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European Journal of Phycology

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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

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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.
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Desmids and diatoms are among the beststudied groups of freshwater algae in tropical regions. Several extensive studies have been made in the Indo-Malaysian and N o r t h 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 examined 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. MATERIALS AND METHODS Papua New Guinea is situated between the equator and 12S, 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, conductivity, 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-MalaysianNorth 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

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 (C) < 15 15-20 20-25 25-30 30-35 > 35 No data < 15 15-20 20-25 25-30 30-35 > 35 No data <15 15-20 20-25 25-30 30-35 > 35 No data < 15 15-20 20-25 25-30 30-35 > 35 No data < 15 15-20 20-25 25-30 30-35 > 35 No data < 15 15-20 20-25 25-30 30-35 > 35 No data < 15 15-20 20-25 25-30 30-35 > 35 No data 0 0 1 64 63 5 10 0 0 2 28 0 0 0 0 1 6 25 4 0 0 0 2 14 10 0 0 0 3 9 11 2 0 0 0 1 5 0 0 0 0 0 I0 5 3 0 0 0 3

Nature 0-500 m P/B P1 Total

Type 84 1 64 49 2 41 -3 1 133 4 35 5 2

K20 (~tS.cm- l) 0-50 50-100 100-200 200--300 300-1000 > 1000 24 39 25 32 22 1

pH < 6.0 6.0~.5 6.5-7.0 7.0-7.5 7.5-8.0 > 8'0 No data < 6-0 6.0-6.5 6-5-7.0 7-0-7.5 7-5-8.0 > 8-0 No data <6.0 6.0-6.5 6.5-7.0 7.0-7.5 7.5-8.0 > 8.0 No data < 6-0 6.0-6.5 6.5-7.0 7.0-7.5 7.5-8-0 > 8.0 No data < 6.0 6.0~.5 6.5-7.0 7.0-7.5 7-5-8.0 > 8-0 No data < 6.0 6.0-6.5 6.5-7.0 7.0-7.5 7.5-8.0 > 8-0 No data < 6.0 6.0-6.5 6.5-7-0 7.0-7.5 7.5-8.0 > 8.0 No data 12 12 34 28 13 16 28 0 0 2 7 8 7 6 2 4 16 8 4 2 0 3 2 16 0 5 0 0 8 0 8 3 2 2 2 0 0 I 4 1 0 0 0 6 9 1 0 0 5

500-1000m

P/B PI Total

20 10 -30

l 26 2 0 3 1 4 3 5 0

0-50 50-100 100-200 200-300 300-1000 > 1000

0 0 13 8 8 1

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I000-1500m

P/B P1 Total

26 1 29 10 2 2 -3 0 36 4 4 5 t

0-50 50-100 100-200 200-300 300-1000 > 1000

7 13 12 2 2 1

1500-2000 m

P/B PI Total

15 1 22 11 2 3 -3 0 26 4 0 5 1

0-50 50-100 100-200 200-300 300-1000 > 1000

10 7 4 4 1 0

2000-2500 m

P/B P1 Total

17 1 23 8 2 0 -3 1 25 4 1 5 0

0-50 50-100 100-200 200-300 300-1000 > I000

12 8 2 2 1 0

2500-3000 m

P/B P1 Total

5 1 1 2 -3 6 4 5

4 1 1 0 0

0-50 50-100 100-200 200-300 300-1000 > 1000

0 5 0 0 0 1

> 3000 m

P/B P1 Total

16 1 22 5 2 3 -3 0 21 4 0 5 1

0-50 50-I00 100-200 200-300 300-1000 > 1000

19 1 0 1 0 0

Altitudinaldistribution of algae
Sout~ hern-hem isphere0-1% Pantropical8.4% Palaeotropical5.4% Indo-Malaysian- 4.5% North Australian Northern-montane 5,9%
(a)

51

Cosmopolitan 77-7%

~Southern-hemisphere
llIHIllllllllllIS Vantrop

ca 6 ,o

0.3%

Cosmopolitan / 71%

~
~
(b)

Palaeotropical 1% 4-.
-

\ILLLLLILLILLllLIIII~ Indo- Malaysian

Northern-montane2-4%

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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-NorthAustralian, palaeotropical,pantropical, southern hemispheric.Absolutenumbers 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 elsewhere (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 littleknown 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 cosmopolitan. Only 76 taxa (19-8%) are noncosmopolitan 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

