DEVONIAN TRILOBITES·

EVOLUTION AND EVENTS

IvoCHLUPAc

CHLUPAc I. 1994. Devonian Trilobites - Evolution and events. [Trilobites devoniens - Evolution et evenernents
stratigraphiques] . GEOBIOS , 27, 4 : 487-505. Villeurbanne le 30.08.1994.

Manuscrit depose le 16.03.1993 ; accepte definitivement le 06.07.1993.

ABSTRACT
Environmental changes emphasized in event-stratigraphic turnouts are thought to have clearly affected the deve-
lopment of Devonian trilobites. After the latest Silurian /Pridolian! and earliest DevonianlLochkovian diversity
fall, the Lochkovian-Pragian Boundary Event triggerred the increased diversity period which lasted with some
changes until the early Eifelian. The general decline in trilobites, marked by stepwise diversity falls, was strongly
accelerated by the Middle Devonian Kacak and Taghanic, and the late Frasnian Kellwasser Events. The progres-
sive Famennian innovations were interrupted by the Hangenberg Event at the Devonian-Carboniferous boundary.
The retreat of Devonian trilobites was stepwise, event-stratigraphic turning points were lethal mostly for families
showing a decline already prior to the events. Stratigraphic levels of changes in trilobite faunas roughly corres-
pond to those recognizable in some other fossil groups (e.g. ammonoids , dacryoconarids , conodonts etc .), although
the effects could have been different. The general retreat of trilobites is believed to have been not caused but only
accelerated by event-turnovers.
KEY-WORDS : TRILOBITES, DEVONIAN , BIOSTRATIGRAPHY, EVOLUTION, EVENTS

RESUME
Les changements du milieu , accentues aux limites des evenements stratigraphiques, ont influence d'une maniere
considerable le developpement des trilobites devoniens. Apres la decroissance en diversite au sommet du Silurien
superieur (Pridoli) et a la base du Devonien (Lochkovien), l'evenement d'age praguien basal a ouvert une periode
de diversite elevee, qui s'est poursuivie avec certains changements jusqu'a l'Eifelien basal. Le declin general des
trilobites, marque par des chutes graduelles en diversite, a ere fortement accelere par les evenements du Devonien
moyen de Kacak et de Taghanic et l'evenement de Kellwasser du Frasnien tardif. Le retablissement graduel au
Famennien a ete interrompu par l'evenement de Hangenberg a la limite Devonien - Carbonifere. Le recul des
trilobites devoniens a ete graduel. Les renversements des evenements stratigraphiques ont ere particulierement
funestes pour les families qui avaient deja amerce leur declin avant les evenements, Les changements dans les
faunes de trilobites se situent a peu pres aux memes niveaux stratigraphiques que ceux qui affectent d'autres
groupes de fossiles (ammonoides, dacryoconarides, conodontes, etc.) ; toutefois chez ces derniers les effets de ces
modifications peuvent etre differents, Le recul general des trilobites n'a pas ete provoque, mais seulement accelere
par les renversements des evenements.
MOTS-CLES : TRILOBITES, DEVONIEN, BIOSTRATIGRAPHIE, EVOLUTION , EVENEMENTS

INTRODUCTION extinctions occurred there, the development of

Trilobites as a worldwide distributed, richly di-
versified, and extensively studied group of Paleo-
zoic fossils are a suitable object for event-strati-
graphic and ecostratigraphic research. Although
the Devonian represented already a period of de-
cline in trilobite evolution some significant
trilobites throughout the Devonian reflects com-
plexity of interactions between organic and inor-
ganic changes.
Devonian trilobites are in advanced state of stu-
dy in many areas of the world. Since their tho-
rough biostratigraphic review by H. Alberti
 488
(1979), a marked progress in stratigraphy, classi-
fication and palaeoecology of trilobites made a
new evaluation topical.
Devonian stratigraphy was defined more precise-
ly by international acceptance of new series and
stage boundaries and improvement of zonal sub-
divisions (review : Oliver & Chlupac 1991). The
concept of Devonian event-stratigraphy, eustatic
sea level changes and cyclicity was introduced
(e.g, House 1983, 1985, 1989 ; Walliser 1984,
1985 ; Johnson et al. 1985 ; Johnson & Sandberg
1988 ; Sandberg et al. 1988 ; Chlupac & Kukal
1986 ; Boucot 1990 ; Bayer & Ghee 1989 ; Schin-
dler 1990). The knowledge of Devonian trilobites
was substantially extended by numerous syste-
matic studies including those of Arbizu (1979),
Eldredge & Branisa (1980), Snajdr (1980, 1987),
G. Alberti (1980, 1983), Lutke (1980, 1990), Pri-
byl & Vanek (1981, 1986), Cooper (1982), Morza-
dec (1983), Smeenk (1983), Thomas & Holloway
(1988), Yuan (1988), Wenndorf (1990), Zhou &
Campbell (1990), Tomczykowa (1991), Feist
(1992), Ellermann (1992). Also some reviews, e.g.
Maksimova (1978), Thomas et al. (1984), Pek &
Vanek (1989), Strusz (1989), Kowalski (1991),
Spassky (1992), and evolution studies (Feist &
Clarkson 1986 ; Feist 1991 ; Morzadec 1992) con-
tributed to the evaluation of Devonian trilobite
development.
The number of recognized genera increased from
220 reported by H. Alberti (1979) to more than
300 discussed in this paper. However, the know-
ledge of Devonian trilobites is far away from
being ideal : the stratigraphic ranges of many
taxa are not sufficiently known, the correlations
of trilobite occurrences with conodont and other
zonal subdivisions are often doubtful or lacking,
except for the Upper Devonian. Furthermore, the
classification also offers many open questions,
particularly regarding diverse approaches to
ranks of taxa (genera-subgenera, subfamilies, to-
pical especially in Proetida),
The present paper is aimed to summarize data
on development of Devonian trilobites on generic
to family levels, with regard to Devonian event-
stratigraphic turning points and trends.
THE SILURIAN-DEVONIAN
BOUNDARY AND THE
LOCHKOVIAN
The level of the Silurian-Devonian boundary as
internationally defined in 1972 (McLaren 1977),
is markedly expressed in the trilobite biostrati-
graphy : very close above the boundary an explo-
sive worldwide distribution of the Warburgella
(Rugulites) rugulosa (ALTH) group is charac-
teristic of the lowermost Devonian persisting
through the whole lower Lochkovian (equivalents
of the Monograptus uniformis Zone, Chlupac et
al. 1972 ; G. Alberti 1975 ; Ormiston 1977). An
abrupt onset of the Warburgella (Rugulites) domi-
nated assemblages in various, mostly calcareous
facies in different continents clearly indicates the
bioevent nature of the Silurian-Devonian bound-
ary level.
The group of species around W. (Rugulites) rugu-
losa, however, has its ancestors in the Silurian
and Warburgella proper has evolved since the lo-
wer Llandovery (Yolkin 1983), stressing the per-
sisting Silurian aspect of lower Lochkovian trilo-
bites. Genera extending from the Silurian into
the Lochkovian are numerous and include e.g.
Coniproetus, Decoroproetus, Lacunoporaspis, De-
coroscutellum, Paralejurus, Weberopeltis, Kettne-
raspis, Leonaspis, Radiaspis, Dudleyaspis, Cera-
tocephala, Dicranurus, Isoprusia, Lobopyge, Tri-
merus, Dipleura, Ananaspis, Bohemoharpes, Cro-
talocephalus, Calymene, Gravicalymene, Sthena-
rocalymene, Otarion, Harpidella, Scharyia. Al-
though a substantial part of these genera belongs
to conservative and long-ranging genera, they ar-
gue for continuous evolution and absence of any
drastic change at the Silurian-Devonian bound-
ary.
The amount of true extinctions close to this level
is low I Prionopeltis, Tetinia, Richterarges among
widely distributed general and the extinction of
the last Raphiophoridae /Raphiophorus/ and so-
mewhat later also Encrinuridae is of lesser im-
portance, as both families were strongly declining
already in the Pridolian.
Innovations within the Lochkovian are marked
especially in proetids, scutelluids and dalmani-
tids - e.g. the onset or richer development of Al-
taepeltis, Spiniscutellum, Gerastos (some subge-
nera), Hollardia, Ranunculoproetus, Ganinella,
Khalfinella, Tropidocare, Taynaella, Tcherheso-
via, Lepidoproetus, Cordania, Crotalocephalina,
Koneprusia, Terranovia, Oinochoe, Corycephalus,
Huntonia, Odontochile, Phalangocephal us, Cry-
phina, Roncellia, Synphoroides, Paciphacops and
Digonus. Some of innovations concern particulary
endemic Synphoriinae, evolving within the Appa-
lachian Province (Ormiston 1972).
The onset and rapid evolution of Asteropyginae
within the siliciclastic-shale development during
the Lochkovian is of great biostratigraphic and
evolutionary value (Morzadec 1983, 1992 ;
 489

