Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | every 1 − 5 for peat swamps, 0.5 MgCha 1 ± − ) compared to lowland 1 , and L. Castro − 4 ). Soil C sequestration values 1 − 83.4 MgCha ± to the atmosphere (Lovelock et al., 0.5 MgNha 2 ± th.edu.au) ffi 2.0 MgNha , A. Vázquez-Lule ± 3 ; 26.4 1015 1016 1 − ; 13.8 1 − ) emissions to the atmosphere (Van der Werf et al., 2 th University, Nathan, 4111, QLD, Australia , C. Tovilla . The Reserve stores 32.5 Mtons of C or 119.3 Mtons of 2 for marshes. C stocks and soil N stocks were in general ffi 1 for , 722.2 7.2 MgCha − 1 − ± yr 1 − 1 − 18.6 MgCha ± , N. S. Santini 1 38.3 MgCha 0.2 MgCha 73.5 MgCha ± ± ± , with mangroves sequestering (via soil accumulation) 27 762 2 This discussion paper is/has beenPlease under refer review to for the the corresponding journal final Biogeosciences paper (BG). in BG if available. Coastal Plant Laboratory, The School of Biological Sciences, TheColegio University of de Queensland, la Frontera Sur, TuxtlaComisión Gutierrez, Nacional Chiapas, para Mexico el Conocimiento y Uso de laComisión Biodiversidad Nacional (CONABIO), de Áreas Naturales Protegidas, Chiapas, Mexico Australian Institute, Gri 2 St Lucia, 4072,3 QLD, Australia 4 Mexico City, Mexico 5 Received: 3 November 2014 – Accepted: 29Correspondence November to: 2014 M. – F. Published: Adame 16 (f.adame@gri January 2015 Published by Copernicus Publications on behalf of the European Geosciences Union. Abstract Deforestation and degradationemissions of to are thesequestration important atmosphere. rates causes Accurate of arerestoration measurements programs carbon needed with dioxide the of aim forwhole for carbon preventing ecosystem incorporating carbon C (C) emissions. stocks wetlands Here, stocks (trees, wewetlands soil assessed into (mangroves, and and downed marshes conservation wood) andReserve and and in soil peat the N swamps) Pacific stocks coast within ofmangroves of riverine on La Mexico. We the Encrucijada also basis estimated Biosphere soil ofhave C soil large sequestration accumulation. rates C We of hypothesizedcompared stocks, that to and riverine lowland wetlands that mangroves.of Riverine upland wetlands 784.5 mangroves had have large larger C stocks C with and a soil mean N stocks larger for upland (833.0 1 and 336.5 mangroves (659.5 CO were 1.3 Carbon stocks and soil sequestration rates of riverine mangroves and freshwater wetlands M. F. Adame Biogeosciences Discuss., 12, 1015–1045, 2015 www.biogeosciences-discuss.net/12/1015/2015/ doi:10.5194/bgd-12-1015-2015 © Author(s) 2015. CC Attribution 3.0 License. 2009). Wetlands have onemangrove of the have highest been deforestationhas lost rates; disappeared in one since the third the pasttherein). of 1800s 50 Because years, the wetlands (Alongi, while are world’s 2002; rich one in McLeodecosystems third carbon et (C), of results deforestation al., saltmarshes or in 2011 disturbance of and large these references emissions of CO year. 1 Introduction Deforestation and ecosystem degradationcause is, of after carbon fossil fuel dioxide combustion, (CO the largest 5 10 15 25 20 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) 1 N, − 0 yr 43 2 ◦ − of organic C, 10 gCm 1 − cient ecosystems ∼ ffi and other financing + (including organic and 1 − ) and freshwater wetlands 1 − , IPCC, 2003). Similarly, soil yr 1 − 2 − that range between 20 and 40 m 400 Mgha < 1018 1017 ciencies in its use have resulted in a cascade of ffi Rhizophora mangle , geomorphological setting and production by organisms such ff cient at accumulating nitrogen (N) in the soil when production ) are higher than those of terrestrial forests ( ffi 1 − yr 2 − (Reducing Emissions from Deforestation and Degradation) have + W). The LEBR comprises an area of 144 868 ha. The Reserve has five coastal 0 26 ◦ In this study, we assessed whole ecosystem C stocks (trees, soil and downed wood) Wetlands, such as mangroves and marshes, are within the most e Wetlands are e The C accumulated in wetlands originates from organic material such as leaves, in height, which are believed to be the tallest of the country (Tovilla et al., 2007). The 92 connected to seven riverof systems. freshwater The LEBR and isforests. estuarine characterized The by wetlands LEBR large including areas hasforest mangroves, one is marsh dominated of by and the trees peatswamp most of extense areas of the region; the environmental problems (Gallowayto et aquatic al., systems, 2004).activity, but rainfall, Wetlands the runo can potential decrease for N N accumulation inputs varies with anthropogenic estimated soil C sequestrationand rates C of mangroves content.ecosystem on We the C predict basis stocks of thatmangroves and soil the with accumulation high riverine strong Csequestration wetlands riverine rates sequestration within influence compared rates. the have toinfluence. We lowland LEBR higher also mangroves, have soil which predict large N, have that a C upland stronger stocks marine and2 soil C Methodology 2.1 Study site The LEBR is located in Chiapas, in the south Pacific coast of Mexico (14 as cyanobacteria and algaeaccumulation (Alongi, 2009; also Reef increases et withCompton, al., foliage 2010; 2003; cover Adame Liao and et etstocks al., wood al., can 2012a). increase 2007), N with (e.g. and increments Hooker given in and that C (Yimer C et and al.,and N 2006). soil cycles N interact stocks ofLa closely, N riverine Encrucijada wetlands Biosphere (mangroves, marshes Reserve and (LEBR) peat swamps) in within the Pacific coast of Mexico. We also programs, accurate estimates of C2011). stocks and sequestration rates are needed (Alongi, 2011). To prevent theas large REDD emissions thatbeen result proposed. from In order to loss, target programs coastal wetlands such within REDD for C processing andecosystems sequestration, storing (Chmura up et toexample, al., three mangroves times 2003; in more Donato C the et than Caribbean terrestrial al., can 2011; store McLeod up et to al., 987 2011). Mgha For inorganic C) (Donato et al.,those 2011; of Adame tropical et al., and 2013). temperate These values forests typically ( exceed while mangroves in the Indo-Pacific store up to 2203 Mgha C sequestration rates of coastal wetlands (210 gCm (20–30 gCm a wetland depends onand its geomorphological setting, flushing, which andorganic influences therefore patterns matter of the (Eyre, tidal importmost 1993; extensive, and Adame and are export et characterized of bymaterial al., mature suspended and 2010). forests sediment with suspended Riverine large and sedimentsuggested amounts wetlands that inputs of are organic riverine (Woodrofe, wetlands one 1992; have one Eyre, of of 1993). the the It largest C has stocks been (Ewel et al., 1998). twigs and rootstidal produced and in riverbacteria situ, flushing. and fungi, The but or buried organic also within2001; the material from Kristensen soil is (Holguin allochtonous et et either al., al., C 2001; degraded Middleton 2008). imported and and The McKee, through processed amount by of C produced and sequestered within exceeds N demand (e.g.anthropogenic Rivera derived Monroy N et and al., ine 1995). In the past century, increased (Chmura et al., 2003; McLeod et al., 2011). 