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Lakes Complexity in a Latitudinal Gradient

Nelson Fern andeza,b,1,, Cristian Villatea,d , Oswaldo Ter ana,d , Jos e a,d d Aguilar , Carlos Gershenson
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Laboratorio de Hidroinform atica, Universidad de Pamplona, Colombia Centro de Micro-electr onica y Sistemas Distribuidos, Universidad de los Andes, M erida, Venezuela c Departamento de Ciencias de la Computaci on Instituto de Investigaciones en Matem aticas Aplicadas y en Sistemas,Universidad Nacional Aut onoma de M exico d Centro de Ciencias de la Complejidad Universidad Nacional Aut onoma de M exico

Abstract Measuring complexity in ecological systems has demanded general formalizations, in order to compare dierent components and ecosystems at dierent scales. We apply formal measures of emergence, self-organization, homeostasis, autopoiesis and complexity to four aquatic ecosystems disposed in a latitudinal gradient. The measures are based on information theory. Variables representing more complex dynamics in the dierent subsystems of lakes were: In the Physco-chemical, variables related with temperature, oxygen, Ph and hydrology. In the Limiting Nutrients, silicates and phosphorous. In the biomass, Piscivorous and planktivorous shes. Lakes Homeostasis were associated with the spatial-temporal changes according with the seaThis is only an example Corresponding author Email addresses: nfernandez@unipamplona.edu.co (Nelson Fern andez), author.three@mail.com (Cristian Villate), oteran@ula.ve (Oswaldo Ter an), aguilar@ula.ve (Jos e Aguilar), cgg@unam.mx (Carlos Gershenson) URL: http://unipamplona.academia.edu/NelsonFern andez (Nelson Fern andez), http://turing.iimas.unam.mx/cgg (Carlos Gershenson)
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Preprint submitted to Ecological Complexity

December 6, 2013

sons. Biomass subsystem seems follow the temporal dynamics of the physicchemical subsystem than limiting nutrients dynamics. Autopoiesis results showa higher degree of independence of photosynthetic biomass over their environment. On the latitudinal gradient from the Arctic to Tropical, North Lowland Lake appears to represent a transition point for complexity values in all subsystems. Our approach shows how the ecological dynamics can be described in terms of information and can increasing our understanding of ecosystems and complexity itself Keywords: Ecological Dynamics,Complex Systems, Information Theory,Self-Organization, Emergence, Complexity,Homeostasis, Autopoiesis. 1. Introduction In the last years, the complexity theory and their associated properties like the self-organization, emergence, criticality have been increasing numbers of applications in ecological systems ?. The study of complexity in ecology has been tried to relate with ecological richness, abundance, and hierarchical structure. As a result, dierent approximations have been explored to develop mathematical formalisms, in order to represent the ecological complexity in particular as ecological indicator (Parrot, 2005). The information theory has been useful for the development of several models of complexity and it is hasbeen used in dierent ways as it can see in Prokopenko et al (2009). In Ecology the formalizations resulting of application of information theory, often relate the highest degree of complexity to random states; others in an opposite way, relates the highest degree of

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complexity with regularity ?. Thus, the meaning, interpretation and applicability of the complexity notion and their associated properties in ecology remains a challenge in ecology ?. As result, general and simple proposals for modeling and measuring complexity in ecology is needed. According with Fern andez et al.(2014) and Fern andez & Gersheson (2014), the description of the complexity in terms of properties like emergence, selforganization, and others related with the self-regulation and autonomy such as homeostasis and autopoiesis is suitable. The importance of these properties is that they comes from the relevant interactions among systems components and generates novel information. Novel information is useful to describe the complex behavior in ecological systems. It can be said that this novel information is emergent, since it is not in the components, but it is produced by interactions. Interactions can also be used by components to self-organize, i.e. produce a global pattern from local dynamics. Interactions are also key for feedback control loops, which help systems regulate their internal states, an essential aspect of living systems(Fern andez et al, 2014). Among multiple ways to describe the state of an ecosystem, the balance between change (chaos) and stability (order) states has been proposed as a characteristic of complexity Langton1990,Kauman1993. This way, we can say that more chaotic systems produce more information (emergence), and more stable systems are more organized. Thus we propose, based on information theory, that complexity can be dened as the balance between emergence and self-organization (Gershenson & Fern andez 2012; Fern andez et. al. 2015). This approach has been applied to some ecological systems (Fern andez et al. 2013*eccs) with good results indicating that ecological

