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Altitudinal distribution of non-cosmopolitan desmids and diatoms inPapua New GuineaWim Vyverman a
a Laboratorium voor Morfologie Systematiek en Ecologie van de Planten, Gent, Belgium
To cite this Article Vyverman, Wim(1992) 'Altitudinal distribution of non-cosmopolitan desmids and diatoms in PapuaNew Guinea', European Journal of Phycology, 27: 1, 49 — 63To link to this Article: DOI: 10.1080/00071619200650071URL: http://dx.doi.org/10.1080/00071619200650071
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Br. phycol. 3". 27:49-63 1 March 1992
Altitudinal Distribution of Non-cosmopolitan Desmids and Diatoms in Papua New Guinea*
By WlM VYVERMAN
Laboratorium voor Morfologie Systematiek en Ecologie van de Planten, K.L. Ledeganckstraat, 35, 9000 Gent, Belgium
About 20% of the diatom taxa and 27% of the desmid taxa from Papua New Guinea have a non-cosmopolitan distribution. A larger number of non-cosmopolitan desmids is confined to the Indo-Malaysian-North Australian region, non-cosmopolitan diatoms mainly being pantropical or northern montane. The altitudinal distribution of the different biogeographical elements was studied. Pantropical, palaeotropical and Indo-Malaysian-North Australian taxa are predominantly confined to lower and medium altitudes, northern montane taxa mainly to high altitudes. The altitudinal range from 1700 to 2500 m seems to be a transition zone between lowland and highland algal floras.
Desmids and diatoms are among the best- studied groups of freshwater algae in tropical regions. Several extensive studies have been made in the Indo-Malaysian and Nor th Australian region (e.g. Krieger, 1932; Behre, 1956; Bernard, 1908, 1909; Croasdale & Scott, 1976; Ling & Tyler, 1986; Thomasson, 1986; Scott & Prescott, 1958, 1961; Hustedt, 1937-1939, 1942; Thomas, 1983; Foged, 1971, 1976, 1978; Podzorski & Hfikansson, 1987; Prowse, 1962; Vyverman, 1989, 1991a,b). Few studies, however, have exam- ined the altitudinal distribution of freshwater algae (Krieger & Bourrelly, 1956). In the present paper the distributional aspects of non-cosmopolitan desmids and diatoms of Papua New Guinea are compared in relation to an altitudinal gradient.
M A T E R I A L S A N D M E T H O D S
Papua New Guinea is situated between the equator and 12°S, east of Indonesia and north of Australia. Climate is regulated by the SE/NW monsoon system. Most of the island receives between 2000 and 4000 mm rainfall year -l, with a maximum between January and April, when NW winds prevail (McAlpine, Keig & Falls, 1983). A mountain chain extending from west to east
*Laing Island Biological Station publication N 212.
0007-1617/92/010049+ 15 $03.00/0
divides the northern and southern lowlands, with peaks up to 4500 m.
Sampling was undertaken between 1986 and 1988 and mainly covered the northern and central part of the island. A wide variety of freshwaters was sampled, ranging from glacial lakes and tarns on the highest mountains, oligotrophic highland lakes, fast-flowing mountain rivers, salt and hot water springs, to dystrophic lowland backswamps and lakes, sediment-loaded rivers and lagoons and meromictic coastal lakes.
For each station several abiotic factors were measured: altitude, water temperature, con- ductivity, pH, Secchi disc transparency (for a full description of the methodology see Vyverman, 1991a,b). Species lists were made for each sample. Each taxon was given a distributional type according to recent literature (Ffrster, 1982; Ruzicka, 1977, 1981; Prescott et al., 1975, 1977, 1981; Croasdale, De Bicudo & Prescott, 1983; Croasdale & Flint, 1986, 1988; Gasse, 1986; Krammer & Lange-Bertalot, 1986, 1988 and numerous smaller publications; for a complete list of references used see Vyverman, 1991a,b). The categories considered here are: cosmopolitan, northern montane (in literature often referred to as preference for temperate and cool climates), pantropical, palaeotropical, Indo-Malaysian- North Australian and southern hemispheric.
