THE ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA (PRIBYL

THE ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA (PRIBYL

ACT
Vo l. 11/
A
PALAEONTOLOGI
I
9
5
CA
8
POLONI
C
A
No.2
FRANCISZEK ADAMCZAK
THE ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA (PRIBYL)
(OSTRACODA)
Abstract. - This p aper deal s w ith resrult s of in ve stiga tio n s on genus K ozl owsk ieZla
( P r iby l) fr om Midd le Devon ian b ed s in localities Wydryszow, Grzegorzowice and
Ska ly (Holy Cr os s Mountains, P oland). The assi gnment of the mentioned genus to
Ulrichia Jones as its sub genus - as advocated by Pribyl - is here r ej ected . On t o,gen y an d v ariab ili ty of descr ibed fo r m s, in cluding some new species, a re a naly zed.
INTRODUCTION
This paper prese nts information ob tain ed during r esearches on
<os tr acod s of the ge nu s K ozl owskiella (Pribyl) from the Middle Devonian
o f the Holy Cross Mts.
'I'h e co pio us m at erial , on which the in v estigations a re ba sed, has been
collect ed by th e w r it er in loc alities of Wydrysz6w, Grzegorzowice and
S kaly in 1955 and 1956 . No ost racod descriptions have thus far been
publis he d f rom th e fi rst t wo of th ese loc alities a nd the material now
co ll ected ther e sh eds som e light on th e evolution of the Middle Devonian
gen us K ozlow sk iella (Pribyl). The ontogeny and va r iations of the here
.d escr ibed species h ave been st udied in detail, h enc e it has been possible
to det ermin e t he mu tu al relations b etween t hese fo rms as well as t he
modifications which th ey ex pe r ience in the course of ontogeny.
. The relation of Ko zlo wskiella to Ulrichia Jones has also been a na lyzed.
Accord ing to A. Pribyl (1953) K ozl ow ski ella is a subgenus of U lrichia .
The present writer, how ever , h as ar r ive d at the concl usion that Ko zlowski ella should be se pa rat ed into a distinct genus and , in regard to its
.dimor ph ism , perhaps similar to the cru minal one, probably placed within
the fa m ily Beyrichiidae.
Th e a ut hor h er e expresses his warmest gratit ude to Prof. Dr. R. Kozlowsk i for th e va lua ble sug gestions and criticism ge ne rously offered
du ri ng t he preparation of the paper. Many t hanks are a lso due to Dr.
Z . Ki elan fo r r eadi ng t he m anusc ri pt an d to Mrs. J . Humnicka for translating the text into E nglish , as we ll as to Miss L. Luszczewska for the
pai ns t aken in doin g th e photographs .
76
FRANCISZEK ADAl\lCZAK
Cordial t ha nks a re also due to Dr. E. N . Polenova of t he VNIGRI
In stitut e in L eningr ad , for her gift of com parative m aterial a nd for facilities offered to inspect t he r ich ost r acod collection from Devonian beds
of the USSR during the writer's visit to Leningrad in 1957.
Th e present work is the second in a se r ies of papers to be d evoted
to Devonian ostracods from the Ho ly Cross Mts .'
TERMINOLOGY
The terms used here for the various m or phological elements of the
carapace are those introduced by 1. Hessl and (1949), R. V. Kessling (1951),
A. Martinsson (1956) and V. J aanusson (1957). F urthermore, the r e arealso some terms introduced into literature by G. Henningsmoen (1954),
and accepted by t h e present writer.
4
Fig. 1.
Ornamental details of
carapace surface; S l anterior sulcus, L 2 median lobe, S2 median
sulcus , L3 posterior lobe, t tubercles, v r velar ri dge.
Fig. 2.
Transverse section of
carapace of fe male K. pr aetubercutata n. sp., simplified ; 1 ve lar
ri dge, 2 adrnarginal r idge, 3 subv ela r area, 4 do rsa l p li ca, 5 hi n ge
gr oove , 6 mu scle scar .
An ad ditional term is t hat of "dark lines" used to designate peculiar
microscopic struct ur e elements of the ca rapace which have been investigated in t hin sections of r epr esen tatives of genus Ko zlowskiella (Pribyl).
1 The firs t p aper : F. Adamcza k , Po tyzygia Giirich, a n Ostracod genus from the
Givetia n of t h e H oly Cross M t s, - A ct a P a laeont, P ol., vol. T, No.1 , pp. 35-48.
O NTOGENY AND EVOLUTION O F
KO ZLO lV SK IELL .4
77
These ar e dark lines vert ica l to t he surface of t h e carapac e and arra nged
to fo rm distinct r eticu lation of th e carapace.
In what the m argi nal va lve elements are conc erned it ha s been
asce r tained t hat the so -called ad marginal ridge is grad ua lly ' t ra nsformed
in to admarginal t ubercles. Both th ese elements are, th erefore, ho mo logous
and m ay b e of some significance for the compari son of re lated for ms
(fig . 1-4).
MATERIAL AND METHODS O F WORK
The mat erial studied in the pres en t pa pe r has been yielded by
marly-argillaceous beds of Middle Devonian age in the Lysog6ry region
of the Holy Cross Mt s. Samples were t ak en in th e localities of Wydrysz6w,
Grzegorzowice and Skaly. Withi n the two first n amed localities the
occurrence is not ed .of fairly comp lete Lo wer Cou vini an (Eifelian) series.
In either of these localities, however, the Couvin ian s eries are represented
by a slightly different facial typ e and, mo st likely , t hey do not belong
Fi g. 3. - K. ko zlowskii (Pribyl), right valve, inside
view (d ia gram somew hat schematic); 1 hi n ge groove,
2 lis t, 3 ve lar ridge, 4 a dm a r gin al tu be rcles.
to exactly the sa m e stra tigraphic horizon. It r at he r seems that the Wydrysz6w complex is olde r than that of Grzegorzowice. That this is so
may, to a certain ex tent, be infe r r ed fr om th e quantitative sh ar e of the
78
FRANCISZEK ADAM C ZAK
three repre sentatives of genus K cziouiskietla. (Pribyl) yielded by t h e two
successions.
The state of preservation 00£ the ostr acod fauna LS not everywhere
uniform. On the whol e quite a numb er of the Wydryszow zones display
F ig. 4. - L on gi tudin a l section of right valve of K . p r ae t u be rc u l a t a n . sp, (sc he m a tic diagram) ; 1 p or e ca n a ls, 2 dark li ne , 3 muscle scar , 4 a d margi nal r id ge ,
S vela r r id ge.
populations in excellent st at e of preservatio n, con t ai ning a fa ir ab und anc e
of representatives of all, ev en the yo ungest ontogenetic st ages . Only
within the b asal parts, t he va lves are scarc e and damaged so that it w a s
poss ible to coll ect b ut som e few specim ens. The Gr zegorzowi ce m at erial
is less complete, since lar ge pa rts of the se ct ion h er e are covered by loess .
Ostracod va lves from the basal po rtions of that section h av e not b een
stu die d at all in vi ew of their unsati sfact ory state of p reservation. The
upper portions of the Grzegorzowice succession h ave yielded a fa irly
r ich fa unal assemblage containing other fo ssil s b esides ostracods. Howev er , w hen comparing the popula t ions, collected from these two sections ,
t he assemb lage fro m Wydrysz6w is pronouncedly the richer one.
With in b oth th ese s ections, the occ urrence is noted, besides ostracods, also of b r yozoans, brach iopods , trilobites and crinoids. Some foram inife rs have been fo un d to o.
ONTOGENY A N D
EVOLUTION
OF KO Z L O W SKI E L LA
79
Material of Givetian age h as been yielded by one horizon only, i. e.
by the so -c all ed brachiopod shales of the Skaly series. The presence of
representatives of genus K ozlo w skiella (P ribyl) has not t h us far been
recorded from eit he r the lower or th e upp er parts of the Skaly section.
Ostracod va lves collected from the brachiopod shales ar e in an excellent
st a te of preservation; they include t he youngest instars too.
An a nalysis , of this material h as shown that we a r e here dealing
with species Ko zlowskiella ko zlow sk ii (P ribyl 1953), closelly allied w ith
the Couvinian forms.
In t he Skaly m aterial the m ajority of specimens are adult forms,
while in that from the Wydryszow section immature valves predominate.
A total of approx . 3000 valves of genus K ozlow skiella, corresponding to
var ious ins tars, ha s been coll ected from material sampled in the three
mentioned localities.
The copious Wydryszow m at erial has yielded some specimens, wi th
carapac es of the younger instar still attached to them . These specimens
no doubt represen t the momen t of moulting an d, unqu estionably, such
immat ure forms may be regarde d as necroco enosis. The isol ated val ves ,
on th e other ha nd , in the writer 's opinion, r epresent mostly moulting
mat erial, in this pa rticular cas e - a pseudocoe no sis . Such observations
can hav e some signifi cance for the st udy of populations.
Sam ples tak en fr om m arl y and marl y-argillaceous rocks were
mac erated with Gl aub er's sa lt. Th e washing r esiduum w as next examined
under binocular mi croscope, us ing a sim plified Tri eb el tray to pick
out the microfossils.
Measurem en ts we re taken by mea ns of an ocula r m icrom et er under
X 30 magnifica tion . To car ry out t he m easurem ent the va lves w ere pl aced
on a glas s plate coate d with a thin wax lay er enabling a convenient
or ient at ion of t he 'valves. This is a time-sparing m ethod, as several
hundred specimens m ay be placed for com par ison on one gl ass plate.
Some ' valves w er e t r eat ed by h ydrofluor ic acid (1. G. Sohn, 1956;
Ch. F . Up shaw and alies, 1957). The fl uor id ization method has produced
good results. Many ornam en t ation a nd str uct ure d etails m ay thus be
st udied in transm itted light withou t m utilating the va lves as in t he case
of thin se ctions (fig. 5). 10% hydrofluoric acid wa s used b y the writer.
Det ached va lves wer e immersed in a wax ves sel fo r a bou t fift een minutes
and then tak en out and washed with water . Car ap ac es, after treatm ent
by acid , in most cases proved to be damag ed, cracked, and unsuitable for
study in transmitted light; w hereas single valves studied in trans mitted
light , und er a X 600 magnif ica tion , r evealed ev en the most m inute struct ural d et ail s.
FRANC ISZEK ADAMCZAK
80
No d iff iculties w ere enco unte red during segregation of copious
material wi th immature forms. Kesling 's method (1953c), war r antin g great
eco nomy of -t im e, is a m ost suit able o ne fo r th is purpose.
Fig. 5. .
K . pra et ubercula t a n . sp., righ t va lve, fl uor id ized; X 40.
O BSERVATIONS ON THE VA R IABILI T Y IN GENUS KOZLOWSK I EL LA
(P RIBY L )
The three new Couvini an species di fferentiated by the writer within
genus K ozl ow skiella (P ri byl) and that d escribed f rom the Skaly (Givetian)
by A. Pribyl (1953) are geneti ca ll y very closely interallied. The fo rms
from the Couvinia n cons titute fa irly w ell differentiated populations
displaying two esse n tia l evol utiona ry trends.
The first trend is r epresented by K ozl ow skiella sim ili s n .sp . This
species is with a prominent 1.3 and with elo ngated valves.
The other trend is r epres ented by Kozlowskiella tuberculata n. sp.,
distinguished by a broad s u bvela r area a nd by th e presence of two tubercles on L3 . I n ad dition, r epresentatives of th is species a re of a size decidedly gre a ter tha n that of t h e r emaining forms. A ssoci at ed wi th t he same
t ren d of evolution is the third species, namely K ozl ow sk iella pra etuberculata n . sp. , charact erized by a te nde ncy to p rod uce t ub ercl es on
L3 (fig. 6).
K. pr aetubercul at a n . sp. will be h ere discussed fi rst, since it is
apparently tha t fro m w hich t he other fo rms have d erived. Within Lower
Couvinian rocks r epresen t ativ es of this s peci es predominate in the m ajority of samples . On the evidence of analyzed material, yielded by
successiv e horizons, a constant tend ency to increase size ha s been ascer-
ONTOGENY AND EVOLUTION OF KOZLOW SKIEL LA
81
c
B
/:·.,
1
"
Fig. 6. -
'
)
A
Mutual relationship between species of gen us
K ozl ow sk ie lla (Pribyl) in Middle Dev o ni an layers of the
H oly Cr oss M ts. ; A K. p raetu berculata n . sp., B K . t u ber cu /at a n . s p. , C K. kozlowskii (Prib y l). D K. simi/is n . sp .
(le ft v a lv e s).
tained as ta k ing pla ce here. Adult form s within the basal parts of th e
se ct ion as a rule do not ex ceed 1.9 mm in length. Within upper portions
of the succession t heir length r anges from 1.8 to 2.15 mm, while forms
with a length from 1.95 to 2.4 mm hav e been found within top horizon s
(fig. 7A , B) . The outline of carapace is not subject to any great er variation s. In particular insta rs the length/height ratio (fig. 8a) shows a mea n
figure of 1.33, indicating a nearly suboval shape of all t hese carapaces.
Immature sp ecimens do not display any importan t va r iations . Differenc es
in , respect to the quantitative per cent participation of Ko zlowskiella
praetubercula ta n . sp., as com pa r ed against those of other here describ ed
species, are ob viously importan t. K. pr aetub ercula ta .n . sp . · (fig. 9) IS
Acta Palaeontologlca P olon lca -
vol. Ill/2
6
Fig. 7. -
A Size of comple te adult carapaces a nd of single left v alv es in K . prae -
tubercu~ata
n. sp, from three different Wydrysz6w horizons. Length of valves in
mm - on ordinate, samples from succes sive la yers - on ab scissa. B Per cen t
figures of adult forms of K. praetuberculata in the faun al composition of three
different Wydrysz6w horizons. Numbers of lef t valves and of com plete carapaces
in per cen t figures - on ordinate, size in mm - on abscissa.
distinctly predominant in samples collected from the lower layers of the
Wydryszow section. In the Grzegorzowice
a, bdc
9
section
(fig. 10), which has yielded relatively
,
I
,, I
less abundant material (approx. 50 speciI i
! ~ ,j
., 8
mens) , the p articip ation of this species is
! ,\/
~
smaller.
