Table Of ContentAMAZONIANA XV (l/2): ll3 - 128 Kiel, Dezember 1998
Floristic composition and structure of two stands of
Senna ret¡culøta differing in age
by
Pia Parolin
Max-Planck-Institute for Limnology, Tropical Ecology Working Group, Postfach 165,
24302 Plön, Germany.
(Accepted for publication: November, 1998).
Abstract
Senna reticulata ('matapasto') is a woody pioneer which colonizes Amazonian whitewater floodplains.
In open areas, especially those which have suffered a strong anthropogenic impacf, Senna reticulata
dominates over other woody and herbaceous species forming large, apparently monospecific stands. In the
present study, stands of Se¿na reliculata which were two and six years old were inventoried in order to
describe differences ofspecies dominance and physiognomy. Senna reticulata dominated in the first years
after establishment, representing 86.4 Vo of all individuals in the two year old plots and forming the canopy
at 4-5 m height. In the six year old plots, Senna reticulata had only 28.4 % of all individuals, and the
canopy at 7-8 m height was formed by l2 species. The vitality of Senna reticulata was low in the six year
old stands. The trees had higher stems but very small crowns with few leaves, enabling the co-occurring
slow growing, long-lived species to take over dominance. This study shows that Senna reticulala, although
considered a noxious woody weed by the local people, has a restricted period ofdominance and represents
the initial phase ofa successional sequence that leads to a diversity comparable to that ofareas which have
not suffered anthropogenic impact.
Keywords: Floodplain, várzea, succession, Senna reticulatø, Amazonia, Neotropics.
ISSN 0065-675511998/1131 @ MPI für Limnologie, AG Tropenökologie, Plön; INPA, Manaus
l13
Introduction height (dbh), tree height, crown length and crown width were recorded, and different parameters were
calculated:
senna reticulata (wlLLD.) IRV/IN & BARN. (caesalpiniaceae) (synonyms cassia
ahelaigtah,t. cIat scsoilao nriezteicsu olapetøn; aLreOaRs iEnN nZutIr ile9n9t lr)i cihs Aam tyapziocnaila pni ownheieterw traeteer offlo modapxliaminusm, c1a2ll emd aabbusonlduaten cdeensity per ha ttoottaall nnuummbbeerr ooff iinnddiivviidduuaallss ppeerr ssppeecciieess, inca tlhceu lastaemdp tleod I phlaots
relative density relation between total number ofindividuals per species and total numbe¡
seasonal várzea (PRANCE 1979). These environments are characterizedby periodical of individuals of all species in all plots
floodings which can last up to seven months and oscillate by l0 m (JUNK l9g9). senna relative frequency percentage of plots in which the species occurs
reticulata occurs throughout Amazonia along the whitewater rivers (DUCKE 1949; DE absolute dominance stem cross section (m2) per ha, calculated by the dbh of the species
MENEZES 1978; KALLIOLA er al. t99l), but is restricted ro the upper levels of the relative dominance percentage of the species' dominance at total m2 of all species per ha
inundation gradient (maximum down to ca. 23 m asl near Manaus) because seedlings IVI species importance value index = relative density + relative frequency +
and mature trees do not tolerate complete flooding. relative dominance
In the first years after establishment, this species forms large monospecific stands,
especially on abandoned pastures or other areas which suffered a strong anthropogenic Mann-Whitney statistical analysis was used to test the differences oftree height, evenness and species
impact and high 'nutrient input' (Fig. l). Its local name 'matapasto' ('pasture killer') is diversity between the two and six year old plots.
related to its fast growth and the capacity to outshade other species. For this reason,
Senna reticulata is not liked by the local farmers. They usually cut and burn it shortly
Results and discussion
before the beginning of inundation in order to eliminate it. Thus, intact stands of higher
ages than l-2 years are difficult to find.
F'loristic composition
Senna reticulata Îrees do not occur inside mature forests. The species is seldom
mentioned in vegetation inventories (e.g. in woRBES et al. 1992), probably because 2246 frees were inventoried, 1549 in the two year old plots and 697 in the six year
these are generally carried out in little to not disturbed areas. Dense Senna reticulata old plots. In total, 32 woody species from 19 families were determined in the two year
stands can be found directly adjacent to sites at which inventories were performed. If old plots. 34 woody species from22 families were determined in the six year old plots.
