Dave Mackas 1 and Jackie King 2
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1 Classification of Zooplankton Life History Strategies Ctenophores Medusae Pteropod s 0.2 Free-spawning Copepods Reproduction PC Euphausiid Chaetognaths Salp/Doliolid -0.6 Sacspawning Copepods Cladoceran Amphipods Mysid Reproduction P C1 Dave Mackas 1 and Jackie King 2 Fisheries & Oceans Canada 1 Institute of Ocean Sciences & 2 Pacific Biological Station
2 Outline of Talk: Motivation for the analysis LHS theory - historical summary Ordination of zooplankton LHS traits Analysis methods Choice of traits Results & Interpretations Comparison to anomaly time series
3 Motivation: Understanding who wins Zooplankton communities include many species & strategies. A few are usually dominant, others always rare, others show intense, unpredictable outbreaks Ave. climatology vs. Ugly(?) Reality
4 Historical Background (1) r vs. K theory (e.g. Pianka 1970, Levins 1968) Investment in reproduction, rapid growth & dispersal vs. Investment in survival and ability to compete (Doesn t t consider age structure within the population, nor constraints imposed by morphology)
5 Historical Background (2) Demographic Theory (e.g. Winemiller & Rose, 1992) Three end-members: Opportunist - quick turnover ( r strategy) Equilibrium high survival rate, especially of juveniles ( K strategy) Periodic fecundity very high, but reached at older age (betting on eventual big win ) (Subsequent analyses identify additional LHS classes: Salmonid, Intermediate )
6 Historical Background: What about zooplankton?? Intensive research on individual traits within a few major taxonomic groups Egg number and survival for freevs. sac-spawning copepods Egg production vs. food availability Brooding Seasonal dormancy & resting eggs So far, little inter-group comparison
7 Our approach: Multivariate ordination of LHS (Principal Coordinates Analysis (PCoA) of species-species distance matrix derived from species-traits matrix, King & McFarlane 2003) to identify: Life-history traits that covary across taxa Taxa that share similar strategies
8 Issues and obstacles encountered: Qualitative diversity of zooplankton LHS is much greater than for fish Within-species plasticity of some traits Information gaps for some life stages (especially survival rate) < often cannot do LxMx integration (Nevertheless, can rank/classify taxa on each trait. This is sufficient for PCoA ordination)
9 Choice of taxa: 36 temperate-subarctic marine spp. (many are present & occasionally dominant in northern California Current System) Free-spawning copepods: Calanus marshallae,, C. pacificus,, C. finmarchicus, Neocalanus plumchrus, Metridia, Temora, Paracalanus, Eucalanus, Rhincalanus Sac-spawning copepods: Pseudocalanus, Pareuchaeta, Eurytemora, Oithona similis Copepods with resting eggs: Acartia longiremis,, A. tonsa,, A. hudsonica, Centropages, Temora, Eurytemora Euphausiids: Euphausia pacifica, Thysanoessa spinifera Hyperiid amphipod: Parathemisto pacifica Gammariid amphipod: Calliopius laevisculus Cladoceran: Evadne Chaetognaths: Sagitta elegans, Eukrohnia Shelled Pteropods: Limacina,, Clio Planktonic tunicates: Salpa fusiformis (salp) Dolioletta (doliolid) Oikopleura (appendicularian) Ctenophores: Pleurobrachia, Mnemipopsis, Beroe Scyphozoan medusa: Aurelia aurita Hydrozoan medusae: Aglantha digitale, Aequorea
10 Reproductive : Log Fecundity r (per day) Lifespan (years) Iteroparous vs. Semelparous Mode of reproduction: Separate sexes? Hermaphroditic? Parthenogenetic? Alternating sexual/vegetative? Parental brooding: Free spawning Retain eggs Retain larvae Three categories of LHS traits: Growth : Log Adult size Somatic growth rate Fraction of lifespan in adult body form Refuges (Depth & Dormancy): Extent of diel migration Ontogenic migration? Dormancy As egg As adult/late juvenile Lipid storage? Benthic stage?