The predominant form is here K. si.: r/
s 7
\. /
n.
sp.
wh ich, on the average, represents
m
ilis
,
i
,,
6
i
49.8 per cen t of the total ' population. K. tubercula ta n. s p. makes up only 4 per cent in
,
5
samples from t he Grzegorwwice s ection, but
it is an interesting form owing to its inter4
mediate position between K. praeturbercu3
J
lata n. sp. and K. ko zlowskii from the Give,'/
ti a n . These three species are characterized by
2
•
a constant ontogenetic trend to produce
\
\
a broad subvelar area (fig. 11, 12), a prominent L3 and admarginal tubercles. A branch
line in the evolution of this species is re14
15 16 l/H
13
presented by K. similis n. sp. (fig. 6).
Fig. 8. - Length/height r atio
The numer ical share of K . tuoerculata
of carapace in va ri ous instars ;
n . sp , within the basal parts of the Wydrya K . pTaetubeTcu~ata n. sp. ,
b K. tubeTcu~ata n. sp., c K.
szow section does not exceed 10 per cent
kozlowskii (Pribyl), d K. si(fig. 9) of the total material in this sample.
mHis n. sp,
I
\
~
\
"
ONTOGENY AN D EVOLU TION O F
KOZLOW SKI E L LA
83
c
B
F
c ..
Fig. 9. - Per cent figures of three spe cies of Kozlows kiella (P ri byl) in the faunal
composition of si x Wydrysz6w horizons ; A-F suc cessi ve horizons, a K. similis
n. sp ., b K. praetuberculata . n. sp., c K. tuberculata n. sp .
Within the following higher layers the numerical participation of this
species increases steadily. In the Grzegorzowice material, however, the
first rank is occupied by K. praetuberculata n. sp. and K . similis n. sp,
(fig. 13) whose numerical participation in the different samples varies
strongly. .Every max imum share of one species (e. g. K. 3imilis n . sp.)
corresponds to the minimum share of another one (K . praeturberculata
n. sp.) . K. tuberculat a n. sp. has but few representatives.
K. kozlowskii, described by Pribyl from Givetian beds within the
Skaly section, apparently constitutes the terminal stage of the here consid er ed evolutionary line. Even at first sight this form strongly differs
from its predecessors. The most obvious difference lies in the smaller
dimensions of its carapace whose length does not exceed 1.8 mm (fig. 14).
ThE' posterior lobe (L3) is prominent with outline resembling that of L3
in K. similis n . sp. There is, however, a fundamental difference between
t hese elements in the two species. The posterior lobe in K. kozlowskii
84
FRANCISZEK A D AM C Z AK
G
J
K
bCJ
c
F ig. 10. - Per cent figures of three species of K ozl ow sk i el l a (Pribyl) in the fau nal
composition of five G r zegor zow ice horizons; G-K successive h or izon s, a K. similis
n . sp ., b K . praetuberculata n. sp ., c. K. tuberculata n . sp,
(Pribyl) has been produced by the gradual evolution of two t uber cless,
which finally m er ge into one prominent lobe (L3), while in K. similis n. sp.
the same element has probably been formed in a saltatory mode, b ecoming constant in the following generations. Thus, t he same detail of carapacial ornamentation has, within the same genus, evolved in two diff eren t
ways and not contemporaneously, finally resulting in specif ic differentiation. This interpretation is also suggested by the ontogeny of the carapace
(text-pl. I), w hich completely explains the gap within the available m ateria l derived from Upper Couvinian an d Lower G iv et ian beds .
The cha nges experienced by the here considered species, d ur ing th eir
evolution, were not, n at ur ally , confined t o one char act e r only, speaking
p recisely, in this case to L3 . The ve lar r idge an d t he for m ation of t he
so -called adm argina l t ubercles on the right va lve of the os t racods are
clo sely correlated with this ornamental d et ail. In specimens of K . prae tube rculata n . sp. the subvelar part is narrow and its wi dth in ad ult individ ua ls does not exceed 0.4 mm: in a ddition, ·t he so-called admar ginal
r idge (r ight valve) has developed along t he free m a rg in of the va lve. The
O NTOGENY AND
4
EVOLUTION O F
K O Z l. OWS K I E L L A
8 f)
d
I
t
3
c~
d
10
mm
I
I
:2
5
I
r
dJ
2
3
6
5
4
7
8
In.stQr~
I
r
Fig . 11. - Cross secti on
of ventral side of righ t
va lv e (schematic di agr a m ); 1 K. praetu bercu lat a n . sp ., 2 K. si milis n. sp., 3 K. tuberculata n. sp ., 4 K. ko zl owsk i i (Pribyl ), r ad m ar gin a l ridge, t a d margin al tube rcles .
Fi g. 12. - Wi d th of s u bv ela r a rea in v arious
instars ; 1 K. praet uoerculata n . sp., 2 K. kozl ows k ii (Pf lbyl), 3 K. si m ilis n. sp ., 4 K. tuber culata n. sp. Width in mm on ordinate,
instars - on abscissa .
100 %
Fig. 13. - Per cent figures of two
species of K ozlowskiella (Pribyl) in
the faunal composition of five Cou vinian horizons in Grzegorzowice ;
d K. praetuberculata n. sp., e K. simitis n. sp .
Per cent fi gu r es of
numbers of individuals - on or dinate, su ccessive horizons - on abscissa.
d
50
e
2
3
4
s
FRANCISZEK ADAMCZAK
86
..
1.2
mm
1.1
•
• ,
, ,, ,, ,
, ,,
"
•
II
• •
,.
• •
,.
• •
,,
•
•
to
.. .
•
•
II
•
(
.9
1.4
1.5
1.6
1.7
1.8
Fig . 14. - Size of a dult carapaces of K. k ozl ow skii
(Pribyl). .He igh t of ca rapaces i n mm - on or di na te,
length - on abscissa. Female sp ecimen s marked by
squares, males - by t r iangles.
subvelar part is considerabl y broader in K. tuberculata n . sp. , being
0.88 nun (fig. 12). On the other hand, however, instead of the admarginal
ridge the appearance is noted in this form of admarginal tubercles which
are markedly larger and better pronounced in K. ko zlowskii (Pribyl)
(fig. 11, 15). In what the wid t h of the subvelar area is concerned, it is
less than that in K. tub ~rculata n. sp., bu t in proportion to the size of
t he cara pac e when w e con sider tha t t h ese a r e forms on e inst ar smaller.
STRUCTURE OF CARAPACE
Ostracod carapaces studied in thin sections pr ovid e interesting
informa tion regarding the mod e of for mation and di fferentiation of the
va r ious d etails of ornamentation . This is particularly so in respect to
th e marginal parts of the carapace and to it s structure since these are
s ub ject to minor though occ asion ally f un d amental modifications and
h ence ma y be of som e taxonomic significance .
In papers on living ostracods , a vaila ble to t he writer, problems
conce r n ed with carapacial st r uctu r e have been discussed by G. W. Muller
(1894), K. Fassbinder (1912) , R. V. K esling (1951) and others. Many observations have also been m ad e on fossil material: B. Zalanyi (1929),
E. Triebel (1941), 1. H essland (1949), S. A. L evi nson (1949), N. Spjeldnaes
(1951), V. Jaanusson (1957) and R. V. Kesling (1957a).
ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA
87
2
03mm
'--~~~~......J!
I
5
~3mm
Fig. 15. - Longitudinal section of right valve of K. kozlowskii (Pribyl) (schematic diagram); 1 pore canals, 2 dark
line , 3 muscle scar, 4 admarginal tubercles, 5 velar ridge,
6 L3, 7 S2, 8 L2.
In the case of species of Kozlowskiella (Pribyl), the author has done
a number of thin slides of mature and immature carapaces. Furthermore,
he has also studied the thin sections of species belonging to other genera,
such as Saccarchites ornatissimus (Giirich) 2, Polyzygia symmetrica Giirich and Polyzygia trigonata Giirich. A "dark line", by Jaanusson (1957)
referred to as the bisecting line, has been noted in these forms in some
parts of the free margin. In species of Kozlowskiella and in S. ornatissimus
it hClJS been ascertained that these lines are regularly arranged over the
other parts of the carapace also (fig . 4, 15). Quite recently, Kesling (1957a)
describes such elements in genus Hibbardia Kesling where, according to
that author's statement, they produce a reticulation readily discernible
in tangential s ections, The occurrence of "dark lines" on carapaces is
not an uncommon phenomenon. In genus Kozlowskiella (Pribyl) they
constitute, a", in Hibbardia Kesling, distinct reticulation also discernible
2 This species has been described by Giirich (1896, p, 383, pl . 14, fig. 3 a, b)
as Primitia ornatissima from Lower Couvinian rocks (zone with Spirifer dombroviensis Giirich) from Dabrowa near Kielce. In view of the close resemblance of
this species to representatives of genus Saccarchites Swartz & Whitmore, there
is more reason to refer it to this genus.
.
88
FRANC I S Z E K ADAMC ZAK
in transmitted light under X 600 magnification of fluoridized valves
(fig . 16). Most likely , thes e details thu s fa r recorded in few forms only,
ar e all of the sam e typ e. This, among other things, s eems to be pointed
out by the ir position . In polarized light th e st r uct ure of the "da r k lines"
0.05 mm
L.-~~~""""'"
F ig. 16. - F ragment of the surface of ca rapace
in K. p rae tu be r cu tat a n . sp . (d ia gram slightl y
simplifie d) ; pc por e ca na l, dt d ark line.
is distingu ishable from tha t of adjacent pa rts solely by smaller dimensions
of calcite crystals. It should be h er e mentioned that these lines widen
out towards the outer su r fa ce of va lves in a fan-like shape and alw ays
occur on the m argin s of reticulation .
In view of t hei r m in ute dimens ions th ese are not pore canals a s
allege d by Jaanusson (1957) and K esling (1957a). In Kozlowskiella (Pribyl)
t heir size does not ex ceed 5 micrones. The occurrence of th e dark belt
may be interpreted by th e existence t he r e, d ur ing lifetime of the ostracod ,
of some organic substance whose presence caused a sligh tly different
mode of formation of the calcite crystals. Valves when dissolved in
hydrochloric or acetic acid do not reveal the presence of any such organic
matter. It seems probable that the dark lines may in some way be
associated with the calcification process of the valves. K. Fassbinder
(1912), who has studied the immature instal'S of the living Cypris pub era,
states that the calcification process starts earliest within the free margin
ONTOGENY AN D E VO L U T IO N OF K OZI.OW S K IELLA
89
area, immed ia tely after moulting. In im mat ure fo rms of Kozlowskiella
(Priby l) t he presence of th e dark line h as been noted already during t he
first instar (fig. 17), b ut within the Iree m argin a rea on ly, no such d ar k
lines bei ng observable in t he r em ai ning par ts of the carapace . In th is
insta r the valves are very fine, wh ile wi t hi n the free margin area
a distinct velar structure is already in ex ist en ce, transected by a dark
line. In the following instars the dark lines graduall y mak e t heir
appearan ce on t he othe r pa r ts of t he carapace as well. Th eir or ientation
is constant from the ventral to the do rsal m argin. By the '6th inst ar the
whole carapacial surface is covered with them. The same de tails of
struct ure have been noted in Sacca rchites orn atissim us (Gurich), bu t they
Fig. 17. - Cross sections of three earliest instars of
K. praetub erculata n. sp . (diagram f igure).
a r e m issing during the earliest instars of this species; the vela r str ucture
being, moreov er , in t hese for ms less prono unced , and t he retic ulation
more delicate. Here the dark lines begin to appear dur ing t he 3rd or 4th
instar, in t he same m an ner as in genus Kozlowskiella (Pribyl) .
Pore cana ls show a most regula r ar ran gemen t with in retic ulation
pi ts (fig . 4, 15, 16). They are funnel s hape d canalets readily discernible
on fl uor idized valves (fig. 5). Th ey are absent fro m the fre e m argin area.
ONTOGENY
Th e p roblem of ontogeny, i. e . of d evelopmental chang es ex perienced
by the carapace in the course of success ive moultings, is one of great
signif icance for phylogenetic studies. Stress has been laid on it in many
pa pers beginning with that by M. Verworn (1887). So far , however, it
was not possible to draw any general phylogenetic conclusions concerning
the described forms on account of the inadequacy of the available mate ri al. Often the described instars were not linked subsequently with
90
FHANCISZEK ADAMCZAK
phylogeny or were investigated from a different aspect. This will be
easily understood upon considering that immature forms are not al w ay s
found in the residuum of the washed material.
4
50
3
5
6
~o
10
350
500
1000
1500
2000
~
F ig. 18. - D ia gr a m figure showing dimensions an d numbers of carapaces dur in g
va ri ous instars in K. praetuberculata n. sp, Numbers of carapaces - on ordinate,
size in micrones - on abscissa .
On evidence of the available material the writer has been able to
identify probably all the instars in the here considered three new species
of genus Kozlowskiella (Pribyl) and in the form previously d escribed by
P ribyl (1953) as Ulrichia (Kozlowski ella) ko zlowski i . Since these are
closely allied species it was possible to clarify their interrelations. In
con necti on with these observations the writer is of the opinion that the
sp eciation may take pl ace either by sudden leaps (K. similis n . sp .) or by
wa y of slow evolutionary ch anges as is illustrated by the carapacial
ontogeny of K. praetuber cu lata n. sp., K . t u ber cul at a n . sp. and K . kozlowskii (Pribyl) (t ext-pl. I).
The numb er of moults within Beyrichiacea has not been exactly
determined , ranging from 6 to 10. Apparently this is controlled by the
same factors as those now acting in the li ving forms (Kesling, 1952a).
In species of genus Kozlowskiella (Pribyl) 9 instars, i. e. 8 moultings,
have been ascertained.
Carapaces of immature forms are as a rule complete, but the yo un gest
in sta rs are very scarcely represented. They are fragile and brittle, often
filled in by calcite. S epar at e valves of instars 5, 6 and 7 a re fairly common.
Consid erable difficulties have been en countered in distinguishing the
A CTA P ALAEO NT OLO G IC A POLO"reA , VOL . I II
F . ADAMCZAK , TEXT-PL. I
i~
l!)
~
((!J
~
G
~)
e:J
C9
..
. di .d I ontogeny 0 f ca ra p a ces
c
• ' . •praetubercul ata
in:• A K
S li gh tl y sc hema tic diagra
ofa tIn
t u bemr c ul
a nIVI
. s pua
., C K . k ozl owski i (Prib yl), D K. slm zlls n . sp .
n. sp ., B K.