RESULTS
A total of 287 samples from 154 different localities was analysed. A list of the sampling
© 1992 British Phycological Society
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50 W. Vyverman
TABLE 1. The table presents following information: number of periphyton/benthos samples, plankton samples and totals; number of samples taken from different types of water body (1, lake; 2, swamp; 3, spring or bog; 4, river; 5, ephemeral water bodies). Temperature (T), conductivity (K20) and pH data are divided into classes. The classes represent all the water
bodies sampled
Temperature Nature Type (°C) K20 (~tS.cm- l) pH
0-500 m P/B 84 1 64 < 15 0 0-50 24 < 6.0 12 P1 49 2 41 15-20 0 50-100 39 6.0~.5 12
- - 3 1 20-25 1 100-200 25 6.5-7.0 34 Total 133 4 35 25-30 64 200--300 32 7.0-7.5 28
5 2 30-35 63 300-1000 22 7.5-8.0 13 > 35 5 > 1000 1 > 8'0 16
No data 10 No data 28
500-1000m
I000-1500m
1500-2000 m
2000-2500 m
2500-3000 m
> 3000 m
P/B 20 l 26 < 15 0 0-50 0 < 6-0 0 PI 10 2 0 15-20 0 50-100 0 6.0-6.5 0
- - 3 1 20-25 2 100-200 13 6-5-7.0 2 Total 30 4 3 25-30 28 200-300 8 7-0-7.5 7
5 0 30-35 0 300-1000 8 7-5-8.0 8 > 35 0 > 1000 1 > 8-0 7
No data 0 No data 6
P/B 26 1 29 < 1 5 0 0-50 7 <6 .0 2 P1 10 2 2 15-20 1 50-100 13 6.0-6.5 4
- - 3 0 20-25 6 100-200 12 6.5-7.0 16 Total 36 4 4 25-30 25 200-300 2 7.0-7.5 8
5 t 30-35 4 300-1000 2 7.5-8.0 4 > 35 0 > 1000 1 > 8.0 2
No data 0 No data 0
P/B 15 1 22 < 15 0 0-50 10 < 6-0 3 PI 11 2 3 15-20 2 50-100 7 6.0-6.5 2
- - 3 0 20-25 14 100-200 4 6.5-7.0 16 Total 26 4 0 25-30 10 200-300 4 7.0-7.5 0
5 1 30-35 0 300-1000 1 7.5-8-0 5 > 35 0 > 1000 0 > 8.0 0
No data 0 No data 0
P/B 17 1 23 < 15 3 0-50 12 < 6.0 8 P1 8 2 0 15-20 9 50-100 8 6.0~.5 0
- - 3 1 20-25 11 100-200 2 6.5-7.0 8 Total 25 4 1 25-30 2 200-300 2 7.0-7.5 3
5 0 30-35 0 300-1000 1 7-5-8.0 2 > 35 0 > I000 0 > 8-0 2
No data 0 No data 2
P/B 5 1 4 < 15 1 0-50 0 < 6.0 0 P1 1 2 1 15-20 5 50-100 5 6.0-6.5 0
- - 3 1 20-25 0 100-200 0 6.5-7.0 I Total 6 4 0 25-30 0 200-300 0 7.0-7.5 4
5 0 30-35 0 300-1000 0 7.5-8.0 1 > 35 0 > 1000 1 > 8-0 0
No data 0 No data 0
P/B 16 1 22 < 15 I0 0-50 19 < 6.0 0 P1 5 2 3 15-20 5 50-I00 1 6.0-6.5 6
- - 3 0 20-25 3 100-200 0 6.5-7-0 9 Total 21 4 0 25-30 0 200-300 1 7.0-7.5 1
5 1 30-35 0 300-1000 0 7.5-8.0 0 > 35 0 > 1000 0 > 8.0 0
No data 3 No data 5
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Altitudinal distribution of algae 51
Sout~ hern-hem isphere 0-1% Pantropical 8.4% Palaeotropical 5.4%
Cosmopolitan 77-7% Indo-Malaysian - North Australian 4.5% Northern-montane 5,9%
( a )
~Sou the rn -hemisphe re 0.3% llIHIllllllllllIS Vantrop ca 6 °,o
Cosmopolitan 71% / ~ Palaeotropical 4-.1%
\ILLLLLILLILLllLIIII~ Indo- Malaysian -
~ Northern-montane 2-4%
( b )
FIG. 1. The percentage composition of Papua New Guinea diatom (a) and desmid (b) floras in terms of biogeographical distribution patterns. Following types are distinguished: cosmopolitan, northern montane, Indo-Malaysian-North Australian, palaeotropical, pantropical, southern hemispheric. Absolute numbers of taxa per biogeographical element are given in the text.
stations and measurements of abiotic factors have been published elsewhere (Vyverman, 1991a,b). Table I summarizes some habitat characteristics of the localities sampled and the nature of the samples in each altitudinal zone.
Biogeographical composition of the desmid and diatom floras
In total 430 diatom and 429 desmid taxa were found. A detailed list of all taxa, their ecology and distribution is published else- where (Vyverman, 1988, 1991a-c, Vyverman & Compare, 1991).
Figure 1 shows the composition of both floras in terms of distributional patterns. The percentages shown only relate to those taxa for which the geographical distribution is known. Excluded from the analysis were unidentified and new taxa as well as little- known taxa with an insufficiently known distribution. For desmids the number of excluded taxa was 47 (11% of the total number) and for diatoms the number was 72
(16.7% of total). Of the 383 diatoms with a known distribution, 307 (80%) are cosmo- politan. Only 76 taxa (19-8%) are non- cosmopolitan and belong to the following types: pantropical (36), northern montane (20), Indo-Malaysian-North Australian (13), palaeotropical (4) and southern hemispheric (3).
Of the 357 desmid taxa with a known distribution, 260 (73%) are cosmopolitan. The remaining 97 (27%) non-cosmopolitan taxa consist of those confined to the Indo-Malaysian-North Australian region (51) followed by those with pantropical (27), palaeotropical (9), northern montane (9) and southern hemispheric (1) distributions.
Tables II and III show the portion of each distribution type per genus. Only genera with at least one non-cosmopolitan taxon in Papua New Guinea are included. A few diatom genera confined to cooler climates or which have their main distribution there, are present in the data set: Diatoma, Diatomella, Peronia. The genera Neidium, Eunotia and Aulacoseira also have high portions of non-
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tJI
1,4
TA
BL
E I
I.
