On the dynamics of exploited populations ofTisbe holothuriae (Copepoda,Harpacticoida)

A total of 75 weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes were exposed to 0.06, 0.3, 1.5, 7.5 or 37.5 μg Cd⁺⁺l^?1, combined with exploitation rates of 10, 30, 50, 70 or 90% under conditions of surplus food supply at 22° C and 30‰ S. An unusually high mortality was observed for some weeks which could not be ascribed to the added amounts of Cd⁺⁺. A change in properties of supplied water is considered responsible for the unintentional perturbation which offered a further opportunity to study the effect of compensating reactions in stress situations. The results obtained clearly indicate an effect of exploitation rate on responses to detrimental influences. A superior performance has been found in populations exploited at higher rates. The findings are discussed with respect to earlier investigations on the population dynamics ofT. holothuriae.     

On the dynamics of exploited populations ofTisbe holothuriae (Copepoda,Harpacticoida)

A total of 30 weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes were exposed to 148 or 222µg Cd⁺⁺ l^–1, combined with exploitation rates of 10, 30, 50, 70 or 90% under conditions of surplus food supply at 22 °C and 30 S. During the observation period, most populations exhibited distinct U-shaped density trends. Addition of Cd⁺⁺ prolonged and reinforced the downward trends present in the initial periods of the experiments. If population densities, both in control and test populations, fell short of a certain value, the age structures shifted in favour of the nauplii. More pronounced reductions in population densities of the test populations did not result in increased elevations of the nauplius fraction. Within 23 weeks (or 20 generations), the effect of Cd⁺⁺ on population density was counteracted by an acclimation process. No relationship could be established between exploitation rate and observed effects. Erroneous assumptions on the adaptability of parametric methods had led to invalid statements on the dispersion of data published in previous papers of this series. These statements have been revised. Possible limitations of the significance of the results obtained are discussed in some detail.     

Influence of three bacteria strains on the population dynamics ofTisbe holothuriae (Copepoda,Harpacticoida)

The influence of three bacteria strains (Helgo 21, NCMB 308 and NCMB 13) on the life cycle ofTisbe holothuriae Humes 1957 was investigated under constant experimental conditions (19–20°C and 38‰ S). For each of the first six experiments, females (F1), whose life history was followed, were obtained from a common mother (F0). Experiment n^o 7 was carried out with adult females (F2) obtained during exp. 4. Females were bred in 50-ml dishes and transferred to a fresh container as soon as they produced a new egg-sac. Offspring (F2: exp. 1–6; F3: exp. 7) from successive egg-layings were counted as soon as they moulted into adults. Larval mortality was estimated by enumeration of the adults issuing from a known number of nauplii that had been fed different bacteria strains. Some difficulties arose in obtaining adults with NCMB 13: it appeared to be necessary to provide this first generation with a small quantity of TetraMin (commercial fish food) in order to initiate reproduction and obtain a second generation. However, in exp. 4, F2 adult females fed NCMB 13 became ovigerous and fertile without needing TetraMin. This result is interpreted as denoting an indispensable period of adaptation to the diet, which probably requires important changes at the enzymatic level. One experiment was carried out with Helgo 21; another one with NCMB 308; two with NCMB 13 (with a unique additional meal of TetraMin to allow reproduction), two with NCMB 13 during larval development, then only tetraMin during adult stage, and one with NCMB 13 alone (with females issued from exp. 4). Data obtained regarding longevity, duration of reproduction period, number of egg-sacs and number of adult descendants show an important variability between bacteria strains. Lower results were almost always obtained with Helgo 21 and NCMB 308. An interesting result lies in the maximum number of egg-sacs: low with Helgo 21 and NCMB 308 (5), but very ample with NCMB 13 (11 in one experiment). In experiment 7, carried out with animals bearing a large coefficient of inbreeding (0.375), the results concerning the maximum number of egg-sacs are good (10) as are the mean number of descendants per female (294: the highest number of the seven experiments). These results indicate that genetic factors involved in the reduction of descendants in ageing laboratory cultures can be suppressed to a certain extent by nutritional factors. The poor results obtained with Helgo 21 and NCMB 308 alone must be considered with caution: after a period of adaptation with an alimentary complement, these strains could perhaps permit an interesting production of copepods.     

