrow flooding. We determine a critical burrow diameter of 3 mm above which burrows are likely to flood unless the entrance is plugged by the burrow's inhabitant(s) prior to tidal inundation. We examine water penetration via the burrow entrance and seepage through the burrow walls. A practical mathematical solution is provided which can be used to examine the extent to which water may penetrate a given animal burrow in the field.     

Individual variation and prior experience affect the discrimination of a brood-parasite by its subsocial beetle host

In birds, host experience can modify response to parasites but nothing similar is known for insects. We studied two desert tenebrionid beetles, the subsocial host Parastizopus armaticeps and its obligate and ubiquitous brood-(clepto)parasite Eremostibes opacus, which mimics the hosts odour spectrum. Nearly 10% of host burrows in the field (n=214) remain unparasitised even after introducing E. opacus experimentally (n=22). In discrimination tests, 7% of naive host beetles eject the parasite, showing individual variation in olfactory discrimination ability. To test for effects of prior breeding experience, naive (first time) or experienced (second time) breeders with or without prior parasite contact were confined with E. opacus at breeding onset, their behaviour recorded and parasite number counted on breeding completion. More first- than second-time breeding pairs completely excluded the parasite and second-time breeders with no prior contact with E. opacus had fewer parasites than any others. These pairs increased burrow-guarding dramatically, mostly during early reproduction, this being the best predictor of parasite number. Prolonged guarding, however, must be traded-off against other brood-care behaviours. Hence, it might pay breeding pairs to take the risk of accepting some parasites breeding within their burrow, and we expect this effect will be more pronounced if poor feeding and digging conditions are taken into account. This might explain why apparently 90% of the P. armaticeps breeding burrows in the field contain at least one parasitic E. opacus.Communicated by D. Gwynne     

Nestmate recognition and the ontogeny of acceptability in the ant,Leptothorax curvispinosus

Summary In social insects, there is often a brief period following eclosion when workers are highly acceptable in alien nests of their own or other species. This study tested for such an acceptance period in the facultatively polygynous ant, Leptothorax curvispinosus, and compared the duration and effectiveness of this period for conspecific and heterospecific introductions. Workers that eclosed and aged for 1–70 h or 30 days in isolation were introduced into either their parental nests (n=24), alien conspecific nests (n=265), or nests of the closely related and biologically similar species, L. longispinosus (n=341). In alien conspecific nests, acceptance was maximal for workers aged 1–12 h at introduction (67.7% not attacked, 75.8% adopted) and gradually decreased until the level of nonaggression (after 60 h) and adoption (after 36 h) were not significantly different from 30-day-old workers (5.9% not attacked, 17.6% adopted). In heterospecific nests, acceptance was maximal for workers aged 1–4 h at introduction (34.8% not attacked, 37.0% adopted) but thereafter was not significantly different from 30-day-old workers (5.6% not attacked, 8.3% adopted). In their parental nests, workers were generally accepted regardless of age (4–56 h posteclosion, 95.8% not attacked, 100% adopted); a result that is consistent with previous research on older workers (38–157 days posteclosion). This study demonstrates an acceptance period that is more effective and of longer duration within than between these species but that, under uniform laboratory conditions, is often not necessary for the integration of workers into their parental colonies. Within colonies, acceptance periods might only be important during relatively brief periods in a colony's life history when eclosing workers produce genetically based nestmate recongition cues that are not already represented in the colony and must be learned by colony members (e.g., during early colony growth or following adoption of queens), or when young workers must acquire environmentally based nestmate recognition cues to achieve and maintain acceptability.     

Learning,retention and coding of nest-associated visual cues by the Australian desert ant, <Emphasis Type="Italic">Melophorus bagoti</Emphasis>

A variety of social insects use visual cues for homing. In this study, we examine the possible factors affecting the learning and retention of nest-associated visual cues by the Australian desert ant Melophorus bagoti and the manner in which such cues are encoded by foraging ants. We placed four prominent cylindrical landmarks around a nest and trained foragers from that nest to a food source. Ants were tested with the landmark array in a distant testing field after (1) a known number of exposures to the landmarks (1, 3, 7 or 15 trials, spread over a period of 1 day, 2 days or ≥3 days) and (2) after a known period of delay (0, 24, 48, 96 or 192 h). The results show that a combination of an increase in training trials and an increase in number of training days affected the acquisition of landmark memory. Moreover, once the landmarks were learnt, they became a part of long-term memory and lasted throughout the ants’ foraging lifetime. To examine visual cue encoding behaviour, ants trained under similar conditions for 4 days were tested with (1) an identical landmark array, (2) landmarks of the same size used in training, but placed at twice the distance from each other, and (3) landmarks whose dimensions were doubled and placed at twice the distance from each other. In conditions (1) and (3), the ants searched extensively at the centre of the four landmarks, suggesting that, similar to the Saharan ant (genus Cataglyphis) and the honeybee, M. bagoti too uses a snapshot to match the view of the landmarks around the nest. But contrary to the snapshot model, in condition (2), the ants did not search extensively at the centre of the landmarks, but searched primarily 0.5 m from the landmark, the distance from each landmark to the nest during training. We discuss how various search models fare in accounting for these findings.     