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dynamics can be described in terms of information. In this context, this papers expand the useful of the application of complexity measuring applying formal expressions of complexity, self-organization, emergence, homeostasis and autopoiesis to the Physico-chemical, nutrients and biomass subsystems to four types of lakes located in a latitudinal gradient (Arctic, North Lowland, North Highland to Tropical), in focus to evaluate the usefulness and benets in ecological systems. 2. Measures The measures applied in this paper have recently developed and compared with other previously proposed in the literature (Fern andez et al., 2012; Gershenson and Fern andez, 2012); more rened measures, based on axioms, have been presented in Fern andez et al., (2013). In general, Emergence refers to properties of a phenomenon that are present now and were not before. If we suppose these properties as nontrivial, we could say it is harder now than before to reproduce the phenomenon. In other words, there is emergence in a phenomenon when this phenomenon is producing information and, if we recall, Shannon proposed a quantity which measures how much information was produced by a process.Therefore, we can say that the emergence is the same as the Shannons information I. Thus E=I Self-organization has been correlated with an increase in order, i.e. a reduction of entropy (Gershenson and Heylighen, 2003). If emergence implies an increase of information, which is analogous to entropy and disorder, selforganization should be anti-correlated with emergence. We propose as the 4

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measure S = 1 I = 1 E. We can dene complexity C as the balance between change (chaos) and stability (order). We have just dened such measures: emergence and selforganization. Hence we propose: C = 4 E S.. Where the constant 4 is added to normalize the measure to [0, 1] For homeostasis H, we are interested on how all variables of a system change or not in time. A useful function for comparing strings of equal length is the Hamming distance. The Hamming distanced measures the percentage of dierent symbols in two strings X and X. As it has been proposed, adaptive systems require a highC in order to be able to cope with changes of its environment while at the same time maintaining their integrity (Langton,1990; Kauman, 1993). If we have X represent the trajectories of the variables of a system and Y represent the trajectories of the variables of the environment of the system, If X had a high E, then it would not be able to produce its own information. With a highS, X would not be able to adapt to changes in Y . Therefore, we propose: A = C(X )/C(Y) . 3. Case Studies The data of dierent lakes models used in this section was obtained using The Aquatic Ecosystem Simulator (Randerson and Bowker, 2008). The model used is deterministic, so there is no variation in dierent simulation runs. All variables and daily data we obtained from Arctic, North Highland, North Lowland and Tropical are shown in Annex A. The criteria for lakes choosing was the location of each lake in a latitudinal gradient from the Polar to the Tropical Zone (Ar-T), where light and 5

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temperature conditions have a high amplitud and variation. 3.1. Arctic Lake (Ar) Arctic lakes are located at the Arctic Polar Circle. Their mean surface temperature is around 3C and their maximum is near to 9C and their minimum is 0C. In general, Arctic lake systems are classied as oligotrophic due to their low primary production, represented in chlorophyll values of 0.8-2.1 mg/m3 . The lakes water column, or limnetic zone, is well-mixed; this means, there are no stratication (layers with dierent temperatures). During winter (October to March), the surface of the lake is ice covered. During summer (April to September), ice melts and the water ow and evaporation increase. Consequently, the two climatic periods (winter and summer) in the Arctic region cause a typical hydrological behavior in lakes. This hydrological behavior inuences the Physico-chemical subsystem of the lake. Limiting Nutrients in the form of nitrates, silicates and carbon dioxide are between 90 and 100% available for phytoplankton in all year. Thus, phytoplankton and periphyton biomass is dominated by planktonic (38.6%) and Periphytic diatoms (45%). For zooplankton, the 91.7% is dominated by herbivorous. At the Benthic Zone, detritivorous invertebrates with a 86.8% of total abundance and piscivorous shes with the 85.8% are the two groups with high dominance in their respectively group. 3.2. North Highland Lake (NH) .

