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Assessment page for species/group

MEASO Biota Assessment pages should be titled: MEASO Biota: Species name/group

The purpose of these assessment pages is to summarise basic information for a species or taxonomic group and how the status and ecology of the group may be changing over time (link here to the framework for assessing biota within the MEASO framework):

  • General Description
  • Autecology (species' ecology: life history, phenology, food, energetics, movement, rate processes)
  • Habitat (physical environment, biological dependencies,Relationships, thresholds and limits )
  • Population (distribution, status and trends in abundance)
  • Synecology (species interactions: food webs, competitors)
  • Assessments of status (critical state of population e.g. IUCN Red List)
  • Contributing authors and experts
  • References

Authors are encouraged to use the spatial partitioning of the marine ecosystem assessment (sectors etc - link here), as well as circumpolar information where available. Assessments of change can include, where possible, historical change, current trends and prognoses for future change. Generalities should be avoided and, instead, replaced with specific quantities of change and parameters, including error/uncertainty.  Qualitative statements are acceptable provided the reasons for the qualitative conclusions are given, along with the scope for the application of the statements, and the uncertainty surrounding them. 

In the case of Parameter Tables, authors are asked to include estimates of parameters and their error, wherever possible. Delete parameters that are not relevant.  Add parameters that are relevant but not included in the tables.  The reference/s estimating the parameters should be cited, along with any statements as to the maturity of the estimates if known (e.g. under development, personal communications, future work is examining.....).  For some species, life history information and parameters are co-opted from areas outside of the subject area, e.g. from outside the Southern Ocean.  This proxy information is reasonable to include.  Please include also the rationale for co-opting these data, and citations.

Biota pages need to be written concisely, well-referenced, and using the uncertainty language of the IPCC. In the first instance, references can be from peer-reviewed literature, reports that are publicly available or references in the grey literature that could be obtained from repositories. If the information is known but the references cannot be sourced in the first instance that put a placeholder for filling in the citation.  This is useful for translating common ideas into this assessment without worrying about the pedigree of the idea in the first instance.

Under each heading is a list of information (in red italics) that is desirable for that section. Please do not change these main headings. If needed, a section can be further subdivided. Please delete the red italics and the instructions window from the page when it is completed (when the page is in edit mode you can select it for deletion).

On photographs, figures and tables:

If these are taken from publications or libraries that require recognition of copyright then we need to secure the necessary permissions to include them on these pages.  Put a placeholder with a reference to the material and then seek permission for use.  Or contact us and we will undertake this process.

All materials taken from libraries or references need to be cited appropriately, including photographs and text from web sites.  See citation instructions (link here)

Other finalised MEASO Biota pages provide good examples of how to populate these assessments.

MEASO Assessment pages should be titled: MEASO Biota: Species name/group

The purpose of these assessment pages is to summarise basic information for a species or taxonomic group and how the status and ecology of the group may be changing over time (link here to the framework for assessing biota within the MEASO framework). Authors are encouraged to use the partitioning of the marine ecosystem assessment (sectors etc - link here) as well as circumpolar information. Assessments of change can include historical change, current trends and/or prognoses for future change. Ideally, assessments will be quantitative but can include qualitative statements as well. Estimates of error and points of critical uncertainty need to be identified where possible.

Final assessments need to be written concisely, well-referenced, and using the uncertainty language of the IPCC. In the first instance, references can be from peer-reviewed literature, reports that are publicly available or references in the grey literature that could be obtained from repositories.

Under each heading is a list of information (in red italics) that is desirable for that section. Please do not change these main headings. If needed, a section can be further subdivided. Please delete the red italics and this window from the page when you have completed it (when the page is in edit mode you can select it for deletion).

Other finalised MEASO Biota pages provide good examples of how to populate these assessments.


