Acta Botanica Sinica

2004, 46 (5): 522532

http://www.chineseplantscience.com

Growth Dynamics and Self-Thinning of the Dominant Populations in the Mangrove Community
LIAO Wen-Bo 1* , LAN Chong-Yu 1 , ZAN Qi-Jie 1 , WONG Yuk-Shan2 , Nora Fung-Yee TAM2
(1. School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China; 2. City University of Hong Kong, Hong Kong, China)

Abst r ac t : Consecutive investigations in 1994, 1996, 1998, and 2002 in the permanent plots established in Futian Mangrove Reserve of Shenzhen, Guangdong Province, revealed that the breaking, drying and death of the individual plants or branches in the mangrove communities were significant and the number of plants in plots covering an area of 200 m 2 was 417, 341, 196 and 132; the average density of population per square meter is 2.08, 1.70, 0.98 and 0.66; the death rate between the interval of two investigations is 18.2%, 42.5% and 32.6% respectively. The individuals of population exhibit an obvious diameter at breast height (DBH) growth. For example, the maximum increment of individual DBH was up to 3.63, 2.45, and 4.52 cm in the dominant populations Aegiceras corniculatum (L.) Blanco (Ac), Kandelia candel (L.) Druce (Kc), and Avicennia marina (Forsk.) Vierh. (Am), respectively. At the same time, growth of individual height was also prominent. In the second investigation, 233 out of the 341 individuals exhibit a height growth; whereas the number under the third and fourth investigations was 127 out of 196 and 74 out of 132, respectively. During a 2 -year interval, the maximum height growth was 1.5 m in Ac, 1.9 m in Kc, and 1.8 m in Am. The biomass also showed a relative change in the mangrove communities. The total biomass of stems and leaves decreased with time but occasionally an increase was found in the second and third investigations and finally a decrease in the fourth investigation. However, the average biomass of the survived individuals usually increased with time. In terms of the total biomass, the results of the four investigations were similar to the previous research in which the data were calculated by the methods of standardized timbers, etc. being 7.57, 8.36, 5.15 and 7.71 kg/m 2 during 1994, 1996, 1998 and 2002, respectively. The abo ve analysis indicates that self-thinnin g of mangrove communities is an important evolutionary process, characterized by drying, breaking, and death of individuals/ramifications. During the process of evolution, fewer ne w seedlings deve loped, and individual height gr owth, DBH growth , bre aking and death from dryness maybe closely related to the composition, structure and density of population in the mangrove communities. Ke y wo rds : mangrove; Aegiceras corniculatum ; Kandelia candel ; Avicennia marina ; self-thinning; growth dynamic Despite the large body of materials and references concerning the research of seashore mangrove in China in the as pect s of s p ecies comp o s it io n an d g eog rap h ical distribution, vegetation types, community structure, population patterns of mang rove plants , an d ecolo gical and physiological feature of mangrove communities (Chang et al., 1957; Lin , 1987; 1997; Chen an d Miao, 1994; Li et a l., 1994; Fang and Liang , 1995; Miao and Chen, 1997; Wong and Tam, 1997), very few studies have been reported on the po pulatio n an d co mmu nity dy namics, and evo lut ion of mangrove. Permanent plo ts can be used to study th e dynamics of plant communities because it is easy for observing the individ ual characters an d life proces s in eco system, i.e. the continuous changing process in time and space; as well as observing the population change and development of communit y in ecos ystem, an d its environmental relation . And mangrove is a community type with higher productivity and high er recession rate, in which the species are of simpler composition, higher ecological dominance, lower diversity index and evenness; as well as with clumped pattern of dominant populatio n distribution (Chen et al., 1994b; Li et a l., 1994). Therefo re, it is o f impo rt an ce t o examin e th e dy namic p ro ces s o f the d ominant po pulat ion s b y des ig ning permanent plot s in the mangrove ecosys tem for lo ngterm eco log ical research. The authors designed a piece of permanen t plot covering an area of 200 m2 an d complet ed an eigh t-y ear s

Received 26 Aug. 2003 Accepted 10 Feb. 2004 Supported by the National Natural Sc ience Foundation of China (30230030, 39800012) and Cooperative Project Futian-CityU Mangrove Research and Development Centre. * Author for correspondence. E-mail: <lsslwb@zsu.edu.cn>.

LIAO Wen-Bo et al.: Growth Dynamics and Self-Thinning of the Dominant Populations in the Mangrove Community

observation. Th e chief o bjective of this st udy was t o investig ate the bio mass growth , individual plant gro wth in height and diameter at breast h eigh t (DBH), and demograph ic change in the mangrove communities, as a basis for further in spection on the pop ulation d ynamics o f the local mangrove.

