Ovigerity,selective predation,and variable diel vertical migration in Euchaeta elongata (Copepoda: Calanoida)

Summary We present a statistical analysis of a previously published (Yen, 1983) but heretofore unanalyzed data set on the vertical distributions and diel vertical migration (DVM) of adult females of the marine planktonic copepod Euchaeta elongata in Dabob Bay, Washington, USA. Non-ovigerous females were strongly migratory on all four dates sampled, residing between 75–175 m during the day and at shallower depths during the night, commonly entering the upper 50 m of the water column. In contrast, ovigerous females were non-migratory or weakly migratory, largely remaining between 100–175 m both day and night, and entering the upper 50 m of the water column only rarely. Thus non-ovigerous females always migrated much more strongly, as measured by both amplitude of migration and the proportion of animals migrating, than did ovigerous females. These results led us to hypothesize that differential susceptibility to visually orienting predators was the cause of these differences in DVM behavior in female E. elongata, and we subsequently undertook an experimental study of the feeding selectivity of the copepod's natural predator, Pacific herring (Clupea harengus pallasi). Pacific herring exhibited a highly significant preference for ovigerous over nonovigerous adult female E. elongata. The demographic consequences of variable DVM in adult female E. elongata were investigated by way of life table analyses. Results indicated that under conditions of thermal stratification of the water column there is a distinct demographic disadvantage (reduced rate of realized population growth) incurred by non-migratory or weakly migratory ovigerous females due to delayed egg development at cooler subsurface temperatures. We conclude that ovigerous female E. elongata remain at depth both day and night to avoid visually orienting predators, and that such behavior must afford the copepod a demographic advantage of no less than a 26% reduction in adult mortality to offset the demographic cost of delayed egg development.     

Reproductive effort and reproductive nutrition of female desert tortoises: essential field methods

I used three innovative, nondestructive field methods (gas dilution,doubly labeled water and radiography) to measure individualenergy and water budgets of wild, female desert tortoises (Gopherusagassizii). With these budgets, I evaluated whether body reserveshelp females produce eggs independent of rainfall and food availability.Female desert tortoises used large seasonal and annual changesin metabolism and body water, protein and energy reserves tosurvive and produce eggs. Although lipid reserves are importantto female desert tortoises, nitrogen or crude protein appearsto be the primary limiting resource for producing eggs. By reducingmetabolic rates 90%, females conserved enough body reservesto produce eggs during extreme drought conditions; this is aneffective bet-hedging reproductive pattern in an extreme andunpredictable environment.     

Steroid hormone profiles and relative body condition of calling and satellite toads: implications for proximate regulation of behavior in anurans

Males of most anuran species (frogs and toads) vocalize to attractmates. However, individuals of many vocal species may also adoptalternative noncalling "satellite" tactics. Satellite malescharacteristically remain in close proximity to calling conspecificsand attempt to intercept incoming females attracted to advertisingmales. Emerson proposed that alternation between calling andnoncalling behavior in anurans is mediated by a reciprocal interactionbetween circulating levels of corticosterone and androgens thatis driven by depletion of energy reserves during vocalization.We tested this hypothesis by examining steroid hormone profilesand the relative body condition of calling and satellite Woodhouse'stoads (Bufo woodhousii) and Great Plains toads (B. cognatus).Consistent with Emerson's hypothesis, callers had significantlyhigher circulating corticosterone levels and were in bettercondition than satellites. However, levels of testosterone anddihydrotestosterone did not differ significantly between satellitesand callers, and we found no evidence that high levels of corticosteronehad an inhibitory effect on androgen production in either species.These data thus support a relationship between corticosteronelevels and depletion of energy reserves during bouts of vocalizationbut suggest that alternation between calling and satellite behaviormay be associated with direct effects of corticosterone on brainvocal control centers. We propose a model that incorporatesrelationships among energy reserves, androgens, corticosterone,and arginine vasotocin-producing neurons in the telencephalonto explain transitions between calling and satellite tacticsin toads.     