22 species were common to the plots of both ages, while l0 species were restricted to
disturbed areas were to be analysed, then the dominating pioneer species would be Salix
humboltiqna, Cecropia latiloba or Cecropia membranaceø (SALO et al. 1986; the two year old plots and 12 to the six year old plots (Tab. l).
LAMOTTE 1992; \VORBES et al. 1992) which occur parallel to the Sennq reticulata The number of species per plot was twice as high in the six year old plots than in
the two year old plots (Tab. 2). The mean number of trees was similar in both plots,
stands, often on longer flooded and less disturbed sites. These pioneer species seldom
occur together (PAROLIN et al. 1995; PAROLIN, in press). and the number of dead trees, which were not included in the calculations, was twice as
In the present study, vegetation inventories were performed in stands of differing high in the two year old plots.
ages dominatedby Senna reticulata (Fig.2,3). The sites were deliberately chosen in Species composition in the two year old plots (Tab. 3) and in the six year old plots
(Tab. 4) differed with respect to density, frequency, dominance and importance value
altered environments where Senna reticulata was obviously the dominant species. The index (IV! of the species. In the two year old plots, Senna reticulølø prevailed with a
aim of this paper is to describe changes in dominance of Senna reticulata with increas-
relative dominance of 93.7 o/o, and an IVI of 280.2. Triplaris surinamensis and Vitex
ing stand age.
cymosa, which follow Senna reticulata in the ranking, had much lower IVI (81.2 and
62.3, respectively). In the six year old plots, Senna reticulata was still the most frequent
Methods species with a relative dominance of 52.9 %o and an IVI of 181.3. In these plots, the
difference to the next species in the ranking of IVI was much smaller ( 151.9 in Platy'
The study areas were located in the floodplains ofthe Amazon (Solimões) River, near the confluence miscium ulei and I15.5 in Ocoteq amara).
with the Rio Negro, in the vicinity of Manaus, Brazil (Fig. 4). According to the number of annual
increment rings measured in the wood ofseveral trees ofdifferent species, and according to the specifica- Forest structure
tions of the local farmers who cut and burn the stands of Senna retículata regularly, the age of the Most trees had diameters at breast height (dbh) between 1 and 6.9 cm in both the
vegetation stands was identified as:2 years age (5 plots: n" 1,2,3, c, d) and 6 years age (2 plots: n'a, b). two and six year old plots (Fig. 5), but mean dbh in the two year ald plots was lower
Three plots (n" 1,2,3) were located on the Fazenda Lira (Costa do Catalão) and four plots (n" a, b, c, d) than in the six year old plots (Tab. 2). The trees in the six year old plots had a wider
were located on the Fazenda Pec (Tena Nova/llha do Careiro). range of diameters, with a maximum of 32.0 cm(Cecropia membranacea), whereas the
Species were identified in the field with the help of José F. Ramos from INPA (lnstituto Nacional de trees in the two year old plots had a maximum of 14.6 cm (Vitex cymosa).
Pesquisas da Amazônia, Manaus) and Leandro V. Ferreira (INPA/Smithsonian Institution, Manaus). Maximum height was 5.5 m (Senna reticulata) in the two year old plots, and 12 m
Identifications were checked in the herbarium of the INPA.
(Platymiscium ulei) in the six year old plots. Mean tree height in the two year old plots
All woody species (including lianas) with dbh ) I cm or a height > I m were inventoried in the seven was significantly lower than in the six year old plots (Tab. 2;U = 0.5, p = 0.034). The
plots of25 x 25 m each. Dead trees were inventoried separately. For each tree, species, diameter at breast
tt4 ll5
canopy in the two year old plots was between 4-4.9 m and was formed mainly by Senna sites analysed in this study, four crown layers could be found, the highest at 23-28 m,
reticulata. Mean crown diameter in the two year old plots was almost half that of the the lowest at < l0 m (WORBES 1983). At another site on the same island, the highest
trees in the six year old plots (Tab. 2). In the six year old plots, most crowns closed at trees were 26 m high, and mean tree height was 5 to l0 m (KLINGE et al. 1995). In
7-8 m, but the distribution of height classes did not show a well defined canopy (Fig. this forest, the dbh of 50 Yo of the trees was above 19 cm. The same was found in a
6). Senna reticulatq and Platymiscium ulei dominated at 7-8 m and formed the main mature várzea forest near Tefé (AYRES 1993). In a mixedwater inundation forest at
canopy. Lago Janauari, 20 km from the study sites, the dbh of most species was below 20 cm,
but exceeded 100 cm in some species (AMARAL et al. 1997).
