GEOLOGIJA 28/29, 9—53 (1985/86), Ljubljana
UĐK 562.02:551.735.736(497.12)(450.2/4) = 20
Upper Carboniferous and Permian mesolobid chonetacean
brachiopods of Karavanke Mountains (Yugoslavia)
and Carnian Alps (Italy)
Zgornjekarbonski in permijski mezolobidni honetacejski brahiopodi
V Karavankah (Jugoslavija) in Karnijskih Alpah (Italija)
Janez Pečar
Graden 2, 61111 Ljubljana
Abstract
In the preface the definition of terms mesolobid and mesolobidity is
given. In Gzhelian of the Karavanke Mountains two distinct forms of rib-
bed mesolobid chonetacean brachiopods were found in stratigraphically
the same beds. This fact gives evidence for parallel evolution of two rib-
bed mesolobid chonetacean stocks. One of them is genus Paramesolobus
Afanasjeva, 1975, the other Capillomesolobus n. gen. Both genera are also
present in the same beds of Kasimovian of the Carnian Alps. The stra-
tigraphical distribution of mesolobid chonetacean genera and species on
world scale is given. Paleobiogeography and migration of genus Capil-
lomesolobus is outlined. New taxa are Capillomesolobinae n. subfam., Ca-
pillomesolobus karavankensis n. gen., n. sp. (Karavanke Mountains, Gzhe-
lian), C. pontebbanus n. gen., n. sp. (Carnian Alps, Corona Formation,
Kasimovian) and C. heritschi n. gen., n. sp. (Karavanke Mountains, Trog-
kofel beds, Sakmarian or Artinskian).
Kratka vsebina
Uvodoma je podana definicija izrazov mezolobiden in mezolobidnost.
V Karavankah sta bili v stratigrafsko istih gželijskih plasteh najdeni dve
obliki rebrastih mezolobidnih honetacejskih brahiopodov. To kaže na vzpo-
reden razvoj dveh rebrastih mezolobidnih honetacejskih vej. Ena od teh
je rod Paramesolobus Afanasjeva, 1975, druga rod Capillomesolobus n. gen.
Oba rodova sta bila najdena tudi v stratigrafsko istih plasteh v kasimo-
viju Karnijskih Alp. Podana je stratigrafska porazdelitev mezolobidnih
honetacejskih rodov in vrst v svetovnem merilu. Kratko je podana paleo-
biogeografija rodu Capillomesolobus. Novi taksoni so Capillomesolobinae
n. subfam., Capillomesolobus karavankensis n. gen., n. sp. (Karavanke,
gželij), C. pontebbanus n. gen., n. sp. (Karnijske Alpe, formacija Corona,
kasimovij) in C. heritschi n. gen., n. sp. (Karavanke, trogkofelske plasti,
sakmarij in artinskij).
2____ 		Janez Pečar
Introduction
Among the chonetacean brachiopods (superfamily Chonetacea) of Upper
Carboniferous and Permian of the Karavanke Mountains (Slovenia) the chone-
tacean forms with ventral median lobe in median sulcus were also found. It
was realised that they belong to two genera. Both genera were also identified
in the Upper Carboniferous of the Carnian Alps. Thus Karavanke and Carnian
Alps are thought to be the important areas for study of the mesolobid choneta-
cean brachiopods.
In the present paper the morphological terms "mesolobid" and "mesolobi-
dity" are used for those Carboniferous and Permian chonetacean brachiopods
in which in the middle of the ventral median sulcus the median lobe or fold
is present.
Investigations of mesolobid chonetacean forms on generic level
The first described mesolobid chonetacean brachiopod species was Chonetes
mesolohus Norwood et Pratten, 1855 from Pennsylvanian of USA. The presence
of median lobe in ventral median sulcus of chonetacean brachiopod was thus
recognised on the species level. On generic level this was done with erection
of genus Mesolohus Dunbar et Condra, 1932. Type species for genus Mesolohus
was Chonetes mesolohus Norwood et Pratten which was originally described and
illustrated as capillate species. Weiler et McGehee then (1933, 109)
pointed out that in the area from which the material of Norwood and
Pratten was derived, there was no capillate form of genus Mesolohus, but
only the smooth one. Because of it Weiler et McGehee proposed that
capillate specimens of Mesolohus are placed in species Mesolohus striatus Weiler
et McGehee, 1933, and that smooth species Mesolohus mesolohus lioderma Dun-
bar et Condra, 1932 is the type species of genus Mesolohus. Ho are (1964,
315) thought that the exterior of Mesolohus mesolohus decipiens Dunbar et
Condra, 1932 is more similar to the original Chonetes mesolohiLs Norwood et
Pratten than the variety lioderma. He proposed the specimen of variety deci-
piens for the neotype of Mesolohus mesolohus. This application was approved
by the International Commission on Zoological Nomenclature. Thus genus
Mesolohus was typically smooth. It was logically to place the ribbed mesolobid
chonetacean forms in another genus. In this way Afanasjeva (1975, 101)
erected the genus Paramesolohus. She placed in this genus also the North
American ribbed mesolobid chonetacean species. But Afanasjeva (1975,
103) and earlier Ivanov et Ivanova (1936, 21) noted that the mesolobid
chonetacean forms from the Russian Platform arc larger and with stronger
ribs than the North American Pennsylvanian species.
The material from the Upper Carboniferous of the Karavanke Mountains
and the Carnian Alps revealed two distinct ribbed mesolobid chonetacean forms
in the same beds. One of these is genus Paramesolohus Afanasjeva, 1975. The
other forms, which is smaller and with finer ribs than the first is Capillome-
solohus n. gen. These facts give evidence for parallel evolution of two ribbed
mesolobid chonetacean stocks (Pecar, 1986, 137) differing not only on the
generic but probably also on the subfamily level. The process of generic de-
signation of the mesolobid chonetacean brachiopods is outlined on fig. 1.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 11
Fig. 1. The development of the generic designation of mesolobid chonetacean
brachiopods
SI. 1. Razvoj rodovne opredelitve mezolobidnih honetacejskih brahiopodov
Previous investigations of mesolobid chonetacean brachiopods in Carnian Alps,
Karavanke Mountains and in nearby countries
Schellwien (1892, 29, 30) described from the Upper Carboniferous of
the Carnian Alps two species now placed in genus ParamesolohiLs; these were
Chonetes lohatw; and Chonetes latesiniiatus. Later S c h e 11 w i e n (1898, 360)
gave to the species C. lohatus the new name C. sinuosus because the name lo-
batiis was preoccupied by C. lobatus Grünewaldt, 1860. Gortani (1905, 538)
described from the Carnian Alps subspecies Chonetes moelleri carnicus. In the
Lower Permian Trogkofel beds of the Karavanke Mountains the mesolobid
chonetacean was identified (S c h e 11 w i e n , 1900, 38) as Chonetes sinuosus.
Later H e r i t s c h (1938, 103) ascribed this form to a new species, but he did
not name it formally. H e r i t s c h (1931, 10) found Chonetes latesinuatus in the
Upper Carboniferous of the Karavanke Mountains. This species was described
also from the Upper Carboniferous of Croatia (Kostić-Podgorska, 1949,
79) and Hungary (R a k u s z , 1932, 59).
Localities, stratigraphy and age
The localities 1 to 5 are in the Karavanke Mountains (Slovenia, northwest
Yugoslavia), the locality 6 in the Carnian Alps (northeast Italy: figs. 2, 3).
Locality 1. Spodnja počivala. Long known fossil locality, situated in Javor-
niški rovt, about 1.8 km to the north of the village of Javornik (the suburb of
the town of Jesenice), 0.55 km N 62» E of the mountain Spik 967 m. The locality
is situated about 100 m along the water pipeline, above the lower crossing of
the latter with the road from Javornik to Javorniski rovt. Dark-grey to black
shale contains rich marine fauna. In intercalated marly shale Rugosofusulina
alpina antiqua (Schellwien) and Archaeolithophyllum missouriensum Johnson
were found indicating the Gzhelian age (Ramovš, 1971, 1389; Hahn et al.,
12		 		Janez Pečar
Fig. 2. Location map of the brachiopod collecting sites
SI. 2. Položajna skica brahiopodnih najdišč
1977, 138). First paleontological contributions from this locality were those of
Heritsch (1919, 1931) and of Rakovec (1931). Later Ramovš with co-
authors made several paleontological articles from this locality (H a h n G. et al„
1977).
Locality 2. Planina pod Golico. At the third pillar of the cableway to Spanov
vrh. About 2.5 km to the north of the town of Jesenice, 1.15 km S 85"E of the
mountain Kogel 1122 m. A small outcrop was discovered by J. Bedič, it is about
2 m to the north of the pillar. The brownish, partially weathered shale is thought
to be of the Upper Carboniferous age, probably Gzhelian.
Locality 3. Planina pod Golico. Near the fifth pillar of the cableway to Spa-
nov vrh. This secondary occurence is about 250 m to the east of the locality
2 and topographically some tens of m higher. The age of the dark-grey shale
with numerous sponges is supposed to be the same as for locality 2.
Locality 4. Planina pod Golico. Between the sixth and seventh pillar of the
cableway to Spanov vrh. Secondary occurrence, about 100 m to the east of lo-
cality 3 and some tens of m topographically higher. The lithology is the same
and the age is supposed to be the same as for locality 3.
It must be mentioriied that detailed stratigraphy of localities 1 to 4 is un-
known, because the beds are not well exposed.
Locality 5. Dolžanova soteska (= Dolžan Gorge). Classical locality of the
Trogkofel Limestone in Karavanke Mountains. About 2.7 km to the northeast
of town of Tržič, 1.07 km S 690E from mountain Samuha 1172 m. The road from
Tržič to Jelendol, about 100 m farther of the second road curve after the road
tunnel, on the east side of the road. Rose-red, reddish and greyish reef lime-
stone in the abandoned quarry is extending as rocky wall along the road, it is
rich in brachiopods and other fossils. This roof limestone is preliminarily named
Unit B in the present paper. Grey shale (preliminarily named Unit A) is under-
lying and the bedded limestone (preliminarily named Unit C) is overlying. Above
Upper Carboniferous and Permian mesolobid chonetacean brachiopods
 13
fossil locality
Fig. 3. Detailed location maps A, B, and C, taken from fig. 2
SI 3. Podrobnejše položajne skice A, B in C, vzete iz si. 2
Unit C occurs probably another unit of reef limestone of similar colour as Unit
B. The lower part of the Trogkofel Limestone in Dolžanova soteska is briefly
stratigraphicaly outlined on fig. 4. This division is based partly on earlier strati-
graphycal statements (Teller, 1903; Ramovš, 1961) and on personal ob-
servations. Among foraminifers Pseudofusidina rakovcci Ramovš et Kochansky-
Devidé and Robustoschwagerina schellwieni (Hanzava) are present in the Trog-
kofel Limestone (Ramovš, 1978, 121). The age is considered to be Sakmarian
(Waterhouse, 1976, 82; Leven, 1980, pl. 3) or Artinskian (B u s e r , 1979,
20). S C h e 11 w i e n (1900) described the brachiopods from this locality.
