Avrils Limnol 21 (3) 1985: 213-220.

Raphe like vestiges in the pennate diatom suborder

Araphidinae ?

H. Lange-BertalotI
R. L e C o h u ?

K e y w o r d s : F r e s h w a t e r , D i a t o m s , Fragilaria, Achnanthes, pore, slit, r a p h e , like-vestiges, s e p t u m , K e r g u e l e n , W y o m i n g .

R a p h e - l i k e v e s t i g e s h a v e b e e n f o u n d r e g u l a r l y i n t w o n e w t a x a w h i c h p r e s u m a b l y b e l o n g to Fragilaria, one o f t h e m l i v i n g
in s e v e r a l l a k e s of t h e K e r g u e l e n I s l a n d s , the o t h e r in t h e Y e l l o w s t o n e N a t i o n a l P a r k , W y o m i n g , U . S . A . T h e l a t t e r o n e
p o s s e s s e s s e p l a - l i k e e l e m e n t s a s i n t e g r a t e d p a r t s o f a g i r d l e b a n d . O t h e r septa-like e l e m e n t s c a n o c c u r i n d i f f e r e n t s p e c i e s
of Fragilaria. T h e y r e p r e s e n t a p p e n d i c e s of the v a l v o c o p u l a , l o c a t e d b e n e a t h t h e t r a n s a p i c a l c o s t a e . S t r u c t u r e s like lin-
king s p i n e s o n m a r g i n a l p a r t s of the v a l v e f a c e o c c u r in a n Achnanthes s p e c i e s . M o s t of t h e s e c h a r a c t e r s s e e m to h a v e
b e e n u n k n o w n in t h e s e t a x a u n t i l recently.

Diatomées pennales avec des vestiges de raphes : un sous-ordre des Araphidînées ?

M o t s c l é s : e a u d o u c e , d i a t o m é e s . Fragilaria, Achnanthes, p o r e , fente, v e s t i g e s d e raphe, s e p t u m , K e r g u e l e n , W y o m i n g .

D e s v e s t i g e s d e r a p h é o n t é t é o b s e r v é s r é g u l i è r e m e n t c h e z d e u x n o u v e a u x t a x o n s a p p a r t e n a n t p r o b a b l e m e n t a u G . Fra-
gilaria. L ' u n d ' e u x a é t é r e n c o n t r é d a n s p l u s i e u r s lacs d e s î l e s K e r g u e l e n , l ' a u t r e d a n s le p a r c n a t i o n a l d e Y e l l o w s t o n e ,
W v o m i n g , E t a t s - U n i s . C e d e r n i e r p r é s e n t e d e s s t r u c t u r e s d e type s e p t u m i n t é g r é e s à l a c e i n t u r e connective, structures
p o u v a n t ê t r e o b s e r v é e s c h e z d i f f é r e n t e s e s p è c e s d e Fragilaria. Elles représentent des appendices d e l a valvo-copula, pla-
cés- a u - d e s s o u s d e s c ô t e s t r a n s a p i c a l e s . U n s y s t è m e d'indentations s e m b l a n t p e r m e t t r e l ' a g r é g a t i o n d e s f r u s t u l e s est s i g n a l é
c h e z u n Achnanthes. J u s q u ' à m a i n t e n a n t , la p l u p a r t d e s c a r a c t è r e s p r é c i t é s s e m b l e n t i n c o n n u s c h e z ces t a x o n s .