Sennø retículatø in stands of two and six years of age
The presence and dominance of Senna reticulata showed clear differences in the
plots of two and six years of age. Mean number of individuals of Senna reticulata was Conclusions
almost three times as high in the younger plots (Tab. 5), which is documented also by
the absolute and relative dominance and IVI of this species in the plots. The results of this study show that the dominance of Senna reticulata was very high
Species diversity was significantly higher in the six year old plots, compared to the only in the first two years after establishment. In the two year old plofs, Senna reticu-
two year old plots (U = 15.0, p = 0.024). The dominance of Senna reticulata decreased latahad86.4% of all individuals and formed the canopy at4-5 m height. In the six
with stand age. The mean density of senna reticulata decreased from 86.4 % to 28.4 %. year old plots, Senna reticulata had only 28.4% of all individuals, and the canopy at 7-
The total number of species per plot increased from 13 to 27 (Tab.2), indicating that 8 m height was formed by 12 species. At this stage, Senna reticulatq still had the
other species gained more importance. highest relative dominance and IVI, but the tree crowns were very small and had only
lf Sennq reticulata is excluded from the calculations, in the two year old plots there few leaves. This enabled the co-occurring slow growing, long-lived species to take over
was an average of 3.1 individuals per species (Tab. 5), compared to 9.2 in six year old dominance as is typical for successional sequences. In young várzea stands in Central
plots. This is emphasized by the evenness value of abundance (Tab. 2, calculated Amazonia, rapidly growing pioneer species dominate (Salix humboldtiana, Cecropia
according to KEEL & PRANCE 1979), which was significantly higher in the older plots latiloba; LAMOTTE 1992; WORBES et al. 1992), and are replaced by slow growing,
(U = 0.00, p = 0.024). The higher evenness indicates that more species have few long-lived species in the following years (WORBES et al. 1992). This shift is often
individuals in older plots. linked to changes of light incidence below the canopy, a relevant factor especially in the
In the two year old plots, 70 %o of the Senna reticulata trees had diameters below 5 case of Senna reticulata which forms an extremely dense canopy in the hrst years and
cm, compared to 46 o/o in Ihe six year old plots (Fig. 7). Maximum dbh was 8.6 cm in only later, after shedding the majority of its leaves, yields suffrcient light to smaller
the two year old plots, and 13 cm in the six year old plots. The canopy in the two year plants.
old plots was formed exclusively by Senna reticulata trees at a height between 4 and 6 Vast areas along the Amazon river are dominated by Senna reticulata as a conse-
m (Fig. 8). In the six year old plots, most Senna reticulata trees were 7-9 m high. quence of human activities. Forest clearcut and subsequent cattle and water buffalo
Maximum height was 5.5 m in the two year old plots, and l0 m in the six year old raising on pastures (OHLY 1985) lead to high light incidence and nutrient input into the
plots. soils which favour the establishment of Senna reticulata as the main pioneer species.
The relation between dbh and height in the Senna reticulata trees differed in the Although considered a noxious woody weed by the local people (DE MENEZES 1978),
plots of different age (Fig. 9). In the two year old plots, also the trees with very low or Senna reticulata has a restricted period of dominance and plays an important role in the
very high dbh reached heights of 4 m, while in the six year old plots the low trees had initial phase of a successional sequence that leads to a diversity which is comparable to
small dbh, and the high trees had high dbh. that of areas which did not suffer anthropogenic impact.
