The centric diatom Puncticulata balatonis (Pantocsek) Wojtal et Budzy½ska, comb. nov., in the...

509

Nova Hedwigia Vol 93 issue 3ndash4 509ndash524Stuttgart November 2011

Article

copy 2011 J Cramer in Gebr Borntraeger Verlagsbuchhandlung Stuttgart Germany wwwborntraeger-cramerdeDOI 1011270029-503520110093-0509 0029-503520110093-0509 $ 400

The centric diatom Puncticulata balatonis (Pantocsek)Wojtal et Budzyfrac12ska comb nov in the plankton ofeutrophic-hypertrophic Rusaraquoka Lake (Western Poland)

Agnieszka Budzynska1 and Agata Z Wojtal2

1 Department of Water Protection Adam Mickiewicz University Umultowska 89 61-614Poznantilde Poland e-mail abudzamuedupl

2 Department of Phycology W Szafer Institute of Botany Polish Academy of SciencesLubicz 46 31-512 Krakoacutew Poland

With 24 figures and 3 tables

Budzyńska A amp AZ Wojtal (2011) The centric diatom Puncticulata balatonis (Pantocsek) Wojtal etBudzyńska comb nov in the plankton of eutrophic-hypertrophic Rusałka Lake (Western Poland) ndashNova Hedwigia 93 509ndash524

Abstract Cyclotella balatonis Pantocsek is probably a common and widely distributed component ofthe phytoplankton in mesotrophic to eutrophic European lakes and rivers However the distributionand ecology of the species are still unclear since the species remained hidden in the broad C radiosa(Grunow) Lemmermann - concept throughout most of the 20th century Cyclotella balatonis was foundto form quite a large population in the plankton of a small and shallow urban eutrophic to hypereutrophiclake (Rusałka Lake Western Poland) The species occurred in the lake throughout the study periodfrom February till December 2007 with its highest number in the winter - spring period The morphologyand fine valve structure of the species are described in the paper documented with light microscope(LM) and scanning electron microscope (SEM) micrographs and discussed along with the speciesecology Additionally Cyclotella balatonis is formally transferred into the genus Puncticulata[P balatonis (Pant) Wojtal et Budzyńska comb nov] and compared with the morphologicallysimilar taxa ndash P radiosa (Lemmermann) Haringkansson P comta (Ehrenberg) Haringkansson and P prae-termissa (Lund) Haringkansson

Key words Bacillariophyta Puncticulata ecology morphology Poland

Introduction

Centric diatoms are an important component of freshwater ecosystems throughoutthe world They often reach high numbers in eutrophic lakes and rivers (Genkal ampKiss 1993) As their occurrence is usually related to the trophic status of inhabitedwater bodies they are particularly useful bio-indicators in monitoring works andpaleolimnological studies (Wunsam et al 1995 Siver 1999 Wojtal amp Kwandrans2006 Rimet et al 2009 Houk et al 2010)

510

Despite some of the members of Cyclotella genus are known for up to 170 years theecology of many species still remains uncertain Moreover there are also manyuncertainties within the genus concerning taxonomy and nomenclature (Hausmannamp Lotter 2001 Haringkansson 2002 Houk et al 2010) Recently some species of thenot-uniform genus were moved to several newly-established genera (short review inHouk et al 2010) In 2002 the genus Puncticulata was established comprisingspecies characterised by concentrically undulated central part of valve face andnumerous scattered or radially arranged areolae and fultoportulae (or only areolaeor only fultoportulae) with P comta as the type (Haringkansson 2002) The genus hasbeen generally accepted (eg Tanaka 2007 Hayashi 2007 Kobayasi et al 2006Cremer amp Koolmees 2010) though some uncertainties are still discussed (Houk etal 2010)

One of the species sharing the above-mentioned characteristics of the genusPuncticulata (though not transferred to the genus till now) is Cyclotella balatonisand its variety C balatonis var binotata described by Pantocsek (1901 p 104 figs319 331 332) from lake Balaton near Sioacutefok in Hungary Originally the specieswas illustrated by three drawings of valves with scattered (Pantocsek 1901 PlateXV figs 319 332) and radially arranged (Pantocsek 1901 Plate XV fig 331)puncta in the middle part of valve face After Pantocsekrsquos description C balatonisremained practically forgotten and was reported only a few times (eg Handmann1913 Hindaacutek amp Hindaacutekovaacute 2008) It was generally considered a synonym ofCyclotella radiosa (eg Krammer amp Lange-Bertalot 1991 fig 62 11 12) followingthe broad species concept that was in use at that time Thus specimens of a broadmorphological range present in a variety of water bodies were identified as C radi-osa Cyclotella balatonis might have also been misidentified as other species of thecomplex such as P praetermissa (syn Cyclotella praetermissa) or Puncticulatacomta (syn Cyclotella comta) Recently as a result of precise study by Houk et al(2010) using SEM Cyclotella balatonis has been resurrected and the morphologicalranges of similar species were discussed However due to the proceeding long periodof taxonomical confusion the species is almost absent in the literature and needsmuch attention particularly in terms of distribution and autecology

In this paper a population of C balatonis in a small and shallow urban eutrophic tohypereutrophic lake (Rusałka Lake Western Poland) is presented with its quantitativecharacteristics along an annual cycle Morphological features of the diatom aredescribed in detail and documented with LM photographs of live specimens andcleaned frustules as well as with the SEM micrographs Cyclotella balatonis wasalso compared with morphologically similar species namely Puncticulata comtaP praetermissa and P radiosa Due to the great morphological similarity of Cyclotellabalatonis to these and other Puncticulata-species the transfer of Cyclotella balatonisto the genus Puncticulata has been proposed

Study site

Rusałka Lake is situated in the city of Poznań Western Poland (N52deg25E16deg52) It was formed in1943 by damming the Bogdanka River and subsequent flooding of its valley The lake covers the area

511

of 367 ha Although its maximum depth is 90 meters the average does not exceed 2 meters As aresult of the morphometry the lake is mainly polymictic with thermal stratification developing only ina small area around its deepest point in summer (Gołdyn et al 2010)

The lake is strongly eutrophic or hypereutrophic according to different criteria (Table 1) Cyanobacterialblooms occur in summer and autumn (Gołdyn et al 2010) Due to its location and surroundingattractive areas (mainly forests) the lake is intensely used for recreation

Materials and methods

Water samples for the phytoplankton analysis were taken at every meter depth at the deepest part of thelake in 2007 once a month from February till October and once in December Quantitative sampleswere preserved with Lugolrsquos solution and analysed with the use of inverted microscope PZO aftersedimentation in cylindrical chambers according to the Utermoumlhl method (Wetzel amp Likens 1991)

