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Jose Abella Gutiérrez
Comité
Dra. Silvia E. Ibarra Obando
Dra. Theresa Sinicrope Talley
Dra. Sharon Herzka Llona
Dr. Stephen Vaughan Smith
Efectos de la herbivoría de las brantas y los florecimientos algales en la comunidad de Zostera marina
Jose Abella Gutiérrez
Comité
Dra. Silvia E. Ibarra Obando
Dra. Theresa Sinicrope Talley
Dra. Sharon Herzka Llona
Dr. Stephen Vaughan Smith
Effects of Brant Herbivory and Algal Blooms on Zostera marina Community
Introduction
Introduction
Seagrasses as Engineers
Introduction
DETRITIC PATHWAY
Chesapeake Bay
Seagrasses as Engineers
Valentine, J.F., y Heck, Jr., K.L., 1999. Seagrass herbivory: evidence for the continued grazing of marine grasses. Mar. Ecol. Prog. Ser., 176: 291-302.
Introduction
Valentine, J.F., y Heck, Jr., K.L., 1999. Seagrass herbivory: evidence for the continued grazing of marine grasses. Mar. Ecol. Prog. Ser., 176: 291-302.
Turtles and Sirenians are important in some
systems.
Change in herbivorous species.
There are others grazers (limpets, sea urchins,
fish, waterfowl).
Seagrass is food
Introduction
Branta bernicla nigricans
Ward et al., 2005
Introduction
Branta bernicla nigricans
Ward et al., 2005
Introduction
Moore et al., 2004
Ward et al., 2005
Herbivory Effects on Seagrass
Change in seagrass architecture
Guano enrichment and “shortcircuiting” of the detritus cycle
Introduction
Herbivory Effects on Seagrass Architecture
“Since defoliation by grazers rarely kills the host plant, it is generally believed that the principal effect of herbivory is to reduce the competitiveness of grazed individuals rather than to cause outright mortality” (Hulme, 1996)
Seagrass
Macroalgae
Epiphytes
Microphytobenthos
Phytoplancton
Fauna
Introduction
Herbivory Effects on Nutrient Cycling
Introduction
Thayer et al., 1982
Nutrient uptake by leaves and rootsOxygen translocated from the leaves is released into the sediments
Nutrient uptake by leaves and rootsOxygen translocated from the leaves is released into the sediments
Herbivory
Decrease of seagrass competitiveness
Increase of light and nutrients available for primary producers
Decrease of epiphytic abundance
…and usually seagrasses regrow!
Introduction
Ward et al., 2003
Bahía Falsa; Oct – 2007
Eelgrass decline in favor of green macroalgae.
Increase of herbivory intensity as a consequence?
Zertuche et al., 2009
2004
Introduction
Burkholder et al 2007
Seagrass Eutrophication
Introduction
Nutrient over-enrichment produces high
biomass algal overgrowth
Seagrasses shaded by macroalgae
Unfavorable biogeochemical alterations
Seagrass Eutrophication
Introduction
Introduction
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Introduction
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Increase of amonium and sulfide.Hypoxia
Introduction
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Increase of amonium and sulfide.Hypoxia
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Increase of amonium and sulfide.Hypoxia
Short shoots are immersed in toxic concentrations. Anoxia
Introduction
Green macroalgae shade seagrasses
Decrease of oxygen translocatedNutrient fluxes reduced
Increase of amonium and sulfide.Hypoxia
Short shoots are immersed in toxic concentrations. Anoxia
Plant dies and bed disappears if conditions persist
Introduction
Objective: Understand plant-herbivory interactions in seagrasses with frequent and continual algal blooms
Hypothesis: Interactions between algal blooms and herbivory will produce a quick shift from seagrass to algal beds
Objetive and Hipothesis
Guano
Defoliation
Algal blooms
Objetive and Hipothesis
Guano
Defoliation
Algal blooms
Defoliation x Guano