Th
e p
erce
nta
ges
an
d
nu
mb
ers
of
tax
a in
eac
h
dis
trib
uti
on
ty
pe
of
dia
tom
g
ener
a fo
un
d
in P
apu
a N
ew
Gu
inea
w
ith
at
le
ast
on
e n
on
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smo
po
lita
n
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on
Ind
o-
Mal
aysi
an-
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rth
ern
S
ou
ther
n
No
rth
To
tal
Co
smo
po
lita
n
mo
nta
ne
hem
isp
her
e P
antr
op
ical
A
ust
rali
an
Pal
aeo
tro
pic
al
Gen
us
N
N
%
N
%
N
%
N
%
N
%
N
%
Ach
nant
hes
18
11
61
.1
2 11
.1
--
--
2 1
1.I
3
16
.7
--
--
Am
phor
a 8
7 8
7-5
.
..
..
.
1 12
"5
--
--
Aul
acos
eira
12
7
58
.3
3 2
5.0
--
--
2
16
.7
..
..
Cal
onei
s 9
8 8
8.9
I
11.1
.
..
..
..
Cym
bella
2
4
18
75
.0
3 1
2.5
--
--
3
12
.5
--
--
--
Dia
tom
a 2
--
--
2 1
00
.0
..
..
..
.
Dia
tom
ella
1
--
--
1 1
00
-0
..
..
..
.
Dip
lone
is
3 2
66
.7
..
..
I
33
.3
--
--
--
Epi
them
ia
8 7
87
.5
..
..
1
12.5
.
..
.
Eun
otia
51
3
7
72
.6
4 7-
8 --
--
10
19
-6
..
..
Gom
phon
ema
16
13
81
.2
..
..
3
18-8
.
..
.
Fra
gila
ria
10
9
90
.0
..
..
1
10
.0
..
..
Fru
stul
ia
7 5
71
.4
--
--
2 2
8.6
.
..
..
.
Nav
icul
a 91
7
9
86
.8
2 2
-2
--
6 6
.6
I 1.
1 3
3.3
Nei
dium
11
8
72
.7
1 9.
1 --
1
9.1
--
--
1 9.
1
Nitz
schi
a 3
2
31
96
-9
..
..
1
3.1
--
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onia
1
--
--
1 1
00
.0
..
..
..
.
Pin
nula
ria
29
2
4
82
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..
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5
17
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palo
dia
7 6
85"7
--
--
1
14.3
.
..
..
.
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is
10
8 8
0.0
.
..
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0.0
1
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0.0
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0-0
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rella
19
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8
4'2
.
..
..
3
15"8
--
--
,,q
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mid
g
ener
a p
er d
istr
ibu
tio
n
typ
e.
On
ly
gen
era
wit
h
at l
east
on
e n
on
- co
smo
po
lita
n
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on
in
Pap
ua
New
G
uin
ea
are
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Mal
aysi
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rth
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ou
ther
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ph
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tro
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stra
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alae
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enu
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N
%
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N
%
N
%
N
%
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TAB
LE I
V.
The
dis
trib
utio
nal
rang
e of
non
-cos
mop
olit
an d
iato
ms
in r
elat
ion
to a
ltit
ude.
The
alt
itud
e co
rres
pond
ing
wit
h 50
%
of t
he o
ccur
renc
es (
med
ian
alti
tude
) is
in
dica
ted
by a
'•'
. F
or t
hat
purp
ose
the
alti
tudi
nal
rang
e w
as d
ivid
ed i
nto
inte
rval
s of
100
m.
The
num
ber
of s
ampl
es i
n w
hich
it
was
fou
nd i
s gi
ven
betw
een
brac
kets
, fo
llow
ed b
y it
s oc
curr
ence
in
rela
tion
to
cond
ucti
vity
(K
20)
and
tem
pera
ture
K20
T
empe
ratu
re
0 N
0z
S.c
m- t
) (o
c)
• I
2 3
km
• •
•
Nor
ther
n M
onta
ne:
Ach
nant
hes
alta
ica
(6)
11-3
8 12
-20
Ach
nant
hes
mar
ginu
lata
(4
) 33
-39
12-1
3 A
ulac
osei
ra d
ista
ns
(46)
7-
272
12-3
3 A
ulac
osei
ra d
ista
ns v
ar.
alpi
gena
(1
4)
7-69
12
-26
Aul
acos
eira
lir
ata
var.
ser
iata
+ la
cust
ris
(5)
7-21
5 12
-26
Cal
onei
s te
nuis
(4
) 15
-62
13-1
8 C
ymbe
lla a
equa
lis
(3)
15-5
4 15
-20
Cym
bella
lun
ata
(72)
7-
283
12-4
0 C
ymbe
lla p
erpu
silla
(2
3)
7-90
0 13
-32
Dia
tom
a hi
erna
le
(7)
13-6
6 12
-18
Dia
tom
a m
esod
on
(5)
13--
66
12-1
8 D
iato
mel
la b
alfo
urni
ana
(1)
126
26
Eun
otia
bid
entu
la v
ar.