Comparison with other floodplain forests
Species diversity was lower in the stands of both ages of this study than in mature Resumo
stands of Amazonian floodplain forests (PIRES & KOURY 1959; TAKEUCHI 1962;
AYRES 1993; KLINGE et al. 1995). Evenness was 0.23 and 0.58 in the analysed two Senna reticulata ("matapasto") é uma árvore pioneira que coloniza áreas inundáveis de água branca na
and six year old stands of Senna reticulalø, respectively, compared to 0.75 in a black- Amazônia. Em áreas abertas, especialmente aquelas que sofrem alto impacto antropogênico, a espécie
water (igapó) forest (KEEL & PRANCE 1979). The number of individuals was higher domina sobre as outras espécies lenhosas e herbáceas, formando agrupamentos aparentemente monoespecí-
and the number of species was lower in the present study than in older forests. Many ficos. No presente estudo, grupos de Senna reticulata de dois e seis anos foram inventariados, a fim de se
species mentioned for mature forests (WORBES 1983, 1986; KLINGE et al. 1995) descreverasdiferençasdedominânciaefisionomiadasespécies. Sennareliculata dominounosprimeiros
anos após o estabelecimento, representando cerca de 86,4 % dos indivíduos nos plotes de dois anos, cujo
occurred also in the present study, although in the young plots these species were
dossel se encontrava a 4-5 m de altura, Nos plotes de seis anos, a porcentagem de Senna reliculata
represented only by small individuals with yery low IVI. reduziu-se para28,4 % de todos os indivíduos, compondo um dossel de 7-8 m de altura, formado por l2
Dbh and height were lower in the two and six year old Senna reticulata stands fhan espécies. Nestes, a vitalidade de Senna reticulata foi menor. Porém, as árvores eram mais altas, tendo
in mature forests. In a mature vârzea forest on the Ilha de Marchantaria, l0 km from the copas pequenas e reduzidas à poucas folhas permitindo assim a dominância de outras espécies de cresci-
lt6 tt7
mento mais lento, mas de longa existência. Este estudo mostra que mesmo sendo considerada uma ',planta WORBES, M. ( 1983): Vegetationskundliche Untersuchungen zweier Uberschwemmungswälder in Zentral-
daninha" pela população local, Senna retículata representa a fase inicial de uma sucessão que leva à amazonien - Vorläufige Ergebnisse. - Amazoniana S(l): 47-65.
formação de uma floresta com diversidade comparável aquelas sem influència antropogênica. WORBES, M. ( 1986): Lebensbedi¡gungen und Holzwachstum in zentralamazonischen Überschwemmungs-
wäldem. - Scripta Geobotanica, Erich Goltze, Göttingen: I12.
WORBES, M., KLINGE, H., REVILLA, J.D. & C. MARTIUS (1992): On the dynamics, floristic subdivi-
Acknowledgments sion and geographical distribution of Yírzea forests in Central Amazonia. - Joumal of Vegetation
Science 3: 553-564.
I thank L'V. Ferreira, J. Adis, W.J. Junk and J. Ohly for their helpful comments on this manuscript,
and the Instituto Nacional de Pesquisas da Amazônia (INPA) for assistance. The freld work in the várzea
sites was made possible by financial support from the INpA/MAX-pLANCK project.
Table l: Species occuning in 2-year old, 6-year old and in both 2- and 6-year old plots (each 25 x 25 m)
References
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l18
119
Table 2: Comparison of 2-year and 6-year old plots of Senna reticulatai floristic composition and forest \
structure (average and standard deviation).
d
d
plot age 2 years 6 years * ?ÉE.=
,{'x^.
oo
mean number of species per plot (625 m'z) 13 x7 27 !4 Ootsoo ËËê 9! e ! o. a I
mean number of trees per plot 310 + 130 349 ! 52 tootr
deveeandn etrseses per plot (%) 0.231 0i .90.06 0.58 5r.4 0.1I \:>o66 EF r
õo
ZÉ
mean dbh (cm) 3.5 !2 5.2!4 !Edõs E>E!
mean tree height (m) 3.5 11 5.0 t2 3.Þ
rn"un iree height (m) of trees with dbh > 5 cm 4.5 x 0.6 7.6 !2 oo(Ég
or(Ë EE
mean cfown area (m2) 2.5 4.t (ËÈ r
!X * å*
Éaoç
flv
r
E> ci
Ë!