All diatoms from the family Stephanodiscaceae were counted as one taxon The relative frequency ofPuncticulata balatonis and other species belonging to this family was then estimated by counting 100frustules in hydrogene peroxide-treated samples using an Olympus BX-60 microscope Measurementsand the micrographs of live diatoms were done with the use of the same LM and AnalySIS programme

A qualitative analysis of the diatom material was performed after removing carbonates (addition of10 HCl to the samples followed by boiling taking place after 24 h for 15 minutes) and organic matter(boiling in 30 H

2O

2 with small amounts of KClO

3) After air-drying the material was mounted in

Naphraxreg Observations of the diatoms were performed with a Nikon Eclipse 80i light microscopeequipped with Differential Interference Contrast The micrographs were taken with a Nikon DS-Fi 1camera SEM analyses were performed on the material coated with gold using a Hitachi S-4700scanning electron microscope SEM micrographs were taken in the Laboratory of Electron and ConfocalMicroscopy at the Faculty of Biology Adam Mickiewicz University in Poznań and in the Laboratoryof Field Emission Scanning Electron Microscopy and Microanalysis at the Institute of GeologicalSciences of the Jagiellonian University in Krakoacutew Density of valve face structures (eg costae) in10 microm in a line tangential to the radius was calculated based on the formula in Genkal (1977)Additionally data on the morphology of Cyclotella balatonis were obtained from the type material(Balaton Lake Siofok samples 156 157) hosted in the Natural History Museum in Budapest (Hungary)

Table 1 Estimation of the trophic state of Rusałka Lake according to Carlson index (1977) and OECD(1982) criteria (acc to Gołdyn et al 2010)

Carlson indexTSI Values in Boundary values Trophic state

Rusałka lake for eutrophy

TSI (TP) 7437 gt70 HypertrophyTSI (ChL) 7171 53ndash70 HypertrophyTSI (SD) 6737 53ndash70 Eutrophy

OECDCharacteristic Values in Boundary values Trophic state


Mean TP (microgl) 96 gt100 EutrophyMean chlorophyll-a (microgl) 347 gt25 HypertrophyMax chlorophyll-a (microgl) 802 gt75 HypertrophyMean Secchi depth (m) 08 lt15 HypertrophyMin Secchi depth (m) 06 lt07 Hypertrophy

512

Water temperature pH and conductivity were measured in situ using YSI 610-DM-meter Watertransparency was expressed as the visibility of the Secchi disc For the statistical graphs Statistica 80software was used

Results

Morphology of Puncticulata balatonis in Rusałka Lake

Puncticulata balatonis (Pantocsek) Wojtal et Budzyńska comb nov Figs 1ndash12 amp 15ndash22

BASIONYM Cyclotella balatonis Pantocsek Die Kieselalgen oder Bacillarien des Balaton p 104 PlateXV figs 319 332 1901

TYPE LOCALITY Balaton Lake near Sioacutefok Hungary

Cells solitary or in short 2ndash7-cell chains (Fig 1) containing multiple discoidchloroplasts The valve diameter ranges between 87 and 210 microm Stria density is15ndash185 in 10 microm with 35ndash47 costae (thickened ribs) in 10 microm Striae are ofunequal length with inserted shorter (Figs 15ndash17) or divided striae (Fig 19) formingan irregularly circular central area (Figs 6 9 12 17) The central area is concentricallyslightly elevated (Figs 15 17) with numerous scattered openings of valve facefultoportulae (externally visible as minor puncta Fig 16) and areolae (externallyvisible as bigger puncta Figs 15ndash17) External openings of the rimopor-tulae arepresent at the end of the shortened striae (Figs 15 16) The internal opening of thesessile rimoportulae has a radially or slightly obliquely positioned slit (Figs 18 20)situated close to internal alveolar openings (towards the valve face margin) Marginalfultoportula openings with two satellite pores are situated (internally) on thethickened costae (shadow lines) separated by 3ndash5 thinner costae Valve facefultoportula openings are internally covered with domed cribra and surrounded bythree satellite pores (Figs 19 20) The morphology of the valve face internal alveolaropenings and the location of internal rimoportula openings correspond well withthe morphology of the specimens from the type material of Cyclotella balatonis(Figs 13 14 amp 21 22 Table 3)

Table 2 Physio-chemical parameters of water of Rusałka Lake in 2007

Temperature pH Coductivity Transparencyndash(degC) (microScm) Secchi depth (m)

20 II 31ndash41 72ndash78 836ndash841 1526 III 49ndash71 79ndash81 866ndash868 1820 IV 134ndash103 79ndash80 no data 1018 V 110ndash156 77ndash82 820ndash840 0819 VI 112ndash232 73ndash84 760ndash878 0918 VII 126ndash240 75ndash83 697ndash754 08520 VIII 145ndash220 75ndash91 627ndash739 0612 IX 144ndash148 83ndash85 646ndash652 0822 X 85ndash89 80ndash83 668ndash680 10503 XII 35ndash38 81ndash82 740ndash744 135

513

Ecological data

The water of Rusałka Lake was alkaline throughout the study period showing highconductivity Water transparency was relatively low ranging from 06 to 18 m ofSecchi depth (Table 2)

Puncticulata balatonis was present in the lake in all the seasons of 2007 It wasfound to form chains only in February March and April The diatom valve sizedistribution pattern varied in subsequent months with the largest average valve sizein February (Fig 23) Its abundance in the lake was also higher in the winter-springperiod (from February till May fig 24) with the maximum value of 35 times 103 cellsml-1 and a biomass of 298 mg l-1 (19 of the total phytoplankton cell number and525 of the total biomass at 6 m depth in March) In other spring samples

Figs 1ndash14 Puncticulata balatonis (Pantocsek) Wojtal et Budzyńska Light microscopy Figs 1ndash12 Specimens from the Rusałka Lake Figs 13ndash14 P balatonis type material sample no 152Fig 1 Short chain of alive cells in girdle view Figs 5ndash6 The same valve at different focus levels OnFigs 6 12ndash13 the characteristic gaps (notae) are marked with an arrow Scale bar represents 10 microm

514

Figs 15ndash22 Puncticulata balatonis (Pantocsek) Wojtal et Budzyńska Scanning electron microscopyFigs 15ndash20 specimens from the Rusałka Lake Figs 21ndash22 P balatonis type material samples no152 amp 156 Fig 15 external view of the rimoportula opening at the end of shortened striae ndash notae