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North highland lake corresponds with a mesotrophic ecosystem in a cool North-Temperate climate (Mean=5.3C). Levels of chlorophyll are between 2.2.-6.2 mg/sec. The surface is covered with ice in winter (end of November, December, January and early February). Ice covering forms a barrier to the wind which minimizes losses of water evaporation while the bottom of the lake remains unfrozen. The water column does not thermostratied and is permanently well mixed whit levels of 50 percent in summer and 90 percent in winter. The maximum ows are in spring and autumn (9.6 m3 /sec) with minimum ow in summer(0.6 m/sec). Evaporation is reduced because their water is more or less cold and vapour-pressure gradients are no large (mean of 9,262 m3 /day ). Retention Time is maximum in summer with 100 days. Oxygen concentration is upper to 10 mg/lt in the three layers. pH mean values are around 7 to 7.3 units, but it moves in a range of 6.7 to 7.8 units from the surface to bottom. The correlation among variables are more seasonal in NH than NL. This means, the period of summer is related with high retention time, higher pH. Winter season is related with the higher levels of oxygen, inow and outow and oxygen. However, there is a more strong correlation of benthic and sediment Oxygen. Limiting Nutrients like nitrates and carbon dioxide are around of 95% available for phytoplankton. Phosphates and Silicates shown variations and less percentage of availability. The former around of 80% and the second one around 95% all year. Biomass composition is dominated by planktonic (46.7%) and benthic (41%) diatoms. Zooplankton composition is almost of herbivorous zooplankton (91.4%), but carnivorous zooplankton reaches a low

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percentage of 8.6. In the group of benthic invertebrates detritivorous dominates with the 87.5%. Fish community is dominated by benthic sh again, but in a high proportion (88.9%). 3.3. North Lowland Lake (NL) . North lowland is an eutrophic lake, located in a warm North-Temperate climate (mean T of 14C). Their primary production expressed in mg/m3 of chlorophyll is around 6.3-19.2. There are four seasons in a year, winter, spring, summer and autumn. In summer, the ow variations between inow and outow fall to 3.5 from 25.2 m3 /sec. Retention time increases to 100 days. The lack of wind and high temperatures (24C), causes the water column thermostratication. Stratication is expressed in generation of two layers. At the border of these layers, temperature changes dramatically (24C Surface to 20.6C in Planktonic layer, to 17.3C in Benthic layer). Water above and below of thermocline do not mix. The warmer water is near the surface and denser water is near the bottom. In winter, there is no ice covering in the surface. Opposite to the summer when the ow is minimum, in spring and autumn the water column overturns (Retention Time of 14 days and Zone Mixing of 100%), causing increases in conductivity. In summer, depletions of oxygen at the three layers are more drastic than Artic lakes (below 8.7 mg/lt). Oxygen is directly correlated with the zone mixing, inow and outow, and inverse correlated with the others parameters, especially with pH and retention time. All limiting nutrients are above of 90% available for phytoplankton in all seasons. According with this availability, phytoplankton and periphyton 8

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biomass composition is dominated by planktonic (47%) and benthic (34.3%) diatoms. This way, 100% of zooplankton composition is reached by herbivorous zooplankton and sh community is dominated by benthic sh (67.6%). 3.4. Tropical Lake (T) The Tropical Lake is a hyper-eutrophic ecosystem (Chlorophyll 19.2 mg/lt) located in a moist Tropical climate, at north of the equator, near to the tropic of cancer with a mean temperature of 25C in the surface layer. Tropical lake has a wet season and a dry season. Higher irradiance conducts to higher temperatures and smaller thermal dierences between layers. For that reason, the water column is permanently warm and stratied. Stratication is by the heat exchange, but it is less stable than stratication in lakes at higher latitudes. Specially, because the wind could have great incidence in the mixing of the water column. Thus, intra-seasonal variations have an eect in thickness of the mixed layer than other morphometrically similar temperate lakes (AES, Lewis**). The maximum ow of water is in the wet season, and minimum ow is in the dry season. Episodes of heat and mixing, aects the nutrient cycling and plankton dynamics. It is highlighted that primary production in tropical lakes is about twice than higher latitudes. Also, it is known that Nitrogen is the more limiting nutrient. The equilibrium among species inside phyto and periphyton communities (around 33% for diatoms, green algae and cyanobacteria) is higher. Zooplankton populations are dominated by herbivorous (90%), benthic by detritivorous invertebrates (84.4%) and shes (87%).

