Type species

Labechia conferta (Lonsdale, Reference Lonsdale and Murchison1839).

Labechia stylophora Nicholson, Reference Nicholson1891a

Figure 3.1, 3.2

Figure 3.

Longitudinal (1) and tangential (2) views of Labechia stylophora Nicholson, Reference Nicholson1891a , from the Middle Devonian of Shaldon, South Devon, UK, holotype NHMUK PI P5987; note the unclear pillars indicated by yellow arrows. Longitudinal (3) and tangential (4) views of Labechiella serotina (Nicholson, Reference Nicholson1886a) from the Middle Devonian of Teignmouth, Devon, UK, holotype NHMUK PI P5988. Longitudinal (5) and tangential (6) views of Clathrodictyon confertum Nicholson, Reference Nicholson1889, from the Middle Devonian of Hope’s Nose Quarry, Torquay, South Devon, UK, NHMUK PI H3891.

Reference Nicholson1891a   Labechia stylophora Nicholson, p. 161, pl. 20, figs. 7, 8.

Holotype

Specimen (NHMUK PI P5987) from the Middle Devonian of Shaldon, in South Devon, UK (Nicholson, Reference Nicholson1891a, pl. 20, figs. 7, 8).

Description

Labechia stylophora in this study is fragmented, but with an overall laminar form. Its skeletal structure consists of pillars with cyst plates between the pillars. In longitudinal section, pillars are thick, radiate slightly from the bottom and emerge to the surface (Fig. 3.1), spacing 2–4 per 2 mm, ranging 0.1–0.3 mm in width. Cyst plates are abundant, imbricating, with widths 0.5–0.7 mm. The galleries are generally crescent-shaped. In tangential section, pillars are round, isolated (Fig. 3.2), but locally rodlike in oblique section. Cyst plates are convex up, and irregularly or circularly connect pillars. Mamelons are conspicuous, 8–10 mm in diameter, manifested by the circularly arranged pillars and cyst plates, with adjacent two mamelons at a distance of 15 mm. Astrorhizal canals are not clear. The microstructure is compact.

Material

One specimen (NHMUK PI P5987) from the Middle Devonian of Shaldon, in South Devon (UK).

Remarks

Compared with diverse Ordovician, Silurian, and Late Devonian Labechia, the Middle Devonian records are rather rare. Nicholson (Reference Nicholson1891a) established this species according to the growth form of cylinder-shaped mamelons that is unique from the other known species.

Genus Labechiella Yabe and Sugiyama, Reference Yabe and Sugiyama1930

Type species

Labechia serotina Nicholson, Reference Nicholson1886a.

Labechiella serotina (Nicholson, Reference Nicholson1886a)

Figure 3.3, 3.4

Reference Nicholson1886a   Labechia serotina Nicholson, p. 15, pl. 2, figs. 3, 4.

Reference Webby and Selden2015   Labechiella serotina; Webby, p. E722.

Holotype

Specimen (NHMUK PI P5988) from the Middle Devonian of Teignmouth, South Devon, UK (Nicholson, Reference Nicholson1886a, pl. 2, figs. 3, 4).

Description

Growth forms are unclear due to the limitation of the thin sections. Latilaminae exist and presumably represent either growth interruptions or growth phases in the specimens; each latilamina is 10–20 mm thick. The skeleton is composed of thick pillars and thin laterally distributed cyst plates. In longitudinal section, pillars are thick, continuous, post-like, closely or loosely distributed (Fig. 3.3), spacing 4–5 per 2 mm, ranging from 0.15–0.3 mm in width. In some areas, the axial zone of pillars shows possibly overlying cone-in-cone banding. Cyst plates are thin, flat, concave up or concave down, generally complete, with rare cases overlapping the adjacent one, spacing 6–8 per 2 mm, 0.02 mm in thickness. Galleries are rectangular or rarely irregular. In tangential section, pillars are circular, but closely or incompletely connected, approximately forming a vermiform skeleton, with rare, isolated ones (Fig. 3.4). The axial zone of certain pillars illustrates lighter or darker areas. Microstructure is compact.

Material

Two specimens (NHMUK PI P5988, OR971), from the Middle Devonian of Teignmouth, in South Devon (UK).

Remarks

Labechiella serotina was originally assigned to the genus Labechia by Nicholson (Reference Nicholson1886a), but Labechia is markedly different from Labechiella in terms of imbricating cyst plates (Yabe and Sugiyama, Reference Yabe and Sugiyama1930). Thus, this species was revised from Labechia into Labechiella. Detailed discussion of the difference between Labechiella and its related genera is provided by Webby (Reference Webby and Selden2015, p. E722–E723).

Order Clathrodictyida Bogoyavlenskaya, Reference Bogoyavlenskaya1969

Family Clathrodictyidae Kühn, Reference Kühn1939

Genus Clathrodictyon Nicholson and Murie, Reference Nicholson and Murie1878

Type species

Clathrodictyon vesiculosum Nicholson and Murie, Reference Nicholson and Murie1878.

Clathrodictyon confertum Nicholson, Reference Nicholson1889

Figure 3.5, 3.6

Reference Nicholson1889   Clathrodictyon confertum Nicholson, p. 154, pl. 18, figs. 13, 14.

Reference Galloway1957   Clathrodictyon confertum; Galloway and St. Jean, p. 94, pl. 1, fig. 2a, b.

Reference Bjerstedt and Feldmann1985   Clathrodictyon confertum; Bjerstedt and Feldmann, p. 1055, fig. 11.7, 11.8.

Holotype

Specimen (NHMUK PI P5981) from the Middle Devonian of Pit Park Quarry, Dartington, South Devon, UK (Nicholson, Reference Nicholson1889, pl. 18, figs. 13, 14).

Description

External forms are unconfirmed because of the fragmented skeleton. The skeleton is formed by densely distributed laminae and short pillars. In longitudinal section, pillars are short, confined to the interlaminar spaces (Fig. 3.5), irregularly distributed, 0.02 mm in width. Laminae are thin, undulating, crumpled, locally continuous, but generally intermittent, spacing 14–16 per 2 mm, 0.02 mm in thickness. Galleries are irregular, circular, or sub-rectangular. In tangential section, laminae are not apparent. Pillars are locally isolated, but commonly connected with other elements (Fig. 3.6). Astrorhizal canals are unclear. The microstructure is compact.

Material

Three specimens from the Middle Devonian of Pit Park Quarry, Dartington (NHMUK PI P5981) and Hope’s Nose Quarry, Torquay (NHMUK PI H3891, P6110), in South Devon (UK).

Remarks

Clathrodictyon confertum is featured by densely packed, thin, wrinkled laminae and short pillars. It closely resembles Clathrodictyon vesiculosum in terms of wrinkled laminae and confined pillars, but C. confertum presents a smaller and closely distributed skeleton (Nicholson, Reference Nicholson1889; Galloway and St. Jean, Reference Galloway and St. Jean1957).

Clathrodictyon ? heathsense Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993

Figure 4.1, 4.2

Figure 4.

Longitudinal (1) and tangential (2) sections of Clathrodictyon? heathsense Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993, from the Lower Devonian Cave limestone of New South Wales, Australia, NHMUK PI H3892. Longitudinal (3) and tangential (4) sections of Petridiostroma delicatulum (Ripper, Reference Ripper1937a) from the Lower Devonian of Rocky Camp Quarry, Buchan, Victoria, Australia, holotype NHMUK PI H3914.

