PDF The Mangrove Community, Aspects of Its Structure, Faunistics and Ecology

LAGUNAS COSTERAS, UN SIMPOSIO. Mem. Simp. Intern. Lagunas Costeras. UNAM- UNESCO, Nov. 28-30, 1967. Mexico, D. F.: 515-536, 5 figs. (1969)

THE MANGROVE COMMUNITY, ASPECTS OF ITS STRUCTURE, FAUNISTICS AND ECOLOGY

K. RUTZLER ':.

ABSTRACT

In a brief survey of the literature dealing with mangroves in the sense of a littoral habitat, the points of focus and results of earlier works are summarized. The various biotopes within the community are classified in accordance with their dominant floristic and faunistic components. It is felt that not enough consideration has been given to the sedentary faun.a which is reaching dominant importance on mangrove roots in certain regions. The limited abundance and peculiarity of the substrate raises interesting problems concerning interspecific relations and dynamics of associations.

A large number of habitats in mangrove swamps are influenced by the sea. These have been greatly neglected by systematists and ecologists in the past. With coordinated international and interdisciplinary cooperation it should be possible, with reasonable effort, to determine the community structure, i. e. the correlation and self-deter- mination of the bioconotic units.

INTRODUCTION

Back to the days of Alexander the Great we can follow the amazement of travellers first confronted with the ghostly evergreen forest raising from the sea. After Nearchus, commander of Alexander's fleet reported on the mangroves he had observed bordering the Arabian Sea, Theophrastus (305 B.C.), stu- dent of Aristotle, gave the first recognizable description of Rhizophora mangle and its habitat. With Linnaeus a period of morpho- logic and systematic bot'lnical studies began in the second half of the eighteenth century.

Because of the peculiar embryology of Rhizophora and the adaptation of mangrove trees to growth in salt w~ter the interest of biologists soon focused on the ecology of mangrove plant communities and associated vegetation. Studies ant their successions and physiology were carried out by Walter and Steiner (1937) for East Africa, by Davis ( 1940) for Florida, by Chapman (1943)

for Jamaica and by Galley et al (1962) for Puerto Rico.

Besides the osmotic properties of Rhizo- jJhora, Avice1tnia, Laguncularia and Sonne- ratia the function of the stilt roots and pneumatophores has also engaged many physiologists until Scholander et al (1955) proved the hypothesis that they served for ventilation of the rest of the root system which is buried in the anaerobic mud.

An important part of· the literature on mangroves has been devoted to practical human interests, from mosquito control (Gilroy and Chwa tt, 1945) to economic exploitation for building timber, fire wood, and tannic acid; not to mention the fact that the roots of the living trees serve as hiding places for fish and as substrate for much desired oysters (Holdridge 1939, Mattox 1949). Mangroves were even plant- ed artificially to avoid erosion of railway embankments (Bowman, 1917) .

.. Smithsonian Institution, Washington, D.c., U.S.A.

- 515 -

516 RUTZLER

This leads to the interesting aspects of their role in geology and sedimentology. The function of the mangroves in this respect has sometimes been denied, but more fre- quently overemphasized. However, it is a fact that during the changes of tides debris accumulates among the roots. This creates, with the aid of massive growing epiphytes, stagnant bodies of water favorable for de- position. In addition, dead leaves and other plant remains pile up and become imbedded.

There is no way for the roots themselves to hold the sediments as has been frequently stated; on the contrary, they might aid des- truction by breaking through .limestone rock [as in the example of the pneumatophores of Avicennia (Howard, 1950)]. Thus, in co- ordination with favorable geomorphological and hydrographical conditions, mangroves are definitely able to help build up land for a protruding terrestrial forest.

Prior to the turn of this century no zoological research had been done in the mangrove community if one neglects the reports of voyagers complaining about mos- quito attacks and praising the palatable oysters and crayfish. Later, the mangrove forest was recognized as a littoral habitat where truly terrestrial and typical marine organisms lived in close relationship and pe- riodically overlaped with the change of tides.

Also, attention was drawn to the fact that the mangrove is a physiographic unit, the principal components of which are organ- nisms; therefore, the problems, are pre- dominantly of a biological nature.