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4. Results 4.1. Complexity as the Balance Between Emergence and Self-organization 4.1.1. Complexity in the Physico-chemical Subsystem At Ar variables related with light in surface, planktonic and benthic zones (SL, PL and BL) have high value of emergence. Variation of light was 10.28 10.44. Meantime, benthic conductivity (BCd, 600.3 40.8 S) and percentage of water mixing between planktonic and benthic zones (ZM, 50 3.53%) have very high self-organization. Remaining variables were classied in very high complexity category, with the exception of two variables associated conductivity (ICd=3896.96 17.29 S and BCd= 600,32 40.53 S) which were ranking in very low complexity category. At NH, light variables increase their ranking to very high emergence category, in consequence its complexity was reduced to low and very low categories. The light variation for all zones was between 12.31 9.31. Meantime BCd (598.2 95.91 S) and ZM (62.05 19.36) variables increase its complexity to high category. At NL, temperature variables increases its level of change to very high. The variation for all zones was between 11.3 6.5. Also light variables (14.6 7.4). Remaining variables had fair to very high emergence. Thus, self-organization, in general is low. As the result of the above facts, variables with very high complexity were relating with hydrology (IO= 10.7 6.5, RT= 61.9 40.4) conductivity (ICd=870 93.5, PCd=1132.6 182.2) and pH in all zones of lake (7.0 14,67 0.2) At T, all Physico-chemical variables have similar levels of regularity (S) and change (E); consequently most variables have high or very high complex10

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Figure 1: Principal Component Analysis for Physico-chemical Subsystem.

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ity with the exception of ZM (16.48 2.29) and Sediment Oxygen (SdO2 = 2 0). Based on Principal Component Analysis (PCA) Figure 1 the ordination of emergence, self-organization, complexity and autopoiesis properties, we can summarize that variables of Physico-chemical subsystem can conform 3 groups. Group 1 including variables related with high changes or emergence as light. Group 2 conformed by variables associated with high regularity like conductivity and zone mixing, and Group 3 Variables expressing high complexity like temperatures, oxygen,pHs, retention time and inow and outow. 11

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4.1.2. Complexity in the Limiting Nutrient Subsystem Limiting nutrients shown at the Arctic high changes in inow silicates (IS= 25014.31 3545,08), carbon dioxide in the inow and planktonic Zones (ICD= 12006.9 1701.8, PCD= 10105.7 979.4). Very high regularity in nitrogen at the 3 layers of lakes (IN,PN and BN= 67.87 13.36); high regularity in silicates (PS= 40550.7 7272.34, BS= 41160.69 7453.5), phosphorous (PP=9.36 1.25, BP = 9.52 1.28) in planktonic and benthic zones. Also, there is high self-organization for planktonic detritus (PDt= 21.26 2.55). In complexity terms very high category was for IS, PP,BP, BCD, PDt and BDt. For the NH, planktonic carbon dioxide (PCD= 9788.5 2119.1) had very high emergence. Inow and benthic carbon dioxide (ICD= 10005.7 1418.2, BCD= 7571.1 3150.3) were in high emergence category. In contrast, variables with very high self-organization were silicates in planktonic and benthic zone (PS= 25257.32 7025.4 BS= 25703.99 7216.8), phosphorous in 3 layers (IP,PP and BP= 7.69 2.26) and nitrogen in inow and planktonic zone (IN and PN= 62.91 15.1); nitrogen in benthic was high self-organization (79.21 9.34). Variables in the very high complexity category were inow silicates (IS), carbon dioxide in inow and Benthos (ICD,BCD), and detritus (PDt,BDt). At the NL, due to an increasing in the emergence of nitrogen (143.4 39.08) and decreasing in the self-organization of detritus (228989.6 245332.9), 13 of the 16 variables of the limiting nutrient components was classied in very high and high complexity categories. Carbon dioxide in planktonic and benthic zones (PCD= 9746.7 1477.7 and BCD= 10888.03