. 'l-"
Figure I - Stratigraphic
~" stages conodont events biotic events scale of Devonian to illus-
'f..'l- zones tr ate chro nostratigraphic
units, conodont zones, main
~ extinctions (incl .
Si.praesulcata jHangenberg E. event-str a tigraphic turnouts
ctyrnenids] and biotic events. An - An-
cy rogna t h us , Anc . A ncyro-
Z de lloides , crist. - cr ista-
« Pa. expansa
tus , E. - event, Eogn.- Eo-
g n a t ho dus , 1. - Icriodu s,
Z FAMENNIAN O. - Ozarkodina, Pa . -
a ~ .annulata· E. ~ inn ovati ons P almatolepis , Ped . - Pesa-
>
W
Po. postera
vis, Pol. - Polygna thus ;
Fb. trachytera Sch. - Schm id togn ath us ,
0 Si. - S ip h onod ella , T. k. -
Po. margini fera +0 first ely menids
To r t odus kockelianus , U.
0:: Po. rhomboidea
- upper, m. - middl e, I. - lo-
W Fb.crenida
wer, 1m. lowermost.
o, end of reefs Echelle stratigraphiqu e du
n.
:::>
Po. tr icnqulcr is
Pa.giqas
11... Kellwasser E. wid e-sccle extinctions Deuonieri indiquant les uni -
tes chronostratigraphiques,
les zones a conodont es, les
fRASNIAN lJ.
An.tr icnqutor is
-"c
4Ja.
... principau x eoenements stra -
m Pol. tigraphiques et biotiques.
~E
\." asymmetricus
I'm
lIIII Frasnes E. +ofirst Mant i coceras
a.
E.9
Z Fb. dispar ilis :JCl>

« Sch.herm.- AJl.crist
E >
._ a.
X"tl
Z GIVETIAN 0
E
a Pol. va r eus '4 Taghan ic E. +- ext i nctions
>
W
0 Pol. ensens is
~ Kacak E. +- extinctions
W T. k . kockel i anus
--J
0 EIFELIAN Lk . a ust ro l is
0 Pol.cos t. costotus • Chotec I inn ovations + ext inctions
E.
L Pol. cost .part i lus
c Pol.cost . potulus
c
'Q;'
e
PoL serotinus i sta rt of ana rcesrrds
Z 0
PoL inversus 4 Daleje E.
-«
EMSIAN--
.;
Pol. 9 ronbergi
Z 0
s:
a ,~
+- f irst goniatites
> N Pol. dehiscens ~ Basal ZlIchov E. +- last monograptids
W
0 Pol. pireneae

PRAGIAN Eogn. s.k ind le i

0::
W Eogn. sulcatus 4 Basal Proq ion E. I innovati ons
I

3: Ped.pesa vis ,
Anc. delta
a LOCHKOVIAN Oeurekcens is .i
start of g lobo \
distri butto n of
--J
dacryocona ri ds
lw . woschmidt i Sit-Dev. Bound. migrations
•
Sm eenk 1983). In the same facies , Acastella (per- An influx of new elements increase d wit hin the
sisti ng fr om the Pridol ian) r apidl y developed, upp er Lochkovian , where it is demonstrabl e even
being useful even for zonal su bdivisions (R. & E. within offshore realms - e.g. the first occurrences
Richter 1954 ; Gandl 1972 ; Tomczykowa 1991). of Reedops, Plagiolaria, Pragoproetu s, Prodreoer-
Its occurrence in the Baton River Formation of mannia, S chizoproetina, Otarionella a .o. which
New Zealand docum ents its very wide distribu- preceded their later ri ch development in Pragian
tion (Wri gh t , commun. in Strusz 1989). to Em sian times.
 490
Generally, the Silurian-Devonian boundary, well
marked in the trilobite distribution, is obviously
more expressed in migrations than in true extinc-
tions, being thus an example of a dispersal event
influenced by opened migration routes. The sub-
sequent Lochkovian is characterized by stepwise
innovations accelerated in the upper Lochkovian
(Monograptus hercynicus Zone) though the "Silu-
rian" aspect of most Lochkovian trilobite faunas
is retained.
PRAGIAN
Close to the lower Pragian boundary as recently
accepted and defined tChlupac & Oliver 1989), an
eustatic sea level lowering, called the Lochko-
vian-Pragian Boundary Event or Basal Pragian
Event (Chlupac & Kukal 1986, 1988) seems to be
of global significance, corresponding to the base
of the Cycle IA of Johnson et al. (1985). This tur-
ning point is clearly reflected in a quick increase
of diversity and onset of new elements in trilobite
faunas.
A typical example is the Pragian type area of
central Bohemia where the diversity increased
from about 25 late Lochkovian trilobite genera to
more than 50 found in the lowest Pragian. The
total diversity of Pragian trilobites reaches about
230 described species and subspecies, in contrast
to about 45 Lochkovian (Chlupac 1983, and new
data). Similar increase in diversity can be seen in
the Pragian of NW Africa (G. Alberti 1969, 1970,
1983). Data from the Carnic Alps (Ellermann
1992), Urals and Novaya Zemlya (Spassky 1992),
Canadian Arctic regions (Ormiston 1968 and re-
newed correlations) and SE Australia (e.g. the
Garra Limestone, Chatterton et at. 1979) also in-
dicate a similar trend.
The increase of Pragian trilobite diversity is best
marked in diverse offshore facies where the fall
of the sea level contributed to widening of shal-
low-water areas with carbonate (even reef) sedi-
mentation favourable for trilobites. The fall of the
sea-level near the Lochkovian-Pragian boundary
within the nearshore clastic facies resulted in
breaks and/or influx of coarser clastics less favou-
rable for trilobites. The transgressive phase of
the Pragian is believed to have positively affected
the development and distribution of trilobites as
clearly indicated by the occurrence of Asteropygi-
nae (Morzadec 1983, 1992) and Homalonotidae
(Wenndorf 1990).
A rapid evolution and diversification is best
shown in proetids which in the Pragian start
their period of maximum diversity lasting until
the early Eifelian. New or markedly richer repre-
sented genera are Pragoproetus, Orbitoproetus,
Simaproetus, Tropidocoryphe, Kegeliella, Dene-
markia, Astycoryphe, Wolayella, Alberticoryphe,
Miriproetus, Pseudodechenella, Nagaproetus,
Crassiproetus, Unguliproetus, diverse subgenera
of Cornuproetus etc. Maximal enrichment can be
observed in scutelluids (Kolihapeltis, Platyscutel-
lum, Arctipeltis, Bojoscutellum, Breviscutellum,
Metascutellum, Cornuscutellum, Microscutellum,
Poroscutellum, Paralejurus, Radioscutellum). A
pronounced diversification occurred in phacopids
iReedops, Prokops, Plagiolaria), dalmanitids - ri-
chly developing Odontochile and allied taxa, in
some Synphoriinae and allied Trypaulitinae (An-
chiopella, Synphoria, Forillonaria, Schoharie),
harpetids (diversity of Lioharpes, Kielania), chei-
rurids (Crotalocephalina, Crotalocephalides, Cro-
talocephalus), lichids (new Echinolichas, Gaspeli-
chas, Perunaspis, Craspedargest, homalonotids
(first Arduenella, Burmeisteria, Scabrellai. This
trend is pronounced even in increasing species di-
versity of more conservative aulacopleurids and
odontopleurids.
The increasing diversity is accompanied by in-
crease in morphotypes. Among types distinguis-
hed by Fortey and Owens (1990 a.b), phacomorph
(Phacopidae), effaced illaeniform (Paratejurus, Di-
pleura), strongly spinose (Odontopleuridae, Ota-
rionellai, pitted fringe-bearing (Harpetidae) are
more common, frequently persisting up to the
end-Emsian.
Demonstrable extinctions near the Pragian base
are rare. They include e.g. the important genus
Acastella occurring in terrigenous clastic facies,
some scutelluids (Spiniscutellum, Decoroscutel-
tum), phacopids tLochhooella), harpetids iBohe-
moharpes) and some other, rather few and mostly
endemic genera.
The development of Pragian trilobites points to a
general improvement of life conditions. This can
be explained by climatic warming, or in case of
the Prototethys regions, by an accelerated shift of
northern shelves of Gondwana together with as-
sociated microcontinents toward the north, i.e.
into the lower latitudes (this corresponds to an
increase in areas of reef buildups and larger pro-
portions of red coloured sediments). Migration
routes along the northern shelves of Gondwana
are indicated in mutual affinities of Australian
and European trilobite faunas of Lochkovian and
particularly Pragian age (Talent et at. 1972,
Chatterton et at. 1979, Holloway & Neil 1982). In
case of the Lochkovian-Pragian (= Basal Pragian)

