The demography of Calanus finmarchicus (Copepoda: Calanoida) in the Middle Atlantic Bight, USA, 1977-2001

The spatial distribution and annual cycle of abundance for thelate-stage copepodites of the calanoid copepod Calanus finmarchicusare described for the shelf and slope waters of the Middle AtlanticBight (MAB), a subregion of the US Northeast large marine ecosystem.Samples were collected with bongo nets from 1977 to 2001 onbroad-scale surveys of the region and from a continuous planktonrecorder (CPR) towed at 10-m depth along a route that traversedthe area. The copepod’s abundance in shelf waters increasedrapidly in early spring and reached maximum levels in May–June.It declined slowly thereafter, until a minor secondary pulseoccurred in late autumn, which was followed by the annual winterlow. Slope water abundance of C. finmarchicus peaked in Apriland was virtually absent there at 10-m depth from July throughthe following February. The two samplers, both portrayed similarhigh interannual abundance variability with no long-term trendevident for the 25-year period. Cluster analysis pinpointedthree shelf areas with similar abundance patterns, one of whichwas a region of high density located in the northeastern offshorewaters. Evidence is presented that indicates the source of thishigh abundance is likely from the populations that overwinterin deep water basins of the Gulf of Maine (GOM). The copepod’sinterannual abundance variability was found to be negativelycorrelated with water temperature and unrelated to fluctuationsof the North Atlantic oscillation (NAO) index.     

High dispersal potential has maintained long-term population stability in the North Atlantic copepod Calanus finmarchicus

The cool-water copepod Calanus finmarchicus is a key species in North Atlantic marine ecosystems since it represents an important food resource for the developmental stages of several fish of major economic value. Over the last 40 years, however, data from the Continuous Plankton Recorder survey have highlighted a 70 per cent reduction in C. finmarchicus biomass, coupled with a gradual northward shift in the species's distribution, which have both been linked with climate change. To determine the potential for C. finmarchicus to track changes in habitat availability and maintain stable effective population sizes, we have assessed levels of gene flow and dispersal in current populations, as well as using a coalescent approach together with palaeodistribution modelling to elucidate the historical population demography of the species over previous changes in Earth's climate. Our findings indicate high levels of dispersal and a constant effective population size over the period 359,000-566,000 BP and suggest that C. finmarchicus possesses the capacity to track changes in available habitat, a feature that may be of crucial importance to the species's ability to cope with the current period of global climate change.     

Temperature-dependent development and growth of Calanus sinicus (Copepoda: Calanoida) in the laboratory

The calanoid copepod Calanus sinicus was reared in the laboratory under excess food conditions, and its development and growth rates were measured at various temperatures. Egg development time (D_H, days) was dependent on temperature (T °C), and was expressed as D_H = 55.3 (T + 0.7)^–1.44. Post-embryonic development followed the equiproportional rule. The stage duration was short in NI and NII, but compensatingly longer in NIII. Between NIV and CII, it was nearly isochronal, and beyond CII, it tended to increase gradually. The time from egg to adult was expressed as D_CVI = 1258 (T + 0.7)^–1.44. The specific growth rate was also temperature-dependent and highest from CI to CIII, intermediate from NII to CI and from CIII to CV, and lowest from CV to CVI. The growth rates of C. sinicus are higher than those of co-occurring small copepods such as Paracalanus, Acartia and Microsetella.     

Parasitic regulation in a population of Boeckella hamata Brehm (Copepoda: Calanoida)

SUMMARY. 1. The impact of parasitism by a fungus Aphanomyces sp. (Phycomycetes: Saprotegniaceae) on a population of Boeckella hamata in Lake Mahinerangi in New Zealand was studied over 20 months. The incidence of infection among adult female copepods exceeded 12% throughout winter (May-October) and was highest in autumn (46%) when population birth rates were depressed by 48.8%. During most of the study, parasitism lowered the rates of population recruitment and growth by more than 10%. These effects on the B. hamata population exceeded those of Aphanomyces on Boeckella dilatata Sars in another study.
2. It is hypothesized that the high incidence of parasitism by Aphanomyces and its importance at times in regulating populations of Boeckella in New Zealand lakes may be an ecological consequence of life in predator-poor ecosystems.     

Susceptibility to ultraviolet radiation in Calanus finmarchicus and Lepeophtheirus salmonis and the adaptive value of external filtering (Crustacea: Copepoda)

Copepods were exposed to UVB with a spectral irradiance maximumat 313 nm and additional visible light. In the holopelagic Calanusfinmarchicus, with a carotenoid concentration of 0.85 µgmg^-1, the LD₅₀ dose was 29 kJ m^-2 whereas in the parasitic copepodLepeophtheirus salmonis the LD₅₀ was 84 kJ m^-2. The abilityto photorepair was demonstrated for both species. The survivaldata were Weibull distributed, and this result is discussedrelative to a mechanistic ‘Weibull model’. Attenuationof UVB (306 nm) was measured in the inner Oslo fjord (K_d = 3.9m^-1). The adaptive significance of behavioural avoidance isassessed in model simulations; the results of the survival experimentsare used in connection with the Morowitz formula. Two caseswere compared; when Calanus was assumed incapable of activeexternal filtering the specimens would be safe if the depthof the mixed layer was deeper than 7 m in the inner Oslo fjord,however, with the ability of behavioural avoidance, Calanuswould be protected if they avoided the upper 1 m. The adaptivevalue associated with behavioural avoidance of toxic radiation,and the implications relative to vertical migration are discussed.     