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2105), and benthic detritus (BDt= 457320.9 126432.4) were categorized as low complexity variables because of their very high emergence. At this point of the gradient Ar-T, complexity of the limiting nutrients subsystem has an important variation in terms of its increasing with respect of Ar and NH levels. This levels continuous its increment due the balance between emergence and self-organization values at the end of the gradient. This way, at the tropic lake a very high levels of complexity for the majority of variables are shown. Only a very high emergence of detritus were the exception. From PCA ordination Figure 2 for limiting nutrients at all lakes, the groups that can be identied were a rst group representing emergence with detritus and carbon dioxide variables. A second group representing selforganization in nitrogen and inow phosphorous. A third group representing complexity variables with silicates and phosphorous in planktonic and benthic zone. 4.1.3. Complexity in Biomass Subsystem At the Ar, self-organization for all groups of phyto and zooplankton species in all zones, were high or very high. Only the low values of emergence of diatoms (PD and BD= 185.4 191.3), cyanobacteria (PCy and BCy= 118.9 169.3) and green algae (PGA and BGA= 164.2 160.6) permits that these photosynthetic organisms reach very high levels of complexity and autopoiesis. This situation continuing in NH in spite of planktonic diatoms (PD= 281.12 209.35) and cyanobacteria (PCy= 162.9 169.5) reached the fair category of emergence. It means, the feature of this two types of lakes is their regularity. In a similar way with limiting nutrient subsystem, when gradient Ar-T 13

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Figure 2: Principal Component Analysis for Limiting Nutrients Subsystem

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reaches the NL point, the dynamics of emergence and self-organization varies in considerable level. Here, the complexity of almost all variables were maximum due the balance in self-organization and emergence. Only chlorophyceas (PCh= 6.2 5.1), benthic detritivorous (BDt= 3.84 71.71) and shes in planktonic and benthic zones (PiF, BF and PF= 0.2 4.51) have very high regularity for all annual cycle. In contrast to the NL, the biomass subsystem in the tropic reects very low complexity due the very high self-organization of the living taxa. Only planktonic and piscivorous shes (PlF= 0.099 0.005; PiF= 0.13 0.67) have very high and high complexity, respectively. From PCA ordination Figure 3, it can be seen that photosynthetic taxa of

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planktonic and benthic zone are more emergent. In addition with planktonic primaries and secondaries consumers, clorophyceas and benthic detritivorous are more self-organized than other taxa.On the other hand, piscivorous and planktivorous shes are more complex.

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Figure 3: Principal Component Analysis for Biomass Subsystem

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4.1.4. Complexity in Latitudinal Gradient Comparing the average of complexity for an annual cycle in Ar-Ttransect as latitudinal gradient, we can see thatNLappears to represent a transition point for complexity values Figure 4. At this point for Physico-chemical subsystem decreasing complexity goes from very high to highcategory. This isby reason of emergence increasing (0.75). Then, at the tropical lake the category of complexity returns to very high level due to the increasing of

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self-organization (regularity in variables) and their consequently emergence reduction.

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Figure 4: Complexity in The Latitudinal Gradient Ar-T

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For limiting nutrients subsystem, complexity goes from high at the Ar to fair in NH; high category is maintaining in NL and T. The transition pointinNL is more evident fromtheiremergence values. Emergence is almost 0.62 (fair category).Also, emergence starts in the low category in Ar,and nish in it Tin the same category. This means, limiting nutrient change to a greater proportion at NL latitudes. For Biomass, the transition at NL point is more evident than other subsystems because complexity values reach the higher category of Ar-T transect (0,74; very high category). For Ar and NH biomass, complexity value was classied in the low category and for T in very low. In terms of mean complexity by subsystem, it is seen that Physico-chemical Limiting Nutrients Biomass. This order corresponding with the autonomy, which means in general biomass is aected in an important way by changes in their environment. However, the dispersion of Biomass is more than the other two (Figure 4) which means that biomass can respond accord16