Feeding,growth and bioluminescence of the heterotrophic dinoflagellate Protoperidinium huberi

Feeding, growth and bioluminescence of the thecate heterotrophic dinoflagellate Protoperidinium huberi were measured as a function of food concentration for laboratory cultures grown on the diatom Ditylum brightwellii. Ingestion of food increased with food concentration. Maximum ingestion rates were measured at food concentrations of 600 g C l^-1 and were 0.7 g C individual^-1 h^-1 (1.8 D. brightwelli cells individual^-1 h^-1). Clearance rates decreased asymptotically with increasing food concentration. Maximum clearance rates at low food concentration were ca. 23 l ind^-1 h^-1, which corresponds to a volume-specific clearance rate of 5.9x10⁵ h^-1. Cell size of P huberi was highly variable, with a mean diameter of 42 m, but no clear relationship between cell size and food concentration was evident. Specific growth rates increased with food concentration until maximum growth rates of 0.7 d^-1 were reached at a food concentration of 400 g C l^-1 (1000 cells ml^-1). Food concentrations as low as 10 g C l^-1 of D. brightwellii (25 cells ml^-1) were able to support growth of P. huberi. The bioluminescence of P. huberi varied with its nutritional condition and growth rate. Cells held without food lost their bioluminescence capacity in a matter of days. P. huberi raised at different food concentrations showed increased bioluminescence capacity, up to food concentration that supported maximum growth rates. The bioluminescence of P. huberi varied over a diel cycle, and these rhythmic changes persisted during 48 h of continuous darkness, indicating that the rhythm was under endogenous control.     

Mating system of the commensal marine isopod Jaera hopeana (Crustacea)

The commensal marine isopod Jaera hopeana Costa, 1853 was reared in the laboratory through 30 generations in the absence of its host, Sphaeroma serratum, to study its development and reproductive behaviour. It was found that adult males take virtually any opportunity to associate with a young conspecific in manca-I stage (first free-living stage) and carry it around in a characteristic position. This male-manca(I) amplexus ends during the manca's first postmarsupial molt (manca-I to manca-II) at an average age of about 9 d after hatching from the marsupium. Entry into amplexus does not alter a manca's molt timing, the duration of the amplexus thus depending exclusively on the manca's age when it joined by a male. Given a choice, males do not prefer manca-I close to molting over those just hatched. With the first postmarsupial molt, long before sexual maturity, female J. hopeana become receptive to mating. Before a female manca is released from amplexus, it is inseminated by its male partner. Sperm transferred to young immature females are stored within special sperm stores until they are needed for fertilization. The male-manca(I) amplexus seems to be a precopula, wherein males wait for their partners to molt and become sexually receptive. Nevertheless, it is inevitable that every second amplexus on average ends unsuccessfully: although manca-I have already been determined irreversibly (and probably genetically) as males and females in equal numbers, adult males are unable to predict which manca-I are potential mates and which are not. Males accept any manca-I as a precopula partner, and even when given a choice they do not prefer female over male manca-I. From the manca-II stage onward, females are continuously receptive to mating for the rest of their life, and mating can occur immediately upon contact of the partners without any significant investment in energy and time (en passant copulation).     

Effects of dusk and dawn on locomotor activity rhythms in the norway lobster Nephrops norvegicus

The endogenous locomotor activity rhythm of Nephrops norvegicus (L.) shows peaks during darkness, around 3 h after dusk. At the peaks, activity occurs mainly within the burrow, although the prawns repeatedly vacate the burrows during the same periods. The phase of the rhythm shifts with a change of light-dark regime and in response to changed timing of dusk when dawn is fixed. Activity is suppressed immediately at dawn, but recommences before complete darkness during gradually reduced light at simulated dusk. This difference in response to light at dusk and dawn indicates phase responsiveness which is characteristic of endogenous rhythms entrained by light. The light-entrained endogenous rhythm is complementary to the rhythm of emergence which, reflected in commercial catches, appears to be controlled by exogenous factors.     