14		 		Janez Pečar
Fig. 4. Stratigraphy of the lower part of Trogokofel Limestone in Dolža-
nova soteska
SI. 4. Stratigrafija spodnjega dela trogkofelskega apnenca v Dolžanovi
soteski
Locality 6. To the west of Monte Corona. About 200 m to the west of the
base of Monte Corona (= Kronalpe = Krone), 0.74 km N 83®W from the top
of Monte Corona 1832 m (V en t u r i n i, 1982, pi, 1), in raised western part of
the mountain saddle. It is in Italy, a few m from the Italian-Austrian border.
Lithologically it is brownish, weathered shale, with lime addition in some pieces
of the sample. The thickness of foss'liferous beds is probably less than 1 m.
They are rich in marine invertebrates. Dark grey shale is imderlying, it is about
20 m thick, appearingly without fossils. Overlying is the shale of thickness of
some m, above which lies the quartz conglomorate up to 5 m thick. There are
faults between the locality 6 and the base of Monte Corona. (Venturini,
1982, pl. 1). The area of the locality fi belongs, according to Selli (1963, 51), to
Formazione del Corona (Corona Formation). Selli (1963, 51) cited the Gzhelian
age for this formation. F r a n c a v i 11 a (1974, 93) ascribed Stephanian A (pro-
bably its upper part) or Stephanian B age to the land flora of his locality 119,
which belongs to Corona Formation in the vicinity of the present locality 6.
Venturini et al. (1982, 311) designated the age of Corona Formation as
Stephanian B which is corresponding with Kasimovian (Rotai, 1978, 11).
During the field work in 1985 it was not possible to find the S c h e 11 w i e n ' s
(1892, 8) bed 6 of his Krone scction: from this important bed he described among
others the mesolobid brachiopod species Chonetes sinnosus vSchellwien, 1898.
It is possible that the locality 6 is very near to Schellwien's bed 6 of his
Krone section both geographically (the horizontal distance to some 200 m) and
stratigraphically.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 15
Preservation of brachiopods
In the Upper Carboniferous localities 1 to 4 and 6 the brachiopods are pre-
served mainly as moulds, rarely as shells. Numerous biota (mostly bryozoans)
are found on the moulds of Upper Carboniferous brachiopods, especially from
locality 2. This means that biota lived inside shell structure of brachiopods,
perhaps after brachiopod death. In the Permian locality 5 the brachiopods are
preserved as shells and as moulds.
Collections
Brachiopods figured herein are reposited at the Katedra za geologijo in pa-
leontologijo, Univerza Edvarda Kardelja v Ljubljani, Ljubljana (KGPL), at
Tehniški muzej Železarne Jesenice, Jesenice (TMJ), and at Museo Friulano di
Storia Naturale, Udine (GPU).
Terminology
It was used that of Muir-Wood (1962, 7; 1965, 412) with addition of
terminology of Cooper et Grant (1975, 1212). According to the termino-
logy of Muir-Wood the following terms are used for ribs: costate, if in
the middle part of the ventral anterior margin there are 15 ribs or less on
10 mm of the shell width; costellate, if in the same area there are 16 to 25 ribs;
capillate, if 26 ribs or more are in the same area.
Measurements, diagrams, statistical analysis, abbreviations
Explication is needed for measurement and calculation of the number of
ribs at the width of 10 mm in the anterior margin of the pedicle valve. In the
middle of the ventral anterior margin the width of 2 to 4 ribs was measured
with the ocular micrometer. From this value the number of ribs per 10 mm
of the shell width was calculated for each specimen with the use of the percent
calculation. For example, the width of 3 ribs in ocular micrometer is 34 micro-
meter units. The micrometer value of particular magnification of a particular
microscope was measured and calculated earlier; in our case it was 0.031746 mm.
Further calculation: 34 . 0.031746 mm = 1.079364 mm. So 3 ribs occur at the
width of 1.079364 mm; with percent calculation we get 3 : 1.079364 = X : 10,
X = 27.7 ribs. The chonetacean specimen is capillate, because it has more than
25 ribs per 10 mm of the ventral anterior margin. It is suitable to prepare the
table of already calculated data for particular microscope magnification, from
which we can get the result at a glance. In the mentioned example the width
of 3 ribs is 34 micrometer units, and it can be read from the table that it
corresponds to 27.7 ribs per 10 mm of shell width.
The number of ribs per 10 mm of shell width is useful for quick quantitative
description of ribs of chonetacean brachiopods, which can be costate, costellate
or capillate. It is useful even for specimens with the whole width less than
10 mm. In the present paper it is thought that the exact method of rib measur-
ing with ocular micrometer is quicker than the counting of the number of ribs
16	Janez Pećar
on the whole specimen and more exact than the counting of ribs on 1, 2 or 5 mm
of the shell width. Many times it is impossible to measure the entire number
of ribs on ventral valve because they are not all preserved, but it is usually
possible to measure 2 to 4 ribs in the middle of the ventral anterior margin.
On the other hand the weak point of this parameter in the present paper is the
absence of correlation between it and the shell length.
In the present paper the independent variable (on the X axis) is the length
of the ventral valve. For each diagram the correlation coefficient (r) for speci-
mens of the particular locality was calculated if the number of specimens was
sufficent. When r was significant (P < 0.05) the regression line was calculated
too (Spiegel, 1975, 263, 270).
In diagrams the following abbreviations are used:
L	length of the ventral valve
W maximal width of the valve
Nr 10 number of ribs per 10 mm of the width, measured in the ventral anterior
margin
Nr total number of ribs of ventral an+erior margin.
Systematic paleontology
Class Articulata Huxley, 1869
Order Strophomenida Öpik, 1934
Suborder Chonetidina Muir-Wood, 1955
Superfamily Chonetacea Bronn, 1862
Family Rugosochonetidae Muir-Wood, 1962
Diagnosis of this family is provided by Hoover (1981, 49) and Arch-
bold (1892, 2).
Capillomesolobinae n. subfam.
Diagnosis: Small, capillate or smooth Rugosochonetidae with ventral
median sulcus, in which ^^he median lobe is present or absent.
Genera assigned to subfamily: Capillomesolohus n. gen. Meso-
lohus Dunbar et Condra, 1932. Tenuichonetes .Ting et Hu, 1978. New undescribed
genus, typified by Neochonetes unhonopUcatus, figured by Barkhatova,
1964, from the Sakmarian Nenets Beds, Sula River, Northern Timan Mountains
(A r c h b o 1 d , 1982, 2) possibly also belongs to this subfamily.
Discussion and phylogenesis: Capillomesolobinae n. subfam.
differs from other subfamilies of family Rugosochonetidae by usual presence
of ventral median lobe in ventral median sulcus. Material in the present paper
reveals that the presence or absence of mesolobidity is somewhat variable
intraspecifically.
Genera Capillomesolohus, Mesolohus and Tenuichonetes are thought to have
phylogenetic connection, in which Capillomesolohus has starting-point po-
sition (fig. 5). This genus appeared in Pennsylvanian (Atokan), probably it
descended of genus Neochonetes (Sutherland et Harlow, 1973, 31),
it lived until Upper Permian (Punjabian). It was a cosmopolitan genus. From
genus Capillomesolohus derived the endemic North American Pennsylvanian
smooth genus Mesolohus and the likewise endemic Chinese Middle Permian
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 17
Fig. 5. The inferred phytogeny of mesolobid chonetacean brachiopods
SI. 5. Domnevna filogeneza mezolobidnih honetacejskih brahiopodov
capillate genus Tenuichonetes. In all these three genera the basic morphological
characteristics are finely capillate or smooth shell and ventral median lobe
in ventral median sulcus (mesolobidity). The possibility that Tenuichonetes
evolved from either Mesolohus or Paramesolobus was stated earlier by A r c h -
bold (1982, 6). In the present paper it is thought that it possibly evolved from
Capillomesolobus. Genus Tenuichonetes has also plications or lobes on lateral
parts of the shell. In the original description of the type species of genus
Tenuichonetes (Chonetes tenuiliratus Chao, 1928) there is neither a description
of mesolobidity nor it can be reliably seen on plate. Mesolobidity of T. tenuili-
ratus was described by J ing et Hu (1978, 125) and of T. plicatiformis
described and figured by Lee (1962, 486). The morphology of the fourth
possible, undescribed genus of this subfamily, typified by Neochonetes unbono-
plicatus, is peculiar: ventral median sulcus in posterior part changes anteriorly
to a swollen fold, separated from the lateral flanks of the valve by a valley on
either side (A r c h b o 1 d , 1982,2).
The material from Upper Carboniferous of Karavanke Mountains and Carni-
an Alps shows clearly the difference between finely ribbed mesolobid chonetace-
an stock (genus Capillomesolobus) and coarsely ribbed stock whose represen-
tatives are also larger (genus Paramesolobiis). Both genera are present in
Karavanke Mountains and in Carnian Alps in the same beds; this fact gives
the first doubtless evidence for two ribbed mesolobid chonetacean stocks
(Pecar, 1986, 137) with parallel evolution. Because of the striking difference
between Paramesolobus and Capillomesolobus they are separated on the sub-
family level in the present paper. It is possible that their mesolobidity is a
phenomenon of convergence. The predecessor of Paramesolobus is unknown.
2 - Geologija 28/29
18		 		Janez Pečar
Fig. 6. Stratigraphie distribution of mesolobid chonetacean brachiopods on the
world scale	" _
SI. 6. Stratigrafska porazdelitev mezolobidnih honetacijskih brahiopodov v svetovh^n
merilu	^
Genus Capillomesolohus n. gen.: 1 C. striatus (Weller & McGehee), 2 C. îhflexus
(Girty), 3 C. depressus (Stevens), 4 C. obsoletus (Sturgeon & Hoare), 5 C. profundus
(Sutherland & Harlow), 6 C. pontebbanus n. sp., 7 C. karavankensis n. sp., 8 Capillome-
solobus sp. (Gobbett, 1963, 120), 9 Capillomesolobus sp. (Yanagida, 1967, 86), 10 Capil-
lomesolobus sp. (Nakamura, 1959, 205), 11 C. heritschi n. sp., 12 Capillomesolobus (?)
sp. (Tazawa, 1976, 184), 13 C. permianus (Cooper & Grant), 14 Capillomesolobus sp.