Introduction are c o m p a r a t i v e l y p o o r l y d e v e l o p e d w h e r e a s in t h e
e m e n d e d Achnanthes bioretii Germain they h a v e
In t w o p r e v i o u s p a p e r s it has been s h o w n that m o r e s i m i l a r i t y w i t h n o r m a l raphes and t h e r e f o r e
raoheless v a l v e s o f different Achnanthes species many p o p u l a t i o n s o r individuals o f this t a x o n h a v e
exhibit slits o f d i f f e r e n t lenght in distal parts o f the been taken f o r a Navicula, in particular Navicnla
v a l v e faces a n d i n a d d i t i o n the p r e s e n c e o f termi- rotaeana, by m a n y authors. M e a n w h i l e w e c o u l d
nal, in one case terminal and central pores (Le Cohu trace out c o m p a r a b l e structures o f different e x t e n t
& M a i l l a r d 1983, K r a m m e r & Lange-Bertalot 1985). in m a n y o t h e r species o f Achnanthes. M o r e o v e r , w e
Less distinct v e s t i g e s w e r e d o c u m e n t e d in Achnan- i n t r o d u c e t w o n e w taxa e x h i b i t i n g raphe-like vesti-
thes linkei Hustedt b y A n d r e w s ( 1981 ) and g e n e r a l l y ges, to all a p p e a r a n c e b e l o n g i n g t o the s u b f a m i l y o f
m e n t i o n e d by S i m o n s e n (1979). T h e s e vestiges w e r e the F r a g i l a r i o i d e a e , h o w e v e r , not without d o u b t t o
discussed as rudiments o r (less probably) precursors Fragilaria.
of true raphes w h i c h o t h e r w i s e w e r e fully d e v e l o -
ped in the o p p o s i t e v a l v e o f each frustule. In all dif- Observations
ferent species of the K e r g u e l e n Island these vestiges

1. Fragilaria maillardii Le Cohu nov. spec. (Fig. 1/3,

I. Botanisches Institut der J . W . Goethe-Universitât, Sies- 18/23, 24)
maverst. 70. Postfach 11 19 32 - 6000 Frankfurl a.M. (R.F.A.).
7. Laboratoire d'Hydrobiologie U.A. 695 C.N .R.S., Université P.
In f o u r lakes (Lac Alicia. Lac d e s K o r r i g a n s , L a c
Sahatier, 118 route de Narbonne. 31062 Toulouse, France. des Sternes, Lac S t u d e r ) o f the K e r g u e l e n Islands in

Article available at http://www.limnology-journal.org

or http://dx.doi.org/10.1051/limn/1985021
214 H. LANGE-BERTALOT E T R. LE COHU (2)