Reference Ripper1937a   Clathrodictyon regulare; Ripper, p. 16, pl. 3, figs. 1, 2.

Reference Ripper1937a   Clathrodictyon convictum; Ripper, p. 19, pl. 3, figs. 4–8.

Reference Webby, Stearn and Zhen1993   Clathrodictyon? heathsense; Webby, Stearn, and Zhen, p. 134, figs. 11C–F, 12A.

Holotype

Longitudinal and tangential sections (NMV P I41798 and NMV P I41799) from the Lower Devonian of Heath’s Quarry, Buchan Caves Limestone, Buchan, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 11C, F).

Description

Specimens were preserved as fragments; therefore, growth forms are unclear. The skeleton comprises short pillars and laterally extensive laminae. In longitudinal section, pillars are short, post-like or spool-shaped, generally confined to one interlaminar space (Fig. 4.1), spacing about 7–9 per 2 mm and 0.05–0.2 mm in width. Laminae are thin, inflected upward and downward, horizontally continuous, spacing about 9–11 per 2 mm and 0.05–0.15 mm in thickness. Galleries are rectangular or locally circular. Syringoporid symbionts, 0.3–0.5 mm in diameter, spacing 2–3 per 2 mm, are associated, as well as spiral-upward tubeworms (Fig. 4.1). In tangential section, pillars are circular or irregular, commonly isolated (Fig. 4.2), 0.1–0.2 mm in diameter. Laminae are irregularly undulating. Syringoporids and tubeworms are circular. Microstructure is compact.

Material

Six specimens from the Lower Devonian of Heath’s Quarry (NHMUK PI H3917, H3918, H3922, H3924), in Buchan; Cave limestone (NHMUK PI H3892) of New South Wales; and Mitchell’s Quarry, Cave Hill, Lilydale (NHMUK PI H3900), all in Victoria (Australia).

Remarks

This species was identified as Clathrodictyon regulare (Rosen, Reference Rosen1867) and Clathrodictyon convictum Ripper, Reference Ripper1937a. However, the type specimens of these two species were revised to Petridiostroma based on their regular skeletons formed by planar laminae and short pillars (Stearn, Reference Stearn1992). The Australian materials, which closely resemble Clathrodictyon, show slightly bent laminae. Webby et al. (Reference Webby, Stearn and Zhen1993) thus reassigned these specimens as Clathrodictyon? heathsense. This study follows the identification of Webby et al. (Reference Webby, Stearn and Zhen1993). It should be noted that the present specimens differ from Stictostroma in possessing compact microstructure, contrasting with the tripartite structure of Stictostroma laminae.

Family Gerronostromatidae Bogoyavlenskaya, Reference Bogoyavlenskaya1969

Genus Petridiostroma Stearn, Reference Stearn1992

Type species

Simplexodictyon simplex Nestor, Reference Nestor1966.

Petridiostroma delicatulum (Ripper, Reference Ripper1937a)

Figure 4.3, 4.4

Reference Ripper1937a   Clathrodictyon convictum var. delicatula Ripper, p. 20, pl. 4, figs. 1, 2.

Reference Webby, Stearn and Zhen1993   Petridiostroma delicatulum; Webby, Stearn, and Zhen, p. 134, fig. 9E, F.

Holotype

Specimen (NHMUK PI H3914) from the Lower Devonian of Rocky Camp (Commonwealth) Quarry, Buchan, Victoria, Australia (Ripper, Reference Ripper1937a, pl. 4, figs. 1, 2).

Description

Growth forms appear to be laminar, but specimens were cut as fragments. The skeleton is composed of short pillars and horizontally continuous laminae. In longitudinal section, pillars are short, post-like or spool-shaped, commonly confined to one interlaminar space, locally superposed across laminae, spacing about 12–14 per 2 mm and 0.05–0.2 mm in width (Fig. 4.3). Laminae are thin, slightly undulating, laterally extensive, spacing about 15–17 per 2 mm and 0.05–0.1 mm in thickness. Galleries are rectangular or circular. Syringoporid tabulate corals are common, with a width of 0.3–0.4 mm, spacing 2–4 per 2 mm. In tangential section, pillars are circular, isolated, 0.05–0.15 mm in diameter (Fig. 4.4). Laminae are irregularly undulating. Syringoporids are circular in tangential sections. Microstructure is compact.

Material

One specimen (NHMUK PI H3914) from the Lower Devonian of Rocky Camp (Commonwealth) Quarry, Buchan, Victoria (Australia).

Remarks

This species is characterized mainly by planar laminae and short pillars, which are close to Petridiostroma rather than Clathrodictyon. Petridiostroma delicatulum has more densely distributed laminae and pillars than Stictostroma clarum (Počta, Reference Počta1894).

Petridiostroma sp.

Figure 5.1, 5.2

Figure 5.

Longitudinal (1) and tangential (2) views of Petridiostroma sp. from the Lower Devonian of Griffith’s Quarry, Loyola, Victoria, Australia, NHMUK PI H3930. Longitudinal (3) and tangential (4) views of Gerronostromaria buchanense (Flügel, Reference Flügel1959) from the Lower Devonian of Commonwealth Quarries, Buchan, Victoria, Australia, NHMUK PI H3913. Longitudinal (5) and tangential (6) views of Actinostroma clathratum Nicholson, Reference Nicholson1886a , from the Middle Devonian of Gerolstein, Germany, lectotype NHMUK PI P5774.

Reference Ripper1933   Clathrodictyon regulare cylindriferum; Ripper, p. 157, pl. 3, fig. 6A, B.

Reference Ripper1937b   Clathrodictyon regulare; Ripper, p. 2, pl. 1, figs. 1, 2.

Reference Ripper1937b   Clathrodictyon aff. chapmani; Ripper, p. 3, pl. 1, figs. 3, 4.

Reference Webby, Stearn and Zhen1993   Petridiostroma sp.; Webby, Stearn, and Zhen, p. 134, fig. 11A, B.

Description

Growth forms are possibly laminar, but poorly preserved. The skeleton comprises short pillars and laterally extensive laminae. In longitudinal section, pillars are short, spool-shaped, generally confined to one interlaminar space, locally superposed, spacing about 7–10 per 2 mm and 0.05–0.2 mm in width (Fig. 5.1). Laminae are thin, gently undulating, horizontally continuous, spacing about 9–11 per 2 mm and 0.05–0.15 mm in thickness. Galleries are round or rarely rectangular. In tangential section, pillars are circular or irregular, commonly isolated, 0.1–0.2 mm in diameter (Fig. 5.2). Laminae are irregularly undulating. Microstructure is compact.

Material

One specimen from the Lower Devonian of Griffith’s Quarry (NHMUK PI H3930), Loyola, Victoria (Australia).

Remarks

Petridiostroma sp. was assigned by Ripper (Reference Ripper1937b) to Clathrodictyon regulare and Clathrodictyon aff. chapmani Ripper, Reference Ripper1933, but C. regulare shows closely spaced laminae of 18 per 2 mm (see Nestor, Reference Nestor1964) and the type specimen of the latter species (Ripper, Reference Ripper1933) was revised into Atelodictyon (Webby et al., Reference Webby, Stearn and Zhen1993). In addition, the planar laminae resemble Petridiostroma. The current species is left in open nomenclature due to the poorly preserved skeleton. It features round galleries and finer texture, which is different from P. delicatulum.