11 Ordination Results
12 PCoA on reproductive traits First 2 coordinates are significant PCo1 PCo2 Percent Variation Eigenvector log Fecundity r (per day) Lifespan (years) Iteroparous/Semelparous Mode of reproduction Parental brooding
13 Ordination of reproductive traits
14 Ordination of reproductive traits High r & fecundity Reproduction mostly sexual Some hermaphrodites Relatively low r Long life span Iteroparous High r, Short-lifespan, Semelparous, Repid vegetative reproduction Low fecundity Iteroparous Parental brooding
15 PCoA on growth traits First 2 coordinates are significant PCo1 PCo2 Percent Variation Eigenvector log Adult size (dry weight) Somatic growth (per day) Proportion juvenile/adult
16 Ordination of growth traits
17 Ordination of growth traits Large adult size, Rapid somatic growth, Diverse morphology or indirect development Intermediate adult size Low daily growth Direct development most of life in adult body form Small adult size, Rapid somatic growth
18 PCoA on refuge traits First 2 coordinates are significant PCo1 PCo2 Percent Variation Eigenvector Extent of diel migration Ontogenic migration Dormancy Storage of lipids Benthic stage Daytime depth
19 Ordination of refuge traits
20 Ordination of refuge traits Benthic stage or resting egg Remain active & near surface Large diel migration Deep daytime depth Ontogenetic migration, Lipid storage Dormancy as adult or late juvenile
21 Scores on lead axes from each PCoA are intercorrelated, mostly due to size dependence of rates
22 Additional 3D views Clusters sharing taxonomic & morphologic similarity: (I) Large vs. small copepods
23 Additional 3D views Clusters sharing taxonomic & morphologic similarity: (II) Large crustaceans vs. Salps/Doliolids
24 Additional 3D views Clusters sharing taxonomic & morphologic similarity: (III) Chaetognaths vs. Pteropods
25 Comparison with Time Series - Initial expectations: Sign of response correlated with LHS? Some LHS favored during some ocean regimes (response to environmental change similar within a LHS cluster) Time scale of response correlated with LHS Rapid decay of temporal autocorrelation by opportunists (=spikey time series) Slow decorrelation by equilibrium and periodic groups (time series contain gradual trends and multi- year fluctuations)
26 Comparison with Time Series- Results Interesting, but more complicated than expectation.. Anomaly time series provide examples of: Anticorrelation within LHS clusters Differences in amplitude & time scale between LHS clusters Differences in time scale within LHS clusters
27 Anticorrelation within LHS clusters Species replacement, driven by shifting faunal boundaries, not by LHS difference
28 Contrasts of sign & time scale within LHS clusters (2) Salp time series definitely spikey Doliolid time series more like the copepods
29 Gradients of amplitude & time scale between LHS clusters Trend and Regime responses within cool water zoogeographic cluster are modulated by reproductive and dormancy strategies
30 Summary PCoA ordination summarized variability among zooplankton life history strategies ALL observed strategies persist locally and are represented ~globally LHS clusters map strongly onto taxonomy, but with some surprises Interannual change (in our region) is dominated by shifting zoogeographic affinity, but modulated by LHS (True elsewhere?)
31 Future Directions Include within-species plasticity of LHS ( point < range description of traits) Improve classification & quantification of multi-phase LHS (sexual-asexual, asexual, active- dormant, planktonic-benthic, direct development vs. metamorphosis) Include more taxa Compare with other regions
32 BIG thanks to the following for advice & data: Mary Arai & Jenny Purcell (medusae( medusae) Carol Lalli (pteropods) Jaime Gomez-Gutierrez, Gutierrez, Tracy Shaw, Leah Feinberg & Bill Peterson (euphausiids( euphausiids) Andrew Hirst (growth rate theory) Xenia Kosobokova (chaetognaths) Moira Galbraith (size & abundance trends)
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34 Spares & extras
35 LHS of Marine Invertebrates (summary from Ramirez-Llorda 2002) Most comparisons have been among benthic taxa Tradeoffs of fecundity vs. other modes of parental investment (egg size, brooding,.) Effects of food availability Gradients with increasing latitude and depth (fewer and larger eggs)
36 Historical Background (3) Fish LHS refinements (e.g. King & McFarlane 2003) Two additional clusters: Salmonid resemble opportunistic but large adults & eggs; split life cycle between ocean & FW Intermediate resemble periodic but shorter life span and lower fecundity
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