515

P balatonis contributed between 63 and 187 to the total phytoplankton numbersand between 21 and 47 to the total biomass From June until October the numberof the diatom did not exceed 300 frustules per 1 ml (Fig 24) and no valve wasfound in September At the beginning of December P balatonis number increasedslightly again

At the time of the bloom of P balatonis chrysoflagellates (Erkenia subaequiciliataSkuja Chrysococcus triporus Matvienko) pennate diatoms (Fragilaria sp) andorcoenobia-forming Chlorococcales (Didymocystis inconspicua Koršikov Tetastrumsp div Scenedesmus sp div Coelastrum microporum Naumlgeli Crucigenia tetrapediaKirchner) were also abundant Other diatoms such as Aulacoseira ambigua (Grunow)Simonsen Cyclotella meneghiniana Kuumltzing C ocellata Pantocsek Cyclostephanosdelicatus (Genkal) Casper C dubius (Fricke) Round Stephanodiscus parvus Stoermeramp Haringkansson Asterionella formosa Hassal and Navicula sp were also observed

Discussion

Morphology of Puncticulata balatonis (Pant) Wojtal et Budzyńska

Puncticulata balatonis shares important morphological characters assigned to thegenus justifying its formal transfer to this genus a slightly concentrically undulatemiddle part of the valve face with scattered or radially arranged numerous areolaewith internally domed cribra and numerous fultoportulae Additionally valves ofP balatonis have internal alveolar openings with marginal fultoportulae located onthicker costae rimoportulae located externally at the top of shortened striae andinternally in the marginal part of the valve face

The morphology of P balatonis valves from Rusałka Lake corresponds well withthe morphology of the specimens from the Pantocsekrsquos type material (Figs 13 1421 22 Table 3) though the diameter range and the number of striae in 10 microm werefound to be different Pantocsek (1901) stated in his diagnosis (p 104) that thevalves of Cyclotella balatonis are 16ndash20 microm in diameter but we have found bothsmaller and larger specimens (Table 3) The wider diameter range in our observationcould be related to a more complete morphological range of the species since thediatom population in Rusałka Lake was abundant However the time of samplingmay have also been of importance as the medium valve size of the species in our

(arrowed) at concentrically elevated valve face Fig 16 external view of the openings of central (right)and marginal (left) fultoportulae (arrowed) Fig 17 external view of the short and long striae of themarginal area Fig18 internal view of the rimoportula opening (arrowed) located close to the valvemargin and internal alveolar opening and central openings of numerous areolae and fultoportulae(arrowed) Fig 19 internal view of the divided thinner costa (marked) and internal opening of centralfultoportula (arrowed) Fig 20 internal view of the marginal fultoportulae located on the thicker costae(arrowed) Fig 21 external view of the valve face with marked notae Fig 22 internal view of thealveolar openings with thicker costa bearing marginal fultoportulae and internal rimoportula openinglocated near the margin (arrowed) Scale bar represents 5 microm for Figs 15 18 amp 21 4 microm for Fig 163 microm for Figs 19 amp 20 and 2 microm for Figs 17 amp 22

516

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Com

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of P

unct

icul

ata

bala

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s an

d m

orph

olog

ical

ly s

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r sp

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F ndash

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ral

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ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

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ata

Pun

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Pun

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ata

bala

toni

sba

lato

nis

bala

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spr

aete

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sapr

aete

rmis

sapr

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sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

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er-

(Lem

mer

-(E

hren

berg

)(E

hren

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)W

ojta

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Woj

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sson

Haringk

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n)H

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udzy

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Bud

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kaas

Cyc

lote

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Haringk

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on

as C

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as C

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tella

our

prae

term

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as C

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tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

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hren

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ntoc

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(Gru

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Ref

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kH

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1951

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104

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02 p

116

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10 p

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2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

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s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

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ndash12

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a s

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yse

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-w

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etw

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Int

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l-ca

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betw

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tene

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d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519

samples declined in subsequent winter-spring months which is a typical consequenceof diatom cell division The number of striae in 10 microm was found to be lower (11 to17 striae in 10 microm) in some specimens from Rusałka Lake than has been originallystated by Pantocsek (1901) It might be related to the striae counting method becauseeven at original Pantocseks drawings (Plate XV figs 319 332) number of striae in10 microm is lower when the Genkalrsquos method (1977) is applied It also remains inaccordance with our observations of the type material (Table 3) and the observationsin Houk et al (2010)

Puncticulata balatonis is morphologically similar to several species of the complexmaking it difficult for a non-specialist to distinguish them The species resemblesP radiosa but differs from it in valve face ultrastructure such as the location of theinternal rimoportula openings The valves of P radiosa have a more coarse andradially wrinkled central area in the valve face whereas P balatonis has a smootherand finely silicified valve face Additionally the radial pattern of the valve facefultoportulae and areolae is very distinct in P radiosa The external rimoportulaopenings of P balatonis are situated at the end of shortened stria well discernible inLM as gaps (notae) whereas according to Houk et al (2010) the external openingsof P radiosa are situated at the end of only slightly shortened striae The differencesare clearly visible from the internal valvae view because P balatonis has valve facerimoportula openings located close to the internal alveolar openings whereas the

Fig 23 The variation in Puncticulata balatonis valve size in Rusałka Lake in subsequent months in2007

520

rimoportula openings in P radiosa are located more close to the central area ofthe valves Such a close location of the internal rimoportula openings to the valvemargin is typical only for P balatonis within the discussed group (Table 3)

Another diatom showing a similar morphology to Puncticulata balatonis is P prae-termissa (Lund) Haringkansson Most of the morphological features are similar forboth species (Table 3) but the later possesses radially elongated relatively largeinternal alveolar openings Moreover the costae bearing marginal fultoportulaeare only slightly thicker than the rest and the internal rimoportula openings arecloser to the central part of the valve face (Table 3) The last feature as well as themarginal stria length enable also the morphological distinction between P balatonisand P comta

Ecology and distribution

The distinction between the species of the Puncticulata radiosa-complex includingPuncticulata balatonis is not easy for a plankton ecologist or other researcher whois not a centric diatom specialist The correct identification requires detailedmorphological studies with SEM andor specialized knowledge That is why apartfrom the long-lasting taxonomical confusion the real distribution of the speciesfrom the complex is uncertain According to Houk et al (2010) Puncticulata balatonis(as Cyclotella balatonis) is one of the most common members of the P radiosa (as

Fig 24 The number of Puncticulata balatonis frustules in the water column of Rusałka Lake in 2007

521

Cyclotella radiosa) complex in Europe found in numerous mesotrophic and eutrophiclakes and slowly running waters in Hungary Germany and Czech Republic Howeverreports of the species are not numerous Till now P balatonis has also never beenreported from Poland

