By Patricia Lane Dalhousie University Goal to illustrate how Dick Levins loop analysis is useful for analyzing complex systems using a marine ecosystem example Rationale we have one ocean which is constantly under threat we need to understand how perturbations affect ecological netw ID: 685950
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Slide1
Excavating the Bare Bones of Community Structure Using Loop Analysis
By
Patricia Lane
Dalhousie UniversitySlide2
Goal
: to illustrate how Dick Levins’ loop analysis is useful for analyzing complex systems using a marine ecosystem example
Rationale: we have one ocean, which is constantly under threat; we need to understand how perturbations affect ecological networks through a myriad of pathways and feedbacks
Although this is a marine example, a similar use of loop analysis could be helpful to understand any complex system with a detailed data set.Slide3
Traditional DescriptiveFood Webs
This diagram is essentially a summary of every feeding relationship known to the author
Certainly
not stable-excessive connectionsPoor sense of functional groups and links that are ecologically relevantCannot locate parameter
inputs
Not
useful for making predictions about perturbationsSlide4
Objectives
To introduce the field and laboratory studies at the Marine Ecosystem Research Laboratory (MERL)
(NOAA Grant to Levins/Lane)
To illustrate ecological skeletons and some of their propertiesTo acknowledge Dick’s considerable influence on this and other workSlide5
1. Methods: Field Studies
10 cruises in Narragansett Bay
Temperature/lightNutrients & water quality parametersPhytoplanktonZooplanktonSlide6
1.
Methods:
Laboratory Studies Nutrient Enrichment ‘Press’ Experiment at MERLTanks: 1.8 m diameter
5.5 m deep with sediments3 controls
6 treatments
1X, 2X, 4X, 8X, 16X, 32X
~sewage effluents = 1X
12.8N, 1.0P,
0.9Si
-inorganic
Simulate Bay mixing, light, te
mperature, residence time,
b
enthic-pelagic coupling
Slide7
1. Methods: Loop Analysis Models
Loop Analysis
Data Preparation
Prepare a qualitative correlation matrixDetermine functional groups and loop variablesCalculate directed changes in data sets by loop variable
Prepare models
Fit loop predictions to data –try to achieve 90-100% accuracy
Created by Dick
Levins
in 1973 -NY Acad. Science paper
Signed digraphs, qualitative analysis that predicts changes in standing stocks of variables and their turnover rates following a parameter change
Also gives measures of network structure and stability
Used theoretically to explore complex systems as well as with data
setsSlide8
Loop Variables (~300 taxonic
units)
Si=silicaN
=nitrogen/phosphorusN2 =organic nitrogenA1=diatomsA2=dinoflagellatesA3=luxury consuming diatoms
A
4
=silica flagellates
A5
=
microflagellates
/monads
Z
1
& Z
3
= copepods
Z2=
copepodids
and
nauplii
Z4 =
cladocerans
M=mollusc larvae
C=
cirriped
larvae
P=polychaete larvae
G=
gammarids
R=rotifers
S = Sagitta elegansD = decapodsMD=medusaeSlide9
2. Results: 9 Field Loop Models (
Annual Cycle: No 2 models identical
)Slide10
Ecological Skeleton (ES) of the Field Loops
[- +] with 9 link types
Variables in a minimum of 3 models
Solid links 2/3 of networksDashed Links
≥ 1/3 of networks
63 feedback loops & 4400 pathways
ES does not appear in nature-too connected, but it is a framework for the community
~ Chemical isomers
Slide11
Core Loops = Ecological Skeleton (ES)
LA provides basic structure of an ecosystem (or any complex system) as the prevalent variables, links and parameter inputs that explain the pattern of changes in standing stocks
Thus, the Ecological Skeleton (ES) is a form of minimal description (bare bones) used
to capture the main aspects of community structure & how parts and whole interact Most ‘nibbling’ in traditional webs is irrelevant – ES helps get rid of the ‘noisy’ detailSlide12
Some Loop Analysis Benefits
Marine food webs
have a 3 tier structure, largely composed of L1 and L2 feedbacks, short negative feedback loops that are stabilizing, but several