ONTOGENY AND EVOLUTION O F
KOZLOWSKI ELLA
91
'immat ure instars in the th r ee Couvinian s pe cies. A t first sight, their
ea r liest instars are practically indistiguishable. The var io us lin es of
ontogeny could b e differentiated by indirect m eans only, e. g. a close
comparative study of t he diff erent inst a rs, with respect to their state
of preservation, h as revealed that in representativ es of K. tuberculata
n. sp. t he colouration of the carapac e is n early always darker that th a t
i n the r emaining species. Such biometric methods as determining th e
·d imensions in the va ri ou s instars, th e growth coefficient etc: did not
prove of any h elp in th e st udy of t his problem. The yo unges t instars
differ very insig nifican tl y in what size is concerned. In order to identify
the various instars K esling's (1953c) m ethods w ere ap plied, theoretically
based on Brooks law. When dealing with large amounts of material
t hi s method has proved extremely valuab le. Furthermore, a graph has
been plotted of immature and adult individ uals (fig. 18), which coincides
-en tir ely with data obtained with the help of a "slid e rule" .
On evidence of such morphological el ements as S2 , L2 and L3 , also
on reticulation , it was possible to d et ermin e the generic position of carapaces already in th e first ins tar. Specific characters make the ir appearance later and stabilize in insta,rs 6 and 7. This, however, is not a lways
t he case. In Ko zlowskiella t u berculata n . sp ., for instance, the specific
position could be assigned already in the 4th instar. Sexual dimorphism
-does not occur before th e last moulting. In some instances, however, on
the size of carapaces (K. praetuberculata n. sp.) we are led to the suggestion that sexual maturity is reached e arlier (in the 8th instar) .
When st udy ing in divid ual on togeny the w r iter h as t aken into cons ider atio n the t wo essen tial trends, m entioned at the beginning of the
p aper , of which t he fi rst is represen ted by K. praetuberculata n . sp. ,
K. tuberculata n . sp. and K. k ozl ow slcii (Pribyl); th e other one - by
K. simili s n . sp.
In K . pr aetuber cul ata n. sp., as in the other species, S2 appears as
ea rly as t he first insta r . L2 and L 3, toget her with re t icula tion , a re still
ind istin ct. I n t he 3rd in st ar, a d elicat e border is o utlined , fo rmed by the
arrangement into two pa rallel ro ws of the r et icu la tion m eshes. In t he
4t h instar S I is o utlined, while in instar 6 or 7 two m inut e t ubercles appear on the posterior lobe with a cons tant tendency to increase
(text-p l. I A) .
K. tu berculata n . sp . displays already in th e fi rst instar we ll dev eloped L2 and L 3, as w ell as S2. The r eticulati on here is at t h at stage
more conspicuous that in t h e previous s pecies. After the s econd moulting
the tubercles on L3 are nearly completely formed (text-pl. I B). SI bec om es outlined in ' the following instar.
92
FRANC ISZEK A DAMCZAK
The last species of t his evoluti ona ry line is the Giv eti an K. ko zlo w skii (P rib yl). Its fi rst instal' is smaller t han in the previously mentioned
spe cies , while t he other ch ar a cter s coincide. Alre ady in the second inst al',
however, two min ute t ubercles m ak e their appea rance on L3 . SI appe ars ,
as usually , in the 4th instal'. Up to the tim e of the 6th or even the 7th
mou lting two tubercles persist on L3. Aft er t he 7th moulting , i. e. in the
8th instal', L3 becomes mo r e promin ent and p roduces a large k nob not
car rying any tubercles on its surface (text-pl. I C).
S2 is that character of ornamentation w hich has fundamen tally
altered the sc ulpture of the carapace of Kozlowskiella (Pribyl). Its formation is reasonably probable during t he youngest instal's only, when
the resistance power of t he carapace agains t cent r ipetal traction is r elatively weakest (adductor muscl e). All additional alterat ions oc cur around
t he sulc us w h ich h as t hus fo rmed, i. e. the formations of lobes L2 a nd
L3, t he en lar ge ment of the subvelar area as well as several m inor chan ges
bro ugh t abo ut by the fo rmat ion of these elements.
The appeara nce of a ne w character was investigated in the differen t
instal's (text-pl. I) by ob ser ving t he mo ment and mode of its fo rmati on
an d its changes in subsequent instal's. In K. praetuberculaui n. sp ., for
instance, t ubercles on L3 m ak e their appearance in later stages of ontogeny; subsequently t his character transgresses onto earlier stages (K. t u berculaui n. sp .) which is probably connected w ith stabilization of t h is
character. These elem ent s, however, a r e not constant in the here consid er ed evolutionary line. In K. kozlowskii (Pribyl) the t ubercles occur
in yo ung instal's only and constitute transitional elements leading to t he
st rongly developed L3 . Furthermore, on ontogeny of the carapace in
K. praetuberculata n . sp. it may be inferred that this species is, in all
probability, the ancestral form from which the other r epresentatives of
genus Kozlowskiella hav e d erived .
SYSTEMATI C DESCRIPT ION
Suborder P alaeocopa Henningsmoen , 1953
Superfamily Beyrich iacea Ulrich & Bassle r , 1923
F amily Beyrich iidae( ?) J on es , 1894
G enus K ozlowskie lla (Pribyl, 1953)
Geno type: Ulrich ia (Kozlowskiella) kozlowskii P ribyl (1953 , p. 241244 , pl. 1, fig . 1-18) .
Diagn osis. - Trilobat e fo rms, hinge line st raig ht . In central part
of carapac e an add uctor muscle scar . Velar r id ge along the free ed ge .
ONTOGEN Y
AND EVOLUTION OF
K O Z L O WS K I E L L A
93
S urface of carapace reticulated. Well pronounced dimorphism.
Representatives : Ko zlowskieHa ko zlowskii (Pribyl, 1953)
Kozlowskiella pra'etuberculata n. sp.
K ozlowskiella tuberc u lat a n . sp.
Ko zlo wskiella similis n . sp.
Remarks. - G enus Kozlowskiella (Pribyl). recorded from Middle
Devonian rocks of Wydrysz6w, Grzegorzowice and Skaly (Lysogora
r egion of th e Holy Cross Mts.) , by Pribyl (1953) described as a n ew subgenus within genus Ulrichia Jones , is by the present writer separ at ed
into a new genus, independent of Ulriclua, and tentatively included into
the family Beyrichiidae. This conclusion is founded on detailed st udies
on the morphology, on togeny and var ia tions of the considered forms ,
grounded chiefly on the dimorphism sim il ar to the cruminal type , as
defined by Jaanusson (1957) .
Kozlowskiella (Pribyl) displays similari t ies with genus Ulrichia ,
whose representatives ar e mentioned by Pribyl, in some morphological
details only. The cardinal difference between Ulrichia and Kozlowskiella
lies in the presen ce in th e latter genus of Sl and S2 , and in th e distinct
se x ual dimorphism occur r ing throughout a ll th e sp ecies of this genus .
An additi onal difference is a w ell de veloped muscle area in th e central
po r tio n of th e va lve . Of the other forms with which K ozlowskiella is
compared by P ribyl , that of Bolbiprimitia Ulrich only di splays a certain
r esemblance; also some other forms s ubseq uent ly described and referred
to here below. Th e assignment of K ozl owskiella to genus Ulrichi a has
also been qu estioned by J aanusson (1957) . He writes that in type of
lob ation this gen us resembles Thomasatia Kay and pro bably for this
reason he places K ozl owskiella (Pribyl) w ith in the family Bassl eratiidae.
T he r eferen ce of gen us K ozlo w sk iella to the B eyrichiidae is no t
beyond doubt in the light of t he latest studies done by K esling (1957a)
an d of cer ta in observations ca r r ie d o ut by t he present w ri t er on his ow n
materia l. A comparison of o ur form w it h ge n us Hibbardia K esling (1953b)
r ev ea ls certain a nalogies . These consist among o thers of t h e ex ternal
resem blance of di morphis m and clo se sim ilar ities in the develo pment of
the vela r r idge and of ornamen tation. The cardin al diff ere nce is t hat
of the o rig in of dimorph ism in K ozl ow sk iella.
Th e origin of gen us Kozlowskiella is not thoro ug hly, kno wn . On the
comparison of certain m orphologic al elements it may be supposed to
h a ve derived f rom form s coming ne ar to Saccarc hit es orn at issim us (Giir ich ), di stinguished by dimo r phism resembling that in K ozlo wskiella.
Sa ccarchit es ornatissimus di splays a moderate amount of r es emblance
to th e species described by Swartz an d Whitmore ,(1956) as S accarc hites
94
FRANCISZEK ADAMCZAK
saccuiaris. Those authors allege that the considered form resembles genus
PhLycti scapha described by Kesling (1953a). Nevertheless, though these
forms do display a certain resemblance, yet the present writer is of the
opinion that the we'll known genus S accarchites has more charact ers in
common with S. ornatissimus (Glirich) than with the representatives of
PhLyctiscapha. Features r esembling those in Phlyctiscapha are t he type
of dimorphism, outline of carapace and the muscle scar.
Not eworthy a re al so forms which in some re spects a re similar to
Saccarchites , in others to K ozlowskieHa. They ha ve been described by
Polenova (1955) from the Volga-Ural province in the Soviet Union ter r itory und er th e name of AparchiteHina decorata. One of these forms in
particular, namely AparchiteHina cf. decorata , kindly presented to the '
writer by Mrs. Polenova , is to some extent a transitionary form. Particularly noteworthy in this spe cies is the muscle scar, developed similarly
as in S accarchit es, whil e the carapacial outline and details of ornamentation com e closer to these characters in Ko zlowskieUa.
It seems reasonably probable that forms such as : Saccarchites saccularis Swartz & Whitmore, 1956, S. ornatissimus (Glirich, 1896) and
AparchiteHina cf. decorata Polenova, 1955 - constitute a morphological
evolutionary line, with S . saccularis as the probable ancestral form , in
view of its poorly differentiated ornamentation and its occurrence within
Upper Silurian rocks. Th e two other species have been recorded from
t he Middle Devonian.
R. Ruedemann in 1909 (fide Ellis & Messina, 1952-57) has described
MacronoteHa jragaria from the Ordovician of the United States of Americ a . This form is an interesting on e on account of its res emblance to
Sa ccarchi tes ornatissimus (Glirich) , as is stated by its au thor. In both
these forms th er e are s imilarities of the outline of carapace, of ornamentation and of muscle sca r . A comparative study of S. ornatissimus with
MacronoteHa hypercala Kesling & Kilgore (1952) points out some interesting conclusion s too. Dr. Keslin g was kind enough to provide the
present writer with an assemblage of Devonian ostracods from North
America. Direct comparisons of the two above mentioned spec ies d id not ,
however, lead him to detect any differences between the immature form
of S. ornatissimus (5th and 6th instars~ and the American species.
Presumably, a detailed st udy of these forms may clarify this problem. Sections would, it is believed, be of special significance. Forms of
S. ornatissimus, including their ontogeny, have been studied by the
writer. He has .also made a number of thin sections of both, the immature
and adult, forms and has arrived at the conclusion that they may bephylogenetically related with Ko zlowskieUa (Pribyl).
ONTOGENY AND EVOLUTION OF KOZLOWSK IELL A
95
Ko zlowskiella praetuberculata n. sp .
(fi g. 19, 20; p l. I , I II)
Holotypus: fe ma le cara pace in pi. I, fi g. 1 a-e.
Stra tu m t yp ieum: Lower Couvinia n, Lys ogo r y region, Holy Cross M ts .
Locus typi eu s: Wydrysz6w.
D erivat i o n om inis : pra etu ber cu l ata s pecies less advanced than K. t ub ereula t a.
Diagn osis. Trilobate, outline of carapace suboval, hinge line
str aigh t ; subvela r area narrow; two sma ll tubercles on L3. Left valve
overlaps the right along the free edg e. S urface reticulated. Delicat e admarginal ridge on subvelar area (ri gh t valve). Distinct di morphism.
Description . - Outline of carapace suboval, preplete; dorsal border
st ra ight . A velar ridge along the free edge. Dorsall y , S l a nd 82 extend
fr om a common base. One of them, 81, follows the antero-ventra l d irection; the other , 82, markedly . longer, is perpendic ular to the do rsa l
Table 1
Kozlowskiella praetuberculata n. sp.
Dimenslons" of carapaces and growth factor in the several instars
.....0
.c:
Instar bliI:: I:
Q)I:
...:l~
1
.c:
-I-<
~o
0I-<U
~~
~.S
~Ol~_
I::fll-<I:
Q)00l1:
...:l't:lE_
V
0.76
VI
1.00
VII
1.32
VIII
1.68
\>
1.92
1.84
0.48
~~
0.28
,·,, 1
0.20
1.
1
0.48
0.7 6
1.32
1.14
• Qu ot ed da ta are modal values,
I
0.08
I
0.12
I
0.14
,
I
~~
1.16
1.28
1.44
0.26
1.07
0.28
1.21
0.34
1.24
1.17
0.40
0.88
1.36
1.20
0.88
~o
0I-< U
1.50
1.38
1.10
1.3 2
1.28
S::0l1::~
1.08
1.25
1.15
1.48
1.40
Ol I:
1.20
0.72
1.20
.- 't:l
.c:
"'1-<
0.18
0.48 :
1.33
1.33
I,
0.52
0.96
1.28
I
1.20
1.26
1.27
1.33
i
0.72
0.96
1.30
1.15
0.40
1.25
I
1.26
1.31
I
1.20
0.60
0.60
I
0.3 0
I
iiS ..... 1:
t5~
1.25
26
0-~Q)Ol
.c;;_
o~
0.26
0.32
0.40
1.25
1.18
(J
I
I:
.c:1:
E-< ~
.~
:r:~
..... ,
.c:
-I-<
~ 0
.!l: ~
0". u
1.05 \
1.23
0.64
I:
Q) I:
0.38
0.52
~o
.~
~~
0.36
0.44
IV
0I-<U
-0
-I-<
1.29 1
1.18
III
~
'"
Q)
'"
I::
.c:
:s::_
-I-<
0.28
0.36
1.22
II
.c:
1.00
0.40
0.40
FRANC ISZEK ADAMCZAK
border. In th e centre of carapace S2 becomes club-shaped and bears the
muscle scar. Lobes mod erately prominent. Posterior lobe (L3) tending
to produce tubercles . On th e right valve a well developed hinge groove,
over the entire surface, with t he exception of the triangular area in the
s ubvela r area , the larger on the left valve, the smaller and more delicate
o ne - th e so-called admarginad ridge - on the right valve.