The reproductive biology of Cyclops vicinus

Investigations of the reproductive biology of Cyclops vicinusrevealed that mated females oscillate between gravid and nongravidreproductive conditions. The gravid condition can be dividedin two recognizable phases: gravid/nonovigerous and gravid/ovigerous,the former phase being shortest. The maturation of new oocytestakes place when the old egg sacs are still being carried; thisensures a rapid clutch succession. Females which remain unmated,extrude few eggs, in no case complete egg sacs, and remain gravidthus conserving oocytes. Females which mate only once, showa similar reproductive pattern (clutch size and clutch succession)to those which remain combined with males, and thus have theopportunity to remate, but tend to produce fewer clutches. Malesare able to fertilize 3–4 females day^–1. Matingcapacity of males is possibly limited by the time needed tofill a new spermatophore. Short-term starvation (5 days) lengthenedclutch-to-clutch periods and diminished clutch size. When thestarvation period started in the gravid/ovigerous phase, a normalclutch was extruded but no new oocytes matured during starvation,indicating that the energy for egg maturations is provided inthe first part of the reproductive cycle.     

Small-scale distribution of macroplankton and micronekton in the Ligurian Sea (Mediterranean Sea) as observed from the manned submersible Cyana

A series of eight submersible dives (the MIGRAGEL I cruise)was made during late April 1986 using the French submersibleCyana to investigate macrozooptankton in the upper 400–700m of the water column. Paired day and night dives were madeat stations 3, 6, 13 and 23 nautical miles off Cape Ferrat,near Villefranche-sur-Mer, France; the distances represent differentareas in the frontal system of the Ligurian Sea. Detailed day/nightvertical distribution data are shown for the most abundant species;these include the narcomedusa Solmissus albescens, teleost fishCyclothone spp., small appendiculanans (primarily Oikopleuraalbicans), large appendicularians (an undescribed oikopleurid),diphyid siphonophores (mostly Chelophyes appendiculata) andan abundant lobate ctenophore. Salps, pyrosomes, amphipods (Phroniumasedentaria), pteropods (Cavolinia inflexa), macroscopic ‘star-like’protozoa and marine snow are also briefly discussed. The coastalzone was dominated by small appendicularians in the upper layers,with other filter feeders including large appendicularians indeeper water—these just above a non-migratory populationof carnivorous Cyclothone. The carnivorous medusa Solmissusalbescens moved throughout the upper 600 m in the course ofits diel vertical migration. Offshore, carnivores were dominantthroughout the water column, with numerous diphyid siphonophoresin the upper layers, and Cyclothone, lobate ctenophores andmacroprotozoa abundant in deeper water. Solmissus was also present,and was more numerous offshore than in the coastal zone.     

The effect of temperature and water quality on the in vitro development and survival of Asellus aquaticus (Crustacea: Isopoda) eggs

The rate of development and degree of survival of Asellus aquaticus eggs outside the marsupium of ovigerous females are affected by water quality and temperature. Eggs were maintained in polluted river water and relatively clean canal water. Developmental rates increase with increased temperature, but survival decreases. Eggs from polluted site ovigerous females survive better in clean water than in polluted water. Eggs from clean site ovigerous females maintained in polluted water have significantly lower survival rates than eggs from the polluted site at all temperatures tested. The developmental rate of clean site eggs is increased significantly in polluted water at 10–25 °C, possibly as a response to the stress imposed upon them.It is suggested that the method outlined might form the basis of a useful bioassay technique for measuring water quality.     

Reproduction and growth of the Arctic hyperiid amphipod Themisto libellula Mandt

Summary The hyperiid amphipod, Themisto libellula, is abundant in Arctic seas and is an important prey for fish, birds and mammals. Characterization of its life cycle has been hampered by the lack of winter collections. In this study, that portion of the hyperiid population inhabiting the upper 30 m of the water column in Frobisher Bay was sampled periodically in winter, spring and summer during five consecutive years. In one year the winter and spring collections included large numbers of ovigerous females. These samples provide new information about timing of reproduction, size at maturity, fecundity, embryonic development, hatching, release and rate of growth of T. libellula in the Arctic. The average length of the ovigerous females was 23.3 mm. This and the summer growth rate of about 6 mm/month indicates that they can reproduce when one year old. Eggs are laid prior to early February, and although some young may be released from the marsupium as early as mid-March, most are liberated during April and May. The fecundity is significantly correlated with female length and an average 23 mm long individual had 275 embryos. The size-specific fecundity is similar to that of hyperiids from temperate and subtropical waters.     