5 5 25 20 15 10 10 25 20 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , ). R D man ff 1.027; = t R. mangle R. mangle ). Aboveground erent ( ff 32 cm for = Max ) were calculated with the formula of Avicennia germinans C and a mean annual maximum of ◦ and a marsh dominated by the grass 1020 1019 P. aquatica and 42 cm for 1 to 1, where negative values indicated areas without ) per site using methodologies described in Kau − 2 Pachira aquatica (Fig. 1; Table 1). We measured whole-ecosystem C stocks in 20 cm quadrants within each of the 6 plots. The wet mass was × was measured at the main branch, above the highest prop root ( erence Index (NDVI) was obtained with ERDAS Imagine. 0.30). We chose the formula by Fromard et al. (1998) because it included ff = Laguncularia racemosa p C, with a mean annual minimum of 19.2 C; mean annual precipitation is 1567 mm (Sistema Meteorológico Nacional – ◦ ◦ R. harrisonii 2284; = The climate of the LEBR is warm, sub humid with most precipitation occurring in Tree biomass was calculated using allometric equations (Table 2). We used the 36.5 Comisión Nacional del Agua, station No. 7320, 1951–2010). 2.2 Site stratification To determine a criteriaSPOT for 5 stratification satellite ofand images the the mangroves with Universal ofNormalized geographical, the Transverse Di geometric area, Mercator we andThe projection used NDVI radiometric system. values two ranged correction, From from each image, the vegetation, values closecondition, to and zero values2006). close indicated NDVI to senescent values 1, vegetation2013) were indicated and or extracted classified green from in according orwas the divided to a in healthy Ruiz-Luna three mangrove classes: stressed vegetation et the coveragebroadly most (Chuvieco, al. map vigorous corresponded (2010). vegetation to was (CONABIO, The Class upland I mangroves.III mangrove (9253 The ha), (11 vegetation 467 which least ha), vigorous which vegetationII broadly was corresponded (6757 Class ha) to had lowland intermediate orvegetation vigor estuarine values were mangroves; of also Class vegetation foundagricultural vigor in areas). (Fig. forests close 1). to Mangroves human with settlements low (e.g. town2.3 and Field and laboratory analyses Sampling was conducted duringstocks December and 2012, soil where ecosystem C C sequestration stocks, rates soil were N measured. We sampled 9 sites: seven trees with a DBH range similar to those found in LEBR (DBH df 2.3.1 Biomass and C stock withinForest trees structure and was shrubs measured atthe each plot diameter through at measurements of 1.3 the m species height and (DBH) of all trees. The diameter of trees of et al. (2014a). Thea plots perpendicular were direction establishedwithin from 25 trees m the and apart water shrubs,stocks along edge. and downed a interstitial At salinity. wood 125 To estimate m each and Ca transect plot, sequestration the natural rates set horizon we soil in marker mangroves, in sampled to profile. weexplained calculate used We C below. soil also C stocks accumulation. sampled The soil detailed methodology N is and Aboveground biomass in the marsh communities waswithin determined through two plant harvest 20cm 9.6 cm for biomass of trees from the peat swamp ( mangrove forests (3 forswamp Class forest I, dominated 2 for by Typha Class dominguensis II and 2 for Class III mangroves), a peat determined in the fieldoven-dried and to then determine a its dry subsample weight. was collected fromformula each by quadrant Fromard et and al. (1998),which which was is obtained for a mangrovesmangroves of location with French Guiana, a with tropical similaret hot al. humid characteristics (1998) climate). than and WeMexico. Day compared those et the The found al. formulas results (1987), of in the Fromard using LEBR latter both obtained (riverine from formulas mangroves were in Campeche, not significantly di six plots (radius of 7 m; 154 m the summer months (June–October).28.2 The mean annual temperature of the region is mangroves of LEBRE support a high(UNESCO, biodiversity, as 2013). well as fisheries and tourist activity 5 5 25 10 15 20 10 15 20 25 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ◦ man ff from the 7.5 cm in ff < o ◦ was determined 100 cm. Soil depth > 2.5 cm but > P. aquatica man et al., 2014a): Status 1 – man et al., 1995, 2011) ff man et al., 2014a). Four 14 m ff ff 7.5 cm in diameter (hereafter “large” > 1022 1021 man et al., 1995). The core was systematically erent sizes (small, large-sound, and large-rotten) ff ff the direction of the main transect, the other three were established 90 Standing dead trees were also included in the tree C stocks estimations. Each dead 60 pieces of down wood of di ff and calculated their specificUsing the gravity specific as gravity the forconverted each to oven-dried group C weight using of a divided wood conversion debris, by factor biomass of its was 0.502.3.3 volume. calculated (Kau and Soil C and N Soil samples forplot bulk using a densityattached peat and to auger nutrient adivided consisting concentration cross into of depth handle were intervals a (Kau of collected semi-cylindrical 0–15, at 15–30, chamber 30–50, each of 50–100 and 6.4 cm-radius was measured usingSamples a of steel-2 m a rodmass known that to volume determine was were bulk insertedwith collected density. Samples in Hydrochloric in were the acid the sievedbefore ground (HCl) field and analyses. homogenized at and to Concentration and of each then eliminate treated organicElemental plot. dried the C Combustion System and to inorganic 4010 N constant carbon (CA, were USA). determined portion using (carbonates) a2.3.4 Costech Soil C