Genus Gerronostromaria Nestor, Reference Nestor2011

Type species

Gerronostroma elegans Yavorsky, Reference Yavorsky1931.

Gerronostromaria buchanense (Flügel, Reference Flügel1959)

Figure 5.3, 5.4

Reference Ripper1937a   Actinostroma contortum; Ripper, p. 14, pl. 2, figs. 3–6.

Reference Flügel1959   Actinostroma buchanense; Flügel, p. 183, pl. 7, fig. 4.

Reference Webby, Stearn and Zhen1993   Gerronostromaria buchanense; Webby, Stearn, and Zhen, p. 130, fig. 9A–D.

Holotype

Specimen (NMV P141758) from the Lower Devonian Buchan Caves Limestone at Heath’s Quarry, Buchan, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 9A).

Description

Growth forms were described as domical or bulbous by Ripper (Reference Ripper1937b). In longitudinal section, pillars are long, passing through several laminae, spacing about 10–12 per 2 mm and 0.05–0.2 mm in width (Fig. 5.3). Laminae are thin, apparently undulating, laterally continuous, spacing about 14–17 per 2 mm and 0.05–0.15 mm in thickness. Galleries are rectangular. In tangential section, pillars are circular and isolated, 0.1–0.2 mm in diameter. Laminae are concentric circular shapes (Fig. 5.4). Mamelons are distinct, 4–5 mm in diameter. Microstructure is compact.

Material

Three specimens (NHMUK PI H3909, H3911, H3913) from the Lower Devonian of Commonwealth Quarry, Buchan, Victoria (Australia).

Remarks

This species was assigned into Actinostroma by Ripper (Reference Ripper1937b) based on the tiny rod-like structures, but colliculi and hexactinellid networks in typical Actinostroma (Fig. 2.3) have not been noticed. Regular networks formed by extensive pillars and laminae justify its assignment to Gerronostromaria.

Order Actinostromatida Bogoyavlenskaya, Reference Bogoyavlenskaya1969

Family Actinostromatidae Nicholson, Reference Nicholson1886b

Genus Actinostroma Nicholson, Reference Nicholson1886b

Type species

Actinostroma clathratum Nicholson, Reference Nicholson1886b.

Actinostroma clathratum Nicholson, Reference Nicholson1886b

Figure 5.5, 5.6

Reference Nicholson1886b   Actinostroma clathratum Nicholson, p. 226, pl. 6, figs. 1–3.

Reference Nicholson1889   Actinostroma clathratum; Nicholson, p. 131, pl. 1, figs. 8–13; pl. 12, figs. 1–5.

Reference Yavorsky1930    Actinostroma clathratum; Yavorsky, p. 475, pl. 1, figs. 1–10.

Reference Chi1940    Actinostroma clathratum; Chi, p. 309, pl. 2, fig. 2.

Reference Lecompte1951    Actinostroma clathratum; Lecompte, p. 99, pl. 1, figs. 1–12.

Reference Galloway1957   Actinostroma clathratum; Galloway and St. Jean, p. 149, pl. 10, fig. 4a, b.

Reference Zukalová1971    Actinostroma clathratum; Zukalová, p. 31, pl. 3, figs. 3–5.

Reference Yang and Dong1979    Actinostroma clathratum; Yang and Dong, p. 31, pl. 9, figs. 1, 2.

Reference Stock1982    Actinostroma clathratum; Stock, p. 669, pl. 3, figs. 5, 6.

Reference Dong, Wang, Zhou, Zhang, Luo, Fu and Huang1989    Actinostroma clathratum; Dong et al., p. 264, pl. 5, fig. 2a, b.

Reference Webby and Zhen1993    Actinostroma clathratum; Webby and Zhen, p. 329, fig. 3A–D.

Reference Krebedünkel1995    Actinostroma clathratum; Krebedünkel, p. 23, pl. 1, figs. 1, 2.

Reference Webby and Zhen1997    Actinostroma clathratum; Webby and Zhen, p. 13, fig. 6A, 6B, 6D.

Reference Stearn, Webby, Nestor and Stock1999    Actinostroma clathratum; Stearn et al., p. 32, fig. 5E, F.

Reference May2005    Actinostroma clathratum; May, p. 134, pl. 1, figs. 1, 2; pl. 27, figs. 3–5; pl. 29, fig. 2.

Reference Salerno2008    Actinostroma clathratum; Salerno, p. 52, pl. 6, fig. 3.

Lectotype

Specimen (NHMUK PI P5774; Stock, Reference Stock and Selden2015) from the Middle Devonian of Gerolstein, Germany (Nicholson, Reference Nicholson1889, pl. 1, figs. 11, 12).

Description

Growth forms are unclear based on thin sections, but Nicholson (Reference Nicholson1886a) noted that they are commonly bulbous or domical. The skeleton comprises vertically dominant pillars and laterally subordinate colliculi. In longitudinal section, pillars are thick, long, continuous (Fig. 5.5), possibly throughout the whole skeleton or at least each latilamina, spacing about 8–11 per 2 mm and 0.1–0.2 mm in width. In some oblique sections, pillars appear to be short, passing through 1–3 laminae. Colliculi are thin, gently undulating, locally cut as isolated circular dots, spacing about 7–9 per 2 mm and 0.05–0.15 mm in thickness. Galleries are rectangular or round. In tangential section, pillars are circular and isolated between the interlaminar spaces, 0.1–0.2 mm in diameter. Colliculi are interconnected in the previously called laminae, forming a hexactinellid network (Fig. 5.6). Microstructure is compact.

Material

Twenty-five specimens from the Middle Devonian of Dartington (NHMUK PI P5708, H3362), Teignmouth (NHMUK PI P5716, P5717, P5719, P5720, P5721), Kingsteignton (NHMUK PI H4335, H4336), Torquay (NHMUK PI H5286) and six unknown locations (NHMUK PI P5718, P2202, P2227, R1217, 25, 15243), in South Devon (UK); Gerolstein (NHMUK PI P5774, P5779, P5787), Sötenich (NHMUK PI P5777), Hebborn (NHMUK PI P5778, P5783, P5786), and Büchel (NHMUK PI P6090), in Germany; and Rough Range (NHMUK PI P4967) near Mt. Krauss, Kimberley (western Australia).

Remarks

According to Nicholson (Reference Nicholson1886a), this species exhibits variability in external form, but the internal structure is constant. It is characterized by regularly developed long pillars and interrupted colliculi. It resembles A. expansum but differs from it in having thinner skeletons.

Actinostroma compactum Ripper, Reference Ripper1933

Figure 6.1, 6.2

Figure 6.

Longitudinal (1) and tangential (2) sections of Actinostroma compactum Ripper, Reference Ripper1933, from the Lower Devonian of Rocky Camp Quarry, Buchan, Victoria, Australia, NHMUK PI H3910. Longitudinal (3) and tangential (4) sections of Actinostroma expansum (Hall and Whitfield, Reference Hall and Whitfield1873) from the Devonian of Rockford, Iowa (USA), NHMUK PI P5803. Longitudinal (5) and tangential (6) sections of Actinostroma fenestratum Nicholson, Reference Nicholson1889, from the Devonian of Lake Manitoba, Manitoba, Canada, NHMUK PI P5601.

Reference Ripper1933    Actinostroma compactum Ripper, p. 153, fig. 5A, B.