UNDER DEVELOPMENT

Description

Microprotozooplankton ciliates are a monophyletic group of protozoans, or unicellular eukaryotes, characterised by the presence of hair-like organelles called cilia or cirri (compound ciliary organelles) used for locomotion and feeding. Ciliates also possess two types of nuclei, including at least one somatic macronucleus and one germline micronucleus. The DNA in the micronucleus is transferred during conjugation (sexual reproduction). Species are characterised by body shape, colour, and size with the majority of species occupying microplankton sizes 20-200 μm. Ciliates consume prey ranging from bacteria, microalgae (eg. diatoms), other protists, metazoans, dissolved organic matter and other organic particles. Several ciliates, predominantly oligotrichs, are mixotrophic and can account for 40-60% of the total alorica ciliate biomass in Southern Ocean regions (Christaki et al. 2008). Ciliates are prey to copepods, pteropods, and other higher trophic organisms.

Ciliates are divided into two groups: aloricate (including oligotrichs) and loricate (including choreotrichs), referring to the absence and presence of an outer shell known as a lorica. In the Southern Ocean, loricate and aloricate ciliate densities often dominate microzooplankton biomass (Strzepek at al. 2005), and can account for 54-64% of total zooplankton composition (Sushin et al. 1986). Based on reports published between 1900 and 2011, 192 species are recorded to exist south of 40°S, however most reports come from data sets originating from summer with very little reference to mesh sizes of nets used and/or water volumes sampled (Dolan and Pierce, 2014). Highest abundances of tintinnids occur at the ice edge during austral summer (Alder and Boltovskoy 1991). When associated with Phaeocystis blooms, ciliate densities have been recorded up to 150,000 ind. m-3 (Davidson and Marchant, 1992).    

Microzooplankton, comprised of both ciliates and heterotrophic flagellates, make up one of three biogenic Fe pools in the Southern Ocean as recognised by field experiments measuring the ability of microzooplankton to regenerate Fe from prey (Hutchins et al. 1993; Bowie et al. 2001). Other biogenic Fe pools include primary producers and mesozooplankton (Bowie et al. 2001). Microzooplankton serve the functional role of predators tightly coupled with prey, comprised of both heterotrophs and autotrophs, in the microbrial foodweb (Landry et al. 1993). Chase and Price (1997) showed that microzooplankton have significant Fe demands, and laboratory studies have shown the importance of their contribution to what has been referred to as the "Ferrous Wheel" for their participation in the high demand and recycling of Fe within the microbial foodweb (Kirchman 1996; Barbeau et al. 1996; Hutchins and Bruland 1994).

Choreotrich ciliates, including tintinnids, are surface water planktonic protists found predominantly in coastal and pelagic ecosystems, and cosmopolitan, as part of temperate, tropical and polar fauna (Pierce and Turner, 1993). Globally there are over 700 species that are identified by the morphological characteristics of their lorica, or shell, into which the ciliate cell can withdraw, which can be tubular or vase-like in shape (Kofoid and Campbell 1929, 1930).

The composition of tintinnid loricae predominantly comprise of proteins, however the relative amounts of all contents (eg. proteins, carbohydrates, lipids, etc) is unknown. The influence of dead loricae on benthic communities and nutrient cycles upon reaching the deep sea is also unknown (Agatha and Simon 2012).

Oligotrich ciliates have prominent oral cilia. Like choreotrich ciliates, oligotrichs form a large proportion of the protozoan biomass, and are important grazers of nanoplankton populations (Paranjape 1987; 1990) and bacteria (Sherr et al. 1989).

Microprotozooplankton can contribute significantly (<7 to <75 %) to total micro- and nanoplankton carbon (Garrison 1991).

 


Taxonomy 

Eukaryota, Diaphoretickes, SAR, Alveolata, Ciliophora, Spirotrichea  (Domain, "unranked", "unranked", Infrakingdom, Phylum, Infraphylum)

Subclass: Choreotrichia:

Order: Tintinnida, Kofoid & Campbell, 1929, 1939

There are 192 tintinnid species recorded between 40°S and 78°S have been parsed into 2 main groups: "Southern Ocean species", species endemic below 40°S, and "Widespread species", species reaching beyond the Southern Ocean extent (Dolan et al. 2012).

