16
PLEASE SCROLL DOWN FOR ARTICLE This article was downloaded by: On: 22 May 2010 Access details: Access Details: Free Access Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37- 41 Mortimer Street, London W1T 3JH, UK European Journal of Phycology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713725516 Altitudinal distribution of non-cosmopolitan desmids and diatoms in Papua New Guinea Wim 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 Papua New Guinea', European Journal of Phycology, 27: 1, 49 — 63 To link to this Article: DOI: 10.1080/00071619200650071 URL: http://dx.doi.org/10.1080/00071619200650071 Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

Altitudinal Algae

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Page 1: Altitudinal Algae

PLEASE SCROLL DOWN FOR ARTICLE

This article was downloaded by:On: 22 May 2010Access details: Access Details: Free AccessPublisher Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

European Journal of PhycologyPublication details, including instructions for authors and subscription information:http://www.informaworld.com/smpp/title~content=t713725516

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

Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf

This article may be used for research, teaching and private study purposes. Any substantial orsystematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply ordistribution in any form to anyone is expressly forbidden.

The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae and drug dosesshould be independently verified with primary sources. The publisher shall not be liable for any loss,actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directlyor indirectly in connection with or arising out of the use of this material.

Page 2: Altitudinal Algae

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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Page 3: Altitudinal Algae

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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Page 4: Altitudinal Algae

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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Page 5: Altitudinal Algae

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Page 7: Altitudinal Algae

TAB

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Page 8: Altitudinal Algae

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opic

a E

unot

ia r

aben

hors

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unot

ia r

aben

hors

tii v

ar.

mon

odon

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unot

ia t

schi

rchi

ana

Eun

otia

zyg

odon

F

ragi

lari

a st

rang

ulat

a G

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onem

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fine

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omph

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gur

var.

tur

ris

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phon

ema

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um

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icul

a la

gerh

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ii N

avic

ula

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var.

int

erm

edia

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icoi

des

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icul

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i N

avic

ula

sem

inul

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s N

eidi

um g

raci

le

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nula

ria

acro

spha

eria

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innu

lari

a ac

rosp

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ia v

ar.

turg

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a P

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ar.

sum

atra

na

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nula

ria

lucu

lent

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ris

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rone

is m

inut

ula

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ah

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icul

a in

vict

a N

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ula

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avic

ula

sem

inul

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s va

t. s

umat

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eidi

um g

raci

le f

orrn

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qual

is

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aysia

n-N

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nant

hes

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var

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a A

mph

ora

subt

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da

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cone

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avic

ula

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auro

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ten

era

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opte

robi

a pe

lagi

ca

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rell

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a Su

rire

lla

subl

inea

ris

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rell

a te

nuis

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13

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40

30

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28

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446

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19

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Page 9: Altitudinal Algae

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

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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

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6 (8

) 11

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0 (5

) 38

13

(4

) 10

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2

(3)

11-1

2 15

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11-3

13

21-2

7 (4

) 24

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25

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(1)

24

26

(1)

131

36

(1)

26

33

(14)

24

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25

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(4)

552-

700

27-3

2 (2

) 63

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25

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14-1

31

36-4

0 (5

) 95

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28

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24-1

80

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0 (7

) 26

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1 (2

) 32

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29-3

0 (1

) ll

21

(1

) 13

1 36

(3

) 16

-131

23

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32-1

73

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) 16

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63

28

(5)

28-9

2 16

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,<

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Page 10: Altitudinal Algae

Stau

rast

rum

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aste

s va

r. p

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rum

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aura

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m o

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ular

e va

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enti

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tum

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aura

stru

m s

onth

alia

num

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auro

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us a

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a qu

adri

spin

ata

form

a ev

olut

a X

anth

idiu

m a

rmat

ure

var.

ang

ulig

erum

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eotr

opic

ah

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mar

ium

ask

enas

yi

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mar

ium

per

fiss

um

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mar

ium

str

iola

tum

var

. no

rdst

edti

i E

uast

rum

moe

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rast

eria

s ze

ylan

ica

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rast

rum

lon

gibr

achi

atum

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aura

stru

m l

ongi

brac

hiat

um

var.

jav

anic

um

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rast

rum

wil

dem

anii

X

anth

idiu

m s

ubtr

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um v

ar.

inor

natu

m

Indo

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aysia

n-N

orth

Aus

tral

ian:

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lost

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m r

ecti

mar

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tum

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osm

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m b

lytt

ii f

orm

a au

stra

licu

m

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mar

ium

bur

kill

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ar.

depr

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m

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mar

ium

cey

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cum

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m d

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m

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mar

ium

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m fr

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anff

var

. ve

rruc

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m m

ikro

n C

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udum

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m o

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var.

orn

atum

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osm

ariu

m p

andu

rifo

rme

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mar

ium

per

igra

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tum

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osm

ariu

m p

seud

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osm

ariu

m s

umat

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m

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mar

ium

tax

icho

ndru

m v

ar.

pulc

hrum

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osm

ariu

m t

jiben

onge

nse

form

a m

inus

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ariu

m t

rach

yple

urum

var

. no

rdst

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i D

esm

idiu

m b

aile

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orm

a te

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onum

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uast

rum

dis

tort

um

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stru

m g

nath

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hor

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ae

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m l

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pita

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men

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stru

m m

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i va

t. t

etra

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trif

orm

e

(I)

160

(6)

16-1

80

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221

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7-

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26

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6 23

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6 36

23

(11)

10

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) 93

28

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80

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9 •

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11

21

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23

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) 7-

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19

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36

36

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28

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28

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29

19

27-3

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3 29

29

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131

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131

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93

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80

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131

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80

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13

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31

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Page 11: Altitudinal Algae

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

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) 24

28

(4)

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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

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• (7

) 25

-700

25

-29

• (7

) 24

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29

-36

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Page 12: Altitudinal Algae

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,

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Page 13: Altitudinal Algae

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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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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