(Coogan, 1960, 291), 15 C. (?) lissarensis (Diener), 16 Capillomesolobus sp. (Hayasaka,
1925, 93), Genus Mesolobus Dunbar & Condra, 1932: 17 M. mesolobus (Norwood &
Pratten), 18 M. lioderma (Dunbar & Condra), 19 M. euampygus (Girty), 20 M. rochel-
lensis R. H. King, 21 M. indistinctus Stevens. New genus, undescribed (typified by
Neochonetes unbonoplicatus, figured by Barkhatova, 1964, cited by Archbold, 1982,
2): 22. Genus Tenuichonetes Jing & Hu, 1978; 23 T. tenuiliratus (Chao), 24 T. plicati-
formis (Lee). Genus Paramesolohus Afanasjeva, 1975: 25 P. sinnosus (Schellwien), 26
P. latesinuatus (Schellwien), 27 P. carnicus (Gortani), 28 Paramesolohus sp. 1 (present
paper), 29 Paramesolohus sp. 2 (present paper), 30 P ivanovae (Afanasjeva), 31 P.
luganicus Aisenverg, 32 P. miaokouensis (Chao)
Stratigraphical distribution of species of Capillomesolobinae n. subfam. and
of genus Paramesolohus is shown on fig. 6. Of great assistance for elaborating
this figure were the data of Afanasjeva (1978) and Archbold (1982).
Capillomesolohus n. gen.
1986 New genus A — Pecar, p. 137.
Type species: Capillomesolohus karavankensis n. sp.
Etymology: Capillus (lat.) = hair, because of capillate shell surface;
mesolobus, because of morphological similarity with genus Mesolohus.
Diagnosis : Small, finelly capillate Rugosochonetidae with ventral me-
dian sulcus in which median lobe is present or absent.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 19
Description : Small, subrectangular to subtrapezoidal, shell width larg-
er than length. Cardinal extremities moderately acute angled to obtuse-angled.
Lateral margins oblique, rounded. Anterior margin medially indented. Surface
finely capillate. Growth lines weak. Hinge spines divergent, near right-angled
to right-angled. Pedicle valve exterior moderately to markedly convex. Median
sulcus originating behind the umbonal region, widening anteriorly, extending
to anterior margin. In sulcus median fold is present or absent. Brachial valve
exterior concave. Moderately to markedly pronounced median fold in which
median sulcus is present or absent.
Pedicle valve interior with short median septum, continuing sometimes an-
teriorly in low ridge. Adductor and diductor scars weak. Inner surface with
radial rows of taleolae which are larger on lateral and anterior margin of
visceral area. On lateral and anterior margin of valve taleolae are small.
Brachial valve interior with short, wide cardinal process, exteriorly quadri-
lobed, interiorly bilobed. Median septum long, high. Anderidia short, thin. Inner
surface taleolate, in anterolateral region of visceral area papillose. Inner surface
of lateral and anterior margins with small taleolae.
Comparison : Capillomesolobus differs from Neochonetes Muir-Wood,
1962 by deeper ventral median sulcus and usual presence of median lobe in it.
It resembles much the smooth genus Mesolohus Dunbar et Condra, 1932 in size
and morphology, differing from it in capillate surface. It is like Paramesolobus
Afanasjeva, 1975 in having often the ventral median lobe in sulcus, but Para-
mesolobus is larger, with coarser ribs and is more frequently without ventral
median lobe than Capillomesolobus. Both genera sometimes have a nodule on
ventral median lobe (pi. 1.14—1.15, 5.5—5.6; Stevens, 1962, pi. 93, figs. 12—
13). Chonetinella Ramsbottom, 1952 differs from Capillomesolobus in absence
of ventral median lobe in sulcus and in coarser ribs.
Species assigned to genus: Mesolohus striatus Weiler et Мс-
Gehee, 1933, North America (Ohio, Illinois, Missouri, Colorado, New Mexico),
Atokan and Desmoinesian. Chonetes mesolohus inflexus Girty, 1927, North
America (Idaho, Colorado River Valley), Pennsylvanian (Atokan?). Mesolohus
depressus Stevens, 1962, North America (Colorado), Atokan. Mesolohus ohsoletus
Sturgeon et Hoare, 1968, North America (Ohio), Desmoinesian. Mesolohus pro-
fundus Sutherland et Harlow, 1973, North America (New Mexico), Desmoinesian.
Capillomesolobus pontebbanus n. sp., Europe (Carnian Alps), Kasimovian. Ca-
. pillomesolohus karavankensis n. sp., Europe (Karavanke Mountains), Gzhelian.
Capillomesolobus heritschi n. sp., Europe (Karavanke Mountains), Sakmarian-
Artinskian. Mesolohus? permianus Cooper et Grant, 1975, North America (West
Texas), Upper Leonardian. ?Chonetes lissarensis Diener, 1897, Himalaya, Pun-
jabian.
Specifically undetermined statements of genus Capillomesolobus: Mesolohus?
sp., Gobbett, 1963, 120, Svalbard (Spitzbergen), Asselian?. Mesolobus me-
solohus (Norwood et Pratten), Nakamura, 1959, 205, Japan (Honshu),
Sakmarian. Chonetes cf. C. latesinuatus Schellwien, Y a n a g i d a , 1967, 86,
Indochina (Thailand), Sakmarian. Mesolohus siniLOSus (Schellwien), Tazawa,
1976, 184, Japan (Honshu), Kazanian?. Chonetinella cf. C. sinuosa Schellwien,
Coogan, 1960, 291, North America (California), Lower Guadalupian. Chone-
tes sinuosus Schellwien, Hayasaka, 1925,93, Japan (Honshu), Punjabian.
20		 		Janez Pečar
Paleobiography and migration: In beds of Atokan age in New
Mexico two species of Neochonetes with intermediate characters between
Neochonetes and Capillomesolobus (the evolution of ventral median lobe) were
described (Sutherland et Harlow, 1973, 26, 27). One of these species,
N. whitei, is stratigraphically the possible predecessor of Capillomesolohus. The
oldest known species of genus Capillomesolohus, C. striatus (Weiler et McGehee,
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 21
Fig. 7. Paleobiogeography of the genus Capillomesolobus
1 North America (USA), 2 North Africa (the possible Upper Paleozoic position of pre-
sent Karavanke Mountains and Carnian Alps), 3 Svalbard (Spitzbergen), 4 Indochina.
5 Central Japan (Honshu), 6 Nepal, 7 West Texas, 8 North California (cratonic block
Sonomia)
A: 1 C. striatus, C. infletus, C. depressus, C. obsoletus, C. profundus; 2 C. pontebbanus
C. karavankensis. B: 2 (#) C. pontebbanus, C. karavankensis; 2 (O) C. heritschi; 3
Capillomesolobus sp. (Gobbett, 1963, 120); 4 Capillomesolobus sp. (Yanagida, 1967, 86);
5	Capillomesolobus sp. (Nakamura, 1959, 205). C: 2 C. heritschi; 3 Capillomesolobus
sp. (Gobbett, 1963, 120); 4 Capillomesolobus sp. (Yanagida, 1967, 86); 5 (#) Capillome-
solobus sp. (Nakamura, 1959, 205); 5 (O) Capillomesolobus sp. (Tazawa, 1976, 184);
6	C. (?) lissarensis; 7 C. permianus: 8 Capillomesolobus sp. (Coogan, 1960, 291). Maps
(Mollweide projection) are modified from Scotese et al. (1979) with addition of data
from C. A. Ross et J. R. P. Ross (1985)
SI. 7. Paleobiogeografija rodu Capillomesolobus
1 Severna Amerika (ZDA), 2 severna Afrika (možni zgornjepaleozojski položaj seda-
njih Karavank in Karnijskih Alp), 3 Svalbard (Spitzbergi), 4 Indokina, 5 osrednja
Japonska (Honshu), 6 Nepal, 7 zahodni Texas, 8 severna Kalifornija (kratonski blok
Sonomia). Karte (Mollweidova projekcija) so spremenjene po Scotese et al. (1979)
z dodatkom podatkov iz C. A. Ross et J. R. P. Ross (1985)
1933) and C. inflexus (Girty, 1927), are from Atokan of North America (fig. 5).
On base of these data it can be concluded that genus Capillomesolohus origina-
ted in Atokan in North America. It became temporarily extinct there in Upper
Desmoinesian (Upper Moscovian). The oldest locality of Capillomesolohus out-
side North America is in the Carnian Alps where C. pontehhanus n. sp. is
present in Kasimovian. So it can be supposed that Capillomesolohus migrated
from the area of its origin in North America (west coast of Pangaea) to the area
22		 		Janez Pečar
of Carnian Alps (east coast of Pangaea). But how this first step of migration
of Capillomesolobus was possible? All possible directions of migration betwen
North America and the east coast of Pangaea were impeded during Atokan and
Missourian (or during Moscovian and Kasimovian) by biogeographic barriers
(fig. 7 A) : (1) The land mass (Pangaea) closed the route to the east; it is thought
that Euramerica and Gondwana joined together before Atokan (possibly as early
as in Chesterian time; C. A. Ross et J. R. P. Ross, 1985, 28), when
Capillomesolobus evolved. (2) Migration toward northeast and then to the east
was somewhat hampered by cold-water currents; this barrier was possibly
not so important because the genus was found also in the Boreal Realm (Gob-
bett, 1963, 120; Afanasjeva, 1978, 105). (3) Migration toward south
was also inhibited by low temperatures and very long route. (4) Deep ocean
closed the route on the west side of Pangaea, but there were also present cra-
tonic blocks (C. A. Ross et J. R. P. Ross, 1985, 27) which possibly had
some role in migration of Capillomesolobus.
Another less probable possibility is that finely capillate mesolobid forms
originated independently in Pennsylvanian of North America and in Carboni-
ferous in Europe.
Further steps of migration of Capillomesolobus are somewhat easier explain-
ed (fig. 7 B and C). Capillomesolobus was present in the Lower and Middle
Permian at cratonic blocks Honshu and Sonomia, respectively. Summarily it
can be said that genus Capillomesolobus was cosmopolitan, present on the east
and west coast of Pangaea and also on cratonic blocks in Panthalassa. In
comparison with it the genus Paramesolobus was restricted both biogeographi-
cally (to the east coast of Pangaea) and stratigraphically (to Upper Carboniferus
and posibly to lower part of Lower Permian). Thus the separation of mesolobid
chonetacean genera Capillomesolobus, Mesolohus and Paramesolobus has im-
portant biostratigraphic and paleobiogeographic implications.
Capillomesolobus karavankensis n. sp.
PI. LI—L25; fig. 8 A, B; fig. 9 A, B, C
1986 New genus A sp. 1 — Pecar, p. 137, 138, pi. 1, figs. 23—28.
Etymology : Karavankensis — after Karavanke Mountains, the area of
the localities of this species.
Types: Holotype: TMJ 1274. Paratypes: KGPL 5356, TMJ 1219, 1212,
1282, KGPL 5411, TMJ 1278.