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ing with the law of required variety to the environmental changes between its complexity range (0.382 0.22). By lake, we can observe that Ar and NL were in the high category and NH and T were in the fair category. In terms of dispersion T ArNHNL (Fig.**). Parametric multiple comparison by means of the test of Tukey shows that, in terms of average complexity, physic-chemical and limiting nutrients did not have signicance dierences (p= 0.85; p0.05) while biomass has signicance dierences with the other two subsystems (*p0.05). On the other hand, ANOVA test shows that there are not signicance dierences among complexity of lakes in the Ar-T transect (p0.05). In ecological terms, the dynamics observed at NL point in the transect ArT could be estimates as a complexity ecotone or complextone (tone, from the Greek tonos or tension). That means that NL point could be considered as a physical transition zone for complexity values among lakes in a latitudinal gradient. Consequently for some variables therein subsystems it is estimates that it could be represents diverse complexity ecoclines or complexcline (cline from Greek: to possess or exhibit gradient, to lean), due their complexity variation. For example for biomass, we can that there is a biocline in the transect Ar-T and in particular for cyanobacteria at the planktonic zone (PCy) we can name a complex cyanocline. From PCA ordination from variables of all subsystems conducts to the conformation of groups of lakes based on emergence, self-organization, complexity and autopoiesis properties. However, we chose the complexity criteria for denition of groups due to complexity relates the regularity and changing

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aspects, and it is the base for autopoiesis calculation. This way, the general ordination shows theNL disjoins of the group conforming by NH-Ar with T. The separation of NL from other lakes is by cause of their variability and load of some variables of biomass and Physico-chemical subsystem. These variables were macrophytes, clorophyceas and planktonic phosphorous and by the low level in complexity of the shes and light variables. NL separation also supports the consideration that NL represents a transition in the values for all properties, marking this location as dierential on the latitudinal gradient from the arctic to tropic. 4.2. Autopoiesis There are two ways for observing autopoiesis (A). The rst one is the A of the each variable. Variables with more complexity than other have a positive A reecting more autonomy. Variables with low complexity than other have negative A, reecting less autonomy. Results of A by variable in each subsystem can be seen in Annex A. In general, variables in categories of very high and high complexity, have more A and they resulting as more autonomous, as well.That means, they have more capacity of adaptation in front the changes of their environment which is constituted by the other subsystem variables. The second one form of determine autopoiesis (A) is among variables of different subsystems, according with the matter-energy ux in ecosystems. It is well-know that photosynthetic living beings depending of solar radiation and nutrients availability as the base for its metabolism process. Also, zooplankton depending of grazing phytoplankton communities. Starting from complexity values of selected variables of Physico-chemical, Limiting Nutrients and Biomass are shown in the Table 1. We compare A 18

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of Biomass related with the their Physico-chemical and Limiting Nutrients environment. A Biomass values for planktonic and benthic zone are depicted in Figure 5.
Table 1: Selected Variables for Phytoplanktonic Biomass Autopoiesis Calculation

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Component Physiochemical Limiting Nutrients Biomass

Planktonic zone Light, Temperature, Conductivity, Oxygen, pH. Silicates, Nitrates, Phosphates, Carbon Dioxide. Diatoms, Cyanobacteria, Green Algae, Chlorophyta. Light, Temperature,

Silicates, N

Diatom

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From the Table 1, we notice that Biomass in T is near to zero in the planktonic zone and zero in the benthic zone. It means that tropical biomass is almost static. We can verify this with their very-high category of selforganization obtained. This implies that any pattern in complexity can be observed in biomass as the result of the inuence of its environment which is represented by its Physico-chemical and Limiting Nutrients subsystem. This case gives a minimal A for all comparisons carried out with the trajectories of Biomass in tropical lake. Values of A1 were reached by photosynthetic living beings located at the benthic zone of Ar,NH and NL in front of Physico-chemical and Limiting nutrients. Also, A1 was reached by Biomass/Limiting Nutrients of planktonic zone at Ar and NH and Biomass/Physico-Chemical of planktonic zone of NL Figure ??. In terms of the Ashbys Law of Requisite Variety(Ashby, 1956), photosynthetic biomass in Polar and Template latitudes have more variety than its environment. More variety is related with more number of states. 19

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More states permit to face on environmental changes. Variety is the result of very high complexity and could be reecting as more autonomy of the taxa therein.