Guano x Algae
Defol x Algae x Guano
Defol. x Algae
Objetive and Hipothesis
Irradiance, temperature, sampling site
Guano
Defoliation
Algal blooms
Defoliation x Guano
Guano x Algae
Defol x Algae x Guano
Defol. x Algae
Objetive and Hipothesis
Materials
and
Methods
Manipulative experiment from Nov-07 to Mar-08
4 seagrass beds: continuous beds, same depth
Materials and Methods
Treatment Simulations
Clipped treatment (2 cm, 3 months):
No differences between plots RM-ANOVA
F(6, 14)=0.123, p=0.99
Fertilizer addition Multicote
Ulva addition
Materials and Methods
Treatment Simulations
Clipped treatment (2 cm, 3 months)
Fertilizer addition Multicote:
DIN: 23.6 (± 6.9) g/mes
DIP: 8.6 (± 2.5) g/mes
Ulva addition
Materials and Methods
Treatment Simulations
Clipped treatment (2 cm, 3 months)Fertilizer addition Multicote
Ulva addition:
2 Kg wet weight
0.876 (± 369.4) Kg
Materials and Methods
Fully Factorial Experimental Design Using Randomized Complete Blocks
Per seagrass bed (site):
1. No treatment
2. Cut of leaves (C)
3. Ulva addition (U)
4. Nutrient enrichment (N)
5. C x U
6. C x N
7. U x N
8. C x U x N
9. Control PVC (K)
n=4, one sample per site
Materials and Methods
Fully Factorial Experimental Design Using Randomized Complete Blocks
Materials and Methods
Field Sampling
Materials and Methods
3 underwater thermistors in 3 sites from Nov 07 to Mar 08
1 light sensor from Nov 07 to Mar 08
5 light sensors during last month (4 sites + land)
Field SamplingNon destructive response variables. Monthly
Materials and Methods
Field SamplingNon destructive response variables. Monthly
Seagrass and algal cover and seagrass density (5)
Materials and Methods
Field SamplingNon destructive response variables. Monthly
Leaf length (10)
Materials and Methods
Field SamplingNon destructive response variables. Monthly
Epiphytes cover (5)
Materials and Methods
Field SamplingNon destructive response variables. Monthly
C and N content
January(0.001 m2) (1)
March (from biomass; 3)
Materials and Methods
Field SamplingDestructive response variables. March 2008
Aboveground biomass (3)
Belowground biomass (3)
Materials and Methods
Laboratory Work
Monthly samples
•Epiphytes cover: Armitage et al. (2005)0 = absent0.1 = one individual < 5% cover0.5 = few individuals < 5 % cover1 = many individuals < 5% cover2 = 5-25% cover3 = 25-50% cover4 = 50-75% cover5 = 75-100% cover
Materials and Methods
Laboratory Work
Biomass samples
AlgaeZostera
roots
Leaves
Clean and freeze-dry
clean
dry
weight
C:N
epiphytes leaves
%C
Materials and Methods
Data Treatment
1.- New variables from data:
• Z. marina growth: monthly increase in total stem length (density x leaf length)
C = (Lt+1-Lt)/Lt*100
• % N, %C and C:N increase: I = (VJanuary-VMarch)/VMarch*100
• Primary producer / Z. marina (i.e. Epiphytes biomass / Z.marina biomass)
2.- No treatment and PVC control plots were merged =
control
3.- Standardization (treatment / control)
Materials and Methods
Data Analysis
3-way ANOVA destructive samples
3-way Repeated Measures ANOVA monthly variables
MRA: Z. marina ~ site, light, Tª, green algae, brown algae and epiphytes
Materials and Methods
Results
and
Discussion
Enviroment: Irradiance
Data from 10 am to 2 pm
Feb-Mar standardized irradiance (site/land):
ANOVA: F = 17.59; p < 0.001
Tukey: C B D A
Site B
Results
Mean ± SE
Environment: Temperature and Upwelling
Temperature: 2-way ANOVA, month and siteMarch warmer (F=48.92; p < 0.01)
Upwelling in March 16th Bakum index = 296 m-3 s-1
Results
Effects of Treatments on Z. marina
Aboveground and belowground biomass were reduced in plots with clipped treatment
3-ANOVA: F = 16.4, p<0.001; F = 10.26, p < 0.01
Abo
vegr
ound
b.
(g m
-2)
Bel
owgr
ound
b.