Boundary Event, we do not deal with expressive-
ly abrupt environmental change but with a peak
of accelerated changes positively influencing par-
ticularly benthic biota.
LOWER EMSIAN (ZLICHOVIAN)
The recently accepted level of the lower Emsian
boundary at the base of the Polygnathus dehis-
cens Zone is not easily discernible, and from the
viewpoint of trilobite biostratigraphy rather pro-
blematic. The former lower Zlichovian boundary
falling within the P. dehiscens Zone is identical
with starting effects of the Basal Zlichov Event
(Chlupac & Kukal 1986, 1988 = Zlichov Event in
House 1989) which shows some innovations in
different faunal groups but is less expressed in
trilobite evolution.
Considering the Zlichovian (=Lower Emsian) as a
whole, stepwise innovations are characteristic,
becoming more pronounced in the higher part
(Polygnathus gronbergi conodont Zone). This is
also the level of the first worldwide distribution
of ammonoids (the Anetoceras fauna, Chlupac
1976 ; Chlupac & Turck 1gR::n Wh()~0 first appea-
C number
·".As:
... :
C ..
C trilobite genera in individual
Do
L. p. D. E.
491
ranee is likely to have followed the sea level rise
after the transgressive Basal Zlichov Event, as
assumed by House (1985, 1989).
The great majority of Pragian trilobite genera
continue into the Zlichovian. This applies to
many proetids, aulacopleurids, homalonotids,
harpetids and phacopids. A marked enrichment
can be seen in Asteropyginae (newly appearing
genera Comura, Delocare, Rhenops, Kayserops,
continuing evolution of Treveropyge, Pseudocry-
phaeus, Metacanthina, Pilletina - Morzadec 1983,
1992). Within the Appalachian Province, the evo-
lution of Synphoriinae continued (e.g. Synphoroi-
des, Synphoria, Roncellia, Anchiopsis, Schoharie)
with a few innovations.
Zlichovian innovations within the offshore trilo-
bite faunas of the Old World Province (mostly
carbonate facies) involve especially an onset or ri-
cher diversification of some Proetida (Schizoproe-
tus, Phaetonellus, Trautensteinproetus, Proetopel-
tis, some subgenera of Cornuproetus Macro-
blepharum, Tafilaltaspis), but concern to a lesser
degree also other groups (radiation of Scabriscu-
tellum, Chotecops). The starting lineage of coarse-
of genera:
30
20 Figure 2 - Representation of
10 Devonian stages. Stages : L. -
5 Lochkovian, P. - Pragian, Z. -
0 Zlichovian, D. - Dalejan, E. -
Eifelian, G. - Givetian, Fr. -
Frasnian, Fa. - Famennian.
Families: S - ScuteJluidae, Pr
- Proetidae, A . Aulacopleuri-
due, B - Brachymetopidae, Ph
- Phacopidae, Da - Dalmaniti-
dae, As - Asteropyginae, C -
Calmoniidae, Ch - Cheiruri-
dae, Ca - Calymenidae, H -
Homalonotidae, Ha - Harpeti-
dae, Li - Lichidae, 0 ·Odonto-
pleuridae. Stratigraphically
uncertain occurrences not in-
cluded. Repartition des genres
de trilobites dans chacun des
etages deooniens. Les presences
stratigraphiques incertaines nc
son.t pas prises en con.sidero-
G. Fr. Fa. tion.
 492
ly sculptured and large-eyed phacopids (Phacops
s. str.) falls also within the Zlichovian - Phacops
(P.) degener BARRANDE is the first known link of
this lineage. When compared with the Pragian,
scutelluids are retreating (6-7 genera in contrast
to 17 Pragian genera).
Although no abrupt change in trilobite fauna is
apparent at the lower Zlichovian boundary, the
continuing sea level rise contributed to decrea-
sing provinciality of Zlichovian trilobites. Mixtu-
res of the Bohemian offshore and Rhenish
nearshore elements become more common, as evi-
denced in the Variscan mobile belt of Europe (ty-
pical examples occur in the Ardenne-Rhenish
area, Harz Mts., Armorican Massif, Moravia, N.
Spain, Turkey and NW Africa). This trend increa-
sed within the late Zlichovian Polygnathus gron-
bergi Zone being considerably accelerated during
the subsequent transgressive Daleje Event.
Persisting wide migration possibilities within the
Old World Province, and particularly along the
northern shelves of Gondwana are shown in affi-
nities of trilobite faunas from Europe, NW. Afri-
ca, Turkey (Haas 1968), and SE Australia (Chat-
terton 1971 ; Talent et al. 1972 ; Chatterton &
Wright 1986).
Total Zlichovian trilobite diversity (Fig. 2) is si-
milar to that of the Pragian, and its decrease in
some groups (Scutelluidae) is compensated by an
increase in other groups (Asteropyginae).
The starting development of rich and distinctive
Devonian trilobite faunas of the Malvinokaffric
Province (sensu Eldredge & Ormiston 1979) de-
pends on a transgressive phase within South
Gondwana, best recognizable in South America
and South Africa, Although the absence of index
zonal fossils (conodonts, ammonoids, dacryocona-
rids) makes correlations difficult, recent concepts
indicate most likely a lower to late Emsian trans-
gression (Melo 1988 ; Theeron & Loock 1988 ; Ra-
cheboeuf 1992), i.e. a process which may reflect
the transgressive trend starting in the Zlichovian
and particularly accelerated in the Daleje Event.
Malvinokaffric Devonian trilobite faunas are dis-
tinguished by dominant and mostly endemic Cal-
moniidae (Calmonia, Pennaia, Probolops, Malvi-
nella, Plesiomalvinella, Typhloniscus, Schizosty-
lus, Tarijactinoides, Bainella, Paracalmonia, Bou-
leia, Parabouleia, Deltacephalaspis, Chiarumani-
pyge etc.) which developed since the Emsian until
the Eifelian together with an increasing influx of
extra-Malvinokaffric immigrants (homalonotids,
phacopids, aulacopleurids, proetids, Elredge &
Ormiston 1979 ; Eldredge & Brania 1980 ; Coo-
per 1982 ; Popp 1985 ; Malo 1988). Species of Me-
tacryphaeus with traceable lineages from Emsiari
up to the lower Givetian allow a closer zonal sub-
division.
UPPER EMSIAN (DALEJAN)
The eustatic sea level rise seems to be markedly
accelerated within the lower Dalejan, reaching its
peak in the P. inversus- P. serotinus conodont Zo-
nes. This is the expression of the Daleje Event
(House 1985, Chlupac & Kukal ; 1986, 1988)
which clearly influenced the marine biota inclu-
ding trilobites.
Numerous innovations can be seen in offshore
carbonate and shale facies : richer diversification
and wider spread of proetids (Macroblepharum,
Cyrtosymboloides, Linguaproetus, Diademaproe-
tus, Phaetonellus, Proetopeltis, Quadratoproetus,
Myoproetus, Unguliproetusi, spinose scutelluids
(Thysanopeltella) , and some phacopids (Struveas-
pis, Illaenula, Eocryphops). Many lineages with
evolutionary species, however, extend from the
Zlichovian which also includes the majority of
conservative aulacopleurids, cheirurids, harpetids
and odontopleurids.
Dalejan offshore trilobites of the Old World Pro-
vince often occur in the tentaculite shale facies,
marked by impoverished benthos and an abun-
dance of planktonic, nectonic and epiplanktonic
forms. Trilobites with suppressed vision or
blindness are characteristic, e.g. Plagiolaria,
Struveaspis, Phyllaspis, Illaenula, Eocryphops,
Piriproetus, blind harpetids. This facies reflecting
anoxic conditions became widely distributed after
the Daleje Event : it is known from Thuringia,
Moravia, N. Spain, Malaysia, Thailand, N. Viet-
nam, South China (discussed in Chlupac 1983),
continuing in some cases up to the Eifelian. Trilo-
bites show here clearly environmentally conditio-
ned reduction of vision which repeated in diverse
stages of trilobite evolution. Not all tentaculite
shales, however, contain trilobite assemblages of
this type and some calcareous shale facies with
demonstrable autochthonous trilobite remains
lack forms with eye-reduction (the Daleje Shale
of the Barrandien as a typical example, Chlupac
1983).
Trilobites with reduced and non- reduced eyes of-
ten occur together in micritic "cephalopod" lime-
stone facies, characterized in the Dalejan by com-
mon goniatites (typical anarcestids), formerly, be-
fore the new Lower IMiddle Devonian boundary
definition, included in the Eifelian. Particularly
in this facies, the evolution of trilobites continues
 493
from the Dalejan into the lowest Eifelian (the Po-
lygnathus costatus partitus Zone) without any
pronounced changes, even on the species level
(Chlupac 1985b).
Dalejan shallow-water and carbonate, mostly
platform deposits exhibit increasing diversity of
dechenellids (Schizoproetus, Praedechenella, Basi-
dechenella a.o.) but these are still behind their
Eifelian peak. Trilobites of Dalejan reef facies are
insufficiently known ; an exception is the Formo-
sa Limestone of S. Ontario (Appalachian Pro-
vince) where Crassiproetinae are characteristic
(Ludvigsen 1987).
Within the terrigenous clastic and mixed clastic-
carbonate near-shore deposits, Asteropyginae dis-
play clear innovations, attaining their maximum
diversity (new : Psychopyge, Breizhops, Turco-
pyge, Kayserops, Greenops, Feruminops, Gourdo-
nia, comp, Morzadec 1983, 1992). Homalonotidae
show a gentle decrease in generic diversity
(Wenndorf 1990), whereas Dalmanitinae are
clearly declining (including Odontochilinae).
Synphoriinae of the Atlantic Province continued
in evolution from the Zlichovian up to the lower
Eifelian without pronounced and abrupt changes;
their relative Malladaiinae were spread even out-
side the Atlantic Province (Cantabrian region,
Arbizu 1979). A special group of mostly endemic
spinose phacopids (Echinophacops, Toxophacops)
developed under conditions of geographic isola-
tion in Central Mongolia (Zhou & Campbell 1990)
and reached in less typical form even the Far
East (Japan, Kaneko 1990).
EIFELIAN
Whereas the proper level of the Lower /Middle
Devonian boundary internationally accepted at
the base of the Polygnathus costatus partitus co-
nodont Zone (Ziegler & Klapper 1985) generally
shows no apparent change in trilobite faunas, a
pronounced turning point falls somewhat higher,
near the base of the subsequent Polygnathus cos-
tatus costatus Zone. This is the level of the Cho-
tec or Basal Chotec Event (Chlupac & Kukal
1986, 1988 ; House 1989), called also the jugleri
Event (after the goniatite Pinacites jugleri, WAL-
LISER 1984, 1985). This level was recognized al-
ready formerly in different areas as a distinctive
biostratigraphic boundary expressed in diverse
fossil groups.
The level of the Chotec Event is marked in off-
shore carbonate facies of Bohemian type in ge-
nerally increasing abundance of formerly less fre-
quent representatives of Aulacopleura, Cyphaspi-
des, Aulacopleurina, Proetopeltie.Koneprusites, Pi-
riproetus, Thysanopeltis, Erbenites, but in only
few evolutionary new genera (some dechenellids,
lichids). Evolution of many Proetidae, Aulaco-
pleuridae, Harpetidae and Odontopleuridae conti-
nued from the Dalejan into the Eifelian without
any pronounced breaks and only innovations at
the species level are confined to the Chotec Event
level. Among important genera, Paralejurus beca-
me extinct at this level.
The increasing amount of shallow-water carbo-
nate platform and epeiric sea deposits of the co-
ral-stromatoporoid facies, often rich in brachio-
pods, exhibits the acme-development of Dechenel-
linae, often persisting into the early Givetian.
This richly diversified group (Dechenella, Pedino-
dechenella, Cyrtodechenella, Humeia, Camsellia,
Monodechenella) developed in different provinces
of the warm climatic zone. The accompanying ele-
ments were the scutelluids of the Scutellum fla-
belliferum group and large-eyed and coarsely
sculptured phacopids of the Phacops schlotheimi-
latifrons group, whose evolution continued from
the late Emsian into the Givetian (Struve 1970,
1990, later superfluous nomenclatory splitting of
this group discussed by Chlupac 1977). The giant
phacopids from Morocco, such as Ph. (Ph.) mega-
lomanicus (STRUVE), belong here too.
Clear innovations occur in Lichidae which con-
tain characteristic bizzare and often endemic
forms (Ceratarges, Eifliarges, Mephiarges, Akan-
tharges, Ceratolichas). The species diversity of
scutelluids is decreasing, though this group also
shows some bizzare and unusual forms (Ancyro-
pyge) widely distributed in shallow-water facies.
Asteropyginae, concentrated in diverse near-
shore facies, are gently declining with many ex-
tinctions below the Eifelian base and only less
frequent innovations (Heliopyge, Bradocryphaeus,
Asteropyge, Neometacanthus ; Morzadec, 1983,
1992). Homalonotidae, Calymenidae, Cheiruridae
and Dalmanitidae are retreating being repre-
sented only by a few genera.
Synphoriinae of the Appalachian Province show a
clear decrease of generic diversity (6 genera
known) and the same trend may be seen in Cal-
moniidae in the Malvinokaffric Realm.
Generally reviewed, the sea-level rise starting
with the Chotec Event was not accompanied by
innovations fully compensating the decline. Mi-
grations and extinctions are characteristic of this
period, but changes are less dramatic and statis-
tically less important (Fig. 3). Following the sea
 494