The limited number of reports on the occurrence of P balatonis may be related to itsmisidentificationmisinterpretation as Cyclotella comta (Ehrenb) Kuumltz (Kiss amp Pająk1994 fig 13) found in the plankton of the eutrophic Vistula River as Cyclotellapraetermissa quadrijuncta (Wunsam et al 1995) or as Puncticulata radiosa (Wojtalamp Kwandrans 2006) found in the eutrophic Rudawa River We are convinced that apart of other records of P comta the most commonly reported species of the complexin Poland (Siemińska amp Wołowski 2003) in fact might concern P balatonis InRusałka Lake the species may have also been mistaken for P comta (asC comta) reported from the lake by Kotlińska (1976) It is also likely that P bala-tonis in Poland may have been misidentified as P praetermissa although the latterspecies was only reported several times including findings in the benthos of CzarneLake and the Kamionka River NE Poland (Rakowska 1996) in the plankton ofseveral lakes of the Drawa National Park NW Poland (Gołdyn amp Szeląg-Wasielewska2004) in a sand-pit lake in Krakoacutew and a spring fed pond in Krztynia in S Poland(Wojtal unpublished data)

Up to now there has also been much confusion around the ecology of the speciesfrom the complex For instance P praetermissa described by Lund (1951) fromlakes with a low organic matter content was later found both in the plankton and thebenthos in a wide variety of natural and man-made lakes including alpine andsubalpine lakes (Wunsam et al 1995 Lange-Bertalot amp Metzeltin 1996 Guumlttingeramp Straub 1998 Tolotti et al 2004) as well as in more eutrophic habitats (egSchmidt et al 2002 Hindaacutek amp Hindaacutekovaacute 2003 Tanaka 2007) Houk et al (2010)cleared up the ambiguities ascribing P praetermissa and P radiosa to oligo-mesotrophic habitats According to the authors P comta (as Cyclotella comta) is tillnow known only from fossil materials P balatonis (as Cyclotella balatonis) inturn has been assigned to meso-eutrophic lakes and rivers consistently with thetrophic character of its type locality ndash lake Balaton Our results correspond with theautecology of P balatonis (as Cyclotella balatonis) proposed by Houk et al (2010)widening even slightly the preferences of the species for the trophic conditionsdetermined by Houk et al (2010) as it was also able to develop a large populationin an eutrophic to hypereutrophic water body

The highest numbers of P balatonis in Rusałka Lake were observed in the winter-spring period The diatom bloom in the spring plankton is a well-known phenomenon(Sommer et al 1986) ndash sometimes also occurring during the winter period (Kiss ampGenkal 1993) ndash and is usually associated with a changing silica andor phosphorusavailability during that period (Lampert amp Sommer 2001) Sinking was also provedto be an essential factor hampering diatom success during the thermal stratificationperiod (Jewson et al 1981 Ferris amp Lehman 2007) explaining probably the seasonalityof the P balatonis growth in Rusałka Lake as its high development finished withthe stratification onset Relatively high numbers of frustules in deeper water layersmight also indicate a significant role of the sedimentation process

522

Conclusions

The morphology of Puncticulata balatonis in Rusałka Lake corresponds well toother species around P comta with the exception of the location of marginalfultoportula openings and the length of the shortened striae (notae) Analysis of themorphological characters enabled the transfer of Cyclotella balatonis into the genusPuncticulata A large population of the species in the eutrophic to hypereutrophicRusałka Lake remains congruent with its tolerance ranges with respect to the trophicstate proposed by Houk et al (2010) The variation in the cell number and averagedimension of frustules indicated the species preference to develop large populationsin spring

Acknowledgements

We thank Prof Ryszard Gołdyn (AMickiewicz University in Poznań) for his inspiration researchsupervision and useful comments to the manuscript We are also grateful to Dr Bart Van de Vijver andtwo anonymous reviewers for their valuable suggestions and to Dr Krisztina Buczko for enabling theanalysis of the type material of Cyclotella balatonis from the Pantocsekrsquos collection at HungarianNatural History Museum Sampling on Rusałka Lake was part of a project financially supported by theDistrict and Municipial Funds of Enviromental Protection and Water Management in Poznań (Poland)

References

CARLSON RE (1977) A trophic state index for lakes ndash Limnol Oceanogr 22 (2) 361ndash369

CREMER H amp H KOOLMEES (2010) Common diatoms (Centrales and Fragilariaceae Bacillario-phyta) of modern and fossil freshwater environments in the Netherlands ndash Nova Hedwigia 90(3ndash4)343ndash381

FERRIS JA amp JT LEHMAN (2007) Interannual variation in diatom bloom dynamics Roles ofhydrology nutrient limitation sinking and whole lake manipulation ndash Water Res 41 2551ndash2562

GENKAL SI (1977) On the method of calculation of some taxonomicaly significant structuralelements of the valve in the diatoms of the family Thalassiosiraceae Lebour emend Hasle (Bacillariophyta)ndash Bot Zhurn 62 848ndash851 (in Russian)

GENKAL SI amp KT KISS (1993) Morphological variability of the diatom Cyclotella atomusHustedt var atomus and C atomus var gracilis var nov ndash Hydrobiologia 269270 39ndash47

GOŁDYN R S PODSIADŁOWSKI K KOWALCZEWSKA-MADURA R DONDAJEWSKAE SZELĄG-WASIELEWSKA A BUDZYŃSKA P DOMEK amp W ROMANOWICZ (2010)Functioning of the Rusałka Lake ecosystem in Poznań (west Poland) ndash Ocean Hydrobiol Stud39(3) 65ndash80

GOŁDYN R amp E SZELĄG-WASIELEWSKA (2004) Changes in the phytoseston of a river-lakesystem in Drawieński National Park ndash Ocean Hydrobiol Stud 33(2) 17ndash28

GUumlTTINGER W amp F STRAUB (1998) Diatoms of Lake Cadagno ndash In PEDUZZI RR BACHOFEN amp M TONOLLA (eds) Lake Cadagno a meromictic Alpine lake ndash Doc Ist ItalIdrobiol 63 57ndash64

HANDMANN R (1913) Die Diatomeenflora des Almfeegebiets ndash In MANZ GJ (ed) Mitteilungendes Mikrologischen Vereins Linz ndash Mikrolog Ver Regensburg (in German)

HAUSMANN S amp AF LOTTER (2001) Morphological variation within the diatom taxon Cyclotellacomensis and its importance for quantitative temperature reconstructions ndash Freshw Biol 46 1323ndash1333