Type locality, type horizon and age: Spodnja počivala (lo-
cality 1), Karavanke Mountains. Dark-grey to black shale. Gzhelian.
Localities and material: Locality 1, Gzhelian, 15 specimens (2
conjoined valves, 9 pedicle valves, 5 brachial valves). Locality 2, Gzhelian, 23
specimens (12 pedicle valves, 11 brachial valves). Locality 3, Gzhelian, 1 speci-
men (pedicle valve).
Diagnosis : Medium size Capillomesolobus with moderately expressed
ventral median lobe and moderately pronounced ventral lateral plications.
Description : Medium size for genus, subrectangular to subtrapezoidal,
shell width greater than length, widest shortly before hinge or at hinge. Cardi-
nal extremities moderately acute-angled to obtuse-angled, ears moderately
Upper Carboniferous and Permian mesolobid chonetacean brachiopods
 23
Fig. 8. Capillomesolobus karavankensis n. gen., n. sp.
A pedicle valve, mould of interior X 8. 1, TMJ 1274, locality 1, Karavanke
Mountains, Gzhelian; B brachial valve, mould of interior X 8. 2, TMJ
1224'1, locality 2, Karavanke Mountains, Gzhelian
SI. 8. Capillomesolobus karavankensis n. gen, n. sp.
A pecljeva lupina, odtis notranjosti X 8. 1, TMJ 1274, najdišče 1, Karavan-
ke, gželij; B ramenska lupina, odtis notranjosti X 8. 2, TMJ 1224/1, najdi-
šče 2, Karavanke, gželij
Fig. 9. Diagrams of the genus Capillomesolobus
A length against maximal width; B length against number of ribs on 10 mm of width
at ventral anterior margin; C length against total number of ribs at ventral anterior
margin
At A the regression line for ( + ) is W = 1.49 + ] .31 L, r = +0.82, P < 0,02
SI. 9. Diagrami rodu Capillomesolobus
A dolžina proti največji širini; B dolžina proti številu reber na 10 mm širine spred-
njega robu pecljeve lupine; C dolžina proti celotnemu številu reber sprednjega robu
pecljeve lupine
Pri A je regresijska premica za (+) W = 1.49 + 1.31 L, r = +0.82, P <0.02
18__ 		Janez Pečar
pronounced. Lateral margins oblique, rounded, anterior margin medialy widely
indented. Surface capillate, in the middle of ventral anterior margin from
52.8 to 73.0 capillae per 10 mm of shell width (fig. 9 A, B, C). Number of capillae
increasing on pedicle valve mostly with bifurcation, on brachial valve mostly
with intercalation. Growth lines weak. On each side 4 to 5 hinge spines, which
are divergent to perpendicular, their angle to posterior margin is 65" to 90".
Pedicle valve exterior rather convex in lateral profile, mostly in the proste-
rior third of the valve length. Median sulcus originating after umbonal region,
moderately deep, widening anteriorly, extending to anterior margin. Median
fold in sulcus, originating simultaneously with sulcus or slightly anterior to
origin of sulcus, widening anteriorly, extending to anterior margin. Lateral
slope gentle, anterior slope steep. Interarea long, narrow, apsacline.
Brachial valve exterior concave, most about middle of valve length. Median
lobe moderately to profoundly exaggerated. In lobe median sulcus present or
absent. Median lobe and median sulcus widening anteriorly, extending to
anterior margin. Lateral and anterior slope gentle. Interarea hypercline.
Pedicle valve interior with moderately long teeth. Median septum short,
extending to one sixth of valve length, posteriorly high, anteriorly lowering and
continuing in low ridge. Adductor scars weakly to moderately pronounced.
Diductors weak, extending to half of the valve length. Inner surface with radial
rows of taleolae, which are larger on lateral and anterior margin of visceral
area. On lateral and anterior margin of valve taleolae are small.
Brachial valve interior with short, wide cardinal process, exteriorly quadrilo-
bed, interiorly bilobed. Alveolus not observed. Median septum long, extending
to three fourth of valve length, wide, high. Prosocket ridges short, narrow,
sockets narrow. Anderidia short, thin. Inner surface taleolate, in anterolateral
region of visceral area short rows of papillae. Inner surface of lateral and
anterior margins with small taleolae.
Comparison: C. karavankensis n. sp. is larger, with more pronounced
ventral median sulcus and more expressed median lobe in it than C. ponteb-
banus n. sp. Exteriorly and interiorly C. karavankensis is similar to C. striatus
(Weiler et McGehee, 1933) from Pennsylvanian of North America, including
shell size and size of capillae; C. karavankensis has slightly more expressed
ventral lateral plications, less hinge spines, which are more perpendicular than
those of C. striatus. C. karavankensis is very similar but slightly less convex
and with slightly less pronounced median lobe than C. heritschi n. sp. It is also
similar externally to Chonetes cf. Ch. latesinuata Schellwien from Sakmarian
of Thailand (Y a n a g i d a , 1967, 86) and to Mesolobus? sp. from Lower
Permian of Svalbard (Gobbett, 1963, 120), but only one specimen from
each of those localities is not enough for efficient comparison. C. karavankensis
is easily distinguished from Paramesolobus sp. 2 with which it is present in
the same beds (pi. 1.24); it is larger and with coarser ribs than Paramesolobus
sp. 2.
Occurrence: Karavanke Mountains, Gzhelian.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 27
Capillomesolobus pontebbanus n. sp.
PI. 2.1—2.25; fig. 9A, B C; fig. 10 A, B
1986 Pecar, p. 137.
Etymology : Pontebbanus — after Pontebba, nearby town.
Types : Holotype GPU 1778 1. Paratypes: GPU 1779 1, KGPL 54031, GPU
1777 1, 1782, 1783, KGPL 5416 1, 5420 1, GPU 1781, 1784, KGPL 5402'2, GPU 1785,
1786, KGPL 5422/1.
Type locality, type horizon and age: To the west of Monte
Corona (locality 6,), Carnian Alps. Brownish shale, Corona Formation. Kasimo-
vian.
Localities and material: Only type locality, 79 specimens (46 pe-
dicle valves, 33 brachial valves).
Diagnosis: Small Capillomesolobus with weak ventral median lobe in
median sulcus.
Description: Small for genus, subrectangular, shell width greater than
length, the widest part at or shortly before hinge margin. Cardinal extremities
from moderately obtuse-angled to acute-angled. Lateral margins oblique, mo-
derately rounded, anterolateral extremities narrowly rounded. Anterior margin
Fig. 10. Capillomesolobus pontebbanus n. gen., n. sp.
A pedicle valve, mould of interior X 9.0, GPU 1779 1, locality 6, Carnian Alps, Corona
Formation, Kasimovian; ß brachial valve, mould of interior X 8.9, GPU 1785, the
same locality as for A
SI. 10. Capillomesolobus pontebbanus n. gen., n. sp.
A pecljeva lupina, odtis notranjosti X 9.0, GPU 1779/1, najdišče 6, Karnijske Alpe,
formacija Corona, kasimovij; B ramenska lupina, odtis notranjosti X 8.9, GPU 1785,
isto najdišče kot pri A
28___ __Janez Pečar
medially indented. Surface capillate, the number of capillae in the middle of
anterior margin of pedicle valve from 55.9 to 79.2 on 10 mm of valve width.
On ventral valve capillae increasing in number mostly with bifurcation, on
dorsal valve mostly with intercalation. Growth lines weak. Hinge spines at least
four on each side, divergent, near right-angled.
Pedicle valve exterior markedly convex in lateral profile, mostly in anterior
half of shell length. In anterior profile narrowly convex, most in the middle
third of valve width. Umbonal slope steep. Wide, shallow median sulcus origi-
nating at about second fifth of shell length, widening anteriorly. In sulcus
median lobe present or absent, widening anteriorly, extending to anterior
margin. Flanks bounding sulcus forming divergent plications. Lateral slope steep,
anterior slope gentle. Interarea long, narrow, anacline to orthocline.
Brachial valve exterior rather concave, mostly on the border between post-
erior and middle third of valve length. More or less expressed median lobe,
widening anteriorly, medially in lobe, sulcus present or absent. Anterior and
lateral slope moderately steep. Interarea long, orthocline.
Pedicle valve interior with little hinge teeths. Median septum short, extend-
ing to one fifth of valve length, narrow, high, continuing anteriorly sometimes
in low ridge, which extends to the middle of the valve length. Adductor scars
weak to marked, posteriorly pointed, anteriorly semicircular, located at median
septum and shortly before it. Diductor scars weak, semicircular, extending to
two thirds of the valve length. Interior surface taleolate, taleolae especially
large in the transition of the lateral slope into ears.
Brachial valve interior with moderatelly large cardinal process, exteriorly
quadrilobed. Alveolus small, conical, shallow. Median septum long, thin, extend-
ing to four fifth of the valve length, posteriorly low, anteriorly higher. Prosocket
ridges short, thin, sockets shallow. Anderidia short, thin. Adductor scars weak.
Interior taleolate, taleolae especially large in the transition of visceral area into
anterior margin.
Compari s ion: C. pontebbanus n. sp. is smaller and with less pronoun-
ced ventral median lobe than C. karavankensis n. sp. or C. heritschi n. sp. Be-
cause of its much smaller size and because of finer ribs C. pontebbanus is easily
distinguished from Paramesolobus sp. 1 which is present in the same beds.
C. pontebbanus has distinct ventral median sulcus, finer ribs and different
interior in comparison with undescribed species of Neochonetes which occurs
in the same beds.
Discussion : Paleobiogeographically it is noteworthy that C. pontebbanus
n. sp. is stratigraphically the oldest known species of genus Capillomesolobus
out of North America (figs. 6, 7 A).
Occurrence: Carnian Alps, Kisimovian.
Capillomesolobus heritschi n. sp.
PI. 3.1—3.9; fig. 9 A, B, C; fig. 11
1900 Chonetes sinuosa Schellwien — S c h e 1 w i e n , p. 38, pl. 9, figs. 17—18.
1938 Chonetes n. sp. — H e r i t s c h , p. 103, pl. 7, figs. 6—7.
1986 New genus A sp. 2 — Pecar, p. 137, 138, pl. 1, figs. 29—32.
Etymology: In honor of Austrian geologist Franz Heritsch who recog-
nized this form as new species.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 29
Types : Holotype: KGPL 5358. Paratypes: KGPL 5423, 5424, 5357, 5425.
Type locality, type horizon and age: Dolžanova soteska (lo-
cality 5), Karavanke Mountains. Secondary occurrence of reddish reef limestone,
Trogkofel Limestone. Sakmarian or Artinskian.