Effect of density on larval development and female productivity ofTisbe holothuriae (Copepoda,Harpacticoida) under laboratory conditions

The harpacticoid copepodTisbe holothuriae has been cultivated in the Helgoland laboratory for more than 20 years. The effects of density on the larval development and the female productivity were studied by comparing two culture systems: (1) enclosed system, and (2) running-water system. In both systems, a nutritious mixed diet ofDunaliella tertiolecta, Skeletonema costatum, and granulatedMytilus edulis was offered. Larval mortality, larval development and female productivity are found to be significantly dependent on both the population density and specificity of the culture system. Increasing density causes higher larval mortality, longer larval development time, and a reduction in female productivity. In comparison with the enclosed system, the running-water system shows decisive advantages: larval mortality is about 20% lower, the rate of larval development is about two days shorter, and there is a very high rate of nauplii production. The sex ratio exhibits high variations, but in general, there is no clear relationship between sex ratio and population density. Nevertheless, when reared in the running-water system, a relatively high percentage of females (>45%) was found at lower densities.     

Studies on the selective agents operating in experimental populations ofTisbe clodiensis (Copepoda,Harpacticoida)

In order to determine the biotic selective factors affecting the fitness of genotypes in the polymorphic speciesT. clodiensis, five types of experimental populations were utilized:

1. Populations started with equal numbers of virgin females and males of the three genotypes (pp, pP, PP) at f(p)=f(P)=0.5 in Hardy-Weinberg proportions;
2. Populations started with equal numbers of heterozygous females and males;
3. Populations started with equal numbers of recessive homozygous females and dominant homozygous females, each fertilized by males of the same genotype;
4. Populations with virgin females and males of the three genotypes at f(p)=0.2, f(P)=0.8 in Hardy-Weinberg proportions;
5. One population as in (4) but with f(p)=0.7 and f(P)=0.3.

The populations were sampled at about three weekly intervals in order to follow the gene frequencies for about fifteen generations. A comparison between group (3) and other groups suggests that interaction between genotypes may affect the survival of the genotypes concerned. Comparison of groups (1) and (2) seems to indicate that sexual selection is also involved. The estimated selective coefficients, which appear to vary in different periods and groups of populations, lead to the conclusion that fitness inT. clodiensis is a function of the genetic structure of the population as well as of the population density. The selection pressure is then determined by a great number of biotic components operating in different stages of the life cycle; and it seems reasonable to conclude that inT. clodiensis, at least under these experimental conditions, a frequency-dependent selective mechanism is involved.     

Zur Dynamik exploitierter Populationen vonTisbe holothuriae (Copepoda,Harpacticoida)

In weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes, effects of exploitation rate on population density, age distribution, growth, as well as numerical and biomass yields were studied under conditions of surplus food supply at 22° C and 30 ‰ S. Increase of exploitation rate lowered the adult plus copepodid density and raised the density of nauplii, thus shifting the age distribution in favour of the nauplii. Total population density was almost equal at exploitation rates of 0.10, 0.50 and 0.70, but lower at rates of 0.30 and 0.90. Experiments, which have not yet been completed, have shown a higher resistance against the toxic effect of cadmium in populations exploited at rates of 0.30 or 0.90; populations exploited at the lower rate displayed a lesser degree of resistance than populations exploited at the rate of 0.90. Variability of densities was least at intermediate exploitation rates, whereas the variability of the age structure and the intrinsic rate of natural increase was less at higher rates, indicating smaller fluctuations in the ratio of the regulating variables mortality and fecundity with increasing exploitation. The assumption is made that an appropriate exploitation rate contributes to the achievement of favourable culture conditions for copepods, which are subjected to intense predation in natural populations. Within the range of the exploitation rates selected, biomass yields of nauplii and adults plus copepodids were approximately proportional to exploitation rate; therefore, the maximum rate sustainable by aTisbe holothuriae population cannot be established. From data on adult numerical yield, however, it may be concluded that exploitation rates exceeding 0.90 will not result in increased productivity.     