TABLE II. The percentages

and

numbers

of taxa in each distribution type of diatom one non-cosmopolitan taxon

genera

found in Papua

New

Guinea

with at least

IndoNorthern Cosmopolitan N % % -. -. -. . . . . . . . 3 12.5 . . . . . . . -. . -. . 18-8 10.0 . 6 1 . . . 1 . . . . . . . . . 14.3 . 1 . . . 5 . 10.0 . 17.2 . . . . 1 1 3 6.6 9.1 . . . . I -1 . . . I0.0 20-0 15"8 . . . ------. . . . . . 1.1 -3.1 . . . . . 3 1 3.3 9.1 -. . . . --2 16.7 . . . . 2 11.I % % % 11 7 . 3 I . -. . . . -. . 2 --. 28.6 . 3 1 -3 2 1 . . 4 . . -2 1 . 1 . -. . . . . . 100.0 . -. 9.1 . . . . -2-2 . . 12.5 100.0 100-0 25.0 11.1 -. 7 8 18 --75.0 --58.3 88.9 61.1 87-5 2 11.1 . -. 3 1 16.7 12"5 . --montane N hemisphere N Pantropical N Australian N Southern MalaysianNorth Palaeotropical N --%

Genus

Total N

18 8

12 9

24 2 1

--

-,,q

3 8 51 13 9 5 79 8 31 -24 6 8 4 16 84'2 80.0 -82.8 85"7 80.0 72.7 96-9 81.2 90.0 71.4 86.8

2 7 37

66.7 87.5 72.6

. . 7-8

I 1 10

33.3 12.5 19-6

16 10 7 91

11 32

1 29 7 10

Achnanthes Amphora Aulacoseira Caloneis Cymbella Diatoma Diatomella Diploneis Epithemia Eunotia Gomphonema Fragilaria Frustulia Navicula Neidium Nitzschia Peronia Pinnularia Rhopalodia Stauroneis Stenopterobia Surirella

19

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TABLE I l i . T h e p e r c e n t a g e s a n d n u m b e r s o f t a x a o f s e l e c t e d d e s m i d g e n e r a p e r d i s t r i b u t i o n t y p e . O n l y g e n e r a w i t h a t l e a s t o n e n o n cosmopolitan taxon in Papua New Guinea are included

Genus 9 37 87 2 21 . I0 . . 1 . . . 0.8 . . . . . --. . . 3.1 -1 . 1 . . . . . 2 1 4 16'6 2'4 3.4 --I 2 7 8.3 4"9 6"0 . .

Total N %

Cosmopolitan N %

Northern montane N

Southern hemisphere N %

IndoMalaysianNorth Pantropical Australian N % N % .

Palaeotropical N

75.1 90"2 74"3 66.7 65.6

---. --

-3 -1 . . . 1 . -3 -. . 1

-2"6 -3.1 5.0 -2.5 -6.3

Actinotaenium Closterium Cosmarium Desmidium Euastrum Ichthyocercus Micrasterias Onychonema Pleurotaenium Staurastrum Staurodesmus Teilingia Xanthidium
1 10 100 31 2 8 50.0 50-0 71.5 83.4 91.2 66.7 50.0 . . -. . . . 5.0 . . -. . . . 3 1 5 1 1 3 1 1 1 9.4 100.0 25.0 50.0 7-1 2.5 2.9 33.3 6.3 1 16 1 6 . 3 . 3 13 2 . 6

12 41 117 3 32 1 20 2 14 120 34 3 16

. 2"4 13"7 33.3 18.8 . 15-0 . 21.4 10.8 5-9 . 37.5

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TABLE IV. The distributional range of non-cosmopolitan diatoms in relation to altitude. The altitude corresponding with 50% of the occurrences (median altitude) is indicated by a ' ' . For that purpose the altitudinal range was divided into intervals of 100 m. The number of samples in which it was found is given between brackets, followed by its occurrence in relation to conductivity (K20) and temperature

K20 0zS.cm- t)

