N. Jb . Ge o l. P aläon t . M h .

1996, H . 5

293- 308

St u t t gar t , M ai 1996

Remarks on the gen era Striatoporella an d “Riphaeolites”
(Devon ian Tabulata)
By Kl e m e n s O e k e n t o r p an d T o n g- D z u y T h a n h
W it h 4 figu r es in t he t ext
e k e n t o r p , K. & T o n g - D z u y T h a n h (1996): Re m ar ks on t h e ge n er a St riatoporella
an d “R iph aeolites” (D e v o n ian Tab u lat a). - N . Jb . Ge o l. P aläon t . M h ., 1996 (5):
2 9 3 - 3 0 8 ; St u t t gar t .

O

A b st r a c t : Re st u d y o f M esofav osites schouppei P o t t h a s t & O e k e n t o r p 1988
(Em sia n - Eife lian b o u n d a r y in SE M o r r o cc o ) an d o f Pach y fav osites pseudoseptatus
O e k e n t o r p 1975 (Em sian o f A st u r i a s/ N Sp ain ) sh ow s t h at M . schouppei h as t o be
assign ed t o St riatoporella R u k h i n 1938 b e cau se o f t h e t yp ical gr ow t h fo r m , t he
p er ip h er al t h icken in gs o f t h e w alls an d t h e d e v e l o p m e n t o f sp in e s. P. pseudosep­
tatus h as b een assign e d t o Pach y fav osites on t h e b asis o f its st r ikin gly t h ick walls.
D iage n e t ic p r o ce sse s m ay r esu lt in se con d ar y m icr ost r u ct u r e s, for e xam p le t he
ar r an ge m e n t o f car b o n at e fiber s. Th er e for e , wall t h ickn ess is u se le ss as a
d iagn ost ic feat u r e. In t h is view, P. pseudoseptatus co r r e sp o n d s ver y well w it h ot h er
r ep r ese n t at ives o f t h e ge n u s Riph aeolites.
Z u sa m m e n f a ssu n g : N e u u n t e r su ch u n ge n v o n Ge n e r a t ab u lat e r Kor alle n u. a. in
H in b lic k a u f m öglich e diage n et isch e M ik r ost r u k t u r - Üb e r p r ägu n ge n er gab en
n eu e Er k en n t n isse ü b e r die sy st em at isch e Z u o r d n u n g v o n M esofav osites schouppei

P o t t h a s t & O e k e n t o r p 1988 au s d e m Gr en zb e r e ich Em siu m - Eife liu m SE
M ar o k k o s u n d v o n Pach y fav osites pseudoseptatus O e k e n t o r p 1975 au s de m
Em siu m v o n A st u r i e n / N Sp an ie n . - Au fgr u n d der ch ar akt e r ist isch en W u ch sfor m ,
der Z u n ah m e der W an d ver d ick u n ge n zu r P er iph er ie h in u n d der Au sb ild u n g von
D o r n e n w ir d M . schouppei der Gat t u n g St riatoporella R u k h i n 1938 zu gesch r ie b en .
- Die au fgr u n d der star k ver d ick t en W än d e b ish er der G a t t u n g Pach y fav osites
zu ge w ie sen e Ar t P. pseudoseptatus erweist sich als Vert r et er v o n R iph aeolites Y a n e t
1955. Die n eu er lich e U n t e r su ch u n g er gab , daß die W an d ve r d ick u n ge n das
Resu lt at
fr ü h d iage n e t isch e r
U m k r ist allisat io n sp r o z e sse
d ar st ellen .
Ke n n ­
ze ich n e n d fü r die ab io ge n e N at u r sin d Kr ist allver gr ư ß er u n gen m it e in h er ge­
h en d er st är ker er A u fh e llu n g u n d die Au sb ild u n g n eu er , u n r egelm äß iger , fäch er ­
för m iger Kr i st allan or d n u n ge n . Diese k ö n n e n au s d e m p r im är e n Sk elet t h er au s­
w ach sen u n d d am it Sk elet t ver d ick u n ge n vor t äu sch e n . Es wird h ier d an n zw ar der
ar ch it ek t on isch e Rah m e n des Skelet t s w eit ge h en d b e ib e h alt e n , je d och in sein en
M aß e n u n d in se in e m m ik r ost r u k t u r e llen Fein b au völlig ü b er p r ägt . D am it
en t fallt die An w e n d u n gsm ö glich k e it der M ik r o st r u k t u r u n d der W an d ve r ­
di ck u n gen fü r sy st em at isch e Be lan ge . Au fgr u n d de r ü b r ige n M e r k m ale er gab sich
so m it die Z u o r d n u n g des sp an isch en M at e r ials zu Riph aeolites.

0 0 2 8 - 3 6 3 0 / 9 6 / 1 9 9 6 - 0 2 9 3 $ 4.00
1 9 9 6 E . Sc h w e iz e r b a r t ’sc h e V e r la g sb u c h h a n d lu n g , D - 7 0 1 7 6 St u t t g a r t


294

Kle m e n s O e k e n t or p an d T o n g- D zu y Th an h


I n t r o d u c t io n
D u r in g a t h r ee m o n t h st ay at th e For sch u n gsst e lle für Kor alle n p aläozo ologie , Un iver sit y M ü n st er / W est f., Ger m an y, th e ju n io r au t h or d isco ­
ver ed som e in t e r e st in g feat u r es in t ab u lat e cor als st or ed in th e colle c­
t ion s o f th e For sch u n gsst elle. Th ese con cer n a sp ecies o f Striatoporella
p r eviou sly d escr ib ed as “M esofav osites” schouppei P o t t h a s t & O e k e n t o r p
(1988) fr om t h e u p p e r Low er D e vo n ian o f M o r r occo as well as the
e n igm at ic gen u s “R iph aeolites”, d escr ib ed b y Ya n e t (1955). To th e latt er
on e we like t o assign t h e species “Pach y fav osites” pseudoseptatus
O e k e n t o r p 1975, fr om th e Em sian o f Ast u r ias, Nor t h er n Sp ain . Rest u dy
o f th e Sp an ish m at er ial sh ow ed sign ifican t d iagen et ic m icr ost r u ct u r es,
w h ich call for a cr it ical d iscu ssio n o f t h e gen u s “R iph aeolites”. Th e ju n io r
au t h or w ish es to expr ess h is gr at it u d e t o th e Deu t sch e Ak ad em isch e
A u st au sch d ie n st (D A A D ) for fin an cin g h is r esear ch st ay in Ge r m an y an d
t o t h e Viet n am ese P r ogr am for Fu n d am e n t al Resear ch in th e Field o f
N at u r al Scie n ce for t h eir kin d su p p or t . Ad d it io n ally , h e t h an k fu lly
r em em b er s t h e For sch u n gsst e lle für Ko r alle n p aläo zo o lo gie o f the
W est fälisch e W ilh elm s- Un iver sit ät , Mü n st er .