523

HAYASHI T (2007) Puncticulata versiformis sp nov and Cyclotella kathmanduensis sp nov(Bacillariophyta) new fossil species from middle Pleistocene lacustrine sediments Kathmandu NepalHilalaya ndash J Phycol 43(2) 304ndash318

HAringKANSSON H (2002) A compilation and evaluation of species in the genera StephanodiscusCyclostephanos and Cylotella with a new genus in the family Stephanodiscaceae ndash Diatom Res 17(1)1ndash139

HINDAacuteK F amp A HINDAacuteKOVAacute (2003) Diversity of cyanobacteria and algae of urban gravel pitlakes in Bratislava Slovakia a survey ndash Hydrobiologia 506ndash509 155ndash162

HINDAacuteK F amp A HINDAacuteKOVAacute (2008) Diversity of cyanobacteria and algae of the NationalNature Reserve Čiacutečovskeacute mŕtve rameno oxbow (Southern Slovakia) ndash Bulletin SBS 30 (1) 11ndash19 (inSlovak with an English abstract)

HOUK V R KLEE amp H TANAKA (2010) Atlas of freshwater centric diatoms with a brief key anddescriptions Part III Stephanodiscaceae A Cyclotella Tertiarius Discostella ndash Fottea 10 (Supplement)1ndash498

JEWSON DH BH RIPPEY amp WJK GILMORE (1981) Loss rates from sedimentation parasitismand grazing during the growth nutrient limitation and dormancy of a diatom crop ndash Limnol Oceanogr26 1045ndash1056

KISS KT amp SI GENKAL (1993) Winter blooms of centric diatoms in the River Danube and in itsside arms near Budapest ndash Hydrobiologia 269270 317ndash325

KISS KT amp G PAJĄK (1994) Seasonal change of diatom s in the plankton of the Vistula riverabove and below the Goczalkowice reservoir Poland ndash In KOCIOLEK JP (ed) Proceedings ofthe 11th International Diatom Symposium San Francisco Aug 12ndash17 1990 Memoirs of the CaliforniaAcademy of Sciences 17 517ndash524 San Francisco California Academy of Sciences

KOBAYASI H M IDEI S MAYAMA T NAGUMO amp K OSADA (2006) H KobayasirsquosAtlas of Japanese diatoms based on electron microscopy 1 1ndash531 Uchida Rokakuho Publishing CoLtd Tokyo

KOTLIŃSKA E (1976) Fitoplankton Jeziora Rusałka In Fitoplankton sztucznych jezior położonychna terenie Poznania 1ndash54 Poznańskie Tow Przyjacioacuteł Nauk Prace Komisji Biologicznej (in Polish)

KRAMMER K amp H LANGE-BERTALOT (1991) Bacillariophyceae 3 Teil Centrales Fragilaria-ceae Eunotiaceae ndash In ETTL H J GERLOFF H HEYNIGH amp D MOLLENHAUER (eds)Suumlsswasserflora von Mitteleuropa 2(3) 1ndash576 Gustav Fischer Verlag Stuttgart Jena

LAMPERT W amp U SOMMER (2001) Ekologia woacuted śroacutedlądowych ndash PWN Warszawa

LANGE-BERTALOT H amp D METZELTIN (1996) Indicators of oligotrophy 800 representativeof three ecologically distinct lake types Carbonate buffered-Oligotrophic-Weakly buffered soft waterndash Iconogr Diatomol 2 1ndash390

LUND JWG (1951) Contribution to our knowledge of British algae ndash Hydrobiologia 3(1) 93ndash100

ORGANIZATION FOR ECONOMIC COOPERATION AND DEVELOPMENT (1982)Eutrophication of Waters ndash Monitoring Assessment and Control Final Report OECD Paris

PANTOCSEK J (1901) Die Kieselalgen oder Bacillarien des Balaton 1ndash112 XVI Tafeln ndash Druckder K und K Hofbuchdruckerei des Victor Hornyaacutenszky Budapest

RAKOWSKA B (1996) Preliminary elaboration of the diatom communities occuring in the benthosof the Kamionka River and Lake Czarne at the Pojezierze Suwalskie (1993ndash1994) ndash Fragm FloristGeobot Polon 3 221ndash238

RIMET F JC DRUART amp O ANNEVILLE (2009) Exploring the dynamics of plankton diatomcommunities in Lake Geneva using emergent self-organizing maps (1974ndash2007) ndash Ecol Inform4 99ndash110

524

SCHMIDT R R PSENNER J MUumlLLER P INDINGER amp C KAMENIK (2002) Impact of thelate glacial climate variations on stratification and trophic state of the meromictic lake Laumlngsee (Austria)validation of a conceptual model by multiproxy studies ndash J Limnol 61(1) 49ndash60

SIEMIŃSKA J amp K WOŁOWSKI (2003) Catalogue of Polish prokaryotic and eukaryotic algaendash W Szafer Institute of Botany Polish Academy of Sciences Krakoacutew

SIVER PA (1999) Development of paleolimnological inference models for pH total nitrogen andspecific conductivity based on planktonic diatoms ndash J Paleolimnol 21 45ndash59

SOMMER U ZM GLIWICZ W LAMPERT amp A DUNCAN (1986) The PEG-model of seasonalsuccession of planktonic events in fresh waters ndash Arch Hydrobiol 106 433ndash471

TANAKA H (2007) Taxonomic studies of the genera Cyclotella (Kuumltzing) Breacutebisson DiscostellaHouk et Klee and Puncticulata Haringkansson in the family Stepanodiscaceae Glezer et Makarova(Bacillariophyta) in Japan ndash Biblioth Diatomol 53 1ndash205

TOLOTTI M D CALLIARI amp F CORRADINI (2004) Interannual variability of the phytoplanktonof Tovel lake (Trentino Italy) ndash Studi Trent di Scienze Naturali Acta Biol 81 Suppl 2 327ndash340 (inItalian)

WETZEL RG amp GE LIKENS (1991) Limnological analyses 1ndash391 ndash Springer-Verlag NewYork Berlin Heidelberg

WOJTAL AZ amp J KWANDRANS (2006) Diatoms of the Wyżyna Krakowsko-Częstochowskaupland (S Poland) ndash Coscinodiscophyceae (Thalassiosirophycidae) ndash Polish Bot J 51(2) 177ndash207

WUNSAM S R SCHMIDT amp R KLEE (1995) Cyclotella-taxa (Bacillariophyceae) in lakes of theAlpine region and their relationship to environmental variables ndash Aquat Sci 57(4) 360ndash381