Dimensions a nd grow t h fa ctor of ca rapaces -
•
1.5
mm
•
•
1.4
1.3
1.2
see t able 1.
•
., ,
•
•
•
•
...
• •
•
•
,
'lll
•
~
• '"
,
,, ,,
,,
,
1.8
'-
1.9
•
20
2.1
2.2
F ig . 19. -
Size of carapaces in a d u lt form s of K . praetuberculata n . sp . H eigh t of ca rapaces in min on
on a bscissa.
or d ina te, length ma rked by sq u a r es, males -
F em ale specimens
by triangles.
Va riation . - Size of va lves varies r ath er st ro ng ly in d ependence on
t he stratigraphic ho r izon fro m which the material was sam pled. Th e
reti culation m eshes are usu ally subpe ntagonaland uniformly dispersed
over t he en ti re s urfa ce, wi th the exce pt ion of the triangular area in the
a ntero -do rsal portion of the ca rapace. Well d eveloped po re canals in t he
ret iculation .
Dimor ph ism . - In fema le specimens, which pr edom inate amo ng
m a tu r e ind iv id ua ls (68 pe r cent), t he ventral valve is distinctly swo lle n
(fig. 19).
ONTOGENY
AND E V O L UTION O F
KOZLOWSKIELLA
97
Ontogeny. - Th e collected immature valves probably r ep r esent comp let e series of instars, It is not excluded that in some cases adult individuals also moult their carapace, t his being s ugges t ed by the great si ze
range of mature va lv es (fi g. 20).
Instar I - Dimensions (in mm): length 0.36, h eigh t 0.28, thickness
0.20. Outline of valve dist inctly preplet e , h inge line straight. L2 a nd L 3,
and median 8 2 al ready w ell m arked. Delicat e velar rid ge ex tend in g a long
the fr ee edge; reticulation st ill rathe~ indist inct.
20
-
o/c
0
-
15
r---
t-I--
10
I--
5
I--
23
4567
8
l
9
i n s rcr s
Fig. 20. Per cen t figure of
ca r apaces of K. praetuberClllata
n. sp. in various in sta rs .
In star II - Dimen sions (in rnm): length 0.44 , height 0.32, thickn ess
0.26. Va lve s till bilobate ; reticulation more distinct; L 3 larger. The r emaining characters un changed.
Inst ar III - Dimensions (in mm) : length 0.52, height 0.40 , thickness
0.30. Reticulation mesh es larger than in earlier instars; L2 and L3 more
pro m ine n t.
Instar IV - Dimen sions (in mm) : length 0.64, height 0.48 , thickness
0.40 . In this instar 81 is o utlined , gently arcuate, somewhat obliquely
directe d fo rward. The edge on L 3 faintly outlined .
In star V - Dimension s (in m m): length 0.76, height 0.60, thickness
0.48 . 8 1 a nd 82 deepened . Ed ge on L3 di st inctly m arked. Rema inin g
cha r act ers unch anged .
Instar V I - Dimensions (in rnm) : len gth 1.00, height 0.72, thickness
0.52. Various ornamentation details are in t his s ta ge quit e distinct. Two
minute t ube rcles a ppear on L 3. A small rib is outlined between 8 2
a nd 83.
Instar VII - Dimensions (in mm): length 1.32 , height 0.96 , thickness ,
0.72. Tubercles on L3 more distinct ; th e remaining characters unchanged.
Ac ta Pataeontotogtca P ol ontca -
vol. IIIj 2
7
FRANClSZEK ADAMCZAK
98
Instar VIII - Dimensions (in mm): length 1.68, height 1.20, thickness 0.88. This instar precedes the mature form. Small tubercles on L3
st ill increase. Other details of ornamentation are those of mature forms . '
Insta l'
Le n gth of
ca r a pace (mm )
I
II
0.31 - 0.36
0.37 - 0.45
0.47 - 0.56
0.59 - 0.68
0.69 - 0.88
0.94 - 1.14
1.17 -1.43
1.48 - 1.82
1.84 - 2.40
III
IV
V
VI
VII
VIII
Ad ult
H ei ght of
carapace (mm )
0.20 0.22 0.32 0.42 0.50 0.62 0.96 1.10 1.32 -
0.28
0.32
0.42
0.55
0.6 3
0.9 6
1.15
1.35
1.50
R em ark s. - Kozlowsk iella pra etub erculata n. sp. di spla ys numerous
ana log ies wit h the oth er species h ere d escribed. Differences between them
consist in d et ails of orna men tation a nd varying size. There are also some
differences in the successive onto genetic st ages . This species is sim il ar
to A parc hi te llina cf. decorata Polenova . Its r esem blan ce to genus Hibbardia K esling is by t he writer r egarded as less st r ik ing, chiefly due to
different type of dimorphism .
K ozlo w skiella t u bercu lat a n. sp.
(fig. 21, 22; pI. II, IV )
Hol otypus: female carapace in pl . II , fig. 3 a-c.
St ratum typicum : L ow e r Co uvinian, Lysogory region, H oly Cross Mts.
Locus typicus: Wy d rysz6w.
D eri v a tio nominis : tube rculata -
two distinct t ubercles on L 3.
Diagnosis . - Trilob ate, outline of va lve pre plete; hinge lin e straight.
Two di stinct t ubercles on L3. A ve lar rid ge along the fr ee edge of carapace. Subvela r area broad . S urface reticula ted. Well defin ed dimorphis m.
Description . - Val v es of this species are of greate r size than in a ny
of t he rem ain in g fo rms of Koz lowskie lla (P rib yl). The posterior lobe (L3)
bears two d istinct t ube r cles constituting one of the specific ch aract ers.
51 and S2 exten d fr om a commo n base in th e dorsal part of th e valve,
separ a ting L1, L2 and L 3. S1 gently a rche d forward , while S2 occupies
a centr al position. Muscl e scar in the middl e of valv e. Subvelar a rea
ONTOGENY AND EVOLUTION OF KOZL OWSKJELLA
99
broad (0.88 mm) , delicate admarginal tubercles on the right valve. Dorsally (right valve) w ell developed h.ix:ge groove, expanded at both ends.
Pore canals in reticulation meshe s. L.~~t "v alv e overlapping the right.
.
'''.\
.'
Dimen sions a nd grow t h factor of carapaces .,-:-s ee table 2.
... . ~ -.,
\'.
.~ .! . : ..' .
T ab l ~ 2
· . f , . .•...
•'J "ot
, ..
" ~ Kozlow ski ella tuberculata d. sp, ,',
Dim ensions* ' or" ca r apaces and g row th fa'~tQr' 'i~ th e seve ral in stars
. '
.....0
J
I
.c
.c
In star 00s:: ~
I ~ £.
I
~ ~ '
I
II
I
I
I
I
III
0.4 0
~_
oB
.'
~tJ
0~
I
I
I
1.20
0.48
,
.c
I
.-
Q)o"' ~
...:l'OE
_
0.32
1.22
V
0.88
VI
1.20
VII
1.60
VIII
1.84
i
0.56
1.36
I
I
1.33
1.15
9
0
I 2.32
I
I
1.96
I
I
I
0.72
1
0.92
I
I
I
I
I
1.80
1.52
I
I
I
I
I
1.28
1.48
II 1.26 [
:r: ~
I
I
I
I
r
.c
.....
I
Ul
Ul
Q)
.c
....
c
~
.!o: _
~B
~'-'
I
I 0.22
~
I
j
'0
tJ
eu~-
E_
I
I
1.33
I
!
1.12 I
I
I
I
1.33 I"
I
1.33
~
tJ
d~
1.00
I
1.50
0.12
2.00
0.24
1.25
0.30
1.20
0.36
1.50
1.37
I
~
~ 0
0 ....
0.08
I
I
.c
....
0.08
1.09 I
I
~
~ ~~£.
d~
I
1.23
0.24
I 0.32
1.18
1.27 I
. 0.32
0.38
1.21
1.26
0.48 I
0.36
I 1.33
1.28 I
0.64
0.48
1.31
1.27
0.64
I 0.84
1.39
1.28
1.08
0.88
1.15 I
1.18
1.00
I 1.28
1.21 I
1.25 i
I
, 1.60
1.48
I
1.02
I 1.44
I
.c~~
......
0 ....
.c~
E-< _
" '6n ~
~
~ 0
.~ ~
d~
! . 0.26 j
f
0.36
0.4 6
0.72
, 0~
1.13
1.28 I
IV
~.
. 8 t; .. .~
Q)~
tlO"'tlO s:: ~ ~ ~
1.16
0.56
' .c .. . ~
....
~~,-.
s:: '
0.54
I 1.13
I
1.11
1.48 II
!
,
0.60
1.46
0.88
0.68
!
Qu oted d at u nre mud al va hi e ... .
V ar iation . - Only t he size of va lves is sub ject to certa in va ri ati ons.
The subvelar len gth/ width r atio va r ies f rom 4.4 to 7.1 (fig. 21).
Dim or phi sm . - In femal e s pecimens th e va lves are vent r ally strongly
swollen. Th e m al e specimens are mo re sle nder; thei r valves consti tute
a minority a mong ad ult sp ecimens.
On t ogen y (fig. 22):
Instar I - Dimen sions (in mm): length 0.40 , h eight 0.26, thickness
0.22. Ou tline subova l; r eticulation d ist inct. L 3 forms a fai rly poi nted
tubercle. Valve monosulcate.
Instar II - D im ensions (in mm) : length . 0.48 , height 0.32, thickness'
0.24. Outline and details of m orphology without changes.
FRANcmZEK ADAMCZAK
100
2
Fig. 21. - Diagram of car ap a ce length /width of
subvelar area ratio: 1 K . praetuberculata n. sp .,
2 K similis n. sp.. 3 K. tuberculata n. sp .
In st ar III - Dim en sion s (in mm) : length 0.56, h eight 0.38, t hi ckness
0.32 . Two t ubercles make th eir appearance on L3 .
. Instar IV - Dimensions (in mm): length 0.72, h eigh t 0.48, thickness
0.38. In this instar 81 is initially o utlined. T he wi dth of the s ubvelar area
hasdoubled.
20 'Yo0
,.--
15
,.-
10
f--
-------=
-,.-
5
Fig. 22. Per cent figure of
carapaces of K. tuberculata n. sp .
in variou s in stars. Per cent figure of the number of in dividua ls - on or di nate, instars
on abscissa .
-
23
4
5
67
8
9
ln~to r,s
Instar V - Dimen sions (in mm) : length 0.88, he ight 0.64 , thickness
0.48. Details of ornamentation more distinctly marked and not subject
to any essential changes during the following instars.
ON TOGEN Y AND EVOLUTION OF KOZLOWSKIELLA
. 101
Instar VI - Dimension s (in mm): length 1.20 , height 0.84 , thickn ess
0.64. No changes.
Inst ar V II - Dim en sions (in mm) : length 1.60 , height 1.08, thickness
0.88. In t his instar t he subvelar area increases considerably. Details of
orna me ntati on h~ve been definitely developed.
Inst ar VIII - Dimensions (in mm): le ngth 1.84, height 1.28, thickness 1.00. This ins tar precedes the matur e form .
Instar
I
II
III
IV
V
VI
VII
VIII
Adu lt
Length of
I ca rapace
I
I
I
-I
0.29 0040 0048 0.56 0.75 1.00 1.38 1.64 1.88 -
(mm)
H e ig ht of
carapace (mm)
0040
0048
0.56
0.72
0.89
1.28
1.60
1.85
2.70
0.22 0.27 0.32 0.37 0.50 0.64 0.87 1.10 1.42 -
0.26
0.3 2
0.38
0.55
0.68
0.84
1.10
1.30
1.80
Remarks. - K ozlo w sk iell a tu bercu lat a n . sp. is a transitionary stage
in th e evolutionary t rend leading f rom K. praetuberculata to K. kozlow sk ii (Pribyl) . A n um ber of morphological characters trac ed t hroughout its on togeny con firm this ass um ption. Tubercles situated on L3 are
of particular significan ce when comparing these forms. In K . tuberculata
n. sp. they make their appearance during t h e earliest stages of ontogeny
(3rd instar) , w hile in K. k ozlowskii - f ro m 2nd instar; in K. praetubercu lata n. sp., wh ich is und oubtedly a more primitive form , they appear
lat er and a re less strong ly d ev elop ed. F ur t h er mor e, in K . tuberculata
n. sp. admargi nal tub ercl es on t he ri ght v alve are equivalent to th e
adm ar ginal ridge in K. praetubercu lata n . sp. (fig. 11).
K ozlow skiell a k oz low skii (Pribyl)
(fig. 23, 24 ; pl. II)
Type spe cies: Ulri chi a (Ko zl ow ski ell a) ko zlowskii
p. 242-244, pI. I, fig . 1-18).
Pribyl
(1953,
Diagnosis. - Th e original diagnosis , as given .by P rib y l (1953, p. 318),
is: "Anterior node placed much lower than t he po st erior node , w hich
us ua lly projects above the dors al margin . There is a de ep f ur row b etw e en
two nod es. Anterior to th e a nterior node there is a lso a furrow directed
obliquely forward. The su rface of the va lves and nodes is coarsely re ti c ula ted".
FRANCISZEK ADAMCZAK
102
Description .
Carapac e trilobate, almost preplete; hinge line
stra ight. Hinge groove on right v alve along the dorsal border. Posterior
lobe (L3) in the shape of a prominent knob. Velar r idge along the free
edge. S1 ex te nds somew ha t oblique ly in r elation to S2 , sepa r ating th e
two lob es. The cen tral part of t he va lve is occupied b y the m edi al furrow (S2), w h ich stre tches to the centr e of th e va lve; it is there swo llen,
s uggesti ng the muscl e sca r. V er y fi ne reticula tion discernible on th e
poste r ior lob e (L3). Admarginal t uber cles d ev eloped on the right va lve
a long t he fr ee edge. Hinge furrow as in t he r em aining species , with
a hinge list on the left va lve.
Dimension an d growth factor of ca rapaces - see table 3.