Figure 3 - Generic di-

L. P. z. D. E. G. Fr. Fa.
versity of individual fa-
milies and selected
'77,
subfamilies of Devo-
Odontcoteuridce V//hfW: '/// nian trilobites. Trypau-
litinae, Malladaiinae,
Acastavinae and Neo-
Lichidae synphoriinae included
under Synphoriinae.
Harpetidae ' / / 'II/I. ' / / / / Filled collumns - strati-
graphically certain rep-
resenta tion, unfilled
collumns - uncertain
occurrences, L.
Calymenidae Lochkovian, P. - Pra-
gian, Z, - Zlichovian, D
Cheiruridae - Dalejan, E. - Eifelian,
G ' - Givetian, Fr, -
Frasnian, Fa. - Famen-
nian. Dioersite generi-
Calmonidae que de chaque famille
et sous-famille de trilo-
bites devoniens. Les
- sous-familles Trypauli-
As t ero pVC] inee '/// tinae, Malladaiinae,
Acastavinae et Neosyn-
Synphoriinae et -
'I'////.. phoriinae. Les colonnes
allied '//~~W'~////J hachurees correspon-
dent aux repartitions
stratigraphiques certai-
Dalmanitinae (other) l'l//' nes, tandis que les co-
~ lonnes uides indiquent
V///...'l///...'/'/// ~ '///.
les incertitudes.
Phacooidae
j?rachymetopidae
dncLScharvinae)

Eodrevermanniinae -
Aulacooleuridae ~ '//// r//// '//.

Cyrtosym bot i na e,
Drevermanniinae
=--
~~/.a~~~
Dechenellinae ,
Warburg ellinae
- 30
Cornupro etinae '//.Ir/ffh~// '///1
20
Tropidocoryph ina e 10
1o 5