Reference Ripper1937a    Actinostroma compactum; Ripper, p. 15, pl. 2, figs. 7, 8.

Reference Webby, Stearn and Zhen1993    Actinostroma compactum; Webby, Stearn, and Zhen, p. 123, figs. 5A–F, 7A–C.

Reference Webby and Zhen1997    Actinostroma compactum; Webby and Zhen, p. 17, fig. 7A, B.

Reference May1999b    Actinostroma compactum; May, p. 99, pl. 1, fig. 1.

Reference May, Huemann and Piller2007    Actinostroma compactum; May, p. 143.

Holotype

Specimen (NMV P141959-60; Webby et al., Reference Webby, Stearn and Zhen1993) from the Lilydale Limestone at Mitchell’s (Cave Hill) Quarry, Lilydale, Victoria, Australia (Ripper, Reference Ripper1933, fig. 5A, B).

Description

External forms are uncertain due to poor preservation. The skeleton comprises vertically long pillars and lateral colliculi. In longitudinal section, pillars are long, continuous, passing through dozens of colliculi (Fig. 6.1), spacing about 10–12 per 2 mm and 0.1–0.15 mm in width. Colliculi are thin, locally oblique, laterally discontinuous, showing isolated circular dots, spacing about 12–14 per 2 mm and 0.05–0.1 mm in thickness. Galleries are rectangular, irregular or round. In tangential section, pillars are circular and isolated between the interlaminar spaces, 0.1–0.15 mm in diameter. Hexactinellid networks occur from 4–5 colliculi interconnecting each pillar (Fig. 6.2). Tubeworms are locally observed, with a diameter of 1 mm. Microstructure is compact.

Material

Two specimens (NHMUK PI H3910, H3923) from the Lower Devonian of Rocky Camp (Commonwealth) Quarry, Buchan, Victoria (Australia).

Remarks

Compared with A. clathratum, A. compactum shows more densely packed and thinner pillars and colliculi.

Actinostroma expansum (Hall and Whitfield, Reference Hall and Whitfield1873)

Figure 6.3, 6.4

Reference Hall and Whitfield1873    Stromatopora expansa Hall and Whitfield, p. 226, pl. 9, fig. 1.

Reference Nicholson1891b    Actinostroma expansum; Nicholson, p. 316, pl. 10, fig. 1, 2.

Reference Kaźmierczak1971    Actinostroma expansum; Kaźmierczak, p. 136, pl. 40, fig. 2a, b; pl. 41, fig. 5.

Reference Stock1984    Actinostroma expansum; Stock, p. 774, fig. 2A–E.

Reference Stearn1996    Actinostroma expansum; Stearn, p. 200, figs. 2.4, 2.6, 3.1, 3.2.

Holotype

Specimen (NYSM 331) from the Upper Devonian (Frasnian) Nora Member of the Shell Rock Formation, Rockford, Floyd County, Iowa, USA (Stock, Reference Stock1984, fig. 2A–D).

Description

External forms are unclear due to poor preservation. Skeleton is composed of vertically thick pillars and lateral colliculi. In longitudinal section, pillars are long, continuous, passing through several colliculi (up to 19), spacing about 6–8 per 2 mm and 0.1–0.25 mm in width (Fig. 6.3). Colliculi are thin, undulating, laterally discontinuous, locally showing isolated circular dots, spacing about 7–10 per 2 mm and 0.05–0.1 mm in thickness. Galleries are rectangular or round. In tangential section, pillars are circular and isolated, 0.1–0.2 mm in diameter. Hexactinellid networks are formed by 4–5 colliculi interconnecting each pillar (Fig. 6.4). Microstructure is compact.

Material

Three specimens from the Upper Devonian: two from Rockford, Iowa (USA) (NHMUK PI P5803, P5807) and another one from Rough Range near Mt. Krauss, Kimberley (western Australia) (NHMUK PI P4463).

Remarks

Actinostroma expansum differs from other species in widely spaced and thicker colliculi and pillars.

Actinostroma fenestratum Nicholson, Reference Nicholson1889

Figure 6.5, 6.6

Reference Nicholson1889    Actinostroma fenestratum Nicholson, p. 146, pl. 17, figs. 8, 9.

Holotype

Specimen (NHMUK PI P6116) from the Middle Devonian of Teignmouth, South Devon, UK (Nicholson, Reference Nicholson1889, pl. 17, figs. 8, 9).

Description

Growth forms were described as domical/bulbous by Nicholson (Reference Nicholson1889, Reference Nicholson1891a). The skeleton consists mainly of long pillars and subordinate colliculi. In longitudinal section, pillars are rodlike, thick, long, continuous, spacing about 5–6 per 2 mm and 0.3–0.6 mm in width (Fig. 6.5). Colliculi are generally cut as isolated circular dots, spacing about 4–5 per 2 mm and 0.1–0.3 mm in thickness. Galleries are irregular. In tangential section, pillars are round and isolated, 0.2–0.5 mm in diameter. Colliculi are not well developed, interconnecting with pillars, forming irregular or vermicular networks (Fig. 6.6). Microstructure is compact.

Material

Four specimens from the Devonian: one from Lake Manitoba (NHMUK PI P5601), in Manitoba (Canada) and three more from Teignmouth (NHMUK PI P5884, P6111, P6116), in South Devon (UK).

Remarks

Actinostroma fenestratum is different from other species by thick, long pillars and poorly developed colliculi and networks. It also resembles Neosyringostroma logansportense (Galloway and St. Jean, Reference Galloway and St. Jean1957), but the latter shows more well-developed lateral skeletons and cellular microstructure.

Actinostroma hebbornense Nicholson, Reference Nicholson1886b

Figure 7.1, 7.2

Figure 7.

Longitudinal (1) and tangential (2) views of Actinostroma hebbornense Nicholson, Reference Nicholson1886b , from the Middle Devonian of Hebborn, Germany, holotype NHMUK PI P5581. Longitudinal (3, 4) and tangential (5, 6) sections Actinostroma stellulatum Nicholson, Reference Nicholson1886b , from the Middle Devonian of Gerolstein, Germany, holotype NHMUK PI P5570.

Reference Nicholson1886b    Actinostroma hebbornense Nicholson, p. 228, pl. 7, figs. 7, 8.

Reference Nicholson1889    Actinostroma hebbornense; Nicholson, p. 137, pl. 16, figs. 9–16.

Reference Lecompte1951    Actinostroma hebbornense; Lecompte, p. 92, pl. 3, figs. 4–6.

Reference Zukalová1971    Actinostroma hebbornense; Zukalová, p. 30, pl. 3, figs. 1, 2,

Reference Neidhardt1985    Actinostroma hebbornense; Neidhardt, p. 16, pl. 1, figs. 5, 6.

Reference Krebedünkel1995    Actinostroma hebbornense; Krebedünkel, p. 30, pl. 1, figs. 7, 8.

Holotype

Specimen (NHMUK PI P5581) from the Middle Devonian of Hebborn, Germany (Nicholson, Reference Nicholson1886b, pl. 7, figs. 7, 8).

