Abstract :
The present review addresses the reproductive strategies of marine calanoid copepods, as affected by their physiological preconditioning, and aims to enhance understanding of their adaptations to specific environmental conditions.
dge about oocyte development and internal gonad structure, especially in relation to feeding conditions, is essential for a complete understanding of the reproductive strategies of the copepods. Therefore, the foci of the review are to identify general patterns in oocyte and gonad development in calanoid copepod species from marine ecosystems worldwide and to elucidate the significance of gonad structures for reproductive strategies.
sis is similar in all copepod species. During maturation, the morphology of the oocytes changes distinctly and, according to oocyte size and appearance of ooplasm and nucleus, five oocyte developmental stages are distinguished. In contrast, the gonad structure and its changes during the spawning cycle differ considerably among species, and these differences are related to specific reproductive traits.
onad morphology types can be distinguished: the Calanus-type, found in species from all over the world with distinctly different life history traits, is apparently most common in calanoid copepods. In this gonad type, most oocyte developmental stages are present simultaneously, and usually many oocytes mature synchronously, all of which are released in one clutch. The gonad structure allows frequent spawning and large clutches, hence, high egg production rates. This may be a preconditioning for exploiting seasonally high food supply. However, the Calanus-type was also found in species producing eggs at lower rates. In the diverticula of Pseudocalanus-type gonads, only two oocyte developmental stages are present and usually fewer oocytes mature synchronously. Accordingly, the egg production rate is generally lower as compared to the Calanus-type, and apparently only this gonad-type is structurally suitable for ovigerity. Species with Pseudocalanus-type gonads are present from polar seas to the tropics, some of them being key species. The Acartia-type was scarce, found in only one species, Acartia clausi. Here all oocyte developmental stages are present, including intermediate stages, but only a few oocytes mature synchronously and are released together. High spawning frequency compensates for the small clutches, and hence egg production rate may be as high as in Calanus-type gonads. In the Aetidius-type gonad, the total number of oocytes in the diverticula is low as is the number of oocytes maturing synchronously. Less is known about the reproductive biology of species with Aetidius-type gonads; however, their distribution and feeding patterns suggest that this type is common in species inhabiting environments of low food availability.
fferences in gonad structures also lead to differences in the egg size:female size ratio, as the space available for each mature oocyte depends on the total number of oocytes. Independent from gonad-type, the eggs are relatively large in species in which the gonads contain only few oocytes, whereas small eggs are produced by species with gonads filled with many oocytes. Since all species carrying their eggs in external sacs until hatching (ovigerous species) have Pseudocalanus-type gonads, the scatter in their egg size:female size ratio is low. The broadcast spawning species are of all gonad-types, and consequently the scatter among them is high.
r factor affecting the timing and magnitude of spawning of calanoid copepods is the energy supply for gonad development. Therefore, part of the review elucidates the role of internal and external resources in fuelling egg production. In many species, freshly assimilated food is transferred into egg material within a short period of time, and clutch size and spawning frequency are the two parameters that allow adjustment of egg production to food availability and temperature. However, internal body reserves may also fuel oocyte development. The extent to which oogenesis depends on ambient feeding conditions varies considerably among species and even within species, apparently reflecting their adaptation to specific environmental conditions. All copepod species continue to release eggs for a short period after the onset of starvation, indicating that a short-term nutrient pool fuels final oocyte maturation. Some species, especially from polar and subpolar regions, are capable of reproducing on internal reserves, either solely or in addition to freshly ingested food, hence, decoupling the reproduction to some extent from the spring bloom. The reproductive strategy of Calanus finmarchicus, which responds with extreme flexibility to variations in food supply, is presented here as an example for the complexity of the reproductive processes in a calanoid copepod.
ordance with their physiological pre-conditioning, the calanoid copepod species exhibit distinctly different reproductive patterns. Polar and sub-polar environments are characterized by strong seasonality in light regime, and hence, phytoplankton concentration. The reproductive activity of many dominant (herbivorous) species peaks during the spring bloom. However, species that utilize internal reserves or food resources other than phytoplankton, reproduce before and after the bloom, or even year-round. In the North Atlantic, presented here as an example for a boreal marine ecosystem, maximum reproduction of the dominant copepod species Calanus finmarchicus occurs during the spring bloom. However, the timing of peak spawning varies between areas, as the onset of the bloom varies with hydrography and climate. The subtropical Gulf of Aqaba, in contrast to other subtropical systems, is characterized by seasonal phytoplankton succession driven by thermal winter convection. These changing environmental conditions, however, did not affect the reproduction of two of the dominant species, Clausocalanus farrani and C. furcatus. In contrast, other calanoid species exhibit reproductive cycles, either related to temperature (Ctenocalanus vanus) or to ontogenetic migration (Rhincalanus nasutus).
formation gained from studies of morphology and reproductive traits contributes to standardization of methods in reproductive studies. Based on detailed knowledge of gonad morphology and its changes during maturation and due to food supply, classification schemata have been developed allowing the identification of females ready to spawn. In a next step, egg production may be estimated from preserved females by assessing clutch size through the number of maturing oocytes in the gonads. This approach, however, is as yet applicable only to the species best studied, Calanus finmarchicus.
esent review shows that morphological studies on gonad maturation processes may provide significant contributions to fundamental ecological questions. Thus, they extend our knowledge of reproductive ecology from simply relating reproductive traits to abiotic and biotic factors toward a mechanistic understanding of how reproduction is regulated in calanoid copepods.