sequestration rates We estimated C sequestration rates asTo the date amount of the C soilhorizon accumulated in that cores, the was soil we profile. clearlyof used identified the a in volcano Santa natural all Maria’svolcano the marker eruptions eruption cores. of in that the This 1902 20th consisted that Century ashand (Volcanic represented of horizon Self, Explosivity one Index is a 1983). of of the 6 As the volcanicdate out remaining four a of ashes ash largest 7, result can Williams be of established the in eruption, the a Mexican Pacific recognizable coast, plinian northwest deposit of of the volcano. known debris) was measured. Largeand downed rotten. wood Wood debris was was separated consideredbroke in rotten apart if two when it categories: visually kicked. appeared≈ sound To decomposed determine and specific gravity of downed wood we collected transects were established ato the center of each plot: the first one established at 45 tree was assigned to one of three decay status (Kau with the equation of Cairnswas et calculated al. using (1997) the for tropical formulaof forests. by Chave Belowground Komiyama et root et biomass al. al.by (2005) (2009) a and (Table wood 2). factor density Treeof of values C 0.48 marshes was for calculated waset from aboveground calculated al., biomass and 2014a). using by 0.39 multiplying a for factor belowground of biomass; C 0.45 content of thedead total trees without biomassStatus leaves, (Kau 3 Status – 2 dead –tree trees dead without status primary trees was or withoutprovided calculated secondary by secondary Fromard branches. as et branches, The al. a and as biomass (1998). for For percentage the dead each of trees total oftotal. the Status dry The 1, total biomass biomass biomass of was biomass minusbiomass trees calculated using of of the Status leaves the biomass 2 (2.8 % values of was of of the calculated the total). leaves, as Finally, total) the the equivalent plus totalthe biomass to main biomass secondary of stem, minus trees branches 2.8 which % of the (equivalent is Status to of equivalent 3 to 18.7 was % the 76.6 calculated %2.3.2 as of the the biomass total. of Downed wood The mass of dead and downed(Van wood was Wagner, calculated with 1968) the planar adapted intersect technique for mangroves (Kau previous transect. The diameter oftwigs, any branches, downed, dead prop woody roots materialwas or measured. (fallen/detached Along stems the of last 5 trees m and of the shrubs) transect, wood intersecting debris each transect van Breugel et al. (2011), while belowground biomass of diameter (hereafter “small” debris)of was the counted. transect From (12 the m second in meter total), to wood the end debris 5 5 10 15 20 25 10 15 20 25 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , é 1 ff − ect and ff erent ff er from fires ff P. aquatica ect and site as the ff (Table 1). with lesser contributions (in sites Panzacola and erences of biomass and C 20–40 m in height) of mean ff ∼ R. mangle R. harrisonii T. dominguensis erences in soil C and N concentrations standard error. ff ± ect of the model. ANOVA models were also ff 1023 1024 ) with tall trees ( 1 − and few trees of ect of the model. Di ff . All the sampling sites were characterized by relatively low tree erences of biomass and C stocks among sites, where plot (nested in ff erences were found, pair-wise comparisons were explored using Sche (Chmura et al., 2003). We could not measure soil C sequestration rates of ff 1 ect of the model. Normality was assessed using Shapiro-Wilk tests. When − ff A. germinans yr 2 − A. germinans, L. racemosa marsh and peat swamp forests, as these vegetation types frequently su 2.4 Scaling up We multiplied mean ecosystem Carea stocks of and each sequestration rates vegetationwe times type used the to the estimated obtainabove), area which the on broadly C the represented budgetmarsh, basis a for we range of determined the from our its upland LBRE. areafrom forest to For the on downland classification coastal the mangroves, mangroves. vegetation (Classes basis For map ofas I, of the from the II “other Pacific auxiliary wetlands” and south categoryis region cartographic II; obtained (CONABIO, (SERIE likely see 2013), as IV; that well INEGI,underestimated, the 2012) as area and the of our(“popales”) marsh the field with area experience. unknown marsh It includesfor C – the stocks. waterholes LEBR, and and The and thus we areasampling could inundated was its only of conducted identify vegetation (844 C peat ha; from Fig. the swampwas stock 1). satellite underestimated. forest Therefore, – images the is the C was forest stock not where of slightly available our peat over swamp forests or and thus, have confounding ash horizons. 2.3.5 Interstitial salinity Salinity was measured with anfrom YSI-30 water multiprobe extracted sensor from (YSI, 30acrylic cm Xylem tube deep. Inc. (McKee The Ohio, et water USA) al., was 1988). obtained with a syringe and an 2.5 Statistical analyses Analysis of Variance (ANOVA) was performed to test di used to test di site) was the random e stocks among mangrove classes (Classes I,and II plot and (nested III), in where site) class was was the the random fixed e e random e significant di post-hoc tests. Analyses were performedJolla, using CA, Prism USA) ver 6.0 andthe (GraphPad SPSS Software, manuscript, La Statistics data are (version reported 20, as IBM, mean New York, USA). Throughout 3 Results 3.1 Forest structure Mangroves of the LEBR were dominated by trees of species, density forests (1213–5370 treesha by depth were also tested with ANOVA, with depth as the fixed e of Teculapa). Only one of our study sites – Las Palmas – was dominated by a di We estimated soil Cdividing the sequestration depth within of the eachmultiplying ash it plot horizon by by of bulk years since density sixgCm the of of volcano C eruption our content. occurred Soil mangrove and C sites sequestration by rates are expressed in DBH of 8–11 cm (Table 1). The peat swamp forest was dominated by had a similar structure than that of mangroves with a tree density of 2469 treesha trees of updominated to by tall 22 grasses m (2–3 m in in height) height of and mean DBH of 14.5 cm. Finally, the marsh was 5 5 10 15 15 20 10 20 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | . . ± ± > 1 1 − − and 67.8 1 − ± 0.07 % ); large 1 0.0001). ± and was − ) similar to 1 1 − p < − 73.5 MgCha 72.6 MgCha ± ± ) while the marsh for Class III), but 1 15.7 MgCha 0.05). The swamp 0.8 Mgha 1 − 8.147; ± ± − ) and largest at Las 1.3 %). Surface soil N = 1 1.7 Mgha p < ± − ± 44.4 MgCha 7,47 ) than lowland mangroves ± F 1 , with a mean biomass of 0.02). The peat swamp had − 1 )( − 1.826; = 1 . 