TOPOGRAPHY AND CLASSIFICATION

There are different types of mangroves de- pending on the various geomorphic features of the coaSt. Plant ecologists have demon- strated that there are distinct topographical successions which are similar in old world and new world mangroves. Pioneer red man- groves (Rhizophora) protrude farthest into the sea or border channels and· lagoons. Ty-

pically, they are followed by black mangrove (Avicennia) - salt marsh associes which are not regularly flooded. Then comes a seldomly flooded transition zone (e. g. Cono- carpus, HibiSCUS) which connects to dry land, frequently rain forest. According to the substrate the mangrove types are clas- sified as reef, sand, mud, and peat mangro- ves. This classification also expresses dif- ferent hydrographical conditions, which are of interest to the marine biologist. Reef and sand mangroves are the most exposed to- wards the open sea with sufficient water exchange to provide a typical marine en- vironment. One of these has been studied closely by the Great Barrier Reef Expedi- tion at Low Isles (Stephenson et

at

1931), and it was found that all the sedentary fauna from mangrove roots, i. e. corals, sponges, hydroids, anemones and ascidians and also the holothurians clearly belonged to the reef fauna proper.

Mud mangroves usually occur near river mouths, thus showing a steep salinity gra- dient from marine to fresh water. This effect was the subject of an exemplary study by Walsh (1967) in Hawaii.

Peat mangroves grow along sheltered shores and in lagoons without fresh water influences other than subsoil water and rain.

This latter type is of great interest for the marine biologist because the environment is such that a typical marine flora and fauna can exist but the factors are so variable and sometimes extreme that the organisms must be strongly selected.

This type of mangrove is rather common in the Western Atlantic and has stimulated the interest of various systematists and eco- logists as can be learned from comprehen- sive studies by Mattox (1949) in Puerto Rico and by Gerlach (1959) in Brazil.

VERTICAL ZONATIONS

As in all coastal marine habitats the ver- tical distribution of communities is deter-

Figure 1. Typical mangrove channels as represented by Bone Fish Creek, East Bimini. On the permanently submerged parts of Rhi=ophora roots limitation of substrate causes spacial competition between sedentary organisms. HT=High tide level. LT=

Low tide level.

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518 RUTZLER

mined by the exposure to air. The supra- littoral is only a narrow zone because there is no wave action causing spray. However, it is extensive since it is mainly formed by the mangrove trees. Some crabs (Aratus, Sesarma) and gastropods (Littorina, N eri- tina) are dominant here closely following the changing sea level.

In the medio-littoral, the substrate and the living habits and morphological and physiological properties of organisms in- fluence the community structure. In order to avoid coverage by mud sedentary organ- isms must be confined to vertical stable substrates like stems, air-roots, and seedlings of the mangrove trees. Because they are ex- posed, however, the danger of desiccation during low tide is greatest. A characteristic intertidal algal community the "Bostry- chietum" has a high osmotic resistance and withstands desiccation. As field and labo- ratory experiments have proven (Biebl, 1962),it is not very resistant to light. Solar radiation, however, is comparatively low because of the - protecting canopy of the mangrove leaves. Animals with closeable shells like oysters (Crassostrea) and barna- cles (Balanus, Chthamalus) or with tubes within which they can retract, such as ser- pulids, remain inactive but protected during the dry period.

On the extended mud flats (the most in- vestigated habitat to date) representatives of the vagrant macrofauna alternate in their activity periods. Fish, such as gobiids and blenniids remain buried in mud or in clumps of algae during low tide. Crabs such as Uca remain in their humid, but air-containing burrows during high tide; others, like Ara- tus, avoid the rising water by climbing trees. The famous mud-skipper (Perioph- thalmus) is probably one of the very few truly amphibious mangrove animals.

Small, or true micro-organisms are, from the viewpoint of their micro-environment, inhabitants of the infra-littoral. During low

tide they have to withstand high or, in (;a~c

of rainfall, low salinities but not desiccation.

They either remain protected in crevices, bore holes in their substrate (mangrove roots, oyster shells), live endophyticly in clumps of algae, live endozoicly in oysters or spDnges, or live interstitially in mud or sand. In these cases the chemical and phy- sical conditions provided by the substrate determine the community structure.