Localities and material: Secondary occurrence (the rubbel of the
abandoned quarry) of reddish reef limestone in type locality, 18 specimens
(5 conjoined valves, 7 pedicle valves, 6 brachial valves). Unit A in locality 5,
1 specimen (pedicle valve). Unit C between two successions of reddish reef
limestone in locality 5, 4 specimens (1 conjoined valve, 3 brachial valves).
Diagnosis : Medium size Capillomesolohus with well developed, posteri-
orly extending umbonal region and with pronounced ventral lateral plications.
Description: Medium size for genus, subtrapezoidal to subrectangular
shell width greater than length, hinge usually the widest part. Cardinal extre-
mities rectangular to slightly acute-angled, ears not marked. Lateral margins
oblique, anterior margin widely indented. Surface capillate, in the middle part
of the ventral anterior margin from 50.7 to 84.7 capillae per 10 mm of shell
width. In posterolateral regions of brachial valve the lateral parts of capillae
turned posteriorly. Hinge spines divergent, their angle to posterior margin is
75" to 85«.
Pedicle valve exterior rather convex in lateral profile, the most about the
middle of the valve length. Umbonal region extending fairly posteriorly to
posterior margin. Median sulcus originating shortly after umbonal region, wi-
dening and deepening anteriorly, extending to anterior margin. Medially in
sulcus median fold widening anteriorly, extending to anterior margin. Lateral
plications bordering median sulcus rather acutely formed. Lateral and anterior
margins steep.
Fig. 11. Capillomesolobus heritschi n. gen, n. sp.
Pedicle valve, mould of interior X 8.7, KGPL 5358, locality 5,
rosered reef limestone, Karavanke Mountains, Trogkofel
Limestone, Sakmarian or Artinskian
SI. П. Capillomesolobus heritschi n. gen., n. sp.
Pecljeva lupina, odtis notranjosti X 8.7, KGPL 5358, najdišče
5, rožnato rdeči grebenski apnenec, Karavanke, trogkofelski
apnenec, sakmarij ali artinskij
30		 		Janez Pečar
Brachial valve exterior concave, most near the middle of valve length. Me-
dian fold ard median sulcus within it are more or less pronounced, widening
anteriorly, extending to anterior margin.
Pedicle valve interior with median septum, which is posteriorly high, ante-
riorly lowering, continuing to median ridge. Muscle scars weak. Visceral area
excavated. Inner surface taleolate, in lateral and anterior parts of visceral area
papillose, on lateral and anterior margin finely taleolate.
Brachial valve interior with short, wide cardinal process, interiorly bilobed.
Median septum high. Alveolus small, shallow, conical. Prosocket ridges short.
Sockets short, wide, shallow. Inner surface taleolate. Papillae in the anterolateral
margin of visceral area, in the middle of each half of visceral area a group
of papillae.
Other exterior and interior structures were not observed.
Comparison: C. heritschi n. sp. is on the average smaller, with more
pronounced ventral lateral plications and with stronger umbonal region than
C. karavankensis n. sp. In the last two mentioned features and more pronounced
median fold it differs from C. pontebbanus n. sp. Stratigraphically interesting
but inefficient is comparison with other Lower Permian forms of the genus
Capillomesolobus which are represented with only one or a few specimens. The
Asselian(?) specimen from Svalbard (Gobbett, 1963, 120) is uncompletely
preserved. C. heritschi has a deeper median sulcus and more acutely formed
lateral plications of the pedicle valve than the specimen from Sakmarian of
Thailand (Y a n a g i d a , 1967, 86). Rather similar to C. heritschi is the form
from Sakamatozawa Series (Sakmarian) of Japan (Nakamura, 1959, 205).
In comparison with Middle and Upper Permian forms of genus Capillomesolo-
bus the species C. permianus (Cooper et Grant, 1975, 1266) is similar to
C. heritschi but the Lower Guadalupian form from California (C o g a n , 1960,
291) and the Punjabian form from Kitakami Mountains of Japan (Hayasaka,
1925, 93) are larger and differently formed.	^
Discussion: Schellwien (1900, 38) which first described this form
from Dolžanova soteska stated that it differs from the Upper Carboniferous
species Chonetes sinuosus Schellwien. But he could not decide for erection of
a new species. Heritsch (1938, 103) thought that this form is a new species
but he did not name it formally.
Occurrence : Karavanke Mountains, Trogkofel Limestone, Sakmarian or
Artinskian.
Subfamily Rugosochonetinae Muir-Wood, 1962
Genus Paramesolobus Afanasjeva, 1975
Type species: Paramesolobus ivanovae Afanasjeva, 1975.
Diagnosis (modified): Medium size, costellate to capillate Rugosocho-
netidae with ventral median sulcus in which median lobe is present or absent.
Comparison : This genus is larger and with coarser ribs in comparison
with Capillomesolobus n. gen., but resembles it because of frequent presence
of the median lobe in the ventral median sulcus. Genus Chonetinella Ramsbot-
tom can be distinguished by the lack of the median lobe in ventral median
sulcus, but the number of ribbs per unit of width can be similar.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 31
Species assigned to genus: Chonetes sinnosus Schellwien, 1898,
Carnian Alps, Spain?, Moscovian-Kasimovian. Chonetes latesinuatus Schellwien,
1892, Carnian Alps, Karavanke Mountains, Hungary, Croatia, China, Kasimo-
vian-Gzhelian, Asselian?. Chonetes latesinuatus miaokouensis Chao, 1928, China,
Spain, Moscovian-Kasimovian, Lower Permian?. Chonetes moelleri carnicus
Gortani, 1905, Carnian Alps, Kasimovian. Paramesolohus sp. 1, Carnian Alps,
Kasimovian (present paper). Paramesolohus sp. 2, Karavanke Mountains, Gzhe-
lian (present paper). Paramesolohus ivanovae Afanasjeva, 1975, Russian Plat-
form, Kasimovian. Paramesolohus luganicus Aisenverg, 1895, Donetz Basin, Ka-
simovian.
Chonetes latesinuatus tsunyiensis Huang, 1932 from the Upper Permian of
China is poorly known and probably does not belong to genus Paramesolohus.
Jing et H u (1978, 126) assigned it to genus Tenuichonetes Jing et Hu. The
stratigraphical distribution of the species assigned to genus Paramesolohus is
shown on fig. 5.
Discussion : The fact that the first three known species of genus Pa-
ramesolohus were described from the Carnian Alps, illustrates the importance
of this area for the study of genus Paramesolohus. These species are: Chonetes
sinuosus (described 1892, the name was corrected 1898), Chonetes latesinuatus
(described 1892) and Chonetes moelleri carnicus (described 1905). During the
work for the present paper the original Schellwien's (1892) specimens
here not available in the museum (Halle, German Democratic Republic) where
S c h e 11 w i e n cited their repository. Also during the present work the type
locality of Chonetes sinuosus in Carnian Alps (bed 6 of the Krone section) was
not found. The type localities of other two species in Carnian Alps were not
examinated during this work. No original or topotype specimen of Chonetes
latesinuatus Schellwien, 1892 is available. This species was described from Spi-
riferenschicht near Monte Carnizza, to the west of Monte Corona. Only two
specimens of Chonetes moelleri carnicus are now available (Gortani's collection
from Monte Pizzul, partially housed in museum in Udine).
It is clear that the revision of species from the Carnian Alps will be signi-
ficant for the study of Paramesolohus. Further field work in this area with
collection of topotype material is needed. The present work reveals that at
least two species of Paramesolohus are present in the Upper Carboniferous of
Carnian Alps and Karavanke Mountains. Earlier mentioned fact preclude the
definitive identification of species of Paramesolohus in the present paper. How-
ever, the need of completeness of presentation of mesolobid chonetacean bra-
chiopods of the area and the need of comparation of Paramesolohus with Capil-
lomesolohus n. gen. make the presentation of data about Paramesolohus neces-
sary. The work on this genus is being continued by the author.
Paramesolohus sp. 1
PI. 3.10—3.21, 4.1—4.12: fig. 12 A, B; fig. 13 A, B, C
1986 Pecar, p. 137.
Locality and material: Locality 6, Corona Formation, Kasimovian,
203 specimens (2 conjoined valves, 124 pedicle valves, 77 brachial valves).
Descript io n : Large for genus, subcircular, shell width greater than
length, hinge widest part. Ears moderately large and nearly right-angled. Sides
32		 		Janez Pečar
Fig. 12. Paramesolobus sp. 1
A pedicle valve, mould of interior X 5.3, GPU 1791 1, locality 6, Carnian
Alps, Corona Formation, Kasimovian; B brachial valve, mould of interior
X 5.0, KGPL 5429 1, the' same locality as for A
SI. 12. Paramesolobus sp. 1
A pecljeva lupina, odtis notranjosti X 5.3, GPU 1791 1, najdišče 6, Kar-
nijske Alpe, formacija Corona, kasimovij; B ramenska lupina, odtis no-
tranjosti X 5.0, KGPL 5429/1, isto najdišče kot pri A
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 33
oblique, concave. Anterior margin slightly medially indented. Surface capillate,
capillae numbering from 29.7 to 45.2 per 10 mm at the middle of the ventral
anterior margin. Number of capillae increasing predominantly with bifurcation
on pedicle valve and predominantly with intercalation on brachial valve.
Growth lines weak. Posterior margin with at least 5 divergent hinge spines on
each side of the beak, they emerge at about 45« to the hinge.
Pedicle valve exterior in lateral profile rather convex, the most convex in
posterior two thirds of the valve length. In anterior profile the valve is the
most convex in the middle third of width. Umbonal region only slightly exten-
ding posteriorly to posterior margin. Median sulcus originating shortly anteri-
orly to umbo, shallow, widening anteriorly. In median sulcus a weak median
fold is present or absent, widening anteriorly. Lateral slope more steep than
anterior. Interarea long, narrow, anacline to apsacline. Pseudodeltidium was
not seen.
Brachial valve exterior moderately convex, in the lateral profile the most
convex on the border between the posterior and middle third of length. Shallow
median fold originating slightly anterior to umbo, widening anteriorly. In
median fold the median sulcus is present or absent, originating simultaneously
as median fold, widening anteriorly. Anterior and lateral slope gentle. Interarea
long, narrow, hypercline. Hilidium was not seen.
Pedicle valve interior with short hinge teeth. Median septum short, thin,
continuing anteriorly in low median ridge, which sometimes becomes again
higher and septum-like anteriorly. Two paramedian ridges can be present in the
middle of the valve. Adductor scars moderately expressed. Diductor muscle
scars tear-like, extending to two fifths of valve length. Visceral area pronoun-
cedly concave. Interior surface taleolate, taleolae larger on anterolateral border
of visceral area and on ears. On lateral and anterior margins taleolae small.