Temperature 0 (oc)

km

Northern Montane: Achnanthes altaica Achnanthes marginulata Aulacoseira distans Aulacoseira distans var. alpigena Aulacoseira lirata var. seriata + lacustris Caloneis tenuis Cymbella aequalis Cymbella lunata Cymbella perpusilla Diatoma hiernale Diatoma mesodon Diatomella balfourniana Eunotia bidentula var. elongata Eunotia bigibba var. pumila Eunotia diodon Eunotia rostellata Navicula gandrupii Navicula stroemii Neidiurn bisulcatum Peronia fibula
(6) (4) (46) (14) (5) (4) (3) (72) (23) (7) (5) (1) (10) (4) (18) (4) (2) (5) (4) (5) (5) (14) (3) - - -- (20) (1) (1) (1) (73) (24) (22) (8) (1) (17) (3) 28-528 131 30 24 15-700 7-260 56--400 62-1"9 352 24-313 23-256 13-32 31 27 31 16-36 19-29 18-31 13-30 28 27-36 27-33 33-62 10-131 39-283 13-16 13-36 12-16 11-38 33-39 7-272 7-69 7-215 15-62 15-54 7-283 7-900 13-66 13--66 126 7-65 12-69 7-85 12-33 33-243 37-283 12-69 11-33 12-20 12-13 12-33 12-26 12-26 13-18 15-20 12-40 13-32 12-18 12-18 26 13-21 13-16 12-24 14-15 13-24 12-29 13-15 15-20

Southern Hemispheric: Frustulia magaliesmontana Frustulia rhomboides var. elongatissima Rhopalodia novae-zealandiae

Pantropicah Achnanthes inflata Achnanthes oblongella Aulacoseira agassizii var. malayensis Aulacoseira ikapoensis var. procera Cymbella muelleri Cymbella spicula Cymbella turnida Diploneis subovalis Epithemia cistula Eunotia camelus Eunotia camelus var. karveernsis

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Pantropicai:
(4) (13) (4) (7) (59) 30-93 56-313 25-78 28-131 25-190

(1)
18o
240-430 (9)

(19)

14-131

m.

Eunotia camelus var. ventricosa Eunotia didyma var. claviculata Eunotia dissimilis Eunotia monodon vat. tropica Eunotia rabenhorstiana Eunotia rabenhorstii var. monodon Eunotia tschirchiana Eunotia zygodon Fragilaria strangulata Gomphonema affine Gomphonema augur var. turris Gomphonema tenerrimum Navicula lagerheimii Navicula lagerheimii var. intermedia Navicula lepidula Navicula muticoides Navicula perrotettii Navicula seminuloides Neidium gracile Pinnularia acrosphaeria Pinnularia acrosphaeria var. turgidula Pinnularia brevicostata var. sumatrana Pinnularia luculenta Pinnularia rivularis Stauroneis minutula
(2) (90) (4) (11) (2) (3) (1) (36) (19) (26) (9) (46) (3) (14) (1) (1) (1) (8) (1) (12) (4) 26-528 446 25-528 24-76 27-36 29 28-30 28-31 - - - 131 90q531 25-75 40-435 62-264 78-528 94 7-528 28-900 26-590 24-269 7-1.8 24-131 7-264 26 14 528

27 27-32 13-30 27-36 23-32 23 28-31 27-40 36 23-31 28-30 18-34 29-33 29-36 30 13-36 27-32 26-36 28-32 21-36 28-36 21-36 33 40 30

~d

-1 0

Palaeotropicah

Navicula invicta Navicula ruttneri Navicula seminuloides vat. sumatrana Neidium gracile forrna aequalis

Indo-Malaysian-North Australian:

Achnanthes crenulata Achnanthes crenulata var. linearis Achnanthes montana var. tropica Amphora subturgida Cocconeis brevicostata Cocconeis cataractacum Navicula curta Nitzschia woltereckii Stauroneis tenera Stenopterobia pelagica Surirella celebesiana Surirella sublinearis Surirella tenuissima
(27) (2) (29) (2) (5) (1) (3) (10) (4) (10) (1) (1) (17) 110-230 94-199 94 210-352 54-98 15 73 184-590 35-830 24-131 181 135 24-85 20-26 19-28 19 28-30 25-31 18 26 25-35 26-30 27-36 30 23 14-29

(JI t~

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TABLE V. The The distributional range of non-cosmopolitan desmids in relation to altitude. The altitude corresponding with 50% of the occurrences (median altitude) is indicated by a ' ' . For that purpose the altitudinal range was divided into intervals o f 100 m. The number of samples in which it was found is given between brackets, followed by its occurrence in relation to conductivity (K20) and temperature (T) km

K20 (~tS.cm-t) . . .

