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.     

The biogeography of three Boeckella species (Copepoda: Calanoida) in New Zealand

The New Zealand distributions of three species of Boeckella (Copepoda, Calanoida), B. triarticulata, B. dilatata and B. hamata are mapped. B. triarticulata is primarily a pond dweller but is also found in reservoirs and shallow lakes. B. dilatata is mainly found in the deeper glacial lakes and ponds in the central region of the South Island and B. hamata has a more widespread distribution in lakes and ponds in the South Island and lower half of the North Island. Differences in temperature optima, food requirements and dispersal ability among the three Boeckella species are related to vicariant events to explain their distribution in New Zealand.     

Morphological differences between subitaneous and diapause eggs of Boeckella triarticulata (Copepoda: Calanoida)

1. Many calanoid copepods produce subitaneous eggs that hatch immediately and diapause eggs that remain dormant for long periods of time. Boeckella triarticulata (Copepoda: Calanoida) produces diapause eggs that hatch asynchronously over an extended period of time and appear identical under light microscopy to those that hatch immediately.
2. We compared the internal morphology of subitaneous and diapause eggs of B. triarticulata that had been prepared for transmission electron microscopy by four methods.
3. Subitaneous eggs had a thin, single-layered shell whereas diapause eggs had a thick, three-layered shell that appeared consistent over 3 months of diapause. Boeckella triarticulata appears to enter diapause at the multicellular stage.
4. Consistent morphological differences between subitaneous and diapause eggs of B. triarticulata suggest that these two egg types also have distinct differences in physiology, and are not merely extremes of a continuum from subitaneous to diapause eggs.     

UV-B-induced damage and photoreactivation in three species of Boeckella (Copepoda, Calanoida)

Solar ultraviolet (UV) radiation poses a threat to most livingorganisms. Aquatic organisms have evolved three basic mechanismsto cope with harmful levels of radiation. Two mechanisms, avoidance(e.g. vertical migration) and photoprotection (e.g. productionof photoprotective compounds that act as filters, antioxidants,etc.), serve to minimize the dose of UV radiation that reachesthe organism's vital structures (DNA, membranes, etc.). Thethird mechanism, repair (e.g. dark repair mechanisms, such asnucleotide excision repair; or photoreactivation mechanisms,such as photoenzymatic repair), serves to repair the damagefollowing UV exposure. Here, we compare the vulnerability toUV-B radiation of three copepod species (Boeckella brevicaudata,Boeckella gibbosa, and Boeckella gracilipes) that occur in lakesthat differ in UV-B penetration and depth. Our aim was to gaininsight into the significance of each of the three mechanismsin different UV-B environments. Results from a 3-day ‘insitu’ incubation in ultra-oligotrophic Lake Toncek showedthat B.gracilipes is highly vulnerable to UV-B and UV-A radiation.In contrast, virtually no mortality was observed in B.gibbosaand B.brevicaudata during the same period. In order to discriminatethe contribution of photoprotection and photoreactivation, thethree species were subsequently exposed in the laboratory toan artificial source of UV-B radiation, both in the presenceand absence of visible radiation (recovery radiation). The photoprotectionpotential (i.e. resistance to UV-B in the absence of recoveryradiation) of B.gracilipes and B.gibbosa was lower than thatof B.brevicaudata. On the other hand, photoreactivation (higherresistance to UV-B in the presence of recovery radiation) wasobserved in B.brevicaudata and B.gibbosa, but not in B.gracilipes.To cope with damaging UV-B levels in nature, B.gracilipes dependsexclusively on the attenuation by the external media (i.e. avoidance).Although B.gibbosa tends to avoid the surface waters of lakes,it also occurs in shallow transparent pools. Most likely itsability to survive in these shallow, high UV environments isdue to its photoreactivation potential. Finally, despite itsoccurrence in highly turbid lakes, B.brevicaudata seems extremelywell suited to cope with UV-B radiation thanks to a combinationof photoreactivation and photoprotection.     