Fam ily Fa v o sit id a e D a n a , 1846
Ge n u s Striatoporella R u

e.p .
e.p .
e.p .

e.p .
e.p.
e.p .


e.p .

1938
1869
1955
1959
1962
1963
1964
1967
1968
1969
1971
1981
1988
1988
1990

k h in

,

1938

St riatoporella R u k h i n , p. 62
St riatoporella - D u b a t o l o v , p. 80
T h am n opora - S o k o l o v , p. 175
Fav osites - H a m a d a , p. 201
Th am n opora - S o k o l o v , 229
Fav osites - S o k o l o v & T e s a k o v , 128

R udak ites L e l e s h u s , p. 46
Squ am eopora P r e o b r a z h e n s k i y , p. 8
Fav osites - C h e k h o v i c h , p. 43
Fav osites - D u b a t o l o v , p. 53
Th am n opora - S o k o l o v in O r l o v , p. 229
St riatoporella - H i l l in M o o r e , R o b i s o n & T e i c
M esofav osites - P o t t h a s t & O e k e n t o r p , p. 76
Squ am eopora - T o n g - D z u y , T h a n h et al., p. 61
Fav osites - D u b a t o l o v , p. 56

h er t

,

p. 546

T y p e s p e c i e s : St riatoporella m ultiporifera R u k h i n . - Th e h olot y p e
o f the
sp ecies is lost . A n e o t y p e h as b e en ch o se n b y V. N. D u b a t o l o v 1969 (Page 80;
№ 109, coll. 3 3 7; Low er D e v o n ia n ; Ko ly m a b asin , N o r t h e a st Sib ir ia. Co n se r v e d
at t h e In st it u t e o f G e o l o gy an d G e o p h y sic s N ov o sib ir sk , Ru ssia).
D i a g n o s i s : Br an ch in g an d n o d u l o se c or allu m wit h coar se cy lin dr ical
ou t gr ow in g. Cor allit e p o ly go n al in t h e t r an sver se se ct io n , dir ect ed lo n gi t u d i ­
n ally in t h e axial zo n e an d r egu lar ly cu r v in g ou t w ar d t o t h e su r face o f t h e


Rem ar k s on th e gen er a St riatoporella an d “R iph aeolites“

295


cor allu m . W alls sligh t ly t h icke n ed t o t h e p e r ip h e r al zo n e . M u r al p or e s m ay be
sit u at ed t o t h e cor n er o f t h e cor allit e s. Se p t al sp in e s, r ep r e sen t ed b y sh o r t , coar se
sp in es an d sq u am u lae , are ir r egu lar ly d e v e lo p e d in t h e axial an d m or e fr eq u en t
in t h e p er ip h er al zo n e . Tab u lae m ay be t h icken ed in t h e pe r ip h e r y o f t he
co r allu m .

Rem ar k s
1. Br an ch in g an d n o d u lo se cor alla o f favosit id s wit h cylin dr ical
ou t gr ow in gs are w idesp r ead in th e U p p e r Silu r ian an d p ar t icu lar ly in th e
Low er D e vo n ian . Alt h o u gh m o st o f t h em h ave b een d escr ib ed as sp ecies
o f t h e gen u s Fav osites, t h eir t axo n o m ic p o sit io n s at t r act t h e at t en t io n o f
p alae on t ologist s. It seem s t h at t h ese for m s sh ow a t r an sit ion fr om
b r an ch in g favosit id s to p ach y p or id s ( T h am n opora) b u t it is d ifficu lt to
fin d d iagn ost ic feat ur es for se p ar at in g Fav osites-like Th am n opora fr om
Thamnopora-Y ike. Fav osites sp ecies ( P r e o b r a z h e n s k i y 1967, H a m a d a 1959)
(com p ar e L e c o m p t e 1936, 1939).
2. De scr ib in g th e gen u s Striatoporella wit h a n o d u lose b r an ch in g
cor allu m L. B. R u k h i n n ot ed its affin it ies wit h favosit id s b u t n ot pach yp or id s. Th e illu st r at ion s in R u k h i n ’ s d escr ip t ion s (1938) are n ot clear
en o u gh an d , u n for t u n at e ly, all h is m at er ial was lost du r in g th e Se co n d
W or ld War. As a r esu lt , t h e gen u s Striatoporella h as b een for got t en for
lon g t im e or was r egar d ed as a ju n io r sy n o n y m o f Th am n opora ( S o k o l o v
1955, 1962).
3. Em p h asiz in g t h e pr ese n ce o f sq u am u lae in t h e per iph er al par t in a
large n u m b e r o f br an ch in g favosit id s в. V. P r e o b r a z h e n s k i y (1967)
est ab lish ed th e gen u s Squam eopora an d p oin t e d ou t its affin it ies t o th e
gen u s Squam eofav osites a n d Em m on sia. Regr et t ably, a s D. H i l l r em ar ked,
t h e pr esen ce o f sq u am u lae in Fav osites hidensis (ch ose n as t ype species o f
Squam eopora by в. V. P r e o b r a z h e n s k i y ) is n ot m e n t io n e d in th e p u b li­
cat ion o f T. H a m a d a (1959). So d iagn ost ic ch ar act er s o f Squam eopora are
still m issin g. In an y case, favosit id cor als wit h cylin d r ical ou t gr ow in gs

