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Some Data on the Testate Amoebae from the Shetland Islands and the Faeröer
Arch. Protistenkd. 136 (1988) : 79-96 VEB Gustav F ischer Verlag J ena
D ep ar t m en t of Botany, R.U.C.A., Antwerp, B elgium ; L abor a t oir e d e Zoologie Genera le et Faunistique, F .S.A .E., Gembloux, B elgi u m
Some Data on t he Testate Amoebae from t he Shetland I slands and t he F aeroer By L . BEYENS, D . CHARDEZ, D. DE BAERE, with t he collaborat ion of P . D E BOCK and E. JACQUES With 6 Figures a n d 2 Pla t es K ey words: T estate a mo eb a e, Ecology, Shetl a n d , Faeroer
Summary Sev enty-fo ur taxa of testate a m oeb ae were fo u nd in 25 aquati c sa m p les from t he Shetlands a nd the F aeroer. Several communities were di sting uished, some of them characteristic for acid, nutrient-poor conditions (the Arcello hem iephaerica and the D iff lug ia qlobulori» communities), while t he D ifflugia laoustrie community is observed in more m esotr ophie ranges. T h ere see ms to b e a geographical shi ft in the importance of the genus D iff lugia b etween Shetland and Faer6er. Fifty- six taxa were obs erved ill 18 mo ss a n d 5 lichen sa m ple s from the same a rc h ipels. Sev eral gro ups of s a m p les were r ecognizable, which were indicative of the moisture co nt en t of the s u bs t rat e. The Assul ina muscorum - Euglypha str iqosa qlobra community see m s to b e typical for dry habitats.
Introduction In 1942 H OFKER published an article on the testate a moe bae he found in some preserved material from t he Faeroer. He stated t hat no previous papers had been written on the subject, whi le as far as we know, no further papers have since appeared. H OFKER dis cusse d nine species in his contribution . Regarding t he Shetlands we have to go back as fa r as the works of WAILES (1912), who published a large num ber of papers on t he British I sles. It is no ex aggerat ion to state t hat so far the t est at e amoebae fa una of these island s have not been the subject of a profound study, altho ugh their geographical posit ion between the European mainland and Iceland (with the problems surroundin g it s exact biogeograp hical posit ion , i.e, partl y arctic or not .i ) justify a close r look. I n 1983 we had the occasion to take a small number of samples from both archipels. The res ults of t heir analyses are presented here.
Material and Methods The lis t of sa m ples and their n ature is presen t ed in Tables 1 a nd 2. Nomenclature of mo sses and lich ens a ccor d ing to CORLEY et a l. (1981), r esp. DAHL a n d KROG (1973). The loca t ions of the sa m pling s ites are indicated on Figs. 1 a nd 2.
80
L. BEYENS et al.
Table 1. List of moss and lichen samples Sample n?
Site n?
Altitude A.S.L.
Date of collection
F value
Nature of sample (L: lichen; M: moss or livermoss)
VIII VIII VI IV V V
L: Ramalina intermedia L: Cladonia arbuscula M: Eurhynchium praelongum M: Mnium hornum M: Hypnum jutlandicum M: Sphagnum lescurii M: Racomitrium lanuginosum M: Sphagnum capillofolium
VII III VII VIII VIII V II IV VIII VII V VIII
L: Peltigera sp. L: Parmelia pastillifera L: Ramalina intermedia Ramalina farinacea M: Schistidium maritimum M: Caliierqonella cuspidata M: Nardia scalarie M: Isothecium myosuroides M: Racomitrium lanuginosum M: Sphagnum subnitens M: Sphagnum lescurii M: Sphagnum papillosum M: Polytrichum formosum M: Philonotis fontana M: Sphagnum subnitens M: Racomitrium lanuginosum
Shetland LSOI LS03 SMOI SM02 SM03 SM04 SM05 SM06
2 6 2 3 5 5 6 6
<2m 40m <2m ± 10m 50m 50m 40m 40m
13.06.1983 14.06.1983 13.06. 1983 13.06. 1983 14.06.1983 14.06.1983 14.06. 1983 14.06.1983
LS05 LS06 LS07
7 7b 7b
120m 20m 20 m
16.06.1983 16.06.1983 16.06.1983
MS07 MS08 MS09 MSI0 MS11 MS12 MS13 MS14 MS15 MS16 MS17 MS18
7 7 7 7b 8 8 9 9
120m 120m 120m 20m 120m 120 m 120m 120m 120m 140m 140m 140m
16.06.1983 16.06.1983 16.06.1983 16.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983
Faerber
10
11 11 11
+
Mosses and lichens have been sampled from different sites, and stored for transport in polythene bags. Twenty grams of the mosses were taken and thoroughly shaken for 10 min in 100 ml distilled water. The suspension was passed through a filter with meshes of 300 I'm, and then sedimented. Ten grams of lichen were washed in 100 ml distilled water, and after removal of the lichen, the suspension was centrifuged. Microscopical analysis of the sediments obtained was done with a reverse microscope. Plankton samples were obtained by filtering a (variable) amount of water through a net with meshes of 25 I'm width. Scraping the bottom with a PVC-bottle yielded the sapropelium. Fixation of the material was carried out using 3 % formalin solution. After centrifugation, microscopic analysis was performed with a reverse microscope. In order to classify our field data we used an agglomerative polythetic classification method based on minimum variance clustering. To investigate the relationships among the species and samples we applied a principal component analysis in which the original data matrix was transformed into a dispersion or S matrix (SEAL 1968; MORRISON 1967; DAQNELIE 1975). The ordination method condenses the mass of raw data in such a manner that a minimum of information is lost. The cumulative percentage of variance of the third principal component gives a good estimate of the information held in only two planes of the hyperspace (Table 3).
Testate Amoebae from the Shetland Islands and the Faeroer
81
Table 2. List of aquatic samples Site/Samples (SW)
Conductivity (pho. crn r-)
Locality
pH
Description of habitat. Water temperature (near surface). Transparency in ern.
1/1-2
Lerwick
4.3
140
2/3-4
Scat Ness
6.4
2,600
3/5-7
Mousa
8.2
490
4/8-10
Mousa
6.6
1,000
small lake, depth (s.p.}: 18 em 20°C, T: 19, subaquatie mosses
5/11-l2
Lerwick
6.7
170
lake, depth (s.p.): 17 em, II °C, T: 21
6/13-15
Bressay
5.7
180
small pool: 300 em X 120 em, depth: 12 em, II °C, T: 30, subaquatic mosses
7/16
Mykines
7.5
2,400
pool: 35 III X 20 m, depth: 3 em, 18 °C
8/17-18
Skopun (Nordara Halsavatn)
5.6
145
lake, depth: 10 em, 11 °C, T: 29
9/19-22
Skopun
5.4
145
bog: 12 III X 10 m, depth: 10 em 11 °C, T: 29. Potamogeton sp.
10/23-24
Skopun (Heimara Halsavatn)
5.3
170
lake, depth: 10 em, II °C, T = 29, Potamogeton sp. and Juncus sp.
1l/25
Skopun
Shetland small pool: 38 ern X 42 em max. depth: 27 em, 9°C, T: 7.5 small lake, depth (sampling point): 20 em, 14°C, T: 13 aquatic macrophytes: Potamogeton sp., Eleocharis sp. small lake, depth (s.p.): 18 ern 20°C, T: 18. Subaquatic mosses
Faeroer
almost dried-up bog: depth: I em
In the text reference is made to the F-classification of JUNG (1936). This is a humidity scale, based on the water content of moss samples, determined as follows: FI = submerged mosses: Fll = free-floating mosses - partly submerged, partly floating at the air/water interface; FIll = very wet - water drips from the sample without pressure: FIV = wet - water drips on the application of only slight pressure; FV = quasi-wet - water drips after moderate pressure is applied; FVI = moist - little water is produced on the application of high pressures; FVll = quasi-dry - only a few drops of water can be squeezed out; FVlll = dryno water is exuded even after high pressures are applied. Hydrogen-ion concentration was measured with the portable Lab-Center pH-meter L 101, fitted with the combined electrode A213. To determine the conductivity, a WTW LF56 met-.r, with aLTA IOO/k electrode was used. Transparency (symbol T in Table 2), has been measured by lowering/bringing up a disc, attached to a ruler, in a black PVC-tube (50 em long, diameter: 55 mm) filled with the water under examination. The value (distance) on the ruler is noted when the disc is not longer visible/becomes visible again. The mean value is noted. 6
Arch. Protistenkd., Bd. 136
82
L.