Variations in the embryonic development times and hatching success of three Boeckella species (Copepoda: Calanoida)

Embryonic development times of Boeckella triarticulata, B. dilatata and B. hamata are compared. Two species, B. triarticulata and B. hamata had widely varying embryonic development times. Variable development times often resulted in greater hatching success.     

Redescription and distribution of Boeckella gibbosa (Brehm, 1935) (Copepoda,Calanoida, Centropagidae)

Thirty-five species of the family Lecanidae are examined from Tripura state in North-Eastern India. Of these, Lecane levistyla and L. scutata are interesting cold-water forms; L. batillifer is an Australasian element; L. acanthinula and L. sinuata are Oriental endemics and L. braumi, L. lateralis and L. simonneae are palaeotropical species. The lecanid fauna also includes the pantropical L. thienemanni and L. sola while the remainder are cosmopolitan and tropicopolitan elements. This revised version was published online in August 2006 with corrections to the Cover Date.     

Boeckella antiqua n. sp. (Copepoda,Calanoida, Centropagidae) from Patagonia

Boeckella antiqua n. sp. from samples collected in an ephemeral pond on the Patagonian plateau is described and illustrated. Diagnostic features of B. antiqua are almost exclusively related to the male fifth pair of legs, females being almost indistinguishable from those of the closely related B. poppei. The two-segmented, spine-bearing left endopod of the male fifth leg suggests that it may be the basal species in the genus.     

Predation,competition, and co-occurrences of Boeckella and Calamoecia (Copepoda: Calanoida) in Western Australia

Patterns of co-occurrences of Boeckella and Calamoeciafound in Western Australia are documented. Patterns are analyzed inrelation to size of organisms and to geographical distributions oforganisms. Deviations from randomness in the number ofmulti-species assemblages suggest that biotic interactionsinfluence co-occurrences, but size relationships in severalco-occurring pairs indicate that competition for resources is nota factor influencing co-occurrence. Laboratory experimentsdemonstrate that the large Boeckella triarticulata consumesimmature stages of the small Calamoecia tasmanica butprovides a partial refuge for C. tasmanica when theplanktivorous fish Gambusia, which consumes Boeckellamore rapidly than Calamoecia, is present. These resultsindicate that co-occurrence patterns may be related to theintensity of predation by fish: in the absence of fish predation,Boeckella can prevent invasion of Calamoecia, whilemoderate predation by fish will reduce Boeckella density,consequently allowing Calamoecia to invade.     

Resource partitioning of Calamoecia lucasi Brady and Boeckella triarticulata (Thomson) (Copepoda:Calanoida) in an Australian reservoir

In Lyell Reservoir the food niches of Calamoecia lucasi and Boeckella triarticulata appear to be substantially separated by food type and/or food size. Large B. triarticulata eat a wider size range of food particles and exhibit greater dietary richness and breadth than do the small C. lucasi, even though the dietary richness and breadth for the two calanoid species fluctuate through time. The dietary overlap is also variable but is generally low, ranging from 0.329–0.752 in terms of Schoener's index.     

Dispersal ability and relative abundance of Boeckella and Calamoecia (Copepoda: Calanoida) in Australian and New Zealand waters

Two calanoid copepod genera, Boeckella and Calamoecia have broadly similar geographic distributions in Australia but Boeckella is more common than Calamoecia. A field survey and literature search indicate that Boeckella is usually the first calanoid to colonize newly formed bodies of water, and Boeckella is more common in temporary pools and in high altitude environments. Boeckella usually produces larger clutches than Calamoecia, increasing its probability of establishment in lakes and ponds; calculations indicate that Boeckella's probability of establishment is far greater than that of Calamoecia. That Calamoecia is common on the North Island of New Zealand may be due to the presence of planktivorous fish; small species are more likely to survive in such environments. Calamoecia clutches may be able to move more often and farther than those of Boeckella but Calamoecia remains rare because its probability of establishment is low. Boeckella may be able to survive temporary pool environments not only because it can produce larger clutches, but also because many species in this genus are highly pigmented, and as such can survive brightly lighted environments. Patterns of abundance of North American diaptomids appear to be consistent with factors affecting those of Australian and New Zealand centropagids.     