the s u b a n t a r c t i c r e g i o n o f the Indian ocean, one of rotundis apicibus. In SEM : marginales spinae in
us found p o p u l a t i o n s of an up to n o w u n k n o w n transapicalium costarum productione. Striae cons-
taxon. M o s t o f the m o r p h o l o g i c a l characteristics titutae simplici ordine a 4 ad 7 circinitas foraminas
point to Fragilaria, in particular to the taxonomi- quae continuant sine intermissu ad limbum. Area
c a l l y p r o b l e m a t i c g r o u p of forms around F. cons­ axialis linearis-elliptica dilatata in media parte.
truens. In L M no significant difference can be found Rimoportulae et porum apicalis area absentes. In api-
to s e p a r a t e it f r o m populations w h i c h a r e usually cale parte axialis areae, in valvarum interiore et
d e t e r m i n e d by m a n y authors as F. construens var. externo, praesentia pororum, saepe procerarum
venter ( E h r e n b e r g ) G r u n o w or as F. elliptica Schu­ rimarum forma ; pori similes vestigiis raphium
m a n n . W i t h o u t having c o n t r o l l e d the type m a t e r i a l observatis apud aliquuos Achnanthes aut recisis
of E h r e n b e r g o r Schumann, r e s p e c t i v e l y , and ramis raphis apud Rhoicosphenia curvata.
w i t h o u t k n o w l e d g e o f the m o r p h o l o g i c v a r i a b i l i t y
of taxa around F. construens, including varieties and 2. Fragilaria loetschertii Lange-Bertalot nov. spec.
t o n n a e , no m o r e distinctive determination seems to {Fig. 13/17, 30/34) and K r a m m e r & Lange-Bertalot
be possible in any case. T h e valves are mounted with 1985 {Fig. 32, 13, 33.1/5, 34.1/2)
linking spines on the m a r g i n s similar to those of the
so-called Fragilaria elliptica d e m o n s t r a t e d by A population of a second new taxon w a s found in
H a w o r t h (1975, S e m Fig. 14/21), somewhat less simi­ the Firehole River of the Yellowstone National Park,
lar to those o f A r c h i b a l d (1983, Fig. 519/522) w h e r e U.S.A. T h e SEM-structure of the specimens is less
the spines are d i r e c t l y situated on the striae (on similar to established taxa o f Fragilaria in contrast
i m e r c o s t a e in S E M ) , and m o r e o v e r the f o r a m i n a w i t h the Kerguelen populations. Beside regularly
b e c o m e l a r g e r n e a r e r to the margins. C o m p a r e o u r o c c u r r i n g raphe-like polar slits on both valves of a
L M and S E M figures on plate 1 and 2 r e p r e s e n t i n g frustule, it is the presence of a peculiar connecting
Fragilaria elliptica in contrast with F. construens band with septa projecting unilaterally into the inte­
var. venter. T h e o u t l i n e of specimens of the n e w r i o r of the frustule. These septa are gradually redu­
taxon v a r i e s f r o m e l l i p t i c a l to linear w i t h a p p r o x i ­ c e d in length from one valve apex to the other. In
mately bluntly rounded apices. T h e striae consist on L M the specimens can hardly be distinguished from
each side of the axial area of s i m p l e r o w s o f * 4-7 Fragilaria pinnata, h o w e v e r , in S E M the distinct
r o u n d f o r a m i n a , r e s p e c t i v e l y areolae, on the valve double (locally three) r o w s of areolae of each tran-
face running w i t h o u t interruption s o m e w h a t o v e r sapical stria are a third m o r p h o l o g i c character dif­
the m a n t l e . T h e axial area is linear-elliptical, m o r e ferentiating this taxon f r o m F. pinnata and other
o r less e n l a r g e d in the central part. N e i t h e r a r i m o - known species of this genus. Frustules in girdle view
portula nor apical pore fields can be detected. H o w e ­ f o r m i n g ribbon-like chains. V a l v e s elliptical to
v e r , e v e r y v a l v e s h o w s n e a r the apices, inside and linear-elliptical, rarely slightly ovate, w i t h broadly
o u t s i d e , apical r u n n i n g * short slits o r a p i c a l l y rounded apices, ca. 6-14 ^ m long, ca. 4-6 ^ m w i d e .
elongated pores, c o m p a r a b l e to reduced raphe bran­ Axial area narrow, linear. Striae coarse, parallel in
ches in the a f o r e m e n t i o n e d rapheless valves of Ach­ the centre, radiate t o w a r d s the apices of the valve,
nanthes o r the t y p i c a l l y shortened ones of Rhoicos- 6-12/10 ^ m . In S E M : linking-spines arranged near
phenia. In s o m e cases a terminal p o r e seems to be the edge on transapical costae. S i m p l e or bifurcate
developed. tips of the spines touching the transapical striae of
an adjacent frustule. Each stria is f o r m e d of dou­
b l e (rarely three) r o w s of foramina. Each valve face
Diagnosis. H o l o t y p u s : Coll. L e Cohu K e r g . K o r r .
of a frustule, outside and inside, with raphe-like
n ° 23.
short slits near the apices. R i m o p o r t u l a e o r apical
T y p e locality : Kerguelen Islands, Lac des p o r e fields absent. Connecting band with septa uni­
Korrigans. laterally in the frustule.
T h i s s p e c i e s is d e d i c a t e d in honor to R. M a i l l a r d ,
diatomist, Evreux.
Diagnosis. H o l o t y p u s : Coll. Lange-Bertalot nor.
In L.M. : Nulla significans differentia inter haec AmNlO.
species et Fragilaria construens var. venter aut Fra­ Type locality : U.S.A., W y o m i n g , Yellowstone Nat.
gilaria elliptica. Valva elliptica vel Linearis cum Park, F i r e h o l e .
(3> RAPHE L I K E V E S T I G E S IN T H E P E N N A T E DIATOM S U B O R D E R A R A P H I D I N E A E 215

River, 5.6.1983, L e g . Ute R u m r i c h . as a g r o u p which has largely been neglected b y taxo-