6.8 MgCha − p 1 = 3.8 Mgha ± − 102.7 MgCha ± 2.8 %, with a minimum value at Las 7,47 ± ± F ) (Table 3). 0.0028), mainly due to large amounts 9.42; 1 6.8 MgCha = − )( = 1 p ± ) and large rotten wood comprised 34.4 % − 1 2,5 − F 27.0 MgCha 1025 1026 39.0 MgCha ± 7.8 % (between 18.4 % in Las Palmas to 87.2 % 0.52 % and a wide range between 0.32 6.86; ± 0.77 %, respectively) and the highest N stock of ± ± = 13.4 and 93.9 ± , respectively) (Table 5). There was a decreasing 52.5 Mgha 1 ± ± − 2,39 304 MgCha ). The mean C stock within downed wood was 29.4 ). The marsh had surface soil C and N concentrations F 13.9 Mgha 1 1 < − − ± 0.28 % de N) and N stocks (18.5 ) compared to lowland mangroves (Class II and III with 15.3 ) had significantly higher C stocks in trees compared to for Class II and 277.5 ± 0.06 % (Santa Chila). Soil C values increased with depth at 1 1 , respectively. Lowest biomass (above and belowground) was − − 1 3.2 Mgha 1 ± − − ± , respectively, Table 3). 5.5 Mgha (Table 4). Down land mangroves (Class III) had the highest biomass 1 15.3 % and 1.05 15.2 Mgha − ± 1 ± ± 172.6 and 268.7 − ± 0.5 Mgha 1.2 %) and a maximum value at Santa Chila (29.1 . ± 1 and 12.3 22.9 Mgha 620 MgCha ± − 1 ± > − 169.6 MgCha 4.0 % C and 1.10 ± 5.8 MgCha 26.0 Mgha ± ± ± erences were not significant due to large soil C values at Esterillo (Class II). When Surface soil C content (%) had a mean of 17.4 Mean C stock in mangrove trees at LEBR was 215.0 The peat swamp forest had soil surface C and N concentrations similar to those of ff (Las Palmas) and 1.30 (Class I; 26.4 Panzacola, remained similar in depth inat Teculapa and Paistalon the and decreased rest in depth Zacapulco of and the Las Palmas) lowland (Table 5). mangroves Finally, (Class soil N II was Esterillo, higher Santa in upland Chila, mangroves and Class III (%) was variable, with a mean of 2.70 1.6 Mgha gradient of C and Nmangroves stocks (Las from the Palmas, Santa most Chila upland mangrove and site Zacopulco, (Panzacola) Table 5). to lowland mangroves (16.3 all wetlands (40.4 (15.6 Palmas (6.2 Mean C stocks of mangroves in LEBR had a mean of 784.5 those of mangroves and peat swamp forests (Table 5). 3.5 Ecosystem C stocks (562.6 Upland mangroves had higher soil C (620.4 di had the lowest soil C stock with 298.3 Lowest C stocks were measured in the most lowland mangroves, i.e. those with Esterillo was nothigher included soil in C the than downland analysis, mangroves upland ( mangroves exhibited significantly 59.4 and C stocks of downed woodof ( downed wood at Zacapulco (102.4 in Esterillo) of the total C stock. Most of the soil C was organic, with a contribution 86 % for all sites (Table 5). Mangrove soil C stock had a mean of 505.9 a similar soil C stock than upland mangroves (620.4 3.3 Downed wood C Downed wood wasconsiderable insignificant in in some marshhad mangrove and sites. a peat The swamp wide amount forests, range although of within it downed sites, was wood from in mangroves 11 to 205 Mgha Small downed wood comprisedsound 10.2 % wood the of 55.4 the % (33.0 of total the biomass total (6.0 (20.4 3.7 MgCha 3.4 Soil C Soil C in mangroves accounted for 65 Santa Chila and Zacapulco ( 38.2 Palmas (706.6 similar throughout uplandEsterillo and ( lowland forests. However, the siteforest Las and marsh Palmas vegetation had and lower vegetation C stocks (95.1 3.2 Tree biomass and C Mean tree aboveground and belowground biomassmeasured of mangroves at was Santa 421.1 Chila (198.8 and 154.3 5 5 15 10 20 25 15 25 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | . ; ) ± 1 1 1 − − − yr , with 1 1 − ; Jones , IPCC, − 1 1 and was . − − man et al., 1 1 − ff − yr 176 MgCha 1 83.4 MgCha − ± ± 0.0 MgCha ± 73.5 MgCha erences among sites 400 MgCha ± ff ). Finally, marshes had 1 < 38.3 MgCha − ). Largest C stocks were 0.2 Mgha ± 1 (Fig. 2, Table 6). ± − 1 . The peat swamp forest had − (Table 8). erences in the geomorphological ff ) and lowest at Las Palmas (640.9 63.6 MgCha 1 − ± e, 1992). As a result, these mangroves 18.6 MgCha ff ± A. germinans 38.3 MgCha ered according to geomorphology; higher N ± ff A. germinans 1028 1027 , Nguyen et al., 2014) and the Dominican Republic 1 erences in N content have also been associated to − 150.3 MgCha ff ± , values similar to those measured in riverine mangroves 1 − erences in C stocks have been related to geomorphological and man et al., 2014b). Our mangrove C stock estimations were ff ff 57.2 MgCha erences in soil C with depth according to geomorphological setting, ff ± ), the forest dominated by 1 , Kau 1 − − In general, upland mangroves within LEBR had higher C stocks than lowland The N stocks within mangroves also di We also found di et al., 2014). Peat swamp forests had similar large C stocks with 722.2 measured at Esterillo (1114.9 mangroves or those closer to the mouth, however di 151.9 MgCha Lowest C stocks were measured at the marshes with 336.5 highest marine influence (Class III; 659.5 for wetlands in LEBR was estimated to be around3.7 32.5 millions of Soil Mg. C sequestration rates Mean soil C sequestration rates in mangroves was 1.3 are likely to beComparatively, young upland mangroves forests are likely with to low haveproductive higher , C low and stocks soil as have they C high areallows mature, and soil C thus, to C low be as C buried. they stocks. grow in a relatively stable environment that 3.6 C stocks of LEBR By extrapolating the mangrovedeteriorated area forests, by we classes estimated20.9 (Classes the millions of I, C Mg. stock II Theof C of and Mg stock and mangrove III) of that marshes forest and of was of peat estimated excluding swamps LEBR to of be at to close least be to 0.6 11.0 millions of of millions Mg (Table 7). The total C stock with an increase in soil Cat with the depth at most the lowland most mangroves. upland mangroves We and suggest a that decrease in di C similar C stocks than mangrove forests (722.2 were also notable. Di biological processes. For example, the location ofsediment a and forest its could burial influence the rate, amountlandscapes as of depositional (Doetterl landscapes et have al., morehave 2012). soil higher C Also, C than mangroves stocks eroding growing compared2011). closer to Forest