Received 25 March 2010 accepted in revised form 31 March 2011

510

Despite some of the members of Cyclotella genus are known for up to 170 years theecology of many species still remains uncertain Moreover there are also manyuncertainties within the genus concerning taxonomy and nomenclature (Hausmannamp Lotter 2001 Haringkansson 2002 Houk et al 2010) Recently some species of thenot-uniform genus were moved to several newly-established genera (short review inHouk et al 2010) In 2002 the genus Puncticulata was established comprisingspecies characterised by concentrically undulated central part of valve face andnumerous scattered or radially arranged areolae and fultoportulae (or only areolaeor only fultoportulae) with P comta as the type (Haringkansson 2002) The genus hasbeen generally accepted (eg Tanaka 2007 Hayashi 2007 Kobayasi et al 2006Cremer amp Koolmees 2010) though some uncertainties are still discussed (Houk etal 2010)

One of the species sharing the above-mentioned characteristics of the genusPuncticulata (though not transferred to the genus till now) is Cyclotella balatonisand its variety C balatonis var binotata described by Pantocsek (1901 p 104 figs319 331 332) from lake Balaton near Sioacutefok in Hungary Originally the specieswas illustrated by three drawings of valves with scattered (Pantocsek 1901 PlateXV figs 319 332) and radially arranged (Pantocsek 1901 Plate XV fig 331)puncta in the middle part of valve face After Pantocsekrsquos description C balatonisremained practically forgotten and was reported only a few times (eg Handmann1913 Hindaacutek amp Hindaacutekovaacute 2008) It was generally considered a synonym ofCyclotella radiosa (eg Krammer amp Lange-Bertalot 1991 fig 62 11 12) followingthe broad species concept that was in use at that time Thus specimens of a broadmorphological range present in a variety of water bodies were identified as C radi-osa Cyclotella balatonis might have also been misidentified as other species of thecomplex such as P praetermissa (syn Cyclotella praetermissa) or Puncticulatacomta (syn Cyclotella comta) Recently as a result of precise study by Houk et al(2010) using SEM Cyclotella balatonis has been resurrected and the morphologicalranges of similar species were discussed However due to the proceeding long periodof taxonomical confusion the species is almost absent in the literature and needsmuch attention particularly in terms of distribution and autecology

In this paper a population of C balatonis in a small and shallow urban eutrophic tohypereutrophic lake (Rusałka Lake Western Poland) is presented with its quantitativecharacteristics along an annual cycle Morphological features of the diatom aredescribed in detail and documented with LM photographs of live specimens andcleaned frustules as well as with the SEM micrographs Cyclotella balatonis wasalso compared with morphologically similar species namely Puncticulata comtaP praetermissa and P radiosa Due to the great morphological similarity of Cyclotellabalatonis to these and other Puncticulata-species the transfer of Cyclotella balatonisto the genus Puncticulata has been proposed

Study site

Rusałka Lake is situated in the city of Poznań Western Poland (N52deg25E16deg52) It was formed in1943 by damming the Bogdanka River and subsequent flooding of its valley The lake covers the area

511







2O











512


Results









513

Ecological data




514


515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

Ext

erna

lE

xter

nal

posi

tion

lo-

tene

d st

ria

area

Ext

erna

lte

ned

stri

ace

ntra

l are

ate

ned

stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

Inte

rnal

ly a

Ext

erna

l ope

-In

tern

ally

aIn

tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











511







2O











512


Results









513

Ecological data




514


515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

Ext

erna

lE

xter

nal

posi

tion

lo-

tene

d st

ria

area

Ext

erna

lte

ned

stri

ace

ntra

l are

ate

ned

stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

Inte

rnal

ly a

Ext

erna

l ope

-In

tern

ally

aIn

tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











512


Results









513

Ecological data




514


515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

Ext

erna

lE

xter

nal

posi

tion

lo-

tene

d st

ria

area

Ext

erna

lte

ned

stri

ace

ntra

l are

ate

ned

stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

Inte

rnal

ly a

Ext

erna

l ope

-In

tern

ally

aIn

tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











513

Ecological data




514


515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

Ext

erna

lE

xter

nal

posi

tion

lo-

tene

d st

ria

area

Ext

erna

lte

ned

stri

ace

ntra

l are

ate

ned

stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

Inte

rnal

ly a

Ext

erna

l ope

-In

tern

ally

aIn

tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











514


515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

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ith sc

atte

red

undu

late

un

dula

te

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late

and

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late

un

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te

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te w

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ith C

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ith n

ume-

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dial

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ooth

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th C

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ous C

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kled

C

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ed C

Fan

d ar

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d ar

eola

ear

rang

ed in

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ope-

punc

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d ar

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eC

F sc

atte

red

and

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and

areo

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scat

tere

d or

open

ings

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e or

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ning

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ange

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ed in

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ays

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atte

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sor

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nal-

in ra

dial

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al p

atte

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ed in

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al ro

ws

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nged

inly

arr

ange

d in

row

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dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

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in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

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the

valv

aem

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nal a

rea

on th

e va

lve

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mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

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ile la

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face

at t

hecl

ose

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mit

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hear

ea c

lose

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t the

face

at t

heva

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gin

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e mar

gin

valv

e mar

gin

um in

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ique

end

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of th

e ce

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d of

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tern

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d of

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sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

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rge

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ilese

ssile

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umte

ned

stri

ash

orte

ned

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tene

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in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

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tene

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n lo

-of

a s

light

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tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

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tene

dte

d be

twee

nin

the

mar

gina

lto

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rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

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e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











515



Discussion





516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

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lls s

olita

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dat

alo

osel

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osel

yor

in s

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ary

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ly in

unite

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ited

unite

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ely

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ains

2(8

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16ndash2

016

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87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

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heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

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entri

cally

conc

entri

cally

with

mor

e or

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late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

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cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

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late

un

dula

te

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te w

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ith C

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ith n

ume-

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dial

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ooth

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th C

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mer

ous C

Far

eola

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us C

F an

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dial

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and

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kled

C

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ed C

Fan

d ar

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ear

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ed in

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eC

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atte

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tere

d or

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ings

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e or

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ning

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ttere

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arr

ange

dar

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ed in

alw

ays

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nged

insc

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red

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dial

row

sor

occ

asio

nal-

in ra

dial

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al p

atte

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rang

ed in

radi

al ro

ws

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nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

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in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

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the

valv

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nal a

rea

on th

e va

lve

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mar

gina

lon

the v

alva

eon

the

valv

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ea n

ear t

hear

ea n

ear t

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ea n

ear t

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sess

ile la

bi-

face

at t

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ose

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mit

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at t

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ea c

lose

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ce a

t the

face

at t

heva

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gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