1 Natural Geographical Condition

The mangrove of Shenzhen Futian-Neilingding National Nature Reserve of Guangdong Province, China, covers an area of 300 hm2, including 13 mangrove plant species, more than 100 semi-mangrove and seashore plant species, and more than 190 bird species (Deng et a l., 1986; Yang and Shen, 1992a; 1992b; Wang et al., 1993; Chen et al., 1994a; Chang et al., 1998). Especially, it is an important bird habitat, where sometimes there are as many as ten thousand birds. Based on Chen s (1994b) and Li s (1994) research, the species diversity index (Shannon-Wiener index) of Futian mangrove communities is 0.78, the evenness ind ex is 0.49, the impo rtance v alue of the do minant p opulation Aeg iceras corniculatum (Ac), Kandelia candel (Kc) and Avicennia marina (Am) is 0.72, 0.19 and 0.09 respectively. The simple composition and structure of the mangrove community are advantageous for further inspecting and observing the relationship between species and community evolution (Chen et al., 1994b; Li et al., 1994). Our selected permanent plot is situated in the northeast Futian Nature Reserve Mangrove in Shenzhen, with a geographical position at about 22 32N and 114 05E, belonging to an area of lower-subtropical o ceanic monsoon climate. The mean ann ual temperature is 22.55 , mean monthly temperature is 15.0 in Jan uary, and 28.72 in Ju ly . The annual p recipitat ion is 1 794.14 mm, and annual daily sunshine is 2 209 h exhibiting an obvious alternation of arid and humid seasons. In the Shenzhen bay and mangrove distribution area, the tide is the semi-diurnal with an average tidal range of 1.36 m, being maximal at about 2.8 m. The average salinity of seawater is < 1.5%, with a pH of 7.6. The physical and chemical properties of soil are as follows: organic compound, 22.3 g/kg; available N, 49.5 mg/kg; available P, 50.9 mg/kg; av ailable K, 951.2 mg/kg; so il salin ity, 2.1%. Among th e clunch deposit, 6.31% of the grain composition is > 0.25 mm, 24.87% of which is 0.250.01 mm and 67.35% of which is < 0.01 mm (Li et a l., 1994; Tam et a l., 1995a; Chang et al., 1998; Huang et al., 2000).

typical mangrove community in the south of Shazui village of Shenzhen bay. Each zone was 10 m in width and 180 m in length extending towards the beach. The zones were 150 m apart. A 0.3 m concrete bat was built on each side-border of A-zone. A-zone was then irrigated and drained with living waste-water at the time after daily ebb-tide weekly for one year. B-zone was served as control. Four permanent plots A-S1 to A-S4 and B-S1 to B-S4 were set up in A- and B-zone respectively. In each zone the four plots (each plot covering an area of 5 m 5 m) extended in sequ ence fro m the bottomland towards the beach, the total plot area was 200 m2 . The first plot A-S1 in A-zone (likewise B-S1 in B-zone) was 20 m away from bottomland. A-S2, A-S3 and A-S4 in Azone (B-S2, B-S3 and B-S4 in B-zo ne) were located in sequence further away from the botto mland. The sp ace between the neighboring plots was identically 10 m. 2.2 Plot investigation Plants with height of over 1.3 m, including all branches with DBH over 1 cm, t hat had branched from the t runk at levels below 1.3 m above-groun d were account ed. Individual plant was identified with plastic-sealed ID (with the code number). The ID cards were tagged on individual tree/ branch at the height of 1.3 m. Measurements included the tree/branch height (H, or length (L)), trunk circumference at breast h eight (expressed as DBH by divid ing th e breast circumference by 3.14), coord inate position of individuals (X, Y); and fro m which all data were compu terized into database (Li et al., 1987; Yu et al., 1993; Wang et al., 1996). The tree/branch h eight was measured at the highes t (the longest) p art and the breast circumference was measured aro und trun k at 1.3 m ab ove-grou nd us ing a measuring tape. Four consecutive invest igations were carried out in January of 1994, 1996, 1998 and 2002. It is preferab le to t ake a 5 m 5 m -s ized s ite as an individ ual plot, since a plot with an area o f 25 m2 usu ally includes three main dominant populations in the mangrove community. In addition, the number of individual plants in the eight plots, ranging from 41 to 65, represented a suitable densit y fo r st udy. Moreover th e po pulation in the commu nity, bein g different from t hose in th e land fo rest, gro w in clusters with luxurian t bran ches from short and stout bases. During the developmental process, the base changes much s lower than the b ranch es. Therefo re it is much easier to inspect the change of populations in mangrove community by measuring the total area at 1.3 m breast height (ABH; ABH = (DBH/2)2 ) of plants/branches than the t otal area at bas e, as the measurement of the latt er is very difficult and could be inaccurate. Furthermore, as the permanent p lots were des igned in the core region of the