Tabl e 3
K ozl ow sk i ella kozlow sk ii (Pri byl)
Dim ensions > of ca r ap aces an d growth fac t or in the seve ral instars
Instar
- ...
.c
....elI-
"c
.c
=: co .::
~ 0
0 ....
f:
....lal _f:
l::
I
....0
I
I
~ ~ ~1
4J o ctl f:
....l'CE
_
... U
(j ~
0.26
0.28
I
1.23
0.36
0.32
0.48
IV
0.58
V
0.72
VI
0.91
VII
1.04
VIII
1.36
Q
1.72
1.6 4
~8
0 ....
I
1.50
I
1.25
1.14
1.10
I
I
i
I
1.16 ,
1.16 ,
I
I
1.00
0.32
1.15
1.25
0.40
0.60
1.3 1 I
1.25
1.36
1.24
1.08
0.88 1,
1.23
1.60
0.32
0.52
1.17
1.66
0.20
1.26
1.25
1.17
1.08
I
1.28
0.80
1.30
1.00
0.12
0.48
1.15
0.92
1.20
0.12
0.3 8
0.68
1.40
1.56
0.94
0.60
0 ....
... o
(j ~
1.25
1.00
1.40
.......
~ 0
0.10
0.28
I
f:
;::'C
ctl f:
ctl l::_
1.40
0.54 '
0.80
~etSetS -
~ E-
I "c
0.0 8
0.2 8
II 1.28
1.33
0.64
(j~
0.20
0.42
1.24
0 ....
... u
ts"6D ~
"c ......
1.11
1
I
.......
~ 0
"c
0.18
I 1.18
1.15
0.48
1.26
-cf:
8 _
0.30 I
0.32
I
.!<: -
.::! 1:
... u
(j~
1.00
1.20
0'
0.22
0. 26
1.33
'ill
I
'"
al
'"
l::
"c
I
1.28
II
I
I .!::~8
0 .....
.~ f:
... u
al f:
::r:_
(j ~
0.50
0.56
* Q uot ed d a rn tire mod al va lu es.
Var iation. - Length of v alv e ranges f rom 1.4 to 1.8 rom and the
length/width ratio from 1.3 to 1.7 (fig. 23).
Dimorphism. In female spec im ens the a nt ero-v entral portions
of the carapace are distinctly swollen. As compared against the remaining species of genus Ko zlowskiella (Pribyl) , this ventral inflation has
been shifted somewhat anteriorly . Furthermore, it has been ascertained
103
O NTOGENY AN D E VOL U TIO N OF KO Z L O W SK I FJL L A
25 %
mm
-
23
I--
15
15
5 r---
13
14
1.5
16
1.7 L/H
2
F ig. 23. - K. k ozlowskii (Pfibyl ),
variation diagram. P er cent fi gure in the com positi on of various groups - on ordinate.
3
4
Fig. 24. - D ia gr a m figure showing cara paces di mensio ns of fou r species of genus
K ozlow sk ielta (Pribyl) in Mi ddle Dev onian layers of H oly Cross Mts. Supposed
changes w ithin the Uppe r Couvinian and
Lower Giv eti a n indicated by broken line :
1 Lower Co uvinian, 2 Uppe r Co uvinian ,
3 Lower Givetian, 4 Upp er Givetian .
o n the available m at er ia l that fema les cons titute 54.7 per cent of all
m atur e forms . He nc e, among Gi vet ian fo rms , t he n umb er of m ale specimens h as increased (fig . 14).
Instal'
I
II
III
IV
V
VI
VII
VIII
Adult
Length of
carapace (mm )
0.28 0.32 0.38 0.49 0.61 0.79 0.93 1.18 1.40 -
0.30
0.36
0.48
0.5e
0.78
0.91
1.12
1.38
1.88
Height of
carapace (mm )
0.21 - 0.26
0.22 - 0.27
0.20 -0.30
0.30 - 0.42
0.38 - 0.54
0.50 - 0.70
0.62 -- 0.80
0.78 - 1.00
0.92 - 1.24
Ontogeny:
Inst ar I - Dimensions (in mm) : length 0.28, height 0.22, thickness 0.18.
Bilobate form ; o utl ine of ca rapace preplete. L3 fairly prominent and sharp
pointed. Delica t e r eticulation on surface of va lves. Velar ri dge along
the free margin.
Instar II - Dimension s (in m m) : length 0.36, height 0.26, thickness
0.20. Two minute t uber cles appear on th e anterior lob e (L3). Remaining
fe atures witho ut changes.
In star III - Dimensions (in mm) : le ngt h 0.48, hei ght 0.30, thickn ess
0.28. L3 a nd t he t uber cles more prominent .
104
FRAN C ISZEK ADAM C Z AK
Instar IV - Dim ension s (in mm): length 0.58, height 0.42 , thickness
0.28. In this instar SI separating the two lobes (Ll and L2) is o utlined.
Instar V - Dimensions (in mm) : length 0.72, height 0.54 , thickness
0.38. L2 and L3 distinctly marked. Trilobation well defined.
Instar VI - Dimensions (in mm) : length 0.91 , height 0.68 , thickness
0.48 . Reticulation uniform throughout the va lve surface.
In star VII - Dimensions (in mm) : length 1.04 , h eight 0.80 , thickness
0.52. In this instar many individuals los e the tubercles on L3 which
beco mes more promin ent .
Instar VIII - Dimensions (in mm) : length 1.36, height 0.88, thickness 0.60. The di sappearance of tubercles on L3 is connected with formation of a more prominent lob e. The r emaining details of ornamentation
a re definitely developed. This in st ar precedes the m ature form .
Remarks. - K ozlowskiella k ozlo w skii (Pribyl) has so far been reco rd ed from t h e Givet ian and within one horizon only , in the so-called
brachiopod shales. This sp ecies has not as y et been found in other sections of t h e sa m e ag e in Holy Cross Mts., b eing r estricted perhaps to one
locality only. It is clo sely a llied w it h Cou vinian s p ecies, its dimensions,
however , a re smaller . During th e evo lutionary line of K. ko zlowskii w e
first o bse rve an incre ase of dimen sions (Couv inia n species) a nd then '
a se nsib le red uction (fig . 24). The limited horizontal and v ertical distri bution of this s pecies suggest t hat it was n earing extinc t ion .
K ozl ow sk iella similis n . sp .
(fi g . 25 ; pl. I)
H ol otypu s: fe male carapace in pl . I , fi g. 3, 4.
Stra tu m t yp icum : Lower Cou vin ian, L y sogory r egion , H oly C r oss Mts .
Locu s t y pi cus : W y d r y.s;z6w .
D erivatio n ominis: si m His - called so to s t re ss its close si milarity to sp ec ies
be l on gin g to t h e ev olu t ion a r y line : K. p raetube rculata n . sp., K. t u bercu l a t a n. so .
a n d K. kozlow sk i i (Pribyl).
Diagnosis. - Trilob at e form with outline a lmost am ple te. P osterior
lobe (L3 ) in t he s ha pe of a prominent knob. Left valve overlapping the
right ; surfac e reticulated; di stinct d imorphism .
Descr iption . - Carapa ces e lon ga ted, trilob at e and r eticul at ed. S l
a nd S2 e x tend from on e common base, sep ara tin g the particular lob es.
L 3 prominent, considerably larger than L1. Along the free edge of the
right valv e a delicate admargin al ridge, with m inute tubercles in some
individuals (t endency to produce admarginal tubercles) . The left va lve
som ew h at larger and overlapping , the right along the free edge. Pore
ONTOGENY AND EVOLUTION OF KO Z l. O W SKI E L LA
105
ca n als de veloped in reticulat ion m esh es. A h in ge f urrow developed on
the right va lve, dilated at both ends , an d a hinge list on the left va lv e.
Dimensions and growth factor of ca rapaces - see table 4.
Var iat ion. - Th e outline of the carapace and t he dimen sions of
L3 are subj ect to certain variations. The length/subvelar area r atio va r ies
fro m 3.3 to 3.7 (fi g. 21).
Dimorphism. - Carapaces of femal e individuals a re som ewhat la r ger
(fig. 25) , v entrally distinctly inflated . Mal e spe cimens constit ute 20 per
cent of the total mature forms.
1.5
mm
•
1.4
.
•
•
• •
1.3
,
..
1.2
1.9
,
,
-
• •
•
20
•
Fig . 25. - Si ze of carapaces in ad ult
fo r ms of K similis n . sp, Heigh t of ca r apaces in m m - on or d inate. leng th
- on absciss a. Female specimens m a r ked by squares, ma les - by tr ian gl es .
21
Ontogeny:
Instar I - Dimensions (in mm) : length 0.36, h eight 0.22, thickn ess
0.16. Ou tlin e s ubova l; 82 d evelo ped approxi mately in the middl e of t h e
carapac e. Reticulation faint.
Inst ar II - Dimen sions (in mm) : length 0.48, h eigh t 0.36 , th ickness
0.24 . In thi s instal' L3 is r ather prominen t. The remaining cha ract er s
unchanged .
Instar III - Dimensions (in mm): l en gt h 0.68 , height 0.44 , t h ickn ess
0.32. Main detail s of ornamentation with ou t cha nges .
Instar IV - Dimensions (in mm) : length 0.86, heigh t 0.56 , th ick ness
0.44 . 81 m ak es it s appearan ce. R eticulat ion more dist in ct.
In st ar V - Dimen sions (in mm): length 1.08, height 0.68, t hi ckn ess
0.52 . 81 and 8 2 ha v e de ep en ed. L 3 prom in ent.
Instar VI - Dimensions (in mm): len gth 1.24 , heigh t 0.88, t h ick ness
0.64. No changes.
Inst ar VII - Dimensions (in mm) ; length 1.44, h eight 1.00. I n t h e
writer's collection this instal' is r epresent ed by one v al ve only . Det ail s
of ornament ation un changed.
In star VIII - Dime nsions (in mm) : length 1.72, h eight 1.12, thickn ess 0.80. A ll details of ornamentation have developed definitely. T his
instal' precedes the m at ur e form .
FRANClliZEK ADAMCZAK
106
T abl e 4
KozlowskieUa si m ilis n . sp,
Dimens ion s" of carapaces and growth factor in th e several in stars
,
I
,,
I
Insta r
I
..c::
Q/)~
I:: ~
Ql-
I
...:l .£
I
-
i 0
..c:: ....
.....
i :S . . . .::
~ 0
0 .....
.... ~
tl _
tll) <tI
I
tll)~
I:: ~ .... ~
Ql 0 <tI ~
...:l'OE ~
0.36
0.4 8
III
0.68
IV
0.86
V
1.08
VI
1.24
VII
1.44
V III
1.72
.~
Ql
"
~
E-<~
tl~
..c::~
... CJ
<tI
"ollQl
..c:::;;~ ~
:0
E~
' - '0 <tI
~
~<tII:: ~
...
~ 0
0 .....
... CJ
tl ~
1.50
0.12
1.3 3
1.33
0.16
1.37
1.50
0.24
0.44
1.33
1.18
0.52
1.29
~
..c::
.....
0.08
0.32
0.88
-0 '-'
1.50
1.21
1.25
0.32
1.12
1.23
0.64
0.36
1.13
1.00
1.13
1.36
1.12
1.15
1.50
1.21
1.3 6
1.24
0.52
0.80
1.12
1.18
1.60
1.48
.~ ~
0.24
0.68
1.20
...
~ 0
0 .....
1.27
1.09
..c::
.....
0.1 6
0.56
0.96
I::
.!<: ~
1.22
1.37
1.16
tl~
0.44
0.88
1.20
... CJ
til
til
Ql
1.63
1.14
1.16
~i:l
0 .....
0.36
0.64
1.15
I .c
0.22
1.27
1.25
~
... CJ
0.56
2.00
1.88
~~
I
I tl~ I :r: f
0 .....
0.44
1.26
.
....
~ 0
1.57 '
1.40
9
I
0.28
1.33
II
I.c
1.20
1.20
0.60
0.54
* Qllotl'd da rn an ' mo d a l ve lu c - .
R em ar k s. - On the ca r a pacial ou tline and various d et ails of ornamen ta tion this species m ay be opposed to t he r emaining r epresentatives
of gen us Kozlowskiella (P ri byl). The ca rd inal feat ures in w hich it diff er s
fro m t he m are: pro m inent lob e (L3) considerably lar ger t ha n that in
K. ko zlowskii (Pribyl); absence of t ube r cles and edges on this lob e dur ing
ontogenic stages; presence of velar r id ge along the f ree m argin , with the
s ubv elar area , as a r ule, not exceeding 0.6 mm in width (fi g. 12). Anot h er
noteworthy element h ere is t he so- called admar ginal ridge (on t he r ig ht
valve) w hich , in a small nu m ber of ind iv id uals, dis pl ay s d istinct ten dency to p ro d uce admarginal t ubercles. On ev iden ce of its ontogen et ic
changes, this s pecies is belie ved independently to attain in the formation
of morpholog ical d etails results sim ilar to those of Ko zlowskiella k oz lowskii.
Paleozoo lo gical Laborato r y
of the Univer sity Of Warsaw ,
\V a rszaw a, Jan uary 1958
ONTOGENY
AND EVOLUTION OF KO Z I.OW SKI EL L A
107
BIBLIOGRAPHY
A DAMCZAK, F. 19'56. Polyzygia Giirich, an Ostracod ge n us from the Give ti a n of
t he H oly Cross Mo u ntains . - A ct a Palaeon t. P ol., 1, 1, 35'-48, Warsz a w a .
BAS SL ER, R S . & KELLETT, B. 1934. Bi bliograph ic in de x of Pa leozoic Os tracoda. Ceol. Soc. A me r., Sp ec. Pa p., I , Wash in gt on.
EL L IS, B.F. & M ESS INA, A. R l'952-1 9a7. Ca talogue of Ostracoda . - Amer. Mus.
N a t . H i st ., N ew York.
FASS B INDER , K. 1912. Be itr a ge zu r Kenntnis der Sii s sw a sse r os tr a cod e n. - Z ool. J b. ,
A bt. A nat. On tog. Ti ere, 32, '533-<a76, Jena.
GDR ICH , G. 1896. Da s Palaeozoicum im P olnisch en Mi t tel geb ir ge . - Ve r ll . Rus s.
K a is. Mi n. Ge s., 32, 1- 339 , S t. Petersb u rg.