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of th

e ce

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len

d of

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it of

the

end

of s

hor-

end

of s

light

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nal o

pe-

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erna

lE

xter

nal

posi

tion

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ria

area

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erna

lte

ned

stri

ace

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l are

ate

ned

stri

ash

orte

ned

stri

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ngs

at th

eop

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gs a

top

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gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

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rnal

ly a

Ext

erna

l ope

-In

tern

ally

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tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











516

Tab

le 3

Com

pari

son

of P

unct

icul

ata

bala

toni

s an

d m

orph

olog

ical

ly s

imila

r sp

ecie

s C

F ndash

cent

ral

fulto

port

ulae

MF

ndash m

argi

nal

fulto

port

ulae

o

urob

serv

atio

n ty

pe m

ater

ial S

iofoacute

k s

ampl

e no

152

our

obse

rvat

ion

base

d on

the

fig

62

11

12

in K

ram

mer

amp L

ange

-Ber

talo

t 199

1 w

hich

sho

ws

spec

imen

s fr

om th

e ty

pe m

ater

ial o

f Cyc

lote

lla

bala

toni

s Pa

ntoc

sek

(as

C r

adio

sa)

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

Ref

eren

ces

Pant

ocse

kH

ouk

et a

lL

und

1951

H

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

lH

aringkan

sson

H

ouk

et a

l19

01 p

104

20

10 p

39

p 9

820

02 p

116

20

10 p

41

2002

p 1

142

010

p 3

720

02 p

113

20

10 p

36

Plat

e X

V

Plat

es 2

69

Figs

1 2

Figs

422

ndashPl

ates

280

Fi

gs 4

15ndash

Plat

es 2

61

Figs

410

ndash P

late

s 25

7 1

ndashFi

gs 3

19

1ndash11

270

42

61ndash

11 2

81

421

1ndash11

262

41

423

258

1ndash1

133

21ndash

6 2

71

1ndash6

1ndash6

1ndash12

263

25

9 1

ndash12

1ndash5

264

1ndash6

260

1ndash8

Col

onie

sno

dat

a2ndash

82ndash

7 ce

lls2ndash

8 ce

lls

shor

tC

ells

solit

ary

no d

ata

mos

tlyce

lls s

olita

ryno

dat

alo

osel

ylo

osel

ylo

osel

yor

in s

hort

solit

ary

rare

ly in

unite

dun

ited

unite

d ch

ains

loos

ely

unite

dsh

ort c

hain

sch

ains

2(8

)ce

llsD

iam

eter

16ndash2

016

ndash22

87ndash

210

8ndash25

8ndash25

8ndash30

7ndash35

7ndash25

8ndash50

5ndash32

(microm

)St

riae

20 p

er 1

0 microm

ca

16

per

15ndash1

85

per

13ndash1

9 pe

r13

ndash15

per

15ndash1

8 pe

r 10

no d

ata

15ndash1

8 pe

r 10

no d

ata

14ndash1

8 pe

r10

microm

of

10 micro

m10

microm

10 micro

mmicrom

of n

earl

ymicrom

of n

early

10 micro

mne

arly

equ

aleq

ual l

engt

heq

ual l

engt

hle

ngth

occ

upy

occu

py 1

2ndash1

4ta

king

abo

ut1

3ndash1

2 of

of th

e va

lvae

12

of th

e va

lve

valv

e fa

cedi

amet

er fa

ce d

iam

eter

diam

eter

517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

Ext

erna

lE

xter

nal

posi

tion

lo-

tene

d st

ria

area

Ext

erna

lte

ned

stri

ace

ntra

l are

ate

ned

stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

Inte

rnal

ly a

open

ings

on

Inte

rnal

ly a

Ext

erna

l ope

-In

tern

ally

aIn

tern

ally

a la

rge

end

of s

hor-

the

end

ofth

e en

d of

alve

olar

sess

ile la

bi-

a hy

alin

e ar

eare

lativ

ely

ning

s on

ala

rge

sess

ilese

ssile

labi

umte

ned

stri

ash

orte

ned

shor

tene

dop

enin

gsum

in ra

dial

at th

e en

d of

larg

e se

ssile

hyal

ine

area

labi

um in

in o

bliq

ue o

rIn

tern

ally

ast

ria

Inte

r-st

ria

Inte

r-an

d ce

ntra

lpo

sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

dob

lique

po-

circ

umfe

rent

ial

smal

l ses

sile

nally

a se

ssile

nally

a sm

all

area

cate

d m

id-

shor

tene

dpo

sitio

n lo

-of

a s

light

lysi

tion

loca

-po

sitio

n lo

cate

dla

bium

clo

sela

bium

ver

yse

ssile

labi

-w

ay b

etw

een

stria

Int

erna

l-ca

ted

betw

een

shor

tene

dte

d be

twee

nin

the

mar

gina

lto

inte

rnal

clos

e to

inte

r-um

clo

se to

alve

olar

ope

-ly

a s

essi

leal

veol

ar o

pe-

stri

a In

ter-

alve

olar

ope

-ar

ea n

ear t

o th

eal

veol

ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











517

Cen

tral

are

a

flat

tofla

t to

slig

htly

flat t

o sl

ight

lyce

ntra

l are

aco

ncen

trica

llyfla

t or s

light

lysl

ight

ly c

on-

dist

inct

ly c

on-f

lat t

o co

n-fl

at to

con

-sl

ight

ly c

on-

conc

entri

cally

conc

entri

cally

with

mor

e or

undu

late

co

ncen

trica

llyce

ntri

cally

cent

rica

llyce

ntri

cally

cent

rica

llyce

ntric

ally

un-

undu

late

un

dula

te

less

regu

larl

yw

ith sc

atte

red

undu

late

un

dula

te

undu

late

and

undu

late

un

dula

te

dula

te w

ithw

ith C

F an

dw

ith n

ume-

arra

nged

or ra

dial

lyce

ntra

lra

dial

lyra

dial

lysm

ooth

smoo

th C

Fnu

mer

ous C

Far

eola

ero

us C

F an

dra

dial

arra

nged

CF

areo

lae

and

wrin

kled

C

Fw

rinkl

ed C

Fan

d ar

eola

ean

d ar

eola

ear

rang

ed in

areo

lae

ope-

punc

taan

d ar

eola

eC

F sc

atte

red

and

areo

lae

and

areo

lae

scat

tere

d or

open

ings

mor

e or

less

ning

s sca

ttere

dor

arr

ange

dar

rang

ed in

alw

ays

arra

nged

insc

atte

red

orra

dial

row

sor

occ

asio

nal-

in ra

dial

radi

al p

atte

rnar

rang

ed in

radi

al ro

ws

arra

nged

inly

arr

ange

d in

row

sra

dial

row

sra

dial

pat

tern

radi

al p

atte

rn(F

igs

8 amp

9)