Feeding of Boeckella gracilipes (Copepoda, Calanoida) on ciliates and phytoflagellates in an ultraoligotrophic Andean lake

The calanoid copepod, Boeckella gracilipes, is the dominantcrustacean zooplankton in South Andean deep ultra-oligotrophiclakes. Combining field and experimental data we explored thefeeding of the copepod and its access to the mixotrophic ciliate,Ophrydium naumanni, in Lake Moreno Oeste (Patagonia, Argentina).Phytoplankton was dominated by nanoflagellates throughout thewater column. Ophrydium naumanni, which accumulates much ofthe chlorophyll a, as do copepodites and adults of B. gracilipes,has a deep distribution during the day, with maximal abundancesaround 30 m depth. Mouth-part morphology analysis of B. gracilipesindicated that the copepod has an omnivorous diet. Laboratoryexperiments showed that B. gracilipes could access O. naumannionly when it is offered as a single food item. However, whennatural phytoplankton and ciliate assemblages (including O.naumanni)are offered, B. gracilipes did not eat Ophrydium and preyedon the oligotrich, Strombidium viride, and phytoflagellateslike Chrysochromulina parva. The range of ingested sizes wasbroad (3.9–33 µm of equivalent spherical diameter)but all selected particles were motile ones with distinctivemovements, which would enhance the copepod particle detection.     

Effects of Salinity and Temperature on Survival and Reproduction of Boeckella hamata (Copepoda: Calanoida) from a Periodically Brackish Lake

The freshwater calanoid copepod, Boeckella hamata Brehm, occursperiodically in Lake Waihola, New Zealand, a shallow, tidally-influencedlake that experiences daily and seasonal fluctuations in salinity,and seasonal changes in temperature. To determine whether theseenvironmental changes might contribute to the copepod's disappearancefrom the lake in summer, we tested the survival and egg productionof B. hamata at a range of salinities (20-3000 mg^–1 Cl)and two temperatures (10°C, 20°C). Adults survived for20 days or more only at salinities     

Population Dynamics of Crustacean Zooplankton in a Mesotrophic Lake,with Emphasis on Boeckella hamata BREHM (Copepoda: Calanoida)

The crustacean zooplankton in Lake Mahinerangi, a polymictic, mesotrophic reservoir, were studied for 19 months. The cladocerans, Bosmina meridionalis and Ceriodaphnia dubia, had unimodal patterns of abundance with peak densities in summer; cyclopoids had a single peak of maximum abundance in autumn and Daphnia carinata showed a bimodal pattern with peak densities in early summer and early winter. The strong seasonality in abundance of zooplankton in Lake Mahinerangi distinguishes this lake from other temperate lakes in New Zealand and may reflect its cool winter temperatures (< 5 °C for 2 months). A calanoid, Boeckella hamata, bred throughout the year and was the dominant species numerically. Instar analysis indicated that there were at least three, and possibly four, generations per year. Instantaneous birth and death rates were low with annual mean values (0.036 and 0.057, respectively) that were 2-3.5 times less than those recorded in a population of the same species in Lake Pounui, but within the range of those recorded in calanoid populations elsewhere in New Zealand. The Boeckella population in Lake Mahinerangi appears to be controlled largely by temperature, parasitism and food, with predation having no, or a negligible, effect.     

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 effects of temperature and food on naupliar development,growth and metamorphosis in three species of Boeckella (Copepoda:Calanoida)

Three species of Boeckella (B. triarticulata, B. dilatata and B. hamata) were reared from hatching to copepodite I (CI) at three naturally fluctuating food levels and three temperatures in a 3 × 3 × 3 factorial design. Development times, lengths and mortality were measured for each species in nine treatments. Temperature had the major effect on development times but food level had the major effect on CI lengths. Mortality varied interspecifically with treatment. The combined effect of temperature and food on development times and lengths was species-specific. There were trade-offs between development time and growth to meet constraints imposed by naupliar metamorphosis. The three species varied in the timing of metamorphosis to CI. B. triarticulata nauplii were age determined, B. dilatata nauplii were size determined and B. hamata nauplii were flexibly age and size determined depending on treatment. Differences in life history parameters and the timing of naupliar metamorphosis are discussed in relation to the distributions of the species in ponds (B. triarticulata), glacial lakes (B. dilatata) and coastal lakes (B. hamata) in the South Island of New Zealand.     