This species is d e d i c a t e d in honor to my collea- n o m i c and m o r p h o l o g i c w o r k and he states that o u r
gue, late Dr. W i l h e l m Lotschert, ecologist in ignorance m a k e s them the weak link in the Penna-
Frankfurt-am-Main. les. M o r e o v e r it should be re-evaluated, w h e t h e r o r
not the b i p a r t i t i o n of the Fragilariceae ( « Diatoma-
Frustula e facie connectivali reclangulares, cate- ceae ») into T a b e l l a r i o i d e a e and the F r a g i l a r i o i d e a e
nas congestas formantes. Valvae ellipticae usque ad is tenable by the traditional criteria like the f o r m a -
lineari-elïipticae, raro leniter claviformes, apicibus tion of septa.
late rotundatis, circiter 6-14 ^m longae, 4-6 ^m latae.
Area axialis anguste linearis. Striae transapicales in Both n e w taxa w h i c h are c o m b i n e d here w i t h Fra-
partibus medianibus valvae parallelae ad polos ver- gilaria — m a y b e in a provisional arrangement — are
sus radiantes, 6-12/10 ^m. In SEM : marginales spi- lacking any r i m o p o r t u l a and apical pore fields. A s
nae in transapicalium costarum productione cum known so far, the first is a species-specific faculta-
simpHcibus aut bifurcis extremitatibus quae contin- tive character in the subfamily. T h e latter, h o w e v e r ,
junt confinis frustulae transapicales strias. Striae seems to be an o b l i g a t o r y one, except for single spe-
constitutae duabus ordinibus (raro 3) foraminarum. cimens a m o n g a population w h e r e it might b e appa-
In valvarum interiore et externo, frons cum brevi- rently absent, rudimentary o r not to discern without
bus rimis in axialis areae extremitate. Nulla rimopor- difficulty.
tida nulla porum apicalis area. Pleura cum septis uno
W i t h respect to septa, w e find in typical popula-
la'ere in frustulae interiore.
tions of Fragilaria pinnata and F. construens f. ven-
ter flat p r o j e c t i o n s of different lengths p r o t r u d i n g
Discussion into the v a l v a r plane o f the frustule (Fig. 39/40). On
the o t h e r end, linking spines which are a v e r y abun-
dant character {though not obligatory) in Fragilaria
A more or less extensive reduction of the branches seemed to be absent in Achnanthes : nevertheless,
o f the raphe is characteristic f o r a f e w species of L e Cohu & M a i l l a r d (1983) o b s e r v e d indentations in
Navicula which regularly d e v e l o p ribbon-like chains the s a m e p o s i t i o n as o n v a l v e s of Fragilaria in Ach-
as for instance, Navicula gallica. Thus, in s o m e nanthes lanceolata (Bréb.) Grun. v a r . lanceolatoides
cases, taxa of Navicula have o r i g i n a l l y been mista- (Sov.) R e i m .
ken for Fragilaria. T h e n e w taxa, h o w e v e r , share
m o r e similarities w i t h Fragilaria than Navicula, N o w w e found a specimen (Fig. 42) of Achnanthes
taking into c o n s i d e r a t i o n all c h a r a c t e r s in L M and exhibiting linking spines or adequate elements in the
S E M . In these cases neither artefacts nor teratolo- same position as on v a l v e s of Fragilaria o r s o m e t i -
g i r a l o r m o r p h o l o g i c a l i r r e g u l a r i t i e s play a role. mes of Navicula or, v e r y rarely o f Pinnularia. T h e
Actually w e have no p o s s i b i l i t i e s to p r o v e , w h e t h e r specific t a x o n o m i c identification is difficult. P r o -
the raphe-like slits can function f o r m o v i n g o r not. bably it b e l o n g s into the very problematic g r o u p o f
O t h e r w i s e , this information is also missing in many p o p u l a t i o n s a r o u n d Achnanthes marginulata
o t h e r cases o f taxa w i t h unquestionable relations-
Grunow.
hin to typical R a p h i d i n e a e as N a v i c u l a c e a e o r Ach-
nanthaceae. If w e w o u l d h a v e found o n l y one popu- T h e general conclusion is that the occurence o f
la' ion, in particular that wi th the a s y m m e t r i c arran- u n c o m m o n (unknown) c o m b i n a t i o n s of c h a r a c t e r s
gement of septa on the connecting band, the idea o f in D i a t o m genera should induce a re-examination o f
an independent new genus o r even a new family recent definitions even of higher taxa.
w o u l d be more plausible. H o w e v e r , the populations
of the K e r g u e l e n s h o w little d i f f e r e n c e f r o m s o m e
infraspecific taxa of Fragilaria construens, except
Acknowledgements
the « apically e l o n g a t e d terminal p o r e s » . T h i s fact
s u D p o r t s m o r e the idea that F r a g i l a r i o i d e a e o r even
Fragilaria possess m o r e m o r p h o l o g i c capabilities W e gratefully acknowledge the help rendered us by Dr. Kurt
Krammer and Mrs Ute Rumbrich. The electron microscopic work
than anticipated by t a x o n o m i s t s up to recently. was carried out by Mr. Manfred Ruppel (Bot. Inst. Frankfurt), R.
Thus, Simonsen ( 1974, 1979) pointed to Araphidineae Le Cohu (Toulouse) and Dr. Kurt Krammer, Dusseldorf (Pl. Ill,
Fig. 1/2).
216 H . L A N G E - B E R T A L O T E T R. L E COHU (4)