The following is a list of tintinnid species endemic to the Southern Ocean (Dolan and Pierce, 2014):




Fig. 1 Examples of tintinnid species typical of the Southern Ocean:  (a) Salpingella laackmanni, (b) Salpingella decurtata, (c) Salpingella faurei, (d) Laackmanniella naviculaefera, (e) Laackmanniella forma prolongata, (f) Amphorellopsis quinquelata, (g) Amphorides laackmanni, (h) Acanthostomella obtusa, (i) Codonellopsis pusilla, (j) Epiplocylcoides reticulata, (k) Codonellopsis gaussi, (l) Codonellopsis gaussi, (m) Codonellopsis gaussi forma globosa, (n) Codonellopsis gaussi forma cylindricoconica, (o) Condonellopsis gaussi forma coxiella, (p) Cymatocylis affinis/convallaria, (q) Cymatocylis affinis/convallaria forma calcyformis, (r) Cymatocylis affinis/convallaria forma subrotundata, (s) Cymatocylis affinis/convallaria forma drygalski, (t) Cymatocylis affinis/ convallaria forma cylindrica. Species found only the Southern Ocean are Laackmanniella naviculaefera (d–e), Amphorellopsis quinquelata (f), Codonellpsis gaussi (k–o) and Cymatocylis affinis/convallaria (p–t). Note the different morphologies shown by the Southern Ocean endemics.


Summary role in ecosystem


In the microbial foodweb there is a tight coupling between prey, comprised of autotrophs and heterotrophs, and microzooplankton predators (Landry et al. 1993). This tight coupling is likely the reason for the constant supply of algal stocks within iron-limited oceanic zones (Strom et al. 2000), such as the HNLC (high nutrient low chlorophyll) waters of the Southern Ocean.

As a functional group, tintinnid ciliates are a member of the microzooplankton (20-200 μm), and can often dominate grazing of small phytoplankton (2-20 μm) among most pelagic ecosystems (Dolan et al. 2012; Karayanni et al. 2005). Tintinnids have also been reported to feed on cyanobacteria (Karayanni et al. 2005). Tintinnids are prey for Southern Ocean pelagic organisms including copepods, krill, mysid shrimp, salps, chaetognaths, larval Antarctic silverfish, and benthic organisms such as octocorals and deep sea isopods (reviewed by Dolan et al. 2012; Brökeland et al. 2010; Buck et al. 1992; Hopkins 1987; Kellermann 1987; Kruse et al. 2009; Lonsdale et al. 2000; Mauchline 1980; Orejas et al. 2003).

 

Autecology


Conjugation is a distinctive feature of the life cycle of ciliates which gamonts (conjugating partners/cells) partially and temporarily fuse, known as gamontogamy, and a bridge forms between their cytoplasms. Micronuclei undergo meiosis, macronuclei disappears, and haploid micronuclei are then exchanged over the bridge. In most ciliates, gamonts separate after conjugation and new macronuclei emerge from micronuclei in each gamont. Fission follows both conjugation and autogamy.

Critical stages of ciliate conjugation:

  • Temporary and partial fusion occurs between two compatible gamonts
  • Micronuclei undergo meiosis, forming four haploid micronuclei per gamont
  • One haploid micronucleus undergoes mitosis; the three remaining micronuclei disintegrate
  • The fused gamonts exchange a micronucleus via cytoplasmic bridge
  • Gamonts separate
  • Syngamy in each gamont occurs where micronuclei fuse to form a new diploid micronucleus (synkaryon)
  • Mitosis occurs another three times, producing eight micronuclei
  • Four of the newly formed micronuclei become macronuclei, leaving the original macronucleus to disintegrate
  • Four identical daughter cells are formed through two rounds of binary fission

Life history 

Parameters

Values

Notes
References

Maximum age




Average adult size

Size fractions range 20 to >60 μm

Klaas (1997) classify size classes as large (>60 μm), medium (40-60 μm) and small (20-40 μm)Klaas, 1997

Age at maturity




Size at maturity




Spawning/breeding area




Spawning/breeding season




Larval/gestation period




Location of recruits




Size of recruits




Overall natural mortality rate


Non-predation natural mortality rates





Migration, movement 

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Diet (foraging and consumption)

In comparison to protists such as oligotrich ciliates and heterotrophic dinoflagellates, Southern Ocean ciliated tintinnids are generally a minor component of the microzooplankton, but can at times be considered major grazers on small phytoplankton (2-20 μm) but only when surface waters are not dominated by larger primary producers, such as diatoms and/or Phaeocystis (Caron et al. 2000; Froneman, 2004; reviewed by Dolan et al. 2012). 