Brachial valve interior with small cardinal process, quadrilobed exteriorly,
bilobed interiorly. Alveolus small, shallow, conical. Median septum long, exten-
ding to three fifth of valve length, thin, posteriorly low, anteriorly becoming
higher and wider, then lowering again. Prosocket ridges moderately long, thin,
sockets shallow. Anderidia thin. Adductor scars weak, semicircular. Inner sur-
face taleolate, especially large taleolae in anterolateral parts of visceral area.
Only scarce taleolae on ears.
Comparison: Paramesolohus sp. 1 is larger than Paramesolohus sp. 2,
has finer ribs, less distinguished ventral median sulcus and less distinguished
ventral median lobe in it. It is likely that Paramesolohus sp. 1 is the same
species which Schellwien (1892, 29) described as Chonetes lohatus and
subsequently (S c h e 11 w i e n , 1898, 360) corrected this name to Chonetes si-
nuosus. The lectotype or neotype for this species are not erected in the present
paper because neither the original Schellvien's material non the topotype ma-
terial are available (locality 6 is probably not the same as Schellwien's bed 6
of Krone section). Comparison with Chonetes moelleri carnicus Gortani, 1905
(two specimens are reposited in museum in Udine, one of them is figured in
Gortani, 1905, pi. 14, fig. 17 a, b) shows that this species might be the same
as Paramesolohus sp 1, but more material from Gortani's locality will be
needed for efficient comparison. Paramesolohus sp. 1 is larger and with finer
ribs than P. ivanovae Afanasjeva, 1975. It is smaller and with finer ribs than
3 - Geologija 28/29
34
	 		Janez Pečar
Fig. 13. Diagrams of genus Paramesolobus
A length against maximal width; B length against number of ribs per 10 mm of width
et ventral anterior margin; C length against total number of ribs at ventral anterior
margin
At A the regression line for (л) is W - 11.97 + 0.80 L, r = + 0.61, P <0.01; for (•)
is W 4.76 + 1.20 L, r - f 0.61, P <0.01. At C the regression line for (•) is Nr =
= 11.61 + 4.53 L, r = + 0.69, P < 0.01
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 35
SI. 13. Diagrami rodu Paramesolobus
A dolžina proti največji širini; ß dolžina proti številu reber na 10 mm širine spred-
njega robu pecljeve lupine; C dolžina proti celotnemu številu reber sprednjega robu
pecljeve lupine
Pri A je regresijska premica za (Л) W = 11.97 + 0.80 L, r = + 0.61, P <0.01; za (•)
je W = 4.76 + 1.20 L, r = + 0.61, P < 0.01. Pri C je regresijska premica za (•) Nr =
= 11.61 + 4.53 L, r = + 0.69, P < 0.01
36		 		Janez Pečar
P. luganicus Aisenverg, 1985. It is similar to the Chinese species Chonetes la-
tesinuatus miaokouensis Chao, 1928. Paramesolobus sp. 1 can be easily distin-
guished from the also mesolobid, but much smaller and more finely ribbed
species Capillomesolobus pontebbanus n. sp. with vvhich it is present in the
same beds.
Paramesolobus sp. 2
PI. 1.24, 5.1—5.23; fig. 13 A, B, C; fig. 14 A, B
1986 Paramesolobus sp. — P e c a r, 137, 138, pi. 1, figs. 14—17.
Localities and material: Locality 1, Gzhelian, 13 specimens (2
conjoined valves, 7 pedicle valves, 4 brachial valves). Locality 2, Gzhelian, 73
specimens (4 conjoined valves, 35 pedicle valves, 34 brachial valves). Locality
4, Gzhelian, 1 specimen (pedicle valve).
Description : Wledium size for genus, subrectangular to subtrapezoidal
in outline, shell width greater than length, widest usually at hinge. Cardinal
extremities moderately acute-angled to obtuse-angled, ears of medium size.
Lateral margins oblique, straight to slightly convex. Anterior margin medially
indented. Surface finely costellate to coarse capillate, in the midle of anterior
margin of adult pedicle valve from 20.5 to 33.4 ribs per 10 mm of shell width.
The number of ribs increasing on predicle valve mostly with bifurcation, on
brachial valve predominantly with intercalation. Some ribs near the posterior
margin less distinct or joined together. Growth lines weak. Posterior margin
with five to seven divergent hinge spines, their angle is from 40" to 80" with
regard to posterior margin.
Pedicle valve exterior in lateral profile moderately convex, the most convex
near middle of the valve length. Umbonal region extending only a little behind
posterior margin. Median sulcus beginning shortly after umbo, widening and
extending to anterior margin, it is deep to shallow. In sulcus median lobe is
present or absent, originating anteriorly to umbo, widening anteriorly and ex-
tending to anterior margin. Flanks bounding sulcus forming two more or less
pronounced plications. Lateral flanks somewhat more steep than anterior flanks.
Interarea long, narrow, anacline to orthocline.
Brachial valve exterior moderately concave, the most near the middle of
the valve length. Low median fold originating after umbonal region, widening
anteriorly and extending to anterior margin. Medially in the lobe shallow sulcus
is present or absent, originating in second third of the valve length. Lateral
flanks gently rising. Interarea long, narrow, hypercline.
Pedicle valve interior with small teeth. Median septum extending to two
fifth of the valve length, thin, lowering anteriorly. Adductor scars inconspicions
to moderately pronounced. Diductor scars semielliptical, extending to the end
of the first third of the valve length. Visceral disc more concave than valve
margins. Interior taleolate, at the anterolateral margins of visceral disc papillose.
Fig. 14. Paramesolobus sp. 2
A pedicle valve, mould of interior X 8.7, TMJ 1227 1, locality 2, Karavanke Moun-
tains, Gzhelian; B brachial valve, mould of interior X 8.0, TMJ 1222, the same
locality as for A
SI. 14. Paramesolobus sp. 2
A pecljeva lupina, odtis notranjosti X 8.7, TMJ 1227'1, najdišče 2, Karavanke, gželij;
B ramenska lupina, odtis notranjosti X 8.0, TMJ 1222, isto najdišče kot pri A
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 37
38		 		Janez Pečar
Brachial valve interior with short, wide cardinal process, exteriorly quadri-
lobed, interiorly bilobed. Alveolus small, shallow, conical. Median septum begins
not before anderidia, it is narrow, low, anteriorly sometimes papillose. Pro-
socket ridges short, fusing with cardinal process. Sockets narrow. Anderidia
short, thin. Anterior adductors weak, posterior adductors inconspicious. Inner
surface taleolate, anterolateral margins of visceral disc papillose.
Comparison: Paramesolohus sp. 2 is smaller, with coarser ribs, more
distinguished ventral median sulcus and more distinguished ventral median
lobe in it than Paramesolohus sp. 1. It is very similar or even identical with
P. ivanovae Afanasjeva, 1975 and with species which Chao (1928, 22) described
from China as Chonetes latesinuatus Schellwien. Paramesolohus sp. 2 is much
smaller from P. luganicus Aisenverg, 1985. It can be easly distinguished from
Capilllomesolohus karavankensis n. sp. which is present in the same beds (pi.
1.24); Paramesolohus sp. 2 is remarkably larger and has much coarser ribs than
C. karavankensis n. sp.
Discussion : This species shows a considerable variability of shell width,
of expression of ventral median sulcus and of median lobe in it. Various transi-
tions between narrow and wide specimens exist, and this variability is thought
to be intraspecific. The statements of Chao (1928, 22) were similar; he united
both two Schellwien's species Chonetes sinuosus (species with pronounced ventral
median lobus) and Chonetes latesinuatus (species with pronounced ventral
median sulcus). Variability of ventral median lobe shows also P. ivanovae
(Afanasjeva, 1975, 105) which was earlier divided in two different species
on the base of presence or absence of ventral median lobe (Ivanov et Iva-
nova, 1936, 20, 22).
Acknowledgements
Prof. Dr. Anton Ramovš, Univerza Edvarda Kardelja, Ljubljana, made
numerous suggestions at work and read the manuscript. I am grateful to Jože
Bedič, Tehniški muzej železarne Jesenice, for field guidance and for loan of
specimens in his care. Thanks to Dr. Giuseppe Muscio, Museo Friulano di Storia
Naturale, Udine, Italy, for use of the museum collections and for loan the
specimens. I extend my thanks to Dr. Corrado Venturini, Istituto di Geologia
e Paleontologia, Bologna, Italy, for field guidance through Upper Paleozoic
stratigraphy in Carnian Alps. Marjan Grm photographed the specimens. The
figures were drafted by Metka Karer except the figures of brachiopods which
were drawn by the author. The English text was corrected by Prof. Dr. Simon
Pire. Renata Pečar typed the manuscript.
Zgornjekarbonski in permijski mezolobidni honetacejski brahiopodi
V Karavankah (Jugoslavija) in Karnijskih Alpah (Italija)
Povzetek
Med zgornjekarbonskimi in permijskimi honetacejskimi (naddružina Chone-
tacea) brahiopodi Karavank (Slovenija) so bile najdene tudi oblike z ventralno
mediano gubo v mediani brazdi. Te oblike pripadajo dvema rodovoma. Oba ro-
dova sta bila ugotovljena tudi v zgornjem karbonu Karnijskih Alp. V priču-
Zgornjekarbonski in permijski mezolobidni honetacejski brahiopodi	39
jočem delu sta morfološka izraza »mezolobidni« in »mezolobidnost« uporabljena
za tiste karbonske in permijske honetacejske brahiopode, pri katerih je na sredi-
ni ventralne mediane brazde prisotna mediana guba.
Prva opisana mezolobidna honetacejska vrsta je bila Chonetes mesolohus
Norwood et Pratten, 1855, iz Pennsylvanije ZDA. Prvi izključno mezolobidni
honetacejski rod je Mesolobus Dunbar et Condra, 1932. Ta rod je bil po pred-
logu H o a r e a (1964, 315) pripoznan kot tipsko gladek. Smotrno je bilo uvr-
stiti rebraste mezolobidne honetaceje v drug rod, kar je napravila Afanas-
jeva (1975) z določitvijo rodu Paramesolobus Afanasjeva. Po mnenju A f a -
n a s j e v e spadajo v ta rod tudi severnoameriške pennsylvanijske mezolobidne
oblike. Nadaljno delitev rebrastih mezolobidnih honetacijskih oblik pa je omo-
gočil material iz gželija Karavank, v katerem sta v istih plasteh ugotovljeni
dve jasno različni mezolobidni rebrasti obliki. Ena od teh je rod Paramesolo-
bus Afanasjeva, 1975. Druga oblika, ki je manjša in z znatno drobnejšimi rebri
od prve, pripada rodu Capillomesolobus n. gen. Ti podatki kažejo na vzporeden
razvoj dveh rebrastih mezolobidnih honetacijskih vej (Pecar, 1986, 137), ki se
med seboj razlikujeta ne le na nivoju rodu, temveč verjetno tudi poddružine.
