Type species

Strophodonta variabilis Calvin, Reference Calvin1878; Independence Shale, Middle Amana Beds, Frasnian; Iowa, USA.

? Douvillinaria sp. indet.

Figure 4.6–4.11

Figure 4.

(1–17) Strophomenide, productide, protorthide, orthide, and pentameride brachiopods from the Laskowa Góra Beds of Górno, Holy Cross Mountains: (1–5) Skenidioides cf. S. polonicus (Gürich, Reference Gürich1896), juvenile shell, GIUS 4-292/5, in dorsal, ventral, lateral, posterior, and anterior views, respectively; (6–11) ?Douvillinaria sp. indet.: (6–10) juvenile specimen, GIUS 4-292/1, in dorsal, ventral, lateral, posterior, and anterior views, respectively; (11) interior of incomplete dorsal valve, GIUS 4-292/2’ (12–16) ?Spinulicosta sp. indet., slightly damaged shell, GIUS 4-292/3, in dorsal, ventral, lateral, posterior, and anterior views, respectively; (17) Devonoproductus sericeus (von Buch, Reference Buch1837), exterior of incomplete dorsal valve, GIUS 4-292/4. (18–20) Schizophoria schnuri prohibita Halamski, Reference Halamski2012, articulated shell, ZPAL Bp 73/1/9a/1, in dorsal, ventral, and posterior views, respectively. (21, 22) Physemella sp. indet., ventral valve, GIUS 4-292/9, interior and exterior views, respectively. Scale bars = 1 mm (1–11, 17); 10 mm (12–16, 18–22).

Figure 4. Long description

The photo plate contains 22 numbered panels arranged in four rows.

Row 1 (Panels 1 to 5): Skenidioides cf. S. polonicus. Small juvenile shells showing dorsal, ventral, lateral, posterior, and anterior views. The shells are semi-circular with fine radial ribbing. Scale bar equals 1 mm.

Row 2 (Panels 6 to 11): ?Douvillinaria sp. indet. Panels 6 to 10 show a juvenile specimen in dorsal, ventral, lateral, posterior, and anterior views, featuring a fan-like shape with prominent radial costae. Panel 11 shows the interior of an incomplete dorsal valve with complex internal structures. Scale bar equals 1 mm.

Row 3 (Panels 12 to 17): ?Spinulicosta sp. indet. and Devonoproductus sericeus. Panels 12 to 16 show a damaged shell in five views, characterized by a bulbous ventral valve with small spine bases. Panel 17 shows the exterior of an incomplete dorsal valve with concentric growth lines. Scale bars for 12 to 16 equal 10 mm; scale bar for 17 equals 1 mm.

Row 4 (Panels 18 to 22): Schizophoria schnuri prohibita and Physemella sp. indet. Panels 18 to 20 show a large articulated shell in dorsal, ventral, and posterior views, exhibiting a smooth, rounded profile with very fine striations. Panels 21 and 22 show the interior and exterior of a ventral valve of Physemella, which is triangular and robust. Scale bars equal 10 mm.

v Reference Baliński, Racki and Halamski2016 ?Douvillinaria sp.; Baliński et al., p. 136, pl. 5, figs. 2–12.

Materials

A subcomplete juvenile articulated shell and a fragmentary dorsal valve, GIUS 4-292/1,2.

Remarks

Brachiopods described and figured by Baliński et al. (Reference Baliński, Racki and Halamski2016) came mostly from the Laskowa Góra Beds of Józefka. A juvenile specimen and a dorsal interior, both not studied in the previous paper, are figured here.

Order Productida Waagen, Reference Waagen1883

Suborder Productidina Waagen, Reference Waagen1883

Superfamily Productoidea Gray, Reference Gray1840

Family Productellidae Schuchert, Reference Schuchert, Schuchert and LeVene1929

Genus Spinulicosta Nalivkin, Reference Nalivkin1937

Type species

Productus spinulicosta Hall, Reference Hall1857; Hamilton Group, Givetian; shores of Cayuga Lake, New York, USA (for justification for this spelling of the type species, see Baliński and Halamski Reference Halamski2013, p. 15).

? Spinulicosta sp. indet.

Figure 4.12–4.16

Materials

A single incomplete articulated shell, GIUS 4-292/3.

Description

The studied specimen is an articulated shell, mostly lacking marginal regions, except small remnants of anterior margin preserved in a single place. Shell probably rounded rectangular in outline, transverse, moderately convex. Preserved length 23.0 mm (estimated length ~24 mm), preserved width 27.1 mm (estimated width ~30 mm), thickness 10.3 mm. Umbo strongly incurved; beak fine.

Ornament of irregularly disposed spines, ~0.8–1.1 mm thick in marginal region, sometimes as if arising from small radial ridges, denser in umbonal region, more widely spaced near the margins; spines along hinge margin present; ventral interarea sublinear. Low undulating rugae, distinct and spaced < 1 mm in umbonal region, evanescent and spaced ~2–2.5 mm near the margins.

The concave dorsal valve has a dent ~11 mm long and 14 mm wide in the median region; it might correspond to the visceral cavity.

Remarks

This single specimen is tentatively identified as a representative of the genus Spinulicosta on account of irregularly disposed spines with radial ridges, although the latter are very weakly developed. Spinulicosta cf. S. spinulicosta was reported from the lower Givetian of Miłoszów (Baliński and Halamski, Reference Baliński and Halamski2023): it is similar in outline, but smaller in size; scarcity of material prevents a more detailed comparison.

Superfamily Linoproductoidea Stehli, Reference Stehli1954

Family Monticuliferidae Muir-Wood and Cooper, Reference Muir-Wood and Cooper1960

Subfamily Devonoproductidae Muir-Wood and Cooper, Reference Muir-Wood and Cooper1960

Genus Devonoproductus Stainbrook, Reference Stainbrook1943

Type species

Productella walcotti Fenton and Fenton, Reference Fenton and Fenton1924; Frasnian; Hackberry, Rockford, Iowa, USA (see details by Muir-Wood and Cooper, Reference Muir-Wood and Cooper1960, p. 177–179).

Devonoproductus sericeus (von Buch, Reference Buch1837)

Figure 4.17

* Reference Buch1837 Orthis sericea von Buch, p. 68.

v Reference Baliński, Racki and Halamski2016 Devonoproductus sericeus; Baliński et al., p. 137, pl. 5, figs. 13–17.

Type specimens

Specimens from von Buch’s collection (syntypes; not illustrated in the original publication), present repository unknown; ‘Kohlenkalkstein’ [Frasnian in reality], Oberkunzendorf [now Mokrzeszów Górny, Silesia, Poland], for detailed stratigraphy, see Halamski (Reference Halamski2013, p. 302–304) and Baliński et al. (Reference Baliński, Racki and Halamski2016, p. 137).

Materials

A single incomplete dorsal valve, GIUS 4-292/4.

Remarks

Devonoproductus sericeus was described by Baliński et al. (Reference Baliński, Racki and Halamski2016) on the basis of a collection coming mostly from the Frasnian. A single, poorly-preserved specimen from the Givetian, not studied in the previous paper, is figured here. Following Baliński et al. (Reference Baliński, Racki and Halamski2016), this species is defined broadly.

Class Rhynchonellata Williams et al., Reference Williams, Carlson, Brunton, Holmer and Popov1996

Order Protorthida Schuchert and Cooper, Reference Schuchert and Cooper1931

Superfamily Skenidioidea Kozłowski, Reference Kozłowski1929

Family Skenidiidae Kozłowski, Reference Kozłowski1929

Genus Skenidioides Schuchert and Cooper, Reference Schuchert and Cooper1931

Type species

Skenidioides billingsi Schuchert and Cooper, Reference Schuchert and Cooper1931; Black River Formation, Sandbian, Upper Ordovician; near the Ottawa River, Québec, Canada.

Skenidioides cf. S. polonicus (Gürich, Reference Gürich1896)

Figure 4.1–4.5

cf. Reference Gürich1896 Skenidium polonicum Gürich, p. 237, pl. 10, figs. 3, 14.

cf. Reference Baliński and Halamski2023 Skenidioides polonicus (Gürich, Reference Gürich1896); Baliński and Halamski, p. 17, figs. 7A–J, Z–BB, 8.

Materials

A single incompletely preserved juvenile specimen, GIUS 4-292/5.

Remarks

The available specimen is a juvenile shell 1.9 mm long and 3.0 mm wide. It can be tentatively referred to Skenidioides polonicus on account of the presence of a dorsal midrib pair (see Baliński and Halamski, Reference Baliński and Halamski2023, p. 17), but this identification is not considered secure due to the young age of the individual (see Baliński et al., Reference Baliński, Racki and Halamski2016, text-fig. 6) and imperfect preservation of the ornamentation, so open nomenclature is used.

Order Orthida Schuchert and Cooper, Reference Schuchert and Cooper1931

Family Schizophoriidae Schuchert and LeVene, Reference Schuchert, LeVene and Pompeckj1929

Genus Schizophoria King, Reference King1850

Type species

Conchyliolithus (Anomites) resupinatus Martin, Reference Martin1809; lower Carboniferous; England (the neotype selected for this species lacks precise locality data; see Bond, Reference Bond1941, p. 290).

Schizophoria schnuri prohibita Halamski, Reference Halamski2012

Figure 4.18–4.20

v* Reference Halamski2012 Schizophoria ( Schizophoria ) schnuri prohibita Halamski, p. 356, figs. 4A–C, E, M, Q, V–Y, BB–EE, 7A–II, 8C, E.

v. Reference Baliński, Racki and Halamski2016 Schizophoria schnuri prohibita ; Baliński et al., p. 145, pl. 10, figs. 21, 22.

Holotype

ZPAL Bp 64/1/1/1 (see Halamski, Reference Halamski2012, for type locality and stratum, and paratypes).

Materials

A single subcomplete arcticulated shell and eight fragmentary specimens, GIUS 4-208/1, 4-292/5–8, 4-294/1-3, and ZPAL Bp 73/1/9a/1, Bp 81/12a.

Remarks

This subspecies was described from the Laskowa Góra Beds of the Józefka quarry (Halamski, Reference Halamski2012). An early Frasnian representative of this genus from Kadzielnia in Kielce was identified as Schizophoria cf. S. prohibita, although it might also represent a new species (Racki, Reference Racki2024).