Survivorship of Cyclops abyssorum tatricus (Cyclopoida, Copepoda) and Boeckella gracilipes (Calanoida, Copepoda) under ambient levels of solar UVB radiation in two high-mountain lakes

We performed in situ experiments during the summer of 1995 and 1996 to assess the potential effect of solar ultraviolet B (UVB) radiation (290-320 nm) on the survival of Cyclops abyssorum tatricus Kozminski and Boeckella gracilipes Daday. These species are numerically dominant within the crustacean zooplankton living in two high-mountain lakes, one located in the Austrian Alps [Gossenkollesee (GKS), 2417 m above sea level, maximum depth 9.9 m] and another in the Chilean Andes (Laguna Negra, 2700 m above sea level, maximum depth 320 m). The copepods were incubated in quartz tubes (1 1) or in quartz tubes wrapped with Mylar D® to exclude most of the UVB radiation. The organisms were exposed at 0.5 m depth for 10-72 h on cloudless days. Both lakes were very transparent to UVB and 10% of the surface radiation at the nominal wavelength of 305 nm was still present at 9.6 m in GKS and at 12.8 in Laguna Negra. These species migrate vertically and have a maximum daytime distribution close to the bottom (C.abyssorum tatricus) or below 15 m depth (B.gracilipes). Both species were red, but the carotenoid concentration was higher in C.abyssorum tatricus than in B.gracilipes (6.5 and 2.3 g mg^-1 dry weight, respectively). UV-absorbing compounds with a maximum absorption at 334 nm were also detected. Cyclops abyssorum tatricus was highly resistant to UVB radiation and no significant lethal effect was observed. Boeckella gracilipes had a mortality 5 times higher in the treatment receiving full sunlight than in the Mylar treatment (3.2%) only when exposed for 70 h. The resistance of B.gracilipes was higher than that reported in the literature for the same species, suggesting the existence of intraspecific differences in UV sensitivity.      

Phytoplankton diet of Arctodiaptomus salinus (Copepoda, Calanoida) in Lake Shira (Khakasia)

A flow-through method was employed to study the algal part of the diet spectrum of the calanoid Arctodiaptomus salinus in the surface layer of the lake and in the phytoplankton biomass maximum zone (10 m deep). Daily consumption rates of this calanoid differed markedly in these layers, being 1 and 11 g ind ^–1 d^–1, respectively. The cyanobacterial taxa, Lyngbya contorta and Microcystis sp., accounted for more than a half of the ration. Nevertheless, the negative Ivlev electivity coefficient indicated that Arctodiaptomus prefers none of these two taxa. The low measured uptakes suggest that the energy expenditure is compensated by other resources, presumably microzooplankton and detritus.     

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.     

Zooplankton Dynamics of Lake Escondido (Rio Negro,Argentina), with Special Reference to a Population of Boeckella gracilipes (Copepoda,Calanoida)

The zooplancton dynamics and the populations of Boeckella gracilipes Daday of Lake Escondido (Rio Negro, Argentina) were studied from March 1987 to March 1988. Rotifers and Cladocerans showed high densities during spring and summer, while copepods were densest in autumn. The vertical distribution study showed that rotifers prefer upper levels of the water column, while cladocerans and copepods prefer lower levels, especially during thermal stratification. Boeckella gracilipes had a sex ratio biassed toward female, though changes in the sex ratio were observed. The clutch size was high in winter when the proportion of ovigerous females was low. The instantaneous birth rates were low during winter and high in summer. This result is related to the thermal cycle of the lake, by the influence of temperature on egg development time. The mortality rates showed a similar pattern, and maximum values were registered in spring and summer.     

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.