The gape size of tintinnids is defined as the diameter of the mouth-end of its lorica, and the lorica oral diameter (LOD) refers to the size of food items ingested (Dolan et al. 2012). Most commonly, the largest size of prey ingested is approximately half the LOD at its longest dimension (Heinbokel 1978); however the most efficient feeding occurs when prey are 25% of LOD (Dolan 2010). They are known to consume pico-, nano-, and micro-phytoplankton (Montagnes, 2013).

There is a considerable difference in the size spectrum of LODs between Southern Ocean species and Widespread species (Fig 2) which is believed to be related to the size spectrum of food items (mostly phytoplankton) being exploited (Dolan et al. 2009; Dolan and Pierce 2014). 

Fig 2 Among the Southern Ocean tintinnids, the assemblage of endemic species is distinct from that of the widespread species in terms of typical oral diameters, presumably reflecting exploitation of different sizes of prey items. The larger-mouthed endemic species are found mostly within the area bordered by the Polar Front (Dolan et al. 2012). Image from Dolan and Pierce, 2014).


Tintinnids feed by generating micro-currents by propelling their cilia at their anterior end, which enables the capturing of prey while also produces forward movement (Montagnes, 2013).

Among microprotozooplankton tested south of 50°S, estimates of grazing pressure based on a specific growth rate of 0.1 day-1 indicate heavier grazing pressure by protozoa >40 μm (larger sized); however grazing pressure based on average clearance rate of 1 μL ind.-1h-1 indicate heavier grazing pressure from smaller (20-40 μm) size classes (Klaas, 1997).

Energetics 

Parameters


Ingestion rate

Estimated grazing of primary production by microprotozooplankton (Klaas, 1997):

  • southern ACC = 11-54 %
  • Polar Frontal zone = 4-56 %

Clearance rate estimates for dinoflagellates and ciliates vary from 0.1 to >20 μL ind.-1h-1 (Capriulo et al. 1991)

Under controlled conditions, values range 0.1 to 26 μl ind.-1h-1 for oligotrichous ciliates (Jonsson 1986)

Clearance rates on phytoplankton by Antarctic protozoa, measured by radioactive isotopes: 0.0-0.2 μL ind.-1h-1 (Lessard and Rivkin, 1986)

Average clearance rate of microprotozooplankton = 1 μL ind.-1h-1 (Garrison and Buck, 1989)

Metabolism

Fecundity


Length-weight relationships


Growth rate

Uniform 0.1 day-1 (Klaas 1997)

Size at age


Population Productivity (average life time)



 

Habitat 



Ciliate protists are found in a range of Southern Ocean and Antarctic habitats, including marine and freshwater (Petz et al. 2007) pelagic environments and brine channels within sea-ice.

Species of tintinnid ciliates of the Southern Ocean have been found associated with particular habitats (Dolan and Pierce, 2014), including polynyas that are predominantly occupied by endemic species Cymatocylis and Laackmanniella (Dolan et al. 2013).

Relationships, thresholds and limits

Present the relationships between the group and physical and biological environmental attributes.  These may be qualitative and/or quantitative relationships, including descriptions/estimates of uncertainties.  Justification will be needed for the qualitative relationships.  Critical thresholds/ranges/non-linear relationships of the group with habitat variables are important to identify if present, including how they will affect the ecology of the species/group.  Express uncertainties in these relationships as confidence intervals, and/or descriptive uncertainties (such as adopted proxies from measurements elsewhere)

Functional responses of taxon (y-axis) to habitat variables (x-axis) are described here (with citations to the evidence).  Parameters for a response type are to be given with their attendant uncertainties/errors/range, with references.