Scutelluidae

L. P. Z. D. E. G. Fr. Fa.

level rise and marine transgressions, the wider
areas of shallow-water carbonate sedimentation
offered suitable life conditions for some new Proe-
tidae (especially Dechenellinae), Scutelluidae and
Phacopidae which show increasing cosmopolita-
nism.
THE EIFELIAN-GIVETIAN BOUNDARY
INTERVAL AND THE GIVETIAN
The Eifelian-Givetian boundary interval is distin-
guished by the Kacak (or otomari) Event as char-
acterized by House (1985), Walliser (1985), Chlu-
 495
pac & Kukal (1986, 1988), Boucot (1990),
Truyols-Massoni (1990). Culmination of this
event falls near the limit of the Tortodus kocke-
lianus kockeZianus-PoZygnathus ensensis Zones,
where effects of the eustatic sea-level rise and
the onset of anoxic conditions affected particular-
ly the level bottom biota of open shelf areas, in-
cluding trilobites.
The Barrandian (Bohemia) can serve as a typical
example, where none from about 50 trilobite spe-
cies of the Eifelian Chotec Formation survived
the invasion of an anoxic regime at the beginning
of the Kacak Shale sedimentation. Only a few
representatives of opportunistic long-ranging ge-
nera (AuZacopZeura, Leonaspis, Chotecops) reap-
peared near the Kacak-Roblin Members boundary
when the life conditions somewhat improved
(Chlupac 1983).
Givetian transgression, starting with the Kacak
Event, lead to a landward shift and a wide distri-
bution of shallow-water shelf, mostly carbonate
facies accompanied by wide-range migrations of
trilobites, clearly documented especially in Aste-
ropyginae (Morzadec 1992), Phacopidae and
Dechenellinae. The inner shelf and platform
areas beyond the reach of ascending cool oceanic
waters were not severely affected by the Kacak
Event, and trilobite assemblages with typical
Dechenellinae, Proetinae (cf. Lutke 1990), Scutel-
luidae of the Sc. flabelliferum-coetatum group
and large-eyed and sculptured phacopids develo-
ped from the Eifelian into the Givetian without
major changes, as shown in Variscan Europe and
the North American Interior (comp, e.g. evolution
of the Phacops rana stock, Eldredge 1972). A
trend of decreasing provinciality within shelf tri-
lobite faunas persists. The spread of the curious
scutelluid Ancyropyge may serve as an example :
it migrated not only within North American
realms (Ormiston 1972) but reached even central
Europe (Moravia, Chlupac 1992).
Also a trend of decreasing trilobite diversity con-
tinued and was clearly accelerated by the Give-
tian Taghanic Event which had a strong impact
on ammonoids (House 1985, 1992) and develo-
pment of North American faunas (Johnson et aZ.
1988). As Feist (1991a) stated, this event was
lethal for lichids, cheirurids and probably even
homalonotids and calmoniids, all inhabiting the
shallow-water neritic realms. Givetian trilobites
of the deep-water and offshore shale facies are so
far known only scarcely which may reflect their
retreat.
Malvinokaffric trilobites show a rapid decline and
only a few genera (Metacryphaeus, Cryphaeoides)
reached the Givetian regarding the recent corre-
lations (de Melo 1988 ; Theron & Loock 1988 ;
Racheboeuf 1992). The regressive trends caused
by south Gondwana tectonic regime and contras-
ting with global eustatic fluctuations played a de-
cisive role.
Generally, the time-span starting with the Kacak
Event and including the whole Givetian shows a
profound fall of taxonomic diversity of trilobites.
The number of totally disappearing families is si-
gnificant (Cheiruridae, Lichidae, probably Calmo-
niidae and Homalonotidae) and the same con-
cerns a number of subfamilies (Synphoriinae,
Trypaulitinae, Acastavinae, Scharyiinae, Eremi-
proetinae, Cyphaspidinae), The decrease at the
generic level includes more than 70% of that of
the pre-Kacak Eifelian, being thus the most im-
pressive interval of trilobite decline within the
Devonian.
THE FRASNIAN AND THE END-
FRASNIAN EXTINCTION
Trilobites developed from the late Givetian into
the Frasnian particularly in the shelf facies wi-
thout marked breaks, and families surviving the
Middle Devonian events reached as far as the
late Frasnian. Innovations are scarce (new Ptero-
paria-group, Drevermanniinae) and the generic
diversity markedly decreases (Feist 1991).
The latest Frasnian global crisis - the Kellwasser
Event (for reviews esp. House 1985, 1989, Sand-
berg et aZ. 1988 ; Schindler 1990) strongly affec-
ted all benthic groups together with trilobites.
The major part of families and subfamilies per-
sisting up to the Frasnian became extinct (Odon-
topleuridae, Harpetidae, Asteropyginae, Scutel-
luidae, Cornuproetinae, Tropidocoryphinae, comp.
Briggs et al. 1988 and especially Feist 1991).
Survivors, such as opportunistic and small-eyed
phacopids (Phacops, Chotecops, Cryphops,
Nephranops) and blind proetids (Drevermannii-
nae) are scarce and confined to the cephalopod
facies.
Allthough the impact of the Kellwasser Crisis on
evolution of trilobites is severe and documented
in sections studied in great detail (e.g. Montagne
Noire-Feist 1990), the total extinction includes fa-
milies which were strongly reduced already prior
to the Kellwasser Event and represented only by
a few genera and species (comp. Feist 1991) : scu-
telluids did not reach the Kellwasser level at all,
late Frasnian aulacopleurids, odontopleurids,
harpetids and Asteropyginae reached this level,
 496
each with a single genus and species . In th ese
cases, the disappearance proper cannot be classi-
fied as a mass extinction.
The shortage of studied sections in the offshore
shale facies makes it impossible to evaluate the
effects of the Kellwasser Event on expressively
pelagic trilobite assemblages.
THE FAMENNIAN AND THE
DEVONIAN-CARBONIFEROUS
BOUNDARY
Famennian trilobite faunas are dominated by
small proetaceans (typical Cyrtosymbolinae, ran-
ged by some authors within the oldest Phillipsii-
dae ) and normal sized Phacopidae. Thanks to ca-
reful studies commenced by the classic mono-
graph of R. & E. Richter (1926), the biostratigra-
phy of Famennian trilobites with regard to cono-
dont zonation is better known than in earlier De-
vonian stages (summari zing reviews in G. & R.
Hahn 1975 ; C. & B. Brauckmann 1986 ; Yuan
1988 ; Feist 1991).
The majority of described trilobites is derived
from offshore cephalopod facies, and only a minor
part is related to carbonate turbidites (Moravian
Karst), shallow-water carbonates (few European
localities, China) and shale deposits.
After the Kellwasser crisis, lower Famennian
Cheiloceras Biozone (Nehdenian Substage) is
characterized by the first, mostly oculate Cyrto-
symbolinae (Cyrtosym bole, Calybole ) of uncertain
origin, blind Drevermanniinae (Formonia, Fri-
thjofia) and small-eyed or blind Phacopidae (Due-
tina, Dienstina, Cryphops, Trimeroeephalus,
Nephranops).
The subsequent Prolobites-Platyclymenia Zone
(Hembergian) shows only a few innovations tCyr-
tosymbole (Franconicabole) typical) but the anoxic
Platyclymenia Event, reflected mostly in the am-
monoid evolution (House 1985, 1989), seems to
have an influence on increasing trilobite radia-
tion within the cephalopod facies near the Hem-
bergian/Dasbergian boundary (new Dianops,
Chaunoproetus, etc.), Particularly important from
the phyletic viewpoints are the first Waribole and
Pseudowaribole which are regarded by some au-
thors as roots of phillipsiid subfamilies richly ra-
diating within the Lower Carboniferous (Hahn &
Brauckmann 1984, 1989 ; C. & B. Brauckmann
1986 ; Fei st 1991).
The final and pronounced radiation of Devonian
trilobites falls into the upper Famennian Clyme-
Br
lIn
Ph
'.
2 LS.\J
50 species
40
30
20
'0
o
I
Cheiloc . Platycl. Clymenia- I
Wockl. I
Figure 4 - Famenn ian tri lobites species diversity plotted in
main ammonoid zones. Cheiloc. - Cheiloceras Zone, Platycl. -
Platyclymenia Zone, Clymenia - Wockl. - Clymenia and
Wocklumeria Zones, Pr - Pro etidae, Ph - Phacopid ae, Br -
Brachymetopidae. Diuersite specifique des trilobites {amen-
niens dans les principales zones d 'ammonoides.
nia and Woeklumeria Biozones where a general
increase in generic and species diversity can be
demonstrated both in proetaceans (new Dreoer-
mannia, Parafrithjofia, Daihuaia, Haasia, Skem-
matopyge, Typhloproetus, Parachaunoproetus, He-
lioproetus, Silesiops, Mirabole, Perliproetus), and
phacopids (richer diversity within Dianops, small-
eyed Cryphops-like phacopids, oculate Phacops
(Ph.) granulatus and Phacops (Omegops) accipi-
trinus groups). The first Brachymetopus and Pu -
doproetus found in shallow-water carbonates
(Feist & Petersen 1993) are pioneers of early
Carboniferous faunas .
Phacops (Omegops) accipitrinus (PHILLIPS) has a
special biostratigraphic value among latest Fa-
mennian phacopids , as it shows a wide geogra-
phical distribution (E u r op e, NW Africa, Asia) and
a lesser facies dependence - apart from the shal-
low-water carbonates it occurs in turbiditic layers