Molecular phylogeny of the Calanoida (Crustacea: Copepoda)

The order Calanoida includes some of the most successful planktonic groups in both marine and freshwater environments. Due to the morphological complexity of the taxonomic characters in this group, subdivision and phylogenies have been complex and problematic. This study establishes a multi-gene molecular phylogeny of the calanoid copepods based upon small (18S) and large (28S) subunits of nuclear ribosomal RNA genes and mitochondrial encoded cytochrome b and cytochrome c oxidase subunit-I genes, including 29 families from 7 superfamilies of the order. This analysis is more comprehensive than earlier studies in terms of number of families, range of molecular markers, and breadth of taxonomic levels resolved. Patterns of divergence of ribosomal RNA genes are shown to be significantly heterogeneous among superfamilies, providing a likely explanation for disparate results of previous studies. The multi-gene phylogeny recovers a monophyletic Calanoida, as well as the superfamilies Augaptiloidea, Centropagoidea, Bathypontioidea, Eucalanoidea, Spinocalanoidea and Clausocalanoidea. The phylogeny largely agrees with previously-published morphological phylogenies, including e.g., enlargement of the Bathypontioidea to include the Fosshageniidae.     

The occurrence of Calanus finmarchicus (Gunnerus) and Calanus helgolandicus (Claus) in the western Irish Sea

The seasonal abundance of Calanus finmarchicus and Calanus helgolandicusin the North Channel and stratified region of the western IrishSea is reviewed using data collected between 1992 and 1996.Both species occur in the western Irish Sea, but were more abundantin the stratified region during spring. Increased abundanceduring May/June was attributed to an increase in copepoditestages. Calanus helgolandicus dominated in both regions, exhibitingspring and autumn peaks in abundance in the stratified region.It is argued that the presence of ripe females and naupliarstages in the stratified region is evidence of an in situ breedingpopulation, rather than advection of individuals from populationcentres outside the Irish Sea. The lack of geographical separationof the two species in the western Irish sea, and reports thatboth species occur in the Celtic Sea and Malin Shelf, limitthe use of either species as indicators of exchange processesbetween the Irish Sea and neighbouring waters.     

Multi-Gene Analysis Reveals a Lack of Genetic Divergence between Calanus agulhensis and C. sinicus (Copepoda; Calanoida)

The discrimination and taxonomic identification of marine species continues to pose a challenge despite the growing number of diagnostic metrics and approaches. This study examined the genetic relationship between two sibling species of the genus Calanus (Crustacea; Copepoda; Calanidae), C. agulhensis and C. sinicus, using a multi-gene analysis. DNA sequences were determined for portions of the mitochondrial cytochrome c oxidase I (mtCOI); nuclear citrate synthase (CS), and large subunit (28S) rRNA genes for specimens collected from the Sea of Japan and North East (NE) Pacific Ocean for C. sinicus and from the Benguela Current and Agulhas Bank, off South Africa, for C. agulhensis. For mtCOI, C. sinicus and C. agulhensis showed similar levels of haplotype diversity (H_d = 0.695 and 0.660, respectively) and nucleotide diversity (π = 0.003 and 0.002, respectively). Pairwise F_ST distances for mtCOI were significant only between C. agulhensis collected from the Agulhas and two C. sinicus populations: the Sea of Japan (F_ST = 0.152, p<0.01) and NE Pacific (F_ST = 0.228, p<0.005). Between the species, F_ST distances were low for both mtCOI (F_ST = 0.083, p = 0.003) and CS (F_ST = 0.050, p = 0.021). Large subunit (28S) rRNA showed no variation between the species. Our results provide evidence of the lack of genetic distinction of C. sinicus and C. agulhensis, raise questions of whether C. agulhensis warrants status as a distinct species, and indicate the clear need for more intensive and extensive ecological and genetic analysis.     

A genetic determination of cryptic species (Copepoda: Calanoida) and their postglacial biogeography in North America

Allozyme electrophoresis of Hesperodiaptomus arcticus (Marsh) sensu lato at subarctic Churchill, Canada revealed significant deviations from random mating in several populations, as shown by the complete absence of certain heterozygous types. A broad geographic survey of 20 enzyme loci in 1077 individuals from 27 ponds at 11 arctic and alpine sites west of Hudson Bay demonstrated that there were three genetically distinct types. Morphological analysis of genetically classified populations uncovered consistent morphological differences that had been ignored previously. These morphological characteristics were used to reclassify preserved museum and private specimens from additional sites from the entire North American distribution of H. arcticus sensu lato. A detailed study of two of the distinct types co-occurring at Churchill showed reproduction timing and habitat differences consistent with reproductive isolation. I concluded that there are three species and each have different distributions. Most regions examined must have been repopulated by dispersal from glacial refugia after deglaciation. The nominal species is apparently restricted to western arctic localities in the vicinity of the Alaskan refugium while the other two putative taxa have dispersed widely into previously glaciated regions.     