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Figure 5:

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The remaining cases of biomass obtain values less than one (A1) and were related with their response in front of Physico-chemical in the planktonic zone of Ar and NH. Also NL biomass response in front of limiting nutrient at the same zone obtain A1. This values between 0.72 and 0.98 shows that their environment changes more than the populations of photosynthetic living beings. As we can see, the weak or almost fair lake-specic response of the biomass, suggest that species involved in this subsystem could be aected in a high proportion in case of strong change events. On the base of above ndings, we thought that relationships evaluated in 20

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biomass of dierent lakes might be evaluated in the context of environmental variability. 4.3. Homeostasis The homeostasis h calculation by comparing the daily values of all variables is a useful indicator for periodic or seasonal dynamics characterization and determination. h can represents the temporal variation of the state of each subsystem in each lake. This situation is more evident in the Physicochemical subsystem of lakes which responds proportionally with the seasonal changes aecting variables like temperature, light and others related with the hydrological cycle. Limiting Nutrient subsystems works in a shorter scale than Physico-chemical subsystem; it seems that nutrients could vary more at week scales than month scales. For all three subsystems, biomass demonstrated has similar scale variation with Physico-chemical subsystem; in special at the T which their biomass variation takes place in several months intervals. The above homeostasis results demonstrates the importance of the temporal timescale because h can vary considerably if we compare states every minute or every month (see homeostasis gures in Annex A). For H, the Table 2 shows average values and standard deviation for all lakes and subsystems. Considering the lakes studied maintains periods without changes, values of H are all in the category of very-high homeostasis. We observe that Biomass and Physico-chemical were more stable in a year than Limiting Nutrients subsystem. In detail, the Ar and NL Biomass were more regulars than NL and T. Physico-chemical subsystem has a similar regularity for all lakes being the lower NL. For Limiting Nutrients, the lower regularity is also for NL and the high for Ar. 21

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Table 2: Homeostasis averages H for Lakes

Lake

Biomass Sd

P hy Chem Sd

Lim N utr Sd

Ar

0.9803620 0.04471970

0.9594521 0.06440536

0.9574551 0.06520360

NL

0.97643440.05434534

0.94301370.0922801

0.91471510.10648702

NH

0.91729450.10604234

0.95739730.07495470

0.94516000.08047069

0.95799180.11550661

0.96493150.05803853

0.94828260.07298964

Global

0.95802070.03578496

0.956198630.01496563

0.94140320.01791861

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5. Discussion 5.1. The ecological sense of the proposed measures. The proposed measures characterize the dierent ecological congurations and dynamics that elements of lakes acquire through their interactions. From simple mathematical expressions, based on probabilistic features, we can capture the properties and tendencies of the ecological systems, considering the scale at which they are described. In contrast with other complex computational methods, our approach permits analyze the properties and tendencies of any variables, in dierent subsystems, for one or more ecosystems. Also, we can change scale and apply the measures there, in order to determine at which scale the richness dynamics is representative. In instance, for Physicochemical subsystem in Ar, previous analysis shows that base 10 is very informative and represents the dynamics as base 8 or 16,34 and 64 (Fern andez et al., 2014). The integration of self-organization and emergence aspects into our C measure has advantages asthe complex dynamics of an ecological system, can be observed as the balance between regularity and change or variability. In this context, it can dene which variable, process or ecosystem is more complex than other as we can observe in The characterization of the behavior of the biotic component, in front of environmental disturbances or environmental variability, can be complemented with the autopoiesis and homeostasis measures. It is an important feature that will be useful for studies of global ecological change. In general we suggest that systems with a higher complexity are more robust while those with a lower complexity are less. 23

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Clarifying that chaos should not be confused with complexity (Gershenson, 2013), we highlight that our measures can distinguish between random and non-random ecological processes or variables. The former is related with very high emergence (high entropy), it implies too many changes and patterns destruction. The second implies very high self-organization (very low entropy); it prevents that complex patterns emerging. For further details, this randomness can be examined in the probability distribution for any process or variables at dierent scales, as well. Besides our measures can be related with the temporal and organizational scales and uctuating environments, they can be related with other ecological aspects like occupancy, movement patterns and numbers of species as show in Fern andez et al (2013). This way, proposed measures can be good complements in status and trends study, in ecological communities. Based on the complexity average values for the 3 subsystem, we can observe that before the NL point, the complexity of limiting nutrients and biomass have an increasing trend; then the values trend is decreasing. Meantime complexity in the Physico-chemical shows a decreasing trend before NL, then at T ischangein an increasing way. This results suggest that there is a the dierential trend of complexity according with the subsystems. Thus, we can not suggest that there is a clear global pattern in the trend of complexity for lakes according with the latitude. 5.2. Complexity and others Measures of Information. Complexity has been correlated with other measures of information like Fisher Information (Prokopenko et al., 2011) and Tsallis Information (Tsallis, 2002; Gell-Mann and Tsallis, 2004). Contrasting C with Fisher Infor24