(g m
-2)
Results
C=clipped; U=Ulva; N=nutrient
C CN CU CUN K N U UN0
10
20
30
40
50
60
C CN CU CUN K N U UN50
100
150
200
250
300
350
Mean ± SE
Effects of Treatments on Z. marinaA
bove
grou
nd b
. (g
m-2)
Bel
owgr
ound
b.
(g m
-2)
Discussion
Study Aboveground Belowground
Valentine and Heck, 1999 40-50% 40-50%
Nacken and Reise, 2000 47 % 43%
Rivers and Short, 2007 100%
This study ~ 70% ~ 40%
C=clipped; U=Ulva; N=nutrient
C CN CU CUN K N U UN0
10
20
30
40
50
60
C CN CU CUN K N U UN50
100
150
200
250
300
350
Mean ± SE
There was no effect of treatments on Z. marina %N, %C or C:N per month
%N decreased from January to March except when Ulva treatment was involved (3 way ANOVA: F=5.51, p<0.05)
Results
Effects of Treatments on Z. marina
January M arch2,5
2,6
2,7
2,8
2,9
3,0
% N
No Ulva Ulva
3
2.9
2.8
2.7
2.6
2.5
Mean ± SE
Results Discussion
Study: (%N or C:N) Aboveground Belowground
Vergés et al., 2008 0=1=3 > 2 0>1>2>3
McGlathery, 1995 no differences
Ibarra-Obando et al., 2004 no differences
Ferson, 2007 no differences
This study no differences
Effects of Treatments on Z. marina
Large seasonal variability (cover, density, # leaves)
Clipped treatment affected cover and density
Results
Effects of Treatments on Z. marina
C=clipped; U=Ulva; N=nutrient Mean ± SE
Cut treatment enhances growth
RM-ANOVA: F=6.01, p<0.01
Results
Effects of Treatments on Z. marina
N o v -D e c D e c -Ja n Ja n-F e b F e b -M a r-1 0 0
0
1 0 0
2 0 0
3 0 0
Relative G
rowth (cm
)
N o c l ip p e d C l ip p e d
Mean ± SE
Discussion
Decrease or increase in density?C = [(D*L)t+1 - (D*L)t] / (D*L)t * 100
Density was reduced DURING treatment but increased AFTER treatment
Effects of Treatments on Z. marina
N o v -D e c D e c -Ja n Ja n-F e b F e b -M a r-1 0 0
0
1 0 0
2 0 0
3 0 0
Relative G
rowth (cm
)
N o c l ip p e d C l ip p e d
Mean ± SE
Discussion
Decrease or increase in density?C = [(D*L)t+1 - (D*L)t] / (D*L)t * 100
Density was reduced DURING treatment but increased AFTER treatment
GROWTH shoots leaves
Moran and Bjorndal, 2005 X
Vergés et al., 2008 X
Valentine et al., 1997 X
Hughes and Stachowicz, 2004 X
Ferson, 2007 X X
This study X X
Ferson, 2007: moderate herbivory > control > high herbivoryThis study: very high herbivory > control
Effects of Treatments on Z. marina
Mean ± SE
Discussion
Seagrasses regrew after 3 events of simulated herbivory in 60 days.
Seagrasses disappeared with 3 – 6 herbivory events (Valentine and Heck, 1991, 1999; Heck and Valentine, 1995; Maciá, 2000)
Ulva addition can reduce seagrass biomass and production (Hauxwell et al., 2001) and density (Nelson and Lee, 2001), but not in this experiment
Unsuccessful enrichment
Effects of Treatments on Z. marina
Discussion
There was no synergistic effect of algal blooms and herbivory (defoliation) on
eelgrass
Maciá, 2000: Interactions between urchin defoliation and macroagal blooms on Thallasia testudinum density but not on biomass
Effects of Treatments on Z. marina
Seasonal influence on brown algae cover
No significant differences between treatments
Effects of Treatments on Green and Brown Macroalgae
% c
ove
r
Bio
mas
s (g
m-2)
Results
March
C=clipped; U=Ulva; N=nutrient. Z. marina; Green A.; Brown A.
Effects of Treatments on Green and Brown Macroalgae
% c
ove
r
Bio
mas
s (g
m-2)
Discussion
High variability:
Ulva clathrata, U. expansa and Dyctiota undulata are floating
macroalgae
Presence of upwelling during March
March
C=clipped; U=Ulva; N=nutrient. Z. marina; Green A.; Brown A.