Temperature 0 (C) .

1 &.

Northern Montane:
(6) (1) (1) (1) (2) (4) (8) (5) (4) (3) (3) (4) (1) (1) (1) (14) (4) (2) (2) (5) (6) (7) (2) (1) (1) (3) (4) (8) (1) (5) (1) 11-313 24-131 24 131 26 24-131 552-700 63-131 14-131 95-553 24-180 26-85 32-84 ll 131 16-131 32-173 16-161 63 28-92 28 21-27 25-36 26 36 33 25-32 27-32 25-36 36-40 28-32 18-30 19-31 29-30 21 36 23-36 25-30 23-30 28 16-30 33 11-12 15-20 7-16 7 16 12 lO 16-39 11-39 38 10-85 19-26 21 26 15 22 12-26 12-20 13 13-22

Actinotaenium adelochondrum Aetinotaenium adeloehondrum var. kriegeri Closterium costatum var. borgei Cosmarium levinotabile Cosmarium monomazum Cosmarium nasutum Cosmarium quadratum Euastrum oblongum Micrasterias crux-melitensis

Southern Hemispheric:
,<

Staurastrum sagittarium

Pantropieal:

Actinotaenium capax var. minus Closterium nematodes Closterium nematodes var, proboscideum Cosmarium depressum var. elevatum Cosmarium nitidulum var. javanicum Cosmarium quadrum Cosmarium seelyanum Cosmarium subglobosum var. sumatranum Cosmarium trachypolum Cosmarium variolatum vat. rotundatum Euastrum praemorsum Euastrum spinulosum Euastrum spinulosum var. inermius lchthyocercus longispinus Micrasterias apiculata var. lacerata Micrasterias foliacea Micrasterias foliacea var. ornata Micrasterias radians Micrasterias tropica Onychonerna laeve var. latum Pleurotaenium minutum var. subattenuatum

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Staurastrum cerastes var. pulchrum Staurastrum orbiculare var. denticulatum Staurastrum sonthalianum Staurodesmus arcuatus Teilingia quadrispinata forma evoluta Xanthidium armature var. anguligerum
(6) (10) (10) (1) (3) 160 16-180 7-221 7-161 131 16 26 23-36 23-34 19-26 36 23 . 22-30 28 19-29 21 23-36 15-29 19 23-32 30--36 (11) (1) (7) (1) (8) (7) (1) (4) (3) 10-180 93 13-180 11 16-131 7-161 13 13-161 28-131

(I)

Palaeotropicah

Cosmarium askenasyi Cosmarium perfissum Cosmarium striolatum var. nordstedtii Euastrum moebii Micrasterias zeylanica Staurastrum longibrachiatum Staurastrum longibrachiatum var. javanicum Staurastrum wildemanii Xanthidium subtrilobum var. inornatum

Indo-Malaysian-North Australian:
g~

Closterium rectimarginatum Cosmarium blyttii forma australicum Cosmarium burkillii var. depressum Cosmarium ceylanicum Cosmarium dubium Cosmarium exasperatum Cosmarium freemanff var. verrucosum Cosmarium mikron Cosmarium nudum Cosmarium otus var. ornatum Cosmarium panduriforme Cosmarium perigranulatum Cosmarium pseudoarmatum Cosmarium sumatranum Cosmarium taxichondrum var. pulchrum Cosmarium tjibenongense forma minus Cosmarium trachypleurum var. nordstedtii Desmidium baileyii forma tetragonum Euastrum distortum Euastrum gnathophorum Euastrum horikawae Euastrum longicolle vat. capitatum Euastrum moebii var. burmense Euastrum moebii vat. tetrachastriforme
(1) (1) (6) (1) (1) (6) (1) (2) (3) (9) (2) (1) (4) (1) (3) (2) (1) (4) (6) (5) (2) (2) (3) (2) 36 36 23-36 29 36 23-32 36 30-33 29 23-29 27 28 26--28 36 23 29 19 27-36 21-26 21-23 29 29-30 29-31 25-27 131 131 25-180 84 131 12-47 131 26-28 25-84 16-18o 30-45 93 56-180 131 18-180 25 13 26-131 7-16 11-16 63-84 28-84 27-63 24-63