BE YENS
et a !'
Fig. 1. Sketch map of the Shetland I sl ands (for sit es 1-6 see T able 2) .
Results and Discussion A. 'I'h e moss - a n d li ch en ha b itats The com pos it ion of t he fauna is give n in Table 4. Accordi ng to the cluste r anal ysis, several gro ups of samples can be dist ingui sh ed (Fig. 3). 1. The Assuiina muscorum - Euglypha strigosa glabra community, includ ing t he sam ples LS03 , LS06, LS07.
T estate Amoebae from the Shetlan d I sl ands and t he F a eroer
83
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F ig . 2. Ske t ch m a p of h e Faerii er (for s ites 7-11 see T a bl e 2).
2. Th e Centrop yxis aerop hila community, which is charact erized by t he dominance of this species. Samples SM03, SM07, SM09. 3. The Oeniropqei« sylvati ca - Phruqamella acropodia community, SMOI, SM02, SM05. 4. The N ebela communit y, where different N ebela spp . (N. collaris and N. tincta) and the qu antitative prese nce of Trin ema lin eare delineate several subunits : T able 3. Cumulative per cen t a ges of varia nce Prin cip al com ponen t
Cu m ulative per cent a ge of variance
1
24.6 37.6 48.3
2
3 6·
84
L.
BEYENS et.
hI.
P late 1. Aquatic sa m pling sites F ig . 1. Fig. 2. Fig. 3. Fig. 4. Fig. 5. Fig. 6.
Sh etl a nd, South Mainl a nd, Lerwick, site 1. Sh etland, So ut h Mainl a nd , Sca t N eKK, Kite 2. Sh etland, Mousa , South Isle, site 3. Sh etland, Mo usa , North I Kle, sit e 4. Sh et land, South Mainland, Sa nd y Lo ch R eservoir, Kite 5. F uer oer, Mykines, site 7.
Testate Amoebae from the Shetla nd I sla n ds lind the F aeroer
P late 2. Aquatic sa m pli ng sites Fig . F ig. Fig . F ig. Fig . Fig.
1. 2. 3. 4. 5. 6.
F a er oer , Sand oy , Skopun , Nordara Halsavat .n, site 8. Site 8: sa m pling p oint. F a eroer , Sa nd oy, Skop un, site 9, situated b et ween the sit es 8 a n d 10. Site 9 : sa m pli ng point. Faeroer, Sa nd oy, S k op u n , H eimara Halsavatn, site 10. F a er oer, Sandoy, Sk o p u n , s ite 11.
85
P ENARD Protop lcuri opsuci« ri dderae CIIARDEZ D ifflugia diatomosus CHARDEZ et GASPAR D iffl u gia bryop hila (PENARD) .TUNG D ifflu gi a tu berculata W ALLICH Difflu gia ylobulosa D UJARDIN N ebela collari s (E HRENBERG) L EIDY N ebela carinata (ARCHER) LEIDY N ebela dentisto ma PENARD N ebela [la bellulwm LEIDY N ebela m ilitari s P ENARD
Gyclop ya:i s p enardi DEFLANDRE 'l'rigonopya:is arcula (LEIDY)
D ECLOITRE
Gentrop ya:is aerop hila v . cornata
(D EFLANDRE) 'I'ROMAS
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CHARDEZ
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MS M S MS MS MS MS MS 'M S MS M S MS M S 07 08 09 10 11 12 13 14 15 16 17 18
+=
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f ound in less than 6 % of the sa m p les ;
She tland
=
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T able 4. Co m positio n of t h e fauna in the m oss a n d li chen sam p les. R 2 1-40 %; 3 = 41-60 % ; 4 = 61-80 % ; 5 = 8 1- 100 %
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~
o F ig . 3. Dendrogram re sulting from the cl us te r ing of the moss a nd the lichen sa m ples with the r ela tive fr equencies of the taxa within the co u nts as variables.
a) The Nebela collaris subunit, samples SM04, SM06 with N. collaris as dominant species. b) a group of sa m ples which all have high scores of Trinema lineare in common: SM12, SM14, SM13. c) The N ebela lincta su bun it , named afte r its most import ant species, SMll, SM17. All t he ot her samples are of m ixed com posit ion , and represent trans it ions between t he d ifferent com munities. A typical example are t he samples LS05 and SMlO, with elements of the Assulina muscorum - Euglypha strigosa community and the Centropyxis aerophila community. To test the ecological validity of these communities, we have used an ordination method. As is shown in Table 3, the information explained by the first axes stays rather low. Only some communities, as recognized from the cluster analys is, are clearly distinguishable in Fig. 4, and the following sequen ce along the axis of Component 2, ranging from low to higher frequencies is found: the A ssulina muscorum Euglyplla strigosa glabra community, the transitional unit ]~S05 - SlVIlO, and a rat.h-
90
L. I
2
I
BEYENS
et al.
I
I
I
I
-
f-
.lSOl
1~
0
SM03 SM17 .SM15 SM16 • .l. ...SM06 SM09 SM12~08 SM04 • ~1 ~ SM14
f-
SM07
•
o
.l.
SM11
ll.SM05
-
ll.SM01 .SM18
.SM10
-1
f-
-2
-
-3
f-
-
-
.lS05
o LS06 o LS03
-
N
C
Q)
c::: 0
o LS07
-
Q,
E
0
U I
-2
I
I
I
I
I
-1
0
1
2
3
Component 1
Fig. 4. Ordination diagram. Component 2 represents the humidity of the substrate, increasing in a vertical direction.
er complex cluster of the other associations. A comparison between this sequence and the observed humidity values reveals that in the lower half only dry samples are present, while dry and wetter ones are found in the upper part. Component 2 can be identified with the humidity of the substrate, which increases in a vertical direction. The meaning of score 1 (x-axis of Fig. 4) is still unclear, mainly because the lack of the appropriate ecological field-measurements. Nevertheless, from the literature we know that in some regions, the Centropyxis aerophila community inhabits the least acidic mosses and lichens while the Assulina muscorum community occurs under more acidic conditions (BEYENS et al. 1986a). We are inclined to believe that Component 1 reflects the acidity of the substrate, but we stress that this is only a tentative conclusion.
Testate Amoe bae fro m the Shetland I sl a n ds and the Faeroe r
91
200
z
150
0
I
Ci.i a:: w
CI.
CI'J
0
a:: w
100
ICI'J ...J
CJ I
Z
:I: I-
~
f-
1
~
'---
"--
~
f-
50
ff-
f-
ff-
o
oS~~~~~~~~~~~g~~~~~8~~~~ ~~~~~~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~
Fig. 5. D end r ogra m r esulting from t he clustering o f the a q uat ic sa m p les.
B. The aqu a ti c samp le s The composition of the fa una is found in Table 5. According to t he clust er analysis, several group s of samples can be disting uished (Fi g. 5). 1. The A rcella hemis phaerica community. SWOl and SW02 . 2. T he Cyphoderia ampulla community, where relatively high frequen cies of this species are found (> 25 % ). SW04, SW24, SWI8. 3. T he Dijjlugia globularis com munity. It is rather difficul t to pin a n ame t o t his cluste r of samp les, since D. globularis is on ly found in half of t hem. H owever, when present, it is the dominant species . It also has the hig hest mean re lative freq uen cy in this cluster . The other samples are linked together e.g . by a "common sp ecies" with relat ively high values (SWI5-SW20: Trinema enchelys, resp . 11.9-1 2.7 % ), or by a com bination of species unique to t hese samples, i.e, Cyclopyxis pirini, Dijjluyia minuia and L esquereusia ylobulosa (SWI9-SW20) . SW14, SW25, SW15, SW20,
SW17, SWI9.
4. The Diifluqia lacustris com m unity, where this species rea ches im port a nt valu es (domina nt in SW06 and SW09 ) be ing com plete ly absent in a ll other samples.
SW06, SW08, SW09.
(EHRENBERG)
-
-
Plagiopv'Cis callula P ENA R D Plagiopya,i.9 decliois 'l'1l01llAS P lagiop y'Cia labiata PENARD D iffluq ia acuminata EHItENBERG
PENARD
Oentropyxis eeorn ia LEIDY Oyelopyxia kahl i DEFLANDRE Oyclopyx i s piri ni G OL E M AN SK Y Trigo n opyx is areula (L~;IDY)
CHARDEZ
R -
-
-
-
-- -
-
It
-
It
-
-
-
-
-
-
- .