rd:
So
QdoF ¡ F F r
dØ= € ts
_c
S¿E
Table 5: Comparison of Senna reticulata in 2-year and 6-year old plots. d9
Êo ts
(Ëo
plot ¡ge 2 years 6 years oNçO ts
'6¡
L.q
FE
-e r
absolute dominance of Senna reticulata (in m2lha) 1.3 0.2 âhbdEo Ê
relative dominance of Senna reticulata 93.7 s2.9 N
IYI oî Senna reticulata 280.2 l8t.3 Rg
mean number of Senna reticulata individuals 268 + 107 99+7 õ9
d(É I
((mSpeeearn nnp alno urt)emtibceurl aotfa inedxicvluiddueadl)s per species per plot 3.1 9.2 too!o üI 6 5Êg o s Fq ö O II ¿ O I E2 åEåÊI Ë € 'Ês 5ËE I Iã I E Ef sE
=o
39
.ÁF>:d ro _6>o ¡ .3rsgul Iã I6b I r)5I3 r9Ë).TI ó56EI bgI öË !I.rs$II) €I I I EÊI .9 tEIÊ EsI8'o=EI olgd I E82sp'têgÊr I5ÉtIor I
xo
Fõ9nt Ê È
aoooÞoD Eã;.,.dô;ì ÈË sðìNEÊ!s Èlt) ôI !Ë tì e.ñÊÊ sð .Ès sÈlrs I 3 .a\ { s! s:ì,5 ÈegËÈe èÈã!Þ o.Fi s{sI rissü tgr N ¡tas
,ô
Fd r
120 l2t
IVI 181.3151.9I 15.5106;7105.7 105.1104.0103.5103,2103.0r022102.0101.8101.6101,4t.310 2t01 101,0101.0 100.7 51.2 50,9 50^8{ô(s0350.450.450.45ô? 50,250.2 ,<lì 50"2 2900.0
e
elativeminanc 52.918.66,64.9A7 1A1.7 1.50,9 1.3 0.6 0"9 0.50,4OA0.50.30,30.3ôt 0,3020.20.2 0-1 0.1 0,10.t0.Ì 0,1 0.1 0.1 0.10.0 100.0
ro
d
*liana. ¡bsolutedominancem'/haìfin 980.1 0.06960,02480_0182 0.0160.00520-00640.00560.00320-00480.00220-00320.0018 0.00160,00140.0020.00120,00r0.0010.00090,00120.00070.00080.00080.00040.00040,00040.00040.00030^00020-00020.00020.0002 0-0001 0.3744
xt). bh
with abundance, tealculations, see dme¡n relativefreouenwIcmì 1005 1001.5 210071001008 1001 2100 210010012100I1002100I100I100I1002100I1001001 1001 l^5100501 500,7501 250501 501 501 I50501.5 501 501I50501 0.550
do Careiro); nations cand absoluteperdensity ha 396464124262052322832 2216t8l614l0t2l0l06t2l0844444222222 1394
(Terra Nova/llha c IVI (for explaand relativeabundâncedensiW 28.4198 33.3o¿8.9l31,9l01"4to3.7)al6?Õt4l6l2t;7n1.681,1 91.3l.l8 1,070.750.960"75o;75 0.460.9 0,7540.60.3220_320.320,320.3I0.1I0,1I0.1I0.1I0.1I0.1 100.0697 I -lÌ1ã' { dõ' 'd<-Þo ãg-. Þ- 5Þ9= ãt o.i fÞ :¿zØ=xñ +"-oo Þooo NoÀ oo Þ oôo
Pe e
zenda minanc bDlot 104 15033 11 3 8l549684 54 J4 J II0 00000220I000I 385
a o
F d
m) on ecies, plot a 9482292'7 l8 1 107634444I324a 6 5422200I0III0 312
5 sp
x ots (each 25 2ofthe dbh mean nmelocal MataDæto MæacaúbaLowoGenipapoIauíMr¡lueúBuiussub¡æcaknbauba Limo¡maCæiøoIapwtneá *cigma Unha de *Cipó TæbímaelaIrnbauba Limoraa 2JmbebaMuaubaAbiuæaAgudæeiËSadinheiraPataiubaPancææa2ãaçaGoiaba Euvi¡aTuimaGoiabaæaTmaqueiraPaicææTaD€rebáLimoræa 3lnsá-acu
d pl cy,
