Short-term population dynamics of Tisbe cucumariae (Copepoda: Harpacticoida)

Population dynamics of a benthic harpacticoid copepod (Tisbe cucumariae) were studied in a seawater-system holding tank during fall-spring, 1979/1980, and January, 1981. These populations were opoortunistic scavengers, especially on the dead bodies of Ciona intestinalis, a solitary tunicate. Copepod population densities, life-history stage structures, sex ratio and percentages of females with egg sacs were determined in the absence or in the presence of dead tunicates (from death through dissolution to final disappearance). Taken together, all the data are consistent with a hypothesis for food-dependent cycling of populations of T. cucumariae through the following 4 density states: (1) a maintenance state of low density, which is the usual condition during the absence of a concentrated food source; (2) a colonization state of moderate density, which results from attraction and immigration to a new food source; (3) a bloom state at the food source, where a high density results from intensive reproduction combined with a tendency of all life stages to remain at the site; (4) a dispersal state of moderate density as the copepods emigrate from the site of the consumed food. Evidence is also presented for a facultative slowing of naupliar development in T. cucumariae in the absence of a concentrated food source such as a dead tunicate. Possible advantages afforded by delayed development are discussed in terms of risk-spreading in a stochastically hostile environment.     

Chromosome number in ten species of the genus Tisbe (Copepoda: Harpacticoida)

Of 10 species of the genus Tisbe, 9 were shown to possess the same haploid number of 12, whereas only 1 has a haploid number of 8. The hypothesis is advanced that n=12 is the primitive condition in the genus Tisbe and that T. biminiensis with n=8, is a recent form.     

Effect of crowding on larval viability in Tisbe holothuriae (Copepoda: Harpacticoida)

Larval Tisbe holothuriae Humes exposed to anionic detergent (0.5 mg l^-1 sodium dodecyl benzene sulphonate) were found to survive better than control larvae. The detergent appears to enhance viability by relaxing inhibitory interactions between nauplii. This conclusion is inferred from the following experimental observations: (1) crowding depresses larval viability; (2) the effect of crowding on viability is indirect, and results from behavioural changes which reduce contact with food; (3) detergent exposure delays or suppresses the behavioural response to crowding. The nature of the interactions which modify behaviour is unknown, but evidence suggests that close encounters between nauplii are important, and that they may lead to the impairment of distance-chemoreception. Naupliar antagonism could have evolved as a mechanism for staggering larval development or as a means of reducing the relative fitness of neighbouring individuals.     

Trade-off between mate guarding and mate attraction in the polygynous great reed warbler

Summary In a Swedish population of the polygynous great reed warbler Acrocephalus arundinaceus, about 40% of the males formed pairs with more than one female. Males sang two completely separated types of song: (1) long song when they tried to attract mates and (2) short shong when they guarded a fertile female. As soon as a male had attracted a female he immediately switched from long to short song and apparently guarded her for at least 3 days. Most males left their female and started singing long several days before her fertile period ended. This behavior probably increased the risk of cuckoldry. By assuming that males of polygynous species maximize their fitness over the whole breeding season (i.e., maximize fitness rate), we predicted longer mate guarding periods when the probability of attracting a second female was low and the risk of cuckoldry was high. These predictions were supported by observations. The probability of attracting a second female decreased during the breeding season and, in accordance, the number of days that males sang long song during the primary female's fertile period was negatively correlated with the time of the season. This trend held also for individual males when comparing mating periods of their primary and secondary female. The increasing rate of male intrusions with season may also have contributed to the increasing mate guarding periods. Offprint requests to: D. Hasselquist     

Tactics, effectiveness and avoidance of mate guarding in the blue-footed booby (Sula nebouxii)

We examined the dynamics and avoidance of mate guarding, by males and females, in the blue-footed booby, in which the two social mates are usually simultaneously present on the territory but each of them is unmonitored by the other for one-quarter of its time. Both sexes were promiscuous and liable to switch mates. Cuckolded individuals did not increase their overall presence on the territory, but in response to the extra-pair (EP) courtships of their mates, both sexes doubled their rate of intra-pair (IP) courtship and sometimes showed aggression. The male or female's presence depressed the social mate's EP activity, but intra-pair courtship had no such effect, tending even to propitiate that EP activity. Similarly, when females responded to their social mates' EP courtship with approach or aggression, disruption of EP activity was short-lived. Promiscuous females modified their diurnal pattern of attendance, as if attempting to sidestep monitoring by their mates, but cuckolded males matched the modification. Both sexes tended to perform their EP activities at a distance when their mates were present, possibly to evade monitoring or disruption by their mates. Male and female boobies cannot monitor their mates continuously, they do little to facultatively adjust their presence on territory to the risk of infidelity, and their immediate responses to overt infidelity have only the briefest impact; but the information they acquire while monitoring their mates may be critical to constraining their mates' infidelity and also to calibrating their own reproductive investment.     