Daily egg production ofTemora longicornis (Copepoda,Calanoida) during winter and early spring in the Marsdiep (Southern North Sea)

Mature females of the copepodTemora longicornis from the Marsdiep tidal inlet were incubated at ambient water temperature during the winter of 1989/1990 to estimate their egg production during 24 hours. Throughout winter this production remained at a level of 0 to 10 eggs per female. In March a steep rise to 60 to 80 eggs coincided with an equally steep rise in chlorophyll-a from 4 to 20 g.l^–1. Excess food given during incubation tended to increase the observed production rate only in October at relatively high temperature. Additive effects of rising food level and temperature on daily egg production were found in spring. In March and April 1988 and 1990 the observed maximum daily egg production per female increased from about 20 at 5°C and 5 g.l^–1 chlorophyll to 70 at almost 10°C and 20 g.l^–1. The egg production in winter gives a prebloom potential to maintain a stock of premature stages ready to utilize the algal spring bloom. Early spawning before April can also contribute to the development of the abundance peak of adults in May–June. Due to growth rates increasing with temperature in the months March and April, most of the adults at their peak density may have hatched from eggs before the period of maximum egg production rate at the end of April. The exceptionally high production rate observed at the end of March may be related to a developingPhaeocystis bloom.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

The freshwater Calanoida (Crustacea: Copepoda) of Thailand

14 Species of freshwater Calanoida in Thailand are reported in this study. The most common species is Neodiaptomus botulifer. Most of these belong to the Southeast Asian species, the remainder are either Indian or East Asian copepods. The zoogeographical demarcation of copepods in Thailand is not very distinct due to its interconnecting riverine and reservoir systems as well as frequent flood. The peculiarity of the Thai Calanoida and their distribution is discussed in a Southeast Asia context. A taxonomic key to the Thai Calanoida is formulated.     

Some reflections on the structure of the mandibleplate ofEurytemora affinis (Copepoda,Calanoida)

The mandibleplates of several specimen of the copepodEurytemora affinis collected from the Westerschelde and the Gironde estuary were studied. Using the Edge index (ITOH, 1970) this species could be classified as an omnivore. Pictures obtained by light and scanning electron microscopy reveal that the mandible teeth ofE. affinis have a molar-like structure, which likely changes in functional morphology with developmental stage, sex and age of the adults. Based on these observations, the desirability to consider the three dimensional structure of copepod mandibles for feeding mode classification is discussed.Communication no. 540 of the Delta Institute for Hydrobiological Research, Vierstraat 28, 4401 EA Yerseke, The Netherlands.     

The China Coastal Current as a driving force for transporting Calanus sinicus (Copepoda: Calanoida) from its population centers to waters off Taiwan and Hong Kong during the winter northeast monsoon period

Calanus sinicus is a copepod with wide geographical distributionin the continental shelf waters of China. Its occurrence inoceans around Taiwan and Hong Kong is limited by temperatureto winter and spring. We present evidence to show that C. sinicusis carried into the coastal waters of Taiwan and Hong Kong frompopulation centers in the Yellow Sea and the East China Seaby the China Coastal Current during the northeast monsoon periodin winter.     

The Malaysian Tropodiaptomus (Copepoda : Calanoida) and its distribution

In Peninsular Malaysia there are three species of Tropodiaptomus namely: T. hebereri Kiefer, T. ruttneri Brehm, and T. vicinus Kiefer. The last species is most common in swampy habitats along the west coast whereas T. ruttneri occurs only in the north and T. hebereri is present in the south. The affinity of the Malaysian tropodiaptomids and their Sunda counterparts is discussed.     

Daily changes in population structure and production of Eudiapomus gracilis (G.O. Sars) (Copepoda, Calanoida) during summer in a shallow lake (Balaton, Hungary)

Population structure and production of Eudiaptomus gracilis(G.O. Sars) have been calculated from daily measurements inLake Balaton for one month during each summer in 1975 and 1977.Fecundity was different in the two years 6.98 ? 1.16 and 10.68? 1.42 in 1975 and 1977, respectively. A total of 80% and 64%of the population were made up of larvae in 1975 and 1977, respectively.The ratio of males to females was 1:1.63 and 1:1.1, respectively.In 1975 it was possible to follow the development of cohorts,19 –20 days in the former year, 17 –20 days in thelatter. In both years of examination 80 – 83% of naupliiand 12–15% of copepodites were eliminated. Daily net biomassproduction was 6.88 mg (dry wt.)/m³ in 1975 and 1.94 mg (drywt.)/m³ in 1977. Ratios of eggs, nauplii and copepodites indaily production was 1:2.4:1.4 in 1975 and 1:1.0:1.0 in 1977.The daily potential biomass production were 34.40 mg (dry wt.)/m³ and 10.28 mg (dry wt.)/m³ and the elimination were 80 and81 wt.%, respectively. P/B ratio — allowing for net productionvalues — was nearly the same in both periods of investigation(0.12 and 0.099 respectively).     

Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.