Order Pentamerida Schuchert and Cooper, Reference Schuchert and Cooper1931

Suborder Pentameridina Schuchert and Cooper, Reference Schuchert and Cooper1931

Superfamily Gypiduloidea Schuchert and LeVene, Reference Schuchert, LeVene and Pompeckj1929

Family Gypidulidae Schuchert and LeVene, Reference Schuchert, LeVene and Pompeckj1929

Subfamily Gypidulinae Schuchert and LeVene, Reference Schuchert, LeVene and Pompeckj1929

Genus Physemella Godefroid, Reference Godefroid1974

Type species

Physemella maillieuxi Godefroid, Reference Godefroid1974; middle Frasnian; Frasnes-lez-Couvin, Belgium.

Physemella sp. indet.

Figure 4.21, 4.22

v. Reference Baliński, Racki and Halamski2016 Physemella sp.; Baliński et al., p. 145, pl. 10, figs. 23–25.

Materials

Four incomplete ventral valves, some embedded in rock, GIUS 4-292/9–12.

Remarks

Poorly preserved specimens tentatively assigned to this species occur in the Laskowa Góra Beds. It has been described in detail on the basis of a better collection from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description.

Order Rhynchonellida Kuhn, Reference Kuhn1949

Superfamily Uncinuloidea Rzhonsnitskaya, Reference Rzhonsnitskaya and Thalmann1956

Family Hypothyridinidae Rzhonsnitskaya, Reference Rzhonsnitskaya and Thalmann1956

Remarks

At present, the taxonomy of the family Hypothyridinidae remains unclear. This is due mainly to historical reasons, as discussed at length by Sartenaer (Reference Sartenaer2003). In summary, Atrypa cuboides Sowerby, Reference Sowerby, Sedgwick and Murchison1840, the type species of the genus Hypothyridina Buckman, Reference Buckman1906, which in turn is the type genus of the family, was described on the basis of a single specimen coming from Devonshire, a region characterized by complex geological structure. The internal structures of the type species and exact age of the type stratum were therefore unknown. As a consequence, on the one hand, Atrypa cuboides came to be interpreted too widely, whereas on the another hand, new genera within the family were proposed without adequate comparison with the genuine Hypothyridina. A systematic revision of the Hypothyridinidae, including Hypothyridina cuboides, was performed by Baxter (Reference Baxter1999), but her results remain unpublished and thus unknown to the most of scientific community; moreover, her nomenclatural proposals cannot be regarded as published for the purposes of zoological nomenclature (ICZN, 1999, Art. 9). In light of the analysis of the material of Hypothyridina cuboides from the type region (Baxter, Reference Baxter1999), the interpretation of Hypothyridina in both editions of the Treatise of Invertebrate Paleontology (Schmidt, Reference Schmidt1965; Savage, Reference Savage and Kaesler2002) must be considered misguided.

The interpretation of Hypothyridina by Baxter (Reference Baxter1999), followed by the present authors, is based on the premise that the type locality of Hypothyridina cuboides, described simply as ‘Plymouth’ in the original publication (Sowerby, Reference Sowerby, Sedgwick and Murchison1840) can be restricted to the Mount Wise Shell Bed (Ussher, Reference Ussher1907; Orchard, Reference Orchard1977), the age of which corresponds to a later part of the Givetian stage. Baxter (Reference Baxter1999) serially sectioned a specimen from a coeval bed in a nearby locality (Lummaton Shell Bed, middle to upper part of the Givetian). This specimen (BM 5839) served as a de facto epitype allowing greater clarity in the diagnosis of the species. This course of action was quite similar to that followed by Halamski and Baliński (Reference Halamski and Baliński2019) in their revision of Spirifera simplex Phillips, Reference Phillips1841, another commonly misunderstood species described initially from Devonshire.

According to Baxter (Reference Baxter1999), the family Hypothyridinidae is characterized by strong variability of external characters, but all its representatives lack the dorsal septum and the septalium. Out of a total of six genera (or eight, including uncertain ones) belonging to the family Hypothyridinidae, three are relatively similar externally, namely Hypothyridina, Glosshypothyridina Rzhonsnitskaya, Reference Rzhonsnitskaya1978, and Tullypothyridina Sartenaer, Reference Sartenaer2003 (= Hemihypothyridina Baxter, Reference Baxter1999, nomen nudum). Even if the proposed distinctions are not entirely clear, because Hypothyridina is said to differ from Glosshypothyridina in “the development of definite dental plates,” but Glosshypothyridina is described as possessing “well expressed dental plates” (Baxter, Reference Baxter1999, p. 381, 399), a workable comparison of the three genera can be devised (see Fig. 5, Table 1).

Figure 5.

Synoptic comparison of internal features of Glosshypothyridina theophili n. sp. with those of the type species of the three genera discussed in the text. (1) Hypothyridina cuboides (Sowerby, Reference Sowerby, Sedgwick and Murchison1840), modified after Baxter (Reference Baxter1999, fig. 96). (2) Tullypothyridina venustula (Hall, Reference Hall1867), after Sartenaer (Reference Sartenaer2003, fig. 1). (3) Glosshypothyridina procuboides (Kayser, Reference Kayser1871), modified after Schmidt (Reference Schmidt1941a, pl. 7, fig. 19; distances between sections not provided in the cited publication). (4) Glosshypothyridina theophili n. sp. (see also Fig. 8). Scale bars = 5 mm.

Figure 5. Long description

The diagram consists of four vertical columns, each labeled at the top with a species name and a panel number. Horizontal scale bars representing 5 mm are placed near the top of each column.

* Column 1: Hypothyridina cuboides. Contains four sections. The first section at 0.4 shows a small dorsal hinge structure. The second at 0.75 shows the development of the shell wall with dashed lines indicating incomplete preservation. The third at 1.35 and fourth at 1.85 show the widening of the internal cavity and small central dental plates.

* Column 2: Tullypothyridina venustula. Contains four sections. The first at 1.35 features a curved arrow pointing to a dark, serrated hinge plate. The second at 1.95 shows a large, circular shell outline with thick lateral walls. The third and fourth sections at the bottom are smaller fragments showing the ventral valve floor.

* Column 3: Glosshypothyridina procuboides. Contains four sections. The first is a small, circular hinge section. The second shows a triangular internal chamber. The third shows a wide, flat-bottomed shell with a central serrated process. The fourth shows the distal expansion of the shell walls.

* Column 4: Glosshypothyridina theophili n. sp. Contains four sections. The first at 1.0 shows the initial hinge. The second at 1.8 includes a rectangular inset box showing a magnified view of a serrated median septum, indicated by a curved arrow. The third at 2.65 and fourth at 3.0 show the broad, winged shape of the mature shell with distinct lateral cavities.

Table 1.

Comparison of three Middle to Upper Devonian genera of hypothyridinid brachiopods: Hypothyridina, Tullypothyridina, and Glosshypothyridina. HCM, Holy Cross Mountains

Table 1. Long description

The table consists of five columns. The first column lists the morphological or taxonomic category, while the subsequent columns provide data for the genera Hypothyridina, Tullypothyridina, and Glosshypothyridina (the latter split into two columns for different species).

* Species: Hypothyridina type species is H. cuboides (Givetian). Tullypothyridina type species is T. venustula (Givetian). Glosshypothyridina includes type species G. procuboides (Eifelian) and G. theophili n. sp. (late Givetian).

* General form of the shell: Hypothyridina is spherical; Tullypothyridina is ellipsoid; Glosshypothyridina is transverse.

* Highest point of the dorsal valve: All three genera have the highest point at or near the anterior commissure.

* Costae form: Hypothyridina costae are not flattened and have no median groove. Tullypothyridina and Glosshypothyridina have flattened costae with a median groove.

* Costae beginning: Hypothyridina starts in the posterior half; Tullypothyridina near the umbo; Glosshypothyridina in the anterior half.

* Dental plates: Hypothyridina and Tullypothyridina have fine, ventrally convergent plates. Glosshypothyridina has strong plates that are either variable in direction or ventrally divergent.

* Cardinal process: Absent in Hypothyridina; wide with many lamellae in Tullypothyridina and Glosshypothyridina.

* Source of morphological data: Baxter 1999 for Hypothyridina; Sartenaer 2003 for Tullypothyridina; Sartenaer 2003, Schmidt 1941b, and the current work for Glosshypothyridina.

* Other species: Hypothyridina includes Frasnian and Givetian species; Tullypothyridina includes Givetian to Frasnian species; Glosshypothyridina includes Eifelian species from Russia and China.

It follows thus that the interpretation of Hypothyridina based on serial sections of a specimen identified as ‘Hypothyridina sp. cf. H. impleta’ (Sowerby, Reference Sowerby, Sedgwick and Murchison1840) by Schmidt (Reference Schmidt1965) and Savage (Reference Savage and Kaesler2002) is erroneous, insofar as the sectioned brachiopod possesses a cardinal process and thus does not belong to the genus Hypothyridina. For the varying interpretations of Glosshypothyridina see below, under that genus.

Genus Glosshypothyridina Rzhonsnitskaya, Reference Rzhonsnitskaya1978

Type species

Rhynchonella procuboides Kayser, Reference Kayser1871; Eifelian (most probably late Eifelian, according to Sartenaer, Reference Sartenaer2003); Eifel, Germany.

Remarks

As observed by Sartenaer (Reference Sartenaer2003, p. 35), serial sections of Glosshypothyridina procuboides, the type species of the genus, figured by Schmidt (Reference Schmidt1941a, pl. 7, fig. 19; refigured by Savage, Reference Savage and Kaesler2002, fig. 766.1c–f) and Schmidt and McLaren (Reference Schmidt, McLaren and Moore1965, fig. 442.2; refigured by Savage, Reference Savage and Kaesler2002, fig. 766.1g–j) clearly do not belong to the same species, insofar as the former sectioned brachiopod lacks a dorsal septum and a septalium that the latter does possess. The interpretation followed here is based on the specimen sectioned in 1941; the original figure has better quality than that republished in the new Treatise and allows discerning a multilamellate cardinal process (see Fig. 5C).

Glosshypothyridina theophili new species

Figures 5.4, 6–8

Figure 6.

Glosshypothyridina theophili n. sp. from the Laskowa Góra Beds (1–25) and Szydłówek Beds (26–35) of Górno, Holy Cross Mountains. Seven shells, in dorsal, ventral, lateral, posterior, and anterior views, respectively: (1–5) GIUS 4-292/17; (6–10) GIUS 4-292/13; (11–15) GIUS 4-292/14; (16–20) holotype ZPAL Bp 73/1/13/1; (21–25) ZPAL Bp 73/1/13/2; (26–30) MWGUW ZI/124/1; (31–35) MWGUW ZI/124/2. Scale bars = 10 mm.