an d t h ick en ed walls in th e p er iph er al par t s o f cor allit es (with or w it h ou t
sq u am u lae ) can be r egar d ed as t h e r epr esen t at ives o f a gen u s d iffer en t
fr om all ot h e r gen er a o f th e fam ily Favosit id ae.
4. O n t h e basis o f ab u n d a n t t ab u lat e sp e cim e n s fr om R u k h i n ’ s
(1938) localit ies in t h e Low er D e vo n ian o f th e Ko ly m a River (N or t h east
Ru ssia), V. N . D u b a t o l o v (1969) h as p r oved th e pr ior it y o f t h e valid
n am e Striatoporella. A cco r d in g t o t h e d et ailed d escr ip t ion o f two sp ecies,
in clu d in g th e t ype- sp ecies Striatoporella m ultiporifera in D u b a t o l o v ’ s
m o n o gr a p h (1969: 8 0 - 82, PI. XLVI, Fig. 1- 4, PI. XLVII, Fig. 1- 6),
essen t ial m o r p h o lo gical t r ait s o f t h is gen u s can be ch ar act e r ized as
follow s:


296

Kle m e n s O e k e n t or p an d T o n g- D zu y Th an h

- N o d u lo se wit h cylin d r ical ou t gr ow in gs an d b r an ch in g cor allu m .
- Cor allit e walls t h icken t ow ar d s th e per iph er al zon e.
- Ir r egular d e ve lop m e n t o f sept al sp in es in t h e axial zon e an d
asso ciat ion o f spin es an d sq u am u la in th e per iph er al zon e.
- Lar ge m u r al por es, so m e t im e s sit u at e d at th e c o m e r o f t h e cor allit es.
Th ese feat ur es were n ot p oin t e d ou t in t h e d escr ip t io n o f D u b a t o l o v ,
b u t on e o f us (T.D.T.) h ad t h e o p p or t u n it y to ob ser ve it in t h in sect ion s
o f th e t ype- sp ecies st or ed in th e In st it u t e o f G e o lo gy an d Ge o p h y sics in
Novosib ir sk . It can also be seen in D u b a t o l o v ’ s illu st r at ion s (1969: PI.
XLVI an d XLVII).
5.
C o m p o sit io n an d d ist r ib u t ion o f Striatoporella. Repr e se n t at ives o f
Striatoporella are w idesp r ead fr om t h e U p p e r Silu r ian t o th e M id d le

D e vo n ian , par t icu lar ly in th e Low er D e vo n ian . Apar t fr om t yp ical
sp ecies d escr ib ed fr om Low er D e v o n ian o f N o r t h e ast Ru ssia, the
follow in g br an ch in g favosit id s sh ou ld be assign ed t o Striatoporella:
- A gr eat n u m b e r o f d escr ib ed br an ch in g favosit id s, in clu d in g th e
gr ou p s o f Fav osites socialis S o k o l o v & T e s a k o v , F. p olaris C h e k h o v i c h , F.
hidensis ( K a m e i ), F. k ozlow sk y i ( S o k o l o v ) a. o.
- D e s c r i b e d s p e c i e s o f Squam eopora, for e x a m p l e
m agn issim a,
vuk h uci ( T o n g - D z u y T h a n h e t a l . 1 9 8 8 ) a . o .

s.

s.

Fig. 1. St riatoporella schouppei ( P o t t h a s t & O e k e n t o r p , 1988). Em sian - Eifelian
b o u n d a r y ; H a m a r Lagh d ad / Tafilalt , SE M o r o c c o .
1: Cr o ss se ct ion o f a cy lin dr ical c o l o n y wit h t h in w alls in axial par t , bu t
t h icke n ed on e s at t h e per ip h er y. N u m e r o u s sp in es an d sq u a m u lae are d e ve lop e d .
- Se ct io n G I M - M A 3 0 —1; X1 .6 .
2: Lo n git u d in al se ct ion o f a cy lin dr ical ou t gr ow t h wit h walls t h ick en in g t ow ar ds
t h e per ip h er y. Pores an d n u m e r o u s sp in es ar e well d e v e lo p e d , cr oss se ct ion s o f
t h e lat t er oft e n ap p e ar in g as dar k d ot s. - Se ct ion G I M - M A 8 8 - 1 ; X 1.6.
3: Cr o ss se ct ion o f t he cy lin dr ical ou t gr ow t h (Fig. 1.2) wit h walls t h ick en in g
t ow ar ds t h e pe r ip h e r y o f t h e colon y. - Se ct io n G I M - M A 8 8 - 1 ; X2 .3 .
4: Lo n git u d in al se ct ion sh ow in g p ar t o f a wall (left ) an d on e o f t h e rare
sq u am u lae . - Se ct ion G I M - M A 8 8 - 1 ; X2 0 .
5: Lo n git u d in al se ct ion wit h a p or e an d st r on g sp in e s, dir ect ed u p w ar d s. Se ct ion G I M - M A 8 9 - 2 ; X2 0 .
6: Lo n git u d in al se ct ion wit h wall p o r e s an d p or e s sit u at ed in t h e cor n er o f the
wall (“M esofav osites”- z p p e aran ce ). - Se ct ion G I M - M A 9 4 - 1 ; X2 0 .
7: Det ail o f a cr oss se ct ion sh ow in g m od e r at e ly t h icken ed walls wit h dist in ct

b o u n d a r y t o t h e cor allit e cavit y. Th e dar k m id d le lin e is still pr eser ved. - Se ct ion
G I M - M A 9 4 - 1 ; X2 0 .
8: Det ail o f a r e cr yst allize d wall. Th e or igin al b o u n d a r y o f t h e wall is lost du e
t o gr ain gr ow t h , p r o d u c in g a saw- like b or d e r wit h ir r egu lar in d e n t at ion s. Th e
lat t er on e s r ep r ese n t scale n o h e d r o n s du e t o cr ystal gr ow t h . - Se ct ion G I M - M A
8 8 - 1 ; X4 8 .