links have 2 or more link types = volatile links (often nonlinearities can be identified)Can identify missing variables (N3 and Ax) and link types (silica-diatoms) that are counter-intuitive, but later explained by biologyCan identify where parameter inputs enter-not always the manipulated oneCan observe how the variables and links change through an annual cycle and how one
community prefigures
another
Loop analysis helps identify what is necessary to measure with great economy of resources (10-15%)
Can compare over ecosystems to measure ES similarities
IN
SUMMARY, LA PROVIDES NEW INFORMATION AND
INSIGHTSSlide13
Comparison of the Field and Lab ESs
Narragansett
Bay(9 cruise dates)
MERL Tanks(135 dates x tanks)Slide14
Some Results
Field
3 large pelagic predators out-competed in MERL tanksPredictions 145 (3 wrong or 2% and 6 zeroes or 4%) 94% correctSystem of small plankton less discrete than in tanksConnectance
: 16%-in line with theoretical predictions of Gardner and Ashby of 13 +/-2 % dividing stable and unstable networksMERL Lab (X-32X nutrient additions)Benthos absorbed most of increased productivity-strong coupling
1046 predictions (12 wrong or 1% and 102 zeroes or 10%) 88% correct
Little response of small algae and copepods up trophic
gradient
Connectance
: higher (19%)-perhaps because of confined space
Limits to enrichment after 4XSlide15
Structural Complexity Calculations
MACRO-UNIVERSE: A
20 variable loop model has 3
N² or 3400 or 7.06 x 10
190
mathematically possible configurations-but most are not biologically reasonable
Lab: 15 variables = 3
225
or 2.25 x 10
107
MICRO-UNIVERSE: 6 x 10
7
to 1.3 x 10
12
for the tanks-biologically reasonable
THE TRUTH “truly” IS THE
WHOLE in the structural and functional complexity of all of these feedback relationships in ecosystems
Stable UnstableSlide16
Ecological Complexity
Dick has called complexity, “the central intellectual problem of our
time.”A complex system cannot be completely described: has at least on incomputable model. (Robert Rosen)Both structural and functional definitions
To advance ecological theory, we need to understand the level of complexity in ecological skeletons with their rich feedbacks, only studying pairwise interactions in natural communities is not enough.Need to develop ecological skeletons for all major kinds of ecosystems Slide17
Emergence
Core models or ecological skeletons (ES) have been computed as summaries of a set of loop models and therefore are calculated as collective properties
Over a number of marine ecosystems, the ES are robust and very similar, transcending space, time, taxonomy, field & lab methods and investigator intuition: EMERGENT???ESs for freshwater ecosystems are different and less robust and have more structural change with nutrient enrichmentAttempts to ‘break out’ of the marine ES have been unsuccessful
Can’t prove one ES is the best or most optimum descriptorSlide18
4. Thank You Rosario
When the MERL work first
started, Dick had a massive heart attack and many of us wondered if he would reach his 55th birthday let alone his 85th.
Although Rosario is not here today, she has to be given a lot of credit.She oversaw Dick’s change of lifestyle, which gave him another 30+ years of great accomplishments, and he gave us innumerable academic gifts, friendships, and life-altering influences.Slide19
4. Thank You Dick
Teaching
-Communities and Ecosystems-Political Ecology -
Sustainability & Global Change 2. Research-Tropical fieldwork & theoretical ecology/Ph.D.-Ecological networks/loop analysis
Political
Work
-
International Development-with a political conscience
-
Cuban
Model-an
alternative to neoliberal
globalization
-H
ow
Cuba could be the First Sustainable
Society?Slide20
Happy
85th
Birthday to an amazing complex system with delightful emergent properties & unique s
elf-organization always exuding creativity and humanity in equal portions.