Travaux cités. Krammer (K.) & Lange-Bertalot (H.). 1985. — Naviculaceae. Neue
und wenig bekannte Taxa, neue Kombinationen und Synonyme
sowie Bemerkungen zu einigen Gattungen. Bibl. Diatomologica
Andrews ( G . W . ) . 1981. — Achnanthes linkei and the originin of
(in press).
monoraphid Diatoms. Bacillaria, 4 : 29-40.
Ross (R.), Cox (E.J.), Karayeva (N.I.), Mann (D.G.), Paddock (T.B.B.),
Anonymous. 1975. — Proposals for a standardization of diatom ter­
Simonsen (R.) & Sims (P.A.). 1979. — An amended terminology
minology and diagnoses. Nova Hedwigia, suppl. 53 : 323-354.
for the siliceous components of the diatom cell. Nova Hedwi­
Archibald (R.E.M.). 1983. — The Diatoms of the Sundays and Great
gia, suppl. 64 : 513-533.
Fish Rivers in the Eastern C a p Province of South Africa. Bibl.
Simonsen (R.). 1974. — The diatom plankton of the Indian Ocean
Diatomologica, 1 : 1-432.
Expedition of R/V Meteor 1964-1965. Meteor Forsch. Ergeb-
H a w o r t h (E.). 1975. — A Scanning electron microscope study of
nisse, ser. D, 19 ; 1-107.
some different frustule forms of the genus Fragilaria found in
Simonsen (R.). 1979. — The diatom system : ideas on phylogeny.
Scottish Late Glacial sediments. Brit, phycol, /., 10 : 73-80.
Bacillaria, 2 : 9-71.
Le Cohu (R.) & Maillard (R.). 1983. — Les diatomées monoraphi-
dées des îles Kerguelen. Annh Limnol., 19 (3) : 143-167.
(5* R A P H E L I K E V E S T I G E S I N T H E P E N N A T E DIATOM S U B O R D E R A R A P H I D I N E A ? 217

11 12 13 14 15 16 17

Plate I : (Fig. 1/17 L M x 1 500 ; Fig. 18/23 SEM x 11000, x 7 000, x 7000, x 6 000, x 6 500, X 7000). Scale LM : 10 m , M