It is difficult to obtain all references on systematics and distribution of mangrove dwelling organisms since they are frequently hidden in some specialized taxonomic work.

However, I have tried to summarize from a literature study all organisms characteris- tic for the different environments in both Eastern and Western mangroves. The resul ts are presented in table 1.

INFRA-LITTORAL TIDE CHANNELS

The proper infra-littoral, since the occur- rence of mangrove trees is limited by water depth, is almost exclusively restricted to a number of channels through which the water of the changing tides flows and to lagoons or ponds devoid of trees. Here the only solid substrate uncovered by mud are the ro- ots of bordering Rhizophora. The stilt roots of these trees extend to deeper water than the stems. At the current-exposed outer sides of the channel bends ("Prallhang") the soil is washed out, thus exposing a wicker- work of mud-roots under a mud-cornice.

This overhang is left in the upper medio- littoral because the water current is very slow near the peak of the tide. Examples of this forma tion can be found In narrow parts of mangrove channels where the current is accelerated such as in Bone Fish --or other creeks in the mangroves of East Bimini, Bahamas (figure 1).1

1Observations from Bimini mentioned here were made during work under Office of Na- val Research Contract NONR 552 (07). I am indebted to Mr. Robert Mathewson, director of the Lerner Marine Laboratory.

Figure 2. ]Jos/rycbieflllll grows on the tntertiebl parts of the red mangrove roots. At high tide numerous fishes find here hiding space.

Figure 3. Tedania ignis is one of the most common mangrove sponges. Because of its toxic properties, which even effect the human skin on touch and because of its flame-red colot it is com-

monly called the fire-sponge.

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TABLE 1 (CONTINUATION)

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TABLE 1 (CONTINUATION)

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TABLE 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITATS

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TAllL.l, 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITATS

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CyprMI'i.l·mlll/Ml'lIlalil ARTHROPODA

(DECAPODA) Veil /baYl'ri Vea p1/gllax rapax Alfibl'lIS bl'leroebaelis Vbogl'bia brasilil'lIsis Scylla sl'rra/a Thalami/a crl'Jla/a Sesar/l/(/ bidl'l1s T halassilla allomala Cliballarius striolatu., ARTHROPODA (ISOPODA)

Ligia altstralil'1lsis

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BV,B LT E,W f. Gerlach, 1958

BV,B LT W Gerlach, 1958

BV,B C W Gerlach, 1958

BV,B HT W Gerlach, 1958

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T ABLE I (CON TIN U A TION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITATS

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<: III

Z _~

<:

"

o

...

..

_<

... Z z :>

-< '!(

Z "-

i

Q

o Z o <

<.

~.~ t ~

~c. ~~ C ;::?

~g ~.~ ~ ⁰ ~~

::iJ= _~~ ... _9~ ...., ~;:

~~ t-..:r·~~ ~]~g

;:::

cQ ... ~'"'... " ' . . . . >- .

o iJ.l Z <....:::l .~::: ~ - ~ ~:: 1:::: ~

~bS :X:~~~~ t-;~'':: ~~"E ~

I.;.l:::-:o:r: u~c-;.;.~l.. '-~o::: <",.,~ O~

III " . . . Z II ~ -:::..E; > II -I t; ~ ~ ^{::< I:::!}

::s

z"

~ II·;: II 1=h I II 0 II II ...

:> <: ~ ::