Description

External forms were noted as domical or bulbous by Nicholson (Reference Nicholson1889). The skeleton is composed of vertically thick pillars and lateral colliculi. In longitudinal section, pillars are rodlike, long, continuous, but commonly appear to be short in oblique sections (Fig. 7.1), spacing about 9–11 per 2 mm and 0.1–0.2 mm in width. Colliculi are generally undulating, commonly cut as isolated circular dots, spacing about 10–14 per 2 mm and 0.05–0.1 mm in thickness. Galleries are rectangular or locally circular. In tangential section, pillars are round and isolated dots between interlaminar spaces, 0.05–0.2 mm in diameter. Colliculi are rodlike, 0.05–0.1 mm in width, interconnecting with pillars, forming hexactinellid networks (Fig. 7.2). Syringoporids and tubeworms locally occur. Microstructure is compact.

Material

Five specimens from the Middle Devonian: two from Teignmouth (NHMUK PI P5579, P5950), one from Torquay (NHMUK PI P6106), in South Devon (UK), and another two from the Devonian of Hebborn (NHMUK PI P5573, P5581) in Germany.

Remarks

According to Nicholson (Reference Nicholson1889), this species is featured by regular hexactinellid networks, slender and densely arranged pillars, and the uncommon occurrence of astrorhizae. It is different from A. clathratum in thinner and oblique pillars. It is distinguished from A. compactum by regular and planar colliculi.

Actinostroma stellulatum Nicholson, Reference Nicholson1886b

Figure 7.3–7.6

Reference Nicholson1886b    Actinostroma stellulatum Nicholson, p. 231, pl. 6, figs. 8, 9.

Reference Nicholson1889    Actinostroma stellulatum; Nicholson, p. 140, pl. 14, figs. 1–8; pl. 15.

Reference Lecompte1951    Actinostroma stellulatum; Lecompte, p. 111, pl. 11, figs. 1–5.

Reference Sleumer1969    Actinostroma stellulatum; Sleumer, p. 34, pl. 21, fig. 1; pl. 22, fig. 2.

Reference Kaźmierczak1971    Actinostroma stellulatum; Kaźmierczak, p. 133, pl. 37, fig. 3a, b.

Reference May1993    Actinostroma stellulatum; May, p. 29, pl. 2, fig. 1.

Reference Krebedünkel1995    Actinostroma stellulatum; Krebedünkel, p. 45, pl. 4, figs. 1, 2.

Reference Méndez-Bedia1999    Actinostroma stellulatum; Méndez-Bedia, p. 124, fig. 2A, B.

Reference Salerno2008    Actinostroma stellulatum; Salerno, p. 60, pl. 9, fig. 1.

Holotype

Specimen (NHMUK PI P5570) from the Middle Devonian of Gerolstein, Germany (Nicholson, Reference Nicholson1886b, pl. 6, figs. 8, 9).

Description

Growth forms are laminar based on limited specimens, but Nicholson (Reference Nicholson1889) also noted the occurrence of bulbous or columnar shapes. The skeleton consists of vertically short pillars and laterally long laminae. In longitudinal section, pillars are short (Fig. 7.3), commonly upward spreading (Fig. 7.4), confined to one interlaminar space, locally superposed, spacing about 9–14 per 2 mm and 0.1–0.15 mm in width. Colliculi are thin, laterally continuous, slightly undulating, spacing about 11–13 per 2 mm and 0.05–0.1 mm in thickness, with a central dark line. Galleries are arch-shaped, rectangular, or round. Mamelons are inconspicuous, 0.2 mm in height. In tangential section, pillars are round and isolated, 0.1–0.15 mm in diameter. Colliculi are curved (Fig. 7.5). Radial processes are small, thin, rod-like, 0.01–0.03 mm in diameter, radiating from pillars and forming regular hexactinellid networks between the inter-laminae spaces (Fig. 7.6). Astrorhizae are distinct, with central and branching canals, 5–7 mm in diameter. Microstructure is compact.

Material

Fifteen specimens from the Middle Devonian of Dartington (NHMUK PI P5943, P5948), Teignmouth (NHMUK PI P5942, P5944, P5945, P5946, P5947), Bishopsteignton (NHMUK PI H4334, H4340) in South Devon (UK), and Gerolstein (NHMUK PI P5570, P5583, P5858, P5860, P5862, P6089), in Germany.

Remarks

As noted by Stock in Stearn et al. (Reference Stearn, Webby, Nestor and Stock1999), Actinostroma stellulatum is different from other species in atypical features, such as short pillars and thin colliculi.

Actinostroma verrucosum (Goldfuss, Reference Goldfuss1826)

Figure 8.1–8.3

Figure 8.

Longitudinal (1) and tangential (2, 3) sections of Actinostroma verrucosum (Goldfuss, Reference Goldfuss1826) from the Middle Devonian of Büchel, Germany, NHMUK PI P5537. Longitudinal (4) and tangential (5, 6) sections of Bifariostroma bifarium (Nicholson, Reference Nicholson1886b) from the Middle Devonian of Büchel, Germany, holotype NHMUK PI P5639.

Reference Goldfuss1826    Ceriopora verrucosa Goldfuss, p. 33, pl. 10, fig. 6a–c.

Reference Bargatzky1881a    Stromatopora verrucosa; Bargatzky, p. 283.

Reference Nicholson1889    Actinostroma verrucosum; Nicholson, p. 134, pl. 16, figs. 1–8.

Reference Lecompte1951    Actinostroma verrucosum; Lecompte, p. 107, pl. 9, figs. 1–8; pl. 10, figs. 1, 2.

Reference Flügel1959    Actinostroma verrucosum; Flügel, p. 190, pl. 6, fig. 4.

Reference Zukalová1971    Actinostroma verrucosum; Zukalová, p. 36, pl. 5, figs. 5, 6.

Reference Mistiaen1980    Actinostroma verrucosum; Mistiaen, p. 185, pl. 2, figs. 7–9; pl. 3, figs. 1–3.

Reference May1993    Actinostroma verrucosum; May, p. 28, pl. 2, fig. 2.

Reference Krebedünkel1995    Actinostroma verrucosum; Krebedünkel, p. 36, pl. 5, figs. 5, 6.

Reference Méndez-Bedia1999    Actinostroma verrucosum; Méndez-Bedia, p. 124, fig. 2C–E.

Reference Salerno2008    Actinostroma verrucosum; Salerno, p. 63, pl. 8, figs. 1–3; pl. 12.

Lectotype

Specimen (BIOB-PAL-Goldfuss-121b) from the Middle Devonian of Bensberg, Germany (Goldfuss, Reference Goldfuss1826, pl. 10, fig. 6).

Description

External forms are unclear due to poor preservation, but Nicholson (Reference Nicholson1889) mentioned the existence of bulbous or domical shapes. The skeleton is composed of vertically thick pillars and lateral colliculi. In longitudinal section, pillars are post-like or spool-shaped, confined to one interlaminar space but systematically superposed, and commonly appear to be short in oblique sections, spacing about 8–10 per 2 mm and 0.1–0.2 mm in width (Fig. 8.1). Colliculi are conspicuously undulating, locally showing isolated circular dots, spacing about 10–12 per 2 mm and 0.05–0.1 mm in thickness. Galleries are circular. Mamelons are conspicuous, 2–3 mm in height. In tangential section, pillars are round and isolated between the interlaminar spaces (Fig. 8.2). Five to seven colliculi connected with each pillar, forming hexactinellid networks, 0.05–0.15 mm in width (Fig. 8.3). Mamelons are observed as concentric circles, at a distance of 3–5 mm from one to another. Microstructure is compact.