elon
gata
(1
0)
7-65
13
-21
Eun
otia
big
ibba
var
. pu
mil
a (4
) 12
-69
13-1
6 E
unot
ia d
iodo
n (1
8)
7-85
12
-24
Eun
otia
ros
tella
ta
(4)
12-3
3 14
-15
Nav
icul
a ga
ndru
pii
(2)
33-2
43
13-2
4 N
avic
ula
stro
emii
(5)
37-2
83
12-2
9 N
eidi
urn
bisu
lcat
um
(4)
12-6
9 13
-15
Per
onia
fibu
la
(5)
11-3
3 15
-20
Sout
hern
Hem
isph
eric
: F
rust
ulia
mag
alie
smon
tana
F
rust
ulia
rho
mbo
ides
var
. el
onga
tissi
ma
Rho
palo
dia
nova
e-ze
alan
diae
Pant
ropi
cah
Ach
nant
hes
infla
ta
Ach
nant
hes
oblo
ngel
la
Aul
acos
eira
aga
ssiz
ii va
r. m
alay
ensi
s A
ulac
osei
ra i
kapo
ensi
s va
r. p
roce
ra
Cym
bella
mue
lleri
C
ymbe
lla s
picu
la
Cym
bella
tur
nida
D
iplo
neis
sub
oval
is
Epi
them
ia c
istu
la
Eun
otia
cam
elus
E
unot
ia c
amel
us v
ar.
karv
eern
sis
(5)
33-6
2 13
-16
(14)
10
-131
13
-36
(3)
39-2
83
12-1
6
(20)
28
-528
13
-32
- -
• (1
) 13
1 31
•
(1)
30
27
• (1
) 24
31
•
(73)
15
-700
16
-36
• (2
4)
7-26
0 19
-29
(22)
56
--40
0 18
-31
-- •
(8)
62-1
"9
13-3
0 (1
) 35
2 28
•
(17)
24
-313
27
-36
• (3
) 23
-256
27
-33
•
Downloaded At: 03:09 22 May 2010
Pant
ropi
cai:
Eun
otia
cam
elus
var
. ve
ntri
cosa
E
unot
ia d
idym
a va
r. c
lavi
cula
ta
Eun
otia
dis
sim
ilis
Eun
otia
mon
odon
vat
. tr
opic
a E
unot
ia r
aben
hors
tiana
E
unot
ia r
aben
hors
tii v
ar.
mon
odon
E
unot
ia t
schi
rchi
ana
Eun
otia
zyg
odon
F
ragi
lari
a st
rang
ulat
a G
omph
onem
a af
fine
G
omph
onem
a au
gur
var.
tur
ris
Gom
phon
ema
tene
rrim
um
Nav
icul
a la
gerh
eim
ii N
avic
ula
lage
rhei
mii
var.
int
erm
edia
N
avic
ula
lepi
dula
N
avic
ula
mut
icoi
des
Nav
icul
a pe
rrot
etti
i N
avic
ula
sem
inul
oide
s N
eidi
um g
raci
le
Pin
nula
ria
acro
spha
eria
P
innu
lari
a ac
rosp
haer
ia v
ar.
turg
idul
a P
innu
lari
a br
evic
osta
ta v
ar.
sum
atra
na
Pin
nula
ria
lucu
lent
a P
innu
lari
a ri
vula
ris
Stau
rone
is m
inut
ula
Pala
eotr
opic
ah
Nav
icul
a in
vict
a N
avic
ula
rutt
neri
N
avic
ula
sem
inul
oide
s va
t. s
umat
rana
N
eidi
um g
raci
le f
orrn
a ae
qual
is
Indo
-Mal
aysia
n-N
orth
Aus
tral
ian:
A
chna
nthe
s cr
enul
ata
Ach
nant
hes
cren
ulat
a va
r. l
inea
ris
Ach
nant
hes
mon
tana
var
. tr
opic
a A
mph
ora
subt
urgi
da
Coc
cone
is b
revi
cost
ata
Coc
cone
is c
atar
acta
cum
N
avic
ula
curt
a N
itzs
chia
wol
tere
ckii
St
auro
neis
ten
era
Sten
opte
robi
a pe
lagi
ca
Suri
rell
a ce
lebe
sian
a Su
rire
lla
subl
inea
ris
Suri
rell
a te
nuis
sim
a
(4)
30-9
3 (1
3)
56-3
13
(4)
25-7
8 (7
) 28
-131
(5
9)
25-1
90
(1)
18o
(9)
240-
430
(19)
14
-131
(2
) 13
1 (9
0)
90q5
31
(4)
25-7
5 (1
1)
40-4
35
(2)
62-2
64
(3)
78-5
28
(1)
94
(36)
7-
528
(19)
28
-900
(2
6)
26-5
90
(9)
24-2
69
(46)
7-
1.8
(3)
24-1
31
(14)
7-
264
(1)
26
(1)
14
(1)
528
27
27-3
2 13
-30
27-3
6 23
-32
23
28-3
1 27
-40
36
23-3
1 28
-30
18-3
4 29
-33
29-3
6 30
13
-36
27-3
2 26
-36
28-3
2 21
-36
28-3
6 21
-36
33
40
30
(8)
26-5
28
27-3
6 •
- -
(1)
446
29
• (1
2)
25-5
28
28-3
0 •
- -
- (4
) 24
-76
28-3
1 •
(27)
(2
) (2
9)
(2)
(5)
(1)
(3)
(10)
(4
) (1
0)
(1)
(1)
(17)
110-
230
94-1
99
94
210-
352
54-9
8 15
73
18
4-59
0 35
-830
24
-131
18
1 13
5 24-8
5
20-2
6 19
-28
19
28-3
0 25
-31
18
26
25-3
5 26
-30
27-3
6 30
23
14
-29
m. ~d
-1
0 (JI
t~
Downloaded At: 03:09 22 May 2010
TAB
LE V
. T
he T
he d
istr
ibut
iona
l ra
nge
of n
on-c
osm
opol
itan
des
mid
s in
rel
atio
n to
alt
itud
e. T
he a
ltit
ude
corr
espo
ndin
g w
ith
50%
of
the
occu
rren
ces
(med
ian
alti
tude
) is
in
dica
ted
by a
'•'
. F
or t
hat
purp
ose
the
alti
tudi
nal
rang
e w
as d
ivid
ed i
nto
inte
rval
s o
f 10
0 m
. T
he n
umbe
r of
sam
ples
in
whi
ch i
t w
as f
ound
is
give
n be
twee
n br
acke
ts,
foll
owed
by
its
occu
rren
ce i
n re
lati
on t
o co
nduc
tivi
ty (
K20
) an
d te
mpe
ratu
re (
T)
K20
T
empe
ratu
re
0 1
2 3
km
N
(~tS
.cm
-t)
(°C
) •
..