2 Materials and Methods

2.1 Plot design In 1994 we selected two zones, A and B, in a natural and

man g ro v e commun ity , t h ey act u ally o ccu pied o nly 20+5+10+5+10+5+10+5 = 70 m among the 180 m from the bottomland extending to the beach. There was still a 3040 m conservation zone (of the same community), and another 30 40 m zo ne o f an nu al h erbaceou s plan t Aca nt hu s ilicifolius L. (Ai), and also a 30 m zone of sporadic distribution zone at the beach side (Wong et al., 1995a). 2.3 Data analysis The following equations were employed in analyses. Index of t he area ABH: ABH-Ind ex = Total of all individual ABH in community or plots / Total area of all community or plots; Biomass of mangrove community: W = a DBH2 H b (W is weight of totalbiomass; DBH is diameter at breast heigh t; H is tree height (length); a and b are constan ts), based on an analysis of Futian mangrove communities, the a and b values are confirmed by Tam et al. (1995b), i.e.: Ac: Log W = 1.496+0.465 Log(DBH2 H) Kc: Log W = 2.814+1.053 Log(DBH2 H) Am: Log W = 2.092+0.529 Log(DBH2 H) where W is the t otal b iomass of t he abo ve-gro und parts (kg), DBH is diameter at breas t heigh t (m), H is h eig ht (length) (m).

3 Results and Analysis

3.1 Characters of communities in the plots In the eight plots A-S1, A-S2, A-S3, A-S4, B-S1, B-S2, BS3 and B-S4, generally speakin g, the vertical stratification of communities was not significant with crown density of about 0.900.96. The d ominant populations examined in 1994 included Ac with a maximal DBH of 7.6 cm, Kc with a maximal DBH up to 12.1 cm, and Am with a maximal DBH of 9.6 cm, and other biennial herbaceous species such as Ai. Th e nu mb er o f Kc and A c were o verwhelmin g in t he mangrove. And Am was a mino r species, but it could become dominant in some sectional areas. Ai was particularly dominant in B-S3 and B-S4, d istrib uted s catteredly o r in clusters; As they exhibited a rapid life cycle only its height and number were recorded. In A - and B-zo ne, acco rding to Chen s (1994b) and Li s (1994) research , th e degree o f co rrelat io n amon g t he d omin an t p op ulat io ns (A c, Kc an d A m) was les s sign ifican t th an t heir complexit y, reflect in g a relative st ab ilit y o f th e man grov e commu nity. In ad dit io n, t he calcu lat ing un it of tot al biomass in the mangrove co mmu nit ies app roximated 12.14 kg/ m2 (Tam et al ., 1995b ). Based on a detailed investigation in 1994, the main characters of the plot communities were as follows: (1) Plo t A-S1 Th e Ac community, comp osed of Ac