Locomotor activity pattern in pairs of the subsocial desert isopod Hemilepistus reaumurii

Locomotor activity rhythm of fresh adult of H. reaumurii was recorded in pairs with ovigerous and with non-ovigerous females. According to double-plotted actograms and waveform curves, results showed the presence of different locomotor patterns. In fact, locomotor profiles of pairs with ovigerous females were mainly bimodal whatever the recording conditions, whereas locomotor patterns were trimodal in pairs with non-ovigerous females. Results showed also that locomotor activity rhythm was more stable in the presence of the synchronizer (nLD cycle) than in constant darkness whatever the recording type. Moreover, the most important stability of the locomotor rhythm was observed under constant darkness and individuals were more active in pairs with ovigerous females than in pairs with non-ovigerous females.     

Effect of body size and sugar meals on oviposition of the yellow fever mosquito,Aedes aegypti (Diptera: Culicidae)

The effects of dietary sugar and body size on the oviposition of Ae. aegypti were studied under laboratory conditions. In female mosquitoes provided with sugar, the start of maximum fecundity was significantly delayed and the oviposition period was longer than in females provided with water. The peak of oviposition was also delayed in sugar‐fed females. Large females oviposited more eggs per day than small females at maximum fecundity and during eight days of observations. Large females also visited significantly more water‐containing cups in their cages per day than small females at maximum fecundity. During the eight days of observations, large females and sugar‐fed females visited more water‐containing cups in their cages than water‐fed small females. Both large females and sugar‐fed females oviposited their eggs at sites higher above the water line than water‐fed small females. These results suggested that large and sugar‐fed female Ae. aegypti mosquitoes had more energy reserves and oviposited their eggs at higher sites, which would lead to a time lag in hatching.     

A water-borne sex pheromone and trail following in the apple snail, Pomacea canaliculata

We investigated whether individuals of the apple snail Pomaceacanaliculata were attracted by conspecifics or follow mucustrails of other individuals. The snails' behaviour was studiedby a series of choice experiments in a T-maze and in Petri dishes.Both males and females chose the side with a snail of the oppositesex significantly more frequently than the control side withouta snail. Males were attracted by water conditioned with femalesmore frequently than unconditioned water, whereas females didnot show a preference for male-conditioned water. Moreover,juveniles were not attracted by water conditioned with a male,a female and a juvenile. These data indicate that males wereattracted by female odour, which contains one or more water-bornesex pheromones. In addition, both males and females follow mucustrails of snails of the opposite sex. Since females also followedtrails of females, the function of trail following is not necessarilyrelated to reproduction. In these experiments P. canaliculatadid not distinguish the direction of the trail. (Received 10 March 2007; accepted 12 July 2007)     

Diurnal vertical migration of Daphnia hyalina Leydig, 1860 (Crustacea: Cladocera) in Lake Bled (Slovenia) in relation to temperature and predation

Diurnal vertical migration (DVM) of Daphnia hyalina in Lake Bled was most intense during summer stratification. The extent of DVM varied with the size of the animal and its reproductive state. Migration distances were shortest in immature specimens and longest in ovigerous females. During daytime, ovigerous females stayed deeper in the water column than females without ova or immatures. The daytime temperature of water at the median depth of the ovigerous females did not exceed 10 °C, even in the warmest season. At night they migrated upward to an environment which was warmer by as much as 9 °C.Laboratory observations indicate that specimen's size and water temperature determine the velocity of passive sinking, such that morning descent of the different groups of Daphnia can be explained by passive sinking alone.Our hypothesis is that the distribution of different groups of D. hyalina in Lake Bled is influenced by two types of predators: fish (Perca fluviatilis L. and Rutilus rutilus (L.)) and larvae of Chaoborus flavicans (Meig.), the latter appearing in the epilimnion during the night. Fish predation has a key-role at the beginning of thermal stratification. Supposing that in spring the gene pool of Daphnia consists of a mix of different genotypes, distributed at different depths during the day, fish predation combined with a presence of fish chemicals favored genotypes with a lower day-depth during the spring/summer period.     