HENN I N GSMOEN, G. 1953a. Classifica tion of P al e ozoic straight- hinged os tracods . N orsk: G eol. T i d sskr., 31 , 1:85-288 , Os lo .
1953a . The M iddle Or do v ician of the Oslo re gi on , N orw a y . 4 : Ostracod a . Ibid em , 32, 35-56 ,.
19I54a. L ow e r Ordovician ostracods from th e Osl o r eg io n, Norwa y .
I bi d em ,
33, 41- 68.
1954b . Uppe r Ordovicia n ostracods fro m th e Oslo region , N or w a y . - I bi d em ,
33, 69- 108.
1 95i4c. Siluri a n os tracods f rom the Oslo r egion , N orway. I : Beyrichia ce a . W ith
a r evisi on of the B eyrichiida e. - Ibi dem , 34, ,115- 711.
1955. A sho rt a ccount of t he ostracod fa mily Beyri chiid a e. - Mic ropaleontolog y, I , 3, 239-246, N ew Y ork.
H ES SL AND, 1. 1949. In vestigati on s of the L ow er Ordovician of the Si ljan d is tr ict ,
Sweden. I : L ow er Or do v ician ostracods of t he Silj an d is trict. - B ull. Ge o l .
I n st. Uni v . Uppsal a , 33, 97- 408, Up psala .
.JAANUSS O N, V . 1957. Mi d dle Ordovicia n os tracodes of Central and S outhern
S w ed en . - I bi d em , 37, 173-442.
KE SL IN G , RV. -1'95,l a. T h e m or p hology of ost racod m olt stages. - Il l. B i oI. Monog r.,
21, 1V3, 1- 324, Ur ban a , Ill.
195,lb. Me chan ical solution of formulas f or growth r ates .
C on t r . Mu s.
P al eon t . Uni v . Michigan , 8, 10, 231-237 , Ann. Arbor.
1951c. Te rminology of ostracod ca r a paces. - Ibidem, 9, 4, 93-171.
1952a. A stud y of Ctenoloculin a cicatricosa (Warthin). - I bi d em , 9, 8, 247-2g0 .
'195I2b. D ou bli n g in size of ostracod car a paces in each mo lt stage.
J.
Pal eon t. , 26, 5, 772- 780, Me n a sh a.
1953a. A beyric hiid ostr a cod from th e Midd le D ev on ia n W a na k a h s ha le of
western Ne w York. - B u ll. Buff. Soc . Na t . Sci., 21, 2, 1'9-24, Buff a lo.
1953b.
A new beyrichiid ostracod f rom t he Mi d d le Devonian Ro ckpor t
Quarry limestone of M ichigan. - C ont r . Mus. Pa leo nt. Uni v. Michigan , IU,
10, 221)229, An n A rbor.
'1953c. A slide rule fo r t he determi nation of in stars in ostr a cod species .
I bid em , 11, 5, 97- 109.
1957a.
A pe e l tech ni q ue f or os tracod ca rapaces an d s tr uc tu r es reve a led
th ere with in H ibbard ia la crimos a (Swartz & Oriel). - Ibid em, 14, 4, 27-4 0.
1957b. The on t oge n y a n d e col og y of Welleria aftonensis Wa r thin, a Midd le
Devonian os t rac od fr om th e Gravel Point formati on of Michigan. - Ibidem,
14, 5, 41-'~ .
1957c. Origin of bey rich ii d os tracod s. Ibidem , 14, 6, 57-80 .
108
FRAN C I S ZE K AD AMC Z A K
KE SL ING, R.V. & KILG ORE , J .E. 195Q. Os t racods of the f amilies L eperditeltida e ,
D r epa n ellida e , Gl yptop leu rid a e , Kloedene ll id ae , Ba ir d iid a e , B a r y ch ili n i da e
a n d T h ilipsurid a e from th e G ensha w' fo r mation of Michigan. - I bi d em , 10,
1, i-rs.
K E S L ING, R. V . & ROGERS, K. J . 1957. Size, lobation , velate st ructu r es a n d ornamentation in some bey richiid ostracods . - J . P al eont. , 31, 5, 997-1009, Menasha.
LEVINSON, S.A. 1 1 9~1. Thin se ctions of Paleozoic Ostraco da a n d their bearing or>
ta x onomy and m orphology. - I bi d em , 25, 5, 533-'560 .
MART INS SON, A . 195,5,. Studies on t he ostracode family Prim itiopsidae. B u ll
G eol. I n st. Uni v . Uppsala, 36, 1-33, Uppsala.
195:6. Ontogeny and d ev e lop m e n t of dimorph is m in some S ilurian os t ra codes.
A s t u dy of t h e Mulde marl fa una of Go tland. - I bid em , 37, 1-42 .
MUL L E R, G .W . 1894. D ie Ostracoden d es G olfes von N ea p el und der a ngrenzende n
M ee resab sch n itte . F aun a und F lora des Golfes von N e a p el , Monograp h ie 21,.
Berlin.
POLENOV A, E. N . 19515 . Ostrakod y d ev on a volgo - u ra lsk o] oblas ti. - T rud y VN I G RI ,
n . ser., 87, Leningrad.
. P RIBYL, A. UJl53 . Skor epat ci p ol skeho str ed nih o d evonu (giv etu) z profilu Grze gorzewice-Skaly w horach S vatokfizskych. Sb orn. U str. U st . G eol. odd.
p al e ont. , 20, 1-11 2, Praha.
S CHM IDT, E .A . 1941. Ostra coden a u s d en Bo hdalec-Schichten und tiber d ie T a xo n om ie d er Beyrichiacea . Abh. Senckenberg. N at uri , Ge s., 454, 1-9 '6,.
Frankfur t a . M .
SMI T H , M .L . 1956. S om e ostracods from t he Middle D evonian Ledyard Wanakah
mem bers of the Lud lowville fo r mation in wes te rn N ew Y ork.
J . Pa l eont. ,
30, 1, 1-8 , M en a sh a.
SOHN, LG. 1956. The transformation of op a q u e calcium carb on a te t o t r a nslucent
ca lcium fluoride in fossil Ostracod a. - Ibi de m , 30, 1, 113-114.
SPJELDNAES , N . 1951. On togeny of Bey r ic hi a jo nes i B oll. - I bi d em , 25, 6, 745-7515.
STOVER, L. E. 19516 . Ostr a code f rom the Wi ndom s hale (H amilton) of wes tern
New York. - I bi d em , 30, 5,11092 -1142 .
SWARTZ, F . M . 1'9'36. R evision of t h e Primatii dae an d Beyrichiida e , w it h new
Ost racoda f rom th e L ow er D ev on ia n of Pennsylv ania.
I bi d em , 10, 7,
54 1-586.
SWARTZ, F . M. & WH IT M O R E , J r., F . C. 1956. Ostracoda of the Siluria n De ck er
and Ma nlius limestones in New J ersey and e as te r n N ew York . - Ibid em ,
30, 5, 102'9-1091.
T R I E BEL , E. '1'9411. Z ur Mo rphologie und O k olo g! e d er f os silen Ostr a cod a , m it Besc h r eibung e iniger neuer Ga ttungen und A r ten . - Sen ck enb ergi an a , 23, 4/6,
294-400, Frankfurt a. M .
UL R I CH, E .O . & BASSLER, R.S. 1923. P aleozo ic Ostra coda , their mor phology,
classification an d occ u rre n ce . - Maryl and Geol. Surv. , Silurian vol., 271- 391,
Baltimore.
UPSHAW, Ch . F. , T O DD, R.G. & AL L EN , B .D . 1957. Fluoridization · of Micro fo ss il s. - J . P aleont. , 31, 4, 793-795 , Menasha.
Zt schr ..
' VE RW O R N , M . 1887. Zu r En t w icklungsgesch ich te der Be y r ichie n .
deut sch. geol. Ge s., 39, 27-311 , -Ber lin.
ZALANYI B . 1929. Morph u- systematische S tu dien tiber f ossil e M usche1krebse. G eol. H u n gar. , ser. palaeont., 5, 1-150, Bu d ape st. ·
ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA
109
FRANCISZEK ADAMCZAK
ONTOGENEZA I EWOLUCJA RODZAJU KOZLOWSKIELLA (pRIBYL)
(OSTRACODA)
Streszc ze nie
Material, na jakim oparto bad ania on toge ne zy i ewolucji rodzaju Ko zlo w skiella (P ri by l), zebrany zost a l w latach 1955-56 w srodkowym dewonie G6r Swieto k r zysk ich, w miejscowosciach Wydrysz6w , Grzegorzowice i Skaly.
Ko zl owskiella op isana zostala p o r az p ierwszy przez A. P ribyla (1953) jako Ul richia (Kozlowskiella). Autor, opie ra jac si e n a morfologii, on to ge ne zie i zmiennosci,
u zria l ten podrod zaj za r odzaj ni e zalezny od Ulri chia i, ze wzgledu na pewne podobiens tw o d o krumin alne go typu dimorfizmu (la c. crumena = torebka) , u m iescil go
w a ru nk ow o w ob rebie rodziny Beyrichiidae.
Na duzym m ate ri ale (ok ol o 3000 egzernplarzy fo r m d oroslych i m lod ocianych)
przesled zo no m . in. r ozw 6j on togenetycz ny w cza si e ko le jnych wylinek -
czyli zrniany, jakim p odlega skoru p k a
trzech n ow y ch ga t u nkow z e ifl u, (Wydrysz 6w i Grze-
go rz ow ice ), a mianowicie:
Kozlowskiella praetu ber cu lat a n. sp .
K ozl ow sk iella tube rcu lata n. sp.
K ozlow ski ell a similis n . sp.
i jed riego opisanego ju z wc zes n ie] przez Pribyla (1953), z rni ejscowosci Skal y (zywet),
ja ko : Ulrichia (Ko zl owskiella) ko zlowskii Pribyl. Za ob serwow ano tez zmiany , jakirn
po d le ga ja op isa ne gatu nki w procesie za r 6w n o on togenezy, jak ewoluc ji w eiflu
zy wecie,
J esl i ch odzi 0 posz czegolne stad ia rozwoju oso bniczego, to zos taly one wydzie lone, op iera ja c s ie na m etod a ch opr a cowa n yc h p rzez R. V . K esliriga (1953c) . A utor
te n -
n a p odsta w ie prawa Brook sa gloszace go, ze p o kazde j wyli nce skorupk a
po wieksza swoje rozm ia ry w stosu nku p r ocentowym, k t 6ry jest w przy blize riiu
.st a ty dla ga t u nku i p lc i, a k t6ry wy nos i 1,26 -
opr ac ow a l tabele w p osta ci cykl6w
Ioga ry trnicz n yc h, od powiada jacych teorety cznej wa r tosci wzrost u d la jedri e go wym ia r u, po wierzch ni i ob je tosci,
W ewolucji r od zaju K ozl ow sk iel la (Pribyl) a u to r w yroznit d wa szere gi ro zwojowe. Jeden reprez en t ow any
K. tuberculata n . sp . -
jest przez K ozlowski ella
pra etuber culata n.sp. -
K. kozlo ws kii (Pribyl) , a drugi przez jeden tylko ga tu ne k -
K. sim ilis n. sp .
Szereg pierwszy w y r ozmon y zostal n a podstawie przeobrazen podobnych element6w m orfologicznych skorupki, kt6re przesledzic mozna takze wsrod form mlodocianych (text-pl. I A-C). Stwierdzono, ze zmianom podlega m. in. plat L3, kt6ry
u formy wyjsciowe] (K. praetuberculata n. sp .) wykazuje na swej powierzchni tenden cje do tworzenia guzk6w juz od 6 lub 7 stadium. U K. tuberculata n. sp. ten sam
FRANC ISZEK ADAMCZAK
110
element rzezby jest wiekszy i pojawia sie na skorupce od 3 stadium. U K. kozl owskif
(P ri b y l), ktora ' za rnyk u ten szereg r oz wojowy (fig. Bc) , guzki na p lacie L3 poj awia ja
si e w 2 s tad ium i zani kaja po B lub 7 wylin ce (text-pl. I C ). W konsekwencj i t ych
przeksztalcen powstaje wydatny plat L3 . Z przeobrazeniamt t ymi skorelowane sa :
szer ok osc p owierzchni subwelarnej , ktora si e powieksza, or a z Iistwa a d rna r gina ln a,
przeksz ta lcaj qca si e w sze reg drobnych brod awek adm argin alny ch .
K . sim ilis n . sp. r eprezentuje drugi sz ereg rozwojowy . Plat L3 u tego ga tu n k u
jest du zy i w zasad zie p od obn y d o L3 u K. koz lows k ii (Pri byl); p oz a t ym ni e wy kazu je on g u zk ow na swej powierzchni. G atunek ten (K. sim ilis n. sp.) p owstal
prawd opod obnie w sz y bkim tempie , przy cz ym zas adni cz e zm ia n y d okonal y sie w e
wczes nych s ta d iach on t ogen e zy . U f orm ml od ociany ch plat t yln y (L 3) jest z r eguly
w ie ks zy, wy datnie jszy , niz u pozostaly ch ga tu n k ow,
Nawia zu ja c d o tyc h prze ob r azen a utor w y su na l w n iose k , ze, w ob reb ie [edne go
ro d za j u, p ow s tan ie po dob nych el ernentow (w yd a t ny plat L3 ) m oz e n astapic badz
na drodze p ow oln y ch , st op ni owych przeksz tal cen [K. praetuberculata n. sp. -
be rcu lata n. sp. -
K. tu
K. kozlowsk ii (Pfibyl j] , badz tez skok ow o (K. similis n . sp.). W w y -
ni ku t ych zmian pows ta je, oczywi scie, nowa jed n ostk a taksonom icz na w
r a n dz e-
ga t u n k u.
Oso b ny r ozdzial w
tej pracy
sta nowi
za gadnienie s tr u ktu ry sk or u pek
zm ia n , jak im one p odlegaja w proce sie ewolu cji, Chodzi tu
0
oraz
tz w . "c ie m ne sm u gi",
s tw ie r dzo n e na ci e n k ic h przekr ojach u rodzajow K ozl ow ski ella (P ri byl) i S accarcltit es
Swartz & Whitmore . Na skorupka ch gatunkow K ozl owsk iella stwi erdzon o je w zd luz,
w oln e go brzegu skorupki, w postaci ciern n e go pasma , or az n a wyniosly ch e le meritach si atki twor zac e] rzezbe. Struktury te p oj a wiaj a sie juz od wczesnych st a d iow
rozwo ju on togenet yc zn e go, p oczatkow o w str efie w olnego brzegu , a p ozn iej , stop n io wo, w po zo st a le j czesc i skorupki (fig. ,115). U Saccar ch it es ornat iss im us (G u ri ch)
"cie mne s rn u gi " p oja wi ajq s ie w p ozriiej sz y ch s ta d iach on to genezy, lecz w id en t y czn y
jak u K ozl ow sk iell a spo so b. Zdaniem a u to ra, nie je st wyk luczone, ze me ga one m iec
pewien zwiazek z procesem ka lcyfi kacji skor upki.