 

Variable

Taxon size/stage impacted

Functional response (icon)

Parameters and uncertainties

Risk areas/regions impacted

Notes



Paste icon here (choose from Functional relationships)
















 


Population



Range and Structure

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In the Atlantic sector, ciliate biomass has been shown to be greater in the southern ACC than around the Polar Front at sampling stations along the 6°S west meridian (Klaas, 1997), and microprotozooplankton biomass along this transect constitute approximately 30 % of total protozoan biomass (Becquevort, 1997). Abundance of larger size fractions (>60 μm) of ciliates decreases along a gradient from southern ACC towards the PF, then disappear north of 49°S (Klaas, 1997).


Placeholder for map

Fig. 3: Tintinnida maps from the Biogeographic Atlas of the Southern Ocean. Locations of all Southern Ocean sites from which tintinnids have been reported (Dolan & Pierce, 2014).

Dynamics

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Tintinnids sampled from the Palmer Antarctica Long Term Ecological Research Area in January 2010 and 2011 measured average abundances of approximately 30-40 ind. m-3 in the upper 100 m of the water column and representing 5-10% of total microzooplankton biomass (Garzio and Steinberg 2013). 


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Synecology



Consumers (predators) 

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Competitors

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Other interactions e.g. disease

Tintinnids have been observed agglutinating (ie. adhering) mineral particles and/or dead biogenic matter (eg. diatom frustules) to their loricae, however reasons for this behaviour remain unknown. Many believe that the type of particles agglomerated to loricae is randomly selected and thought to reflect the predominant concentrations of such particles found within the adjacent environment (Winter et al. 1986; Henjes and Assmy 2008). Tintinnids possessing loricae without particles attached are considered "hyaline" whereas those with particles attached are "agglomerated" (Agatha et al. 2013). Armbrecht et al. (2017) reported the first observations of East Antarctic tintinnids Laackmanniella naviculaefera and Codonellopsis gaussi agglutinating live diatoms Fragilariopsis curta, F. cylindrus, F. pseudonana and F. rhombica to their loricae. Species such as Laackmanniella are believed to suck out and ingest the protoplasts of diatoms before agglomerating the empty frustules to their loricae (Gowing and Garrison, 1992).

Human Impacts


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Assessments of Status


IUCN Red List

Add the following table or simply say 'None of these species has been assessed for the Red List'

Year of classification:'None of these species has been assessed for the Red List'
Red List Category & Criteria:
Assessment Justification: include description and link to the web site


Other

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References



A list of references referred to on this page.

Please use Ecology style, for more information and examples see: 

https://besjournals.onlinelibrary.wiley.com/hub/journal/13652745/author-guidelines

Agatha, S. & Simon, P. (2012). On the nature of tintinnid loricae (Ciliophora: Spirotricha: Tintinnina): a histochemical, enzymatic, EDX, and high-resolution TEM study. Acta Protozoology, 51, 1-19.

Agatha, S., Laval-Peuto, M. & Simon, P. (2013). The tintinnid lorica. In J.R. Dolan, D.J.S. Montagnes, S. Agatha, D.W. Coats & D.K. Stoecker (Eds.) The Biology and Ecology of Tintinnid Ciliates: Models for Marine Plankton (pp. 17-41). West Sussex, UK: John Wiley & Sons Ltd. 

Alder, V. & Boltovskoy, D. (1991). Microplanktonic distributional patterns west of the Antarctic Peninsula, with special emphasis on the Tintinnids. Polar Biology, 11, 103-112. doi: 10.1007/BF00234272

Armbrecht, L., Eriksen, R., Leventer, A. & Armand, L.K. (2017). First observations of living sea-ice diatom agglomeration to tintinnid loricae in East Antarctica. Journal of Plankton Research, 39(5), 795-802. doi: 10.1093/plankt/fbx036

Barbeau, K., Moffett, J.W., Caron, D.A., Croot, P.L. & Erdner, D.L. (1996). Role of protozoan grazing in relieving iron limitation of phytoplankton. Nature, 380, 61-64. doi: 10.1038/380061a0