and in shale facies (here demonstrably autochtho-
nous e.g. at Velbert, Germany). A distinct envi-
 497
ronmental tolerance of some phacopids apparent
during their earlier evolution (Chlupac 1977) is
demonstrable even in their latest development.
The Late Famennian trilobites show an increase
of morphotypes in both two dominant families
(Proetidae, Phacopidae) : phacomorph, effaced
ilaenimorph (Helioproetus, Dianops) and athe-
loptic (Drevermanniinae) forms, commonly ac-
companied by miniaturisation (Cyrtosymbolinae).
Trilobites at the Devonian-Carboniferous bound-
ary exhibit a radical change, confined to the level
of the Hangenberg Event, which is marked by the
onset of widely distributed, probably transgres-
sive Hangenberg Shale facies followed by a re-
gressive pulse. Late Famennian Cyrtosymbolinae
disappear together with all phacopids. Particular-
ly, the abrupt extinction of phacopids, which de-
veloped since the early Silurian for more than 80
My, is of special interest, as their widespread dis-
tribution and diversity in the late Famennian
shows neither degeneration nor decline.
The earliest Carboniferous assemblages contain
the widely distributed proetaceans Belgibole
abruptirhachis (RICHTER & RICHTER), followed by
Semiproetus (MacroboleJ drewerensis (RICHTER &
RICHTER) and representatives of Liobolina, Cyrto-
proetus, Diacoryphe, Globusia etc. of clearly Car-
boniferous aspect (trilobites of the boundary in-
terval are discussed in more detail by Brauck-
mann et al. 1993).
A great similarity in change of trilobite faunas at
the Devonian-Carboniferous boundary, demons-
trated even in distant regions (Europe-China), re-
flects an abrupt and global turning point which
affected both shallow and deeper marine realms
at least within the warm climatic zone. Although
some similarity with the bioevent at the Silurian-
Devonian boundary exists (e.g. in unusually wide
distribution of some proetaceans), the effects of
the Hangenberg Event were more severe and
lethal for most trilobites, even though survivors
at the generic level do exist (Pseudowaribole, Wa-
ribole, Dreoermannia, Brachymetopus, and Pudo-
proeius).
COMPARISON WITH
DEVELOPMENT OF SOME OTHER
FOSSIL GROUPS
Many taxa, among widely distributed and strati-
graphically evaluated Devonian fossil groups,
particularly ammonoids, conodonts, dacryocona-
rid tentaculites, and to a lesser degree also some
other groups (graptolites, corals) offer reliable
data comparable with trilobite development.
DEVONIAN AMMONOIDS sensitively reflect environ-
mental changes and event-stratigraphic turnouts,
starting with the Emsian (reviews : House 1985,
1989a,b). The appearance of goniatites falls in
the transgressive phase after the Basal Zlichov
Event and their first wide distribution within the
late Zlichovian (Chlupac & Turek 1983) corres-
ponds to a lesser pronounced increase in trilobite
diversity, accompanied by decreasing provinciali-
ty. The Daleje transgressive Event shows a simi-
lar effect on trilobites ; and in goniatites this
event is emphasized by important innovations
(onset of Anarcestidae and Agoniatitidae).
The level of the Chotec Event has a pronounced
effect in the evolution of both groups, but innova-
tions are more pronounced in the goniatites (esp,
within Pinacitidae, Anarcestidae and Agoniatiti-
dae), than in trilobites, where mostly changes on
species-level occurred.
The Kacak Event, lethal for many trilobites, is
marked in ammonoids by the widespread occur-
rence of Cabrieroceras, Tornoceratidae and the
first Sobolewiidae. The Givetian Taghanic Event-
a further step in the trilobite decline, is also mar-
ked by extinctions of ammonoids (Agoniatitidae,
Pinacitidae, Maenioceratitidae, Sobolewiidae, de-
crease in Anarcestidae), but it was followed by
important innovations (multilobed Pharcicerati-
dae, Gephuroceratidae - House 1989b) absent in
trilobites.
The Kellwasser Event deeply affected ammonoids
and trilobites and only a few of these survived
this episode. However, both groups clearly show a
declining trend prior to the event (House 1985,
1989b ; Becker 1986).
The stepwise increase in trilobite diversity within
the Famennian has even better expressed coun-
terparts in ammonoid evolution - the onset of
Cheiloceratidae in the lower Famennian is close
to that of Cyrtosymbolinae, the explosive radia-
tions of Clymeniina and Gonioclymeniina roughly
correspond to trilobite innovations after the an-
nulata Event and in the Wocklumeria Zone. The
Hangenberg Event, close before the Devo-
nian/Carboniferous boundary, was lethal for most
Famennian ammonoids and trilobites.
Generally, the levels of changes in Devonian am-
monoids and trilobites correspond well to each
other, though in trilobites the stepwise retreat
starting within the Middle Devonian is in con-
 498
trast to a much more diverse picture of innova-
tions and extinctions within ammonoids.
CONODONTS. The rapid evolution of Devonian co-
nodonts as summarized by Ziegler and Lane
(1987) shows trends and changes comparable
with trilobite development at different stratigra-
phic levels.
The level of the Basal Pragian Event shows a
contrasting effect : a marked diversity increase in
trilobites and the beginning of a low diversity in-
terval in conodonts roughly corresponding to the
Eognathodus sulcatus to Polygnathus pireneae
Zones.
The high diversity interval in conodont develo-
pment starting with the Polygnathus dehiscens
Zone and persisting up to the P. ensensis Zone
only partly corresponds to the persisting high di-
versity int erval of trilobites throughout the
Zlichovian to early Eifelian time, the limits of
changes being not fully correlatable (the base of
the P. dehiscen s Zone is unexpressive in trilob i-
tes, the level of the Kacak Event is less pronoun-
ced in conodonts).
The conodont extinction event close to the base of
the Polygnathus varcus Zone, and the high diver-
sity interval starting in the upper part of the P.
varcus Zone and persisting up to the Frasni an
have no clear analogue in trilobite evolution and
diversity, which show a markedly declining
trend.
The FrasnianlFamennian crisis - the Kellwasser
Event strongly affected conodonts and trilobites
in a comparable manner (cp. Sandberg et al.
1988; Schindler 1990), although the succeeding
innovations in conodonts already within th e up-
per part of the Palmatolepis triangularis Zone
are much more extreme and earlier than those in
trilobites . The Middl e and late Famennian high
diversity intervals in conodont evolution have
only a lesser marked analogy in the trilobites.
The Hangenberg Event, close to the Devonian-
Carboniferous boundary, exhibits quite compara-
ble effects on both groups, among which only few
surviving stocks exist (cp. Walliser 1984 ; Ziegler
& Lane 1987 for conodonts).
DACRYOCONARID TENTACULITES. Evolution of this
typical Devonian and biostratigraphically evalua-
ted group (reviews : Alberti 1979, 1987 ; Lutke
1979, 1985) allows some comparisons to be made.
The first stage of worldwide distribution and rich
radiation of dacryoconarids within the late
Lochkovian has only lesser expressed analogy in
the increase of trilobite diversity. The marked
rich radiation of trilobites after the Basal Pra-
gian Event has a contrasting counterpart in the
low-diversity episode of Pragian dacryoconarids,
though their global distribution is impressive (the
Nowakia acuaria group ).
Late Pragian and Zlichovian transgressive phases
are reflected in radiations of dacryoconarids
which reached their peak of abundance around
and after the transgressive Daleje Event, having
only a weak analogy in an increased diversity of
trilobites within the Upper Emsian (Dalej an).
Starting with the Chotec Event, the development
of both groups shows a declining trend in diversi-
ty : The Kacak Event, though contributing to a
wide distribution (th e Nowakia otomari group ),
shows only a few innovations in dacryoconarids,
and the retreat continues in the Frasnian which
is a clear extinction interval (N owakiidae and
Styliolinidae disappear). The only flourishing
group - the Homoctenidae were drastically redu-
ced as late as during the Kellwasser Crisis, shor-
tly preceding the total extinction of dacryo cona-
rids within the early Famennian (discussion in
Schindler 1990, Hladil et al. 1992 ).
GRAPTOLITES. Generally strongly declining Devo-
nian Graptoloidea (review in Jaeger 1988) show
some analogy with the development of trilobites
particularly close above the Silurian- Devonian
boundary : the world-wide distribution of the in-
dex Monograptus uniformis PRIBYL corresponds
to that of the Warburgella rugulosa group of
proetids. The spread of Monograptus hercynicus
PERNER falls within the interval of increasing
late Lochkovian trilobite diversity, whereas the
Basal Pragian Event has contrasting effect, r esul-
ting in a progressive restriction of graptolites and
a marked diversity increase in trilobites . Th e ex-