Na p od sta w ie p e wny ch an al ogii r zezby i stru k t ury sk orup k i a u to r przypuszcza,
ze fo rm wy jsc io w ych dla r od zaju K ozl owski ella n al e zaloby p oszukac w obrebi e ga t unkow rodza j u Sacca rchites. J e d yn y przedstawiciel tego ro dza j u, zna ny d ot y ch czas
7
Polski, opisany zostal p r ze z G. G liricha (1953) jako Pr imit ia orn at issi ma. P r zy
blizszym po row n a n iu okazalo sie , ze ga t u ne k ten w yka zu je duzo a na l og ii z p r ze d stawicielami rodzaju S accarchites i z tego powod u u m ieszczono go w ob rebie tego
wlasn ie r odzaj u.
Morfologiczn ie bi orac, stanowisk o p osrednie rni edzy ty mi r odzaj arn i za jm u je
A?JCl rchitellin a P olenov a . S zcz eg olnie jeden ga tu nek tego r odza j u, A . cf. d ecorata ,
k t ory ot r zy m alem od Dr E . N . P ol enowej w 1957 r ., wyk azuje duzo a n a logi! z ob u
wymienion ymi rodzajami. D ef in it y w n e rozstrzy gniecie tego zagadnienia wyrnaga
j ed n a k gr u n townych studiow porownawczych, zwlaszcza struktury skorupki.
ONTOGENY
AN D E VO LUTION O F
KOZLO W SKIELLA
I II
O B J AsN IEN IA DO I L US T R AC JI
Fig. 1 (p , 76)
Elementy r zezby p ow ierzchni skor u p ki ; S l bruzd a p rzednia , L2 pl a t srodkowy ,
S 2 b ru zda me dialna, L 3 plat tylny , t gu zki, vr lis twa wela r na.
Fig. 2 (p. 76)
P r ze k r 6j poprzeczny p r zez skorupke samicy K praetuberculata n . sp., u proszczo n y; 1 lis twa welarna, 2 lis twa a d marginalna, 3 a rea subwelarna, 4 fald d ors a ln y ,
5 bruzda zawiasowa, 6 od ci sk miesnia .
Fi g. 3 (p. 77)
K. ko zl ow skii (P riby l), skoru pka prawa od strony wewnetrznej (rysunek nieco
sch em a t yczny ); 1 b r u zda zawiasowa , 2 listwa, .1 li s tw a w e la r na, 4 brodawki a dm argi n a lne.
Fig . 4 (p. 78)
P rzekr6 j p odlu zn y pr zez skorupk e prawa K. ' praetu be rculata n . sp . (r y sune k
sc h em a ty czn y ); 1 k analy p orowe , 2 ciernna smuga , 3 odcisk mies nia , 4 listwa a dmargina l na, 5 li s t w a we larna.
Fig. 5 (p . 80)
K. praet ub erculata n . sp., skorupka prawa sfluoryd yzow an a; X 40
F i g. 6 (p . 8 1)
Stosunki pokrewienstwa mied zy gatunkami rodzaju K ozlow sk iella (Pribyl)
w srodkowyrn dewon ie G 6r S w i et okrzy sk ich : A K. praetuberculata n . sp., B K . tu ber cul ata n. sp. , C K. k ozlowskii (P ri by l), D K. sim ilis n . sp. (sk o r u p k i lew e).
Fi g. 7 (p . 82)
A - W ie lk osc skorupek d oroslych (k om pl etn y ch i lewych p oj edyn czych) K . pra etu be rcu lata n . sp. w trzech r oznych pozioma ch w Wydryszowie. Na osi r zedn y c h dlugosci skor u p ek w mrn, na osi odc i et ych - pr6by z kolejnych warstw.
B - Procentowy u dzial form dorosly ch K . praetuberculata w trzech rozny ch
p oziomach w Wydryszowie. Na osi rzednych - li czba skorupek lewych i k omple tnych w %°/", na os i od cietych - wielkosc w mm.
Fig. 8 (p , 8 2)
S t osunek dlugosci d o wysok osci skorupki w p oszcze g6ln y ch stadiach ; a K. praetuberculata n . sp. , b K. tube rcul ata n. sp., c K. ko zlowskii (Pribyl), d K . sim ilis n . s p .
Fig. 9 (p . 83)
Procentow y u d zia l t r zech gatunk6w Ko zlowskiella (P ri b yl) w sze sciu pozioma ch
Wydryszowa ; A-F k olej ne p ozi omy, a K. sim ilis n . sp ., b K . praetu bercu lat a n . sp .,
c K . tube rculata n. sp.
Fig. 10 (p . 84)
P rocentowy u d zial trze ch g atunk6w K ozlowskiella (P rib y l) w pieciu pozioma ch
Gr zegorzowic; G- K kolejn e poziomy, a K . sim ili s n . sp., b K . praetuberculata n . sp.,
c K. tu berculata 'n . sp.
FRANCISZEK ADAMCZAK
112
Fig. 11 (p. 85)
Przekr6j poprzeczny przez cZE:SC wentralna skorupki prawej (rys. schemat.) ;
1 K. pra etuberculata n. sp. ; :2 K . similis n. sp., 3 K. tuberculata n. sp. , 4 K. ko zlowskii
( P ri by l), r li stwa admar gin a ln a, t brodawki adrnarginalne.
Fig. 12 (p . 85)
Szerokosc powierzchni subwelarnej w poszczegolnych s tadia ch rozwoju osob ni czego: 1 K. p raetub ercul ata n. sp., 2 K . k ozl owsk ii (Pribyl), 3 K. si milis n . sp .,
4 K. tu bercu lata n. sp . N a osi r z ed n y ch - szerokosci w mm, na osi odcietych stadia .
Fig. 13 (p. 85)
P rocentowy u d zial d w och gatu n k6w K ozl ow skiella (Pribyl) w pi eciu rozn ych p oziomach e iflu w Grzegorzowicach ; d K. prae tu ber cu l at a n . sp ., e K. si mi/is n. sp . N a
{lsi rzedny ch - ilosc osobnik6w w OfoD/D, na osi odc ietych - kolej ne p ozt om y.
Fig. 14 (p . 86)
Wymiary skor upek form d ojrzaly ch K. k ozl ow sk ii (P ri b y l). Na osi rz ed nych d lu gosc. K w adra cik a m i ozn ac zono samice,
t r ojka ta m i - sa mc6w.
wysokosc w m m , na os i odcie tyc h -
Fig. 15 (p . 87)
P r ze k r 6j podluzn y przez skorupke p r a w a K. k ozlowsk i i (P ri b y l) (rysunek sch e m a tyc zny ) ; 1 ka naly p orowe , 2 ciemna smuga, 3 od ci sk m iesn ia , 4 brod awki a dm a r gi naln e, .5 li stw a wela rn a, 6 L3, 7 S2, 8 L 2.
Fi g. 16 (p. 88)
Fra gmen t p ow ie r zch ni s k or u p k i K . praetuberculat a n . sp. (rys u n ek n ie co sc he matycz ny); pc k ana l po rowy, dl ciem na sm u ga .
F ig . 17 (p. 89 )
Prze kr oj e p oprzeczn e p r zez tr zy pi erws ze s ta d ia K . praetube rcu lat a n. s p . (ry
s unek sc hem a tycz ny ).
F ig. 18 (p . 90)
Grafi czne p r zed sta w ie n ie w y m ia r 6w i lic zby sko r u pek w p os zczeg6lnych sta d iac h rozw oju osobnic zego K. praetube rc ulata n . sp . Na osi rz ednych - lic zba, n a
os i odcie tyc h - w ym ia r y skorupek w m ik r on a ch .
F ig . 19 (p. 96 )
Wymia r y skorupek for m do j rza lych K. p r aetubercu la ta n . sp . Na osi rzed nych dlugose. K wadra cikami ozn a czo n o samice,
wysokosc w mm, na osi odc ie tych t r ojka ta m i - samc6w .
F ig. 20 (p . 97)
Procentowy udzi al skorupek K . praetuberculata n . sp . w p osz czeg6ln ych s ta d iach .
Fig. 21 (p . 100)
Graficzne przedstawienie stosunku dlugosci skorupki do szerokosci p owierzchni
s u b w e la r ne j; 1 K. praetuberculata n . sp ., 2 K . simi/is n . sp ., 3 K . tuberculata n . sp.
ONTOGENY AND EVOLUTION OF KOZLOWSKIELLA
113
Fig. 22 (p . 100)
Proeentowy ud zia l sk onup ek K . tubereulata n . sp, w poszczegolny ch s ta di ae h .
Na osi rzednych - li ezb a osob n ikow w "!oOfo, na osi od cietych - st adia .
Fig. 23 (p . 103)
K. koz lowsk ii (Priby l), diagram zm ien n osc i, N a os i rz edn ych udz ia l w p osz czeg ol n y ch k lasae h .
p r oeentow y
Fi g. 24 (p . 103)
Grafiezne prze dsta w ienie w yrniar ow sko r u pek 4 gatunkow rodzaju K ozlowskieU a
(Priby l) w srod kowy rn dewonie Gor Swietokrzyskich. Linia przerywan a zaznaezon o
p rz y pu s zeza lne zmia ny wie lk osci w gornyrn eiflu i dolnym zywecie ; 1 d olny eifel,
2 go r n y ei f el , 3 d olny zywet, 4 gor n y zywet.
Fi g. 25 (p. 105)
Wymiary skorupek fo r m d ojrzal yeh K. si m ilis n . sp. N a os i r zedny ch - w y s ok osc w mm, na osi odc ietych - dlugosc, Kwadracikami oznaezo no sa m iee, tr oj k ata m i - samcow .
Te xt-PI. I (p . 90191 )
Rysunek (n ie eo se he ma tyezn y ) rozwoju osobniezego skorupek ; A K. p r aetu be r.ell /at a n . sp ., B K. tub erculata n . sp ., C K. kozlowskii (Pribyl), D K. si m ili s n. sp .
PI. I
K ozl ow skiella praetuberculata n . sp .
Fig. 1. H ol ot y p , skor up ka sa m iey: a le w a skoru p ka , b s tro na d or salna , c st rona
w e n tr a lna ; X 23.
Fig. 2. Skorupk a sa mc a: a lewa skorupka, b s trona w e ntra l na ; X 23.
Ko zl owsk i ella si m i/is n. sp .
Fig. 3. H ol ot y p , skorupka sarnicy : a lewa skorupka , b stron a d orsaln a , c st r ona
w e n tr a ln a ; X 23. 5. Plat L2 na prawej skorupee u szkodzony.
Fig. 4. Skoru p k a sa mea : a le w a skoru p ka , b stro na d or sal n a; / 26.
PI. II
K ozl ow sk i el la kozlowskii (Pribyl)
F ig. 1. Skorupk a sa miey : a le w a skor u p ka, b strona d orsal na , c strona wen t r a lna ; X 23.
Fig. 2. Skorupka sa mea : a lew a sk orupk a , b s tro na w e n tralna ; X 23.
K ozlo ws kiell a tu berculat a n . sp .
Fi g. 3. H ol ot y p , skor u p ka sam icy : a le w a skor u p ka, X 2'2; b s tr on a d orsalna ,
c s tr on a wen traln a p a raty p u, X 23.
Fig. 4. Skorupk a sa mea : a lew a sk orupka , b strona wentra lna ;: X 22.
PI. III
K ozlow sk ie l la praetu bercu l ata n. sp., X 32
Fig.
Fig.
Fig.
1- 8. Stadia mlodociane I-VIII
9.
Skorupka samea.
10.
Skorupka samiey.
A c t a Palaeon t ol o glca Polon lca -
vol. IIIj 2
8-
F R AN CISZEK AD AM C Z AK
114
P I. IV
K ozl ow sk ie Zla tube r cul at a n. sp ., X 32
Fig.
Fig.
Fi g.
1-8. Stadia ml od oci an e I-VIII
9.
Skorupka sam ca.
10.
Skorupka samicy.
q,PI\HI..(HWE K I\ AJ;lM4I\K
OHTO fEHE3 H 3BOJIl0lJ.HH POAA KOZLOWSKIELL A (P RIBY L) (OSTRA COD A }'
Pe310Me
Marepaaor,
lroJ ,l \Or Oj)l' M
Me lf lUIBOCTb IO ip<l!,l\a
IJIPOH3BD,nHJlHCb IHiCCJIe ,l\OBaHHH IHa A
OHToreHe30M
H 1li3-·
K ozl owskiell a (Pfibyl), 6bIJI co6paJH B 1'9155-56 ronax H3 cpemte ro-
lleBOllia COOI~PIKHORIiX f o p
B uvreC'\1l1OCTIDC BbI\lWblIIIYS, DpiKeropmOBHI..\e H CKaJIbI .
K ozlo wskiell a 6bIJla snepasre OIJJHcaHl..l iITpJHJH6bI JleM (A. Pribyl, H~5i3 ) KaJR Ulrichi a.
( KozlowskieZla). OnHJPa'HChH:l Mopr!lOJlOrlHI, oirrorenese H H3MeWW"BOCT;i
pon , a
3HaJI IB IHleH IHe3 a BH,CIHUVlbI H
AH MOiIXPH3Ma
(crumena
1H3 - a a CXOAC11SClJ AO T.
JIaT. c.yMJRa) nOMeCTHJI
ee
aarop
ThpH-·
Ba3. crtPYMHlH3JJ1b1HOro 'J1HTIa
YC.JlOBHO B
rrpen enax
ceMeHC11Ba'
Beyrichiidae.
H a 6oJl bIIIOM MaTepIHalJle (OROOO 3000 3 R3€1MTIJIHIPOB MOJIOAbIX HB3p0CJ1 bLX rpOPM)
rrpoonear en :Me *J(y npotIHJM onroreaeaac raeroa Ip<1JROBm JlU ,B retI€lHHe osepenusix JlHHelR npyca ( B bI~P bIIllY;B iii I{pjHeropiKOBHI..\e):
-r.e.