Figure 6. Long description

The photograph consists of 35 individual panels organized into seven rows, labeled 1 through 35. Each row represents one specimen.

* Row 1 (1–5): Specimen G I U S 4-292/17.

* Row 2 (6–10): Specimen G I U S 4-292/13.

* Row 3 (11–15): Specimen G I U S 4-292/14.

* Row 4 (16–20): Holotype Z P A L Bp 73/1/13/1.

* Row 5 (21–25): Specimen Z P A L Bp 73/1/13/2.

* Row 6 (26–30): Specimen M W G U W Z I/124/1.

* Row 7 (31–35): Specimen M W G U W Z I/124/2.

Within each row, the five panels show the shell from different perspectives:

1. The first column shows the dorsal view, revealing a triangular to sub-pentagonal outline with fine radial ribs near the anterior margin.

2. The second column shows the ventral view, highlighting the pedicle valve.

3. The third column shows the lateral view, illustrating the biconvexity and profile of the valves.

4. The fourth column shows the posterior view, focusing on the hinge line and umbo.

5. The fifth column shows the anterior view, emphasizing the uniplicate frontal commissure and the depth of the ribs.

White scale bars representing 10 mm are located at the bottom left of each row. The specimens vary in preservation, with some showing clear ribbing and others appearing more weathered or partially crushed.

Figure 7.

Glosshypothyridina theophili n. sp. from the Laskowa Góra Beds of Górno, Holy Cross Mountains. (1–3) Grooved costae: (1, 3) ZPAL 73/1/13/4: (1) on ventral tongue; (3) on dorsal flank of the shell; (2) ZPAL Bp 73/1/13/5, on dorsal flank of the shell. (4) Anterior view of the partial internal shell mold (shell, ZPAL Bp 73/1/13/2), showing the empty spaces left by the dissolution of the dorsal marginal spines at the anterior commissure. (5, 6) GIUS 4-292/14, showing partially preserved, long ventral marginal spines (arrows) running along dorsal flank costae: (5) lateral view of the partial internal shell mold; (6) detail. Scale bars = 5 mm (1–5); 1 mm (6).

Figure 7. Long description

The figure consists of six panels numbered 1 through 6.

* Panel 1 at the top left shows a close-up of grooved costae on a ventral tongue. A white scale bar representing 5 mm is at the top right.

* Panel 2 at the top right displays grooved costae on the dorsal flank of the shell. A 5 mm scale bar is at the bottom left.

* Panel 3 in the middle left shows grooved costae on the dorsal flank of specimen Z P A L B p 73/1/13/4. A 5 mm scale bar is at the bottom left.

* Panel 4 in the middle left, below panel 3, provides an anterior view of a partial internal shell mold. It highlights a row of vertical, elongated empty spaces at the anterior commissure where dorsal marginal spines have dissolved. A 5 mm scale bar is at the top right.

* Panel 5 on the middle right is a lateral view of a partial internal shell mold. A white rectangle outlines an area of interest, and a 5 mm scale bar is at the bottom left.

* Panel 6 at the bottom is a high-magnification detail of the area boxed in panel 5. Six small white arrows point to thin, long ventral marginal spines that run along the dorsal flank costae. A 1 mm scale bar is at the bottom left.

Figure 8.

Transverse serial sections of Glosshypothyridina theophili n. sp. from the Laskowa Góra Beds of Górno, Holy Cross Mountains, through shells GIUS 4-292/15 (1) and GIUS 4-294/4 (2). Distances in mm from the tip of the ventral umbo. Scale bars = 5 mm, unless otherwise labeled.

Figure 8. Long description

The diagram is divided into two horizontal panels.

Panel 1 contains eight primary cross-sections of shell G I U S 4-292/15. The sections are arranged in two rows. The top row shows sections at 0.55, 0.85, 1.5, 1.75, and 1.9 millimeters. The bottom row shows sections at 2.2, 2.75, and 3.6 millimeters. Two inset boxes provide 1 millimeter scale magnified views of the hinge and dental structures for the 1.75 and 1.9 millimeter sections. As the distance increases, the shell outline expands from a small circular shape to a wide, flattened biconvex form with complex internal crural plates.

Panel 2 contains eleven primary cross-sections of shell G I U S 4-294/4 arranged in four rows.

* Row one includes distances 0.3, 0.5, 1.0, 1.45, 1.6, and 1.8 millimeters.

* Row two includes distances 2.0, 2.3, and 2.65 millimeters.

* Row three and four show fragmented sections at 3.0 and 3.2 millimeters.

Two inset boxes with 1 millimeter scale bars show detailed views of the cardinal process and hinge plates for the 1.8 and 2.0 millimeter sections. The internal structures show a progression from simple apical plates to elaborate, curved brachial supports.

Scale bars representing 5 millimeters are located at the top right of panel 1 and bottom right of panel 2.

v. Reference Skompski, Baliński, Szulczewski and Zawadzka2018 Hypothyridina sp.; Skompski et al., p. 625, fig. 9M, N.

Type specimens

Holotype, articulated shell, ZPAL 73/1/13/1 (Fig. 6.16–6.20). Paratypes: ~20 articulated shells from the Laskowa Góra Beds, mostly somewhat deformed (crushed), collection numbers ZPAL Bp 73/1/13 and GIUS 4-292/13–22, 4-294/4; five complete and five fragmentary shells from the Szydłówek Beds, ZPAL 81/13a, MWGUW ZI/124. Type locality: Eastern wall of the Józefka quarry (former Mt. Józefka) near Górno, Holy Cross Mountains, Poland. Type stratum: Laskowa Góra Beds; upper part of the Givetian.

Diagnosis

Glosshypothyridina with relatively large shell, a nearly straight cardinal margin, and strong dental plates.

Occurrence

Józefka, quarry and roadcut (Laskowa Góra Beds and Szydłówek Beds).

Description

Shell transversely subhexagonal in outline, typically ~30–34 mm wide (maximum recorded width 38.7 mm), wider than long (width-to-length ratio 1.27–1.53, most often ~1.3), approximately as high as long, strongly dorsibiconvex; maximal width at approximate midlength; cardinal margin nearly straight, lateral margins rounded, anterior margin truncated, anterior commissure uniplicate. Ventral valve convex in umbonal region, flattened on flanks, with a low, flat-bottomed sulcus in the anterior half. Tongue high, broad, occupying 0.52–0.77 of the valve width, forming an angle of ~70^o with the commissural plane, bordered by subvertical deflexions of the commissure, distally recurved posteriorly. Dorsal valve strongly convex, maximal height at anterior commissure, with a rounded fold appearing at approximate one-third of valve length; dorsal umbo exceeding the ventral one in larger specimens.

Ornamentation of low, thick, rounded costae developed some distance from smooth umbonal regions, separated by sublinear furrows; 8–15 on the tongue and 13–18 per lateral flank, with 0 or 1 parietal costae. In well-preserved shells, each ventral median and dorsal lateral costa is subdivided longitudinally by a median linear groove in proximity of the commissure (Fig. 7.1–7.3). Dorsal median and ventral flank costae become flattened near the commissure.

Etymology

In honor of Teofil Żebrawski (5 April 1800–5 February 1887), Polish geodesist, biologist, archeologist, mathematician, cartographer, architect, and bibliographer.

Zoobank lsid

urn:lsid:zoobank.org:act:7F5AD518-6C09-4F0B-82EE-C1207D6AD96A

Measurements

Mean values and ranges are given in the description. For the complete set of raw data, means, standard deviations, and variation coefficients, see Appendix.

Remarks

The studied brachiopods are included in the superfamily Uncinuloidea and the family Hypothyridinidae on account of their cuboid shape, numerous anteriorly flattened costae, the presence of a wide cardinal process, and lack of a dorsal septum and septalium (Savage, Reference Savage and Kaesler2002, p. 1092, 1127). The detailed comparison of the new species with the type species of Hypothyridina, Glosshypothyridina, and Tullypothyridina is given in the Table 1; it can be said in summary that the presence of a cardinal process precludes the assignment of the discussed brachiopod to Hypothyridina, whereas the strong and ventrally divergent dental plate separate it from both Hypothyridina and Tullypothyridina. Conversely, no major (genus-level) difference of our brachiopod with Glosshypothyridina can be found.

In comparison with Glosshypothyridina procuboides, the new species is larger (30–35 mm in width compared to values for G. procuboides reported as 25–30 mm in the Anti-Atlas by Halamski and Baliński, Reference Halamski and Baliński2013, p. 267; Halamski et al., Reference Halamski, Baliński and Koppka2022a, fig. 16; or even only up to 20 mm in the Eifel according to Höflinger and Jung, Reference Höflinger and Jung2020, p. 107), has a much less angular, nearly straight cardinal margin, and its dental plates are stronger.

The material at hand, sparse and often deformed, is not sufficient to allow a quantitative assessment of the variability. Nonetheless, there are minor discernible differences between the samples from the Laskowa Góra Beds (type stratum; N = 13; Fig. 6.1–6.25) and the overlying Szydłówek Beds (N = 6; Fig. 6.26–6.35). The shells from the latter are larger (width 32.0–38.7 mm, compared to 21.2–36.1 mm in the sample from the Laskowa Góra Beds), slightly less high (thickness-to-length ratio 0.90–1.01 compared to 0.94–1.17 in the type stratum), have a slightly wider tongue (width of the tongue-to-width ratio 0.56–0.77 compared to 0.52–0.69) and more costae in the tongue (13–15 compared to 9–11). These phenotypic differences could correspond either to biological evolution over a period of the order of a million years, or else to ecological differentiation between habitats characterized by dissimilar life conditions.

Superfamily Camarotoechioidea Schuchert, Reference Schuchert, Schuchert and LeVene1929

Family Septalariidae Havliček, Reference Havlíček1960

Genus Phlogoiderhynchus Sartenaer, Reference Sartenaer1970

Type species

Uncinulus arefactus Veevers, Reference Veevers1959; lower Frasnian; Fitzroy Basin, Western Australia.

Phlogoiderhynchus polonicus (Roemer, Reference Roemer1866)

Figure 9.2

Figure 9.