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mation we observed that C is smoother, so it can represent dynamical change in a more gradual fashion. Fisher information has a much higher steepness in comparison with C. On the other hand, test of Tsallis information for Ar lakes shows that it follows self-organization patterns in one cases and others emergence patterns. This results generate a dicult to establish a signicant correlation with C for Tsallis measure, is increased with its variable scale and determination for the optimal q choice. It seems that q=2 is the value in which some correlations can be appear. In the context of the physics, an interesting point is that dierent measures of entropy are used for describe dierent probability distribution. Shanon entropy is logarithmic, and it is appropriated for the phenomena with exponential distribution. Tsallis entropy has a power model, and it is appropriated for phenomena with power distribution. It has been found that critical phenomena considered by someones as complex, usually have power law distribution referred as self-organized criticality (Per Bak, **). Consequently, Tsallis information has been recommended as complexity measure (**). However, we consider that in itself Tsallis entropy might not be a complexity measure, in particular for its q parameter dependence and sensitivity. Tsallis information could be a more a general description applicable to several phenomenon, previous their distribution inspection and knowing. As a sample of this situation, the results of application of Tsallis entropy to Physico-chemical subsystem can be observed in the Annex **

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6. Conclusions Based on Information Theory we dene complexity as a type of balance a balance between change (emergence) and regularity/order (self-organization). The balance is reached in terms of their autonomy (autopoiesis) and (homeostasis). It can be seen that variables, subsystem or system with a homogeneous distribution have higher values of emergence whilevariables with a more heterogeneous distribution have a higher self-organization. For the two additional properties in lakes studied, Homeostasis values coincide with the variation of dierent seasons according with the latitudinal location of lakes. Autopoiesis values show a higher degree of independence of biological components over their environment. There are dierent ways to describe the state of an ecosystem and the dynamical of species therein. Measures of emergence, self-organization, homeostasis, autopoiesis and complexity can complement the description of ecosystems and species dynamics. They could be viewed as ecological indicators at dierent scales and have high potential for comparative analysis among ecosystems. In fact, the complexity analysis can be focused in either particular system components, or a subsystem of the whole, oraecosystem as unity. For example, we can observe the complexity of the predatory-prey cycles related with the movement decisions and foraging behaviors in contrasting with the vegetation patterns. In this sense, our measure can contribute with the interpretation of the six types of Complexity (spatial, temporal, structural, process, behavioral, and geometric- cloehle, 2004). A

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7. Complexity for Each Component


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Figure 6: Complexity in Physico-chemical Subsystem for an Arctic Lake.

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Figure 7: Complexity in Physico-chemical Subsystem for a North Higland Lake.

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@ArticleEinstein, author = Albert Einstein, title = Zur Elektrodynamik bewegter K orper. (German) [On the electrodynamics of moving bodies], journal = Annalen der Physik, volume = 322, number = 10, pages = 891921, year = 1905, DOI = http://dx.doi.org/10.1002/andp.19053221004

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Figure 8: Complexity in Physico-chemical Subsystem for a North Lowland Lake.


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Figure 9: Complexity in Physico-chemical Subsystem for a Tropical Lake.


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Figure 10: Complexity in Limiting Nutrients Subsystem for an Arctic Lake.

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Figure 11: Complexity in Limiting Nutrients Subsystem for a Noth Higland Lake.
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Figure 12: Complexity in Limiting Nutrients Subsystem for a Noth Lowland Lake.
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Figure 13: Complexity in Limiting Nutrients Subsystem for a Tropical Lake

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Figure 14: Complexity in Biomass Subsystem for an Arctic Lake.


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Figure 15: Complexity in Biomass Subsystem for a North Higland Lake


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Figure 16: Complexity in Biomass Subsystem for a North Lowland Lake.

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Figure 17: Complexity in Biomass Subsystem for a Tropical Lake.

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