Material

Ten specimens from the Middle Devonian of Teignmouth (NHMUK PI P5538, P5940), in South Devon (UK) and Sötcnich (NHMUK PI P5535, P5540, H3243), Gerolstein (NHMUK PI P5536), Büchel (NHMUK PI P5537, P5539, P5544), Borbach (NHMUK PI P5545), in Germany.

Remarks

Actinostroma verrucosum is different from other species in having short but superposed pillars, prominent colliculi, and mamelons.

Genus Bifariostroma Khalfina, Reference Khalfina and Ivaniya1968

Type species

Actinostroma bifarium Nicholson, Reference Nicholson1886b.

Bifariostroma bifarium (Nicholson, Reference Nicholson1886b)

Figure 8.4–8.6

Reference Nicholson1886b    Actinostroma bifarium Nicholson, p. 231, pl. 6, figs. 4, 5.

Reference Nicholson1889    Actinostroma bifarium; Nicholson, p. 136, pl. 13, figs. 3–7.

Reference Kaźmierczak1971    Actinostroma bifarium; Kaźmierczak, p. 135, pl. 38, figs. 2, 3; pl. 39, figs. 1, 2.

Reference Zukalová1971    Actinostroma cf. bifarium; Zukalová, p. 38, pl. 6, figs. 3–5.

Reference Krebedünkel1995    Actinostroma bifarium; Krebedünkel, p. 32, pl. 2, figs. 1, 2.

Reference Stearn, Webby, Nestor and Stock1999    Bifariostroma bifarium; Stock in Stearn et al., p. 33.

Reference Stock and Selden2015    Bifariostroma bifarium; Stock, p. E770, fig. 426.

Reference Wolniewicz2016    Bifariostroma bifarium; Wolniewicz, p. 240, figs. 10D, 11A, B.

Holotype

Specimen (NHMUK PI P5639) from the Middle Devonian of Büchel, Germany (Nicholson, Reference Nicholson1886b, pl. 6, figs. 4, 5).

Description

External forms appear to be laminar but were prepared as fragments. The skeleton is composed of regular networks. In longitudinal section, pillars dominate, encompassing two types, some vertically extensive, thick, spacing 4–5 per 2 mm, 0.2–0.5 mm in width (Fig. 8.4). Others are short, post-like or spool-shaped, commonly confined to one interlaminar space, locally superposed across laminae, spacing about 5–7 per 2 mm and 0.05–0.15 mm in width. Colliculi are thin, spacing about 6–8 per 2 mm and 0.05–0.15 mm in thickness. Galleries are rectangular. In tangential section, two types of pillars are round in the interlaminar spaces, but show radiating colliculi across spaces, forming complex interconnected networks (Fig. 8.5, 8.6). Microstructure is compact.

Material

Five specimens from the Middle Devonian of Teignmouth (NHMUK PI P5636, P5641, P5642), in South Devon (UK) and Büchel (NHMUK PI P5639, P5645), in Germany.

Remarks

This species was previously identified as Actinostroma bifarium, but it shows a high resemblance to Bifariostroma in possessing two types of radiating pillars, which is different from the one type of radiating pillar of Actinostroma. Therefore, it was revised to Bifariostroma (Stock, Reference Stock and Selden2015; Wolniewicz, Reference Wolniewicz2016).

Order Stromatoporellida Stearn, Reference Stearn1980

Family Stromatoporellidae Lecompte, Reference Lecompte1951

Genus Stromatoporella Nicholson, Reference Nicholson1886a

Type species

Stromatopora granulata Nicholson, Reference Nicholson1873.

Stromatoporella arachnoidea Nicholson, Reference Nicholson1886b

Figure 9.1, 9.2

Figure 9.

Longitudinal (1) and tangential (2) views of Stromatoporella arachnoidea Nicholson, Reference Nicholson1886b , from the Middle Devonian of Büchel, Germany, holotype NHMUK PI P6063. Longitudinal (3) and tangential (4) views of Stromatoporella solitaria Nicholson, Reference Nicholson1892, from the Middle Devonian of Gerolstein, Germany, NHMUK PI P6037. Longitudinal (5) and tangential (6) views of Clathrocoilona curiosa (Bargatzky, Reference Bargatzky1881a) from the Middle Devonian of Büchel, Germany, NHMUK PI P6051.

Reference Nicholson1886b    Stromatoporella arachnoidea Nicholson, p. 237, pl. 8, figs. 1, 2.

Holotype

Specimen (NHMUK PI P6063) from the Middle Devonian of Büchel, Germany (Nicholson, Reference Nicholson1886b, pl. 8, figs. 1, 2).

Description

External forms are laminar. The skeleton consists of short pillars and laterally long laminae. In longitudinal section, pillars are short, spool-shaped, confined to interlaminar spaces, not superposed, spacing about 6–7 per 2 mm and 0.1–0.5 mm in width (Fig. 9.1). Laminae are thin, horizontally continuous, slightly undulating, spacing about 8–9 per 2 mm and 0.05–0.1 mm in thickness. Galleries are round or laterally elongate, with abundant oblique dissepiments. In tangential section, pillars are commonly isolated or irregular. Ring pillars rarely occur (Fig. 9.2). Laminae are undulating or serve as concentric circles in the mamelons. Microstructure is fibrous or cellular.

Material

Two specimens from the Middle Devonian of Büchel (NHMUK PI P6063) and Gerolstein (NHMUK PI P6064), in Germany.

Remarks

The rare occurrence of ring pillars in Stromatoporella arachnoidea derives from upward branching pillars, and it is different from typical ring pillars of S. granulata, which are derived from upwardly inflected laminae.

Stromatoporella solitaria Nicholson, Reference Nicholson1892

Figure 9.3, 9.4

Reference Nicholson1892   Stromatoporella solitaria Nicholson, p. 210, pl. 7, fig. 4; pl. 27, figs. 4–7; pl. 2, figs. 9, 10.

Reference Lecompte1951     Stromatoporella solitaria; Lecompte, p. 173, pl. 23, figs. 6, 7.

Reference May1993    Stromatoporella cf. solitaria; May, p. 33, pl. 5, fig. 2.

Holotype

Specimen (NHMUK PI P6035) from the Middle Devonian of Gerolstein, Germany (Nicholson, Reference Nicholson1892, pl. 27, figs. 4–7). It appears that although a lectotype was erected by May (Reference May1993), the existence of a holotype has since been confirmed. Therefore, the lectotype designation is no longer necessary and can be set aside.

Description

Growth forms of illustrated specimens are fragmented, but Nicholson (Reference Nicholson1892) noted the laminar shapes. The skeleton consists of regular networks formed by short pillars and laterally long laminae. In longitudinal section, pillars are short, spool-shaped, inflecting upward and downward to laminae, with central tubes, confined to interlaminar spaces, not superposed, spacing about 5–7 per 2 mm and 0.05–0.3 mm in width (Fig. 9.3). Laminae are thin, locally interrupted, slightly undulating, tripartite with a central light zone and surrounding dark zones, spacing about 6–7 per 2 mm and 0.1–0.2 mm in thickness. Galleries are round or laterally elongate, with sparse oblique dissepiments. Mamelons develop, 4 mm in height. In tangential section, pillars are commonly isolated or irregular. Ring pillars are abundant (Fig. 9.4). Laminae are undulating or concentric around mamelons. Microstructure is ordinicellular, tripartite, or cellular.