Rhynchonellide brachiopods from the Laskowa Góra Beds of Górno, Holy Cross Mountains. (1, 3–6) Coeloterorhynchus dillanus (Schmidt, Reference Schmidt1941b), articulated shell, GIUS 4-292/32, in lateral, dorsal, ventral, anterior, and posterior views, respectively. Note probably syn vivo auloporid incrustation. (2) Phlogoiderhynchus polonicus (Roemer, Reference Roemer1866), articulated shell, ZPAL 73/1/11a/1, in dorsal view. Scale bars = 10 mm.

Figure 9. Long description

The photo consists of six panels labeled 1 through 6 against a black background.

* Panel 1 (top left): A lateral view of a Coeloterorhynchus dillanus shell, showing a triangular profile with small, tube-like auloporid incrustations attached to the surface.

* Panel 2 (top right): A dorsal view of a Phlogoiderhynchus polonicus shell, featuring a wide, fan-like shape with distinct radial ribbing. A white scale bar representing 10 mm is positioned below it.

* Panel 3 (middle left): A dorsal view of the Coeloterorhynchus dillanus shell, showing a prominent central fold and more tube-like encrustations. A white scale bar representing 10 mm is located above this panel.

* Panel 4 (middle right): A ventral view of the same shell, showing a smoother surface with a deep sulcus and fewer encrustations.

* Panel 5 (bottom left): An anterior view of the shell, highlighting the zig-zag commissure where the two valves meet, heavily encrusted with branching auloporid tubes.

* Panel 6 (bottom right): A posterior view of the shell, showing the hinge area and the rounded umbo with a slightly textured surface.

* Reference Roemer1866 Camarotoechia? polonica Roemer, p. 676, pl. 13, figs. 9, 10.

v. Reference Baliński, Racki and Halamski2016 Phlogoiderhynchus polonicus; Baliński et al., p. 147, pl. 11, figs. 1–5.

v. Reference Skompski, Baliński, Szulczewski and Zawadzka2018 Phlogoiderhynchus polonicus; Skompski et al., p. 625, text-fig. 9O.

Type specimens

Roemer’s original type material is lost, so the specimen MGUWr s.n., figured by Gürich (Reference Gürich1896, pl. 7, fig. 6) and refigured by Biernat and Szulczewski (Reference Biernat and Szulczewski1975, pl. 24, fig. 2) has been selected as the neotype (Biernat and Szulczewski, Reference Biernat and Szulczewski1975, p. 204).

Materials

Twelve variously preserved shells, GIUS 4-208/2, 4-292/23–31, and ZPAL Bp 73/1/11a/1, Bp 81/14.

Remarks

Phlogoiderhynchus polonicus, considered as a regional index species for the lower Frasnian, is present in the Szydłówek beds (Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018) and the Wietrznia beds (Baliński et al., Reference Baliński, Racki and Halamski2016). The reader is referred to the two already published descriptions.

Superfamily Pugnacoidea Rzhonsnitskaya, Reference Rzhonsnitskaya and Thalmann1956

Family Pugnacidae Rzhonsnitskaya, Reference Rzhonsnitskaya and Thalmann1956

Genus Coeloterorhynchus Sartenaer, Reference Sartenaer1966b

Type species

Coeloterorhynchus tabasensis Sartenaer, Reference Sartenaer1966b; middle and upper Frasnian, Upper Devonian; Iran.

Coeloteorhynchus dillanus (Schmidt, Reference Schmidt1941b)

Figure 9.1, 9.3–9.6

* Reference Schmidt1941b Pugnax acuminatus dillanus Schmidt, p. 284, pl. 1, figs. 2–4.

v. Reference Baliński, Racki and Halamski2016 Coeloteorhynchus dillanus; Baliński et al., p. 147, text-figs. 9, 10, pl. 11, figs. 6–30, 32.

v. Reference Skompski, Baliński, Szulczewski and Zawadzka2018 Coeloteorhynchus dillanus; Skompski et al., p. 625, text-figs. 9A–J, P, 13.

Type specimens

Holotype, SMF XVII 962n, from Frasnian limestone contained in Famennian breccia, Langenaubach, Lahn-Dill Syncline, Germany (Schmidt, Reference Schmidt1941b, fig. 4). For paratypes, see Schmidt (Reference Schmidt1941b, p. 284, figs. 2, 3).

Materials

More than 20 articulated shells, GIUS 4-292/32, 4-294/5-14, and ZPAL Bp 73/3/12/1, Bp 81/9,10,15,16.

Remarks

This species is present in the Laskowa Góra, Szydłówek, and Wietrznia beds of Józefka. The reader is referred to detailed descriptions published in the previous papers dealing with the Józefka section (Baliński et al., Reference Baliński, Racki and Halamski2016; Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018).

Order Atrypida Rzhonsnitskaya, Reference Rzhonsnitskaya1960

Suborder Atrypidina Moore, Reference Moore, Moore, Lalicker and Fischer1952

Family Atrypidae Gill, Reference Gill1871

Subfamily Atrypinae Gill, Reference Gill1871

Genus Kyrtatrypa Struve, Reference Struve1966

Type species

Atrypa (Kyrtatrypa) culminigera Struve, Reference Struve1966; Junkerberg Formation, middle Eifelian; Hillesheim, Eifel, Germany.

Remarks

The distinction between Atrypa Dalman, Reference Dalman1828 and Kyrtatrypa was discussed at length by Halamski et al. (Reference Halamski, Baliński and Kondas2024, p. 233–235), because the discovery of Kyrtatrypa pauli Halamski and Baliński in Halamski et al., Reference Halamski, Baliński and Kondas2024 showed that differences between the two genera were less clear-cut than when only Kyrtatrypa culminigera, the type species of the latter genus, had been considered. The discovery of Kyrtatrypa stanislai new species (below) blurs the discussed distinction even more, insofar as the new species possesses minute dental nuclei. Dental nuclei were heretofore considered diagnostic for Atrypa (Copper, Reference Copper and Kaesler2002, p. 1396; Halamski et al., Reference Halamski, Baliński and Kondas2024, p. 235). However, careful examination of the serial sections of Kyrtatrypa culminigera (see Copper, Reference Copper and Kaesler2002, fig. 947e) reveals the presence of similar minuscule dental nuclei comparable to those in the species described below. As a consequence, the diagnosis of the genus Kyrtatrypa must be emended as follows: prominent, large, solid teeth, but minute dental nuclei may sometimes be present (otherwise as of Copper, Reference Copper and Kaesler2002). Conversely, only medium-sized to large dental nuclei should be considered diagnostic for Atrypa.

Kyrtatrypa stanislai new species

Figures 10.6–10.30, 11, 12

Figure 10.

Atrypide brachiopods from the Laskowa Góra and Szydłówek beds of Górno, Holy Cross Mountains. (1–3) Desquamatia globosa jozefkae Baliński in Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018, articulated shell, ZPAL Bp 81/23, from the Szydłówek Beds, in dorsal, ventral, and posterior views, respectively. (4) Davidsonia cf. D. enmerkaris Halamski in Baliński et al., Reference Baliński, Racki and Halamski2016, fragmentary ventral valve, GIUS 4-292/52, from the Laskowa Góra beds, in dorsal view. (5) Spinatrypina (Exatrypa) explanata (von Schlotheim, Reference Schlotheim1820), articulated shell, GIUS 4-208/3, from the Laskowa Góra beds, in ventral view. (6–30) Kyrtatrypa stanislai n. sp. from the Szydłówek Beds of Górno, Holy Cross Mountains, five shells, in dorsal, ventral, lateral, posterior, and anterior views, respectively: (6–10) ZPAL Bp 73/3/1/2; (11–15) holotype ZPAL Bp 73/3/1/1; (16–20) MWGUW ZI/124/3; (21–25) MWGUW ZI/124/4; (26–30) MWGUW ZI/124/5. Scale bars = 2 mm (4); 10 mm (1–3, 5–30).

Figure 10. Long description

The photo contains 30 numbered views of fossilized brachiopod shells arranged in rows against a black background.

* Row 1: Specimens 1 through 5. Views 1 to 3 show a single shell of Desquamatia globosa jozefkae in dorsal, ventral, and posterior views. View 4 shows a fragmentary ventral valve of Davidsonia cf. D. enmerkaris. View 5 shows a ventral view of Spinatrypina explanata.

* Rows 2 through 7: Specimens 6 through 30. These rows display five different shells of Kyrtatrypa stanislai n. sp., with each shell shown in a standard five-view sequence: dorsal, ventral, lateral, posterior, and anterior.

* Row 2 and 3 (top right): Views 6 to 10 represent specimen Z P A L Bp 73/3/1/2.

* Row 3 (bottom left) and 4 (top right): Views 11 to 15 represent the holotype Z P A L Bp 73/3/1/1.

* Row 4 (bottom left) and 5 (top right): Views 16 to 20 represent specimen M W G U W Z I / 124 / 3.

* Row 5 (bottom left) and 6 (top right): Views 21 to 25 represent specimen M W G U W Z I / 124 / 4.

* Row 7: Views 26 to 30 represent specimen M W G U W Z I / 124 / 5.

The shells are generally rounded or sub-triangular with distinct radial ribbing patterns. Scale bars are provided on the left side of rows 1, 2, 3, 4, 5, and 7, and below specimen 4 and 5, indicating 2 mm for specimen 4 and 10 mm for all others.

Figure 11.

Transverse serial sections of Kyrtatrypa stanislai n. sp. from the Szydłówek Beds of Górno, Holy Cross Mountains, through shells ZPAL Bp 73/3/1/12 (1) and ZPAL Bp 73/3/1/13 (2). Distances in mm from the tip of the ventral umbo. Scale bars = 5 mm.

Figure 11. Long description

Panel 1 contains thirteen sections of shell Z P A L B p 73/3/1/12. The sequence begins at the top left with section 1.1, showing a small circular profile. As the distance increases through 1.6, 2.3, 2.9, and 3.6, the shell expands into a wide, winged shape with complex internal structures including a central cavity and dental plates. A dotted horizontal line separates the top row from the middle. Sections 4.2, 4.8, 5.2, and 6.6 show the gradual thinning of internal supports. The bottom two rows, sections 7.9, 8.6, 9.3, 10.3, and 16.6, show the shell becoming a broad, shallow V-shape with small internal punctae or ridges appearing in the final section.