tinction of graptoloids close to the PragianlEm-
sian boundary has no apparent counterpart in
trilobite development.
SOME OTHER GROUPS. The development of Devo-
nian rugose corals as reviewed by Oliver (1990)
for the Eastern Americas realm, shows analogy
with trilobites in the "Silurian" aspect of Lochko-
vian corals, followed by a strong change and in-
novations correlatable with the Basal Pragian
Event, which triggered the onset of elements
characterizing Devonian coral faunas up to the
Kellwasser crisis. The coral turnover within the
Polygnathus costatus costatus Zone can be corre-
lated with the Basal Chotec Event (or if delayed
also with the Kacak Event). The Taghanic Event
was reflected in rugose corals by markedly more
 499
innovations which contrast with generally decli-
ning trilobite diversity. The Kellwasser Event at
the FrasnianlFamennian boundary, was fatal for
most corals and trilobites.
CONCLUSIONS
The development of trilobites clearly reflects en-
vironmental changes and event-stratigraphic
turnovers throughout the Devonian.
Lowered trilobite diversity persists from the mid-
Silurian into the Lochkovian. The Silurian-Devo-
nian boundary interval exhibits no drastic chan-
ges in trilobite evolution with only minor extinc-
tions but more important migrations.
The Pragian age is distinguished by markedly in-
creased trilobite diversity. Although this was pre-
ceded already by the late Lochkovian radiation,
the diversity rise was triggered by the Basal Pra-
gian (=Lochkovian-Pragian Boundary) Event of
regressive character, which widened the shallow
marine realms suitable for trilobites.
The overall trilobite diversity shows no strong
drop from the Pragian until the early Eifelian.
The transgressive events (particularly the Daleje
Event) were accompanied by moderate changes in
trilobite faunas but extinctions were mostly com-
pensated by innovations. A trend of decreasing
provinciality is apparent within the Old World
Realm. The development of Malvinokaffric trilobi-
tes was controlled at its beginning by transgres-
sive, probably eustatic phases within the Emsian
but later controlled by the South Gondwanan tec-
tonic regime.
The rapid decline of trilobites was markedly trig-
gered by the transgressive Kacak Event being ex-
pressed in general diversity fall and extinctions
not compensated by innovations. This trend con-
tinued through the Givetian and became rapidly
accelerated in the Kellwasser Crisis near the
Frasnian/Famennian boundary.
Famennian innovations, which involved only
proetids (Cyrtosymbolinae, Drevermaniinae) and
phacopids, show a stepwise progressing trend
which was abruptly interrupted by the Hangen-
berg Event close before the Devonian-Carbonife-
rous boundary. A few surviving genera gave rise
to Tournaisian radiations of Proetida.
Comparisons with selected other fossil groups, es-
pecially ammonoids, conodonts, dacryoconarid
tentaculites etc., show that changes in develo-
pment fall mostly in correlative levels, though
the effects were variable.
Analogous effects are confined to persisting "Silu-
rian" aspect of Lochkovian faunas (trilobites, co-
nodonts, corals), late Lochkovian innovations (co-
nodonts, trilobites, very marked in dacryocona-
rids), increase of diversity after the Basal Pra-
gian Event (trilobites, corals) and the trend of ra-
ther persistent high diversity up to the early Ei-
felian (ammonoids, conodonts, dacryoconarids,
trilobites). The Kacak Event is marked by a pro-
found decrease in diversity of trilobites with a si-
milar but less extreme effect on dacryoconarids,
ammonoids and corals (compensated by innova-
tions in the last two groups). The Kellwasser and
Hangenberg crises were generally drastic tur-
nouts with extinctions detectable in most groups.
Stepwise innovations within the Famennian have
analogues in diversity increase of trilobites, am-
monoids and particularly conodonts.
Contrasting effects on different faunal groups
were observed especially at the level of the Basal
Pragian Event (diversity decrease in conodonts
and dacryoconarids, increase in trilobites and co-
rals). This may be explained by the sea level fall
which positively affected the distribution and ra-
diation of shallow-water benthic (but not pelagic)
biota. Innovations contrasting with general de-
cline of trilobites are in corresponding levels, es-
pecially in Middle to Late Devonian conodonts,
ammonoids and rugose corals.
The explanation of changes in trilobite develo-
pment during the Devonian lies in climatic shifts
related to changes of the sea level and current
circulation markedly accelerated in event- strati-
graphic turnouts. The cause of general trilobite
retreat, however, was evidently not dependant on
events proper but had a reason functioning
throughout a long period, i.e. probably the hy-
drosphere development since the Ordovician.
Acknowledgements - The author thanks Dr. R Feist
(Montpellier), Dr. B. Chatterton (Edmonton) and P.
Morzadec (Rennes) for a critical rewiew of the
manuscript, and Dr. RM. Owens (Cardiff), Dr. J.N.
Spassky (St. Petersburg) and Dr. B.G. Waisfeld
(Cordoba) for valuable comments and suggestions. Mrs.
H. Bryndova, Dr. V. Kachlik, and Ing. P. Hofman
helped in technical preparations of the manuscript.
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APPENDIX
Data base of Devonian trilobite families, subfami-
lies and genera considered in the paper
Abbreviations : L - Lochkovian, P - Pragian, Z -
Zlichovian, D - Dalejan, E - Eifelian, G - Give-
tian, Fr - Frasnian, Fa - Famennian.
SCUTELLUIDAE Richter & Richter, 1925
Altaepeltis MAKSIMOVA, 1968 L?PZD
Ancyropyge CLARKE, 1891 P?EG
Arctipeltis MAKSIMOVA, 1968 PZDE
Bojoscutellum SNAJDR, 1958 P
Boreoscutellum PRIBYL & VANEK, 1971 LP?
Breuiscutellum SNAJDR, 1966 PZDE
Bubupeltina SNAJDR, 1991 D
Cornuscutellum SNAJDR, 1960 P
Decoroscutellum SNAJDR, 1958 L
 503
Dentaloscutellum CHATTERTON, 1971 Z
Eoscutellum MAKSIMOVA, LP?
Hidascutellum KOBAYASHI & HAMADA, 1977
Kobayashipeltis MAKSIMOVA, 1978 D?E?
Kolihapeltis PRANTYL & PRIBYL, 1947 PZD
Kotisopeltis VANEK, PEK & KUPKovA, 1991 L
Metascutellum SNAJDR, 1960 PZ?
Microscutellum SNAJDR, 1960 LPE
Paralejurus HAWLE & CORDA, 1847 LPDE
Platyscutellum SNAJDR, 1958 L?PZDE
Poroscutellum SNAJDR, 1958 P
Radioscutellum SNAJDR, 1972 P
Scabriscutellum RICHTER & RICHTER, 1956
P?ZDEG?
Scutellum PUSCH, 1833 EGF
Spiniscutellum SNAJDR, 1960 LP
Tenuipeltis LUTKE, 1965 G? F
Thysanopeltella KOBAYASHI, 1957 DE
Thysanopeltis HAWLE & CORDA, 1847 E
Unicapeltis MAKSIMOVA, 1977 ZD
Uraloscutellum MAKSIMOVA & ANCIGIN, 1978 P
Weberopeltis MAKSIMOVA, 1959 LPZ?D?E?
PROETIDAE Hawle & Corda, 1847
Proetinae Haw1e & Corda, 1847
Boliuiproetus ELDREDGE & ORMISTON, 1979 ZD
Coniproetus G. ALBERTI, 1966 LP
Dohmiella LUTKE, 1990 EG
Erbenites PRIBYL, 1964 D?E
Gerastos GOLDFUSS, 1843 LPZDEG
Hamiroproetus SNAJDR, 1977 PZ
Kegeliella LUTKE, 1990 PZ
Kolymoproetus MAKSIMOVA, 1978 P
Longiproetus CAVET & PILLET, 1958 EG
Myoproetus SNAJDR, 1976 Z?DE
Orbitoproetus PILLET, 1969 PZDEG
Podoliproetus SNAJDR, 1980 PZ
Pragoproetus SNAJDR, 1977 P
Proetina PRIBYL, 1946 P
Proetocephalus LUTKE,1977 ZID
Pudoproetus HESSLER, 1963 Fa
Simaproetus SNAJDR, 1960 P
Unguliproetus ERBEN, 1951 PZDE
Ryckholtia SNAJDR, 1980 L
Crassiproetinae OSMOLSKA, 1970
Crassiproetus STUMM, 1953 L?PZDEG
Hedstroemia PRIBYL & VANEK, 1978 L
Mannopyge LUDVIGSEN, 1987 D
Monodechella STUMM, 1953 G
Decoroproetinae ERBEN, 1966
Decoroproetus PRIBYL, 1946 LPZ
Eremiproetinae G. ALBERTI, 1967
Eremiproetus RICHTER & RICHTER, 1919 PZDEG
Koneprusites PRIBYL, 1946 Z?DE
Miriproetus SNAJDR, 1977 P
Nagaproetus SNAJDR, 1977 PE
Phaetonellus NovAK, 1890 ZDEG
Richteraspis G. ALBERTI, 1970 Fr
Trautensteinproetus PILLET, 1969 ZDE
Tropidocoryphinae PRIBYL, 1946
Alberticoryphe ERBEN, 1966 PZD?
Astycoryphe RICHTER & RICHTER, 1919 PZDEG
Bojocoryphe SNAJDR, 1976 P
Denemarkia PRIBYL, 1946 P
Erbenicoryphe FEIST & ClARKSON, 1989 Fr
Hollardia G. ALBERTI, 1964 LP
Phyllaspis RICHTER, 1863 PZ?D
Pribylia ERBEN, 1951 P
Prionopeltis HAWLE & CORDA Ubasel
Pterocoryphe FEIST, 1976 Fr
Pteroparia RICHTER, 1913 Fr
Ranunculoproetus SNAJDR, 1977 LP
Tropidocoryphe NovAK, 1890 PZDEG
Wolaye(la ERBEN, 1966 PZD?
Cornuproetinae Richter, Richter & Struve, 1959
Buchiproetus PILLET, 1969 ZDE
Cornuproetus RICHTER & RICHTER, 1919 LPZDEG
Cyrtosymboloides G. ALBERTI, 1967 ZDE
Dalejeproetus SNAJDR, 1977 LPZD
Ignoproetus SNAJDR, 1977 ZE
Lepidoproetus ERBEN, 1951 LPD
Macroblepharum G. ALBERTI, 1964 ZDE
Linguaproetus H. ALBERTI & RUCHHOLZ, 1964 DIE
Parapalpebralia PILLET, 1972 D
Perakaspis KOBAYASHI & HAMADA, 1972 DE?
Piriproetus ERBEN, 1952 DE
Prodreuermannia G. ALBERTI, 1969 LPZ
Proetopeltis PRIBYL, 1965 ZDE
Richteraspis G. ALBERTI, 1969 Fr
Quadratoproetus G. ALBERTI, 1967 D
Sculptoproetus ERBEN, 1951 LPZDE
Vicinoproetus G. ALBERTI, 1967 P
Xiphogonium HAWLE & CORDA, 1847 PZ?D
Warburgellinae OWENS, 1973
Parawarburgella MAKSIMOVA, 1975 L
Taynaella SPASSKY, 1989 L