-
-
+
-
-
-
-
+ -
It
-
-
-
-
-
--
-
I
-, '
-
-
-
_..
4
-
-
R
R
+-
-
E
-
--
-
-
-
2
-
-
---
-
-
I
-
-
.-
-
1
-
-
-
-
-
-
-
-
-
-
-
5
_..
-
-
-
-
-- -
-
+ - -
-
._-
-
-
-
--
-- -
-
---
--
-
-
S
12
P
11
._- ---- -
I
-
S
10
+- - -
-- - - -
--
--
-
-
-
R
-
-
-
-
--- --
-
.)
-.,
-
- - --
--- -
-
-
I{
-
-- -
-
R
-
-
- - R
-
-
..
_-
-
-- -
-
--
2 It
[)
-
-
-
-
P
09
-
S
14
-
-
-
-
-
-
-
-
-
-
-.
-
-
-
-
-
-
-
-
-_.-
-
-
1
E
15
-
-
'-
-
H.
-
-
-
-- -
-
-
-
-
-
-
-
.-
-- +-
-
P
13
-
-
-
-
-
2
-
-
It
-
-
-
-
-
2
-
-
+- n
-
-
--
-
R
It R
R
R
It It
-
-
+
-
- -
.- .
.-
--
-
-
R
-
-
-
.-
-
21
-
-
--
'-
-
-
-
P
23
'-
. -
-
- - -
_.-- -
'-
-
---
2
3
22
SEE
20
-
-
R R
-
-
-
-
H
-
-
-
-
It
-
-
P
19
-
-
-
--
-
-
+- -
-
-
-- _.- -
-
--
-
-
-
-
-
S
18
+- --
-
-
P
- - It
-
-
-
S
17
-
-
-
+
It
-
--
-
-
-
-
S
24
-
E
08
--
Oentropyxia aerophila DEFLANDRE Oentropyxia aerophila v. minuta -
VAN QYE
Oentropyxis acul eata v. m inima
STEIN
Oentropyxia acul eata
DEFLANDHE
Areella vulgaris EHHENBERG AreeUa vulgaris f. reqularis
STEPAN~JK
Areella rotundata f. u ndulata
DEFLANDRE
Arcella roturulata P r,AYFAIR Arcella rotu n data v , sten ostom a
DEFLAND ltE
Arcella gibbo.9a P~: NAltI> A rcella hemi sphaerica PERTY Arcella hemisphaeri ca f . undulata
PLAY~'Am
Arcella diecoid es v . scuielliformie
S
07
SSP
06
P
05 16
04
02
01
+-
It
R
+
-
-
It
-
-
S
SW 25
.- - - - - - - - - - - - - -
21-40 %
F ueroer
11-20%; 2
SW SW SW SW SW SW SW SW SW
= 6-10 %; 1
Sh etlund
+-
SWSWSWSW SWSWSWSWSWSWSWSWSWSW
Table 5. Com position o f the fauna in the a q uat ic sam ples . R = found in less th an 6 % o f the samples; 3 = 41-60 % ; 4 = 61 -80 % ; 5 = 81-100 % . P = planetonic ; S = saprop eli c; E = ep ip hy t ic sample
r"
e,
ttl
'"e-
7\
t'l
~
l'J
I:l:l
~
-
PLAYFAl lt
---
-
-
R
--
--
-
-
.-
- _ ow
-
- . -
_
.
_
~
-
.._.-
- --
-
--
-
-
---
-
.-
-
-
-
-
-
- --
-
-
-
--'-
-
-
'- .
-
--
-
-
-
-
-
-
-_.
-
-
-
-- -
_.
-
-
-- -- -
--- -
---
-
-
_
-- - -
. _ --
-
--
_.
I
-
-
H -
- - - + - - - -- --- -- - - -- - - -.. _. - - _.- - - -. _ . - - - - - _. - -- -. - - - - - -~
_. _ .-
- - - - - - - --
- - - -
1
-
-
- -- -
_
----
- - ,_.• -- - -- -
2 -
2 _.- -
I
- --- - - - - -- - -- + - - - --- - - ..- - - "-.. -- - - -- - - - -_.. - -- -. - - - -. -. - - _. -- - - -
- --
:~
-- --_. - -- -
--
-.
-
-
- - e -- -- -
_._- _ . --- -
--- - -- -R - --
--
-
-
- - .-- - ---.. - - - -. --
'-
D ifflugia oentricosa DEFLA NDR E -DifJlugia tliffici iis T H OM A S D ifflugia diff1:eilis eurvata CHARDEZ Difflugia leidy W AI L E S Poniiqulasia eompressa RHUlIIB L E R Pontigulasia incisa R H UMB L E R Nebela dentistoma PE NARD N ebela collaris L E ID Y _..• N ebela f labellulum L E ID Y N ebela lagenijormis P E NARD
.._-
-
.-
---
-- -
- _..
_. _ ~
- -
-
2
1
-
-
-
- - -
-
_.
.-
J~
-
-- -
pulex PENARD paulii OGD EN pyriforme lla CHARD E Z rub esoen s PENAR D rotunda OGDEN scalp ellum P E N ARD tub erculata ,"V ALLIe H iubercu lat a v , sp haerica
--
--
- - .- -
.. -
_
-- R - - -
DiJflugia Difflugia Difflugia Difflugia DifJlugia DiJJlugia D ifflugia
DiJJl1~gia
T HOMAS
.TUNG
labiosa ' V Al LES minuta R AMPI parva ( THOMAS) O GDE N p enardi ( P E N AJtD)
&
HOPKINSON
Difflugia Difflugia D iJflugia D 1j f lugia
GAUTH.·L.
Difflugia lacus tris ( P E NARD) D iffl uqia Lu cida PENAR D D if f Tiugia lin eari s (PE N AR D )
LfHDY
Diff lugia eleqans v , bico rnis .TUNG D ifflugia gla ns PE N AR D Diff lugia qlobu lari« (WALL l e H )
- -."-
-
- - _. - - - -- - - - - - -- - - ..-
-- -
DELFANDJtfJ
DEFLANDJtE
-
--
+
Diff lugia eleqans v . a nqu sta ta
--
I
-
n
-
(PENA RD)
D ifflugia eleqa ns P E N ARD Diffluqia clegan s v . teres
.TUNG
Difflugia bryopMla 1
R
R
-
H
R
+ --
-
I R -
--
-
-
--
'-
-- -
-
-
-
-
2
+
-
R
-
H
R
R
R
R
It
R R R H R
-
-
-
-
---
I-t
-
-
-
R
-l-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
~ -- -
+ -
-
- - - -
-
-
-
-
-
-
-
-_.
1
-
-
-
-
-- -
-
-_.• -
-
-
-
.-
-
-
-
-
-
-
-
-
-
-
-
-
--- .
-
-
---
-
-
-
-
-
-
-
-
-
-
-
-
-
-
_. -
-
-
-
-
-
-
-
-
-
2
--
-
- - -- - - - R - + -
-
-
R
-
+
-
-
-
-
-
-
-
-
- -
-
H
- - R - - I _.. - -- -- -- -- - - - - --- -- -- - - -- - - - .- - - -- - - - - - - - - - - - - - - - - -
-
-
1
-
+ R I H - - 2
-
-
-
I
+
- -
- - -- - -- --
-
-
3
~
~
w
0'
l'.J
'" '..," "...
I-.j
:0-
~
...
;:I
'"
ta
::sp .
I>'
s,
H
0-
;:I
I;
0;-
<1>
:0-
IJJ
:3
0
::j>
~
0'
<'l>
0
~
<'l>
0;~ 0;-
'"
1-3 <1>
& THOlllAS
(HE RTWIG
& Lessun) HOl'KINSON
P hruqan ella aeropodia
LEIDY
T'rin ema comp lanat u m P E NAR D Sphenoderia minuta DEF LAN D R E Cyphoderia ampulla (EHR E N BE R G)
LEIDY
Euglypha compressa CART E R A ssulina muscorum GREEFF T'rinema lineare PENARD T'rin ema enchelys (EHRENBERG)
'VAlLES
Euglypha strig osa v . muscorum
LEIDY
Euglypha laevis PERTY Euglypha rotunda W AILES Euglypha strigosa (E H R E NBERG)
(EHU~:NB~:RG) PEIlTY
Euglypha acanth ophora
BUTSCHLI
Lesquereusia modesto R H UM BLER L esquereusia spiralis (E H R E NBE R G)
GAUTHn~Il-L .