Panel 2, located at the bottom, contains eight sections of shell Z P A L B p 73/3/1/13. The top row shows sections 0.65, 0.9, 1.3, 1.7, 2.05, and 2.4. These progress from a small solid circle to an arched shape with developing internal hinge structures. The bottom row contains section 3.2 on the right and 4.0 on the left, showing the fully expanded, wide-winged transverse profile of the brachiopod shell. Scale bars representing 5 mm are present in both panels.

Figure 12.

Biometric comparison of the lower Frasnian Kyrtatrypa stanislai n. sp. from Górno with the closest species, the middle Givetian K. pauli Halamski and Baliński in Halamski et al., Reference Halamski, Baliński and Kondas2024, from Laskowa and Błonia Sierżawskie, Holy Cross Mountains: (1) thickness as a function of length; (2) tongue width as a function of shell width; (3) width as a function of length.

Figure 12. Long description

Three scatter plots arranged vertically, labeled 1, 2, and 3. A legend at the bottom identifies three data groups: black circles for K yrtatrypa stanislai from Józefka, dark gray triangles for K. pauli from Laskowa, and white triangles for K. pauli from Błonia Sierżawskie.

* Panel 1: Thickness in millimeters on the y-axis (0 to 35) versus Length in millimeters on the x-axis (0 to 50). Data shows a positive linear correlation. White triangles are clustered at the lower end (length 12 to 30, thickness 5 to 20). Dark gray triangles overlap and extend further (length 20 to 45, thickness 15 to 27). Black circles are concentrated at the upper end (length 30 to 43, thickness 21 to 30).

* Panel 2: Tongue width in millimeters on the y-axis (0 to 30) versus Width in millimeters on the x-axis (0 to 60). Dark gray triangles are scattered widely between width 22 and 52. Black circles form a tighter cluster between width 32 and 42, with tongue widths ranging from 15 to 27 millimeters.

* Panel 3: Width in millimeters on the y-axis (0 to 60) versus Length in millimeters on the x-axis (0 to 50). This plot shows a strong linear relationship for all groups. White triangles occupy the lower-left (length 12 to 28, width 12 to 28). Dark gray triangles span the middle to upper range (length 20 to 46, width 22 to 52). Black circles are tightly grouped in the upper-middle section (length 31 to 43, width 31 to 42).

Type specimens

Holotype, ZPAL Bp 73/3/1/1 (Fig. 9.16–9.20); paratypes, ZPAL Bp 73/3/1 and MWGUW ZI/124; mostly relatively well-preserved articulated shells. Type locality: Józefka quarry near Górno, Holy Cross Mountains, Poland. Type stratum: brachiopod-rich lens within the Szydłówek Beds, lowermost Frasnian.

Diagnosis

Large atrypine, possessing a distinctly convex ventral valve, teeth with minute dental nuclei, uniplicate anterior commissure with rather broad and often relatively high-rounded tongue, and rather coarse radial ornamentation.

Description

Shell rounded to shield-shaped in outline, dorsibiconvex, typically ~33–37 mm long, 34–39 mm wide, and 23–27 mm thick; approximately as long as wide: width-to-length ratio (0.91–)1.01–1.05(–1.31), mean 1.04, N = 30, usually markedly convex: thickness-to-length ratio (0.58–)0.68–0.72(–1.01), mean 0.71; N = 29. Maximum width at approximate posterior one-third of the length. Anterior commissure uniplicate; tongue, moderately wide, making (0.38)–0.53–0.59(–0.66), mean 0.55, N = 28, of the width of the shell, often high, rounded (parabolic-shape). Ventral valve distinctly convex in umbonal region, sometimes flattening on flanks, but only rarely being completely flat near the lateral margins; frills not observed; beak small, appressed to the dorsal umbo. Dorsal valve arched in anterior view, mostly high; beak hidden by the ventral valve.

Ornament of radial ribs, mostly 5 or 6 per 5 mm at anterior margin, 9 or 10 per 5 mm at 5 mm from the umbo.

Interior of both valves with thickened umbonal region. Ventral valve with thick pedicle callist; deltidial plates absent; teeth thick, solid, but with minute dental nuclei, barely visible in some transverse sections; median muscle scars narrowly incised in umbonal and posterior region, widening anteriorly.

Dorsal interior with deep sockets and distinct accessory middle socket ridge; socket plates thick, separated by deep cardinal pit; cardinal process short, lining socket plates and cardinal pit posteriorly; median myophragm thick and distinct; crura fibrous, laterally directed, spiralia with nine whorls in one of the sectioned specimens (Fig. 10.1).

Etymology

In honor of Stanisław Skompski (b. 1952; Latin form of the forename: Stanislaus; genitive: Stanislai), in recognition of his contributions to the geology of the Holy Cross Mountains; in particular, he collected a part of the described material of K. stanislai n. sp., including the holotype.

Zoobank lsid

urn:lsid:zoobank.org:act:87D32925-9B56-404E-96DD-DD5BBF6B5DC7

Measurements

Mean values and ranges are given in the description and the remarks. For the complete set of raw data, means, standard deviations, and variation coefficients, see Appendix.

Remarks

The studied brachiopods are included within Kyrtatrypa on the basis of teeth with minute dental nuclei (see Remarks under the genus), the uniplicate anterior commissure, and the distinctly (even if not strongly) convex ventral valve. Frills were not observed, which is likely due to the specimens having been extracted from hard limestone.

Kyrtatrypa stanislai n. sp. is close to older (late middle Givetian) K. pauli in shape and ornamentation. It differs therefrom in being higher (W/T mean 0.71, maximum 1.01; in K. pauli 0.60 and 0.79, respectively), having a less straight hinge line, and possessing rudimentary dental nuclei (fully absent in K. pauli, see discussion above). Moreover, the ventral valve of K. stanislai n. sp. is higher than that of K. pauli, which can often be nearly flat (Halamski et al., Reference Halamski, Baliński and Kondas2024, fig. 4M, R). The tongue of K. stanislai n. sp. is mostly rounded, whereas in K. pauli it is often subtrapezoidal (flattened in its top part). These differences, in addition to the stratigraphic separation of the two samples, are judged sufficient for warranting a species-level distinction, although it should be stressed that they can be appreciated if a sufficiently large sample is analyzed; single shells can sometimes be difficult to identify.

The studied species is known from the type locality and stratum only. Three other Frasnian species of Kyrtatrypa have been reported in the literature; all three are relatively easily distinguished from K. stanislai n. sp. Kyrtatrypa brandonensis (Stainbrook, Reference Stainbrook1938) from Iowa is distinctly wider than long, has a lower shell, and in particular, the ventral valve is lower; the ornament is somewhat denser. Kyrtatrypa teicherti (Coleman, Reference Coleman1951) from the Frasnian of Western Australia is similar in shape, but much smaller. The late Frasnian K. barnimi Halamski, Reference Halamski2013 from the Sudetes is globose.

Subfamily Spinatrypinae Copper, Reference Copper1978

Genus Spinatrypina Rzhonsnitskaya, Reference Rzhonsnitskaya1964

Type species

Spinatrypina margaritoides Rzhonsnitskaya, Reference Rzhonsnitskaya1964; Pragian; Siberia, Russia.

Subgenus Spinatrypina ( Exatrypa ) Copper, Reference Copper1967

Type species

Terebratulites explanatus von Schlotheim, Reference Schlotheim1820; Refrath beds, Frasnian; Refrath, Paffrath Syncline, Bergisches Land, Germany.

Spinatrypina ( Exatrypa ) explanata (von Schlotheim, Reference Schlotheim1820)

Figure 10.5

* Reference Schlotheim1820 Terebratulites explanatus von Schlotheim, p. 263.

. Reference Racki1993a Spinatrypina robusta Copper, Reference Copper1967; Racki, p. 312.

v. Reference Baliński, Racki and Halamski2016 Spinatrypina (Exatrypa) explanata ; Baliński et al., p. 151, text-fig. 12, pl. 15, figs. 1–33 [ubi syn.].

Type specimens

Lectotype MB.B s.n., selected by W. Struve, published and figured by Copper (Reference Copper1967, pl. 20, fig. 2), from Refrath beds (Frasnian), Paffrath Syncline, Bergisches Land, Germany.

Materials

More than 60 mostly imperfectly preserved articulated shells, GIUS 4-208/3–23, 4-292/32–51, 4-294/15–36.

Remarks

This species, common in the Laskowa Góra Beds, but present also in the Szydłówek Beds and the Wietrznia Beds, was described on the basis of > 80 articulated shells (all the three stratigraphic levels combined) by Baliński et al. (Reference Baliński, Racki and Halamski2016) as Spinatrypina (Exatrypa) explanata. The reader is referred thereto for description, biometric data, and illustrations. However, in an earlier contribution brachiopods from this level (Kielce region localities, see below) were identified as Spinatrypina (Exatrypa) robusta Copper, Reference Copper1967 (Racki, Reference Racki1993a). At present, no straightforward solution of this dilemma can be proposed: brachiopods from Górno agree nearly perfectly to Spinatrypina explanata in terms of planar shape, but are thicker (Baliński et al. Reference Baliński, Racki and Halamski2016, p. 151); on the other hand, Spinatrypina robusta is of similar thickness as brachiopods from Górno, but less wide (width-to-length ratio ~1.1). According to Copper (Reference Copper1967, p. 136), both species have restricted and disjunct stratigraphic ranges in the Refrath beds (topmost lower to lower part of middle Frasnian; Ma et al., Reference Ma, Ebbighausen and Becker2015): Spinatrypina (Exatrypa) explanata is present in their lower part (Steinbreche horizon, F2a) and Spinatrypina (Exatrypa) robusta in their upper part (Saalermühl horizon, F2b). The representatives of Spinatrypina (Exatrypa) from Górno are apparently morphologically rather uniform from the upper part of the Givetian to the lower Frasnian.

Provisionally, Spinatrypina (Exatrypa) robusta, a less wide and more biconvex variant, is included in the intraspecies variability of Spinatrypina (Exatrypa) explanata. In view of an apparently relatively long stratigraphic range of the brachiopod sample from Górno, such a solution, despite its imperfection, seems more satisfactory than describing a new species.

Subfamily Variatrypinae Copper, Reference Copper1978

Genus Desquamatia Alekseeva, Reference Alekseeva1960

Type species

Atrypa (Desquamatia) khavae Alekseeva, Reference Alekseeva1960; lower Eifelian; Urals, Russia.