..
&
. •
•
Nor
ther
n M
onta
ne:
Act
inot
aeni
um
adel
ocho
ndru
m
Aet
inot
aeni
um
adel
oeho
ndru
m v
ar.
krie
geri
C
lost
eriu
m c
osta
tum
var
. bo
rgei
C
osm
ariu
m l
evin
otab
ile
Cos
mar
ium
mon
omaz
um
Cos
mar
ium
nas
utum
C
osm
ariu
m q
uadr
atum
E
uast
rum
obl
ongu
m
Mic
rast
eria
s cr
ux-m
elit
ensi
s
Sout
hern
Hem
isph
eric
: St
aura
stru
m
sagi
ttar
ium
Pant
ropi
eal:
Act
inot
aeni
um
capa
x va
r. m
inus
C
lost
eriu
m n
emat
odes
C
lost
eriu
m n
emat
odes
var
, pr
obos
cide
um
Cos
mar
ium
dep
ress
um v
ar.
elev
atum
C
osm
ariu
m n
itid
ulum
var
. jav
anic
um
Cos
mar
ium
qua
drum
C
osm
ariu
m s
eely
anum
C
osm
ariu
m s
ubgl
obos
um v
ar.
sum
atra
num
C
osm
ariu
m t
rach
ypol
um
Cos
mar
ium
var
iola
tum
vat
. ro
tund
atum
E
uast
rum
pra
emor
sum
E
uast
rum
spi
nulo
sum
E
uast
rum
spi
nulo
sum
var
. in
erm
ius
lcht
hyoc
ercu
s lo
ngis
pinu
s M
icra
ster
ias
apic
ulat
a va
r. l
acer
ata
Mic
rast
eria
s fo
liac
ea
Mic
rast
eria
s fo
liac
ea v
ar.
orna
ta
Mic
rast
eria
s ra
dian
s M
icra
ster
ias
trop
ica
Ony
chon
erna
lae
ve v
ar.
latu
m
Ple
urot
aeni
um m
inut
um v
ar.
suba
tten
uatu
m
(6)
7-16
19
-26
(1)
7 21
(1
) 16
26
(1
) 12
15
(2
) lO
22
(4
) 16
-39
12-2
6 (8
) 11
-39
12-2
0 (5
) 38
13
(4
) 10
-85
13-2
2
(3)
11-1
2 15
-20
(3)
11-3
13
21-2
7 (4
) 24
-131
25
-36
(1)
24
26
(1)
131
36
(1)
26
33
(14)
24
-131
25
-32
(4)
552-
700
27-3
2 (2
) 63
-131
25
-36
(2)
14-1
31
36-4
0 (5
) 95
-553
28
-32
(6)
24-1
80
18-3
0 (7
) 26
-85
19-3
1 (2
) 32
-84
29-3
0 (1
) ll
21
(1
) 13
1 36
(3
) 16
-131
23
-36
(4)
32-1
73
25-3
0 (8
) 16
-161
23
-30
(1)
63
28
(5)
28-9
2 16
-30
(1)
28
33
,<
Downloaded At: 03:09 22 May 2010
Stau
rast
rum
cer
aste
s va
r. p
ulch
rum
St
aura
stru
m o
rbic
ular
e va
r. d
enti
cula
tum
St
aura
stru
m s
onth
alia
num
St
auro
desm
us a
rcua
tus
Teil
ingi
a qu
adri
spin
ata
form
a ev
olut
a X
anth
idiu
m a
rmat
ure
var.
ang
ulig
erum
Pala
eotr
opic
ah
Cos
mar
ium
ask
enas
yi
Cos
mar
ium
per
fiss
um
Cos
mar
ium
str
iola
tum
var
. no
rdst
edti
i E
uast
rum
moe
bii
Mic
rast
eria
s ze
ylan
ica
Stau
rast
rum
lon
gibr
achi
atum
St
aura
stru
m l
ongi
brac
hiat
um
var.
jav
anic
um
Stau
rast
rum
wil
dem
anii
X
anth
idiu
m s
ubtr
ilob
um v
ar.