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o~

TABLE V. cont.
1 2 3

km

N -- - (4) (2) (4) (1) (6) (4) (1) (1) (4) (3) (1) (15) (1) (2) (1) (19) (2) (14) (9) (4) (2) (i) (4) (5) (11) (7) (7) 16-160 32-131 14-131 28 11-32 16-63 28 24 16-161 16-46 552 10-161 28 94-118 63 26-211 30-92 12-160 13-173 78-700 24-32 25 10-160 24-118 12-84 25-700 24-131 23-30 30-36 30-40 30 21-30 23-25 28 28 25-26 23-29 27 21-31 30 30-32 25 27-36 27-28 19-29 19-33 29-32 30 29 22-26 29-30 19-30 25-29 29-36

K20 Temperature 0 (laS.cm- ~) (C)

Micrasterias lux Micrasterias mahabuleshwarensis vat. reducta Micrasterias subincisa Pleurotaenium coroniferum var. multinodulosum Pleurotaenium kayei Pleurotaenium ovatum var. tumidum Staurastrum contectum Staurastrum ehrenbergianum var. rostratum Staurastrum ensiferum Staurastrum freemanii Staurastrum graeile forma kriegeri Staurastrum gutwinskii Staurastrum gutwinskii var. evolutum Staurastrum javanicum Staurastrum protectum var. rangoonense Staurastrum playfairi Staurastrum pseudozonatum Staurastrum wildemanii var. majus Stauraslrum zonatum vat. majus Staurodesmus gibberulus Staurodesmus gibberules var. mucronatus Xanthidium acanthophorum var. raciborski Xanthidium apiculatum Xanthidium hastiferum var. javanicum Xanthidium multicorne Xanthidium sexmamillatum var. pulneyense Xanthidium subtrilobum

Altitudinal distribution of algae


5O 4O

59

50
2O I0 2 "S E
Z

0 0,1 7O 6O 50 40 5O 0.5
1 1,5 2 2,5 3 5-5

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2O I0
o

0.1

0.5

1,5

2.5

5.5

Altitude (1000 m. a.s.l.) Northern-montane PalaeotropJcal ~ Pantropical

F7777 Indo-Mala ysian- Australian ~

F16. 2. Numbers of northern montane, pantropical, palaeotropical and Indo-Malaysian-NorthAustralian diatom (a) and desmid (b) taxa along the altitudinal gradient.

cosmopolitan taxa. A m o n g desmids, noncosmopolitan taxa are mainly found in the genera Cosmarium, Euastrum, Micrasterias, Pleurotaenium and Xanthidium. N o 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 hemispheric 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,

60

W. Vyverman

Surirella tenuissima), the pantropical palaeotropical 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 affinities 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 altitude, 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 IndoMalaysian-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 IndoMalaysian-North Australian desmids were found below 100 m [Fig. 2(b)]. F r o m 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 cosmopolitan distribution. The degree of cosmopolitanism is comparable to other tropical regions. In the Lake Chad region, for