Desquamatia globosa jozefkae Baliński in Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018

Figure 10.1–10.3

v* Reference Skompski, Baliński, Szulczewski and Zawadzka2018 Desquamatia globosa jozefkae Baliński in Skompski et al., p. 626, text-figs. 7, 8, 14, 15.

Type specimens

Holotype, ZPAL Bp 81/1 (Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018, fig. 7Z–DD) from the Szydłówek Beds of Józefka (Givetian-Frasnian boundary beds). For paratypes, see Skompski et al. (Reference Skompski, Baliński, Szulczewski and Zawadzka2018, p. 626).

Materials

More than 150 articulated shells, ZPAL Bp 81/1/2–8, 17–19, 23–25 and GIUS 4-208/24–27.

Remarks

This subspecies was described on the basis of > 150 articulated shells present in a fossiliferous lens in the Szydłówek Beds of the Józefka quarry (Skompski et al., Reference Skompski, Baliński, Szulczewski and Zawadzka2018).

Suborder Davidsoniidina Copper, Reference Copper1996

Superfamily Davidsonioidea King, Reference King1850

Family Davidsoniidae King, Reference King1850

Genus Davidsonia Bouchard-Chantereaux, Reference Bouchard-Chantereaux1849

Type species

Davidsonia verneuillii Bouchard-Chantereaux, Reference Bouchard-Chantereaux1849; upper Eifelian; Germany.

Davidsonia cf. D. enmerkaris Halamski in Baliński et al., Reference Baliński, Racki and Halamski2016

Figure 10.4

cf. Reference Baliński, Racki and Halamski2016 Davidsonia enmerkaris Halamski in Baliński et al., Reference Baliński, Racki and Halamski2016, p. 153, pl. 12, figs. 16–26.

Type specimens

Holotype, ZPAL Bp 73/1/19/1, from the lower Frasnian Wietrznia Beds (sets 21 and 22) of the Józefka roadcut. For paratypes, see Baliński et al. (Reference Baliński, Racki and Halamski2016, p. 153).

Materials

A fragmentary ventral valve, GIUS 4-292/52.

Remarks

Davidsonia enmerkaris has been described in detail on the basis of a rich collection from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description. The only specimen from the Laskowa Góra Beds is a small fragment of ventral valve with characteristic spiral grooves; it is tentatively referred to the discussed species, because the dorsal valve is not known. It can be noted Davidsonia enmerkaris is the youngest known representative of the suborder Davidsoniidina.

Order Athyridida Boucot, Johnson, and Staton, Reference Boucot, Johnson and Staton1964

Suborder Athyrididina Boucot, Johnson, and Staton, Reference Boucot, Johnson and Staton1964

Superfamily Athyridoidea Davidson, Reference Davidson1881

Family Athyrididae Davidson, Reference Davidson1881

Subfamily Helenathyridinae Dagys, Reference Dagys1974

Genus Biernatella Baliński, Reference Baliński1977

Type species

Biernatella polonica Baliński, Reference Baliński1977; Palmatolepis gigas Biozone, Frasnian; trench Z-4 south of Dębnik near Cracow, southern Poland.

Biernatella ovalis Baliński, Reference Baliński1995

Figure 13.19–13.23

Figure 13.

Athyridide, spiriferide, and spiriferinide brachiopods from the Laskowa Góra Beds of Górno, Holy Cross Mountains. (1–4) Cyrtina cf. C. arkonensis Ehlers and Wright, Reference Ehlers and Wright1975, incomplete shell, GIUS 4-292/76: (2–4) dorsal, lateral, and posterior views, respectively; (1) detail of the delthyrium. (5–7) Echinocoelia parva Baliński in Baliński et al., Reference Baliński, Racki and Halamski2016, articulated shell, GIUS 4-292/62, in dorsal, lateral, and posterior views, respectively. (8–12) Echinocoelia sp. indet., complete shell, GIUS 4-292/67, in dorsal, ventral, lateral, posterior, and anterior views, respectively. (13–17) Ambothyris sp. indet., complete shell, GIUS 4-292/54, in dorsal, ventral, lateral, posterior, and anterior views, respectively. (19–23) Biernatella ovalis Baliński, Reference Baliński1995, slightly damaged shell, GIUS 4-292/53, in dorsal, ventral, lateral, posterior, and anterior views, respectively. (24–28) Emanuella aff. E. takwanensis (Kayser, Reference Kayser and von Richthofen1883), articulated shell, GIUS 4-292/72, in dorsal, ventral, lateral, posterior, and anterior views, respectively. (18, 29–33) Indospirifer sp. indet.; slightly damaged and partially exfoliated shell, GIUS 4-294/37: (29–33) dorsal, ventral, lateral, posterior, and anterior views, respectively; (18) detail showing micro-ornamentation preserved on a patch of the primary shell layer. (34) ?Reticulariopsis sp. indet., fragmentary ventral valve, GIUS 4-292/75, in exterior view. Scale bars = 1 mm (1–22), 2 mm (23–28), 5 mm (29–34).

Figure 13. Long description

The plate contains 34 numbered black and white photographs of fossilized brachiopods.

- Top row (1 to 7): Cyrtina cf. C. arkonensis (1 to 4) showing a detailed delthyrium and dorsal, lateral, and posterior views of an incomplete shell. Echinocoelia parva (5 to 7) shows an articulated shell in dorsal, lateral, and posterior views.

- Second row (8 to 12): Echinocoelia species indet. showing a complete shell in dorsal, ventral, lateral, posterior, and anterior views.

- Third row (13 to 17): Ambothyris species indet. showing a complete shell in dorsal, ventral, lateral, posterior, and anterior views.

- Middle left (18): A high-magnification detail of micro-ornamentation on the primary shell layer of Indospirifer.

- Fourth row (19 to 23): Biernatella ovalis showing a slightly damaged shell in dorsal, ventral, lateral, posterior, and anterior views.

- Fifth row (24 to 28): Emanuella aff. E. takwanensis showing an articulated shell in dorsal, ventral, lateral, posterior, and anterior views.

- Bottom rows (29 to 34): Indospirifer species indet. (29 to 33) showing a partially exfoliated shell in dorsal, ventral, lateral, posterior, and anterior views. Item 34 is a fragmentary ventral valve of ?Reticulariopsis species indet. in exterior view.

Scale bars are provided for each group: 1 mm for items 1 to 22, 2 mm for items 23 to 28, and 5 mm for items 29 to 34. The shells vary from smooth and rounded to heavily ribbed and triangular.

Reference Baliński1995 Biernatella ovalis Baliński, p. 142, figs. 7A–C, G–M, O–Q, S, T, 8A.

Type specimens

Holotype, ZPAL Bp XXXVIII/25-1, from the Szydłówek Beds of Bocianek (Baliński, Reference Baliński1995, fig. 7G–K). For paratypes, see Baliński (Reference Baliński1995, p. 138, 142).

Materials

A single shell with damaged ventral valve, GIUS 4-292/53.

Occurrence

Upper Givetian of the Holy Cross Mountains (Szydłówek, nowadays Bocianek, suburb of Kielce, type locality; Józefka).

Remarks

External features, e.g., a very small, smooth shell and a small ventral interarea, of the specimen from Józefka indicate that it represents the genus Biernatella—athyridids with double spires. It is externally undistinguishable from Biernatella ovalis from the lower part of the Szydłówek Beds (upper Givetian), cropping out at Bocianek, ~13 km west-northwest of Józefka. Both forms have an elongate suboval shell outline, with the maximum width of the shell located more anteriorly than in all other species of the genus.

Order Spiriferida Waagen, Reference Waagen1883

Suborder Spiriferidina Waagen, Reference Waagen1883

Superfamily Adolfioidea Sartenaer, Reference Sartenaer1966a

Family Echinospiriferidae Ljaschenko, Reference Ljaschenko1973

Genus Indospirifer Grabau, Reference Grabau1931

Type species

Spirifer padaukpinensis Reed, Reference Reed1908; Middle Devonian, probably Eifelian; Padaukpin, Northern Shan States, Myanmar (Burma).

Indospirifer sp. indet.

Figure 13.18, 13.29–13.33

Materials

A single shell, GIUS 4-294/37, with somewhat crushed anterior region, strongly decorticated, but with shell matter preserved in patches.

Description

Shell 29.3 mm wide, ~17 mm long, 16.7 mm thick, ventribiconvex; transverse with a long hinge margin, only slightly shorter than the maximum shell width at approximately one-third of the dorsal valve length; cardinal extremities narrowly rounded, lateral margins gently rounded, anterior margin truncated, anterior commissure uniplicate.

Ventral valve deeper than dorsal valve; interarea distinct, ~5 mm in height, apsacline, concave, with distinct longitudinal striation perpendicular to the hinge line; beak erect; delthyrium encompassing an angle of 52°, open but bordered by subvertical ridges; sulcus rather shallow, originating at the beak, laterally defined by bordering plications, anteriorly extended into a distinct tongue that is deflected almost at a right angle. Dorsal valve with a distinct fold appearing at the beak, bordered by a pair of distinct furrows.

Shell ornamentation of strong, rounded plications, 9 or 10 simple on each flank, four in sulcus, and five on fold, a single subdivision observed in the sulcus and on the fold; new plicae in the sulcus emerging from the plicae bounding the sulcus. Micro-ornament visible only on small patches of preserved shell matter (Fig. 12.16) consisting of concentric growth lines intersected by subradial capillae and sparsely distributed granules; capillae with a tendency to converge along the summits of the plicae.

The single available specimen reveals the presence of extrasinal dental plates and ctenophoridium.

Remarks

Only one subcomplete specimen of this species has been found and it is assigned to Indospirifer due to its plicate shell with a dorsal sulcus and a ventral fold, subradial capillae, and the presence of extrasinal dental plates and ctenophoridium. In these characters, it resembles the type species of the genus, I. padaukpinensis Reed, Reference Reed1908 from the Middle Devonian (reported as Eifelian, but not certain, see above) of Myanmar (Anderson et al., Reference Anderson, Boucot and Johnson1969) and perhaps also the lower Givetian of the Kuznetsk Basin (Russia; Izokh et al., Reference Izokh, Popov, Sobolev and Shcherbanenko2022), but differs mainly in a wider shell and a less inclined ventral interarea. Due to the damage to the anterior part of the studied shell, it is difficult to confirm the presence of the characteristic subradial arrangement of the median plicae that occurs in the type species. The specimen from Poland bears a strong external resemblance to I. exfoliata Cooper and Dutro, Reference Cooper and Dutro1982 described from the Frasnian Sly Gap Formation in New Mexico (Cooper and Dutro, Reference Cooper and Dutro1982), but the latter has a lower, more concave and more strongly inclined ventral interarea. Indospirifer orestes (Hall and Whitfield, Reference Hall and Whitfield1872) from the Frasnian Martin Formation in Arizona (Beus, Reference Beus1978) differs more clearly from the specimen studied here in its smaller size and a more elongated shell ornamented with fewer plicae.