L esquereueia globulosa
P~~ NARD
Quadrulella symmetrica W AL LleH Hyalosph enia subflava CASH H eleopera p etricola v , ame thys tea
I?:MW
N ebela penardiana DEFLAN D R E N ebela tincta (LEIDY) A W E R I N T -
Table 5 . continued
R
2
R
07
E
'T
08
P
09
S
R
10
E
5
11
P
12
S
2 1
3
13
P
It
-
14
S
R It
It It
R
It
It
15
E
+
- - -
+
+
It
2
-
S
18
-
R
-
10
P
21
22
-
H
H
It
-
-
SEE
20
24
3
+
S
25
I R
R R
S
-- +
23
P
17
06
S P
P
01) 16
01 S
02
SS
01
P
SW SW SW SW S W SW SW SW SW SW
._ - - - - - - - - - - - - -
Faeroer
S hetland
SW SW SW S W SW SW SW SW S W SW SW SW SW SW
~
~
.,..
~
t'J
><
t'J
tli
t-<
;e
Testate Amoebae from t he Shetland I sl an d s and the Faeroer
95
FAEROER
SHETLAND ARCELLA CENTROPYXIS DIFFLUGIA
f-I-------,
EUGLYPHA
NEBELA TRINEMA
o
20
40 %
o
20
40 %
F ig. 6. Histograms indicating the mo st important genera, a ccor d ing to their number of spe cies. This is ex pre ssed for each genus as a per centage o f the total n umber o f s pe cies .
5. The Tr inema lineare community, cha racteriz ed by the dominance of this species. Although this species occurs in other commu nit ies, it never rea ches suc h high va lues (> 40 %) t here. SW05, SWlO, SW21, SW16, SW23. A compa rison of these communities with the measured hydrogen-ion concent ra t ion and conduct ivit y proves that on ly two communities have limited ecological ranges. The Arcella hemisphaerica community is restricted to oligotrophic conditions, which is consist ent with most other dat a (SCHl)NRORN et al, 1965). Likewise, t he Di fflugia globularis communit y is only found in acid, nutrient-poor con dit ions. The Difflugia lacustris community on the other hand is observed in t he intermediate conductivity ranges, indicating a rather meso trophic occurrence. In this case t he habitats proved to be alka line to weakly acidi c. The wide cond uct ivit y ra nge occupied by t he Cyplwderia ampulla community and t he Trinema lineare community r ules out their usefulness in an ecological characterizat ion of the waters . At most sites both plankt oni c and sapropelic samples were taken, and it turned out, as one could expect, t hat in small basins both kinds of samples were similar. T his was obviously not t he case with the materia l from la rger basins.
Biogeographical as pects There is still a marked differen ce bet ween the aquat ic and t he terrestrial (moss lichen ) comm un it ies. As is known from literat ure (BEYENS et al. 1986 b), the differ ence becomes less evident in t he arct ic regions . On the other ha nd a diminution in t he importance of the Difflugia genus is already noticeable if we compare t he Shetland and the Faeroer records (F ig. 6). Thus the compos ition of t he fa una of t he lat t er archipe l already approaches t hat of I cela nd , and seems to hold an int erm ed iat e position between the moderate and the more boreal faunistic assemblages.
Note on some speci es Two spe cies, so far on ly recorded once , have been ob served again .
96
L. BEYENS et al., Testate Amoebae from the Shetland Islands and the Faeroer
Cyclopyxis pirini GOLEMANSKY Discribed by GOLEMANSKY and TODoRov (1985) from a mountain lake in Bulgaria. The observed specimens correspond well with the original description, and are mostly covered with diatoms, and have a sligthly invaginated aperture, surrounded by large irregular particles. They were encountered in aquatic samples (SWI9, SW20) from the Faerber.
Protoplagiopyxis ridderae CHARDEZ First observed by CHARDEZ (1969) in moss samples from Iceland, this species has been found again in the moss SM05 (Racomitrium lanuginosum) from Shetland. Diameter of shell: 66-73/-U11.
Acknowledgements We wish to thank Mrs. S. POOTERS, Mrs. A. TORFS, Mr. F. NEEFS and Mr. R. NEEFS for their technical assistance. We are also much indebted to Pro£. D. K. FERGUSON for correcting the English text. One of us (L. B) received a grant from the National Science Foundation (NFWO-Belgium) for the fieldwork involved.
Literature BEYENS, L., CHARDEZ, D., and DE LANDTSHEER, R., with collaboration of DE BOCK, P., and JACQUES, E. (1986a): Testate Amoebae Populations from Moss and Lichen Habitats in the Arctic. Polar BioI. 5: 165-173. - - - with collaboration of DE BAERE, D. (1986b): Testate Amoebae Communities from Aquatic Habitats in the Arctic. Polar BioI. 5: 197-205. CHARDEZ, D. (1969): Contribution it la Faune Thecamoebicnne de L'Islande. Bull. Inst. R. Sci. nat. Belg. 45: 1-16. CORLEY, M. F. V., CRUNDWELL, A. C., DULL, R., HILL, M. 0., and SMITH, A. J. E. (1981): Mosses of Europe and the Azores: an annoted list of species, with synonyms from the recent literature. J. Bryo!. 11: 609-689. ' DAGNELIE, P. (1975): Analyse statistique it plusieurs variables. Gembloux. DAHL, E., and KROG, H. (1973): Macrolichens of Denmark, Finland, Norway and Sweden. Oslo Bergen Trornso. GOLEMANSKY, B., and TODOROV, M. (1985): Thecamoebiens du sol de la Montagne Vitocha (Bulgaria). Acta Zool. Bulgarica 29: 50-63 (in Russian). HOFKER, J. (1942): Amoebida Testacea. Zoology of the Faroe, 1: 1-8. •JUNG, W. (1936): 'I'hekamoben urspriinglicher lebender deutscher Hochmoore. Abh. Landesmus. West£. 7: 1-87. MORRISON, D. F. (1967): Multivariate statistical methods. New York. SCHONBORN, W., FLOSSNER, D., und PROFT, G. (1965): Die Limnologische Charakterisierung des Profundals einiger norddeutscher Seen mit Hilfe von Testaceen-Gemeinsehaften. Limnologica (Berlin) 3: 371-380. SEAL, H. L. (1968): Multivariate statistical analysis for biologists. London. WAILES, G. H. (1912): Freshwater Rhizopoda from the Hebrides Orkney and Shetland Islands and Western Scotland with description of new species. Scott. Naturalist: 59-65. Authors' addresses: L. BEYENS, P. DE BOCK and E ..JACQUES, Leerstoel Algemene Plantkunde (Department of Botany), RUCA Groenenborgerlaan 171, B - 2020 Antwerpen, Belgium; D. DE BAERE, Department of Biology, U.LA., Universiteitsplein 1, B - 2610 Wilrijk, Belgium; D. CHARDEZ, Laboratoire de Zoologie Generale et Faunistique, Facult.e des Sciences Agronomi· ques de l'Etat, B - 5800 Gembloux, Belgium.
D ep ar t m en t of Botany, R.U.C.A., Antwerp, B elgium ; L abor a t oir e d e Zoologie Genera le et Faunistique, F .S.A .E., Gembloux, B elgi u m
Some Data on t he Testate Amoebae from t he Shetland I slands and t he F aeroer By L . BEYENS, D . CHARDEZ, D. DE BAERE, with t he collaborat ion of P . D E BOCK and E. JACQUES With 6 Figures a n d 2 Pla t es K ey words: T estate a mo eb a e, Ecology, Shetl a n d , Faeroer
Summary Sev enty-fo ur taxa of testate a m oeb ae were fo u nd in 25 aquati c sa m p les from t he Shetlands a nd the F aeroer. Several communities were di sting uished, some of them characteristic for acid, nutrient-poor conditions (the Arcello hem iephaerica and the D iff lug ia qlobulori» communities), while t he D ifflugia laoustrie community is observed in more m esotr ophie ranges. T h ere see ms to b e a geographical shi ft in the importance of the genus D iff lugia b etween Shetland and Faer6er. Fifty- six taxa were obs erved ill 18 mo ss a n d 5 lichen sa m ple s from the same a rc h ipels. Sev eral gro ups of s a m p les were r ecognizable, which were indicative of the moisture co nt en t of the s u bs t rat e. The Assul ina muscorum - Euglypha str iqosa qlobra community see m s to b e typical for dry habitats.