Rem ar k s on th e gen er a St riatoporella an d “R iph aeolites”

297

- Th e gen u s R udak ites L e l e s h u s (1964) fr om Low er D e v o n ian o f Cen t r al
Asia m igh t be also r egar d ed as a you n ge r sy n o n y m o f Striatoporella.
Ju d gin g fr om L e l e s h u s ’ s d escr ip t io n (1964) ap ar t fr om th e b r an ch in g
sh ap e o f th e cor allu m th e d ist in gu ish in g t rait o f R udak ites is t h e p oor
d e ve lop m e n t o f sep t a, w h ich occu r on ly in t h e per ip h er y o f th e cor allu m
in th e for m o f ob t u se sp in es. Th e lat t er are sim ilar ly ob ser ved in th e t ype
sp ecies o f Striatoporella ( D u b a t o l o v 1969: 80- 81).


298

Kle m e n s O e k e n t o r p an d T o n g- D zu y Th an h

- M esofav osites schouppei d escr ib ed by P o t t h a s t & O e k e n t o r p (1988)
fr om th e Em sian - Eife lian b o u n d ar y st r at a o f SE M o r o c c o r epr esen t s a
ver y in t er est in g species o f Striatoporella.

N o t e s o n St riat op orella sch ouppei

New ob se r vat ion s in well pr eser ved p olyp ar ies st or ed in th e
For sch u n gsst elle
fü r Ko r alle n p aläo zo o lo gie , Un iver sit y M ü n st er
(Ger m an y), sh ow t h at th e m ain m o r p h o lo gical elem en t s o f t h ese cor als
co r r e sp on d to t h e d iagn osis o f Striatoporella. Th ese are (1) br an ch in g an d
n o d u lo se cor alla w ith cylin d r ical ou t gr ow in gs (Fig. 1.1- 1.3), (2)
t h icke n ed walls t ow ar d th e sur face o f t h e cor allu m (Fig. 1.1- 1.3;
com p ar e Fig. 1.7) an d (3) rare sq u am u lae (Fig. 1.4) associat ed with
coar se sp in es, especially in th e per iph er al zo n e s o f th e cor allu m (Fig.
1.2, 1.5).

Fig.

2. R iph aeolites pseudoseptatus ( O e k e n t o r p 1975). Fe r r on es Cr in o id e a n
l im e st on e , Em sia n ; En se n ad a de Ban u gu e s, Ast u r ias, N or t h e r n Sp ain .

1: Lo n git u d in al se ct ion sh ow in g m o d e r at e ly t h icken ed walls an d zon alit y . Se ct io n G I M - B2 - 3 2 2 .2 ; X2 .2 .
2: Lo n git u d in al se ct ion t h r ou gh an e n cr u st in g colon y . Th e cor allit e s gr ew alo n g
t h e su b st r at e b efor e b e n d in g u pw ar d s. - Se ct io n G I M - B2 - 1 6 .1 ; X1 .3 .
3: In t h e b asal par t o f t he se ct ion cor allit es ar e t h in - w alled b u t b e co m e
ext r em ely t h ick on t op . - Se ct io n G I M - B2 - 3 2 4 .1 ; X2 .2 .
4: Cr o ss se ct ion wit h m o st cor allit es h avin g ext r em ely t h icke n ed w alls. Part ly,
d iagen et ic car b o n at e fills t h e w h ole cavit y. - Se ct ion G I M - B2 - 3 2 4 .2 ; X2 .2 .
5: Lo n git u d in al se ct ion sh ow in g t h e zo n alit y o f wall t h ick en in gs; t h e cor allit es
o f t h e b asal par t s r em ain t h in - w alled . - Se ct io n G I M - B2 - 3 2 7 .4 ; X2 .4 .
6: Det ail o f a lon git u d in al se ct ion sh ow in g t h e spir al t u be o f a c o m m e n sa l w or m ,
H elicosalpin x ast u rian a O e k e n t o r p 1969. - Se ct io n G I M - B2 - 3 2 2 .2 ; X4 5 .
7: Det ail o f a cr oss se ct ion . Th e p o l y go n al cor allit e s are lim it ed b y t h ick walls o f
k n o t t y ap p e ar an ce . Th e se “p a r at r ab e cu la” are cau se d b y di age n esis. Th e d ot s in
t h e lar ge cor allit e s (low er r igh t ) are cr oss- cu t - spin es. - Se ct ion G I M - B2 - 1 6 .3 ;

X 45.
8: Det ail o f a lon git u d in al se ct ion . Th e w alls in a zo n e o f t h icke n in g sh ow
calcit ic fiber s in fan - like ar r an ge m en t . In t h e u p p e r par t t h e walls are t h in an d
m ay r ep r e sen t t h e or igin al t h ickn e ss (co m p ar e wit h Fig. 4.5). - Se ct ion G I M - B2 327.4; X 45.
9: En lar ged cr oss se ct ion sh ow in g diffe r en t wall st r u ct u r es an d t h ickn esses. In
t he low er p ar t , so m e o f t h e t h in walls st ill sh o w t h e dar k m id d le lin e. In t he
u p p e r par t t h e walls are t h icken ed b y coar ser an d lon ge r fiber s ar r an ged in
fan - syst em s (co m p . Fig. 4.4). Th e lat t er in d icat e s a di age n et ic or igin , for w h ich
t h e ligh t en ed co lo r o f t h e car b o n at e m at er ial is an ad d it ion al e vid en ce. - Se ct ion
G I M - B2 - 3 2 4 .1 ; X27.


Rem ar k s on th e gen er a St riatoporella an d “R iph aeolites*

299

D ist in gu ish in g feat ur es o f t h is species ar e: 1. Th e p r ofu se
d e ve lop m e n t o f sept al spin es (Fig. 1.1- 1.2). In t h e axial zo n e , t h ey are
n u m e r ou s an d r ep r esen t ed as lon g an d acu t e elem en t s (Fig. 1.5). In the
p er iph er al zon e , som e sq u am u lae (Fig. 1.4) an d squ am u lae- lik e spin es
occu r in asso ciat ion wit h ob t u se sp in es. 2. In ad d it io n to th e t ypical
m u r al p or es, m an y por e s are sit u at ed at th e edge o f th e walls (Fig. 1.5,
Fig. 1.6), in th e cor n er o f cor allit es, e. g. in M esofav osites.