Choosy females and indiscriminate males: mate choice in mixed populations of sexual and hybridogenetic water frogs (Rana lessonae, Rana esculenta)

For several decades, behavioral ecologists have studied theeffects of the environment on the behavior of individuals;but only fairly recently they have started to ask the reversequestion: how do the behavioral strategies of individuals affectthe composition and dynamics of populations and communities?Although intuitively obvious, this feedback from individualto higher levels is difficult to demonstrate, except in systemswith exceptionally fast and marked responses of the populationsto the behavior of its members. Such a system exists in sperm-dependentspecies. In European water frogs, for instance, successfulreproduction of a hybrid species (R. esculenta, genotype LR)requires mating with one of its parental species (R. lessonae,genotype LL), except in the rare cases where hybrids are triploid.The sexual host LL, however, should avoid matings with the sexual parasite LR, because the resulting LR offspring willeliminate the L genome from their germ line. In this studywe investigate how this conflict is solved. Since water froghybrids come in both sexes, rather than as females only likein other sperm-dependent systems, we performed the tests withboth females and males. One individual was given a choice betweentwo individuals of the opposite sex, one an LL and the otheran LR. In both species, females showed the predicted preferencefor LL males, whereas males did not discriminate between LLand LR females. On the individual level, we interpret the sexdifference in choosiness by the lower costs from mating withthe wrong species (LR) and the higher benefits from matingwith large individuals in males than in females. In "normal"species, male preference for large (i.e. more fecund) femalesis advantageous, but in this system such a choice can resultin mating with the larger LR females. With respect to the structureand dynamics of mixed populations, we discuss that the observed female preference is consistent with the higher mating successof LL males found in nature. Hence, mate female choice is astrong candidate for a mechanism promoting coexistence of thesperm-dependent hybrid and its sexual host. This confirms predictionsfrom previous theoretical models.     

Body condition changes of sand martins (Riparia riparia) during breeding, and a comparison with fledgling condition

The body condition of adult sand martins during the breeding season and a sample of fledglings was investigated by carcass analysis and by examination of live birds in the field. Fat scoring of live individuals was a reliable predictor of an individual's lipid content in most instances. Body mass changes during breeding were related to changes in the size of birds' lipid reserves, pectoral muscle mass, body water, and to development of the reproductive system. In both males and females, reserve lipid declined between the onset of breeding and nestling rearing. Pre-breeding males had significantly greater pectoral muscle masses than did nestling rearing females. The potential significance of protein in pectoral muscles as an energy reserve during breeding was small compared with lipid reserves. By comparing potential energy reserves in body lipid and protein with daily energy expenditure, it was calculated that neither incubating nor nestling rearing adults could survive a day of normal activity without feeding. The selective premium on adults optimizing their use of time and energy for self-maintenance behaviour during breeding is therefore likely to be great.     

Distribution and habitat preferences of two grapsid crab species in Mar Chiquita Lagoon (Province of Buenos Aires,Argentina)

Cyrtograpsus angulatus andChasmagnathus granulata (Grapsidae) are the two dominant decapod crustacean species in the outer parts of Mar Chiquita Lagoon, the southernmost in a series of coastal lagoons that occur along the temperate Atlantic coasts of South America. Distribution and habitat preferences (water and sediment type) in these crab species were studied in late spring. There is evidence of ontogenetic changes in habitat selection of both species. Recruitment ofC. angulatus takes place mainly in crevices of tube-building polychaete (Ficopomatus enigmaticus) “reefs” and, to a lesser extent, also in other protected microhabitats (under stones). In the latter, mostly somewhat larger juveniles were found, suggesting that these are used as a refuge for growing individuals. Adults are most frequently found on unprotected muddy and sandy beaches.C. angulatus was found in all parts of Mar Chiquita Lagoon, including freshwater, brackish, and marine habitats.C. granulata, in contrast, was restricted to the lower parts of the lagoon, where brackish water predominates and freshwater or marine conditions occur only exceptionally. It showed highest population density on “dry mud” flats and inSpartina densiflora grassland, where it can build stable burrows and where high contents of organic matter occur in the sediment. Such habitats are characterized by mixed populations of juveniles (including newly settled recruits) and adults, males and females (including a high percentage of ovigerous). Unstable “wet mud” as well as stony sand were found to be inhabited by chiefly adult populations, with only few ovigerous females. In “dry mud” flats, the proportion of males increased vertically with increasing level in the intertidal zone, showing a significantly increasing trend also in their average body size. These observations may be explained by higher resistance of males, in particular of large individuals, to desiccation, salinity, and temperature stress occurring in the upper intertidal. However, an opposite, or no such, tendency was found in the distribution of ovigerous and non-ovigerous females, respectively. With increasing distance from the water edge, salinity increased and pH decreased significantly inC. granulata burrows, whereas temperature showed no consistent tendency within the intertidal gradient. A highly significant linear relationship (r=−0.794; P<0.001) between salinity and pH in water from crab burrows is described. This regression line is significantly different from one that had been observed in water from the lagoon, indicating consistently lower pH values at any salinity level in burrow water. This is interpreted as a result of crab and/or microbial respiration.     