solely, contained a total of 17 clusters with 54 main branches. Since A c grows in clu sters and all main branches usually grow directly from the root (or base), therefore t hey were recorded as the same coordination, but for those branches grown above 1.3 m, from a main trunk they were accounted as the same individual plant. (2) Plot A-S2 The Ac + Kc community contained five clusters of Ac with 41 branches and two clusters of Kc with eight branches. (3) PlotA-S3 The Ac community contained 19 clusters with 41 branches. (4) Plot A-S4 The Ac + Kc community contained 21 clusters of Ac with 43 branches and one cluster of Kc with four branches. (5) Plot B-S1 The A c + Kc commun ity co ntained 27 clu sters of A c wit h 48 branches and four clusters of Kc with 16 branches. (6) Plot B-S2 The Ac + Kc + Am community contained 16 clust ers o f Ac with 50 branches, th ree clusters of Kc with 10 branches, and one plant of Am (one branch). (7) Plot B-S3 The Ac + Kc + Am community contained two clusters of Ac with 16 branches, six clusters of Kc with 12 branches, six clusters of Am with nine branches, and the Ai included nine clusters with height ca. 1.5 m and six clusters with height ca. 2 m. (8) Plot B-S4 The Kc + Ac + Am community contained nine clusters of Ac with 13 branches, 20 clusters of Kc with 47 branches, two clusters of Am with six branches, and Ai more than 200 single branches. 3.2 Growth dynamics of population 3.2.1 Growth of population Based on the four consecutive investigations in January of 1994, 1996, 1998, and 2002, the individualchange of A- and B-zone is shown in Table 1. From Table 1, it is shown that in A-zone, the population Ac exhibited the highest branch death rate being 10.6% in 1994 and up to 36.9% in 1998, and that of Kc was minimal as no death was foun d in 1996, and only o ne branch died (8.3%) in 1998. In B-zone, the branch-death rate was stillthe highest in the Ac population, accounted for 31.7% in 1996 and 73.2% in 1998. Eighty-six out of 126 individual plants survived in 1996 and 23 su rvived in 1998. Branch death rate of Am population ranked the second highest, being 31.2% in 1996, 45.4% in 1998, and 33.3% in 2002. The death rate of Kc was high er than that in A-zone, accoun ted fo r 14.3% in 1996, 23.6% in 1998 and 20.0% in 2002. 3.2.2 DBH growth As shown in Table 2, in t he 1994 inv estigation o f A-zone, th e DBH of the Ac popu lation ranged 1.467.64 cm with an average incremen t of 0.37,

LIAO Wen-Bo et al.: Growth Dynamics and Self-Thinning of the Dominant Populations in the Mangrove Community Table 1 Demographic change in mangrove plots under different investigations*

Year / Individuals 1994 1996 Second death rate** 1998 T hird death rate** A-zone Ac 179 160 10.6% 101 36.9% Kc 12 12 0 11 8.3% TI 191 172 10.0% 112 34.9% B-zone Ac 126 86 31.7% 23 73.2% Kc 84 72 14.3% 55 23.6% Am 16 11 31.2% 6 45.4% TI 226 169 25.2% 84 50.3% T I in A- and B-zone 417 341 18.2% 196 42.5% *, number in 2000 not available; **, the death rate was calculated by using number of individuals in the f ormer n u mb e r . Ac , Aegiceras corniculatum ; Am, Avicennia marina ; Kc, Kandelia candel; T I, total individuals.

2002 Fourth death rate** 63 37.6% 11 0 74 33.9% 10 56.5% 44 20.0% 4 33.3% 58 31.0% 132 32.6% investigation as the cardinal

Table 2

Individual diameter at breast height (DBH) growth in mangrove plots under different investigations*

DBH Increment Increment Increment 1994 1996 1998 2002 (cm) (cm)** (cm)** (cm)** A-zone Ac TI 179 160 101 70 Maximum 7.64 8.82 3.63 9.49 1.43 10.35 4.68 Minimum 1.46 1.43 0.29 0.80 0.35 2.99 0.51 Average 4.15 4.58 0.37 5.12 0.22 5.80 0.49 SD 1.569 1.878 0.478 1.821 0.304 1.835 0.990 Kc TI 12 12 11 11 Maximum 12.10 12.68 1.27 13.12 0.76 14.45 6.18 Minimum 3.66 3.82 0.16 6.53 0.19 7.64 0.92 Average 7.42 8.18 0.76 9.08 0.50 10.31 1.23 SD 2.215 2.331 0.276 2.059 0.208 2.262 0.789 B-zone Ac TI 126 86 23 10 Maximum 6.37 7.80 2.55 8.12 0.70 9.04 0.96 Minimum 0.99 1.27 0.41 2.55 0 1.91 0.80 Average 3.02 3.46 0.27 4.35 0.11 4.28 0.38 SD 1.075 1.262 0.416 1.437 0.160 1.892 0.764 Kc TI 84 72 55 43 Maximum 10.19 10.99 2.45 11.78 1.46 14.01 11.46 Minimum 1.43 2.23 0.35 2.55 0.06 3.92 0.79 Average 3.78 4.81 0.74 5.48 0.38 6.52 1.01 SD 1.500 1.553 0.590 1.779 0.395 2.028 1.114 Am TI 16 11 6 3 Maximum 9.55 14.08 4.52 14.65 1.27 16.15 2.55 Minimum 3.03 3.09 0.16 6.37 0.06 9.14 0.70 Average 5.10 6.66 1.00 9.55 0.31 12.21 1.42 SD 2.066 3.602 0.825 3.233 0.527 3.181 0.819 *, all dead plants and branches were excluded from statistical analyses; **, positive increment > 0, zero increment = 0, negative increment < 0 (T hat the stem of individuals dur ing the process of branch drying and breaking shrank slightly results in a negative increment of the DBH growth and leads to ultimate death). Abbreviations are the same as in Table 1. Population