302

Kle m e n s O e k e n t or p an d T o n g- D zu y Th an h

2. In cr oss se ct ion , th e t h in (e.g. Favosites-M k z) walls n or m ally sh ow a

clear dar k m id d le lin e an d t wo fr am in g zo n e s o f dar k grey, fin e- gr ain ed
car b on at e (Fig. 4.4). Car b o n at e fiber s h ave r arely been ob ser ved . Th e
b or d er t o t h e viscer al ch am ber s is u su ally p r on o u n ce d . Som e t im e s th e
m id d le lin e h as been r ecr yst allized t o for m a b r igh t e n ed zon e with
in d ist in ct b or d er s. W all t h ickn esses in t h e Fav osites-like par t s r an ge
bet w een 0.14 to 0.18 m m .
3. In Cleistopora-like par t s o f th e co lo n y t h e walls are p ar t ially
t h ick en ed, r an gin g fr om 0.1 t o 0.33 m m , an d are n or m ally ver y ligh t an d
coar sely fib r ou s. Th e fiber s are ar r an ged in fan s o f ir r egular sh ap e an d
ext en sion w it h ou t an y u n ifor m it y (Figs. 2.8, 2.9, 3 .1 - 5 , 4.1- 6). W it h in
th e fan s a fin e lam ellat ion o f t h e fiber s can be n ot iced . Th is lam ellat ion
is du e t o th e d iagen et ic gr ow t h o f cr yst als. Th e sam e feat ur e h as been
ob ser ved an d d escr ib ed by O e k e n t o r p (1989) in Pleist ocen e scler act in ian s fr om th e Sin ai P en in su la, Egypt . Th e b o u n d ar y t o th e viscer al
ch am b er s gen er ally var iab le b u t m ost ly saw- like in sect ion , e. g. the
in d e n t at ion s, are cau sed by t h e m o d e o f gr ow t h o f th e cr yst als r esp. the
fan s (Figs. 3.3, 3.4, 4 .1 - 3 , 4 .5- 6). Th ese t h ick en in gs are d e ve lop e d in
m or e or less r egu lar zo n e s an d on the sam e level t h r ou gh ou t th e colon y.
Ad d it io n ally , t h ey alt er n at e w it h dar ker zon e s, w h ich are oft en t h in n er
t h an th e ligh t er on es (Fig. 3.2).

Fig.

3 . R iph aeolites pseudoseptatus ( O e k e n t o r p
1 9 7 5 ) . Fe r r on es Cr in o id e a n
l im e st on e , Em sia n ; En se n ad a de Ban u gu e s, Ast u r ias, N or t h er n Sp ain .

1: Lo n git u d in al se ct ion t h r ou gh a zo n e o f t h icke n ed wall. Th e fiber s are ar r an ge d
in fan - syst e m s. - Se ct io n G I M - B2 - 3 2 2 .2 ; X2 0 .
2: Var iable t h ick en in g o f t he w alls in a lo n git u d in al se ct ion . Part ly t h e walls are
ext r e m ely sw ollen b y di age n et ic fan - syst e m s. - Se ct ion G I M - B2 - 1 6 .1 ; X2 0 .

3: En lar ge d wall wit h sever al fan s o f coar se an d e lo n gat e d fibr es in lon git u d in al
se ct ion . Th e fan s are t h e so calle d “St ir n z o n e n ”, e.g. ar ch es o f r ecr yst allizat ion
fr o n t s, p r ovin g t h e di age n et ic n at u r e o f t h ese st r u ct u r es - (Fig. 4.2, b u t also Figs.
4.1, 4.3, 4.4, 4.5, 4.6). - Se ct ion G I M - B2 - 1 6 .5 ; X4 5 .
4: En lar ge d lon git u d in ally - cu t wall sh ow in g a very clear fib r ou s fan , w h ich h as
r e p laced t h e pr im ar y w all st r u ct u r e. M or e ove r , t h e fiber s grew b e y o n d t h e
lim it at ion o f t h e p r im ar y wall in t o t h e cavit y o f t h e cor allit e , givin g t h e wall a
clu b - sh ap e d ap p e ar an ce w it h an ir r egu lar b o r d e r (Co m p a r e Fig. 4.6). - Se ct ion
G I M - B2 - 1 6 .5 ; X 45.
5: A clu b - sh ap e d t h icken ed par t o f a wall b u ilt o f coar se an d e lo n gat e d fibr es,
b e co m e s m u ch t h in n e r an d again t h ick en ed . Th e t h in par t o b v io u sly r epr esen t s
t h e p r im ar y wall (Co m p a r e Fig. 4.5) - Se ct ion G I M - B2 - 1 6 .1 ; X4 5 .
6: Recr y st allized wall an d t ab u la. Th e in fillin g o f t h e cor allit e cavit y con t ain s
som e coar se cr y st als o f d o lo m it e , w h ich sh ow t h at di agen esis o f t h e skelet on
b e gan ver y ear ly, e. g. b e fo r e t h e cor als were e m b e d d e d in t o t h e se d im e n t s, u n d e r
t h e in flu e n ce o f m e t e or ic wat er. - Se ct ion G I M - B2 - 1 6 .6 ; X4 5 .


Rem ar k s o n t h e gen er a St riatoporella an d “R iph aeolites”

303

4.
Cr o ss sect ion s also sh ow p ecu liar it ie s o f th e wall st r u ct u r e, even
t h ou gh det ails (such as th e fan syst em s) are lackin g. Th e t h icke n ed walls
look par t ially n o d u lo se , r esu lt in g in a so- called “p ar at r ab ecu lar ” m icr o ­
st r uct ur e ( T o n g - D z u y 1966, 1988 an d Fig. 2.7) or in a “coe n e n ch y m e ”like ap p ear an ce. Th is r esem bles sim ilar st r u ct ur es in sp e cim e n s o f th e
Silu r ian gen u s Thecia, e. g. Thecia (T h eda) ex p atiata ( L o n s d a l e 1839)