The energetic costs of tail autotomy to reproduction in the lizard Coleonyx brevis (Sauria: Gekkonidae)

Summary Energy reserve utilization and energy budgets were compared in tailed and tailless adult female Coleonyx brevis. Carcass, fat body and caudal energy reserves were used for vitellogenesis; mass and energy content (cal/mg and/or cal/reserve) of each were significantly lower at oviposition than at the initiation of vitellogenesis. Total energy reserves accounted for 53% of the reproductive energy investment in tailed females compared to 29% in tailless females. Tailed females had over twice as many reserve calories for egg production than tailless females. Caudal energy reserves represented 60% of the total reserves of tailed females and were one-third greater than the total energy reserves of tailless females. To produce a clutch of eggs both tailed and tailless females supplemented energy reserves with net metabolizable energy that was available after metabolic costs were paid. Tailless females had a significantly greater rate of food ingestion and more net metabolizable energy available for reproduction than tailed females, yet allocated significantly fewer calories/day to reproduction than tailed females, primarily because of the loss of caudal reserves. Reproductive efforts of tailed and tailless females were equivalent. However, the loss of caudal reserves resulted in the production of eggs that were significantly lower in mass and energy content (cal/mg and cal/egg) than when caudal reserves were used. Results empirically support the hypothesis that reproduction has energetic priority over tail regeneration in short-lived, iteroparous species with a low probability of future reproductive success.     

Population biology and breeding cycle of the burrowing shrimp Callichirus seilacheri (Decapoda,Callianassidae) from the eastern tropical Pacific

Year-round continuous reproduction in tropical regions is an established paradigm in marine ecology. In this study, we tested this paradigm using the ghost shrimp Callichirus seilacheri from the tropical eastern Pacific as a model species. We also examined size-frequency distribution, sex ratio, and recruitment cycle to contribute to the biological knowledge of this species. To this end, a total of 456 individuals of C. seilacheri were collected during 12 months of sampling. Population structure was symmetrical for both sexes, and the overall sex ratio did not differ from evenness. Males outnumbered females in smaller size classes, though, revealing a potential sex-dependent mortality in small individuals. The breeding pattern followed the well-marked seasonal regime of the region, with ovigerous females registered during the rainy season. While natural variation in the seawater temperature had no influence on reproduction of this species, changes in water salinity possibly triggered the appearance of egg-bearing females in the population. Recruitment occurred throughout the year but was more intense during the dry season, following the appearance of ovigerous females. The adaptability of the life cycle of C. seilacheri to the seasonal climate provides further evidence that reproduction in tropical species is not always continuous.     

Flight energetics in relation to sexual differences in the mating behaviour of a mayfly,Siphlonurus aestivalis

Summary Mayflies (Ephemeroptera) are known to have short adult life-spans. Adults are unable to feed, and they utilize reserves stored during their aquatic larval stage. Energy reserves (fat, glycogen, and free sugars) of mature larvae, subimagoes and imagoes of both sexes of Siphlonurus aestivalis Eaton were compared. All the stages of both sexes had low glycogen and free sugar contents, and the only significant change occurred during the transformation of the mature larva to subimago when almost all the reserves of free sugars were used up. Glycogen and free sugars may serve as energy sources permitting individuals to swim and fly out of the water during emergence. Fat made up most of the energy reserves of mature larvae and was the main source of energy used during the final development of both sexes. Young adult males had high fat reserves which they used as a source of energy for their swarming flights. In contrast, females did not seem to use a significant amount of fat for flight. This difference is probably related to the different mating strategies of the sexes in this species. Males perform long flights waiting for females, whereas females perform only brief flights to mate and reproduce.     