Tcherkesouia SPASSKY, 1989 L
Tropidocare CHLUPAc, 1971 L
Warburgella REED, 1931 L
Dechenellinae PRIBYL, 1946
Basidechenella RICHTER, 1912 ZDE
Benesouella CHLUPAc, 1969 E
Borealia MAKSIMOVA, 1977 ZD
Camsellia ORMISTON, 1976 E
Chauffouraspis PILLET, 1972 Z?D?
CZauiboZe HAHN & BRAUCKMANN Fr
Cyrtodechenella RICHTER & RICHTER, 1950 EGFr?
Dechenella KAYSER, 1880 EG
Dechenellurus MAKSIMOVA, 1960 LPD
Deltadechenella ORMISTON, 1967 E
Eocyrtosymbole PILLET, 1972 ZD?
Fuscinipyge ORMISTON, 1972 E
Ganinella YOLKIN, 1968 LPZDE?
Humeia ORMISTON, 1975 E
Khalfinella YOLKIN, 1968 LPZDE
Lacunoporaspis YOLKIN, 1966 LZDE?
Monodechenella STUMM, 1953 EG?
Ormistoniella COOPER, 1982 DE
Paradechenella RICHTER, 1912 DEG
Pedinodechenella ORMISTON, 1967 E
Praedechenella MAKSIMOVA, 1953 PZD
Pseudodechenella PILLET, 1972 PZDEG?
Schizoproetina MAKSIMOVA, 1977 LPZD
Schizoproetoides ORMISTON, 1967 PE?
Schizoproetus RICHTER, 1912 ZDEG
Vitaella H. ALBERTI, 1976 Fa
Xenodechenella PEK & VANEK, 1991 D
Drevermanniinae MAKSIMOVA, 1960
Bapingaspis YUAN, 1988 Fr Fa
Chaunoproetus RICHTER & RICHTER, 1926 Fa
Daihuaia YUAN, 1988 Fa
 504
Drevermannia R. RICHTER, 1909 Fa
Formonia R. & Z. RICHTER, 1927 Fa
Frithjofia PRIBYL & VANEK, 1978 Fa
Palpebralia R. & E. RICHTER, 1927 Fr
Pontipalpebralia Liitke, 1968 Fa
Parafrithjofia Yuan , 1988 Fa
Cyrtosymbolinae Hupe, 1953 (and Archegoninae G.
Hahn & Brauckm ann, 1984)
Calybole RICHTER & RICHTER, 1926 Fa
Cyrtosy mbole RICHTER, 1913 Fa
Devonocoryph e YUAN , MS Fa
Haasia YUAN, 1988 Fa
Helioproetus RICHTER & RICHTER, 1919 Fa
Mirabole OSM6LSKA, 1962 Fa
Pseudowaribole HAHN & HAHN, 1967 Fa
Pusillabole H. ALBERTI , 1973 Fa
Skemmatopyge RICHTER, 1919 Fa
Typhloproetus RICHTER, 1913 Fa
Waribole RICHTER & RICHTER, 1926 Fa
Subfam. incerta:
Perliproetus RICHTER & RICHTER, 1926 Fa
AULACOPLEURIDAE Angelin , 1852
Aulacopleu rinae Angelin, 1852
Aulacopleura HAWLE & CORDA, 1847 PDEG
Cyphaspis BURMEISTER, 1843 PZDEGF r
Harpidella McCoy, 1849 LPZE?G?
Otarion ZENKER, 1833 LPZDEG?
Otarionella WEYER, 1965 LPZDE
Otarionides G. ALBERTI, 1969 P
Pseudotrinodu s KOBAYASCHI & HAMADA, 1971 DE?
Cyphaspidinae Pribyl, 1947
Cyp haspides NovAK, 1890 PZDEG
BRACHYMETOPIDAE Pran tl & Pribyl, 1950
Brachymetopinae PRANTL & PRIBYL, 1950
Asiagena MAKSIMOVA, 1978 L
Brachymetopus McCOY, 1847 Fa
Cordani a CLARKE, 1892 L P
Mystrocephala WHITTINGTON, 1960 DEG?
Tschernyshewella TOLLY, 1899 P?
Scharyiinae Osmol sk a , 1957
Scharyia PRIBYL, 1946 LPZDE?
RORRINGTONIIDAE Owen s & Hammann, 1990
Eodrevermanniinae Hupe , 1953
Aulacopleurina PRIBYL, 1949 DE
Baillielloides KOBAYASHI & HAMADA, 1972 DE
Eodrevermannia PRIBYL, 1946 L?PZDE
Erbenaspis G, ALBERTI, 1966 LPZD
Moravocoryph e PRIBYL & VANEK, 1978 D
PHACOPIDAE Ha wle & Corda, 1847
Phacopinae Ha wle & Cord a , 1847
A nanaspis CAMPBELL, 1967 L
Boeckops CHLUPAc, 1971 PZD
Chotecops CHLUPAc, 1971 ZDEGFa
Cryphops RICHTER & RICHTER, 1926 FrFa
Dianops RICHTER & RICHTER, 1923 Fa
Eocryphops RICHTER & RICHTER, 1931 DE
Kainops RA.'V!SKOLD & WERDELIN , 1991 LP
Lochkovella CHLUPAc, 1972 L
Nephranops RICHTER & RICHTER, 1926 G?FrFa
Pacipha cops MAKSIMOVA, 1972 LPZDE
Phacop s EMMRICH, 1839 PZDEGFrFa
Plagiolaria KEGEL, 1952 LPZDE?
Prokops CHLUPAc, 1971 L?P
Reedops RICHTER & RICHTER, 1925 LPZD
Rhinoreedops MAKSIM OVA, 1978 Z
S igna tops PRIBYL & VANEK, 1971 D
Tr imerocephalus McCoy, 1849 Fa
Viaphacops MAKSIMOVA, 1972 PZDE?G?
Ph acopidellinae Delo, 1935
Afrops G. ALBERTI, 1983 LIP
Altaesajania MAKSIMOVA, 1978 E
Dienstina RICHTER & RICHTER, 1931Fa
Ductina RICHTER & RICHTER, 1931 Fa
Illaenula CHLUPAc, 1977 DE?
S tru veaspis G. ALBERTI, 1966 DE
?Blanodalma nites KOBAYASHI & HA.\1ADA,
1972 DIE?
Echinoph acopinae ZHOU & CAMPBELL, 1990
Echinophacops ZHOU, 1983 D
Rhinophacops KANEKO, 1990 D
Toxophacops ZHOU & CAM PBELL, 1990 D
DALMANITIDAE Vogdes, 1890
Dalmanitinae Vogdes, 1890
Corycephalus HALL & CLARKE, 1888 L
Devonodontochil e SNAJDR, 1985 Z
Fran covichia BRANISA & VANEK, 1973 ZD
Gamod enaspis BRANISA & VANEK, 1973 ZD
Huntonia CAMPBELL, 1977 LP
Neoprobolium STRUVE, 1958 L?D
Odontochile HAWLE & CORDA, 1847 LPZD
Phalango cephalu s CAMPBELL, 1977 L
Reussiana SNAJDR, 1987 LP
Roncellia LESPERANCE, 1975 LP
Aste ropyginae Delo, 1935
Alcaldops ARBIZU, 1979 EG
A rgentopyge BALDIS , 1972 L?
Aste ropyge HAwLE & CORDA, 1847 DE
Bradocrypha eus HAAS & MENSINK, 1970 EGFr
Breizhops MORZADEC, 1983 D
Com ura RICHTER & RICHTER, 1926 ZD
Delocare STRUVE, 1958 ZD
Echinopyge HAAs, 1968 D
Feruminops HAAs . 1968 Z?D
Gourdonia PILLET, 1954 DE
Greenops DELO, 1935 DEGF
Heliopyge HAAs & MENSINK, 1970 EGFr
Hexacosta FARSAN, 1981 G
Kayserops DELO, 1935 ZD
Metacanthina PILLET, 1954 PZD
Neocalmonia PILLET, 1964 F
Neometacanthus RICHTER & RICHTER, 1948 E G
Paracryphaeus GANDL, 1972 LPZD?
Pilletina HAAs , 1970 PZD
Protacanth ina GANDL, 1972 LPZ
Pseudocrypha eus PILLET, 1954 LPZ
Psychopyge TERMIER & TERMIER, 1950 D
Radiopyge FARSAN , 1981 GFr
Rhenops RICHTER & RICHTER, 1943 ZD
S anidopyge Haas , 1968 EG?
Treveropyge STRUVE , 1958 LPZD
Synphoriinae Delo, 1935
Anchiopella REED, 1907 PZDE
Fenestraspis BRANISA & VANEK, 1973 D
Odontocephalus CONRAD, 1840 ZDE
Sy nphoria CLARKE , 1900 PZDE
Synphoroides DELO, 1940 LPZD
Trypaulitinae Lesp eranc e, 1975
Chacomurus BRANISA & VANEK, 1973 D
Coronura HALL & CLARKE , 1888 ZDE
Forillonaria LESPERANCE, 1975 P
 505
Schoharie LESPERANCE, 1975 PZDE
Trypaulites DELO, 1935 ZDE
Malladaiinae Arbizu, 1978
Malladaia OEHLERT & OEHLERT, 1896 P?ZD
Furacopyge ARBIZU, 1978 D
Acastavinae Struve, 1958
Acastava RICHTER & RICHTER, 1954 PZDE?
Acastella REED, 1925 L
Pelitlina HAAs, 1968 ZD
Neosynphoriinae Hupe, 1955
Centauropyge HAAs, 1968 E
Cryphina OEHLERT & OEHLERT, 1889 PZ
CALMONIIDAE Dela, 1935
Calmoniinae Dela, 1935
Australops BALDIS, 1972 L?
Bainella RENNIE, 1930 ZD
Bouleia KOZLOWSKI, 1923 ZOE
Calmonia CLARKE, 1913 ZD
Chiarumanipyge BRANISA & VANEK, 1973 0
Cryphaeoides DELO, 1935 EG
Deltacephalaspis ELDREDGE & BRANISA, 1980 ZO
Kozlowskiaspis BRANISA & VANEK, 1973 ZD
Malvinella WOLFART, 1968 Z?DE
Metacryphaeus REED, 1907 OEG
Oosthuizenella Cooper , 1982 0
Palpebrops LIEBERMA."l', EDGECOMBE &
ELDREDGE, 1991 Z?D
Parabouleia ELDREDGE, 1972 LZ
Paracalmonia STRUVE, 1958 DE
Pennaia CLARKE, 1913 Z?DE
Plesiomalvinella LIEBERMAN, EDGECOMBE &
ELDREDGE, 1991 DE
Probolops DELO, 1935 D
Punillaspis BALDIS & LONGOBUCCO, 1977 EG?
Renniella COOPER, 1982 D
Schizostylus DELO, 1935 ZOE
Tarijactinoides SUAREZ SORUCO, 1971 Z
Tibagya STRUVE, 1958 DE
Typhloniscus SALTER, 1856 DE?
Vogesina WOLFART, 1968 ZDE
Acastinae Dela, 1935
Acaste GOLDFUSS, 1843 LP
Acastellina RICHTER & RICHTER, 1954 DE
Acastoides DELO, 1935 LPZD
Morocconites STRUVE, 1989 D?
Phacopina CLARKE, 1913 L?PZDE?
CHEIRURIDAE Salter
Cheirurinae Salter, 1864
Crotalocephalides G. ALBERTI, 1967 P
Crotalocephalina PRIBYL & VANEK, 1964 L P
Crotalocephalus SALTER, 1853 LPZDEG
Didrepanon LANE, 1971 L
Mezocrotalus MAKsIMOVA, 1978 L
CALYMENIDAE Milne Edwards, 1840
Calymeninae Milne Edwards, 1840
Calymene BRONGNIART, 1822 LPZ?D?
Gravicalymene SHIRLEY, 1936 LPZ?D?
Nipponocalymene KA.~EKO, 1985 E?
Sthenarocalymene SIVETER, 1976-LP
HOMALONOTIDAE Chapman, 1890
Homalonatinae Chapman, 1890
Arduenella WENNDORF, 1990 PZ
Burmeisterella REED, 1918 PZ
Burmeisteria SALTER, 1825 ZD
Digonus GURICH, 1909 LPZD
Dipleura GREEN, 1832 LPZDEG
Parahomalonotus REED, 1918 LPZD
Scabrella WENNDORF, 1990 PZD
Trimerus GREEN, 1832 LPZD
HARPETIDAE Hawle & Carda, 1947
Harpetinae Hawle & Corda, 1847
Bohemoharpes VANEK, 1963 L
Harpes GoLDFUSS, 1839 EGFr
Helioharpes PRIBYL & VANEK, 1981 LPZDEG
Kielania VANEK, 1963 LPZDEG
Lioharpes WHITTINGTON, 1950 PZDEG Fr?
Reticuloharpes VANEK, 1963 ZDEG
LICHIDAE Hawle & Corda, 1847
Lichinae Hawle & Corda, 1847
Oinochoe THOMAS & Holloway, 1988 L
Echinolichinae Phleger, 1936
Ceratolichas HALL & CLARKE, 1888 E
Echinolichas GURICH, 1901 PZOE
Gaspelichas CLARKE, 1907 P
Terataspis HALL, 1863 E
Trochurinae Phleger, 1936
Acanthopyge HAWLE & CORDA, 1847 LPZDEG
Akantharges PHLEGER, 1936 E
Ceratarges GURICH, 1901 DEG?
Craspedarges GURICH, 1901 P
Eifliarges RICHTER & RICHTER, 1917 E
Lobopyge PRIBYL & ERBEN, LPZDE
Mephiarges RICHTER & RICHTER, 1930 E
Nipponarges KANEKO, 1984 E
Perunaspis PRIBYL, 1949 PZDE
Radiolichas REED, 1923 O?E?G
Richterarges PHLEGER, 1936 L?
Terranovia MAKSIMOVA, 1977 LPZD
ODONTOPLEURIDAE Burmeister, 1843
Odontopleurinae Burmeister, 1843
Diacanthaspis WHITTINGTON" 1941 L?PZ?D?
Dudleyaspis PRANTL & PRIBYL, 1949 LPZD
Kettneraspis PRANTL & PRIBYL, 1949 LPZDEG
Leonaspis RICHTER & RICHTER, 1917 LPZDE?
Radiaspis RICHTER & RICHTER, 1917 L?PZDEGF
Miraspidinae RICHTER & RICHTER, 1917
Ceratocephala WARDER, 1838 LPZDE
Ceratocephalina WHITTINGTON, 1956 LP
Ceratonurus PRANTL & PRIBYL, 1949 LPZDE?
Dicranurinae PRANTL & PRIBYL, 1949
Dicranurus CONRAD, 1841 LP
Kaneprusiinae VANEK & PEK, 1987
lsoprusia BRUTON, 1966 DEG
Koneprusia PRANTL & PRIBYL, 1949 LPZDEG
Laethoprusia RAMSKOLD, 1990 L?P
I. CHLUPAC
Department of Geology
Charles Univ ersity
Albertov 6
128 43 Praha 2, Czech Republic