Rim

o-2

ndash3 in

the

(1)2

ndash5 in

the

1ndash2(

3) in

the

no d

ata

at le

ast o

ne

1ndash5

in th

eat

leas

t one

at

leas

t one

in

at le

ast o

ne

at le

ast o

ne

port

ulae

mar

gina

lm

argi

nal

mar

gina

l

Inte

rnal

lyon

the

valv

aem

argi

nal a

rea

on th

e va

lve

the

mar

gina

lon

the v

alva

eon

the

valv

aear

ea n

ear t

hear

ea n

ear t

hear

ea n

ear t

hea

sess

ile la

bi-

face

at t

hecl

ose

to li

mit

face

at t

hear

ea c

lose

tofa

ce a

t the

face

at t

heva

lve

mar

gin

valv

e mar

gin

valv

e mar

gin

um in

obl

ique

end

of s

hor-

of th

e ce

ntra

len

d of

sho

r-lim

it of

the

end

of s

hor-

end

of s

light

lyE

xter

nal o

pe-

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erna

lE

xter

nal

posi

tion

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ria

area

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erna

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ned

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ace

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l are

ate

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stri

ash

orte

ned

stri

ani

ngs

at th

eop

enin

gs a

top

enin

gs a

tca

ted

betw

een

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rnal

ly a

open

ings

on

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rnal

ly a

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erna

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tern

ally

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tern

ally

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rge

end

of s

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the

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e en

d of

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olar

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a hy

alin

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ely

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umte

ned

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e en

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e se

ssile

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ine

area

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um in

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ue o

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tern

ally

ast

ria

Inte

r-st

ria

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r-an

d ce

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sitio

n lo

-a

slig

htly

labi

um in

slit

at th

e en

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lique

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circ

umfe

rent

ial

smal

l ses

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ssile

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all

area

cate

d m

id-

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tene

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n lo

-of

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light

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tion

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n lo

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ay b

etw

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erna

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ted

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d be

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the

mar

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lto

inte

rnal

clos

e to

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se to

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ope

-ly

a s

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ar o

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stri

a In

ter-

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olar

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-ar

ea n

ear t

o th

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ar o

pe-

nal a

lveo

lar

inte

rnal

alv

e-ni

ngs

and

labi

um in

ning

s an

dna

lly a

ses

-ni

ngs

and

cent

ral a

rea

ning

s w

ithop

enin

gs

olar

ope

ning

sce

ntra

l are

aob

lique

to c

ir-ce

ntra

l are

asi

le la

bium

cent

ral a

rea

radi

ally

or

with

radi

ally

with

radi

ally

cum

fere

ntia

lw

ith a

slit

ori-

obliq

uely

po-

or o

bliq

uely

or o

bliq

uely

slit

posi

tion

ente

d ob

lique

-si

tione

d sl

itpo

sitio

ned

posi

tione

d sl

itly

to n

earl

ysl

itci

rcum

fere

n-tia

llyIn

tern

alsm

all

cost

aesm

all

cost

aesm

all

cost

ae

radi

ally

radi

ally

elon

-ra

dial

ly e

lon-

smal

l co

stae

smal

l co

stae

smal

l co

stae

smal

l co

stae

alve

olar

bear

ing

MF

bear

ing

MF

bear

ing

MF

elon

gate

dga

ted

cos

tae

gate

d c

osta

ebe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fbe

arin

g M

Fop

enin

gscl

early

larg

ercl

early

larg

ercl

early

larg

erco

stae

bea

-be

arin

g M

Fbe

arin

g M

Fcl

earl

y la

rger

clea

rly la

rger

clea

rly la

rger

clea

rly

larg

erth

an th

in ri

bsth

an th

in ri

bsth

an th

in ri

bsri

ng M

Fcl

early

larg

ersl

ight

ly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

than

thin

ribs

clea

rly la

rger

than

thin

ribs

than

thin

ribs

than

thin

ribs

518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











518

Lak

e B

alat

onM

esot

roph

ic

Eut

roph

ic

Plan

kton

of a

no d

ata

Plan

kton

or

no d

ata

Plan

kton

of

no d

ata

foss

il di

atom

near

Sioacute

fok

eutr

ophi

chy

pere

utro

-nu

mbe

r of

litto

ral o

ligo

olig

om

eso-

lake

s re

ser

ph

ic re

serv

oir

lake

s po

orm

esot

roph

ictr

ophi

csl

owly

run

-in

org

anic

lake

sm

ainl

y su

b-ni

ng w

ater

sm

atte

r and

alpi

ne la

kes

One

of t

heso

me

quar

rym

ost c

omm

onpo

ols

inE

urop

ean

Eng

lish

Lak

esp

ecie

s of

the

Dis

tric

tC

-rad

iosa

com

plex

Tab

le 3

con

tinue

d

Cyc

lote

llaP

unct

icul

ata

Pun

ctic

ulat

aC

yclo

tella

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

Pun

ctic

ulat

aP

unct

icul

ata

bala

toni

sba

lato

nis

bala

toni

spr

aete

rmis

sapr

aete

rmis

sapr

aete

rmis

sara

dios

ara

dios

aco

mta

com

taPa

ntoc

sek

(Pan

tocs

ek)

(Pan

tocs

ek)

Lun

d(L

und)

(Lun

d)(L

emm

er-

(Lem

mer

-(E

hren

berg

)(E

hren

berg

)W

ojta

l et

Woj

tal e

tH

aringkan

sson

Haringk

anss

onm

ann)

man

n)H

aringkan

sson

Haringk

anss

onB

udzy

ńska

Bud

zyńs

kaas

Cyc

lote

llaH

aringkan

sson

Haringk

anss

on

as C

yclo

tella

as C

yclo

tella

our

prae

term

issa

as C

yclo

tella

com

taba

lato

nis

obse

rvat

ion

Lun

dra

dios

a(E

hren

berg

)Pa

ntoc

sek

(Gru

now

)K

uumltzi

ngL

emm

erm

ann

519




520






521





522

Conclusions


Acknowledgements


References











523



















524











519




520






521





522

Conclusions


Acknowledgements


References











523



















524











520






521





522

Conclusions


Acknowledgements


References











523



















524











521





522

Conclusions


Acknowledgements


References











523



















524











522

Conclusions


Acknowledgements


References











523



















524











523



















524











524











The centric diatom Puncticulata balatonis (Pantocsek) Wojtal et Budzy½ska, comb. nov., in the...

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Transcript of The centric diatom Puncticulata balatonis (Pantocsek) Wojtal et Budzy½ska, comb. nov., in the...