inor
natu
m
Indo
-Mal
aysia
n-N
orth
Aus
tral
ian:
C
lost
eriu
m r
ecti
mar
gina
tum
C
osm
ariu
m b
lytt
ii f
orm
a au
stra
licu
m
Cos
mar
ium
bur
kill
ii v
ar.
depr
essu
m
Cos
mar
ium
cey
lani
cum
C
osm
ariu
m d
ubiu
m
Cos
mar
ium
exa
sper
atum
C
osm
ariu
m fr
eem
anff
var
. ve
rruc
osum
C
osm
ariu
m m
ikro
n C
osm
ariu
m n
udum
C
osm
ariu
m o
tus
var.
orn
atum
C
osm
ariu
m p
andu
rifo
rme
Cos
mar
ium
per
igra
nula
tum
C
osm
ariu
m p
seud
oarm
atum
C
osm
ariu
m s
umat
ranu
m
Cos
mar
ium
tax
icho
ndru
m v
ar.
pulc
hrum
C
osm
ariu
m t
jiben
onge
nse
form
a m
inus
C
osm
ariu
m t
rach
yple
urum
var
. no
rdst
edti
i D
esm
idiu
m b
aile
yii f
orm
a te
trag
onum
E
uast
rum
dis
tort
um
Eua
stru
m g
nath
opho
rum
E
uast
rum
hor
ikaw
ae
Eua
stru
m l
ongi
colle
vat
. ca
pita
tum
E
uast
rum
moe
bii
var.
bur
men
se
Eua
stru
m m
oebi
i va
t. t
etra
chas
trif
orm
e
(I)
160
(6)
16-1
80
(10)
7-
221
(10)
7-
161
(1)
131
(3)
16
26
23-3
6 23
-34
19-2
6 36
23
(11)
10
-180
22
-30
• (1
) 93
28
•
(7)
13-1
80
19-2
9 •
(1)
11
21
• (8
) 16
-131
23
-36
• (7
) 7-
161
15-2
9 •
(1)
13
19
• (4
) 13
-161
23
-32
.•
(3)
28-1
31
30--
36
•
36
36
23-3
6 29
36
23
-32
36
30-3
3 29
23
-29
27
28
26--
28
36
23
29
19
27-3
6 21
-26
21-2
3 29
29
-30
29-3
1 25
-27
(1)
131
(1)
131
(6)
25-1
80
(1)
84
(1)
131
(6)
12-4
7 (1
) 13
1 (2
) 26
-28
(3)
25-8
4 (9
) 16
-18o
(2
) 30
-45
(1)
93
(4)
56-1
80
(1)
131
(3)
18-1
80
(2)
25
(1)
13
(4)
26-1
31
(6)
7-16
(5
) 11
-16
(2)
63-8
4 (2
) 28
-84
(3)
27-6
3 (2
) 24
-63
g~
O K
Downloaded At: 03:09 22 May 2010
o~
TAB
LE V
. co
nt.
N
K20
T
empe
ratu
re
0 (l
aS.c
m- ~
) (°
C)
• 1
2 3
• •
• km
Mic
rast
eria
s lu
x M
icra
ster
ias
mah
abul
eshw
aren
sis
vat.
red
ucta
M
icra
ster
ias
subi
ncis
a P
leur
otae
nium
cor
onif
erum
var
. m
ulti
nodu
losu
m
Ple
urot
aeni
um k
ayei
P
leur
otae
nium
ova
tum
var
. tu
mid
um
Stau
rast
rum
con
tect
um
Stau
rast
rum
ehr
enbe
rgia
num
va
r. r
ostr
atum
St
aura
stru
m e
nsif
erum
St
aura
stru
m fr
eem
anii
St
aura
stru
m g
raei
le f
orm
a kr
iege
ri
Stau
rast
rum
gut
win
skii
St
aura
stru
m g
utw
insk
ii v
ar.
evol
utum
St
aura
stru
m ja
vani
cum
St
aura
stru
m p
rote
ctum
var
. ra
ngoo
nens
e St
aura
stru
m p
layf
airi
St
aura
stru
m p
seud
ozon
atum
St
aura
stru
m w
ilde
man
ii v
ar.
maj
us
Stau
rasl
rum
zon
atum
vat
. m
ajus
St
auro
desm
us g
ibbe
rulu
s St
auro
desm
us g
ibbe
rule
s va
r. m
ucro
natu
s X
anth
idiu
m a
cant
hoph
orum
var
. ra
cibo
rski
X
anth
idiu
m a
picu
latu
m
Xan
thid
ium
has
tife
rum
var
. jav
anic
um
Xan
thid
ium
mul
tico
rne
Xan
thid
ium
sex
mam
illa
tum
var
. pu
lney
ense
X
anth
idiu
m s
ubtr
ilob
um
(4)
16-1
60
23-3
0 •
(2)
32-1
31
30-3
6 •
(4)
14-1
31
30-4
0 •
(1)
28
30
• (6
) 11
-32
21-3
0 •
(4)
16-6
3 23
-25
-- •
--
(1)
28
28
• (1
) 24
28
•
(4)
16-1
61
25-2
6 •
(3)
16-4
6 23
-29
• (1
) 55
2 27
•
(15)
10
-161
21
-31
• (1
) 28
30
•
(2)
94-1
18
30-3
2 •
(1)
63
25
• (1
9)
26-2
11
27-3
6 •
(2)
30-9
2 27
-28
• (1
4)
12-1
60
19-2
9 •
(9)
13-1
73
19-3
3 •
(4)
78-7
00
29-3
2 •
(2)
24-3
2 30
•
(i)
25
29
• (4
) 10
-160
22
-26
• (5
) 24
-118
29
-30
• (1
1)
12-8
4 19
-30
• (7
) 25
-700
25
-29
• (7
) 24
-131
29
-36
•
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Altitudinal distribution of algae 59
2 "S
E Z
5O
4O
50
2O
I0
0 0,1 0.5
7O
6O
1 1,5 2 2,5 3 5-5
50
40
5O
2O
I0
o 0.1 0.5 I 1,5 2 2.5 5 5.5
Altitude (1000 m. a.s.l.)