THE ECOLOGY OF THE HONG KONG LIMPETS CELLANA GRATA (GOULD 1859) AND PATELLOIDA PYGMAEA (DUNKER 1860): REPRODUCTIVE BIOLOGY

Cellana grata is large, relatively long lived (3 years), withone winter breeding season each year which can be correlatedwith major seasonal climatic and hydro-logical events, e.g.,winter reduction in temperature and higher seas. C. grata ischaracterised by a relatively low value of mean annual reproductiveeffort (males: 23%; females: 18%). Patelloida pygmaea is small,short lived (1 year), with one spring breeding season each yearand devotes a large proportion of metabolic energy for reproduction.P. pygmaea showed a higher value of mean annual reproductiveeffort (males: 46%; females: 31%) than C. grata. Mature gonads of Cellana grata first appeared in June and itreached a peak maturity (100%) in September, remaining so untilDecember, i.e., the maturation of the gonads coincided withincreasing water temperatures, with spawning associated witha decrease in water temperature. Gonadal development by Patelloidapygmaea began in either July or August and reached a peak inDecember, remaining so until a major peak in spawning took placebetween April and May, i.e., gonad maturation coincided witha decline in water temperature, with spawning following thesubsequent increase. Limpet gonadal cycles are likely gearedto temperature and food supply. Spawning occurred between Novemberand December, when wave heights are typically 1 m higher thanin summer. (Received 1 December 1992; accepted 25 June 1993)     

Reproductive biology of the isopod Excirolana braziliensis at the southern edge of its geographical range

A full analysis of the reproductive biology of the isopod Excirolana braziliensis Richardson 1912 was conducted in a sandy beach of Uruguay, located at the southernmost edge of its distributional range in the Atlantic Ocean. Reproductive and recruitment periods of E. braziliensis were concentrated in austral summer. Females with oostegites appeared in November, whereas total biomass, individual sizes and fecundity of ovigerous females peaked between December and January. These concurrent traits were responsible for the significant peak of juveniles in January. The size at maturity was 9.88 mm. Four embryonic developmental stages were described and identified: mean length linearly increased from stages I to III, whereas dry weight exponentially decreased from stages I to IV. The high reproductive output (0.41–0.58), reported for the first time in this isopod, exceeds the rates documented for other isopods. Reproduction of E. braziliensis at the southern edge of its range is semelparous: females produce one brood during the reproductive season, exhaust their energy reserves during incubation, and probably die at the end of the reproductive season. A macroscale comparison suggests that E. braziliensis at the southern edge of its range counteracts its narrow reproductive period by a short incubation period with larger individual mature female and embryo sizes, higher fecundity and a higher percentage of ovigerous females than in subtropical and tropical populations. These extreme reproductive indicators could be attributed to the internal retention of embryos that assures offspring survival, coupled with a high adaptation capability to environmental variations across its range.     

The effect of adult feeding on lipid and protein reserves in African armyworm, Spodoptera exempta, moths before and during reproduction

ABSTRACT. Newly-emerged Spodoptera exempta (Walker) (Lepidoptera: Noctuidae) moths contain high levels of lipid held largely in the abdominal fat body, the quantity depending on the larval feeding conditions. There is a positive relationship between weight-related lipid content and moth weight, which is consistent for female but not for male moths, suggesting that larval feeding conditions producing large individuals allow the accumulation of quantitatively disproportionate lipid reserves. Male and female moths have comparable levels of abdominal protein.
Changes in the water content in starved moths or ones provided with distilled water or sucrose solution show that while starved individuals die rapidly from desiccation, water-fed moths regulate their water contents between narrow limits which are higher for females than for males. Sucrose-fed moths maintain higher, more variable water contents probably due to the phagostimulatory effect of the sugar.
Reproducing and unmated moths are able to supplement their lipid and, to a lesser extent, their protein reserves following carbohydrate uptake.
During both larval and early adult stages, the capacity to accumulate lipid reserves in excess of those apparently required for reproduction, suggests that these reserves also provide the main fuel for the prolonged flights of which migratory individuals are known to be capable.