>

^Vl II· u h ~" :> II

Z:;: l'< ...C::OC::O-l:Q <::t>-Jl) ,,~::< <_

- - - - -

MOLLUSCA

(GASTROPODA) ¹⁵

Alderia 'Uda BV HT W Gerlach, 195 8

Pyra::lIs palltstris BV HT F Stephenson et a.l, 1931

Telesco/lilt m tclcsco/,ill m BV HT E Stephenson rf ai, 1931

MOLLUSCA

(BIVALVIA) ^5-t-

Area tu.berclIlosll BS 1'11' W f. Gerlach, 1958

Pel/illil pllilltam BV,n I-lT E Stephenson et a.!, 1931

PISCES 5-1-

Gobillcllu.s smaragdus BV,B lIT W Gerlach, 1958

Pcr;ophtbalmlls koe!rrlltcri BV,13 HT E Stephenson et ai, 19.11

REPTILIA

Crocodyllls poroSIlS BV C E f. Gerlach, 1958

INFRA-L lTTORAL MANGROVETREE DIATOMS

++

^{ns - E,W Stephenson et ai, 1931}

SOLID (stem, stilt and muJ CYANOPHYTA

++

^BS

^-

^{E,W' Stephenson et ai, 1931}

roots washed ou t CHLOROPHYTA 5

by current) Caulerpa BS - E,W Stephensonet ai, 1931

Acetabular;a BS - W Margalef, 1962

ValOllia BS - W Margalef, 1962

PCIt;cilllls BS - W Riitzlec, unpuhl.

TABLE 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITATS

...l<

"

o ~

"" I'<

5

~

~

~

""

o ~

~ !;;

I'< ~

>< :>

"" <I>

~o ~

""

~ <

0: "

:> ""

""< <I>_~

Z :>

'"

0:<

o..J

~ <

"" z z

:>

< <

z

~

i

Q

o Z

Q <

PHAEOPHYTA Ectoca.rplls Padina

Dictyota divaricata RHODOPHYTA

Laurel/cia obfllsa PORIFERA

Tedal/ia igl/is 1rcillia fascicu.lata Chol/drilla '/I1tcula Dysidea fragilis Sigmadocia caerulea Al/thosigl71i'lla Farial/s MY"iastra

Bie1ltl/a microst)'l"

Deslltacella ;al/ia Spol/gia officil/alis Spirastrclla purpurea CNIDARIA

(ANTHOZOA) Bartholomea alllltliata Condylactis

<I> I'<

c; :>

U⁰

\OJ 0:

I'< '-' V>

~

.

o ~ Z

> 0

" 0 ><

\OJ 0: <

"l " ""

~ Z 0:

::> < ~

z ;; I'<

8+

6+

28+

2

.... O¢

~ ~

'E

~':'::

.z

..

^~

;.

'"

_{"') - be}

.

t .;: ~ .::! .E

~~~:E5

:I: ~ ~

E

^t:

<.,;)...t)~ ^{... ;;::}

~ ~ ~ 'n ~

:i'QCO_CO

BS BS BS

B5

BS BS BS BS BS BS BS BS BS BS BS

BS

BS

a. '"

~ ~ ~ ~

~ ...

~ --t:! "" ;;:

>- .~ :l :;::

t: ~...§

5

~ II II "II

""!-o!-o

~::r::>-lU

C C C C C C C C C C C

C C

'"

.... ;.

s

^~

... be

be ::

~ ~

::! ~

;::

>< ::

.,. :: ...

...

...

'"

I'< '" ....

0( i::; ~

~ ~ ~

@

II II

"'-'J~

E,W W W

W

W W W W W W W W W E E

W

W

0: '"

0-,;::t:;,::.,

~

II

< -

Stephensonet ai, 1931 Margalef, 1962 Biebl, 1962

Biebl, 1962

Hechtel, 1965 Riitzler, unpubl.

Riitzler, unpubl.

Hechtel, 1965 Hechtel, 196 5 Riitzler, unpubl.

Riitz[er, unpubl.

Laubenfels, 1950 Laubenfels, 1950 Stephenson et a.l, 1931 Stephenson cf ai, 1931

Glynn, 1964 Riitzler, unpub!.

l'ABLE 1 (CONTINuAtiON)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF' MANGROVE SWAMp HABiTATS

"

~

_{~ ~}

::l _~

i'l f- I'< <

o ~

i'l '"

'";- 0'1;:>

;.. '"

I'<

o i'l

i'l f-

oe ;;i

;:> ;...

f- '"

< <Xl

Z ~

<

"

o-'_I'<

;... Z<

Z ;:>

<: <

Z I'<

s: ""

o Z

Q <

'"

'"

i'l ;:>

8

~

'" <.:1 I'< '" •

o i'l Z

> 0

" 0 ><

i'l " <:

<Xl <.:1 f-

~ ^{Z "}

;:> <: i'l

Z ~ '"

... co

::: .~

~ ~

c

~.~ ~ ^Q

~ ~ ... 0 ~ l'I

i-~""'-~~~~::::

~ .~ .~ ~.; ^u.l ~ ~ ~

<

":§

~ ~

g

~ ^--.t:! ~ ^.:::

:z: ~ ;:: >:: ... >< ,;;0 ;:l ...