Introduction In 1942 H OFKER published an article on the testate a moe bae he found in some preserved material from t he Faeroer. He stated t hat no previous papers had been written on the subject, whi le as far as we know, no further papers have since appeared. H OFKER dis cusse d nine species in his contribution . Regarding t he Shetlands we have to go back as fa r as the works of WAILES (1912), who published a large num ber of papers on t he British I sles. It is no ex aggerat ion to state t hat so far the t est at e amoebae fa una of these island s have not been the subject of a profound study, altho ugh their geographical posit ion between the European mainland and Iceland (with the problems surroundin g it s exact biogeograp hical posit ion , i.e, partl y arctic or not .i ) justify a close r look. I n 1983 we had the occasion to take a small number of samples from both archipels. The res ults of t heir analyses are presented here.
Material and Methods The lis t of sa m ples and their n ature is presen t ed in Tables 1 a nd 2. Nomenclature of mo sses and lich ens a ccor d ing to CORLEY et a l. (1981), r esp. DAHL a n d KROG (1973). The loca t ions of the sa m pling s ites are indicated on Figs. 1 a nd 2.
80
L. BEYENS et al.
Table 1. List of moss and lichen samples Sample n?
Site n?
Altitude A.S.L.
Date of collection
F value
Nature of sample (L: lichen; M: moss or livermoss)
VIII VIII VI IV V V
L: Ramalina intermedia L: Cladonia arbuscula M: Eurhynchium praelongum M: Mnium hornum M: Hypnum jutlandicum M: Sphagnum lescurii M: Racomitrium lanuginosum M: Sphagnum capillofolium
VII III VII VIII VIII V II IV VIII VII V VIII
L: Peltigera sp. L: Parmelia pastillifera L: Ramalina intermedia Ramalina farinacea M: Schistidium maritimum M: Caliierqonella cuspidata M: Nardia scalarie M: Isothecium myosuroides M: Racomitrium lanuginosum M: Sphagnum subnitens M: Sphagnum lescurii M: Sphagnum papillosum M: Polytrichum formosum M: Philonotis fontana M: Sphagnum subnitens M: Racomitrium lanuginosum
Shetland LSOI LS03 SMOI SM02 SM03 SM04 SM05 SM06
2 6 2 3 5 5 6 6
<2m 40m <2m ± 10m 50m 50m 40m 40m
13.06.1983 14.06.1983 13.06. 1983 13.06. 1983 14.06.1983 14.06.1983 14.06. 1983 14.06.1983
LS05 LS06 LS07
7 7b 7b
120m 20m 20 m
16.06.1983 16.06.1983 16.06.1983
MS07 MS08 MS09 MSI0 MS11 MS12 MS13 MS14 MS15 MS16 MS17 MS18
7 7 7 7b 8 8 9 9
120m 120m 120m 20m 120m 120 m 120m 120m 120m 140m 140m 140m
16.06.1983 16.06.1983 16.06.1983 16.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983 17.06.1983
Faerber
10
11 11 11
+
Mosses and lichens have been sampled from different sites, and stored for transport in polythene bags. Twenty grams of the mosses were taken and thoroughly shaken for 10 min in 100 ml distilled water. The suspension was passed through a filter with meshes of 300 I'm, and then sedimented. Ten grams of lichen were washed in 100 ml distilled water, and after removal of the lichen, the suspension was centrifuged. Microscopical analysis of the sediments obtained was done with a reverse microscope. Plankton samples were obtained by filtering a (variable) amount of water through a net with meshes of 25 I'm width. Scraping the bottom with a PVC-bottle yielded the sapropelium. Fixation of the material was carried out using 3 % formalin solution. After centrifugation, microscopic analysis was performed with a reverse microscope. In order to classify our field data we used an agglomerative polythetic classification method based on minimum variance clustering. To investigate the relationships among the species and samples we applied a principal component analysis in which the original data matrix was transformed into a dispersion or S matrix (SEAL 1968; MORRISON 1967; DAQNELIE 1975). The ordination method condenses the mass of raw data in such a manner that a minimum of information is lost. The cumulative percentage of variance of the third principal component gives a good estimate of the information held in only two planes of the hyperspace (Table 3).
Testate Amoebae from the Shetland Islands and the Faeroer
81
Table 2. List of aquatic samples Site/Samples (SW)
Conductivity (pho. crn r-)
Locality
pH
Description of habitat. Water temperature (near surface). Transparency in ern.
1/1-2
Lerwick
4.3
140
2/3-4
Scat Ness
6.4
2,600
3/5-7
Mousa
8.2
490
4/8-10
Mousa
6.6
1,000
small lake, depth (s.p.}: 18 em 20°C, T: 19, subaquatie mosses
5/11-l2
Lerwick
6.7
170
lake, depth (s.p.): 17 em, II °C, T: 21
6/13-15
Bressay
5.7
180
small pool: 300 em X 120 em, depth: 12 em, II °C, T: 30, subaquatic mosses
7/16
Mykines
7.5
2,400
pool: 35 III X 20 m, depth: 3 em, 18 °C
8/17-18
Skopun (Nordara Halsavatn)
5.6
145
lake, depth: 10 em, 11 °C, T: 29
9/19-22
Skopun
5.4
145
bog: 12 III X 10 m, depth: 10 em 11 °C, T: 29. Potamogeton sp.
10/23-24
Skopun (Heimara Halsavatn)
5.3
170
lake, depth: 10 em, II °C, T = 29, Potamogeton sp. and Juncus sp.
1l/25
Skopun
Shetland small pool: 38 ern X 42 em max. depth: 27 em, 9°C, T: 7.5 small lake, depth (sampling point): 20 em, 14°C, T: 13 aquatic macrophytes: Potamogeton sp., Eleocharis sp. small lake, depth (s.p.): 18 ern 20°C, T: 18. Subaquatic mosses
Faeroer
almost dried-up bog: depth: I em
In the text reference is made to the F-classification of JUNG (1936). This is a humidity scale, based on the water content of moss samples, determined as follows: FI = submerged mosses: Fll = free-floating mosses - partly submerged, partly floating at the air/water interface; FIll = very wet - water drips from the sample without pressure: FIV = wet - water drips on the application of only slight pressure; FV = quasi-wet - water drips after moderate pressure is applied; FVI = moist - little water is produced on the application of high pressures; FVll = quasi-dry - only a few drops of water can be squeezed out; FVlll = dryno water is exuded even after high pressures are applied. Hydrogen-ion concentration was measured with the portable Lab-Center pH-meter L 101, fitted with the combined electrode A213. To determine the conductivity, a WTW LF56 met-.r, with aLTA IOO/k electrode was used. Transparency (symbol T in Table 2), has been measured by lowering/bringing up a disc, attached to a ruler, in a black PVC-tube (50 em long, diameter: 55 mm) filled with the water under examination. The value (distance) on the ruler is noted when the disc is not longer visible/becomes visible again. The mean value is noted. 6
Arch. Protistenkd., Bd. 136
82
L.
BE YENS
et a !'
Fig. 1. Sketch map of the Shetland I sl ands (for sit es 1-6 see T able 2) .
Results and Discussion A. 'I'h e moss - a n d li ch en ha b itats The com pos it ion of t he fauna is give n in Table 4. Accordi ng to the cluste r anal ysis, several gro ups of samples can be dist ingui sh ed (Fig. 3). 1. The Assuiina muscorum - Euglypha strigosa glabra community, includ ing t he sam ples LS03 , LS06, LS07.
T estate Amoebae from the Shetlan d I sl ands and t he F a eroer
83
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F ig . 2. Ske t ch m a p of h e Faerii er (for s ites 7-11 see T a bl e 2).
2. Th e Centrop yxis aerop hila community, which is charact erized by t he dominance of this species. Samples SM03, SM07, SM09. 3. The Oeniropqei« sylvati ca - Phruqamella acropodia community, SMOI, SM02, SM05. 4. The N ebela communit y, where different N ebela spp . (N. collaris and N. tincta) and the qu antitative prese nce of Trin ema lin eare delineate several subunits : T able 3. Cumulative per cen t a ges of varia nce Prin cip al com ponen t
Cu m ulative per cent a ge of variance
1
24.6 37.6 48.3
2
3 6·
84
L.
BEYENS et.
hI.
P late 1. Aquatic sa m pling sites F ig . 1. Fig. 2. Fig. 3. Fig. 4. Fig. 5. Fig. 6.