those to development closer the and to water forest thestocks edge landward productivity in edge also taller (Kau and influence more C productive stocks, forests with (Adame larger et al., 2012b). and biological forces acting inthe riverine C mangroves distribution explain within the the variation sedimentof in column. the C Downland mangroves stocks estuary close and to aredirectly the exposed mouth struck to frequent by changes tropical in storms hydrology, sedimentology (Woodroo and are stocks were measured inreceive upland high compared N to inputslowland downland mangroves due probably forests. to receive Upland lower agricultural Nnutrients mangroves activity loads than as in oceanic riverine the water water. hasmicrobial catchment Di usually (UNESCO, activity lower 2014); suchflushing as in bacteria mangroves and (Alongi, 1988). protozoans, Higher which nitrification are and in denitrification and turn lower linked to tidal the lowest ecosystem C stock with 336.5 similar among upland and lowland mangroves. Lowest values (0.4 4 Discussion The riverine wetlands measureddouble in than this study those had measured large in C stocks, terrestrial which forests were almost (typically were measured in the site dominated by in Vietnam (762.2 2003). Mangroves hada a maximum of mean 1115 MgCha ecosystem C stock of 784.5 higher than those in karstic settings in Yucatan, Mexico (mean of 663 (853 MgCha Adame et al., 2013) and those in Northwest Madagascar (367–593 MgCha 5 5 20 10 10 15 25 15 20 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | . ) 2 2 1 A. − yr 1 − R. mangle (Las Palmas) setting emissions within ff ) and highest in Terminos 1 − A. germinans ) than wood density of 3 − ), and are similar to those measured 1 − ; Lovelock et al., 2014). Long-term soil 1 yr − 1 − yr 1030 1029 1 − emissions to the atmosphere. Another example is to 2 2.6 millions tons of C or 20.6 % of the emissions of the cult to obtain, thus the values obtained in this study ffi ∼ is lower (0.67–0.99 gcm M. F. Adame designed the project, leaded the field campaign, collected ), which dominated all other sites. Wood density is a major predictor , which is equivalent to the annual emissions of approximately 9143 , respectively. Every year between 500 and 4500 ha of marshes are 3 1 erences in C and N stocks between upland and downland mangroves, 2 − − erent. This forest had the highest tree biomass, lowest soil C and ff ff A. germinans. can be due to the lower C wood content that is buried in the soil. Wood A. germinans ective way to reduce C emissions (Siikamäki et al., 2012), while protecting ff The C stocks and sequestration values shown in this study can help provide Besides the di Ecosystem C stocks were associated to soil C stocks, as this component contributes Mangroves in riverine deltas are the most extensive and highly developed forests incentives intothroughout the similar reforestation wetlandare and ecosystems. very conservation For susceptiblecommunication, projects example, 2013). to With marsh of firethat the and damage every this C hectare swamp stocks during of reserve calculated marsh forests the or in and peat dry this swamp study, forest season we that (L. is can burned estimate Castro, emits 1237 personal ton CO Degradation of wetlands inupriver the dredging, region fires, due hydrological topotential modifications, C increased and storage sediment of illegal loads these harvesting derived wetlands. threaten from the use the C stocks provided in this study to negotiate for o The wetlands of LEBR store about 32.5 MtonC, which is equivalent to 119 Mton CO and 2650 ton CO burned within the reserve (L.an Castro, annual personal mean communication, emission 2013), of state which of results Chiapas in (based onused emissions to reported emphasize by the IEA, importanceits 2001). of This large managing information C can fires stocks be in and the avoid LEBR CO in orderthe to maintain country orimport abroad. C For credits from instance,et forests al., California Chiapas, 2011). the USA, To include state mangrovesa has and where cost-e other signed this wetlands study an in takes similar agreement agreements place could to (Morris be the data and wrote theand manuscript. N. participated S. in Santini designedcampaign, writing the collected and project, performed data preparation datageographical and analysis of information contributed the system to manuscript. datathe the C. manuscript. and Tovilla manuscript. L. map, participated Castro A. performedmanuscript. in participated Vázquez-Lule data in field prepared field analysis the campaign, and collected contributed data to and contributed to the biodiversity (Adame et al., 2014)provide. and the various ecosystem servicesAuthor that contributions. mangroves , Campeche, Mexico (6.5 MgCha N fixation rates could furtherremains explain to low be N tested. stocks in lowland mangroves; however, this it stands out that the mangrove forest dominated by are within thewith lowest range values of in Rookery those Bay, reported Florida (0.2 in MgCha the review by Chmura et al. (2003), to most ofsoil C the sequestration C rates in measured in mangroves mangroves (Donato of LEBR et (0.4–1.8 MgCha al., 2011; Adame et al., 2013). The was notably di germinans (0.810–1.05 gcm of stored Csoil in (Flores wood and biomass Coomes,dominated and by 2011), could and thus, explain the the low low C values stocks of in C the buried mangrove forest in the lowest C sequestration rates measured in thisdensity study. of Lowest C stocks in soils of in Moreton Bay, Australia (0.8 MgCha (Woodroofe, 1992). Thus,riverine the wetlands, results which inworld. are Additionally, this likely the LEBR to study isC be contribute an monitoring important one to in location of Mexico, for the which the C C studies includes most as a budgets C it flux is of rich tower the for ecosystems base monitoring of in daily the C variations. are valuable forexample, we estimations can estimate ofis that 27 762 C MgCyr the sequestration sequestrationMexicans rate (using of emissions rates by the country of mangrove from soil IEA, mangrove 2011). of LEBR forests. For C sequestration rates are di 5 5 15 20 25 30 10 15 10 25 20 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | erent ective ff ff th University and ffi erent kinds of mangroves forests provide di 1032 1031 ff man, J. B., Medina, I., Gamboa, J. N., Torres, O., Caamal, J. P., We want to thank the Australian Rivers Institute at Gri ff man, J. B., Murdiyarso, D., Kurnianto, S., Stidham, M., and Kanninen, M.: ff goods and services, Glob. Ecol. Biogeogr. Lett., 7, 83–94,assessments, 1998. Methods in and Evolution, 2, 214–220, 2011. ground biomass andOecologia, dynamics 115, of 38–53, 1998. mangrove ecosystems: new data from French Guiana, Microb. Ecol., 15, 59–79, 1988. 