Postcopulatory mate guarding by vocalization in the Formosan squirrel

The Formosan squirrel, Callosciurus erythraeus thaiwanensis, emitted different vocalizations in response to terrestrial and aerial predators and snakes. Each vocalization caused nearby individuals to adopt a different type of anti-predator behaviour. In mating bouts, males produced two types of loud calls: precopulatory calls, emitted before copulations, and postcopulatory calls, emitted after copulations. The latter continued for 17 min on average. The estrous female and other males attending the mating bouts stopped moving during the postcopulatory call, so that the calling male was able to tend the female without interruption. The sound characteristics of anti-terrestrial-predator and postcopulatory calls recorded in the captivity were compared, and none of the ten characters of duration and frequency measured differed between the two calls. Playback experiments also showed that responses to the sounds in two different contexts, escape behaviour and defensive immobility, did not differ. The similarity between anti-predator and postcopulatory calls is discussed with reference to the possibility of manipulation and other explanatory hypotheses.     

Vertical distributions of Calanus finmarchicus and C. helgolandicus (Crustacea: Copepoda)

The vertical distributions of the spring populations of Calanus finmarchicus (Gunnerus) and C. helgolandicus Claus are described and compared. The differences we observed between the two species have probably confused the understanding of the vertical distribution and development of the populations of Calanus spp. in the shelf seas around the United Kingdom where the species occur together. The results imply that these two congeneric species have different behaviour patterns which minimise interspecific competition where the species have sympatric distributions. C. finmarchicus has its younger development stages overlying the older stages in the water column. In C. helgolandicus the converse is true; i. e., the majority of the populations of Stage I and II copepodites of the first spring generations are found below the thermocline. It is also suggested that the different behaviour patterns lead to different feeding regimes and strategies.JONSDAP Contribution No. 52     

Elaphoidella grandidieri (Harpacticoida: Copepoda): demographic characteristics and possible use as live prey in aquaculture

In freshwater ecosystems, rotifers and cladocerans are ideal prey for fish larvae whereas copepods, due to their purported low growth rate and predatory tendency, are not. We recently isolated the parthenogenetic Elaphoidella grandidieri (Gueme et Richard, 1893) a benthic freshwater harpacticoid, from a fish farm in the State of Morelos, central Mexico and tested its potential as a live prey organism for larval vertebrates. Population growth and life table demography experiments were conducted, in 100 ml recipients with 50 ml of test medium on a diet of Scenedesmus acutus at a density of 1.0 X 10(6) cell ml(-1); the former on live algae alone while the latter on live algae as well as detritus. We also conducted experiments to document the prey preference for this copepod by the larval Ameca splendens (Pisces: Goodeidae) and Ambystoma mexicanum (Amphibia: Ambystomatidae), fed the rotifer Plationus patulus, the ostracod Heterocypris incongruens, and the cladocerans Moina macrocopa and Daphnia pulex. Elaphoidella grandidieri is relatively easy to maintain under laboratory conditions, reaching densities (copepodites and adults) of more than 10,000 l(-1). The generation time ranged between 30-45 days, depending on the diet. The net reproductive rate was as high as 60 nauplii female(-1) day (1). Population growth rates ranged between 0.03 and 0.11 d(-1), live algae being the superior diet compared to detritus. Both predators showed no preference for E. grandidieri, but in the absence of alternate prey they consumed 80% of the harpacticoids offered. The data have been discussed in relation to the potential of E. grandidierias live food for aquaculture.     