304

Kle m e n s O e k e n t o r p an d T o n g- D zu y Th an h

Fig. 4. R iph aeolites pseudoseptatus ( O e k e n t o r p 1975). Fe r r on es Cr i n o i d e a n
lim e st o n e , Em sia n ; En se n ad a de Ban u gu e s, Ast u r ias, N o r t h e r n Sp ain . Dr aw in gs
o f wall st r u ct u r es t o d e m on st r at e m or e clear ly t h e d iffer en t ap p e ar an ce o f m i c r o ­
st r u ct u r es an d t h e var iab le t h ickn esses o f t h e w alls. Th e t h ickn ess is cau se d by
r e o r gan iz at io n o f t h e or igin al m icr o st r u ct u r e con t r olle d b y t h e p o ly p s. Th e
r e or gan isat ion r esu lt s in gr ain gr ow t h , b r igh t e n in g o f t h e car b o n at e an d fan -like
cr yst al gr ow t h o f t h e “r e b u ilt ”, or n ew aggr egat e s. D u r in g t h ese p r o ce sse s t he
fibe r s grew b e y o n d t h e lim it at ion o f t h e p r im ar y walls r esu lt in g in an ir r egu lar
b o r d e r an d ext r em e t h icken in g. Th e or igin al wall m icr o st r u ct u r e an d wall
t h ickn ess is r e t ain ed o n ly locally, for in st an ce in Fig. 4.5 (u p p e r par t ) or Fig. 4.4
(low er par t ). - Figu r e 4 n o t t o scale.


Rem ar k s on th e gen er a St riatoporella an d “R iph aeolites”
d e sc r ib e d
O

ek en t o r p

w h ere

th e

305

f i g u r e d b y N i c h o l s o n , 1879 o r A m s d e n , 1949 ( s.

1970: p i . 1, f i g . 2 a n d 3; c o m p a r e f u r t h e r p i s . ) . A s i n Thecia,

an d

d iage n e t ic

m ic r o st r u c t u r e s

h ave

been

m o d ifie d

by

th e

e x i st e n c e o f se p t a , si m i l a r c o n d i t i o n s , e. g. t h e d e v e l o p m e n t o f sp i n e s,
c a u se d a c o m p a r a b le m ic r o st r u c t u r e in

Riph aeolites.

In cr oss sect io n s, a gr an ular m icr ost r u ct u r e can be ob se r ved
d e p e n d in g on t h e d ir ect ion o f fiber s in t h e fan - syst em s in lon git u d in al
cut s. Th e m at er ial ap p ear s dar k gr ey in t h in walls b u t ligh t er in th e r ecr y­
st allized par t s (Figs. 3.2, 3.5). Th is colo r ch an ge is a clear evid en ce for a
d iagen et ic or igin . Th e m id d le lin e is oft en ob scu r e or par t ially
br ow n ish ; b u t a dar k m id d le lin e can be fo u n d , w h ere t h in walls h ap p en
to be pr eser ved .

Th ese feat u r es, e sp e cially th e ligh t an d coar se fib r o u s fan - sy st em s
co m b in e d w ith t h eir ir r egu lar it y an d th e ir r egu lar b o r d er in g to th e
viscer al ch am b er s, are clear ly th e r esu lt s o f p ost - m or t em d iagen et ic
p r o ce sse s. Th e ap p ear an ce o f m icr ost r u ct u r al d et ails (sh ap e o f th e fan s;
th e d im e n sio n s o f fib e r s; th e in cr eased t r an sp ar en cy o f th e car b o n at e ;
in d e n t at io n o f th e b o r d er s, et c.) cor r e sp o n d s to o b se r v at io n s in
P leist ocen e cor als fr om th e Sin ai P e n in su la/Egy p t ( O e k e n t o r p 1989)
an d P er m ian r u gose cor als fr om In d o n e sia ( O e k e n t o r p 1980, 1984).
M or eover - esp ecially in P erm ian cor als b u t also in t h ose fr om ot h er
p e r io d s - th e zo n alit y o f t h ick en in g is co n sp icu o u s. It h as t o be c o n si­
d er ed as a p r ogr essive ph ase- lik e “r e cr y st allisat io n ” d u r in g ear ly d iage ­
n esis in sh allow m ar in e en v ir on m en t s, r esu lt in g in so- called “Stirn fr o n t e n ” (S c h o u p p e & S t a c u l 1966; O e k e n t o r p 1980, 1984; P o t t h a s t
1992). Sh allow m ar in e c o n d it io n s an d in flu en ce o f m et eor ic w ater
d u r in g fo ssiliz a t io n is evid en ced by th e exist en ce o f d o lo m it e cr y st als in
th e cavit y in fillin gs as w ell as in cem en t s (Fig. 3.6).
Accor d in g t o t h ese ob ser vat ion s an d t h eir in t e r pr et at ion , t h e t h ick e­
n in gs are secon d ar y, d iagen et ic feat ur es r at h er t h an skelet al pr oper t ies
o f t axo n om ic sign ifican ce. Th er efor e, “Pach y fav osites” pseudoseptatus
O e k e n t o r p , 1975 can n o lon ger be con sid er ed as a r epr esen t at ive o f the
gen u s Pach y favosites, b u t r esem bles t h e gen u s Riph aeolites, w h ose wall
st r uct ur e is t h e sam e as in th e Sp an ish m at er ial.
A m o n g t h e few kn ow n species o f Riph aeolites, t h e on e n ear est t o the
t ype sp ecies (R . sok olov i) is t h e Em sian for m fr om No r t h er n Sp ain
d escr ib ed as Pach y favosites pseudoseptatus (O e k e n t o r p 1975: 6 8 - 7 2 ; PI.
XI I , Fig. 4; PI. XI I I , Fig. 4; PI. XIV, Fig. 2- 3). Th e sim ilar it y o f t h ese two
sp ecies is clear ly exp r e sse d in t h e cr u st ose, lam ellar sh ap e o f the
cor allu m , t h e m an n e r o f r ap id ly ch an gin g cor allit e sh ap e an d th e wall
st r uct u r e. Th eir sim ilar it y is also expr essed even in th e coexist en ce with
2 0 * N. Jb . Geol. P alaon t . Mh . 1996