Material

Ten specimens from the Middle Devonian of Gerolstein (NHMUK PI P6034, P6035, P6036, P6037, P6038, P6039, P6040, P6047, P6048), in Germany and Teignmouth (NHMUK PI P6041), in South Devon (UK).

Remarks

Abundant ring pillars in Stromatoporella solitaria are formed by spool-shaped hollow pillars, which is different from ring pillars of S. granulata by upward inflected laminae.

Genus Clathrocoilona Yavorsky, Reference Yavorsky1931

Type species

Clathrocoilona abeona Yavorsky, Reference Yavorsky1931.

Clathrocoilona curiosa (Bargatzky, Reference Bargatzky1881a)

Figure 9.5, 9.6

Reference Bargatzky1881a    Stromatopora curiosa Bargatzky, p. 285.

Reference Nicholson1886c    Stromatoporella curiosa; Nicholson, p. 8, pl. 1, figs. 1–3.

Reference Nicholson1892    Stromatoporella curiosa; Nicholson, p. 213, pl. 28, figs. 1–3.

Reference May1993    Clathrocoilona (Clathrocoilona) curiosa; May, p. 40, pl. 6, fig. 1.

Reference Krebedünkel1995    Stromatoporella curiosa; Krebedünkel, p. 93, pl. 12, figs. 5, 6.

Reference Avlar and May1997    Clathrocoilona (Clathrocoilona) curiosa; Avlar and May, p. 109, pl. 1, fig. 2.

Reference Salerno2008    Clathrocoilona curiosa; Salerno, p. 66, pl. 10, fig. 1.

Holotype

Specimen from the Middle Devonian of Eifel, Germany (Bargatzky, Reference Bargatzky1881a, p. 285).

Description

Growth forms are thin laminar, locally encrusting solitary rugose corals. The skeleton is composed of regular networks formed by short pillars and extensive laminae. In longitudinal section, pillars are post-like, confined to interlaminar spaces, rarely superposed, spacing about 5–8 per 2 mm and 0.1–0.2 mm in width (Fig. 9.5). Laminae are thin, long, tripartite with a central light line and surrounding dark zones, spacing about 5–7 per 2 mm and 0.1–0.2 mm in thickness. Galleries are subrectangular, round, or laterally elongated. In tangential section, pillars are round, and isolated (Fig. 9.6). Laminae are undulating or concentric. Mamelons locally occur. Microstructure is tripartite or ordinicellular.

Material

Four specimens from the Middle Devonian of Büchel (NHMUK PI P6051, P6052, P6094) and Gerolstein (NHMUK PI P6054), in Germany.

Remarks

This species was originally assigned to Stromatoporella by Nicholson (Reference Nicholson1886b), but no ring pillars are observed. The thin, encrusting skeletons and tripartite laminae, however, closely resemble Clathrocoilona.

Clathrocoilona spissa (Lecompte, Reference Lecompte1951)

Figure 10.1, 10.2

Figure 10.

Longitudinal (1) and tangential (2) sections of Clathrocoilona spissa (Lecompte, Reference Lecompte1951) from the Middle Devonian of Gerolstein, Germany, NHMUK PI P5735. Transverse (longitudinal) (3) and tangential (4) sections of Dendrostroma oculatum (Nicholson, Reference Nicholson1886a) from the Middle Devonian of Büchel, Germany, holotype NHMUK PI P6073 (this sample is the one used by Stearn, Reference Stearn and Selden2015b, fig. 4.36).

Reference Lecompte1951    Stromatoporella spissa Lecompte, p. 187, pl. 27, figs. 1–4.

Reference Kaźmierczak1971    Stromatopora spissa; Kaźmierczak, p. 92, pl. 21, fig. 2.

Reference Zukalová1971    Clathrocoilona spissa; Zukalová, p. 56, pl. 15, figs. 1, 2.

Reference Flügel1974    Clathrocoilona spissa; Flügel, p. 165, pls. 24, 26, 27.

Reference Mistiaen1980    Clathrocoilona spissa; Mistiaen, p. 196, pl. 7, figs. 3–9.

Reference Cockbain1984    Clathrocoilona spissa; Cockbain, p. 25, pl. 11.

Reference Mistiaen1985    Clathrocoilona spissa; Mistiaen, p. 96–101, pl. 6, figs. 6–8.

Reference Mistiaen and Brice1988    Clathrocoilona spissa; Mistiaen, p. 174.

Reference May1993    Clathrocoilona (Clathrocoilona) solidula spissa; May, p. 38, pl. 7, fig. 2; pl. 8, figs. 1, 2.

Reference May1999c    Clathrocoilona (Clathrocoilona) solidula spissa; May, p. 127.

Reference May2005    Clathrocoilona (Clathrocoilona) solidula spissa; May, p. 180, pl. 13, figs. 2, 3.

Reference Salerno2008    Clathrocoilona spissa; Salerno, p. 74, pl. 11, fig. 2.

Reference Huang, Liang, Wang, Kershaw, Jeon, Li and Qie2022    Clathrocoilona spissa; Huang et al., p. 718, fig. 6.

Holotype

Specimen (7164) from the Middle Devonian of Dinant Basin, Belgium (Lecompte, Reference Lecompte1951, pl. 27, fig. 1).

Description

Growth forms are uncertain due to fragmented specimens. The skeleton consists of densely packed thick pillars and laminae. In longitudinal section, skeletons are varied. In some areas, laminae and pillars can hardly be differentiated. Pillars and laminae are thick, and irregularly amalgamated. Gallery shapes are irregular, locally with abundant dissepiments. In the other areas, skeletons comprise regular laminae and pillars (Fig. 10.1). Pillars are spool-shaped, confined to interlaminar spaces, spacing 4–5 per 2 mm, 0.1–1 mm in width. Laminae are thick, locally with dark axial lines and the coated light zones, spacing 4 per 2 mm, 0.2–0.5 mm in thickness. In tangential section, skeletons are amalgamated, with local pillars round (Fig. 10.2), subtriangular, or irregular, isolated, 0.1–1 mm in diameter. Astrorhizal canals are common, 0.1–0.2 mm in width, with sparse dissepiments. Microstructure is compact or tripartite.

Material

Four specimens from Middle Devonian of Büchel (NHMUK PI P6053), Gerolstein (NHMUK PI P5735), and Sötenich (NHMUK PI P5736, P6091) in Germany.

Remarks

Clathrocoilona spissa is different from C. curiosa in having irregular skeletons formed by densely packed, thick pillars and laminae. It is revised from specimens previously assigned to Stromatoporella damnoniensis and S. curiosa by Nicholson (Reference Nicholson1886b, Reference Nicholson1892).

Genus Dendrostroma Lecompte, Reference Lecompte1952

Type species

Idiostroma oculatum Nicholson, Reference Nicholson1886a.

Dendrostroma oculatum (Nicholson, Reference Nicholson1886a)

Figure 10.3, 10.4

Reference Nicholson1886a    Idiostroma oculatum Nicholson, p. 101, figs. 14, 15.

Reference Nicholson1892    Idiostroma oculatum; Nicholson, p. 225, pl. 29, figs. 10, 11.

Reference Lecompte1952    Dendrostroma oculatum; Lecompte, p. 320, pl. 61, fig. 1.

Reference Stearn and Selden2015b    Dendrostroma oculatum; Stearn, p. E784, fig. 436a, b.