ATP concentration and adenylate energy charge of tropical zooplankton from waters inside the great barrier reef

The adenylate energy charge ratio (EC) of 8 species of zooplankton ranged from 0.70 to 0.89. Eucalanus subcrassus had significantly (P<0.05) lower EC when captured during a 30-min tow than during a 3-min tow, indicating stress at capture. The concentration of ATP in the 8 species ranged from 9.1 to 36.0 g/mg protein and from 0.17 to 15.7 g/mg dry weight. Excluding a low value for the ctenophore Pleurobrachia sp., the average concentration was 8.1 g ATP/mg dry weight, which is equivalent to an approximate C/ATP ratio of 50.Contribution from the Australian Institute of Marine Science, Australia, and the Institute of Marine Biology, University of Bergen, Norway     

Effect of temperature and food on the heart beat,ventilation rate and oxygen uptake of Mytilus edulis

The heart beat, ventilation rate and oxygen uptake of Mytilus edulis L. were measured simultaneously, in response to changes in temperature and food level. There was no thermal acclimation of heart-beat frequency or amplitude to temperatures from 5° to 25°C. Oxygen consumption and ventilation rate acclimated to 10°, 15° and 20°C, but not to 25°C. Starvation reduced the rate of oxygen uptake and heart-beat frequency to a standard level and, in response to food, the ventilation rate and oxygen consumption immediately increased to an active level. Feeding was maintained after the initiation of active metabolism, and during the following 10 days the heart-beat frequency gradually increased to the level characteristic of fed individuals. There was no direct correlation between the rate of oxygen consumption and heart rate, and an apparent absence of a close nervous coupling between ventilation rate and heart rate in M. edulis.     

Digestive carbohydrases from the hepatopancreas of two Antarctic euphausiid species (Euphausia superba and E. crystallorophias)

The hepatopancreatic extracts of Euphausia superba Dana and E. crystallorophias Holt and Tattersall collected from the Antarctic. Ocean during January 1985, are most effective in hydrolysing substrates containing (13)--glucosidic linkages. Three enzymes appear to be involved in the depolymerization of (13)--D-glucans in the euphausiid diet: (1) an exo-(13)--D-glucanase, (2) an endo-(13)--D-glucanase and (3) a -D-glucosidase. The glucanases have a pH optimum of 5.4, a temperature optimum of 50°C and are optimally extracted in bistripropane buffer, pH 7.2. Levels of (13)-D-glucanase in laboratory-cultured E. superba are inversely affected by food availability, and activities double after starvation for 12 d. The increase is due mainly to higher activities of exo-acting glucanases. -Amylase and endo-(14)--D-glucanase (cellulase) activity are also present in the extracts in addition to glycosidase activity against a range of p-nitrophenyl substrates (-and -D-glocose, - and -D-galactose, -D-xylose, - and -D-mannose). Digestive activity against several acidic polysaccharides, including the acidic mucilage polysaccharide of the ice diatom Stauroneis amphioxys, is minimal and is not induced when the polysaccharide is present in the diet of E. superba. These results indicate that some, but not all, components of the algal material in the euphausiid diet can be hydrolyzed and assimilated.     

Mate choice and fitness in a hybrid frog: Rana esculenta females prefer Rana lessonae males over their own

Summary The evolution and maintenance of female choice based on purely genetic differences is still a controversial issue, not only for theoretical reasons, but also because of the practical difficulty of demonstrating the fitness consequences of preferences and heritability of and genetic variability in the chosen traits. We argue that hybrid systems (broadly defined) offer suitable models for studying mate choice according to genetic differences. We present such a study for European water-frogs of the hybridogenetic Rana lessonae/Rana esculenta complex (L/E complex). R. esculenta, originally a hybrid between R. lessonae and R. ridibunda, eliminates the L genome premeiotically and only produces eggs and sperm containing only the R. ridibunda (R) genome. Consequently, the hybrid will only persist when it lives and mates with R. lessonae in mixed populations where it can regain the lost L genome. In such mixed populations, there is strong selection against E x E matings. because these will produce no viable offspring. We tested whether females of the hybrid R. esculenta do indeed avoid their own R. exculenta males and choose males of the parental species R. lessonae instead. Eleven E females were offered a simultaenous choice between one L and one E male. Females exhibited a significant preference for L males that was determined by the type of male, rather than by its size or activity. This choice is in the direction predicted from genetics. The question of why L males agree to mate with E females, but L females only rarely mate with E males, is answered by a sexual asymmetry in the cost/benefit ratios of mating with the wrong type and the right size. Our results are consistent with the mating pattern found in natural populations, but further studies are needed to show that female choice really causes this pattern.Correspondence to: H.-U. Reyer     