Becquevort, S.(1997). Nanoprotozooplankton of the Atlantic sector of the Southern Ocean in early spring: biomass and feeding activities. Deep-Sea Research II, 44, 355-373. doi: 10.1016/S0967-0645(96)00076-8

Bowie, A.R., Maldonado, M.T., Frew, R.D., Croot, P.L., Achtergerg, E.P., Mantoura, R.F.C. & Boyd, P.W. (2001). The fate of added iron during a mesoscale fertilisation experiment in the Southern Ocean. Deep-Sea Research II, 48, 2703-2743. doi: 10.1016/S0967-0645(01)00015-7

Brökeland, W., Guomundsson, G. & Svavarsson, J. (2010). Diet of four species of deep-sea isopods (Crustacea: Malacostraca: Peracarida) in the South Atlantic and the Southern Ocean. Marine Biology, 157, 177–187. doi: 10.1007/s00227-009-1308-9

Buck, K.R., Garrison, D.L. & Hopkins, T.L. (1992). Abundance and distribution of tintinnid ciliates in an ice edge zone during the austral autumn. Antarctic Science, 4, 3–98. doi: 10.1017/S0954102092000038

Capriulu, G.M., Sherr, E.B. & Sherr, B.F. (1991). Trophic behaviour and related community feeding activities of heterotrophic marine protists. In P.C. Reid, C.M. Turley & P.H. Burkill (Eds.) Protozoa and their role in marine processes, NATO ASI Series, Vol G25 (pp. 219-265). Belrlin, Germany: Springer. 

Caron, D.A., Denett, M.R., Lonsdale, D.J., Moran, D.M. & Shalapyonok, L. (2000). Microzooplankton herbivory in the Ross Sea, Antarctica. Deep-Sea Research II, 47, 3249-3272. doi: 10.1016/S0967-0645(00)00067-9

Davidson, A.T. & Marchant, H.J. (1992). The biology and ecology of Phaeocystis (Prymnesiophyceae). In F.E. Round & D.J. Chapman (Eds.) Progress in Phycological Research, Vol 8 (pp. 1-45). Bristol, UK: Biopress.

Dolan, J.R., Ritchie, M.E., Tunin-Ley, A. & Pizay, M.D. (2009). Dynamics of core and occasional species in the marine plankton: tintinnid ciliates in the north-west Mediterranean Sea. Journal of Biogeography, 36, 887-895. doi: 10.1111/j.1365-2699.2008.02046.x

Dolan, J.R. (2010). Morphology and ecology in tintinnid ciliates of the marine plankton: correlates of lorica dimensions. Acta Protozool, 49, 235-344. 

Dolan, J.R., Pierce, R.W., Jin Yang, E. & Young Kim, S. (2012). Southern Ocean biogeography of tintinnid ciliates of the marine plankton. Journal of Eukaryotic Microbiology, 59(6), 511-519. doi: 10.1111/j.1550-7408.2012.00646.x

Dolan, J.R. & Pierce, R.W. (2014). Tintinnid ciliates of the marine plankton, In C. De Broyer & P. Koubbi (Eds.) Biogeographic Atlas of the Southern Ocean (pp. 254-259). Cambridge, UK: Scientific Committee on Antarctic Research.

Froneman, P.W. (2004). Protozooplankton community structure and grazing impact in the eastern Atlantic sector of the Southern Ocean in austral summer 1998. Deep-Sea Research II, 51, 2633-2643. doi: 10.1016/j.dsr2.2004.09.001

Garrison, D.K. & Buck, K.R. (1989). Protozooplankton in the Weddell Sea, Antaractica: abundance and distribution in the ice-edge zone. Polar Biology, 9, 341-351. doi: 10.1007/BF00442524

Garrison, D.L. (1991). An overview of the abundance and role of protozooplankton in Antarctic waters. Journal of Marine Systems, 2, 317-331. doi: 10.1016/0924-7963(91)90039-W