Superfamily Ambocoelioidea George, Reference George1931

Family Ambocoeliidae George, Reference George1931

Subfamily Ambocoeliinae George, Reference George1931

Genus Ambothyris George, Reference George1931

Type species

Spirifera infima Whidborne, Reference Whidborne1893; Givetian; Lummaton, Devon, England.

Ambothyris sp. indet.

Figure 13.13–13.17

Reference Baliński and Halamski2023 Ambothyris sp.; Baliński and Halamski, p. 61, figs. 43A–Y, 44.

Materials

One complete and three almost complete shells, four shells damaged to varying degrees, GIUS 4-292/54–61.

Occurrence

A similar, if not conspecific form was described recently by Baliński and Halamski (Reference Baliński and Halamski2023) as Ambothyris sp. indet. from the lower Givetian beds at Miłoszów (Holy Cross Mountains, Poland).

Remarks

This rare brachiopod from Józefka shows great external similarity to Ambothyris sp. indet. described by Baliński and Halamski (Reference Baliński and Halamski2023) from the lower part of the Polygnathus timorensis Biozone of the Skały Formation at Miłoszów. Full discussion of taxonomic issues related to this form and comparisons with other similar species can be found in the above-mentioned paper.

Genus Echinocoelia Cooper and Williams, Reference Cooper and Williams1935

Type species

Echinocoelia ambocoelioides Cooper and Williams, Reference Cooper and Williams1935; Tully Formation, Givetian; New York, USA.

Remarks

This genus is represented in the studied material by two species. Echinocoelia parva Baliński in Baliński et al., Reference Baliński, Racki and Halamski2016 was described from early Frasnian strata in the Józefka section. In that contribution (Baliński et al., Reference Baliński, Racki and Halamski2016), the material from both Givetian and Frasnian strata was treated as a single species. The Frasnian Echinocoelia parva and the Givetian Echinocoelia sp. indet. are like each other in reaching similar size and having a similar ratio of the convexities of both valves, but differ in the outline of the valves (maximum width at approximate midlength of the dorsal valve in Echinocoelia parva, anteriorly in Echinocoelia sp. indet.). All specimens figured by Baliński et al. (Reference Baliński, Racki and Halamski2016) are from the Frasnian and represent the genuine Echinocoelia parva.

Echinocoelia parva Baliński in Baliński et al., Reference Baliński, Racki and Halamski2016

Figure 13.5–13.7

v*p Reference Baliński, Racki and Halamski2016 Echinocoelia parva Baliński in Baliński et al., Reference Baliński, Racki and Halamski2016, p. 157, pl. 17, figs. 8, 10–17, pl. 18, figs. 1–16.

Type specimens

Holotype, ZPAL Bp 73/1/23/1, from the lower Frasnian Wietrznia Beds of the Jóżefka roadcut (Baliński et al., Reference Baliński, Racki and Halamski2016, pl. 18, figs. 12–16). For paratypes, see Baliński et al. (Reference Baliński, Racki and Halamski2016, p. 157).

Materials

Five shells, GIUS 4-292/62–66.

Remarks

This species occurs sparsely in the Laskowa Góra Beds. It has been described in detail on the basis of a rich collection from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description.

For comparison with Echinocoelia sp. indet., see Remarks under the genus.

Echinocoelia sp. indet.

Figure 13.8–13.12

vp Reference Baliński, Racki and Halamski2016 Echinocoelia parva Baliński in Baliński et al., Reference Baliński, Racki and Halamski2016; p. 157, non pl. 17, figs. 8, 10–17, pl. 18, figs. 1–16.

Materials

Five shells, GIUS 4-292/67–71.

Description

Shell smooth, very small, reaching 3.9 mm in length, ventribiconvex, in the outline of a rounded triangle, with the greatest width slightly anterior to shell midlength. Ventral valve strongly convex, about twice as deep as the dorsal valve, with a massive umbo; sulcus not developed; interarea apsacline, curved, delthyrium open, beak erect to incurved. Dorsal valve weakly convex, with a broad and gentle median depression resulting in a slightly unisulcate anterior commissure.

Interior unknown. Micro-ornamentation poorly preserved, but concentric growth lines well discernible in places with preserved primary shell layer.

Remarks

The characteristic prominent, thick umbo of the ventral valve and the greatest width of the shell located rather anterior to the midlength, give the specimens studied a somewhat pear-shaped outline. This differentiates the described form from all other species in the genus Echinocoelia. The sample discussed here appears to represent a new species or subspecies, but no further attempts at taxonomic assessment have been made due to limited material.

For comparison with Echinocoelia parva, see Remarks under the genus.

Subfamily Rhynchospiriferinae Paulus, Reference Paulus1957

Genus Emanuella Grabau, Reference Grabau1923

Type species

Nucleospira takwanensis Kayser, Reference Kayser and von Richthofen1883; Middle Devonian; possibly Daguan (Takwan), Yunnan Province, southern China (see Hou et al., Reference Hou, Chen, Rong, Ma, Zhang, Xu, Su, Xian, Zong, Rong, Jon, Shen and Zhan2017, p. 458).

Emanuella aff. E. takwanensis (Kayser, Reference Kayser and von Richthofen1883)

Figure 13.24–13.28

aff. Reference Kayser and von Richthofen1883 Nucleospira takwanensis Kayser, p. 84, pl. 10, figs. 2, 2a–h.

v Reference Baliński, Racki and Halamski2016 Emanuella aff. takwanensis; Baliński et al., p. 158, text-fig. 14, pl. 18, figs. 17–31.

Materials

Two almost complete shells and a single fragmentary ventral valve, GIUS 4-292/72–74.

Remarks

This species occurs sparsely in the Laskowa Góra Beds. It has been described in detail on the basis of a rich collection from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description.

Superfamily Reticularioidea Waagen, Reference Waagen1883

Family Reticulariidae Waagen, Reference Waagen1883

Subfamily Reticulariopsinae Gourvennec, Reference Gourvennec1994

Genus Reticulariopsis Fredericks, Reference Fredericks1916

Type species

Spirifer (Reticularia) dereimsi Œhlert, Reference Œhlert1901; Emsian; Santa Lucia, León Province, Spain.

? Reticulariopsis sp. indet.

Figure 13.34

v Reference Baliński, Racki and Halamski2016 ?Reticulariopsis sp.; Baliński et al., p. 159, pl. 14, figs. 21–33.

Materials

Two fragmentary ventral valves, GIUS 4-292/75 and ZPAL Bp 73/1/25a/1.

Remarks

This species occurs sparsely in the Laskowa Góra Beds and in the Szydłówek Beds. It has been described in detail on the basis of a sample from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description.

Order Spiriferinida Ivanova, Reference Ivanova1972

Suborder Cyrtinidina Carter and Johnson in Carter et al., Reference Carter, Johnson, Gourvennec and Hou1994

Superfamily Cyrtinoidea Fredericks, Reference Fredericks1911

Family Cyrtinidae Fredericks, Reference Fredericks1911

Genus Cyrtina Davidson, Reference Davidson1859

Type species

Calceola heteroclita Defrance, Reference Defrance1824; Pragian; Néhou, Normandy, France.

Cyrtina cf. C. arkonensis Ehlers and Wright, Reference Ehlers and Wright1975

Figure 13.1–13.4

cf. Reference Ehlers and Wright1975 Cyrtina arkonensis Ehlers and Wright, p. 160, pl. 2, figs. 1–10, pl. 3, figs 1–20.

v Reference Baliński, Racki and Halamski2016 Cyrtina cf. arkonensis; Baliński et al., p. 160, pl. 19, figs. 1–19.

Materials

Two fragmentary shells and three incomplete ventral valves, GIUS 4-292/76–80.

Remarks

This species occurs sparsely in the Laskowa Góra Beds. It has been described in detail on the basis of a richer collection from the Wietrznia Beds (Baliński et al., Reference Baliński, Racki and Halamski2016); the reader is referred thereto for description. Difficulties in finding a meaningful taxonomy for the discussed genus were stressed by Halamski and Baliński (Reference Halamski and Baliński2013, p. 295), Gourvennec (Reference Gourvennec2019, p. 119), Halamski et al. (Reference Halamski, Baliński and Koppka2022a, p. 61), and Jansen (Reference Jansen2025, p. 217).

Order Terebratulida Waagen, Reference Waagen1883

Suborder Terebratulidina Waagen, Reference Waagen1883

Superfamily Cryptonelloidea Thomson, Reference Thomson1926

Family Cimicinellidae Stehli, Reference Stehli and Moore1965

Genus Cimicinoides Anderson, Boucot, and Johnson, Reference Anderson, Boucot and Johnson1969

Type species

Cimicinoides struvei Anderson, Boucot, and Johnson, Reference Anderson, Boucot and Johnson1969; Eifelian; Padaukpin, Northern Shan States, Myanmar (Burma).

? Cimicinoides sp. indet.

Figure 14

Figure 14.

?Cimicinoides sp. indet. from the Laskowa Góra Beds of Górno, Holy Cross Mountains: (1–5) almost complete shell, GIUS 4-292/81, in dorsal, ventral, lateral, posterior, and anterior views, respectively; (6) posterior region of the shell in dorsal view; (7) endopunctate shell exterior. Scale bars = 5 mm (1–5); 1 mm (6, 7).

Figure 14. Long description

A multi-panel photo series of a fossilized brachiopod shell.

Panel 1 shows the dorsal view of the almost complete shell, which is roughly diamond-shaped with a pointed apex and a weathered central patch.

Panel 2 shows the ventral view, displaying a similar shape with a slightly more uniform, pitted surface.

Panel 3 shows the lateral view, revealing the biconvex profile of the two valves meeting at a sharp commissure.

Panel 4 shows the posterior view, highlighting the hinge line and a small circular pedicle opening at the apex.

Panel 5 shows the anterior view, illustrating the curved edge where the valves meet.