Sh etl a nd, South Mainl a nd, Lerwick, site 1. Sh etland, So ut h Mainl a nd , Sca t N eKK, Kite 2. Sh etland, Mousa , South Isle, site 3. Sh etland, Mo usa , North I Kle, sit e 4. Sh et land, South Mainland, Sa nd y Lo ch R eservoir, Kite 5. F uer oer, Mykines, site 7.
Testate Amoebae from the Shetla nd I sla n ds lind the F aeroer
P late 2. Aquatic sa m pli ng sites Fig . F ig. Fig . F ig. Fig . Fig.
1. 2. 3. 4. 5. 6.
F a er oer , Sand oy , Skopun , Nordara Halsavat .n, site 8. Site 8: sa m pling p oint. F a eroer , Sa nd oy, Skop un, site 9, situated b et ween the sit es 8 a n d 10. Site 9 : sa m pli ng point. Faeroer, Sa nd oy, S k op u n , H eimara Halsavatn, site 10. F a er oer, Sandoy, Sk o p u n , s ite 11.
85
P ENARD Protop lcuri opsuci« ri dderae CIIARDEZ D ifflugia diatomosus CHARDEZ et GASPAR D iffl u gia bryop hila (PENARD) .TUNG D ifflu gi a tu berculata W ALLICH Difflu gia ylobulosa D UJARDIN N ebela collari s (E HRENBERG) L EIDY N ebela carinata (ARCHER) LEIDY N ebela dentisto ma PENARD N ebela [la bellulwm LEIDY N ebela m ilitari s P ENARD
Gyclop ya:i s p enardi DEFLANDRE 'l'rigonopya:is arcula (LEIDY)
D ECLOITRE
Gentrop ya:is aerop hila v . cornata
(D EFLANDRE) 'I'ROMAS
Oentrop gtcie sylvatica
D EI'LANDRI;
-
-
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+ -
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-- -
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L S LS LS 05 0 6 07
- - -
-
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MS 06
-
Arcella roturula ta PLAYFAIR Arcella vulgaris EHR ENBERG Gentrop ya:is aerophila DEFLANDRE Gentrop ya:is aerophila v , sp ha qnicola DEFLANDRE Oentrop ptci» discoides (PENARD) -
CHARDEZ
Arcella arenaria GREEF A rcella arenaria f . comp ressa
MS 05
L S LS 01 03
MS 04
S hetlnnd F a eroer
11-20 % ; 2
F aer ber
6-10 %; 1
MS M S MS MS MS MS MS 'M S MS M S MS M S 07 08 09 10 11 12 13 14 15 16 17 18
+=
MS 03
f ound in less than 6 % of the sa m p les ;
She tland
=
MS M S 01 02
T able 4. Co m positio n of t h e fauna in the m oss a n d li chen sam p les. R 2 1-40 %; 3 = 41-60 % ; 4 = 61-80 % ; 5 = 8 1- 100 %
~
".,..
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to<
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THOMAS
THOMAS
&
&
Go.L .
--
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\VAILE S
LEIDY
Assulina muscorum GREEFF 1 A ssu lina seminulum (EHREN BERG) -
L E IDY
Euglyph u com p ressa CARTER Placocista sp in osa (CARTER )
+-
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WAILES
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Euglypha stri qosa f. heterospina ( P E NAH O) \VAILES et PENARD Euglypha stri qosa v . muscorum
-
-
--
--
-
(EHREN BERG)
WAILES
-
--
-
-
+
-- R - -
Euglypha strigosa f. glabra
LEIDY
Euglypha rotunda Euglyplul stri gosa
T H OMAS
L esqu ereusia modesta v , tru ncata
RnUMB LER
L esqu ereu sia motlesta
GAU T III E lt · L n !;V R E
L esqu ereusia gibbosa
S CU OND ORN
H eleop era sylva tica v , i nf lata
B ON NET
-
-
-
Ileleop era p etricola v, humicola
PENARI>
-
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--
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It -
-
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2
+-
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N ebela marginata PENAR D N ebela tubu losa PENARD N ebela ti ncta (LEIDY) A WEHfN '.rZEW N ebela tin cta f. stenostom a JU:NG N ebela tincta v , major DEFLANDRE N ebela tincta v . rotunda PENAR D N ebela parvu la CASH Quadrulla symmetrica W ALLleH fIyalosphenia su bf lava CAS H fIyalosphenia j irovici STEPANE K lIeleop era p etricola LEIDY H eleop era p etri cola v . amethqstea -
+-
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'l'rin ema lin eare P ENAlt D T'rinema complanatu m P ENARD I Coryth ion dubiu m 'L'AItANEK Coryth ion du b'ium v . orbicularis + P ENAItO Corythion dibium f. m in ima + CHARDEZ Corythion dubium v . gigas THOMAS Sphen9deria ovoid ea .r UNG Cyp hoderia ampulla (E HltENBERG) LEIDY Phryganella aero-podia (HERTWIG I et LESSER) H OPKINSON
2
04
03
02
01
MS
MS
Sh etl a n d
MS MS
T able 4. co nti nued
-
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MS MS MS MS MS MS MS MS MS MS MS MS 07 08 09 10 I I 12 13 14 15 16 17 18
Faer ber
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Shet land F a er ber
~
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....
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Testate Amoebae from the Shetland I sl a n d s ami the "F aeroe I'
89
150
z
0
en a: w Q.
en 3 a: 100 w
r
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o F ig . 3. Dendrogram re sulting from the cl us te r ing of the moss a nd the lichen sa m ples with the r ela tive fr equencies of the taxa within the co u nts as variables.
a) The Nebela collaris subunit, samples SM04, SM06 with N. collaris as dominant species. b) a group of sa m ples which all have high scores of Trinema lineare in common: SM12, SM14, SM13. c) The N ebela lincta su bun it , named afte r its most import ant species, SMll, SM17. All t he ot her samples are of m ixed com posit ion , and represent trans it ions between t he d ifferent com munities. A typical example are t he samples LS05 and SMlO, with elements of the Assulina muscorum - Euglypha strigosa community and the Centropyxis aerophila community. To test the ecological validity of these communities, we have used an ordination method. As is shown in Table 3, the information explained by the first axes stays rather low. Only some communities, as recognized from the cluster analys is, are clearly distinguishable in Fig. 4, and the following sequen ce along the axis of Component 2, ranging from low to higher frequencies is found: the A ssulina muscorum Euglyplla strigosa glabra community, the transitional unit ]~S05 - SlVIlO, and a rat.h-
90
L. I
2
I
BEYENS
et al.
I
I
I
I
-
f-
.lSOl
1~
0
SM03 SM17 .SM15 SM16 • .l. ...SM06 SM09 SM12~08 SM04 • ~1 ~ SM14
f-
SM07
•
o
.l.
SM11
ll.SM05
-
ll.SM01 .SM18
.SM10
-1
f-
-2
-
-3
f-
-
-
.lS05
o LS06 o LS03
-
N
C
Q)
c::: 0
o LS07
-
Q,
E
0
U I
-2
I
I
I
I
I
-1
0
1
2
3
Component 1
Fig. 4. Ordination diagram. Component 2 represents the humidity of the substrate, increasing in a vertical direction.
er complex cluster of the other associations. A comparison between this sequence and the observed humidity values reveals that in the lower half only dry samples are present, while dry and wetter ones are found in the upper part. Component 2 can be identified with the humidity of the substrate, which increases in a vertical direction. The meaning of score 1 (x-axis of Fig. 4) is still unclear, mainly because the lack of the appropriate ecological field-measurements. Nevertheless, from the literature we know that in some regions, the Centropyxis aerophila community inhabits the least acidic mosses and lichens while the Assulina muscorum community occurs under more acidic conditions (BEYENS et al. 1986a). We are inclined to believe that Component 1 reflects the acidity of the substrate, but we stress that this is only a tentative conclusion.
Testate Amoe bae fro m the Shetland I sl a n ds and the Faeroe r
91
200
z
150
0
I
Ci.i a:: w
CI.
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0
a:: w
100
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f-
1
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f-
50
ff-
f-
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oS~~~~~~~~~~~g~~~~~8~~~~ ~~~~~~~~~~~~~~~~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~
Fig. 5. D end r ogra m r esulting from t he clustering o f the a q uat ic sa m p les.