S E M : 1 ^m.
F i g . 1/3 : Fragilaria maillardii n o v . s p e c . ; 4/7 : Fragilaria elliptica S c h u m a n n { L i n k i n g spines discernible directy o n the
p u n c t a t e s t r i a e ) ; 8/10a : Fragilaria construens f. venter ( E h r e n b e r g ) H u s t e d t ; 10b/12 : Fragilaria pinnata E h r e n b e r g
(12 : v a l v o - c o p u l a w i t h f a i n t l y d i s c e r n i b l e s e p t a - l i k e a p p e n d i c e s ) ; 13/17 : Fragilaria loetschertii n o v . s p e c , ; 18/23 : Fra­
gilaria maillardii n o v . s p e c . (18/20 : v a l v e o u t s i d e w i t h l i n k i n g s p i n e s o n t r a n s a p i c a l c o s t a e a n d w i t h e l o n g a t e d t e r m i ­
nal p o r e s ; 21/23 : v a l v e i n s i d e w i t h s o m e w h a t m o r e e l o n g a t e d r a p h e - l i k e v e s t i g e s ) .
 218 H. L A N G E - B E R T A L O T ET R. L E COHU (6)

P l a t e I I : ( S E M X 5 000, c a . X 6 600, c a . X 11 000, c a . X 7 00O, c a . X 7 000).

F i g . 24 : Fragilaria maillardii n o v . s p e c , ( v a l v e i n s i d e ) ; 25/26 : Fragilaria construens f. venter (25 : v a l v e o u t s i d e w i t h

l i n k i n g s p i n e s a r r a n g e d o n t r a n s a p i c a l c o s t a e a n d s m a l l a p i c a l p o r e f i e l d ; 26 : v a l v e i n s i d e w i t h c l o s i n g s t r u c t u r e s
o n u n i f o r m l y l a r g e a r e o l a e ) ; 27/29 : Fragilaria elliptica (27 : v a l v e o u t s i d e w i t h l i n k i n g s p i n e s c h a r a c t e r i s t i c a l l y a r r a n ­
g e d o n m a r g i n a l i n t e r c o s t a e ; 28/29 : v a l v e i n s i d e w i t h closing s t r u c t u r e s o n a r e o l a e w i t h d e c r e a s i n g size t o w a r d s t h e
axial area).
(71 RAPHE LIKE VESTIGES I N T H E P E N N A T E DIATOM SUBORDER ARAPHIDINEAE ? 219

I'lme I I I : ( S E M x 10 000, x 24 000, ca. x 15 000, c a . x 5 000, c a . x 13 000).

Fig. 30/34 : Fragilaria loetschertii n o v . s p e c . (30 : a d j a c e n t f r u s t u l e s in g i r d l e v i e w w i t h g i r d l e b a n d , linking s p i n e s o n
t r a n s a p i c a l c o s t a e , d o u b l e r o w s o f f o r a m i n a , t e r m i n a l r a p h e - l i k e slits p r e s e n t , r i m o p o r t u l a a n d special a p i c a l p o r e
fields a b s e n t ; 31 : e n l a r g e d a p i c a l s t r u c t u r e s ; 32734 : valves inside w i t h t e r m i n a l slits a n d unilateral asymmetric septa-like
elements respectively).
220 H. LANGE-BERTALOT ET R. L E C O H U (8)

Plate IV : (SEM ca. x 4 800. ca. x 11 000, ca. x 6 000, ca. x 11 000, ca. x 8 000, ca. x 9 000, ca. x 4 000, ca. x 5 000).
Fig. 35/39 : Fragilaria pinnala (35 : « Opephora-martyï »-form with comparatively wide valve dimensions and linking
spines on transapical costae, the fragment right below represents the same population as figure 38 : 36 : comparati-
vely small pinnata-form with grill-shaped foramina ; 37 : other pinnata-population with grid-like foramina and linking
spines on intercostae comparable with Opephora pacifica ; 38 : other population with linking spines on transapical
costae ; 39 : frustule fragment, valvocopula above with septalike appendices juxtaposed to the transapical costae) ; 40 :
presumable Fragilaria construens f. venter (generally same structures as in figure 39) ; 41 : Achnanthes aff. marginu-
lata (normal rapheless valve without linking-spines) ; 42 : Achnanthes aff. marginulata (rapheless valve with marginal
linking spine-like elements).