in oln 6-year nsity, frequen familv Cæsaloiniacæe PaÞd¡onaceaeLauæeRubiaøeA¡emcæPapilionaceae Fabacæe ec¡opiaceaeC Rutac@oOch¡aceae0upho¡baceæMimosacæe aesaloiniaceaeC Bignoniaceaeflippoc¡ataceaePolvqonæeæCæropiaceaeRutaceaeSolaacæBombacaceae SapotacaaoLeguinosæFlacorrtiacøeFlacourtiacøeCæsaloiniaceaeMlrtæaeAmonææAquifoliææeMvrtacqeRutaceaeMimowæAnacadiaceaeFlacourtiaceæMimosæae
o e
4: ble compositiSpecies relative dand abs'olute 5DeCrcS Sdtu rcticillalauleiPlalymiscium amaraOcotea amøicanaCenioa ttrocamm iauariErvlhrina fuscaOrmosia sp.Iatilobazcropia Zanthoxylum compactumOurat¿a sp.glandulosmSapium lnøa Dùnclafa 4nabidaea sp. Salacia sp. rcropia membramceaZanthorylum sp.øilinoSolanum Pæudobombu munzubaglomerataPouteria undet. Legum.Laetia corymbulosaactlealaCasearia Schizolobium sD.Psidium acutan%lumaotandra seud lyp P p Ie h b o inundafaIIex Euzenia sp.Fdgata compaclumoolvolullaEnlata ,ondias luteaintermediûmXylosoma Inøa so. TOTAI, Ë?l{ñ'.<!.:o Ê- o-ob .dô o ?lÞNoÞ--Þo pèoÞp 9
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N)NJ N)u)
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m
/r-
0
t
0 5
10km
Fig. 4:
Location ofthe study areas: Fazenda Lira on Costa do Catalão (l) and Fazenda Pec on Ilha do Careiro (2).
Fig. 3:
Six year old stand of Senna reticulata on Fazenda Pec/Ilha do Careiro.
t24 125
40 E 2 years
I 60 E 2 years
6 years
I
30 50 6 years
^\ 40
Ø 20 o\
(l)
c) Ø 30
L o
o
10 9! 20
10
0
Éo Oc\\¡ O+ \ \Oo\ O6 \ Oo \ ON \ O$ \ O\o\ Oæ\ OO\ O(\ \ O* r O\o\ Oæ\ Oo \ Ocl\ 0 q q q q q
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O\6r)F-O\c.lr)f-O\ (\ ôl C.ì C'.1 a.l cô -. r¡i r; ñ ¡ q qI
diameter classes [cm] o\ m
diameter classes [cm]
Fig. 5:
Diameter classes of all trees inventoried in Senna reticulata stands of 2 and 6 years age. n = I 567 trees in Fig. 7:
plots of 2 years, n = 701 trees in plots of 6 years. Diameter classes ofSe¿r¡ø reticulata trees inventoried in stands of2 and 6 years age. n = 954 trees in plots
of 2 years, n = 196 trees in plots of 6 years.
50 E 2 years
I 50 E 2 years
6 years
40 I
6 years
40
s30
t30
(t)
o
o 20 Uo)
c) 20
10
10
0
q q q q q q q
q-; ôl cô o+\ \o F- æ o\ cì 0 q q q q o\ o\ q q
ñ m + .o Þ- æ; O. .: ñ ô.I¡ ó s ro ÈI- æoI o\
-.'; ôi di + .o (-- oo I
-.¡
height classes [m]
height classes [m]
Fig. 6:
Height classes of all trees inventoried in Senna reticulqta stands of 2 and 6 years age. n = 1355 trees in Fig. 8:
plots of 2 years, n = 701 trees in plots of 6 years. Height classes of Senna retîculata trees in stands of 2 and 6 years age. n = I 134 trees in plots of 2 years,
n = 196 trees in plots of 6 years.
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