0.22 and 0.49 cm in 1996, 1998 and 2002 respectively. Comparatively that of the Kc population ranged 3.6612.10 cm wit h an av erag e increment of 0.76, 0.50 an d 1.23 cm respectively, indicat ing a more rapid g rowt h o f t he Kc population. In the B-zone, the average DBH of Ac, Kc and Am population in 1994 was 0.996.37 cm, 1.4310.19 cm, and 3.03 9.55 cm, increasing 0.27, 0.74 and 1.00 cm in 1996, 0.11, 0.38 an d 0.31 cm in 1998, and 0.38, 1.01 and 1.42 cm in 2002 respectively. Their growth was obviously increased in the

order of Am > Kc > Ac. 3.2.3 Hei ght growth Th ere was an obvious gro wth in height in the main population of the mangrove community and that branch breaking, blight or death leading to negative growth was also apparent (Table 3). From 1994 to 1996, the height growth of Ac increased 1.5 m in A-zone and that of Kc increased 1.9 m in B-zone. Further analys is showed significant growth difference among the dominant populations Kc, Ac and Am in A-zone an d B-zo ne. Du ring t he fou r in vest ig at io ns , th e to tal

 Table 3 Individual height growth in mangrove plots during eight years*

Height Increment Increment Increment 1994 1996 1998 2002 (m) (m)** (m)** (m)** A-zone Ac TI 179 160 101 63 Maximum 5.800 6.500 1.500 6.500 1.300 6.200 0.800 Minimum 1.400 0.800 4.000 0.600 3.800 1.300 2.500 Average 4.180 4.180 0.044 4.470 0.164 4.400 0.230 SD 0.992 1.565 0.884 1.480 0.726 1.465 0.259 Kc TI 12 12 11 11 Maximum 5.500 6.300 1.100 6.400 0.200 6.500 0.600 Minimum 4.400 5 0.300 5.100 0 5.100 0.400 Average 5.000 5.700 0.650 5.880 0.127 6.000 0.164 SD 0.387 0.496 0.215 0.485 0.065 0.563 0.277 B-zone Ac TI 126 86 23 10 Maximum 5.200 5.900 1.800 6.900 1.000 5.800 1.000 Minimum 1.300 0.600 3.900 2.600 1.100 1.500 3.900 Average 3.570 3.630 0.044 4.490 0.070 4.050 0.590 SD 0.933 1.332 0.959 0.903 0.370 1.283 1.500 Kc TI 84 72 55 44 Maximum 5.100 5.400 1.900 6.400 1.200 6.700 3.000 Minimum 2.200 0.600 3.500 3.500 0.500 1.300 2.800 Average 3.620 4.190 0.399 4.720 0.253 5.820 1.057 SD 0.820 0.992 0.872 0.632 0.266 1.079 1.254 Am TI 16 11 6 4 Maximum 5.700 6.700 1.800 6.900 0.600 6.200 0.500 Minimum 1.300 1.400 2.600 4.000 2.500 4.500 0.700 Average 3.850 4.680 0.364 5.400 0.717 5.700 0.025 SD 1.165 1.942 1.314 1.178 1.309 0.850 0.499 *, dead plants and branches were excluded from statistical analyses through all investigations; **, positive increment>0, zero increment = 0, negative increment <0. Abbreviations are the same as in Table 1. Population

number of individuals decreased from 417 t o 341, 196 and 132 res pectively. In the second inv estigation, among 341 individuals 286 exhibited positive DBH growth, 22 individuals had n o gro wt h and 33 h ad negative growt h (DBH const riction). However, in th e third investig ation, 137 of the 196 individuals exhibited positive DBH growth, 47 had no growth and 12 had negativ e growth. As for t he height growth , in 1996, 233 of the 341 individu als had increased height growth, 20 had no growth and 88 had negative height growth. From 1996 to 1998, 145 individuals died, including 68 from the 88 with negative height growth counted in 1996. And among the rest 196 survived individuals, 127 had increased height growth, 39 ceased growth and 30 had negative height growth. 3.3 Self-thinning of population and succession of communities Death of branches in mangrove plants initiated from the apex, expanded to the base and finally broke off. A common phenomenon is that firstly the branch stopped growing in length and then in DBH. Many of the breaking and drying branches migh t become s omewhat sto uter near the t ree crown and then shrank and eventually died and broke off.