B. The aqu a ti c samp le s The composition of the fa una is found in Table 5. According to t he clust er analysis, several group s of samples can be disting uished (Fi g. 5). 1. The A rcella hemis phaerica community. SWOl and SW02 . 2. T he Cyphoderia ampulla community, where relatively high frequen cies of this species are found (> 25 % ). SW04, SW24, SWI8. 3. T he Dijjlugia globularis com munity. It is rather difficul t to pin a n ame t o t his cluste r of samp les, since D. globularis is on ly found in half of t hem. H owever, when present, it is the dominant species . It also has the hig hest mean re lative freq uen cy in this cluster . The other samples are linked together e.g . by a "common sp ecies" with relat ively high values (SWI5-SW20: Trinema enchelys, resp . 11.9-1 2.7 % ), or by a com bination of species unique to t hese samples, i.e, Cyclopyxis pirini, Dijjluyia minuia and L esquereusia ylobulosa (SWI9-SW20) . SW14, SW25, SW15, SW20,
SW17, SWI9.
4. The Diifluqia lacustris com m unity, where this species rea ches im port a nt valu es (domina nt in SW06 and SW09 ) be ing com plete ly absent in a ll other samples.
SW06, SW08, SW09.
(EHRENBERG)
-
-
Plagiopv'Cis callula P ENA R D Plagiopya,i.9 decliois 'l'1l01llAS P lagiop y'Cia labiata PENARD D iffluq ia acuminata EHItENBERG
PENARD
Oentropyxis eeorn ia LEIDY Oyelopyxia kahl i DEFLANDRE Oyclopyx i s piri ni G OL E M AN SK Y Trigo n opyx is areula (L~;IDY)
CHARDEZ
R -
-
-
-
-- -
-
It
-
It
-
-
-
-
-
-
- .
-
-
+
-
-
-
-
+ -
It
-
-
-
-
-
--
-
I
-, '
-
-
-
_..
4
-
-
R
R
+-
-
E
-
--
-
-
-
2
-
-
---
-
-
I
-
-
.-
-
1
-
-
-
-
-
-
-
-
-
-
-
5
_..
-
-
-
-
-- -
-
+ - -
-
._-
-
-
-
--
-- -
-
---
--
-
-
S
12
P
11
._- ---- -
I
-
S
10
+- - -
-- - - -
--
--
-
-
-
R
-
-
-
-
--- --
-
.)
-.,
-
- - --
--- -
-
-
I{
-
-- -
-
R
-
-
- - R
-
-
..
_-
-
-- -
-
--
2 It
[)
-
-
-
-
P
09
-
S
14
-
-
-
-
-
-
-
-
-
-
-.
-
-
-
-
-
-
-
-
-_.-
-
-
1
E
15
-
-
'-
-
H.
-
-
-
-- -
-
-
-
-
-
-
-
.-
-- +-
-
P
13
-
-
-
-
-
2
-
-
It
-
-
-
-
-
2
-
-
+- n
-
-
--
-
R
It R
R
R
It It
-
-
+
-
- -
.- .
.-
--
-
-
R
-
-
-
.-
-
21
-
-
--
'-
-
-
-
P
23
'-
. -
-
- - -
_.-- -
'-
-
---
2
3
22
SEE
20
-
-
R R
-
-
-
-
H
-
-
-
-
It
-
-
P
19
-
-
-
--
-
-
+- -
-
-
-- _.- -
-
--
-
-
-
-
-
S
18
+- --
-
-
P
- - It
-
-
-
S
17
-
-
-
+
It
-
--
-
-
-
-
S
24
-
E
08
--
Oentropyxia aerophila DEFLANDRE Oentropyxia aerophila v. minuta -
VAN QYE
Oentropyxis acul eata v. m inima
STEIN
Oentropyxia acul eata
DEFLANDHE
Areella vulgaris EHHENBERG AreeUa vulgaris f. reqularis
STEPAN~JK
Areella rotundata f. u ndulata
DEFLANDRE
Arcella roturulata P r,AYFAIR Arcella rotu n data v , sten ostom a
DEFLAND ltE
Arcella gibbo.9a P~: NAltI> A rcella hemi sphaerica PERTY Arcella hemisphaeri ca f . undulata
PLAY~'Am
Arcella diecoid es v . scuielliformie
S
07
SSP
06
P
05 16
04
02
01
+-
It
R
+
-
-
It
-
-
S
SW 25
.- - - - - - - - - - - - - -
21-40 %
F ueroer
11-20%; 2
SW SW SW SW SW SW SW SW SW
= 6-10 %; 1
Sh etlund
+-
SWSWSWSW SWSWSWSWSWSWSWSWSWSW
Table 5. Com position o f the fauna in the a q uat ic sam ples . R = found in less th an 6 % o f the samples; 3 = 41-60 % ; 4 = 61 -80 % ; 5 = 81-100 % . P = planetonic ; S = saprop eli c; E = ep ip hy t ic sample
r"
e,
ttl
'"e-
7\
t'l
~
l'J
I:l:l
~
-
PLAYFAl lt
---
-
-
R
--
--
-
-
.-
- _ ow
-
- . -
_
.
_
~
-
.._.-
- --
-
--
-
-
---
-
.-
-
-
-
-
-
- --
-
-
-
--'-
-
-
'- .
-
--
-
-
-
-
-
-
-_.
-
-
-
-- -
_.
-
-
-- -- -
--- -
---
-
-
_
-- - -
. _ --
-
--
_.
I
-
-
H -
- - - + - - - -- --- -- - - -- - - -.. _. - - _.- - - -. _ . - - - - - _. - -- -. - - - - - -~
_. _ .-
- - - - - - - --
- - - -
1
-
-
- -- -
_
----
- - ,_.• -- - -- -
2 -
2 _.- -
I
- --- - - - - -- - -- + - - - --- - - ..- - - "-.. -- - - -- - - - -_.. - -- -. - - - -. -. - - _. -- - - -
- --
:~
-- --_. - -- -
--
-.
-
-
- - e -- -- -
_._- _ . --- -
--- - -- -R - --
--
-
-
- - .-- - ---.. - - - -. --
'-
D ifflugia oentricosa DEFLA NDR E -DifJlugia tliffici iis T H OM A S D ifflugia diff1:eilis eurvata CHARDEZ Difflugia leidy W AI L E S Poniiqulasia eompressa RHUlIIB L E R Pontigulasia incisa R H UMB L E R Nebela dentistoma PE NARD N ebela collaris L E ID Y _..• N ebela f labellulum L E ID Y N ebela lagenijormis P E NARD
.._-
-
.-
---
-- -
- _..
_. _ ~
- -
-
2
1
-
-
-
- - -
-
_.
.-
J~
-
-- -
pulex PENARD paulii OGD EN pyriforme lla CHARD E Z rub esoen s PENAR D rotunda OGDEN scalp ellum P E N ARD tub erculata ,"V ALLIe H iubercu lat a v , sp haerica
--
--
- - .- -
.. -
_
-- R - - -
DiJflugia Difflugia Difflugia Difflugia DifJlugia DiJJlugia D ifflugia
DiJJl1~gia
T HOMAS
.TUNG
labiosa ' V Al LES minuta R AMPI parva ( THOMAS) O GDE N p enardi ( P E N AJtD)
&
HOPKINSON
Difflugia Difflugia D iJflugia D 1j f lugia
GAUTH.·L.
Difflugia lacus tris ( P E NARD) D iffl uqia Lu cida PENAR D D if f Tiugia lin eari s (PE N AR D )
LfHDY
Diff lugia eleqans v , bico rnis .TUNG D ifflugia gla ns PE N AR D Diff lugia qlobu lari« (WALL l e H )
- -."-
-
- - _. - - - -- - - - - - -- - - ..-
-- -
DELFANDJtfJ
DEFLANDJtE
-
--
+
Diff lugia eleqans v . a nqu sta ta
--
I
-
n
-
(PENA RD)
D ifflugia eleqa ns P E N ARD Diffluqia clegan s v . teres
.TUNG
Difflugia bryopMla 1
R
R
-
H
R
+ --
-
I R -
--
-
-
--
'-
-- -
-
-
-
-
2
+
-
R
-
H
R
R
R
R
It
R R R H R
-
-
-
-
---
I-t
-
-
-
R
-l-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
~ -- -
+ -
-
- - - -
-
-
-
-
-
-
-
-_.
1
-
-
-
-
-- -
-
-_.• -
-
-
-
.-
-
-
-
-
-
-
-
-
-
-
-
-
--- .