Zur Dynamik exploitierter Populationen vonTisbe holothuriae (Copepoda,Harpacticoida)

Zusammenfassung 1. An wöchentlich mit Raten von 0,10; 0,30; 0,50; 0,70 oder 0,90 exploitierten Populationen des harpacticoiden CopepodenTisbe holothuriae Humes wurde unter konstanten Bedingungen (22° C, 30 S) die Abhängigkeit der Reproduktionsleistung, des Geschlechtsverhältnisses, der Entwicklungsdauer und der Überlebenszeit von der Exploitationsrate untersucht.2. Es lassen sich drei Klassen von Eisäcken unterscheiden: (a) Eisäcke mit Entwicklung aller Eier bis zum schlüpfenden Nauplius, (b) Eisäcke mit Entwicklung nur eines Teils der Eier und (c) Eisäcke, aus denen keine Nauplien hervorgehen. Eine Abhängigkeit von der Entnahmerate besteht bei den Anteilen an Eisäcken, bei denen aus allen Eiern oder aus keinem Ei Nauplien hervorgehen.3. Der Anteil an Eisäcken, aus denen keine adulten Individuen hervorgehen, ist bei allen Entnahmeraten größer als der Anteil, aus dem keine Nauplien schlüpfen. Der Unterschied entsteht durch Eisäcke, bei denen unmittelbar nach dem Schlupf sämtliche Nauplien sterben.4. Die Zahl der pro 100 Eisäcke im Mittel produzierten Eier, Nauplien und adulten Individuen ist abhängig von dem Mittel der mittleren Populationsdichten zwischen den Entnahmen.5. Bei allen Befunden wird die durch die Exploitationsrate beeinflußte mittlere Populationsdichte zwischen den Entnahmen als die unmittelbarer wirkende Einflußgröße angesehen.6. Bei höherer Entnahmerate beziehungsweise geringer mittlerer Populationsdichte zwischen den Entnahmen werden im Mittel pro Eisack mehr Eier, Nauplien und adulte Individuen produziert als bei geringer Entnahmerate beziehungsweise höherer mittlerer Populationsdichte zwischen den Entnahmen. Es scheint die Tendenz zu bestehen, unabhängig von der Entnahmerate einen durch die Kulturbedingungen als Führungsgröße vorgegebenen Sollwert der Biomasse zu erreichen und konstant zu halten.7. Die Sterberate der Nauplien ist bei der Entnahmerate von 0,90 herabgesetzt. Das Reproduktionspotential wird allerdings selbst bei dieser Entnahmerate nur etwa zur Hälfte ausgenutzt.8. Außer bei der Entnahmerate von 0,90 werden im Mittel pro Eisack mehr Embryonen gefunden als Nauplien schlüpfen. Die relative Häufigkeit der Eier und Eisäcke, in denen keine Embryonen gefunden werden, zeigt keine Anhängigkeit von der Entnahmerate beziehungsweise der mittleren Populationsdichte zwischen den Entnahmen.9. Bei isolierten zeigt sich unmittelbar nach der Isolierung eine Abhängigkeit der Frequenz der Eisackbildung von der Entnahmerate beziehungsweise der mittleren Populationsdichte zwischen den Entnahmen. Es scheint, als erführen die Individuen durch die Populationen, aus denen sie stammen, eine Prägung, die in der Isolation allerdings bald verlorengeht.10. Das Geschlechtsverhältnis ist abhängig von der Entnahmerate beziehungsweise der mittleren Populationsdichte zwischen den Entnahmen. Bei der Entnahmerate von 0,90 überwiegt die Zahl der , bei der Entnahmerate von 0,10 die der .11. Bei isolierten Brutsätzen konnte eine Abhängigkeit der embryonalen Entwicklungsdauer von der Entnahmerate beziehungsweise der mittleren Populationsdichte zwischen den Entnahmen beobachtet werden. Eine Beeinflussung der Überlebenszeit adulter sowie der Geschwindigkeit der Entwicklung vom geschlüpften Nauplius bis zum adulten Individuum konnte bei den isolierten Brutsätzen nicht festgestellt werden.12. Das mittlere Gewicht eines Individuums im adulten oder Copepoditstadium und die mittlere Länge des adulten stehen in inverser Beziehung zur mittleren Populationsdichte zwischen den Entnahmen.