Potek rodovne opredelitve mezolobidnih honetacijskih brahiopodov je prikazan
na si. 1.
Najdišča 1 do 5 so v Karavankah (Jugoslavija), najdišče 6 je v Karnijskih
Alpah (Italija; si. 2, 3).
Najdišče 1. Spodnja počivala. Je v Javorniškem rovtu pri Jesenicah. Naj-
dišče je okrog 100 m vzdolž cevovoda nad njegovim spodnjim križiščem s cesto.
V temnem skrilavcu je bogata morska nevretenčarska favna. V vmesnih lapor-
nih plasteh najdeni Rugosofusulina alpina antiqua (Schellwien) in Archaeolitho-
phyllum missouriensum Johnson kažeta na gželijsko starost (Ramovš, 1971,
1389; Hahn et. al., 1977, 138).
Najdišče 2. Planina pod Golico. Pri tretjem oporniku žičnice na Spanov vrh.
Najdišče je odkril J. Bedič. Rjavkasti skrilavec je zgornjekarbonske starosti,
verjetno pripada gželiju.
Najdišče 3. Planina pod Golico. Pri petem oporniku žičnice na Spanov vrh.
Starost temno sivega skrilavca je verjetno enaka kot pri najdišču 2.
Najdišče 4. Planina pod Golico. Med šestim in sedmim opornikom žičnice
na Spanov vrh. Sekundarno najdišče enake litologije in verjetno enake starosti
kot najdišče 3. Pripomniti je treba, da podrobna stratigrafija najdišč 1 do 4 ni
znana, ker so plasti slabo razgaljene.
Najdišče 5, Dolžanova soteska nad Tržičem. Klasično najdišče trogkofelskih
plasti v Karavankah, v opuščenem kamnolomu 100 m severno od druge ser-
pentine nad cestnim predorom. Gmota rdečkastega in sivkastega grebenskega
apnenca z bogato morsko nevretenčarsko favno je v tem času preliminarno
imenovana enota B. Nad njo je plastnati apnenec (enota C), nad njim je ver-
jetno še ena enota rdečkastega grebenskega apnenca. Stratigrafija spodnjega
dela trogkofelskih plasti je prikazana na si. 4. Trogkofelske plasti spadajo
v sakmarij (Waterhouse, 1976, 82; Leven, 1980, tab. 3) ali artinskij
(Buser , 1979, 20).
Najdišče 6, Zahodno od Monte Corona (= Krone). Je 200 m zahodno od
spodnjega dela te gore, v nekoliko dvignjenem zahodnem delu gorskega sedla.
Litološko gre za rjavkasti, prepereli skrilavec, ponekod s primesjo apnenca.
40		 		Janez Pečar
Najdišče pripada formaciji Corona, starostno gre za kasimovij (Venturini
et al., 1982, 311).
V diagramih so uporabljene naslednje okrajšave: L = dolžina pecljeve lu-
pine, W = največja širina lupine, Nr 10 = število reber sprednjega robu pecljeve
lupine. V nadaljevanju sledi opis novih nadvrstnih taksonov.
Sistematska paleontologija
Družina Rugosochonetidae Muir-Wood, 1962
Capillomesolobinae n. subfam.
Diagnoza: Male, kapilatne ali gladke Rugosochonetidae z ventralno
mediano brazdo, v kateri je mediana guba prisotna ali odsotna.
Rodovni sestav poddružine: Capillomesolohus n. gen. Mesolohus
Dunbar et Condra, 1932. ?Tenuichonetes Jing et Hu, 1978. Novi, neopisani rod,
tipiziran 2 Neochonetes unhonopUcatus, upodobljen v Barkhatovi, 1964,
iz sakmarijskih plasti Nenets, reka Sula, Severno timansko gorovje (Arch-
bold 1982, 2), morda tudi pripada tej poddružini.
Razpravljanje in filogeneza: Capillomesolobinae n. subfam, se
razlikuje od ostalih poddružin družine Rugosochonetidae po običajni prisotnosti
ventralne mediane gube v ventralni mediani brazdi. Rodovi Capillomesolohus,
Mesolohus in Tenuichonetes so po pričujoči shemi v sorodstvenem odnosu, pri
čemer je izhodiščni rod Capillomesolohus (si. 5). Stratigrafsko porazdelitev vrst
nove poddružine Capillomesolobinae in rodu Paramesolohus je prikazana na
si. 6.
Capillomesolohus n. gen.
1986 New genus A — Pecar, p. 137
Tipska vrsta: Capillomesolohus karavankensis n. sp.
Etimologija: Capillus (lat.) = las, zaradi kapilatne površine; mesolo-
bus, zaradi morfološke podobnosti z rodom Mesolohus.
Diagnoza : Majhne, drobno kapilatne Rugosochonetidae z ventralno me-
diano brazdo, v kateri je mediana guba prisotna ali odsotna.
O p i s : Majhen, subpravokoten do subtrapezoiden, širina večja od dolžine.
Sprednji rob mediano zajeden. Površina drobno kapilatna. Prirastnice šibke.
Sklepne bodice divergentne, blizu pravokotnih do pravokotne. Zunanjost peclje-
ve lupine izbočena. Mediana brazda se navzpred širi. V njej je prisotna ali
odsotna mediana guba. Zunanjost ramenske lupine vbočena, v mediani gubi
prisotna ali odsotna mediana brazda.
Notranjost pecljeve lupine s kratkim septumom. Mišični odtisi šibki. No-
tranja površina taleolatna, taleole so večje na lateralnem in sprednjem robu
visceralnega predela. Na stranskem in sprednjem robu so taleole majhne.
Notranjost ramenske lupine s kratkim, širokim sklepnim izrastkom, navz-
ven štirirežnjatim, navznoter dvorežnjatim. Mediani septum dolg, visok. Ande-
ridija kratka, tanka. Notranja površina taleolanta, v anterolateralnih delih
visceralnega predela papilozna. Notranja površina stranskih robov in sprednje-
ga robu z majhnimi taleolami.
Primerjava: Capillomesolohus se razlikuje od rodu Neochonetes po
globlji ventralni mediani brazdi in po običajni prisotnosti mediane gube v njej.
Upper Carboniferous and Permian mesolobid chonetacean brachiopods__ 41
Po obliki je močno podoben rodu Mesolohus, od njega se razlikuje po rebrasti
površini. Capillomesolobus je manjši in ima drobnejša rebra kot Paramesolobus.
Paleobiogeografija in migracija: Rod Capillomesolobus se
je najverjetneje razvil iz rodu Neochonetes v atokiju (pennsylvanij) Severne
Amerike. Tam je začasno izginil v desmoinesiju. Najstarejše znano izvenameri-
ško nahajališče rodu Capillomesolobus je v Karnijskih Alpah, kjer je v kasimo-
viju ugotovljen C. pontebbanus n. sp. Domnevati je torej možno, da se je rod
razvil v Severni Ameriki, torej na zahodni obali Pangee. Nato je migriral na
vzhodno obalo Pangee, na področje sedanjih Karnijskih Alp in Karavank. Ta
prvi del migracije rodu Capillomesolobus je zaradi biogeografskih barier težko
pojasniti (si. 7 A).
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44		 		Janez Pečar
Plate 1 — Tabla 1
Capillomesolobus karavankensis n. gen. n. sp.
1—11, 20, 21, 25 Locality 1, Gzhelian. 1, 2 pedicle valve, mould of exterior X 1 and
X 2, paratype KGPL 5356; 3—5 pedicle valve, mould of exterior X 1 and X 2,
mould of interior X 2, holotype TMJ 1274; 6—9 pedicle valve, exterior and partial
mould of interior, ventral view X 1 and X 2, anterior X 2, side X 2, paratype
TMJ 1219; 10, 11 pedicle valve, mould of interior, ventral X 2, side X 2, paratype
TMJ 1212; 20 brachial valve, mould of exterior X 2, paratype TMJ 1282; 21 bra-
chial valve, mould of interior X 3,5, paratype KGPL 5411; 25 cardinal process and
brachial valve, mould of exterior X 7, paratype TMJ 1278
12, 13, 18, 19, 22—24 Locality 2, Gzhelian. 12, 13 pedicle valve, mould of exterior X 1
and X 2, KGPL 5412; 18 pedicle valve, mould of interior X 2, KGPL 5414; 19
brachial valve, mould of exterior X 2, KGPL 5415; 22 brachial valve, mould of
interior X 2, KGPL 5413; 23 brachial valve, mould of interior X 2, TMJ 1224 1;
24 at left lower corner brachial valve, mould of exterior of the preceding speci-
men X 2, TMJ 1224/1. At the upper part of 24 is Paramesolobus sp. 2, pedicle valve,
mould of interior X 2, TMJ 1224/2
14—17 Locality 3, Gzhelian. Pedicle valve, exterior, ventral view X 1 and X 2,
posterior X 2, side X 2, TMJ 1265
1—11, 20, 21, 25 Najdišče 1, gželij. 1, 2 pecljeva lupina, odtis notranjosti X 1 in X 2,
paratip KGPL 5356; 3—5 pecljeva lupina, odtis zunanjosti XI in X 2, odtis
notranjosti X 2, holotip TMJ 1274; 6—9 pecljeva lupina, zunanjost in delni odtis
notranjosti, ventralno X 1 in X 2, sprednje X 2, stransko X 2, paratip TMJ 1219;
10, 11 pecljeva lupina, odtis notranjosti, ventralno X 2, stransko X 2, paratip
TMJ 1212; 20 ramenska lupina, odtis zunanjosti X 2, paratip TMJ 1282; 21 ramen-
ska lupina, odtis notranjosti X 3,5, paratip KGPL 5411; 25 sklepni izrastek in
ramenska lupina, odtis zunanjosti X 7, paratip TMJ 1278
12, 13, 18, 19, 22—24 Najdišče 2, gželij. 12, 13 pecljeva lupina, odtis zunanjosti X 1
in X 2, KGPL 5412; 18 pecljeva lupina, odtis notranjosti X 2, KGPL 5414; 19 ra-
menska lupina, odtis zunanjosti X 2, KGPL 5415; 22 ramenska lupina, odtis no-
tranjosti X 2, KGPL 5413; 23 ramenska lupina, odtis notranjosti X 2, TMJ 1224.1;
24 v levem spodnjem kotu ramenska lupina, odtis zunanjosti predhodnega pri-
merka X 2, TMJ 1224 1. V levem zgornjem kotu 24 je Paramesolobus sp. 2, pec-
ljeva lupina, odtis notranjosti X 2, TMJ 1224 2
14—17 Najdišče 3, gželij. Pecljeva lupina, zunanjost, ventralno X 1 in X 2, zadajšnje
X 2, stransko X 2, TMJ 1265
Upper Carboniferous and Permian mesolobid chonetacean brachiopods_ ^
46		 		Janez Pečar
Plate 2 — Tabla 2
Capillomesolobus pontebbanus n. gen., n. sp.