331–349, 2002. uncertainties of sequestration potential, Environ. Sci. Policy., 14, 462–470, 2011. a worldwide wood economics spectrum, Ecol. Lett., 12, 351–366,tidal, 2009. saline wetland soils,2003 Global Biogeochem. Cy., 17, 1111,, doi:10.1029/2002GB001917 the world’s upland forests, Oecologia, 111, 1–11, 1997. de suelo y vegetación(2005), escala de 1 la : 50 zona 000. CONABIO, costera Mexico, 2013. asociada ageomorphic los controls manglares, on Región soilBiol., Pacífico organic 18, Sur C 2218–2232, 2012. pool composition and C stabilization,Mangroves Glob. among Change the2011. most carbon-rich forests in the tropics,and Nat. nearshore Geosci., coastal zone, 4, Aust. J. 293–297, Mar. Fresh. Res., 44, 845–866, 1993. areas for restoration offirst, ecosystem 2014. services, Conserv. Biol., doi:10.1111/cobi.12391, online mangrove forests along a86, 21–30, gradient 2010. of geomorphological settings, Estuar. Coast.extensive Shelf. cyanobacterial mats S., in an63, arid 1323–1650, subtropical 2012a. coastal wetland, Mar. Freshwater. Res., Terrestrial-marine connectivity: patternssediments determined of by terrestrial analysis1492–1502, 2012b. of soil glomalin related carbon soil deposition protein, Limnol. inReza, Oceanogr., coastal 57, M.,within and the Herrera-Silveira,doi:10.1371/journal.pone.0056569, 2013. karstic J. landscape A.: Carbon of stocks the of Mexican tropical Caribbean, coastal PLoS wetlands One, 8, e56569, Flores, O. and Comes, D.Fromard, F., Puig, A.: H., Mougin, Estimating E., Marty, G., the Betoulle, J. L., wood and Cadamuro, density L.: Structure, above- of species for carbon stock Alongi, D. M.: Bacterial productivity andAlongi, microbial D. biomass M.: Present in state tropical and future mangrove of sediments, theAlongi, world’s D. mangrove M.: forests, The Environ. EnergeticsAlongi, Conserv., of 29, Mangrove D. Forests, Springer, M.: Dordrecht, 2009. Carbon paymentsChave, J., for Coomes, D., mangrove Jansen, S., conservation: Lewis, ecosystem S.Chmura, L., G. constraints Swenson, L., N. Anisfeld, and G., S. C., and Cahoon, Zanne, D. R., A. and E.: Lynch, Towards J. C.: Global carbonChuvieco, sequestration S. in E.: Teledetección ambiental,Cairns, España, Ariel, M. 2006. A., Brown, S., Helmer, E. H.,CONABIO and (Comisión Baumgardner, Nacional G. para el A.: Conocimiento Root y biomass Uso allocation de in la Biodiversdiad). Mapa deDoetterl, uso S., Six, J., Wesemael, B. V., and Van Oost, K.: Carbon cycling in erodingDonato, landscapes: D. C., Kau Eyre, B.: Nutrients in the sediments of aEwel, K. tropical C., north-eastern Twilley, R. R., Australian and estuary, Ong, catchment J. E.: Di Adame, M. F., Neil, D., Wright, S. F., and Lovelock, C.Adame, E.: M. F., Sedimentation Reef, within R., and Granham, A., among Holmes, G., and Lovelock, C.Adame, E.: M. Nutrient exchange F., of Wright, S. F., Grinham, A., Lobb, K., Reymond, C.Adame, E., and M. Lovelock, F., C. Kau E.: Acknowledgements. to the Mexican Fund forla the Naturaleza Conservation (FMCN), of A.C.) Nature and (Fondoby the Mexicano USDA the para Forest la Service. US Conservación Funding Agency de quantification for the of for study wetland International was carbon provided Development stocks.their We contributions (USAID/Mexico) wish of as data to collection part thankCommission in the of the for participants field. a of Natural We workshop this areFrausto Protected workshop grateful Leyva on for and for Areas field Richard support (CONANP) BirdseyNavy, to for Mexico and the logistic and National to support. the Finally, Vanessa Nationalwhich we Valdez, Comission thank contributed the for with Juan Secretariat Knowledge their Manuel of andterm data the Use monitoring”. from the of project Biodiversity “Mexican (CONABIO) mangroves, current state and long References Adame, M. F., Hermoso, V., Perhans, K., Lovelock, C. E., Herrera, J. A.: Selecting cost-e 5 5 30 10 25 15 20 15 20 25 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | and Spartina , Front. Ecol. 2 Rhizophora mangle 1034 1033 sets in California climate policy, Resources for the ff , Am. J. Bot., 75, 1352–1359, 1988. enhanced ecosystem carbon and nitrogen stocks in the Yangtze Estuary, China, man, J. B., Cummings, D. L., Ward, D. E., and Babbitt, R.:man, Fire J. in the B., Brazilian Heider, Amazon: C., Cole, T. G.,man, B. Dwire, J., K. Donato, A., D. C., and and Donato, Adame, D. M. C.: F.: Ecosystem Protocols for carbon the measurement, monitoring man, J. B., Heider, C., Norfolk, J., Payton, F.: Carbon stocks of intact mangroves and ff ff ff ff alternifolia Ecosystems, 10, 1351–1361, 2007. subsurface processes in keepingBay, Queensland, pace Australia, with Ecosystems, sea 14, level 745–757, 2011. rise in intertidalContemporary wetlands of rates Moreton mangroves of and saltmarshes of carbon South771, East 2014. sequestration Queensland, Australia, and Estuar. Coast., 37, vertical 763– Schlesinger, accretion W. H., of andunderstanding sediments Silliman, of B. the in R.: role of A vegetated blueprint coastal for blue in carbon: sequestering toward CO an improved Avicennia germinans formation in Belizean island forests, J. Ecol., 89, 818–828,challenges 2001. of internationalFuture, forests Issue Brief, o 11–12, 2011. forest in Mui Ca Mau National Park, Vietnam, CATENA, 121,patterns, 119–126, extent 2014. and current723, condition 2010. of Northwest Mexico mangroves, Wetlands, 30, 717– 2010. mangrove ecosystems: a review, Aquat. Bot., 89, 201–219, 2008. Environ., 9, 552–560, 2011. concentrations and redox potentials near the aerial roots of Asner, G. P., Cleveland,Porter, C. J. C., H., Green, Townsend,future, A. P. Biogeochemistry, A., 70, R., 153–226, Holland, and 2004. E. Vorosmarty, C. A., J.: Karl, Nitrogenproductivity, D. conservation, cycles: M., past, and Michaels, presentFertil. rehabilitation A. Soils, and of 33, F., 265–278, mangrove 2001. ecosystems: an overview,first Biol. century after agricultural abandonment, Ecol. Appl., 13, 299–313,suelo 2003. y vegetación serie IV, escala 1 : 250 000, INEGI,Land-Use Mexico Change, City, Mexico, and 2012. Forestry, editedKruger, D., by: Pipatti, Penman, R., J., Buendia, Gytarksy,for L., M., Global Miwa, Environmental Hiraishi, K., Strategies, T., Ngara, Kanagawa, Krug, T., Japan, Tanabe, T., K., 2003. and Wagner, F., Institute Ecological variability and carbonMadagascar, stock Forests, estimates 5, of 177–205, mangrove 2014. ecosystems in Northwestern 1. Biomass, nutrient pools, and1995. losses in slashed primary forests, Oecologia, 104, 397–408, stocks of Micronesian mangrove forests, Wetlands, 31, 343–352, 2011. and reporting of structure,la biomass and medición, carbon monitoreo stockslos in y manglares,), mangrove forests reporte Working (ProtocoloIndonesia, paper para de 2014a. 117, la Center estructura, for biomasa International ycarbon Forestry emissions reservas Research, arising de from Bogor, 518–527, their carbono 2014b. conversion de in the Dominican Republic, Ecol.weight Appl., of 24, mangroves, J. Trop. Ecol., 21, 471–477, 2005. Kau Komiyama, A., Poungparn, S., and Kato, S.: Common allometric equations for estimate the tree Lovelock, C. E., Bennion, V., Grinham, A., andLovelock, C. Cahoon, E., D. Adame, R.: M. F., The Bennion, role V., Hayes, of M., O’Mara, surface J., and Reef, R.,McLeod, Santini, E., N. Chmura, G. S.: L., Bouillon, S., Salm, R., Bjork, M., Duarte, C. M., Lovelock, C. E., Middleton, B. A. and McKee, K. L.: Degradation ofMorris, mangrove tissues D. and F., implications for Richardson, peat N., Riddle, A.:Nguyen, Importing T. T., Dung, climate L. V., Nhuan, mitigation: M. T., and the Omori,Ruiz-Luna, K.: potential Carbon storage and A., of a tropical Cervantes-Escobar, mangrove A., and Berlanga-Robles,Reef, R., Feller, C.: I. C., and Assessing Lovelock, C. E.: Nutrition distribution of mangroves, Tree Physiol., 30, 1148–1160, Kristensen, E., Bouillon, S., Dittmar,Liao, C., T., Luo, Y., and Jiang, L., Marchand, Zhou, X., C.: Wu, X., Organic Fang, C., Chen, carbon J., and dynamics Li, B.: in Invasion of McKee, K. L., Mendelssohn, I. A., and Hester, M. W.: Reexamination of pore water sulfide Galloway, J. N., Dentener, F. J., Capone, D. G., Boyer, E. W., Howarth, R. W., Seitzinger, S. P., Holguin, G., Vazquez, P., and Bashan, Y.:Hooker, The T. D., role Compton, J. of E.: Forest ecosystem sediment carbon microorganisms andINEGI nitrogen (Instituto in Nacional accumulation de during Estadística the the y Geografía) Conjunto de datosIPCC, vectoriales Intergovernmental de Panel uso del on Climate Change: Good Practice Guidance for Land Use, Jones, T. G., Ratsimba, H. R.,Kau Ravaoarinorotsihoarana, L., Cripps, G., andKau Bey, A.: Kau 5 5 20 25 30 10 15 10 15 20 25 30 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ected by ff (100 %) 23, last ac- (12.3 %) (83.2 %) (25.1 %) (10.6 %) (13.9 %) + (96.9 %) (94.5 %) (100 %) (97.5 %) (87.7 %) (87.6 %) (68.9 %) not assessed. = MEX = R. mangle R. mangle T. dominguensis R. mangle R. mangle A. germinans R. mangle L. racemosa A. germinans L. racemosa P. aquatica R. mangle A. germinans diameter at breast height. = all&code = emissions from forest loss, Nat. 2 ) Salinity (ppt) Dominant species 1 − maximum; DBH = 1036 1035 Paistalon 25 9.9 (0.9) 2035 (134) n.a. Teculapa 30 7.5 (1.0) 2761 (398) 19.3 (5.3) Las Palmas 28 7.9 (0.4) 5370 (388) 28.9 (0.6) Sta Chila 22 9.9 (0.6) 2371 (157) 37.5 (0.6) Characteristics of sampling sites within La Encrucijada Biosphere Reserve (LEBR). e, C.: Mangrove sediments and geomorphology, in: Tropical Mangrove Ecosystems, ff Marsh 3 – – n.a. Mangroves SiteIII Max height (m) DBH Panzacola (cm) Tree density (treesha III Esterillo 40 10.5Peat (1.1) swamp Zacapulco n.a. 1213 (278) 8.8 n.a. (1.0) 8.8 (0.8) 3346 (148) 22 n.a. 1765 (274) 37.6 14.5 (5.3) (0.9) 7.6 (0.4) 2469 (301) 0.0 (0.0) Geosci., 2, 737–738, 2009. Guatemala, J. Volcanol. Geoth. Res., 16, 33–56, 1983. John Wiley & Sons, American Geophysical Union, Washington DC, USA,topographic 2010. aspect and vegetation344, in 2006. the Bale Mountains, Ethiopia, Geoderma, 135, 335– of nitrogen and sedimentCoast in Shelf S., a 40, fringe 139–160, mangrove 1995. forest in Terminoscarbon Lagoon, dioxide Mexico, emissions Estuar. from, mangrovedoi:10.1073/pnas.1200519109 loss, 2012. P. Natl. Acad. Sci. USA, 109, 14369–14374, conagua.gob.mx/ocgc/ (last access: 12 September 2014), 2014 (in Spanish). Gómez, O. R.: InformeGobierno final: Inventario del forestal Estado de deCiencia los y Chiapas, bosques Tecnología Consejo de de Chiapas, manglar Nacional LaboratorioColegio del de de de Soconusco, Ecología La Ciencia de Frontera y Sur, Manglares Tapachula, Tecnología, y Chiapas, Consejo Zona México, Costera, 2007 de El (in//www.unesco.org/mabdb/br/brdir/directory/biores.asp?mode Spanish). cess: 12 November 2014, 2013. secondary forests: decisions andForest uncertainties Ecol. associated Manag., 262, with 1648–1657, allometric 2011. biomass models, Jackson, R. B., Collatz, G. J., and Randerson, J. T.: CO The mangrove classes(Class III) represent and a were derived gradient from a from NDVI (normalized upland vegetation index). (Class n.a. I) to lowland mangroves Table 1. Values are shown as mean (standard error). Max Williams, S. and Self,Woodro S.: The October 1902Yimer, plinian F., Ledin, S., eruption and Abdelkair, A.: of Soil organic Santa carbon and Maria total nitrogen stocks Volcano, a Rivera-Monroy, V. H., Day, J. W., Twilley, R. R., Vera-Herrera, and Coronado-Molina, F.:Siikamäki, Flux J., Sanchirico, J. N., and Jardine, S. L.:Sistema Global Meteorológico Nacional economic Comisión potential Nacional for del Agua, reducing SMN,Tovilla, available H. at: C.,http://www. Salas, R. L. R., De la Presa, P. J. C., Romero,United E. I. Nations B., Ovalle, Educational, E. F., Scientific and andVan Breugel, M., Ransijn, Cultural J., Craven, D., Bongers, Organization, F., and Hall, J. available S.: Estimating carbon at: stock in Van derhttp: Werf, G. R., Morton, D. C., DeFries, R. S., Olivier, J. G. J., Kasibhatla, P. S., 5 25 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | belowground = ) 1 − ) and total carbon (C) in 1 − Fromard et al. (1998) diameter at breast height. Wood = 1.11 ) ) ) C (MgCha ) Komiyama et al. (2005) 2.22 2.22 1 lnAGB) Cairns et al. (1997) − 2.22 × R aboveground biomass; BGB D ( (DBH (DBH = lnDBH Van Greugel et al. (2011) × × × × ) ) ) 1038 1037 0.8836 2.5 + 0.899 0.899 0.899 2.4 2.295 2.6 R + D DBH (0.67 (0.60 (0.84 DBH × × 1.0587 × × × × − 2.514 Aboveground Belowground − 0.199 0.199 0.199 Exp( 0.1282 0.140 0.1023 ======) within