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On the dynamics of exploited populations ofTisbe holothuriae (Copepoda, Harpacticoida)III. Reproduction, sex ratio, rate of development and survival time

In weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes, effects of exploitation rate on production of eggs, embryos, nauplii and adults, as well as frequency of egg-sac formation, rate of development of embryos and nauplii and survival time of adult females were studied under conditions of surplus food supply at 22° and 30 S. Three classes of egg sacs could be distinguished: (a) Egg sacs in which all eggs developed until hatching of the nauplii, (b) egg sacs in which only part of the eggs developed, and (c) egg sacs with eggs that did not hatch at all. The fraction of egg sacs giving rise to no adult individual is larger at all exploitation rates than the fraction of egg sacs giving no nauplius. This difference is caused by the fact that some egg sacs produce nauplii, all of which die shortly after hatching. The mean number of eggs, nauplii and adults produced per egg sac depends on the mean average population densities between exploitations. In general, the mean average population densities between exploitations are considered to be a more immediate causal agent than the removal rate. At a higher removal rate or a lower density between exploitations, more eggs, nauplii and adults are produced per egg sac than at a lower removal rate or at a higher density. There seems to be a tendency to reach and to keep constant a set point of biomass, rather than population size, the culture conditions being the controlling variable. Death rate of nauplii is lowered at the exploitation rate of 0.90, and even at this highest removal rate only half of the eggs gave rise to adult individuals. Except at the exploitation rate of 0.90, more embryos per egg sac are found on the average than hatched nauplii, implying death of a certain number of embryos during development. The relative frequency of eggs and egg sacs in which no embryos could be detected shows no dependency upon the removal rate or mean population density between exploitations. In isolated females, an inverse relationship has been found between the frequency of egg-sac formation and the average population density between exploitations shortly after isolation. There seems to be some imprinting by the populations in which the specimens are living. Sex ratio is influenced by the removal rate or the average population density between exploitations in such a way that at the removal rate of 0.90 there is a surplus of males while at the rate of 0.10 a surplus of females is found. In isolated broods an inverse relationship between embryonic developmental rate and average population density between exploitation could be established. No influence of population density on rate of development of juvenile stages and survival time of adult females could be observed in equally isolated broods. The average weight of an individual in the adult or copepodid stage, and the average length of an adult female are inversely related to the average population density between exploitations.

     

Zur Dynamik exploitierter Populationen vonTisbe holothuriae (Copepoda,Harpacticoida)

In weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes, the dynamics of population development and the effect of the rate of water renewal were studied under conditions of surplus food supply, constant temperature (22° C) and salinity (30 ‰). At exploitation rates of 10, 30, 50, 70 or 90% per week, the populations maintained stable mean populations sizes. None of the populations established stationary equilibria. Reduced rates of water renewal resulted in a reduction of mean population density, in the course of which the density of adults plus copepodids was diminished to a lesser degree when low, as opposed to high, percentages of specimens were removed. The density of nauplii was distinctly reduced at all exploitation levels. The age structure shifted in favour of the adults and copepodids and was subject to higher variability at moderate exploitation rates and lowered rate of water renewal. To avoid and to remove difficulties inherent to current concepts of population dynamics, the populations were regarded as a component of a feedback control system, and an attempt has been made to describe the observed processes by analogy to technical control systems. According to not yet completed series of experiments, populations undergoing higher exploitation rates seem to compensate more succesfully for perturbations of their environment, than populations exploited at lower rates.     

Effects of seven diets on the population dynamics of laboratory culturedTisbe holothuriae Humes (Copepoda,Harpacticoida)