Effect of ionizing radiation on haemoglobin of marine lamellibranchs

The effect of ionizing radiation on the iron-linked protein (haemoglobin) of the marine lamellibranchs Anadara granosa (Linn.) and Cardita antiquata (Lam.) from Bombay waters, India is discussed. Purified haemoglobin solutions were exposed to a ⁶⁰Cobalt source delivering a dose of about 4,600 rad/min, at the sample irradiation point. Radiation damage or degradation was measured spectrophometrically by studying changes in the absorption spectra following irradiation in the presence and absence of oxygen. Exposure to ionizing radiation in general caused a decrease in absorption of both haemoglobins, irrespective of location, viz extra-versus intracellular, at Soret (412 m), (540 to 42 m) and (574 to 76 m) peaks, and an increase in absorption at 510 and 630 m. Upon exposure to a higher dose, O₂Hb of C. antiquata showed an increase in absorption at the protein peak (280 m); exposure to a lower dose, however, resulted in decreased absorption. Furthermore, the changes following irradiation were dependent upon the initial state of the pigment. Oxyhaemoglobin, when exposed to radiation, oxidized to hemiglobin, and hemiglobin reduced to oxyhaemoglobin. The extracellular haemoglobin of high molecular weight (3x10⁶) of the false cockle C. antiquata was found to be extremely radio-resistant, whereas intracellular haemoglobin of low molecular weight (74,000) of the arcid clam A. granosa was highly radiosensitive, since it could not be exposed to doses exceeding 18,000 r.     

Co-operative breeding by the Australian Bell Miner Manorina melanophrys Latham: A test of kin selection theory

Summary A population of Bell Miners Manorina melanophrys cosisted of two social units (clans), each consisting of one or more monogamous paris and numerous helpers. Individuals of a breeding pair occupied one home range, while non-breeding individuals occupied partially overlapping home ranges. Attendants (i.e. parents and helpers) showed clear preferences as to whom they aided (fed) and how much aid they provided. A significant correlation was found between the amount of aid given by an attendant and the genetic relationship between the attendant and the recipient. This provides strong quantitative evidence for the theory of kin selection. Only 14% of eggs laid produced juveniles which lived longer than 3 months. It is suggested that the extremely high juvenile mortality favours individuals which help relatives, even while such individuals have dependent offspring of their own.     

Interactions between light and temperature on the physiological ecology ofGracilaria tikvahiae (Gigartinales: Rhodophyta)

Main effects and interactions of light and temperature on rates of growth (), net photosynthesis (P_s), and dark respiration (R) of the red seaweedGracilaria tikvahiae were investigated in outdoor, nutrient-replete continuous-flow seawater culture chambers. Below 15°C,G. tikvahiae did not grow and between 15° and 30°C, both main effects and interactions of light and temperature on and P_s were significant, which explains the occurrence of this alga as a summer annual in its northern range. Temperature interacted with light (I) through its influence on the vs I and P_s vs I curves. The initial slope of the vs I curve, , the light saturation intensity, I_s, and maximum growth rate, _max, were all significantly lowerat 15°C compared to 20°, 25°, or 30°C. Maximum values of _max, the P_s:R ratio and the net photosynthesis:gross photosynthesis ratio (P_s:P_g) all occurred at 25°C, suggesting that this is the best temperature for growth ofG. tikvahiae. Values for P_max increased up to 30°C, indicating that the temperature for maximum growth and net photosynthesis are not the same forG. tikvahiae. Significant photoinhibition of growth and photosynthesis at full incident sunlight (I₀) occurred at 15°C but not at 20°, 25°, or 30°C. Steele's equation fit the 15°C vs I data best, whereas the hyperbolic tangent function fit the 20°, 25°, and 30°C data best. Main effects and interactionof light intensity and temperature on rates of R were also significant (P<0.001). R was highly intercorrelated with and P_s (0.86r0.94), indicating that R inG. tikvahiae is primarily regulated by growth rate and not temperatureper se. Environmental factors that regulate growth, such as light intensity, exert a great influence on R inG. tikvahiae.     