Macroalgae as habitat patch islands for Scutellidium lamellipes (Copepoda: Harpacticoida) and Ampithoe tea (Amphipoda: Gammaridae)

Scutellidium lamellipes and Ampithoe tea live in patches formed by the brown alga Pelvetia fastigiata. The distributions and abundance of both crustaceans were examined on P. fastigiata obtained during 1975 and 1976 from 7 collecting and 6 experimental sites in the mid-intertidal zone at La Jolla, California (USA). At each collecting site, the abundance of each crustacean generally increases with increasing plant size. Following defaunation and transplantation of algae, abundances of both crustaceans on individual plants appear to be governed by equilibria determined by in situ reproduction, immigration and emigration. Immigration rates of both animal species decrease significantly with increasing distance from areas occupied by P. fastigiata. During short-term experiments, neither crustacean was able to establish colonies on defaunated plants placed across a 30 m P. fastigiata-free zone and both species would have to reproduce to persitst on plants transplanted to that site. More frequent immigration and reproduction (and, for S. lamellipes at least, decreasing emigration rates with increasing patch size) would increase abundances and decrease probabilities of local extinction on larger and/or less distant plants. However, even within sites with isolated plants, the smallest plants often are not inhabited by either crustacean. The densitics of both species differ with plant size and distribution. Densities of S. lamellipes are greatest within a large aggregation of P. fastigiata, whereas those of A. tea are greatest on moderate-sized isolated plants at mid-tide levels.     

The influence of sexual selection and predation on the mating and postcopulatory guarding behavior of stone crabs (Xanthidae,Menippe)

Summary Postcopulatory mate guarding in crustaceans traditionally has been viewed as a behavioral mechanism that prevents predation on the soft post-molt female. This study tests the effects of sexual selection and predation on the postcopulatory guarding durations of male stone crabs, Menippe mercenaria, M. adina, and their hybrid. Male stone crabs were held with a pre-molt female, and either another adult male stone crab, an intermolt female, or a male blue crab, which corresponded to intermale competition, control, and predation treatments, respectively. The mating behavior of the heterosexual pair was recorded with a time lapse video system and the durations of copulation and postcopulatory guarding were measured. Males guarded longer in the intermale competition treatment than either the control or predation treatments. In the competition treatment, agonistic encounters occurred between the males at the den containing the female and several mate takeovers occurred. Females survived the predation treatment in trials in which the guarding durations were the longest, whereas females were eaten by the blue crab in trials with the shortest guarding durations. Sexual selection appears to be important in maintaining postcopulatory mate guarding in stone crabs.     

Molecular phylogeny and evolution of the pelagic copepod genus Neocalanus (Crustacea: Copepoda)

Portions of the mitochondrial genome (ca. 4 kb), encoding three protein-coding (COI, ND4L, ND6) and two ribosomal RNA (srRNA, lrRNA) genes, were sequenced for all six currently recognized species, plus one form, of the pelagic calanoid copepod genus Neocalanus. In Neocalanus gracilis, the ND6 gene was not found in the sequenced portion of the mitochondrial genome. Unambiguously aligned sequences were subjected to Bayesian, maximum-likelihood, maximum-parsimony, and neighbor-joining analyses using Eucalanus bungii as an outgroup. The resultant tree topologies from these four methods were congruent, robust, and all nodes were supported by high bootstrap values and posterior probabilities of 92–100%. Two tropical and subtropical species (N. gracilis and N. robustior) occupied the most basal position, and a subantarctic (N. tonsus) and three subarctic Pacific species (N. cristatus, N. plumchrus, and N. flemingeri) diverged subsequently. Transequatorial dispersal of the ancestral population during glaciations is suggested for this pattern of speciation, in which sister clades exhibited antitropical distributions. Although the area of ocean is much broader in the subantarctic than the subarctic Pacific, a higher number of species occur in the subarctic Pacific (three) than the subantarctic (one). The possibility that marginal seas, such as Japan Sea and Okhotsk Sea, function as natal areas for the divergence of species is discussed.     

Assessments and decisions in Metellina segmentata (Araneae: Metidae): evidence of a pheromone involved in mate guarding

Assessments and decision-making underlying the initiation of mate guarding in a common web-building spider, Metellina segmentata, are examined in a series of field and laboratory studies. Adult males do not build webs but wander in search of females and mating opportunities. Adult males then wait at the edge of the webs of females and guard them prior to courtship and mating. Guarded females were heavier, larger and carried more mature eggs than solitary females. An active process of information gathering is apparent from introductions of males to the webs of females. Males make accurate assessments about female quality, even in the absence of the resident female. Cues involving web architecture are not used. Males may assess pheromonal cues on the web of the female in deciding whether to guard or abandon a female.     