Northern-montane ~ Pantropical
PalaeotropJcal F7777 ~ Indo-Mala ysian- Australian
F16. 2. Numbers of northern montane, pantropical, palaeotropical and Indo-Malaysian-North Australian diatom (a) and desmid (b) taxa along the altitudinal gradient.
cosmopolitan taxa. Among desmids, non- cosmopolitan taxa are mainly found in the genera Cosmarium, Euastrum, Micrasterias, Pleurotaenium and Xanthidium. No desmid genera exist with a predominantly northern montane distribution. In contrast, the genus Ichthyocercus occurs only in the tropics.
Distribution of non-eosmolmlitan taxa in relation to altitude
In Tables IV and V the distributional range of the non-cosmopolitan diatoms and desmids is plotted vs. altitude. In Fig. 2 the numbers of taxa per biogeographical type (with the exclusion of the southern hemi- spheric type) are plotted vs. altitude. For each taxon its complete range of occurrence
was taken into account, even if it was not found in a given altitudinal interval within its range.
Of the 20 northern montane diatom taxa, 14 were confined exclusively to water bodies sampled above 1700 m. Aulacoseira distans, Cymbella lunata and C. perpusilla occurred over a wide altitudinal range from sea level up to over 3500 m with a median altitude around 2000 m. Diatomella balfourniana, Navicula gandrupii and N. stroemii were found from about 800 m to over 3500 m. Two of the three southern hemispheric taxa were only observed in mountain lakes. Frustulia rhomboides var. elongatissima was found over the entire altitudinal range. With a few exceptions (e.g. Cyrnbella muelleri, Eunotia dissimilis, Cocconeis cataractacum,
Downloaded At: 03:09 22 May 2010
60 W. Vyverman
Surirella tenuissima), the pantropical palaeo- tropical and Indo-Malaysian-North Australian elements are mainly found in the lowlands or in waters up to 2000-2400 m altitude. The conductivity ranges show no clear relationship with distribution type.
The changes in the biogeographical affini- ties of the flora are illustrated in Figure 2(a). The pantropical element shows a steep drop above 2300 m, the Indo-Malaysian-North Australian element drops more gradually from 1200 m on. The palaeotropical element was only found up to 500 m. In all three categories the greatest number of taxa occurred in the lowlands below 100 m. The number of northern montane taxa increases gradually with alti- tude, with a maximum above 2400 m.
No northern montane desmids were found in lowland waters. Six of them occurred no lower than 1500 m and the remaining taxa no lower than 1900 m. The single southern hemispheric taxon Staurastrum sagittarium was found in a high-altitude lake. Pantropical, palaeotropical and Indo- Malaysian-North Australian taxa are mainly found from sea level up to 1800-2300 m. Only Onyehonema laeve var. latum and Staurastrum longibrachiatum also occurred at higher altitudes. With a few exceptions, most taxa were found at low to medium conductivities.
Like the diatoms, the largest number of pantropical, palaeotropical and Indo- Malaysian-North Australian desmids were found below 100 m [Fig. 2(b)]. From 100 m, they show a more or less even distribution up to 1700 m; at higher altitudes their numbers decrease, reaching a minimum between 2300 and 2500 m. The distribution of northern montane desmids shows two peaks, one at 2000 and one at 3200 m.
DISCUSSION
Most of the desmids and diatoms in Papua New Guinea have a world-wide or cosmo- politan distribution. The degree of cosmo- politanism is comparable to other tropical regions. In the Lake Chad region, for
example, the cosmopolitan and subcosmo- politan element amounts to 71% of the algal flora (Compare & Iltis, 1983).
Of the desmids with a known distribution in our data set, 26.3% are confined to tropical regions; of diatoms, this is the case for only 16"3%. The northern montane element represents 2.4% of the desmids and 5.9% of the diatoms. This is consistent with the published literature in which relatively few desmids have been described as prefer- ring cooler water whereas northern montane diatoms are much more numerous. In addi- tion, the desmid genus Ichthyocercus has a pantropical distribution. Conversely, diatom genera like Diatoma, Peronia and Tabellaria seem to have their main distribution in more temperate regions. Furthermore, most non- cosmopolitan diatoms belong to genera known to oligotrophic habitats. In contrast, genera known from more eutrophic waters, such as Amphora, Nitzschia, Fragilaria, Epithemia and Rhopalodia, have relatively more taxa with a worldwide distribution.