6hIJI
1H3MeHeJ-I1HH, HOTOpbI M
IIOABep-
'!1Pex IH 0i9 bIX BHAOB f13 -:3iJ.rpeJIbCIRoro·
K ozl ows kiella praetuberculata n. sp,
K ozl owskie lla t uberculata
n.
sp.
K ozl owskiella si m ilis n. sp.
Ii
omroro
IlIHAa
(iH:H Be l'Cllm H
B bI JIH
'JlIp y c)
H3 yq BiH bI
otrroreneaa
onacasnroro
[lOA
YJHe
pannsure
IHJa3BalllHe.Vl
U3MeHJeHH.H;,
IRa,KHM
TI p IKH6 bI JleM
(19 53) 1li3 IM BCT'HlCJC11H ORaJIbt
Ulrichia (Kozlowskiell a)
[l0AOOpraIO TruI
OiIliHCWHiHble
kozlo w sk ii
BHJAbI
IB
P fibyl.,
nponeceax
H 3 BOOIOI..\HH 18 '!1eIQeHlHe 3H¢eJlH IH JKHBeTa.
LI TO R:aJCaeTCH OT,l\eJI bHbLX CTMeH IlJIH AllBHiAY aJI b OOro Ipao.!lIliTlliI , 0lIIH 6bIJm iBbI,l\e Jl€lH bI l}{a OOHOBa'H1l:H M eTOAQB o6pa6oTCliHlHlb IX K ec.JIHH roM (R.
OOH'OBaIHIH'H 3<1JI«)HlCl.
Bpy;IOC:a
(Brooks) , COrJlOOHO
xoropcay
V.
Kesling,
paaxeps; B IIJIp oI..\e Hl1HOIM COOTHOIIIelllHH 1 nOCJIe KCltiK~OH .rnHlHbRH, 3TO'I' asrop
n6e.rrH
B
B:HAe
JIompmpMHQeOOHX
~HRJIOB,
1953c). H a
.p<lJOOBHI IJRal yBe.lLJNHBaer
OTOOtIalOI.I.I;1iX
TeOpeTHtIOOROH
csoa
06pa60TaJI
e eJDH'LH He'
POCT<lJ ~!'fl o n a o ro tH3M€lpeHlHJH ~HJalJlp. MH:HbI), IlOOOpXHOCTtH H 06oe~m .
1
CooTHOIII€lHHe <Yl'O, ' I]pIH6JIIH3iH'J1eJIbiHO nOCTOJlIlI1HO ~TJ:H RHJ\a H
n ona , PClJBllIO
1,26.
ONTOGENY AN D EVOLU T I ON O F
B 3BOJIIO~HH I~a Ko zl owsk iella
no
K . praetu berculata
esomouaonnue
K. t ub erc ul ata
n . sp . -
(Priby l), 'JI:pyroii TOJIb KO O~I1 liM RH~o.Vl -
K. ko zlowskii
sp, -
115
aBTOp BbI~e.llJlliJI ABa
(Pfibyl)
PJI,AbI. O~ ·H3 HiiX npencraenea I9l~AaMH:
KOZ LOWS K I ELLA
K. similis n . sp .
O e p Bbl H PH~ 6bliJl Bbl.r\e.leH ,Ha OCHOBaJHIHII CX~I1 bI X fllpeoQpaJ30B3JHlI ii M 0IXPoJIOtmf-
lfeCKII..X
3 JIeMe HTOB rpaK'03IIII bl , A a IOIlUiX>CfI
(TeKCT-IUI.
I A - C) . Bsino
La
(K. praet u bercul ata
'IIIM
JIOIlacT b
Ha'IlillJl3iH C
6
sp.)
l o b e) ,
,KOTOpaH
TeHAeH I.\1II0
JIrpoRBJIR eT
K OTOp aiH
aaersrxa er aror
nOH BJIH IOTCH '8 0 2-oH CTall\ HJII
B Ipe3YJI bTa~
c
H
<pOPM
rtp o-
nc -reaaior
iIlJpeo6pa30B8IHlIaMH
<PoP;\lbl
TOT 'Ke 3JIe.' l e IlT oKyJl hIIryp bl
rpM
3 BOJIl OU.IIOHiHbIH
rrOCJIe
3111IOC JlIpe06pa3 0 B<lJHIHiiBoo H%Ka eT
YlIOMJJIlIYTblMH
HCXO~HOH
y
66JIhlllllX ,pa3MepoB III 'II OJI,B.1H€lJ'CH ,Ilk1 'pwKOBI!Hre 11Ia 'I,HJHa H C 3-eH
(Pr i byl ),
IOHbIX
06pa30BaHIIH 'I1a n ooopx!IIOC11H 6yropHoo yme
7 ClJ'8J~HJII. Y K. tuberculata n . sp .
.H M
open a
:r<lilnlte
yC T8JHOBJIeHO, 'Iro 113;\le He INUL.VliII IIoAOOpraeTC:H Me?f\.AY
(posterior
no
IIlip oc.1 e.AJIT b
COC'1'OHT :
6
CTa'~II. K. kozlowskii
6 C) ,
(eplIr.
U JIH 7 IJ"IiHH blKll
(T eKCT -ff1J1.
La
I C) .
La. B IKO~pe.JIH u.lIlI
B blPa3IHTeJ1b1HUH
WHJplIIIa
6yropKil IHa
noaepxnocm
'oy6 B€ J]ffp HOti
(s u b vel a r are a) , HOTOPaH yBeJIll'I'HBCl,eTOH, H Cl'J\:MClipM IIaJIbiHatH ,Ka H.1lm (a d margi nal rid g e) ,
HOTOP<lR
rrpeepa utaeroa
B IpaA MeJI~llac Cl~Map[1IUIJClJIblHIhLX 6opO.rl<lillc IK
(adrna rgina l
tu-
b ercle s) .
,npe~cTaBJI H€'I'
K . si milis
II LB 'iI'PHHl.I,HIIO 'COCO,ll ll<lJ C
aropoii
ropsos, BllA K. similis n. sp.
La
3 Bomou.lloHH blH :PHA .
L a 'Y K. ko zlowskii
(Pribyl) ; Ha
aroro
y
ee
13OOHl",tK Berp0HllH10 6b!JCl1PO, IIlIpH'Ie..'1
lMeHbl 'f ip0II30 W JIH B IpaHHHX CTa:.rI;llHX OHTODeHe381.
-BllAa 60JI blllaH
'IlO OOP~HICl C1111 H€'I'
6y -
o cno easre nepe-
Y IOH!bIX <PoP ;\l 3MIIH'JI .JI0IIaCTb La'
H<lff\ npaBH JIO, 6 0 JIbWe ill 'BbI 1pa31f"reJI l>lIe e 'I eM y OCT<lJJI b HlhIX J!ill):\OB.
Ha
OOHOBaJlflIH
3TJLX
rrpecopaaoaaaa a aarop
B b~BOA,
:/(e JIaeT
'ITO
oOpaI3O BaHllIe
npen enax onaoro pona .IlIOmeT IlIpOHCOCo-
CXO;{HIhIX SJIeMe Ill1'03 ( BbUp aGlITeJI b HaH La~ B
~I:lTb JIiH60 JIY11€M Me~JIeH>HbLX Il~IleH>lIbIX
berculata n. sp . - K . tub erculata n.
HO
(K . simili s
n.
sp . -
n peoopaao saean (Kozlowskiella pr aetuK . kozlowskii (Pfi bylj) , JIIl60 OKa 'I~oo6pa.3­
sp.). B ipeayn srare STIIX ilI3MeHeHllI H r..03lDIIJlm e T O'IeBII;AHo ,HOBaR
rfCllHCOHOMll'I ecKa'H €ll\HiHllI u.a B Ip8iHDe BlIAa .
r viaaa CT<lJTbiII noonmuena ,BOIJIPO'C'Y Cl1p J'IR'J'Yip bl IpaiK OBIIH iII 113MeHeHJIH;\l,
fi OTOP bl M OIIIIl nonsepraiorca B npouecce 3BO JI101lilI1I. A e.rro snecs 'B ,,~H!bIX rton ocax "
(dark line) 'KOHOTa m poBallllI blX na roHi~X paapeaax ypDAOB K ozlo w sk iella (Pribyl)
II Saccarchites Swartz & Withmore. Ha Ip<lJKO:BIIIH8JX Bll~O B K ozl owskiella OHllI 6blJlH
OTl!\eJIbHaR
yC TaHOBJIeHbl iB~OJI h 'ClIloOO,ll HOI'O iKpaH IpaiKOBIIIHbl n
l jalOIl.I,HXOH S.JIeMeHTax CeT~1l
oopaaytouief
;B IIAe
CKy .TJ blnTYiP Y .
TeJ,mOH IlOJIOCbl ill 'lila
a11lI
s sr-
Cl1P'YK11Y'P bl n OBlBJ1HIOTCfl
'Ha ttIIlIlaH C IpaHHIlIX CTa:JlillH ouroreaeravecxoro pa3BH'IlIIiH, [lepOOIIl1.'lCliJI I>HO B WHe CBO-
6o~Horo
ecpaa,
ornatissim us
OtITOreHe3a,
a ::>aT€LVl, n o crerrenao, B ocran sao a '1aCTll PCliKOBllJlIbl (etm r.
( G iir i c h )
HIO Ta ,KIIM
"Te M Hble
me oopaa ocr W U\
I!CK.JIIO'IaeTCH B03 MOII\HlOCTh CBH3H
KOBlllI'bl .
.rrOJIOCbl"
n OHB.JIHlOTCtH
,B
60JIee
y K ozlow sk iell a. ITo
STOro HBJleHIlIR C npoueocov
:Il03 ~HlliX
15). Y S .
craansx
MH€llilHlO a s ropa
He
K a~l b ~HqlilIiKau. II.II p a-
FRANCISZEK ADAMCZAK
116
Ha
OOHOBaW lIH ;lle KOTO p b I X :3.1e 1ieHTOB CiKJyJIb nTYipb I
U peo./¥lOJIaJI'aeT , 'ITO HCX~HbIX <t>opM .IVl H
yp ~
8 !fil(OIS IPO~U
flj:Je ,nc T aBWr e .'l b
aa rop
OJIeIlY e T H OKa n ,
Saccarchites Swartz & Whitmore. ~HHOI1BeIHIH bI H , no OH.X n o p
:3TOl'O p ~ a iH3 BeCTlHiblii iH3 ,TIOJI billIlH 6 blJI OIliHC<lJH f lO P HXOM (G. Gtirich,
19'53i) n on IHa3BaJll1He~1 Primitia
(-11\Oro iBH;J,a
H C'I1PYih-ryp bI Ip a KO BH'H
'POJl,<1J K ozl owskieHa (P ri b y l)
or n atissi m a.
OK OOUJI OCb , 'I TO 011 .n poHBJIHe T
rL P H
~UHOro
oOJIe e-
BlFHU"raTeJllilF()tM
1H3y 'Ielt :111
<l<H1aJI OI1J1H C np e,ncTaBH re:I€l~I IP OJP
Saccar ch i t es 'H no aroii npH'I H'lle 011 ;J,OJIOOOll 6 bIT b n o neuien B rrpenenax H1M eHllO
:11101'0 poaa .
C
MOpepOJIOrHY OOKO i'r
'PO~<l!MlI 3Ull lHiMaCT
TO'IK;H
3peHIHH,
Apar chit el lina
A . d . deco r ate , npe.n;oCT <lJB JI€lHlF bl i i
",moro
allla'JIOnH:H. IJlO 0 11110 1llell IHI O iI\
u p o MenilyTO 'lHy lo
0601H iM .lJbllU e HClJ3IBaIIlUl bIIM
~1€o}h:1lY
n03HLlM JO
Polen ova. B OCoOeH HlC)CT H
3 BTO:PY E. H. IIo.: le Ho 30 ii
O~H
B
1957
ponaoi .
B Joq
:31101'0
:3rn ~1H
PO;1,11 ,
roay, lliPO~BJIHeT
OKOII'lafl'a'l b:HOO
paapemenae :3TO I1: np06: I€ Mb I 'l'p e o y e T o :J/lIWCO OCll IOBUTeJIlm blX c p amliHTCJl lill hl X HOClTlC ;J,o BaHlHH, ocoeeerao IlHlJi.\ C TpyIICTY'j}C)i'r p allW BWHbl.
PLAT E S
EXPLANATIONS OF PLATES
PI. I
Kozlowskiella praetuberculata n. sp,
Fig.
Fig.
l.
2.
Holotype, fem a le : a left valve, b dorsal view, c ventral view; X 23.
Male : a left valve, b ventral view; X 23.
Kozlowsk iella si m ili s n. sp. :
Fig.
3.
Fig.
4.
Holotype, female: a left valve , b dorsal view, c ventral view ; X 23.5.
L2 , on the right valve, d amaged.
Male: a left v alve, b ventral v iew; X 26.
PI. II
K ozl ow sk iella k ozl ow skii (Pribyl)
Fig.
Fig.
2.
Fig.
3.
Fig.
4.
l.
Female: a left valve, b dorsal view, c ventral view; X 23.
Male: a left valve, b ventral view ; X 23.
K ozlowskiella tuberculata n. sp ,
Holotype, female: a left valv e, X 22; b dorsal view, c v e n tr a l view of
para type, X 23.
M ale : a left v alve, b ventral view; X 22.
PI. III
Kozlowskiella praetuberculata n. sp ., X 32
Figs. 1-8. Instars I-VIII.
Fig.
9.
Male.
Female.
Fig. 10.
PI. IV
K ozlowskiella tuberculata n. sp. X 32
Figs. 1-8. I ns ta r s I-VIII.
Male.
Fig.
9.
Female.
Fig. 10.
A CTA PALAEONTOLOGICA POLONICA. VOL . III
F . ADAMCZAK . PL . f
ACTA PALAEONTOLOGICA POLONICA , VO L . III
F . ADAM CZAK, PL . II:
AC T A PALAEONTOLOGICA POLONICA . VO L . III
F . ADAMCZAK, PL. III
ACTA PALAEONTOLOGICA POLONICA, VOL. III
F. ADAMCZAK. PL. IV