306

Kle m e n s O e k e n t o r p an d T o n g- D zu y Th an h

specific sy m b io n t s in th e for m o f co m m e n sal w or m s. H ow ever , Riph aeolites pseudoseptatus st an d s ou t by its pecu liar sept al st r uct ur e. In th e
ad u lt st ages o f cor allit es, d isjoin t ed , r idge-like sept a are ob ser ved in
ad d it ion t o coar se, spin e- like se pt a ( O e k e n t o r p ) 1975: Pl. XI I , Fig. 4; PI.
XIV, Fig. 1). By t h ese d ist in gu ish in g m o r p h o lo gical elem en t s, R . pseu do­
septatus differ s fr om all k n ow n sp ecies o f Ripbaeolites. M or e over , the
Sp an ish sp ecies d iffer s fr om t h e t ype- sp ecies by th e sm all size o f th e
cor allit es an d th e t h in walls.
W h ile a det ailed d escr ip t ion an d illu st r at ion o f R ipbaeolites pseudosep­
tatus h as been p r ovid ed by O e k e n t o r p ( 1 9 7 5 ) , t h is p u b licat ion ad d s the
illu st r at ion o f som e im p or t an t m or p h ological elem en t s t h at im p r ove
th e d iagn osis o f t h e species.

Re fe r e n c e s
, V. N . (1969): Tab u ljat y i b io st r at igr afija n izh n e go D e v o n a Se ve r n ogo
Vost o k a SSSR . - In st it u t G e o lo gii i Ge ofizik i, 70: 179 p p , 67 P L; N o v o si ­
b ir sk (N au k a). [Tabu lat a an d Biost r at igr ap h y o f t h e Low er D e v o n ia n o f
N o r t h East e r n U SSR ]
D u b a t o l o v , V. N . & S p a s s k i y , N . Y. A. (1964): N e k ot or y e n ovy e ko r ally iz d e vo n a
So v ’et sk ogo So y u sa. - [In :] St r at igr afich eskiy i geogr afich esk iy o b z o r
d e von sk ik h k o r allo v SSSR : 112- 137, pi. 1 - 1 1 ; M o sk v a (Ak ad e m ia N a u k
SSSR ). [Som e n ew cor als fr om D e v o n ia n o f t h e Sov ie t U n io n ]
F l ü g e l , E. (1978): M ik r ofazielle U n t e r su c h u n gsm e t h o d e n vo n Kalk en . - 454 p p .,
68 figs., 57 t b ls., 33 p is.; Be r lin - H e id e lb er g- Ne w Yor k (Spr in ger ).
H a m a d a , T. (1959): O n t he t axo n o m ic p o si t io n o f Fav osites h iden sis an d its
D e v o n ia n age. - Ja p . J. Ge o l. G e o p h y s., 30: 2 0 1 - 2 1 3 , pi. 16.

H i l l , D. (1981): Tab u lat a. - [In :] T e i c h e r t , C . (Ed .): Tr eat ise on In ver t ebr at e
P ale on t o lo gy . F. Coe le n t e r at a, Su p p l. 1.2: 4 3 0 - 7 6 2 ; Bo u l d e r / C o l . Law r e n ce / Kan s.
L e l e s h u s , V. L . (1964): N o vy e d ev on sk iy r o d R udak ites (Tab u lat a) iz Tadzh ikist an a. - P ale on t . Z h u r n al, 1964 (4): 4 5 - 4 8 ; M o sk v a (Ak ad e m ia N a u k SSSR ).
[Ne w D e v o n ia n ge n u s R udak ites (Tab u lat a) fr o m Tadzh ik ist an ]
O e k e n t o r p , K l . (1969): Ko m m e n sa l i sm u s b ei Favosit id e n . - M ü n st er . For sch .
Ge o l. P aläon t ., 12: 16 5 - 2 1 6 , 10 A b b ., Taf. 1 3 - 1 6 ; Mü n st er .
- (1970): Re visio n des Ge n u s Th ecia E d w a r d s & H a i m e 1849 (Co e le n t e r at a,
Tab u lat a). - N . Jb . Ge o l. P aläon t . A b h ., 137 (1): 103- 173, 12 A b b .; St u t t gar t .
- (1975): Be sch r e ib u n g u n d Sy st e m at ik d e v on isch e r Favo sit id ae Ast u r ie n s
u n d Be o b ac h t u n ge n zu r Bioge ogr ap h ie n or d sp an isch e r Ko r alle n fau n e n . Mü n st er . For sch . Ge o l. P aläon t ., 37: 1- 12 9, 10 A b b ., 1 Tab ., 14 Taf.;
Mü n st er .

D

u ba t o l o v


Rem ar k s o n th e gen er a St riatoporella an d “R iph aeolites“

307

K l . (1980): Ar agon it u n d D ia ge n e se bei ju n gp a l äo z o i sch e n Kor allen .
O e k e n t o r p , K l . (Ed .): Fe st sch r ift P r ofe ssor Dr. A l e x a n d e r v o n
S c h o u p p e . - M ü n st e r . For sch . Ge ol. P aläo n t ., 5 2 : 1 19 - 2 39, 25 figs., 15 p ls.;
Mü n st er .
- (1984): Ar agon it e an d D ia ge n e sis in P er m ian Cor als. - [In :] O l i v e r , W. A.
jr. et al. (Ed s.): Recen t Ad van ce s in t h e P ale o b io lo gy an d G e o l o gy o f t h e
Cn id ar ia. (P r o ceed in gs o f t h e Fou r t h In t e r n at ion al Sy m p o si u m on Fossil
Cn id ar ia an d Ar ch ae ocy at h id s an d St r o m a t o p o r o id s). - P alae o n t ogr ap h ica
Am e r ican a, 5 4 : 2 8 2 - 2 9 3 , 12 figs., 2 p is.; It h aca, N.Y.