~~~~ i-o~C:::

<.:1 -,=> ... ;:l -

11-

^<:>

~1111:~i1 ~

II n

::: ;::. Vl

II

^u r,

..<"'OQ:I ... "'O < >-.1v

"

"

^;:.

;:. <:>

c ...

... ;;0 co -:: ~

~ ;:

:::: :::

>- ::: ~

:z: ... ...

~ ~ ~

~ ~ ~

g ~ II

"'''<~

" "

~~

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II

< - CNIDARIA

(HYDROZOA)

Myriollcma amboil/clIsc Eudendrill1n

ANNELIDA (POLYCHAETA)

Sabellasttlrtc magl/ific"

Spirorbis MOLLUSCA

(BIVALVIA)

Brachydol/tes rcrUI"l/lIs ECHINODERM AT A

(OPHIURIDEA) Ophiotrix al/glllat"

ECHINODERMATA (HOLOTHURIOID.) StichoplIs badiol/otlls Holotburia pal-va

14+

-l

2

BS BS

BS BS

BS

BS

liS BS

C C

c:

C

C

C

C C

E, W W

\'if

\'if

W

\'if

W W

Stephenson ct ai, 1931 Riitzler, unpubJ.

Glynn, 1964 .Margalef, 1262

Glynn, 1964

Glynn, 1964

Glynn, 1964 Glynn, 1964

TABLE 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITATS

....<

"

o ~ t- I'<

t- >-

~ I-

~ ~~

o "

w~

I'< I:Q

>- ;:J t- '"

'"

o '"t-

"

<^t-

~;:J

'"

ROCKS AND GRAVEL (in channels, cleaned from sediments by currents)

SESSILE MACRO- BENTHOS

0(

"

o..J

'"

⁰⁽

t- Z Z ;:J

0( 0(

Z '"

;: Q

~ Z

Cl 0(

TENTACULATA (BRYOZOA)

Aetea al/guil/a 13ugula lIerilil/a Caulibl/gllia ar/l/al"

TUNICATA (ASCIDIACEA)

Polieilor olh'aef'lls Eetril/ascidia IlIrbil/a/,1 AseMia nigra

Clal'elilla oblol/gata MieroeO.ll1l1lS exasperall/s Botryills

Didf111111/1I/

PORIFERA Adoeia toxia SIJirasl reI/a 1111rpurea Dysidea tragilis ARTHROPODA

(DECAPODA)

Synalpheu.s brc/liearpus

'" Q.;

"' ;:J

U0

"' "

'"

~

:.:.. In •

o ~ Z

;> 0

~ ~ ~

I:Q ~ t-

liE Z ~

;) 0( ~

Z ;;; I'<

5+

10+

10+

.... "0

L~ .~

tj,"':::; ..J:::l

, ;. ~ ~

~.~ r...~.;:;