La Encrucijada Biosphere Reserve. Values are shown as mean BGB BGB AGB AGB AGB BGB BGB 1 − SiteMangroves PanzacolaTeculapa Biomass 383.6 (Mgha (153.6)Paistalon 127.9Esterillo 342.4 (47.6) (87.0)Sta 391.6 Chila (87.0) 234.0 (92.3) 118.3Zacopulco (20.4) 621.3 140.0 (310.9)Las (25.4) Palmas 210.5 198.8 (49.4) (13.4) 303.5 203.1 (76.5) (85.1) 242.6 706.6Peat (51.6) (172.6) swamp 93.9 (3.8) 377.4 127.8 (182.4) (29.9) 268.7 162.2 (52.5) (27.3)Marsh 195.5 (48.3) 440.0 43.5 (103.1) 132.1 (6.8) (7.8) 76.5 (11.6) 95.1 (15.7) n.a. 38.2 (5.8) ) values used for calculating belowground biomass were obtained from Chave diameter above highest prop root (cm); DBH 3 − = sp. lnAGB R Allometric equations used to calculate aboveground and belowground biomass Aboveground biomass, belowground biomass (Mgha D A. germinans L. racemosa Aboveground biomass R. mangle A. germinans L. racemosa Belowground biomass R. mangle P. aquatica Pachira Table 3. vegetation (MgCha (standard error). biomass; Table 2. (kg) of mangrove and peat swamp trees.density AGB (gcm et al. (2009). Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) 1 − 2.5 > ) 1 − ) 1 7.5 cm). Values are shown > ) (MgCha 7.5 cm) C stock 1 − > ) N stock (Mgha 1 − ) of downed wood at La Encrucijada 1 − Sound Rotten 1040 1039 7.5 cm) Large wood ( < 50 0.8 (0.3) 0.04 (0.01)50 83.4 (34.7) 16.7 (5.2) 3.5 (1.3) 1.02 (0.39) 379.850 (68.8) 26.4 (5.6) 4.7 (0.9) 0.24 (0.03) 114.1 (21.1) 6.8 (0.7) 50 17.1 (3.9) 0.71 (0.17) 389.450 (21.1) 16.4 (1.3) 16.1 (3.4) 0.56 (0.11) 479.350 (44.6) 2.6 (0.2) 14.8 (1.7) 0.49 (0.07) 317.750 (83.8) 11.7 (3.3) 8.5 (1.5) 0.34 (0.06) 247.2 (61.2) 12.7 (2.1) 50 26.8 (1.4) 1.04 (0.07) 451.650 (30.0) 17.5 (1.3) 15.9 (3.1) 0.67 (0.12) 421.2 (29.5) 18.4 (1.4) 15–3030–50> 1.7 (0.4) 1.2 (0.2) 0.09 (0.03) 0.07 (0.01) 20.3 (3.1)15–30 28.0 (5.5)30–50> 19.2 (5.9) 30.0 1.3 (7.2) 1.18 (0.2) (0.41) 1.5 1.69 (0.2) (0.35) 70.3 (26.1)15–30 105. (21.8)30–50> 6.9 3.6 (1.8) (1.2) Total 13.0 6.8 (3.0) (1.4) 0.42 (0.08) 0.65 (0.17) 32.0 (6.0) 113.8 (19.2) 5.8 (0.8) 2.8 (0.3) 298.3 (39.0) 18.5 (1.7) Total15–3030–50> 19.4 (4.0)Total 13.0 (4.0) 0.82 (0.16) 0.50 (0.13) 63.0 (6.5)15–30 69.6 (9.0)30–50> 21.7 (4.2) 2.6Total (0.2) 16.5 0.91 (4.1) (0.17) 613.6 2.9 (32.2) (0.1) 0.65 66.7 (0.15) (8.2)15–30 88.1 (14.3)30–50 27.2 (2.0) > 23.2 (2.4) 3.1Total (0.4) 12.0 3.1 (2.6) 1.08 (0.6) (0.12) 613.6 (23.6) 0.45 (0.09) 47.2 (5.6)15–30 71.9 (8.8)30–50 25.4 (1.5) > 11.8 (3.7) 2.8Total (0.3) 3.9 (1.6) 0.58 (0.20) 732.2 3.4 (53.8) (0.4) 37.9 0.18 (5.6) (0.06) 45.5 (10.9) 13.6 (1.1) 3.8Total (1.5) 393.0 1.3 (128.8) (0.5) 16.9 (5.7) Total 380.1 (68.6) 15.5 (4.3) 174.8 (41.9) 9.1 (1.7) 614.6 (85.7) 40.4 (5.5) 15–3030–50> 14.6 (3.7)Total 21.0 0.76 (2.8) (0.19) 0.92 37.9 (0.13) (7.1)15–30 73.6 (7.8)30–50> 20.1 (3.9) 1.9 (0.3) 8.8 (3.7) 0.76 (0.21) 3.5 (0.5) 68.6 0.37 (6.7) (0.16) 59.4 (11.8) 2.6 (0.5) 634.0 2.4 (25.7) (0.5) 26.5 (1.1) ) (Mgha 1 ) and C stocks (Mgha − 1 − Las Palmas 0–15 6.2 (1.2)Zapotón 0.32 (0.07) 43.1 (5.5) 0–15 16.3 2.8 (5.5) (0.3) Tular 1.05 (0.32) 59.5 (15.13) 0–15 3.6 (0.1) 15.6 (4.0) 1.10 (0.28) 38.3 (7.9) 3.0 (0.5) Paistalon 0–15 22.3 (4.4)Esterillo 0.82 (0.15) 91.6 (7.5) 0–15 20.4 3.6 (3.7)Santa (0.4) Chila 0.95 (0.18) 0–15 98.1 (6.6) 29.1 (1.3) 1.30 (0.06)Zacapulco 4.8 (0.4) 66.1 (6.2) 0–15 12.4 (2.9) 3.1 (0.4) 0.58 (0.15) 49.6 (8.1) 2.9 (0.6) SitePanzacola 0–15 Depth (cm) C (%) 16.6 (1.5) 0.88Teculapa (0.08) N (%) 71.0 (4.2) 0–15 C stock (Mgha 14.8 3.6 (4.0) (0.2) 0.78 (0.20) 64.3 (9.1) 3.7 (0.4) (Mgha Biomass (Mgha Soil carbon (C) and nitrogen (N) concentrations (%), and soil C and N stocks (Mgha PanzacolaTeculapa 5.8 (1.0)Paistalon 10.3Esterillo (2.8) 5.3Sta (1.1) ChilaZacapulco 5.5 (0.9)Las 6.6 Palmas (1.9) 4.4 (1.3)Zapoton 4.4 (0.9) 79.8 (24.0) 14.0 (4.4) 1.4 9.2 (0.6) (1.5) 7.7 (2.7) 3.4 (1.3) 43.5 0.5 (15.5) (0.4) 5.8 5.7 (3.0) (1.7) 88.9 (26.7) 11.9 (3.0) 34.1 (11.1) 4.5 111.5 (1.4) (45.2) 10.8 5.7 (3.5) (2.1) 9.4 (2.2) 102.4 (27.0) 3.0 (1.6) 5.3 (0.4) 11.5 (1.9) 22.1 (6.6) 20.4 (6.2) 12.5 (2.8) Site Small wood ( 7.5 cm), and large sound and large rotten wood (diameter erent depths (0–150 cm) of wetlands from La Encrucijada Biosphere Reserve. Values are < ff Table 5. at di shown as mean (standard error). Table 4. Biosphere Reserve. Wood debris was calculated separately for small wood (diameter and as mean (standard error). Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) 1 − emisions 2 emissions (Millions of Mg) within 2 844 0.6 2.2 (ha) (Millions of Mg) 1042 1041 TeculapaPaistalon 743.6 (42.2) mean 865.6 (55.1) Santa Chila 840.2 (50.1) 536.6mean (88.8) Las Palmas 825.8 (54.4) 640.9 (114.8) mean 659.5 (18.6) 32 62545 543 11.0 32.5 40.3 119.3 > ∼ ∼ Class II Esterillo 1,114.9Class (150.3) III Zacapulco 678.1 (115.7) Class IIClass III 6757 11 467 5.6 7.6 20.5 27.8 VegetationMangrove Class I Panzacola Site 911.6 (74.5) C (MgCha Mangrove meanPeat swampMarsh 784.5 (73.5) 722.2 (63.6) 336.5 (38.3) Mangroves Class IPeat swamp 9253Marsh 7.8TOTAL 28.5 Vegetation Area C stock CO Ecosystem C stocks for wetlands of La Encrucijada Biosphere Reserve. Values are C stocks (Millions of Mg) and their equivalent in CO wetlands of La Encrucijada Biosphere(I, Reserve, II and Mexico. III) Mangroves which were represented classified a in gradient from Classes upland (Class I) to lowland (Class III) mangroves. Table 7. Table 6. shown as mean (standard error). Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) 1 − ) of mangroves within La yr 1 1 − − yr 1 − (Mgha 1044 1043 Santa ChilaLas Palmas 1.3 (0.1) 0.4 (0.0) TeculapaPaistalon 1.4 (0.1) 1.7 (0.1) MEAN 1.3 (0.2) Class IIClass III Esterillo Zacapulco 1.8 (0.1) 1.5 (0.0) Mangrove class SiteClass I Soil C sequestration rate Panzacola 1.0 (0.1) Sampling sites within La Encrucijada Biosphere Reserve, Chiapas, Mexico. Soil carbon (C) sequestration rates (MgCha Figure 1. Mangroves werecorresponded classified a range according ofa the sites marsh were from also NDVI upland sampled. to (see lowland mangroves. Sect. A peat 2) swamp in forest and three classes, which Table 8. Encrucijada Biosphere Reserve, Mexico. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | (soil at (b) ) of riverine wetlands (mangroves, peat 1 − 1045 (trees and shrubs and down wood) and belowground (a) Aboveground erent depths and roots) carbon stocks (MgCha ff Figure 2. di swamp and marshes) withinIII) La represent a Encrucijada range Biosphere from Reserve. upland Mangrove to lowland classes forests. (I, II and