A large number of desmids (15.7%) in the data set has an Indo-Malaysian-North Australian distribution. The existence of such an element in the flora has been demon- strated by several authors (e.g. Krieger, 1932; Behre, 1956; Thomasson, 1986). With diatoms the Indo-Malaysian-North Australian element is much less distinct (4.5% of the taxa with a known distribution type), the pantropical element being more important (8.4%). In fact, many of Hustedt's (1937-1939, 1942) proposed types of endemisms have been falsified because of incorrect taxonomy and information lacking on diatom distribution patterns. Recent and more extensive collecting has revealed that many of the species mentioned by Hustedt have a wider distribution then previously thought. Nonetheless, a number of clear-cut taxa, like Achnanthes erenulata and Surirella celebesiana, indicate the existence of a distinct Indo-Malaysian-North Australian element in diatoms. The different numbers of Indo-Malaysian-North Australian and pantropical taxa suggest that some differ- ences must exist in the dispersal capacities
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Altitudinal distribution of algae 61
and/or speciation rate of diatoms and desmids. However, very little is actually known about dispersal mechanisms of microalgae.
Wind dispersal (Round, 1981: 357; John, 1986: 158), rather than dispersal by birds, seems to be more likely in the case of pan- tropical taxa, as most migratory routes follow a N-S direction.
For the palaeotropical element, a some- what similar distribution is shown by rotifers (Dumont, 1983). Large-scale air circulation patterns over the Indian Ocean region might act as an important transport agent. An alternative hypothesis is that climatic and vegetational changes during the Pleistocene may have formed temporal links between Africa and Asia due to the presence of suit- able biotopes (Dumont, 1983).
Bird migration (Proctor, 1966; Atkinson, 1972, 1980) may be invoked as a more important factor than wind dispersal to explain the Indo-Malaysian-North Australian element in Papua New Guinea. Indeed, although few data exist on migration of water fowl in this part of the world (Beehler et al., 1983; Parish, 1989), pre- liminary data suggest that the South-East Asian continent, the Sunda Islands, the Philippines and North Australia lie on the most important routes for migrating birds in this part of the world. In addition, a number of Indo-Malaysian-North Australian taxa has been found in more temperate regions of China and Japan. A similar feature was also observed by Coesel, Duque & Arango (1988), confirming the idea that migratory birds may be responsible for the dispersal of neotropical desmid species to temperate North America.
The distribution of the different biogeo- graphical elements in Papua New Guinea shows a distinct relationship with altitude and water temperature (Tables IV and V). Indo-Malaysian-North Australian, pan- tropical and palaeotropical elements amongst diatoms and desmids are in Papua New Guinea mostly confined to the lowlands and mountains below about 1700- 2500 m, indicating their warm-stenothermic
(Uherkovich, 1983: 304) nature. The greatest diversity of these elements was found in the black- and mixed waters of the Sepik flood- plain at an altitude below 50 m. Many of them, however, are very rare in the study area and were only found in small numbers. Several taxa seem to have a wider amplitude with respect to temperature, such as Cymbella muelleri, Eunotia dissimilis and Staurastrum longibrachiatum.
Above 1700-2500 m, northern montane taxa largely replace the tropical flora elements. Most northern montane desmids and diatoms are mainly found in the high- lands, which points to their cold-steno- thermic nature (Round, 1981: 382). Gasse, Tailing & Kilham (1983: 19) suggested that suitable, oligotrophic conditions are more important than low temperatures. In the northern hemispheres, where most of the taxonomic and ecological studies on algae have been carried out, such habitats are increasingly restricted to high-latitudes and montane regions which might explain the northern montane distribution of these taxa. This hypothesis is only partially supported by our data, as in the case of Cymbella lunata, C. perpusilla and Aulacoseira distans, which were found along the entire altitudinal gradient. Oligotrophic habitats, however, are widespread both in the highlands and lowlands of Papua New Guinea, as indicated by the condictivity data (Table I) and the scarce literature data (Chambers et al., 1987). It thus seems that at least a consider- able number of the observed northern montane diatoms and desmids are indeed cold-stenothermic.
The major changes in the distribution of the different biogeographical elements indi- cate that the altitudinal range between 1700-2500 m is a transition zone between the tropical aquatic environments of the lowlands and those of the more temperate and cool highland region. This seems also to be true for aquatic macrophytes, as Chambers et al. (1987) found that Papua New Guinean lakes at 1700 m altitude had a distinct lowland flora, which was absent at higher altitudes.
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62 W. Vyverman
However, zooplankton communities of some lakes between 1700 and 2570 m were dominated by species widespread in the Australasian region (Chambers et al., 1987) while taxa from more temperate climates were only found at higher altitudes.
A C K N O W L E D G E M E N T S
The expeditions to Papua New Guinea were financed by the Fund for Collective Scientific Research (N 32.9006.86). The author has a grant as Senior Research Assistant of the National Fund for Scientific Research (Belgium). I am greatly indebted to Prof. Dr P. Van der Veken (R.U.G.) , Dr P. Compfre (National Botanical Garden, Belgium) and Dr E. Coppejans (R.U.G.) for critically reading the manuscript. Many thanks to D. Gees for typing the text and to two anonymous referees for their useful comments on the manuscript.
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(Accepted 14 October 1991)
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