The process of branch-death could extend slowly down to 1.3 m above ground. The dryin g, breaking and dying of branches of mangrove population is a natural self-thinning process as well as an ecological phenomenon of forest gap formation which is closely related t o population co mpositio n, commu nity cons truction, age construction and nature geograph ical position. In the Futian mangrove region of Shenzhen City, the mangrove population was of a cluster growth. Both Ac and Am had abu ndant branches, thus, their branch death obvio usly could affect space ch ange, alt hough the base diameter o f the plants showed relat ively little chang e as being expressed in the disappearance of only nine coordinat ions in A-zo ne but 17 co ordinations in B-zone wh ich was inves tigat ed in 1996, reflecting a ch ange of gro und stratum indeed. Moreover, the forest gap formation of mangrove community was different from th at of the land forest community such as the lower sub-tropic monsoon evergreen broadleaved forest in South China, which generally resulted from death of various populations. In contrast, the self-thinning proces s in th e mangro ve popu lat ion, becaus e of tho se

LIAO Wen-Bo et al.: Growth Dynamics and Self-Thinning of the Dominant Populations in the Mangrove Community

plants with ab undant branches and bigger base diameter, was usually carried out by continuous drying and breaking of branches. The forest commun ity in Shenzhen mang rove Nature Res erve, in part icular, h as develo ped into a mat ure and stable one, where the young plants of the abundant populations did not occur, and the seedlings were also seldom found, it was likely that the natural renew may be attributed to the formation of forest gap after the death of individuals. Nevertheless, if the formation of forest gap and self-thinn ing was related t o in d iv id u al d eath from n at ural sen escence, the colon ized dist ribu tion of p opulation in mangrove community would b ecome randomized, and the communities will eventually undergo a stage of decay (Chen et a l., 1994b ; Li et a l., 1994). An increas e of dry ing and breakin g branches an d th e sparsit y of youn g trees and seedlings in the community indicate th at this community was at a special stage, or a s ub-climax stage. On the other hand, based on a recent research (Zhong et al., 1996), the age of Shenzhen mangrove community could rough ly be abo ut 20 years, and it was ap proved th at t he ev olut ion cycle of mangrove commu nity co uld be rather rapid and short. 3.4 Dominance of communi ties and their change in biomass The dominance of communities were usually expressed by the total ABH of the populations (Table 4). From 1994 to 1996, the DBH of the main populations in

A- and B-zone was obvio usly increased, whereas much more decreased by 1998 indicating a marked strengthening of self-thinn ing activity. In the development process of mangrove community, the total individuals were obviously deceased, while the ABH of each individual was increased during the eight years, thus it was considered that the selfthinn ing was mainly res ulted from the individual diminution and in turn reduction of population density. The selfthin ning p rocess varied among three p opulations of the community, characterized by an individual d iminutio n of Ac and increment of ABH (no obvious change of number) of Kc in the A-zone and apparent individual diminution of bot h Ac and Kc in t he B-zone, as if there was less DBH change of Ac and Kc in the B-zone than in A-zone. Apparently there was a negative correlation between the change of ABH and individual number in Ac of A-zone as wellas in Ac and Kc of B-zone. As for theAm in B-zone, there was an increase of ABH and n o obvious d ecreas e of individ ual numb er, because unlike Ac, it has a major trunk with few branches. Generally speaking, taking the land forest ecosystem as an example, a maximum ABH index and a minimum plant or branch density may give rise to a higher community value. In fact, the evaluating indexis related to various methodology and validation of the fo rest community. Fo r example, an increase of the ABH and ABH-index indicates a growt hclimax o r -s ub climax of th e co mmu nity fo llowed b y a declination trait and eventu ally an trend to declin e, an

Table 4
A-zone

Change of total area at breast height (ABH) in mangrove plots during the eight years
Year / Individuals Ac