Dec Jan Feb M ar0
1
2
Cover Index
N o c l ip p e d C l ip p e d
Large seasonal influence on epifaunal, green and red algae cover.
Clipped treatment affected total biomass, but not its relationship with Z. marina. Also affected red algae cover
Effects of Treatments on Epiphytes
C CN CU CUN K N U UN0
1
2
3
4
5
6
7
8
9
Epiphyte B
iomass (g m
-2) Pneophyllum confervicola
Results
C=clipped; U=Ulva; N=nutrient Mean ± SE
Dec Jan Feb M ar0
1
2
Cover Index
N o c l ip p e d C l ip p e d
Effects of Treatments on Epiphytes
C CN CU CUN K N U UN0
1
2
3
4
5
6
7
8
9
Epiphyte B
iomass (g m
-2) Pneophyllum confervicola
Discussion
Settlement of epiphytes in less than 14 days (Borum, 1987)
Differences in settlement patterns across groups (Borowitza et al., 1990)
C=clipped; U=Ulva; N=nutrient Mean ± SE
Effects of Enviroment on Eelgrass Characteristics
Irradiance, green algae biomass, site and epiphyte biomass were related with some Z. marina
characteristics
Results
Z. marina R2 Irradiance Green algae Site 1 Site 2 EpiphytesAbovegroun B. 0.848 -24.39 -17.61 -15.39 12.72 62.67Belowground B. 0.444 -0.499 -0.11 0.178
Density 0.842 -246.28 -194.47 294.01Leaf Length 0.56 10.312 -10.36 8.44
# leaves 0.499 -0.071 -0.04
GREEN ALGAE:
ANOVA: addition of Ulva did not affect Z. marina biomass
MRA: Negative relationship between Ulva and Z. marina biomass
Review (Young, 2009):
•Worlwide: 32 studies, 29 reported eelgrass decline
•Pacific Northeast: 4 studies, 2 reported effects on eelgrass
Algal bloom in San Quintin (Jan – Feb 2009) produced eelgrass decline but regrew (Jul – 09)
High resistance of seagrasses to change (i.e. Sfriso et al., 1989; Venice lagoon)
Discussion
Effects of Enviroment on Eelgrass Characteristics
Conclusions
Large seasonal and spatial variability
Simulated herbivory affected eelgrass, but seagrass reserves allowed the recovery of the bed
The decrease in Z. marina aboveground biomass produced a parellel decrease in epiphyte biomass
Although no effect of Ulva on eelgras was found, there was a negative relationship between green
macroalgae and eelgrass
Future investigations
It is necessary separate herbivory effects during grazing and after grazing
More research is needed to understand the relationship of N content in seagrasses and grazing preferences by
herbivores
More studies of eutrophication are needed in estuaries affected by upwelling
More research that explores the interactions between algal blooms and herbivory is necesary
Agradecimientos
Esta tesis corresponde a los estudios realizados con una beca otorgada por la Secretaría de Relaciones Exteriores del Gobierno de México.
El trabajo fue financiado por el proyecto de UC-MEXUS 622-215 (O0C053): Efecto de las brantas sobre las comunidades de pastos marinos en Bahía de San Quintín
AgradecimientosGracias a mi comité
A Drew Talley
A Juan Guerrero y Ana Salazar
A Hector Atilano y Miriam Poumian
A Victor Camacho y Pepe Zertuche
A los hermanos Aguilar
A aquellos que me acompañaron al campo voluntariamente: Tiago, Doris, Marta, Annelise, Vania, Daniela, Lluis, Berta, Julian, Mariana, Yuca, Karla, Brenda, Luis, Mónica y Tomás. Así como a los trabajadores de San Quintín que nos echaron una mano para cortar el “zacate marino”
A aquellos que me ayudaron en el laboratorio voluntariamente o en servicios sociales. Entre otros, Elsa, Araceli, Venecia, Raul y Filipo. Que ahora no recuerde los nombres de algunos de ellos no significa que les esté menos agradecido.
A Gabi
Y a toda la gente de CICESE...
Muchas gracias
por demostrarme que uno puede sentirse como en casa incluso en el extranjero