1—25 Locality 6, Corona Formation, Kasimovian. 1—3 pedicle valve, mould of exte-
rior X 1 and X 4, mould of interior X 4, holotype GPU 1778 1; 4, 5 pedicle valve,
mould of exterior X 1 and X 4, paratype KGPL 5403 1; 6—10 pedicle valve, mould
of exterior X 1 and X 4, mould of interior, ventral, anterior and side X 4, para-
type GPU 1779/1; 11 brachial valve, mould of exterior X 4, paratype GPU 1781;
12, 13 pedicle valve, mould of exterior X 1 and X 4, paratype GPU 1777 1; 14, 15
pedicle valve, mould of interior, ventral and side X 4, paratype GPU 1782; 16—18
pedicle valve, mould of interior, ventral, anterior and side X 4, paratype GPU
1783; 19 pedicle valve, mould of interior X 4, paratype KGPL 5416 1; 20 pedicle
valve, mould of interior X 4, paratype KGPL 5420/^1 ; 21 brachial valve, mould
of interior X 4, paratype GPU 1784; 22 brachial valve, mould of interior X 4,
paratype KGPL 5402 2; 23 brachial valve, mould of interior X 4, paratype GPU
1785; 24 brachial valve, mould of interior X 4, paratype GPU 1786; 25 brachial
valve, mould of interior X 4, KGPL 5422/Í1
1—25 Najdišče 6, formacija Corona, kasimovij. 1—3 pecljeva lupina, odtis zunanjosti
X 1 in X 4, odtis notranjosti X 4, holotip GPU 1778 1; 4, 5 pecljeva lupina, odtis
zunanjosti X 1 in X 2, paratip KGPL 5403 1; 6—10 pecljeva lupina, odtis zunanjo-
sti X 1 in X 4, odtis notranjosti, ventralno, sprednje in stransko X 4, para-
tip GPU 1779 1; 11 ramenska lupina, odtis zunanjosti X 4, paratip GPU 1781; 12,
13 pecljeva lupina, odtis zunanjosti XI in X 4, paratip GPU 1777 1; 14, 15 pec-
ljeva lupina, odtis notranjosti, ventralno in stransko X 4, paratip GPU 1782; 16—18
pecljeva lupina, odtis notranjosti, ventralno, sprednje in stransko X 4, paratip
GPU 1783; 19 pecljeva lupina, odtis notranjosti X 4, paratip KGPL 5416 1; 20
pecljeva lupina, odtis notranjosti X 4, paratip KGPL 5420 1; 21 ramenska lupina,
odtis notranjosti X 4, paratip GPU 1784; 22 ramenska lupina, odtis notranjosti X 4,
paratip KGPL 5402 2; 23 ramenska lupina, odtis notranjosti X 4, paratip GPU
1785; 24 ramenska lupina, odtis notranjosti X 4, GPU 1786; 25 ramenska lupina,
odtis notranjosti X 4, KGPL 5422 1
Upper Carboniferous and Permian mesolobid chonetacean brachiopods
 47
48		 		Janez Pečar
Plate 3 — Tabla 3
Capillomesolobus heritschi n. gen., n. sp.
1—9 Locality 5, Trogkofel Limestone, Sakmarian or Artinskian. 1, 2 pedicle valve,
mould of interior X 1 and X 2, paratype KGPL 5423; 3—6 pedicle valve, partial
exterior and interior, ventral X 1 and X 2, posterior and side X 2, holotype KGPL
5358; 7 brachial valve, mould of exterior X 2, paratype KGPL 5424; 8 brachial
valve, mould of exterior X 2, paratype KGPL 5357; 9 brachial valve, mould of
exterior X 2, paratype KGPL 5425
Paramesolohus sp. 1
10—21 Locality 6, Corona Formation, Kasimovian. 10, 11 pedicle valve, mould of ex-
terior X 1 and X 2, GPU 1787 1; 12 brachial valve, mould of exterior X 2, GPU
1788 1; 13, 14 pedicle valve, mould of exterior X 1 and X 2, GPU 1789/1; 15 bra-
chial valve, mould of exterior X 2, KGPL 5426 1; 16, 17 pedicle valve, exterior
X 1 and X 2, KGPL 5400; 18, 19 pedicle valve, mould of exterior X 1 and X 2,
GPU 1775 1; 20, 21 pedicle valve, mould of interior, ventral and anterior X 2,
KGPL 5427 1
Capillomesolobus heritschi n. gen., n. sp.
1—9 Najdišče 5, trogkofelski apnenec, sakmarij ali artinskij. 1, 2 pecljeva lupina,
odtis notranjosti XI in X2, paratip KGPL 5423; 3—6 pecljeva lupina, delna zu-
nanjost in notranjost, ventralno XI in X 2, posteriorno in stransko X 2, holotip
KGPL 5358; 7 ramenska lupina, odtis zunanjosti X 2, paratip KGPL 5424; 8 ra-
menska lupina, odtis zunanjosti X 2, paratip KGPL 5357; 9 ramenska lupina, odtis
zunanjosti X 2, paratip KGPL 5425
Paramesolohus sp. 1
10—21 Najdišče 6, formacija Corona, kasimovij. 10, 11 pecljeva lupina, odtis zunanjo-
sti X 1 in X 2, GPU 1787 1; 12 ramenska lupina, odtis zunanjosti X 2, GPU 1788.1;
13, 14 pecljeva lupina, odtis zunanjosti XI in X 2, GPU 1789 1; 15 ramenska lu-
pina, odtis zunanjosti X 2, KGPL 5426 1; 16, 17 pecljeva lupina, zunanjost X 1 in
X 2, KGPL 5400; 18, 19 pecljeva lupina, odtis zunanjosti X 1 in X 2, GPU 1775 1;
20, 21 pecljeva lupina, odtis notranjosti, ventralno in sprednje X 2, KGPL 5427 1
Upper Carboniferous and Permian mesolobid chonetacean brachiopods
 49
4 - Geologija 28 '29
50		 		Janez Pečar
Plate 4 — Tabla 4
Paramesolobus sp. 1
1—12 Locality 6, Corona Formation, Kasimovian. 1 pedicle valve, mould of interior
X 2, GPU 1790 1; 2, 3 pedicle valve, mould of interior, ventral and posterior X 2,
GPU 1791 1; 4—6 pedicle valve, mould of interior, ventral, anterior and side X 2,
GPU 1792; 7 pedicle valve, mould of interior X 2, KGPL 5428 1; 8 brachial valve,
mould of interior X 2, GPU 1776 2; 9 brachial valve, mould of interior X 2, GPU
1793 1; 10 brachial valve, mould of interior X 2, GPU 1794; 11 brachial valve,
mould of interior X 2, KGPL 5429 1; 12 cardinal process and umbo of brachial
valve, mould of exterior X 8, GPU 1795
1—12 Najdišče 6, formacija Corona, kasimovij. 1 pecljeva lupina, odtis notranjosti
X 2, GPU 1790 1; 2, 3 pecljeva lupina, odtis notranjosti, ventralno in zadajšnje
X 2, GPU 17911; 4—6 pecljeva lupina, odtis notranjosti, ventralno, sprednje in
stransko X 2, GPU 1792; 7 pecljeva lupina, odtis notranjosti X 2, KGPL 5428 1;
8 ramenska lupina, odtis notranjosti X 2, GPU 1776 2; 9 ramenska lupina, odtis
notranjosti X 2, GPU 1793 1; 10 ramenska lupina, odtis notranjosti X 2, GPU
1794; 11 ramenska lupina, odtis notranjosti, KGPL 5429 1; 12 sklepni izrastek in
umbo ramenske lupine, odtis zunanjosti X 8, GPU 1795
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 51
52		 		Janez Pečar
Plate 5 — Tabla 5
Paramesolobus sp. 2
1—4, 9, 15—23 Locality 2, Gzhelian. 1—3 pedicle valve, mould of exterior X 1 and
X 2, mould of interior X 2, TMJ 1227 1; 4 brachial valve, mould of exterior X 2,
TMJ 1248; 9 brachial valve, mould of exterior X 2, TMJ 1245; 15 juvenile pedicle
valve, mould of interior X 2, KGPL 5430; 16 juvenile pedicle valve, mould of in-
terior X 2, KGPL 5431; 17 pedicle valve, mould of interior X 2, TMJ 1242; 18
pedicle valve, mould of interior X 2, TMJ 1223; 19 pedicle valve, mould of interior
X 2, TMJ 1252; 20 brachial valve, mould of interior X 2, TMJ 1229; 21, 22 bra-
chial valve, mould of interior X 2, cardinal area X 8, TMJ 1222; 23 cardinal
process, mould of exterior X 8, TMJ 1245
5—8 Locality 3, Gzhelian. 5—8 pedicle valve, exterior, ventral X 1 and X 2, anterior
and side X 2, KGPL 5432
10—14 Locality 1, Gzhelian. 10 brachial valve, mould of exterior X 2, KGPL 5433;
11—14 pedicle valve, exterior, ventral X 1 and X 2, anterior and side X 2, TMJ
1214
1—4, 9, 15—23 Najdišče 2, gželij. 1—3 pecljeva lupina, odtis zunanjosti XI in X 2,
odtis notranjosti X 2, TMJ 1227 1; 4 ramenska lupina, odtis zunanjosti X 2, TMJ
1248; 9 ramenska lupina, odtis zi^nanjosti X 2, TMJ 1245; 15 mladostna pecljeva
lupina, odtis notranjosti X 2, KGPL 5430; 16 mladostna pecljeva lupina, odtis no-
tranjosti X 2, KGPL 5431; 17 pecljeva lupina, odtis notranjosti X 2, TMJ 1242;
18 pecljeva lupina, odtis notranjosti X 2, TMJ 1223; 19 pecljeva lupina, odtis no-
tranjosti X 2, TMJ 1252; 20 ramenska lupina, odtis notranjosti X 2, TMJ 1229;
21, 22 ramenska lupina, odtis notranjosti X 2, kardinalni predel X 8, TMJ 1222;
23 sklepni izrastek, odtis zunanjosti X 8, TMJ 1245
5—8 Najdišče 3, gželij. 5—8 pecljeva lupina, zunanjost, ventralno X 1 in X 2, sprednje
in stransko X 2, KGPL 5432
10—14 Najdišče 1, gželij. 10 ramenska lupina, odtis zunanjosti X 2, KGPL 5433;
11—14 pecljeva lupina, zunanjost, ventralno X 1 in X 2, sprednje in stransko X 2,
TMJ 1214
Upper Carboniferous and Permian mesolobid chonetacean brachiopods	 53