<~:Q~~

~~~~~_{~ ~ ~}

-

II 11. ¹

~

;::. '" II II

CQ::>:l_CQ

BS BS BS

BS BS BS BS BS BS"

BS

BS BS BS

BV, I

Q

~ ~ ~

~~] ~

I'< ' " .;::

>- .~;:l ';::

r-",,:,:..s

c

~ II II "II

t-h h

~::r:>.JU

C C C

C C C C C C C

C C C

C

"

"

..

~_...

::

_"0

"0:::

::: ~

~ ~

~ ::: t

'"

...

I'< .. ~

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@

II II

(J1.:.l~

\'II W W

E W W

\XI

\XI W W

E E E

w

"'

t-

" 0(

o "

I t-

t- _~

;:J ;:J 0( ' "

Osburn, 1940 Osburn, 1940 Osburn, 1940

f. Gerlach, 1958 Mattox, 1949 Mattox, 1949 Glynn, 1964 Glynn, 1964 Mattox, 1949 RUtzler, unpubl.

Stephenson et ai, 1931 Stephensonet a,l, 1931 Stephensonet ai, 1931

Glynn, 1964

TABLE 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFER ENT TYPES OF MANGROVE SWAMP HABITATS

-J

..

<: 0

'"

o ~ ~

1-0 I'<

'"

1-0 >- >- :3 ^{1- '-}

..

₀

E:

;;J ~

" ^"

:J 1-0 1-0_<: '"⁰¹

Z ^:J'"

<:

!-- Z Z ::J

<: <:

Z ..

i

0

o zo ""

~ ~ '-:..

~ ⁵ 1~ .~ ~ ~

v _O ,,~ -c, ~ ,.:""

.. ~ ~ ~.;.~].~ § _{~ ~ ~} ;. _{~ ~}

o ;; ~ ^<~ ~

:s g

~ ~ .~ ^J: ~ ~

~ ^{0 ><} !: t _~

t ;-.

>-.~

5

~ ^c.. ~ ~~

-~~ Q~~"""':: t:-.:-

c

~l:S:::t

~

z""

~

II II .:;"" E II II '"

,,"J

II

?"" Cl :::;> '"

II II

v

t:

^h

I) 8 .11 ;;:,.

z~~ ..J~~_CQ <:...l...-JV ~..:.J.-

~'"-.

'- II

++ ++

(HYDROZOA)

1+

EV,B C

I C

I C

BV C

C MOBILE

(EPIPHYTIC) ALGAE

SAND, MUD

TENTACULATA (BRYOZOA)

BlIglIl" I1I'riti",r ARTHROPODA (DECAPODA)

Ubogrbia BACTERIA PROTISTA CNIDARIA Cassiopr"

ASCHELMINTHES (NEMATODA)

Sabtrtieri" c1at'irtrllda ARTHROPODA

(COPEPODA) NUoer., bypel'itis ARTHROPODA

(OSTRACODA) Mesoeytbere elol/galll ARTHROPODA

5-1-

2+

26

BS

c

C

c

\'\I Osburn, 1940

E Macnae, 1963

'W Margalef, 1962

\'If Margalef, 1962

\'If Riitzler, unpubl.

\'(. Gerlach, 1958

\'If Gerlach, 1958

\'If Gerlach, 1958

TABl.E 1 (CONTINUATION)

MARINE ORGANISMS CHARACTERIZING DIFFERENT TYPES OF MANGROVE SWAMP HABITA'rS

\VI Rodriguez, 1963 W R iitzler, unpubL

\XI Rodriguez, 1963 W Riirzler, unpubL

W Davis and Williams, 1950 W Mattox, 1949

W Margalef, 1962 W Glynn, 1964 E,W Yor. au thors

5

^~

Ii. I-

:.:l <:

0

" "

~ :J ^l-

i'. ~ ^~

,..

_:J

I- 7.;

'"

PELAGIC

"

"

3

Ii.

"

I- ;z.

;z. :J

" '"

~

'"

~ Cl

o 7.;

o "

(DECAPODA) Calillrc/rs sapitlm Pal/ulirlls MOLLUSCA

(GASTROPODA) Polymrsoda arc/a/(, Tridachia crisp,,/a COPEPODE NAUPLll, ETC.

DIA TOMS, ETC.

CILIATES ARTHROPODA

(ANOSTRACA) PISCES, YAR.

Ar/I'II/ia salil/a

'" 0.,

\.:.l ~

U0

I.:.J: ~

e; ~

5

^\.lJ__ i0 '" 0 x

r..:.: 0::: -<

r!l <:l I-

~ Z .,J

;:) -:r:: ~

Z ~ ~

4+

4-1-

50+

.10+

++

I 10j-

... 0.,

~ ~ .~

~.~ ~

;:....

.

, ...,

-

^~'

~ .~ ^I,.) .~ .~

o:l-Q :.i:1 ."';; :il

< ""'.lo ~ ^- c

~ t :: t: ~

~ ~ r:... ~

'"

^{~ '" ;-:}

%

II II .:; """

~ ^{;::. Vl}

II II

...l~~ ... :::::::

BV

BY

IW BV

g '-

....'"'::: ~ ~;::

~ ~ ~

g

c.. ~ ::::

e,., ~ ...

t

~~ ~ ~

~

_{, h h}

II II "II :<:r:-.Jv

C C

C C

'"

'-

,.

~~

,

'"

.... ....

'"

^'

.... ~

::::: :

>- ... ^l....

:r: ;.: ~

::. ~ t

~ ~ ;:,.

~ ~,~

<:l"-J~

~ ~ o~1;:'::'

~

II

"'-

---_._---_._---_._.---_

^..

_----_._---

Figure 4. Mass development of Cbolldril/a .,,1/cllla on mangrove roor prohibiting any orher sedentary organisms to settle.

Figure 5. On the shaded backside of dominating Ircillia !asciclliata four different species of sponges occur epizoicly.

'""

'"

N

;;d

c::

-1N r<

tT1

;;d

The bottom of these channels is covered with a thin layer of mud containing shell fragments and diatoms, which is inhabited by a rich microfauna. The underlaying peat is rich in H2S and organic carbon (Kor- nicker, 1958) and therefore poorly inha- bited. In areas more exposed to tidal cur- rents there are occasional stands of seagrass (Thalassia, CYlIlodocea) and green algae (PeniciIl1ls, Call1crjJa).