- (1989): Diage n e sis in cor als: Sy n t axial ce m e n t s as evide n ce for
p o st - m or t e m skelet al t h icke n in gs in P alae ozoic cor als. - M e m . Asso c.
Au st r al. P alae on t ., 8: 169- 177, 6 figs.; Br isb an e.
P o t t h a s t , I. (1992): Sh or t - t e r m pr ogr essive ear ly di age n esis in d e n sit y b an d s o f
r ece n t cor als: Porites co lo n ie s, M au r it iu s I slan d , I n d ian O c e a n . - Facies, 27:
10 5 - 1 1 2 , 3 p is.; Er lan gen .
P o t t h a s t , I. & O e k e n t o r p , Kl . (1987): Ein e Fa v o sit id e n fau n a au s d e m E m si u m /
Eife liu m des H a m a r Lagh d a d , Tafilalt (SE M ar ok k o). - Mü n st er . For sch .
Ge o l. P aläon t ., 66: 57- 94, 4 A b b ., 6 Taf.; Mü n st er .
P r e o b r a z h e n s k i y , B. V. (1967): Zn ach e n ie zo n aln y k h yav le n iy v skelet e t abu lyat om o r fn y k h kor allov. - P ale on t . Z h u r n al., 1967 (3): 3 - 8 ; M o sk v a (Akad .
N a u k SSSR ). [Sign ifican ce o f zo n al feat u r es in t h e ske le t on o f t ab u lat e
cor als]
R u k h i n , L. B. (1938): N iz h n e p ale o z o isk ie ko r ally i st r o m at o p o r o id e i ver k h n ey
ch ast i b asse y n a R . Koly m y. - [In :] M at e r ialy p o izu ch e n iy u Ko ly m sk o go I n d igir sk ogo Kr aya., ser. 2, Ge o l. i ge o m o r f., 10: 119 p p ., 28 p is.;
Le n in gr ad , M o sk v a. [Low er P alae oz oic cor als an d st r o m at o p o r o i d s o f t h e
u p p e r par t o f t he Ko ly m a- I n d ir ga River basin ]
Sc H O U P P ii, A. v. & S t a c u l , P. (1966): M o r p h o ge n e se u n d Bau des Sk elet t e s der
P t er ocor allia. - P alae on t o gr ap h ica Su p p l ., 1 1 :- 1- 1 8 6 , 132 figs., 8 t bls., 6
p ls.; St u t t gar t .
S o k o l o v , B. S. (1955): Tab u lyat i p a l ae o z o y a e v r o p e ysk oy ch ast i SSSR . Vveden ie.
O b sh ch ie v o p r o sy sist em at ik i i ist or ii r azvit iy a t ab u ly at . - Tr u d y V N I G R I ,
85: 527 p p ., 82 t ext - fig., 90 pl .; Le n in gr ad . [P alaeozoic Tab u lat a o f t he
Eu r o p e a n par t o f t h e U SSR . I n t r od u ct io n t o t h e gen er al st u d y o f t h e syst em at ics an d d e v e lo p m e n t o f t h e Tabu lat a]
- (1962): P odk lass Tab u lat a. - [In :] O r l o v , Y u . (Ed .): O sn o v y P ale on t o logii.
2. Gu b k i, Ar k h eot siat y , Kish e c h n o p o lo st n y e , Ch e r vy : 1 92- 26 5, pls. 1 - 18 ;
M o sk v a (Akad . N a u k SSSR ). [Su b class Tab u lat a - [In :] Fu n d am e n t als o f
P alae on t ology , 2, Sp o n ge s, Ar ch ae ocy at h id s, Coe le n t e r at e s an d W or m s]
S o k o l o v , B. S. & T e s a k o v , Y u . I. (1963): Tab u lyat y p a l e o z o y a Sib ir i. - 188 p p ., 11
t ext - figs., 21. p l.; M o sk va, Le n in gr ad (Ak ad e m ia N a u k SSSR . Sib ir skoe
ot d e len ie I n st it u t Ge o l o gii i Geofizik i). [P alaeozoic Tab u lat a o f Sibir ia]

T o n g - D z u y T h a n h (1966): N o u v e a u x gen r e et sou s gen r e ch ez les Coe le n t e r es
t ab u lat o m o r p h e s du D e v o n ie n du N o r d Viet n am . - Act a Scie n cia. Viet n am
Se ct ion Biol. Ge ogr . Ge o l., 1: 2 2 - 3 2 , 2 t ext - figs., 1 p l.; H a n o i.
T o n g - D z u y T h a n h ; N g u e n D u c K h o a ; K h r o m y k h , V . G.; N g u y e n H u u H u n g ;
N g u y e n T h e D z a n & Ta H o a P h u o n g (1 9 8 8 ): S t r a t ig r a fia i t se le n t e r a t y
O ek en torp,

- [In :]

D e v o n a V ie t n a m a , 2 T se le n t e r a t y . - 2 4 8 p p ., 6 4 p l. [D e v o n ia n st r a t ig r a p h y
a n d c o e le n t e r a t a o f

Viet n am ,

2

Coe le n t e r at a]


308

K. O e k e n t or p an d T o n g- D zu y T h an h , Rem ar k s

Ya n e t , F. E. (1956): Klass A n t h o z o a , G r u p p a H e lio lit id a , O t r y ad Fav o sit ace a. [In :] K i f a r i s o v a , L. D .; M a r k o v s k i y , B. P. & R a d c h e n k o , G . P. (E d s.):
M at e r ialy p o p a le o n t o lo gii. N ovy e se m e ist va i r ody. - Tr u d y V SE G E I , n . s.,
12: 3 0 - 3 5 , p ls. 8 - 9 ; Le n in gr ad . [C lass A n t h o z o a , G r o u p H e lio lit id a . O r d er
Fav o sit ace a. [In :] C o n t r ib u t io n t o P alae o n t o lo gy . New fam ilie s an d n ew
gen er a.]

Revid ier t es


M an u sk r ip t

bei

der Tü b in ge r Sch r ift le it u n g
19. D e z e m b e r 1995.

A n sc h r ift e n

e in ge gan ge n

am

d e r V e r fa sse r :

e k e n t o r p , For sch u n gsst elle fü r Ko r al l e n p al ä o z o o l o gi e , Ge ol.- P aläon t .
u n d M u se u m , W est fälisch e W ilh e lm s- Un ive r sit ät , P fer d egasse 3,
D - 4 8 1 4 3 M ü n st e r / W e st f.; T o n g - D z u y T h a n h , D e p ar t m e n t o f Ge o logy , Viet n am
N a t i o n a l Un iv e r sit y H a n o i , 90 N gu e n Tr ai Ro a d , D o n g- D a. H a n o i , Viet n am .

K

l em en s

I n st it u t

O