The harpacticoid copepodTisbe holothuriae was collected from Saronicos Gulf (Greece) and reared under constant laboratory conditions. In order to study the effects of food on the population dynamics, seven diets were tested: the seaweedUlva; five artificial compound feeds: the liquid Fryfood^® (Waterlife), a powder ofMytilus, yeast, soya andSpirulina, respectively; and a mixed diet consisting ofUlva and Fryfood. The life cycle parameters (mortality, sex ratio, generation time, offspring production) were measured, and the demographic variables [mean generation time (T), net reproductive rate (Ro), and intrinsic rate of natural increase (r_m)] were determined. As to their efficiency regarding population dynamics, the diets ranked as follows: (1)Ulva+Fryfood, (2),Ulva, (3) Fryfood, (4)Mytilus, (5) soya, (6) yeast, and (7)Spirulina. In this order they cause a progressive increase of both larval mortality and generation time, a progressive decrease of sex ratio, number of offspring per egg sac, number of egg sacs per female and, consequently, of Ro and r_m. The observed differences between diets were most pronounced with respect to offspring production. Of the compound diets, those containing animal extracts were more efficient than those containing vegetable materials.Ulva plays an important role in the nutrition ofT. holothuriae, favouring offspring production as well as larval survival, development and pigmentation.Ulva in combination with Fryfood led to a greater copepodid survival and offspring production. This mixed diet proved to be the most favourable for rearing the Greek population oft. holothuriae, resulting in an efficient intrinsic rate of natural increase (r_m=0.304) of the population.     

Dry weight and chemical composition (CHN) in relation to population density of cultivatedTisbe holothuriae (Copepoda,Harpacticoida)

The harpacticoid copepodTisbe holothuriae was cultivated at different densities in a running-water system. Dry weight and chemical composition (CHN) of both females and egg-sacs have been determined. Dry weight decreased significantly with increasing density from 3,48 to 1.94μg in egg-sacs, and from 10.80 to 6.58μg in females. Increasing density in both egg-sacs and females results in decrease of carbon, nitrogen and hydrogen contents, expressed as a percentage of dry weight of egg-sacs and females, respectively. Carbon decreases from 52.56 to 50.57% (egg-sacs) and 45.80 to 39.95% (⧫), nitrogen from 12.63 to 11.94% and 10.72 to 9.73% and hydrogen from 9.08 to 7.78% and 7.47 to 6.17%. The dry weight and elemental compositions of the egg-sacs varied in accordance with that of the females, however, a higher percentage of elemental content was observed in egg-sacs. The energy equivalents were calculated from the carbon content. This indicates that more energy was transferred from the maternal body into egg-sacs. The ratio of carbon to nitrogen did not show any marked variations; no clear relation appeared between C:N and density, indicating a relatively constant chemical composition in both females and egg-sacs.     

Effect of crowding on nauplii production during mating time inTisbe clodiensis andT. holothuriae (Copepoda,Harpacticoida)

Tisbe clodiensis andT. holothuriae females and males were maintained at different densities, from sexual maturity to the extrusion of the first egg sac, in order to study the effect of crowding on the number of nauplii produced by the first egg sac. They were tested in pure cultures at densities of 1, 2, 4, 6 and 8 couples per 20 ml of sea water, and in mixed cultures (half of the individuals belonging to each of the two species) with 2, 4, 6 and 8 couples per 20 ml, without food limitation. At the highest crowding level, and forT. holothuriae only, the effect of daily renewal of the culture water was examined. The mean number of nauplii per female decreases with increased density. A significant linear regression of number of nauplii on number of couples was found. The regression slopes appear to be the same for both species, regardless of whether they are cultured singly or together. The daily renewal of water diminishes the effect of crowding. The possibility that a complex chemical compound, produced by the animals, which enables the latter to perceive and to react to crowding, is discussed.     

Effects of copper sulphate on Tisbe holothuriae Humes (Copepoda) and development of tolerance to copper

A study was made of the toxicity of copper to the marine copepod Tisbe holothuriae Humes and the development of increased tolerance due to acclimation. In sublethal concentrations, copper causes a prolongation of maturation time and a reduction in offspring production. These effects increased proportionally to the copper concentration used, while the respiratory rate (measured at the highest concentration used: 0.008 mg l^–1 Cu) increased. A tendency for higher tolerance due to acclimation was also observed. The copper induced delay in maturation time observed in the F₂ generation, became less pronounced from F₃ onwards. In animals exposed at 0.008 mg l^–1 Cu the maturation time of F₃ was not statistically different from that of untreated animals. The LC50 48 h also presented an increase due to acclimation in animals exposed at 0.004 (F₅) and 0.008 mg l^–1 Cu (F₃ and F₅ generations), but this increase was not statistically significant.     