-
-
---
-
-
-
-
-
-
-
-
-
-
-
-
-
-
_. -
-
-
-
-
-
-
-
-
-
2
--
-
- - -- - - - R - + -
-
-
R
-
+
-
-
-
-
-
-
-
-
- -
-
H
- - R - - I _.. - -- -- -- -- - - - - --- -- -- - - -- - - - .- - - -- - - - - - - - - - - - - - - - - -
-
-
1
-
+ R I H - - 2
-
-
-
I
+
- -
- - -- - -- --
-
-
3
~
~
w
0'
l'.J
'" '..," "...
I-.j
:0-
~
...
;:I
'"
ta
::sp .
I>'
s,
H
0-
;:I
I;
0;-
<1>
:0-
IJJ
:3
0
::j>
~
0'
<'l>
0
~
<'l>
0;~ 0;-
'"
1-3 <1>
& THOlllAS
(HE RTWIG
& Lessun) HOl'KINSON
P hruqan ella aeropodia
LEIDY
T'rin ema comp lanat u m P E NAR D Sphenoderia minuta DEF LAN D R E Cyphoderia ampulla (EHR E N BE R G)
LEIDY
Euglypha compressa CART E R A ssulina muscorum GREEFF T'rinema lineare PENARD T'rin ema enchelys (EHRENBERG)
'VAlLES
Euglypha strig osa v . muscorum
LEIDY
Euglypha laevis PERTY Euglypha rotunda W AILES Euglypha strigosa (E H R E NBERG)
(EHU~:NB~:RG) PEIlTY
Euglypha acanth ophora
BUTSCHLI
Lesquereusia modesto R H UM BLER L esquereusia spiralis (E H R E NBE R G)
GAUTHn~Il-L .
L esquereueia globulosa
P~~ NARD
Quadrulella symmetrica W AL LleH Hyalosph enia subflava CASH H eleopera p etricola v , ame thys tea
I?:MW
N ebela penardiana DEFLAN D R E N ebela tincta (LEIDY) A W E R I N T -
Table 5 . continued
R
2
R
07
E
'T
08
P
09
S
R
10
E
5
11
P
12
S
2 1
3
13
P
It
-
14
S
R It
It It
R
It
It
15
E
+
- - -
+
+
It
2
-
S
18
-
R
-
10
P
21
22
-
H
H
It
-
-
SEE
20
24
3
+
S
25
I R
R R
S
-- +
23
P
17
06
S P
P
01) 16
01 S
02
SS
01
P
SW SW SW SW S W SW SW SW SW SW
._ - - - - - - - - - - - - -
Faeroer
S hetland
SW SW SW S W SW SW SW SW S W SW SW SW SW SW
~
~
.,..
~
t'J
><
t'J
tli
t-<
;e
Testate Amoebae from t he Shetland I sl an d s and the Faeroer
95
FAEROER
SHETLAND ARCELLA CENTROPYXIS DIFFLUGIA
f-I-------,
EUGLYPHA
NEBELA TRINEMA
o
20
40 %
o
20
40 %
F ig. 6. Histograms indicating the mo st important genera, a ccor d ing to their number of spe cies. This is ex pre ssed for each genus as a per centage o f the total n umber o f s pe cies .
5. The Tr inema lineare community, cha racteriz ed by the dominance of this species. Although this species occurs in other commu nit ies, it never rea ches suc h high va lues (> 40 %) t here. SW05, SWlO, SW21, SW16, SW23. A compa rison of these communities with the measured hydrogen-ion concent ra t ion and conduct ivit y proves that on ly two communities have limited ecological ranges. The Arcella hemisphaerica community is restricted to oligotrophic conditions, which is consist ent with most other dat a (SCHl)NRORN et al, 1965). Likewise, t he Di fflugia globularis communit y is only found in acid, nutrient-poor con dit ions. The Difflugia lacustris community on the other hand is observed in t he intermediate conductivity ranges, indicating a rather meso trophic occurrence. In this case t he habitats proved to be alka line to weakly acidi c. The wide cond uct ivit y ra nge occupied by t he Cyplwderia ampulla community and t he Trinema lineare community r ules out their usefulness in an ecological characterizat ion of the waters . At most sites both plankt oni c and sapropelic samples were taken, and it turned out, as one could expect, t hat in small basins both kinds of samples were similar. T his was obviously not t he case with the materia l from la rger basins.
Biogeographical as pects There is still a marked differen ce bet ween the aquat ic and t he terrestrial (moss lichen ) comm un it ies. As is known from literat ure (BEYENS et al. 1986 b), the differ ence becomes less evident in t he arct ic regions . On the other ha nd a diminution in t he importance of the Difflugia genus is already noticeable if we compare t he Shetland and the Faeroer records (F ig. 6). Thus the compos ition of t he fa una of t he lat t er archipe l already approaches t hat of I cela nd , and seems to hold an int erm ed iat e position between the moderate and the more boreal faunistic assemblages.
Note on some speci es Two spe cies, so far on ly recorded once , have been ob served again .
96
L. BEYENS et al., Testate Amoebae from the Shetland Islands and the Faeroer
Cyclopyxis pirini GOLEMANSKY Discribed by GOLEMANSKY and TODoRov (1985) from a mountain lake in Bulgaria. The observed specimens correspond well with the original description, and are mostly covered with diatoms, and have a sligthly invaginated aperture, surrounded by large irregular particles. They were encountered in aquatic samples (SWI9, SW20) from the Faerber.
Protoplagiopyxis ridderae CHARDEZ First observed by CHARDEZ (1969) in moss samples from Iceland, this species has been found again in the moss SM05 (Racomitrium lanuginosum) from Shetland. Diameter of shell: 66-73/-U11.
Acknowledgements We wish to thank Mrs. S. POOTERS, Mrs. A. TORFS, Mr. F. NEEFS and Mr. R. NEEFS for their technical assistance. We are also much indebted to Pro£. D. K. FERGUSON for correcting the English text. One of us (L. B) received a grant from the National Science Foundation (NFWO-Belgium) for the fieldwork involved.
Literature BEYENS, L., CHARDEZ, D., and DE LANDTSHEER, R., with collaboration of DE BOCK, P., and JACQUES, E. (1986a): Testate Amoebae Populations from Moss and Lichen Habitats in the Arctic. Polar BioI. 5: 165-173. - - - with collaboration of DE BAERE, D. (1986b): Testate Amoebae Communities from Aquatic Habitats in the Arctic. Polar BioI. 5: 197-205. CHARDEZ, D. (1969): Contribution it la Faune Thecamoebicnne de L'Islande. Bull. Inst. R. Sci. nat. Belg. 45: 1-16. CORLEY, M. F. V., CRUNDWELL, A. C., DULL, R., HILL, M. 0., and SMITH, A. J. E. (1981): Mosses of Europe and the Azores: an annoted list of species, with synonyms from the recent literature. J. Bryo!. 11: 609-689. ' DAGNELIE, P. (1975): Analyse statistique it plusieurs variables. Gembloux. DAHL, E., and KROG, H. (1973): Macrolichens of Denmark, Finland, Norway and Sweden. Oslo Bergen Trornso. GOLEMANSKY, B., and TODOROV, M. (1985): Thecamoebiens du sol de la Montagne Vitocha (Bulgaria). Acta Zool. Bulgarica 29: 50-63 (in Russian). HOFKER, J. (1942): Amoebida Testacea. Zoology of the Faroe, 1: 1-8. •JUNG, W. (1936): 'I'hekamoben urspriinglicher lebender deutscher Hochmoore. Abh. Landesmus. West£. 7: 1-87. MORRISON, D. F. (1967): Multivariate statistical methods. New York. SCHONBORN, W., FLOSSNER, D., und PROFT, G. (1965): Die Limnologische Charakterisierung des Profundals einiger norddeutscher Seen mit Hilfe von Testaceen-Gemeinsehaften. Limnologica (Berlin) 3: 371-380. SEAL, H. L. (1968): Multivariate statistical analysis for biologists. London. WAILES, G. H. (1912): Freshwater Rhizopoda from the Hebrides Orkney and Shetland Islands and Western Scotland with description of new species. Scott. Naturalist: 59-65. Authors' addresses: L. BEYENS, P. DE BOCK and E ..JACQUES, Leerstoel Algemene Plantkunde (Department of Botany), RUCA Groenenborgerlaan 171, B - 2020 Antwerpen, Belgium; D. DE BAERE, Department of Biology, U.LA., Universiteitsplein 1, B - 2610 Wilrijk, Belgium; D. CHARDEZ, Laboratoire de Zoologie Generale et Faunistique, Facult.e des Sciences Agronomi· ques de l'Etat, B - 5800 Gembloux, Belgium.