1994 1996 1998 2002 Total individuals 179 160 101 64 TABH (cm 2 ) 2 635.09 2 885.81 2 256.90 1 793.90 AABH (cm 2 )* 14.72 18.04 22.35 28.51 Kc Total individuals 12 12 11 11 TABH (cm 2 ) 560.66 677.53 744.63 958.00 AABH (cm 2 ) 46.72 56.46 67.69 87.09 Subtotal Total individuals 191 172 112 75 TABH (cm 2 ) 3 195.76 3 563.34 3 001.54 2 753.90 B-zone Ac Total individuals 126 86 23 10 TABH (cm 2 )** 1 014.87 913.49 317.84 169.54 AABH (cm 2 )*** 8.05 10.62 16.43 16.95 Kc Total individuals 84 72 55 44 TABH (cm 2 ) 1 088.84 1 440.73 1 432.12 1 606.15 AABH (cm 2 ) 12.96 20.01 26.04 36.50 Am Total individuals 16 11 6 4 TABH (cm 2 ) 377.40 485.41 470.98 492.21 AABH (cm 2 ) 23.59 44.13 78.50 123.05 Subtotal Total individuals 226 169 84 58 TABH (cm 2 ) 2 481.11 2 839.63 2 280.94 2 267.90 *, negative c orr elat ion of tota l indiv iduals ( P < 0.05 ); **, sign ificant po sitive c orr elat ion of tot al indiv iduals ( P < 0.05 ); * **, high ly signific ant ne gat ive co rre lation of tot al individua ls ( P < 0.0 1). AABH, a ver age ABH; TABH, to tal ABH.

LIAO Wen-Bo et al.: Growth Dynamics and Self-Thinning of the Dominant Populations in the Mangrove Community Table 5 Comparison of population density in mangrove plots during eight years*
1994 Population density 2.16 1996 Population density 2.00 1998 Population density 1.56 2002 Population density 0.88

Year / Individuals A-S1 A-S2

Ac 54 50 39 22 Ac 41 36 23 20 1.96 1.36 1.24 1.12 Kc 8 8 8 8 A-S3 Ac 41 1.64 34 1.36 14 0.56 7 0.28 A-S4 Ac 43 40 25 14 1.88 1.76 1.60 0.68 Kc 4 4 3 3 B-S1 Ac 48 35 4 2 2.56 2.00 0.68 0.60 Kc 16 15 13 13 B-S2 Ac 50 34 15 8 Kc 10 2.44 10 1.80 9 1.00 7 0.64 Am 1 1 1 1 B-S3 Ac 16 8 4 0 Kc 11 1.44 11 1.08 10 0.76 6 0.36 Am 9 8 5 3 B-S4 Ac 12 9 0 0 Kc 47 2.60 36 1.88 23 0.92 18 0.72 Am 6 2 0 0 T I** in A- and 417 2.08 341 1.70 196 0.98 132 0.66 B-zone *, in the different plots of 5 m 5 m in A- and B-zone, density change (individuals/m 2 ) was the result of population difference. Abbreviations are the same as in Table 1.

mangrove community. Comparing with the results in Table 1, the death rate was positiv ely related to the popu lation density. 4.2 Dynamics of population growth in A-zone and B-zone Based on the resu lts demonstrated in Tab les 14 and Fig.4, the difference in the dynamics of population growth was also obvious. In 1995, plots in A-zone was experimentally treated with living waste-water irrigation and drainage after ebb-tide at weekly interval for one year, receiv ing a total volume equaling 2 600 m3 in the plots o f ca. 1 800 m2 (A-zone, an area of 180 m 10 m). B-zone was served as control. By comparing the two kinds of plots, we might find some evidence whet her t he mangrov e forests and mangrove plants could develop some anti-pollution endurance that is of important significance in ecological and environment conservation. 4.2.1 Comparison of population mortality (Table 1) In A-zone, th e death rat e was 10.0% in 1996 an d 34.9% in 1998, and was 25.2% and 50.3% in B-zone, in dicating a higher death rate in the latter. Furthermore, the treatment of waste-water irrigation and drainage in A-zone exert ed no impact on its population mortality. In further analysis, the death rate of population in A-S1, A-S2, A-S3 and A-S4 of Azone in 1996 was 7.4%, 10.2%, 17.1% and 6.4% respectively, with an average of 10.0%; and that of B-S1, B-S2, B-S3 and B-S4 o f B-zo n e was 21.9%, 26.2%, 25.0% an d 27.7% respectively with an average of 25.2%. Noticeably, B-zone