example, the cosmopolitan and subcosmopolitan 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 preferring cooler water whereas northern montane diatoms are much more numerous. In addition, 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 noncosmopolitan 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 demonstrated 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 differences 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 pantropical taxa, as most migratory routes follow a N-S direction. For the palaeotropical element, a somewhat 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 suitable 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), preliminary 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 biogeographical elements in Papua New Guinea shows a distinct relationship with altitude and water temperature (Tables IV and V). Indo-Malaysian-North Australian, pantropical and palaeotropical elements amongst diatoms and desmids are in Papua New Guinea mostly confined to the lowlands and mountains below about 17002500 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 floodplain 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 highlands, which points to their cold-stenothermic 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 considerable number of the observed northern montane diatoms and desmids are indeed cold-stenothermic. The major changes in the distribution of the different biogeographical elements indicate 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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Zealand Desmids. IL University Press, Botany Division, Christchurch. CROASDALE, H. SCOTT, A. M. (1976). New or otherwise interesting desmids from northern Australia. Nova Hedwigia, 27: 501-596. DUMONT, H. J. (1983). Biogeography of rotifers. Hydrobiologia, 104: 19-30. FOGED, N. (1971). Freshwater diatoms in Thailand. Nova Hedwigia, 22: 267-369. FOGED, N. (1976). Freshwater diatoms in Sri Lanka ACKNOWLEDGEMENTS (Ceylon). Bibl. Phycol. 23: 1-113. The expeditions to Papua New Guinea were FOGED, N. (1978). Diatoms in Eastern Australia. Bibt. Phycol. 41: 1-243. financed by the Fund for Collective Scientific Research (N 32.9006.86). The author has a grant FfRSTER, K. (1982). Das Phytoplankton des Siisswassers. 8 : l. Conjugatophyceae: Zygnematales und as Senior Research Assistant of the National Desmidiales (excl. Zygnemataceae). BinnenFund for Scientific Research (Belgium). I am gew/isser 16, Stuttgart. greatly indebted to Prof. Dr P. Van der Veken GASSE, F. (1986). East African diatoms. Taxonomy, (R.U.G.), D r P. C o m p f r e (National Botanical ecological distribution. Bibl. diatomol., 11: 1-202. Garden, Belgium) and Dr E. Coppejans (R.U.G.) GASSE, F., TALL1NG,J. F. & K1LHAM,P. (1983). Diatom for critically reading the manuscript. M a n y assemblages in East Africa: classification, thanks to D. Gees for typing the text and to two distribution and ecology. Rev. Hydrobiol. Trop., 19: 3-34. anonymous referees for their useful comments on HUSTEDT, F. (1937-39). Systematische und 6kologische the manuscript. Untersuchungen fiber die Diatomeenflora von Java, Bali und Sumatra nach dem Material der Deutschen Limnologischen Sunda-Expedition. REFERENCES Tell I. Systematischer Tell, Tell II. Allgemeiner ATKINSON,K. M. (1972). Birds as transporters of algae. Tell, Arch. Hydrobiol. SuppL, 15: 131-506, Br. phycol. J., 7: 319-321. 638-790. ATKINSON, K. M. (1981). Experiments in dispersal of HUSTEDT, F. (1942). Siisswasser-Diatomeen des indophytoplankton by ducks. Br. phycol. J., 15: malayischen Archipels und der Hawaii-inseln. Int. 49-58. Revue ges. Hydrobiol. Hydrogr, 42: 1-152. BEEHLER, B. M., THANE, K., PRATT, C. & ZIMMERMAN, JOHN, D. M. (1986). The Inland Waters of Tropical West G. (1983). Birds of New Guinea. University Press, Africa. Advances in Limnology 23. Arch. Princeton. Hydrobiol., Stuttgart. BEHRE, K. (1956). Die Siisswasseralgen der Wallacea- KRAMMER, K. & LANGE-BERTALOT,H. (1986). SiissExpedition. Arch. Hydrobiol., Suppl. 23: 1-104. wasserflora yon Mitteleuropa. Vol. 2, part 1. BERNARD, CH. (1908). Protococcacbes et Desmidides Naviculaceae. Gustav Fischer, Stuttgart. d'Eau Douce Rdcoltbes ~ Java. Batavia. KRAMMER, K. & LANGE-BERTALOT,H. (1988). SiissBERNARD, CH. (1909). Sur Quelques Algues wasserflora yon Mineleuropa. VoL 2, part 2. Unicellulaires d'Eau Douce Rbcoltbes dans le Nitzschiaceae, Epithemiaceae, Surirellaceae. Domain Malais. Dfpartement de l'Agric, aux Gustav Fischer, Stuttgart. Indes-Nferlandaises, Buitenzorg. KRIEGER, W. (1932). Die Desmidiaceen der Deutschen CHAMBERS,M. R., KYLE, J. H., LEACH,G. J., OSBORNE, Limnologischen Sunda-Expedition. Arch. HybroP. L. & LEACH, D. (1987). A limnological study biol., Suppl., 11: 129-230. of seven highland lakes in Papua New Guinea. KRtEGER, W. & BOUP~ELLY,P. (1956). Desmidiacres des Science in New Guinea, 13: 51-81. Andes du Venezuela. Ergebn. Deutsch. Limnol. COESEL, P. F. M., DUQUE, S. R. & ARANGO,G. (1988). Venezuela-Expedition 1952, 1: 141-195. Distributional patterns in some neotropical LING, H. U. & TYLER, P. A. (1986). A Limnological desmid species (Algae, Chlorophyta) in relation Survey of the Alligator Rivers Region. Part II." to migratory bird routes. Rev. Hydrobiol. trop., Freshwater Algae, Exclusive of Diatoms. 21: 197-205. Australian Government Publications Service., COMPARE, P. & ILTIS, A. (1983). The phytoplankton Canberra. (Qualitative composition of the algal flora). In McALPINE, J. R., KEIG, G. & FALLS,R. (1983). Climate Lake Chad. Ecology and Productivity of a of Papua New Guinea. CSIRO and ANU Press, Shallow Tropical Ecosystem (Carmouze, J.-P., Canberra. Durand, J.-R. & Lfvfque, C., editors), 145-152. PARISH, D. (1989). Population estimates of water birds Junk, The Hague. using the East Asian/Australasian flyway. CROASDALE,H., DE BICUDO, C. E. & PRESCOTT,G. W. Interwader Annual Report: 8-13. (1983). A Synopsis of North American Desmids. PODZORSKI,A. C. & HAKANSSON,H. (1987). Freshwater Part IL Desmidiaceae. Placodermae. Section 5. and marine diatoms from Palawan (a Philippine University of Nebraska Press. island). Bibl. Diatomol., 13: 1-244. CROASDALE,H. & FLINT, E. A. (1986). Flora of New PRESCOTT,G. W., CROASDALE,H. T. & VINYARD,W. C. Zealand Desmids. L Ward, Wellington. (1975). A Synopsis of North American Desmids.