Tisbe (Copepoda: Harpacticoida) species from the Lagoon of Venice. I. Seasonal fluctuations and ecology

Distribution and abundance of Tisbe species were studied throughout a period of 3 years at 3 stations in the Lagoon of Venice (Italy). At 2 other stations samples were taken occasionally, as also at the Lido station (open sea). Twelve species may be considered as common inhabitants of the lagoon, and 3 species appear to be occasional transients, compared to the 9 species found at the Lido. A comparison of the physical data shows that the stations were similar in temperature and salinity, but differed primarily in pH values, sediment characteristics, algal substratum and faunal benthic community. The fact that remarkable differences in species composition and distribution of Tisbe could be observed between the various parts of the lagoon and the lagoon and the open sea, indicates that temperature and salinity are not so important for species diversity as are biotic factors. A particularly strong difference existed between samples taken at the bottom and from pilings, which constitute a very special biotope for the benthic communities in the lagoon and seem to be a preferred habitat for T. lagunaris and T. cucumariae. The data suggest that there is a certain interaction between T. holothuriae and the other species. Whenever T. holothuriae was abundant, the other species were rare, but T. clodiensis, T. dobzhanskii (Stations 1 and 2) and Tisbe sp. (Station 3) attained high relative abundance whenever T. holothuriae decreased considerably in number. T. holothuriae represents certainly the hardiest species of the lagoon, displaying a great ability to cope with stress conditions, even those present in the interior lagoon where strong fluctuations in salinity, temperature, pH and oxygen prevail.     

Scanning electron microscope study of the modified spine characteristic for males of some Tisbe (Copepoda: Harpacticoida) species

The scanning electron microscope proved to be an ideal instrument for examining the species-specific spine (on the endopodite basal segment of male P2) in the group of sibling species Tisbe holothuriae, T. pontina and T. battagliai. Three other species: T. gracilis, T. cucumariae and T. biminiensis belonging to the gracilis group are included in the present study. A possible functional significance of the spine is discussed.     

Brood size and chemical composition of Pareuchaeta norvegica (Crustacea: Copepoda) in Loch Etive,Scotland

Brood size and chemical composition (carbon, nitrogen, and ash) of Pareuchaeta norvegica in Loch Etive, Scotland have been determined. From 23 to 37 eggs were found in each egg mass (mean=30 eggs). The eggs are oval; the mean size of the longer axis is 438 m. Variation in egg mass (as dry weight) is related to increase in dry weight of the maternal body. an increase of 1 mg in maternal body dry weight produces an increase of 0.03 mg dry weight in the egg mass. Carbon content increases with increasing body dry weight both in pre-spawning and spent females, but nitrogen content decreases with increasing body weight. Thus, the carbon:nitrogen ratio increases with increasing body dry weight. The carbon content of an egg mass ranges from 61.4 to 65.3%, and nitrogen content from 9.6 to 13.2% of the total dry weight. Some examples of partitioning of chemical components of the maternal body into the egg mass are also described.     

Characteristics of suspensions of Kuwait oil and Corexit 7664 and their short-and long-term effects on Tisbe bulbisetosa (Copepoda: Harpacticoida)

A technique for preparing seawater suspensions of Kuwait oil and Corexit was developed. The resulting hydrocarbon concentrations were analysed by gas-chromatography and spectrofluorometric methods and the stability of the suspensions with time was determined. It was established that the suspensions have an effective stability from Days 3 to 15 after preparation, since in this period the concentration varies within a relatively narrow range. Adult female Tisbe bulbisetosa appeared to be quite tolerant of this type of hydrocarbon suspension, in short-term experiments, especially considering that the concentrations used in the bioassays were about 200 times higher than those measured in a relatively polluted area of the lagoon of Venice. Long-term effects on number of eggs produced, number of nauplii and hatching success for females of the third and fourth generations, subject to continuous exposure, were negligible compared with controls.     

Tisbe pori n. sp. (Copepoda: Harpacticoida) from the Mediterranean coast of Israel and its cultivation in the laboratory

Tisbe pori n. sp., from the Mediterranean coast of Mikhmoret, Israel, was cultivated in the laboratory and is described. The relative food values of 9 species of marine algae were evaluated according to the copepods' development time (length of period from hatching to reproductive maturity). Individuals fed on Dunaliella tertiolecta or D. primolecta developed the fastest. The rate of feeding of the adults on Dunaliella primolacta was determined. In the temperature range 17° to 31.5°C, the shortest development time (7 days) was at 26° to 28°C. The range of salinity tolerated by adults was 26 to 48.