How a subsocial intertidal beetle,Bledius spectabilis,prevents flooding and anoxia in its burrow

Summary All stages of Bledius spectabilis Kratz (Staphylinidae) dig wine-bottle shaped burrows in the intertidal Salicornia zone. The adult female lays her eggs around her burrow and, by remaining with them, prevents both flooding by the tide and anoxia. Flooding is prevented by an ever-ready burrow which exploits a surface tension effect and by blocking the burrow once the tide comes in. The burrow has a narrow neck (about 2 mm in diameter) leading to a living chamber 5 mm in diameter. Tides over artificial tubes in agar showed that a critical minimum neck diameter of 2–3 mm prevented sudden flooding, giving time to block the neck with mud. Blocking took about four minutes. Ensuring the burrow is reopened at each low tide is a vital role of the brooding female in anaerobic or impermeable soils. Field models of burrows became anoxic in 4 days, much less than the 4 week long egg stage. A lab model system with anaerobic agar and calculations of oxygen uptake and diffusion supported this conclusion. Mortality of orphaned larvae may be much lower, however, in burrows within large Bledius aggregations because of mitigating good drainage and soil aeration: larval mortality from physical causes in these burrows did not increase over 3 weeks but without the mother, 14% of the eggs were attacked by mould, and two burrows were taken over by a predatory carabid, Dichierotrichus gustavi, and all larvae eaten. The surface tension effects of small air-filled openings may be used by small air-breathing animals in many intertidal habitats. Like Bledius, other intertidal animals, including large ones, may block their burrows at high tide to keep them full of air. Maternal care, in particular the combination of behaviours which protect the brood from the tide and anoxia, enables this airbreathing insect to colonise the inhospitable habitat of the intertidal saltmarsh.     

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

Lack of social entrainment of free-running circadian activity rhythms in the Australian sugar glider (Petaurus breviceps: Marsupialia)

Summary Social influence on circadian activity rhythms was investigated in the nocturnal Australian marsupial Petaurus breviceps. The activity of two and two was recorded electroacoustically and observed by an IR television camera in LD 12:12 (10¹:10^-1lx) and in LL (10^-1lx) when housed isolated and in pairs (+). In LD-entrained animals the average duration of locomotion, orientation movements, grooming and total activity is influenced by social housing, and individually different activity patterns are harmonized. In constant illumination conditions, however, the members of a pair are not mutually synchronized and free-run with different circadian periodlengths. Therefore social influence on the LD-entrained activity rhythm in Petaurus should be interpreted as social masking rather than direct influence on the circadian system.     

Identification and quantitative analysis of the amino acids present in protein of the brown shrimp Penaeus aztecus

In order to better formulate an artificial shrimp diet, the protein composition of shrimp was assessed and the essential amino acids determined. Penaeus aztecus were treated to remove lipids, carbohydrates and other interfering substances, and the protein was hydrolyzed to its constituent amino acids. The amino acids were then quantitatively analyzed by gasliquid chromatography as their trimethylsilyl derivatives. The essential amino acids were then determined. Shrimp were injected with ¹⁴C-labeled glucose and analyses performed to determine which amino acids then exhibited radioactivity, i.e., were synthesized from the labeled glucose. Those amino acids which were not manufactured from the glucose were categorized as essential.     

Influence of three sediment types on copper toxicity to the polychaete Neanthes arenaceodentata

Adult male Neanthes arenaceodentata were exposed to 0.10±0.015 mg l^-1 copper in the seawater of a continuous-flow bioassay system in the presence of a sand, a mud, a mixture of the sand and mud, and no sediment, to assess the influence of sediment type on Cu-induced mortality. The sediment type did influence mortality. The time to 50% mortality was 7.8 days without sediment, 36.5 days with sand, 54.5 days with the mixture, and 50.0 days with mud. There was no threshold Cu body burden that caused death. The mean Cu concentration per gram of N. arenaceodentata (24 h after death) without sediment was 270 g, in sand 994 g, in the mixture 1047 g, and in mud 1464 g. The differences in the toxic responses are discussed.     

Rates of seasonal sediment reworking in Axiothella rubrocincta (Polychaeta: Maldanidae)

Sediment reworking rates of Axiothella rubrocincta (Johnson, 1901) (Polychaeta: Maldanidae) were measured in situ in Tomales Bay, California (USA), from August, 1969 through July, 1970. On the average, each adult worm (approximate fresh weight=1 g) reworks about 5 g dry sediment d^-1 at a mean temperature of 13.4°C and a mean salinity of 31.8.Reworking rates are positively correlated with temperature and salinity, and negatively with sediment organic carbon, sedimentation rates and grain size. An inverse correlation exists between sediment reworking rates (g dry sediment g^-1 wet wt of worm d^-1) and g wet weight of worm. Sediment parameters deseribing unworked sediments are not significantly different from those for fecal sediments. Although these data suggest this species to be a non-selective deposit-feeder, it is more likely that it is faculatively selective.