had a higher and more even death rate among plots, while A-zone showed great disparity in death rate of population, averaged from 6.4% to 17.1%. The analysis suggested that the deat h process could mainly be related to the s pecies, branch composition and population construction. 4.2.2 Comparison of DBH growth The average DBH increment of Ac in A-zone was 10.36% in 1996 and 11.79% in 1998; and that of Ac in B-zone was 14.09% in 1996 and 22.65% in 1998 (Table 2). Th e average increment o f DBH growth of Kc in A-zone was 10.24% in 1996 and 11.00% in 1998; an d t h at o f B-zo n e was 26.56% an d 13.99% resp ectively. Both Ac and Kc exhib ited a dominant DBH growth in B-zone. Moreover, in B-zone Am also showed a remarkable average increment of DBH growth of 30.59% in 1996 and 43.39% in 1998. 4.2.3 Comparison of height (length) growth The average gro wt h increment of heigh t o f A c in A -zo ne was 0.95% (negative growth) in 1996, and 6.94% in 1998; and was 1.69% and 22.72% respectively in B-zone. The average increment of heig ht growth o f Kc was 13.10% in 1996 and 3.16% in 1998 in A-zone and 14.80% and 12.65% res pectively in B-zone (Table 3). Obviously the height growth of both Ac and Kc in B-zone was more significant. Likewise the Am population also had significant height growth in Bzone, being 21.56% and 15.38% respectively. On the whole, the more obvious change in B-zone was related to the original status (density, ABH, absolute height

of plant or branch, etc.) in the population and community. In th e first investig ation in 1994, t he average DBH o f Ac was 4.15 cm in A-zo ne an d 3.02 cm in B-zone s howin g a differen ce of 1.13 cm, and the latter was 37.42% less than the former. In respect of average height, Ac was 4.18 m in A-zone and 3.57m in B-zone respectively with a difference of 0.61 m, then the height of Ac in B-zone was 17.09% less than that in A -zone. Ev idently po pulations of B-zon e exhibited much more growth potential both in DBH and height. At the s ame time, the individual death rate in B-zone was higher, indicating that pollu tion may no t be the ess ential cause for the disparity between the two zones. On the other hand , one might co nsider if was te-water irrig ation might provid e so me nourishment for th e vegetatio n co uld be considered. Some recent studies (Tam et al., 1995a; Wong et a l., 1995a; Huan g et a l., 2000) hav e revealed t hat the to tal organic C an d P in s oil had no sign ificant chan ge before and aft er waste-water discharge in A-zone and the leaf conten ts of t otal organic C, P, and N were similar as well.

(4) Nonetheless, the potential physiological impacts of the environmen tal pollution to the mangrove community merits further consideration and investigation. References:
Chang H-T, Chen G-Z, Liu Z-P, Zhang S-Y. 1998. Studies on Futian M angrove Wetland Ecosystem, Shenzhen, Guangdong Province. Guangzhou: Guangdong Science and Technology Press. 1204. (in Chinese) Chang H-T, Zhang C-C, Wang B-S. 1957. M angrove community of Laizhou peninsula, Guangdong Province, China. Acta Sci Nat Univ Sunyatseni , 3(1): 122145. (in Chinese w ith English abstract) Chen G-Z , M iao S-Y , Tam F-Y , Wong Y-S. 1994a. Effect of s ynthet ic w ast ew at er on eco-p hy s iological indexes of Kandelia candel seedlings: a preliminary report. Chin J Appl Ecol , 5: 221224. (in Chinese with English abstract) Chen G-Z , Li M -S, Lan C-Y , Li S-H , Chen X-R, Liu Z-P, Tam F-Y , Wong Y-S. 1994b. Research on coenology of mangrove

5 Conclusion
An 8-year consecutive and comprehensive survey and analysis of the typical mangrove plots in Shenzhen Futian Mangrove Nature Reserve had demonstrated that the total number of individuals and population density in mangrove community d ecreased pro gres sively with relevant ind ividual death, branch-breaking and growth of DBH, height as well as biomass. This indicates that this mangrove commun ity has been developing with a pos itive pro cess of evolution characterized by the following features: (1) Self-thinning of the Shenzhen Futian mangrove community is an important evolutionary process characterized by frequent plant and branch death, drying and breaking. But the clu stered develo pmen t of mang rove plants has much less impacts on positional change of ground surface. (2) Growth in DBH and height of mangrove population was quite significant. Generally speaking, the total aboveground biomass usually decreased, nevertheless, the average individual biomass was increased. For example, the average individual biomass of Ac in A-zone was 3.18, 3.50, 4.98 and 4.46 kg /p lan t in 1994, 1996, 1998 an d 2002 res pectively ; that of Kc was 16.78, 23.2, 28.66 an d 38.50 kg/ plant resp ect iv ely ; an d t hat o f A m in B-zo n e was 11.13, 17.06, 25.73 and 34.12 kg/plant respectively. (3) The change in growth of height and DBH, individual and branch deat h is direct ly related t o the original st ate (co mposition and st ructu re con struction) of po pulat ion during the evolutionary process of the community.

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(Managing editor: HAN Ya-Qin)