Effects of light (photoperiod,spectral composition) on the population dynamics of Tisbe holothuriae Humes (Copepods,Harpacticoida)

The harpacticoid copepod Tisbe holothuriae Humes was reared for several generations under different photoperiods (white light) and spectral quality (continuous illumination). The main life cycle parameters were measured and demographic variables were determined. Development time was retarded under green wavelengths and at photoperiods different from LD 12:12, especially under continuous white light. Total body length of females and males was shortest under LD 12:12 and longest under constant dark. Photoperiods different from LD 12:12 prolonged the interval between broods of egg sacs. Short photoperiods caused higher abortion rate, while long photoperiods retarded the maturation time of the egg sacs. Red wavelengths stimulated the abortion rate. Photoperiod LD 12:12 and blue wavelengths increased the production of offspring and the life span of females. Highest intrinsic rates of natural increase were estimated under LD 12:12 (rm =0.304) and blue wavelengths (rm = 0.254). These light conditions proved the most favourable for rearing Tisbe holothuriae.     

On Gaussia Wolfenden (Copepoda,Calanoida, Metridinidae)

Aspects of the female genital segment with and without attached couplers and enclosed spermatophores are studied of Gaussia specimens collected in the SW Atlantic and SE Pacific. A new species found in the SW Atlantic, Gaussia asymmetrica, is proposed.     

On the dynamics of the hanuman langur populations of jodhpur (Rajasthan,India)

The city of Jodhpur (26°18′N, 73°8′E) supports a population of about 900 hanuman langurs (Presbytis entellus) divided into 24 bisexual troops and 12 all-male bands in an area of 60 km². This population has been censused from 1968 to 1978. Over this period the population of bisexual troops has remained stable around 700, while the population of all-male bands has increased from 160 to 230 individuals. The bisexual troops show a tendency towards a reduction in population growth rate with increasing troop size, with troops over 50–60 tending to split. Very small troops may grow by large scale immigration. Although a number of male changes and mortality through infanticide have been recorded, there is no evidence of a regular periodicity in the occurrence of initial, growth and mature phases in the life history of a bisexual troop. Unlike the bisexual troops, the all-male bands show no tendency towards a reduction in growth rate with the increase in band size, but show a continuous growth of band size over the study period. Langurs of Jodhpur rely heavily on cultivated fields for their sustenance. This cultivation has been on increase over the study period, and since the males invade cultivation more readily, they may have been able to take fuller advantage of these increasing resources and affect a population increase that has not been possible for the bisexual troops.     

On the ecology of marine cyclopoid copepods (Crustacea, Copepoda)

The goal of these studies was to improve our understanding ofhow Oithonidae and Oncaeidae can exist in nearly every partof the ocean. In particular, it was intended to obtain quantitativeinformation on reproduction rates and the longevity of adultfemales, and on feeding and growth rates of Oncaeidae. Whenfeeding at relatively high food levels, early copepodids ofOncaea mediterranea ingested     

On Neodiaptomus schmackeri (Poppe & Richard, 1892) (Crustacea,Copepoda)

Neodiaptomus schmackeri and related species are redescribed. The synonymy of this species with N. strigilipes and N. handeli is discussed.     

On the foraging and feeding ability of Oithona davisae (Crustacea,Copepoda)

Growth experiments with macrophytes revealed anomalies in the growth and development of Najas intermedia. Further analyses showed that these anomalies were caused by differences between the male and female plants. To examine the gender-related differences, the results of two field experiments and a phenological monitoring programme were analysed. The field experiments were conducted in four different lakes in Southern Germany. Over the course of 2 years, eight different sediments were used to research the interaction between the gender and the growth of N. intermedia plants. The phenological monitoring programme, on the other hand, was focused on the development of three different populations of N. intermedia in three Southern German lakes. The results of the experiments revealed significant differences in the gender-related development: Depending on the environmental conditions, male plants grew between 20 and 40% faster than female plants. Additionally, the male plants developed flowers earlier in the vegetation period and died before the female plants. As a result, male and female plants coexisted in the same lake only for a short period of time. This might be a strategy to reduce the competitive pressure between the genders and to gain an advantage over rival species.