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.

CROASDALE,H. & FLINT, E. A. (1988). Flora of New

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Part I1. Desmidiaceae: Placodermae, Section 1. University of Nebraska Press. PRESCOTT,G. W., CROASDALE, T. & VINYARD, C. H. W. (1977). A Synopsis of North American Desmids. Part IL Desmidiaceae: Placodermae, Section 2. University of Nebraska Press. PRESCOTT,G. W., CROASDALE, T., VINYARD,W. C. t~ H. BICUDO, D. (1981). A Synopsis of North American Desmids. Part IL Desmidiaceae: Placodermae, Section 3. University of Nebraska Press. PROCTOR, V. W. (1966). Dispersal of desmids by waterbirds. Phycologia, 5: 227-232. PROWSE,G. A. (1962). Diatoms of Malayan freshwaters. The Gardens Bulletin, Singapore, 19: 1-80. ROUND, F. E. (1981). The Ecology of Algae. Cambridge University Press. RUZ1CKA, J. (1977). Die Desmidiaceen Mitteleuropas. 1(1). Schweizerbart, Stuttgart. RUZICKA, J. (1981). Die Desmidiaceen Mitteleuropas. 1(2). Schweizerbart, Stuttgart. SCOTT, A. M. & PRESCOTT, G. W. (1958). Some freshwater algae from Arnhem Land in the Northern Territory of Australia. Rec. Amer.-Austr. Sci. Exp. Arhnhem Land, 3: 9-136. SCOTT, A. M. & PRESCOTT,G. W. (1961). Indonesian Desmids. Hydrobiologia, 17: 1-132. THOMAS, D. P. (1983). A limnological survey of the Alligator Rivers Region. I. Diatoms (Bacillariophyceae) of the region. Supervising Scientist Alligator Rivers Region, Res. Rep., 3: 1-137.

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THOMASSON, K. (1986). Algal vegetation in North Australian Billabongs. Nova Hedwigia, 42: 301-378. UHERKOVlCH, G. (1984). Phytoplankton. In The Amazon. Limnology and Landscape Ecology of a Mighty Tropical River and its Basin (Sioli, H., editor), 295-310. Junk, The Hague. VVWRMAN, W. (1988). Three new diatom taxa from the Central Highlands of Papua New Guinea. Diatom Research, 3: 259-264. VYWRMAN, W. (1989). Diatoms (Bacillariophyta) from Mount Giluwe (Southern Highlands Province, Papua New Guinea). Bull. Soc. Roy. Bot. Belg., 122: 61-80. VWERMAN, W. (1991a). Diatoms from Papua New Guinea. Bibl. Diatomol., 22: 1-221. VVV~RMAN, W. (1991b). Desmids from Papua New Guinea. Bibl. Phycol., 87: 1-200. VVVERMAN, W. (1991c). Distribution and ecology of periphytic diatom assemblages from 136 Papua New Guinean freshwaters. Hydrobiologia, in press. VYVERMAN, W. & COMPARE, P. (1991). Pinnularia sakulensis spec. nov. and Rhopalodia tholulata spec. nov., two new diatoms (Bacillariophyta) from Papua New Guinea. Belg. Journ. Bot., 124: 27-32.

(Accepted 14 October 1991)

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