Gowing, M.M. & Garrison, D.L. (1992). Abundance and feeding ecology of larger protozooplankton in the ice edge zone of the Weddell and Scotia Seas during the austral winter. Deep-Sea Research Part A, Oceanographic Research Papers, 39, 893-919. doi: 10.1016/0198-0149(92)90128-G

Heinbokel, J.F. (1978). Studies on the functional role of tintinnids in the southern California Bight. I. Grazing and growth rates in laboratory cultures. Marine Biology, 47, 177-189. doi: 10.1007/BF00395638

Henjes, J. & Assmy, P. (2008). Particle availability controls agglutination in pelagic tintinnids in the Southern Ocean. Protist, 159, 239-250. doi: 10.1016/j.protis.2007.10.003

Hopkins, T.L. (1987). Midwater food web in McMurdo Sound, Ross Sea, Antarctica. Marine Biology, 96, 93–106. doi: 10.1007/BF00394842

Hutchins, D.A., DiTullio, G.R. & Bruland, K.W. (1993). Iron and regenerated production: Evidence for biological iron recycling in two marine environments. Limnology and Oceanography, 38, 1242-1255. doi: 10.4319/lo.1993.38.6.1242

Hutchins, D.A. & Bruland, K.W. (1994). Grazer-mediated regeneration and assimilation of Fe, Zn and Mn from planktonic prey. Marine Ecology Progress Series, 110: 259-269. doi: 10.3354/meps110259

Kirchman, D.L. (1996). Microbial ferrous wheel. Nature, 383, 303-304. doi: 10.1038/383303a0

Jonsson, P.R. (1986). Particle size selection, feeding rates and growth dynamics of marine planktonic oligotrichous ciliates (Ciliophora, Oligotrichina). Marine Ecology Progress Series, 33, 265-277. doi: 10.3354/meps033265

Karayanni, H., Christaki, U., van Wambeke, F., Denis, M. & Moutin, T. (2005). Influence of ciliated protozoa and heterotrophic nanoflagellates on the fate of primary production in the northeast Atlantic Ocean. Journal of Geophysical Research, 110, C07S15. doi: 10.1029/2004JC002602

Kellermann,  A. (1987). Food and feeding ecology of postlarval and juvenile Pleurogramma antarcticum (Pisces; Notothenioidei) in the seasonal pack ice zone off the Antarctic Peninsula. Polar Biology, 7, 307–315. doi: 10.1007/BF00443949

Klaas, C. (1997). Microprotozooplankton distribution and their potential grazing impact in the Antarctic Circumpolar Current. Deep-Sea Research II, 44(1-2), 375-393. doi: 10.1016/S0967-0645(96)00080-X

Kofoid, C.A. & Campbell, A.S. (1929). A conspectus of the marine and freshwater Ciliata belonging to the suborder Tintinnoinea, with descriptions of new species principally from the Agassiz expedition to the Eastern Tropical Pacific 1904-1905. California, USA: University of California. 

Kofoid, C.A. & Campbell, A.S. (1939). Reports on the scientific results of the expedition to the Eastern Tropical Pacific, in charge to Alexander Agassiz, by US Fish Commission Steamer "Albatross" from October 1904 to March 1905, Lieut. Commander LM Garrett, USN commanding 37. The Ciliata: The Tintinnoinea. Bulletin of the Museum of Comparitive Biology,  84, 1-473.

Kruse, S., Jansen, S., Krägefsky, S. & Bathman, U. (2009). Gut content analysis of three dominant Antarctic copepod species during an induced phytoplankton bloom EIFEX (European iron fertilization experiment). Marine Ecology, 30, 301–312. doi: 10.1111/j.1439-0485.2009.00284.x

Landry, M.R., Monger, B.C. & Selph, K.E. (1993). Time-dependency of microzooplankton grazing and phytoplankton growth in the subarctic Pacific. Progress in Oceanography, 32, 205-222. doi: 10.1016/0079-6611(93)90014-5

Lessard, E.J. & Rivkin, R.B. (1986). Nutrition of microzooplankton and macrozooplankton from McMurdo Sound. Antarctic Journal of the United States, 21, 187-188. 

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Christine WeldrickIMAS/ACE CRC






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