The intertidal parts of stilt roots and seedings are covered with a thick layer of bostrychictllIn (figure 2). No oysters or barnacles are present in the Bimini man- grove. Just below low tide level, heavy clus- ters of sponges cover the stilt roots, down to the point where the bottom sediments set a limit. Dominant are the species Tedania ignis, lrcinia fasc.'clliata and Chondrilla 1111C1lla (figures 3, 4). The only space com- petitors are the tunicate Ectcinascidia tttr- binata, some hydroids, mainly E1Idendri1t11l sp, and Myrio1lellla sp, and the anemone CVlldylactis sp.

Behind a curtain of stilt roots the above mentioned mud-caves are located; there again, on the exposed mud-roots of Rhizo- fJhora, sponges dwell abundantly. The do- mina ting species are Anthosig II'tl'1la 'variam, Myriastra sp, and Haliclona sp.

SPACLAL COMPETITION

The' very restricted a'vailability of sub- strate suggests studies on the spacial compe- tition of sponges and other sedetttaria. Ear- lier observations on isolated hard-bottom structures have shown (Riitzler, 1965) that the lack of substrate can be overcome by certain species which are able to settle and survive 'on other species through specialized morphological and chemical adaptations. A comparable situation can be found on the mangrove roots where numerous small spon- ge crusts, hydroids, and some bryozoans grow on the dominant primary settlers. Also a horizontal zonation according to light can be observ.ed, the epizoa preferring the semi-

LAGUN AS COSTERAS, UN SIMPOSlO 533- obscure surfaces which are faced toward the·

mud bank (figure 5).

Nothing is known yet about the effect of tannic acid and other chemical substances.

precipitated by the mangrove roots on the settlement of larvae. Other limiting factors are undoubtedly food supply and physical action of the tide current. The latter is naturally strongest in the undercuts. Sponges' of a certain size originating there are frequently torn loose, leaving space for new successions. A study concentrating on these·

environmental factors and their biocoenotic consequences is currently under way with, the aid of an instrument combination re- cently asserribled (Forstner and Riitzler~

1969). Main emphasis is given to long-term in situ measurements of micro-climatic fac- ters (mainly light quality and quantity~

water turbulences, temperature, salinity and oxygen), with the aim of identifying ana arranging micro-environments according to·

their ecological significance.

ENVIRONMENTAL AFFINITIES

The question arises finally, to which other habitats the mangrove community can be related. In fact, there are four environment complexes superimposed.

The tree tops are strictly terrestrial~

usually inhabited by the flora and fauna of a tropical rain forest. Lower branches and parts of the stem are comparable with the supra-littoral and pneumatophores, parts of the stilt roots and bases of tree stems with the medio-littoral of unexposed shaded rocky coasts. The mud flats are true marine medio-Iittoral soft bottom environments, which have been compared with the shoals of moder:lte latitudes, such as the North.

German U\'V'attenmeer" (Gerlach, 1958).

. The channels and lagoons, even in the- middle of the swamp, represent a marginal biotope. The muddy bottom with decompos- ing plant remains underneath and the com- position of the sedentary organisms growing on plant structures as sole substrate indicate-