A white scale bar representing 5 millimeters is positioned between panels 3 and 4.

Panel 6 is a close-up of the posterior region in dorsal view, providing a detailed look at the pedicle opening and the surrounding shell structure.

Panel 7 is a high-magnification view of the endopunctate shell exterior, showing a dense pattern of microscopic pits or pores across the surface.

White scale bars representing 1 millimeter are located at the bottom of panels 6 and 7.

Materials

A single, almost complete shell, GIUS 4-292/81.

Description

Shell deltoidal in outline, 11.5 mm long, 10.9 mm wide, and 5.1 mm thick, ventribiconvex; posterolateral margins almost straight, anterior margin narrowly rounded, giving the shell a slightly rhomboidal outline; anterior commissure rectimarginate. Ventral valve more than twice as deep as the dorsal valve; beak suberect; palintrope prominent, occupying approximately one-sixth of the shell width; delthyrium closed by deltidial plates pierced by a large hypothyrid foramen. Dorsal valve shallow, without perceptible fold.

Shell macroscopically smooth, shell substance densely punctate; punctae ~180–240 per mm². Interior unknown.

Remarks

The studied specimen reveals strong external resemblance to Cimicinoides struvei described from the Eifelian of Myanmar (Anderson et al., Reference Anderson, Boucot and Johnson1969). Both forms have similar shell size and shape, and a similarly prominent ventral palintrope and hypothyrid foramen. Differences include a slightly wider and less thick shell in the studied specimen.

Differences with other entirely smooth Devonian terebratulides are as follows: Centronella Billings, Reference Billings1859 has a mesothyrid to submesothyrid foramen (Cloud, Reference Cloud1942, p. 72); Oriskania Hall and Clarke, Reference Hall and Clarke1893 has a sulcate commissure (Cloud, Reference Cloud1942, p. 69); Elmaria Nalivkin, Reference Nalivkin, Nalivkin and Verber1947 apparently has a low dorsal fold (Nalivkin, Reference Nalivkin, Nalivkin and Verber1947, p. 133), but this genus is poorly known; Cryptonella Hall, Reference Hall1861 has a submesothyrid foramen (Jin and Lee, Reference Jin and Lee2006, p. 2024); Newberria Hall in Whiteaves, Reference Whiteaves1891 is larger (Cloud, Reference Cloud1942, p. 95) and has a more incurved beak. Because of the above-mentioned similarities and differences, the studied brachiopod is tentatively included in Cimicinoides, although evidently lack of knowledge about the internal structure makes this assignment provisional.

Glosshypothyridina

The genus Glosshypothyridina is represented by two undisputed species. The type species, G. procuboides, was described from the upper Eifelian of the Eifel and is also present in the lower Givetian (timorensis Biozone) of the Holy Cross Mountains (Baliński and Halamski, Reference Baliński and Halamski2023) and in the lower or middle Givetian of the Anti-Atlas (Halamski and Baliński, Reference Halamski and Baliński2013). Its other possible occurrences include the middle part of the Middle Devonian in Belgium and England (Sartenaer, Reference Sartenaer2003 and references therein), but these reports require confirmation. The species described here, G. theophili n. sp., is known solely from the upper Givetian of a single site of the Holy Cross Mountains (Fig. 15.1).

Figure 15.

Paleogeographic distribution of Hypothyridina, Glosshypothyridina, and Tullhypothyridina. Due to significant uncertainties in taxonomy, distribution is shown twice: (1) with confirmed occurrences only; (2) with all unconfirmed reports. See text for further explanation. Background map modified after Scotese and McKerrow (Reference Scotese and McKerrow1990), Golonka et al. (Reference Golonka, Krobicki, Pająk, Van Giang and Zuchiewicz2006), and Murphy et al. (Reference Murphy, van Staal and Collins2011).

Figure 15. Long description

A two-panel paleogeographic map set. A legend at the bottom identifies three symbols: a black square for Glosshypothyridina, a black triangle for Hypothyridina, and a black circle for Tullhypothyridina. Shading indicates deep sea in light gray, shallow sea in white, and land in dark gray.

Panel 1, labeled at the top left, shows confirmed occurrences only. The map features the supercontinents Euramerica and Gondwana separated by the Rheic Ocean, all surrounded by the Panthalassa ocean. Occurrences are concentrated in the tropical and subtropical regions of Euramerica and the northern margin of Gondwana. Specifically, one circle is in western Euramerica, one triangle is in central Euramerica, and three squares are clustered near the Rheic Ocean margin.

Panel 2, labeled at the middle left, shows all reports including unconfirmed ones. This map displays a much wider distribution. In addition to the points in Panel 1, there are several triangles and squares spread eastward across the shallow seas of northern Gondwana and into the eastern Panthalassa margins. The distribution forms a broad latitudinal band across the equatorial region of the globe.

On the basis of general similarity in external and internal characters between the two species mentioned above, it can be safely assumed that Glosshypothyridina theophili n. sp. is an evolutionary descendant of the earlier G. procuboides. This speciation event occurred in place, namely on the southern shelf of the eastern part of Euramerica (Laurussia), or in other words in an area corresponding to present-day Europe. Glosshypothyridina procuboides spans at least three million years, from the late Eifelian (~387 Ma; Becker et al., Reference Becker, Marshall, Da Silva, Gradstein, Ogg, Schmitz and Ogg2020) to the later part of the early Givetian (~384 Ma). After the Taghanic Event, it was replaced by G. theophili n. sp. in the late Givetian (~380–379 Ma), although details here are unclear, because no representatives of the genus are known from any precisely dated strata immediately following the Taghanic Event (contrary to the very precisely dated replacement of Atrypa subtrigonalis Biernat, 1964 by Kyrtatrypa pauli; Halamski et al., Reference Halamski, Baliński and Kondas2024).

Two further species included by Baxter (Reference Baxter1999) in this genus are Hypothyridina batschatensis Rzhonsnitskaya, Reference Rzhonsnitskaya and Markowski1968 and Hypothyridina vulgaria Rzhonsnitskaya, Reference Rzhonsnitskaya and Markowski1968 (including both var. typica and var. coronatiformis Rzhonsnitskaya, Reference Rzhonsnitskaya and Markowski1968) from the Eifelian of the Kuznetsk Basin and two additional ones, included tentatively, Rhynchonella procuboides var. lungtungpeensis Kayser, Reference Kayser and von Richthofen1883 and Hypothyridina lungtungpeensis magna Chen in Xu et al., Reference Xu, Wan and Chen1978 are from the Middle to Upper Devonian of China (Fig. 15.2). According to Sartenaer (Reference Sartenaer2003), definite systematic placement of each of them requires confirmation. Based on the data available so far, it can be concluded that Glosshypothyridina is a genus found only in the Old World Realm (sensu Boucot and Blodgett, Reference Boucot, Blodgett, Brunton, Cocks and Long2001; see Jansen et al., Reference Jansen, Halamski, Mottequin and Linnemann2025, p. 841).

Tullypothyridina and Hypothyridina

Concerning Tullypothyridina, Baxter (Reference Baxter1999) included seven (sub)species within this genus (Hemihypothyridina of Baxter, Reference Baxter1999, nomen nudum), all from the USA, namely: Hypothyridina venustula robusta Cooper and Williams, Reference Cooper and Williams1935; Hypothyridina venustula multicostata Cooper and Williams, Reference Cooper and Williams1935; Rhynchonella intermedia Barris, Reference Barris1879; Hypothyridina intermedia magniventra Stainbrook, Reference Stainbrook1942; Rhynchonella emmonsi Hall and Whitfield, Reference Hall and Whitfield1877; and Hypothyridina emmonsi rotunda Cooper and Dutro, Reference Cooper and Dutro1982. Three others were tentatively included: Hypothyridina bifurcata Brice, Reference Brice1982 from Arctic Canada, Hypothyridina calva Markovsky in Ljaschenko, Reference Ljaschenko1959a from Russia, and Hypothyridina hunanensis Wang, Reference Wang1956 from China. Sartenaer (Reference Sartenaer2003) accepted solely Rhynchonella intermedia from Iowa as a member of Tullypothyridina (Fig. 15.1) but estimated that the presence of the genus in Europe is likely (Fig. 15.2) on the basis of some specimens figured by Phillips (Reference Phillips1841). So, Tullypothridina might be either a genus present solely in the Eastern Americas Realm (sensu Boucot and Blodgett, Reference Boucot, Blodgett, Brunton, Cocks and Long2001; formerly the Appalachian Province, Boucot et al. Reference Boucot, Johnson and Talent1969; see Jansen et al., Reference Jansen, Halamski, Mottequin and Linnemann2025, p. 841) or present in both Eastern Americas and Old World realms.

On the contrary, as for Hypothyridina, besides its European type species (Fig. 15.1), Baxter (Reference Baxter1999) included two other species in this genus—Hypothyridina magister Belanski, Reference Belanski1928 from Iowa and Hypothyridina margarita Veevers, Reference Veevers1959 from Western Australia—whereas seven other taxa from both the Old World and Eastern Americas realms were included tentatively. These uncertain taxa include: Hypothyridina cuboides latusa Ljaschenko, Reference Ljaschenko1960 and Hypothyridina contracta Ljaschenko, Reference Ljaschenko1960, both from the Volgo-Uralian region; Hypothyridina linglingensis Wang, Reference Wang1956 from Hunan; Hypothyridina cameroni Warren, Reference Warren1944 from the Mackenzie River region; Hypothyridina mesodevonica Ljaschenko, Reference Ljaschenko1969 from the Russian Platform; Hypothyridina cuboides var. crassicostata Nalivkin, Reference Nalivkin1951 from the southern Urals; and Hypothyridina florens Crickmay, Reference Crickmay1963 from Mackenzie River region (see Baxter, Reference Baxter1999). It is thus likely that this genus was indeed cosmopolitan (Fig. 15.2), as was often previously considered on the basis of its wide understanding (Savage, Reference Savage and Kaesler2002; see criticism by Sartenaer, Reference Sartenaer2003, p. 36), but it must be emphasized that at present such a statement should be considered as a working hypothesis, requiring confirmation through careful study of internal characters of all the presumed representatives all over the world. As a minimum requirement, it is recommended that wide and narrow understanding of the species Hypothyridina cuboides be carefully distinguished (see, e.g., Jansen et al., Reference Jansen, Halamski, Mottequin and Linnemann2025, fig. 26.9.Ah).