Holotype

Specimen (NHMUK PI P6073) from the Middle Devonian of Büchel, Germany (Nicholson, Reference Nicholson1886a, figs. 14, 15).

Description

Growth forms are dendroid, 7–10 mm in diameter. In longitudinal section, the skeleton consists of short pillars and arched laminae. In the central zone, skeletons are featured by an axial tube, 0.5–0.8 mm in diameter. In the peripheral zone, skeletons are well differentiated (Fig. 10.3). Pillars are short, post-like or spool-shaped, confined to interlaminar spaces, not superposed, spacing 4–5 per 2 mm and 0.1–0.5 mm in width. Laminae are laterally extensive, thick, slightly wavy, locally tripartite with central light line and surrounding dark zones, spacing 8–9 per 2 mm and 0.1–0.3 mm in thickness. Galleries are elongate. Tabulate coral symbionts are common, 0.5 mm in width. In tangential section, networks are formed by short pillars and concentric laminae. The central zone presents an axial tube, with coral symbionts spread outward. In oblique section, pillars are isolated (Fig. 10.4). The microstructure is compact to fibrous.

Material

Two specimens from the Middle Devonian of Büchel (NHMUK PI P6073, H4777) in Germany and one from the Devonian of Dartington (NHMUK PI P6115) in South Devon (UK).

Remarks

This species was previously assigned to Idiostroma by Nicholson (Reference Nicholson1886a, Reference Nicholson1892), but the occurrence of isolated and confined pillars is distinguished by vermiform and superposed pachysteles of Idiostroma.

Genus Stictostroma Parks, Reference Parks1936

Type species

Stictostroma gorriense Stearn, Reference Stearn1995.

Stictostroma clarum (Počta, Reference Počta1894)

Figure 11.1, 11.2

Figure 11.

Longitudinal (1) and tangential (2) views of Stictostroma clarum (Počta,1894) from the Lower Devonian of Commonwealth Quarries, Buchan, Victoria, Australia, NHMUK PI H3912. Longitudinal (3) and tangential (4) views of Stictostroma damnoniensis (Nicholson, Reference Nicholson1886b) from the Middle Devonian of Teignmouth, South Devon, UK, holotype NHMUK PI P6042.

Reference Počta1894    Clathrodictyon clarum Počta, p. 152, pl. 18, figs. 7, 8.

Reference Ripper1937a    Clathrodictyon clarum; Ripper, p. 21, pl. 4, figs. 3, 4.

Reference Webby, Stearn and Zhen1993    Petridiostroma clarum; Webby, Stearn, and Zhen, p. 132, figs. 10A–F, 31A.

Reference May1999a    Stictostroma clarum; May, p. 174.

Reference May2005    Stictostroma clarum; May, p. 176, pl. 40, fig. 1; pl. 43, fig. 1; pl. 11, fig. 2.

Holotype

Specimen from the Middle Devonian of the Prague Basin, Czech Republic (Počta, Reference Počta1894, pl. 18, figs. 7, 8).

Description

External form is unclear based on fragmented skeletons. The skeleton consists of laterally extensive laminae and short pillars. In longitudinal section, pillars are short, post-like, generally confined to one interlaminar space, rarely superposed (Fig. 11.1), spacing about 6–8 per 2 mm and 0.02–0.1 mm in width. Laminae are thin, planar, laterally continuous, spacing about 8–10 per 2 mm and 0.02–0.1 mm in thickness. Galleries are laterally elongate or circular. In tangential section, pillars and laminae are isolated dots and undulating (Fig. 11.2), respectively. Syringoporid tabulate corals are 0.3–0.4 mm in width, spacing 3–4 per 2 mm. Coarse ordinicellular structure is observed.

Material

Two specimens (NHMUK PI H3912, H3926) from Lower Devonian of Commonwealth Quarries, Buchan, Victoria (Australia).

Remarks

This species was originally grouped with Clathrodictyon and Petridiostroma. May (Reference May1999a, Reference May2005) redescribed the type material and reassigned it into Stictostroma; this study follows this decision.

Stictostroma damnoniensis (Nicholson, Reference Nicholson1886b)

Figure 11.3, 11.4

Reference Nicholson1886b    Stromatoporella damnoniensis Nicholson, p. 237, pl. 8, figs. 3, 4.

Reference Nicholson1892    Stromatoporella damnoniensis; Nicholson, p. 207, pl. 27, figs. 8, 9.

Holotype

Specimen (NHMUK PI P6042) from the Middle Devonian of Teignmouth in South Devon, UK (Nicholson, Reference Nicholson1886b, pl. 8, figs. 3, 4).

Description

External form is unclear, but Nicholson (Reference Nicholson1886b) noted domical or bulbous shapes. The skeleton is composed of networks formed by short pillars and intermittent laminae. In longitudinal section, pillars are post-like or spool-shaped, confined to interlaminar spaces, not superposed, spacing about 4–6 per 2 mm and 0.1–0.4 mm in width. Laminae are long but interrupted, spacing about 6–8 per 2 mm and 0.1–0.2 mm in thickness (Fig. 11.3). Galleries are round, irregular, or laterally elongated. In tangential section, pillars are round, isolated (Fig. 11.4). Laminae are undulating or concentric, with circular holes inside. Mamelons locally occur, 8 mm in width. Microstructure is fibrous and coarsely ordinicellular.

Material

One specimen from the Middle Devonian of Teignmouth (NHMUK PI P6042) in South Devon (UK).

Remarks

This specimen was assigned to Stromatoporella due to the occurrence of possible ring pillars, but these circular structures are parts of laminae. Lecompte (Reference Lecompte1951) revised the paratype of this species into Clathrocoilona crassitexta (Lecompte, Reference Lecompte1951). The holotype from UK, however, shows more relationships with Stictostroma due to coarsely ordinicellular microstructure.

Stictostroma saginatum (Lecompte, Reference Lecompte1951)

Figure 12.1, 12.2

Figure 12.

Longitudinal (1) and tangential (2) views of Stictostroma saginatum (Lecompte, Reference Lecompte1951) from the Middle Devonian of Teignmouth, South Devon, UK, NHMUK PI P5699. Longitudinal (3) and tangential (4) views of Stictostroma socialis (Nicholson, Reference Nicholson1892) from the Middle Devonian of Gerolstein, Germany, NHMUK PI P6095; note the ordinicellular microstructure indicated by the yellow arrow in (3). Longitudinal (5) and tangential (6) views of Tubuliporella calamosa (Ripper, Reference Ripper1933) from the Lower Devonian of Mitchell’s Quarry, Cave Hill, Lilydale, Australia, holotype NHMUK PI H3898.

Reference Lecompte1951    Stromatoporella saginata Lecompte, p. 171, pl. 22, figs. 5–7; pl. 23, figs. 1–3.

Reference Stearn1983    Clathrocoilona cf. saginata; Stearn, p. 549, fig. 5G–H.

Reference Cockbain1984    Clathrocoilona saginata; Cockbain, p. 25, pl. 10, figs. A–D.

Reference Mistiaen1985    Stictostroma saginatum; Mistiaen, p. 115, pl. 8, figs. 6–11.

Reference Mistiaen and Brice1988    Stictostroma saginatum; Mistiaen, p. 176, pl. 21, figs. 5–9.

Reference Mistiaen1999    Stictostroma saginatum; Mistiaen, p. 36, pl. 3, figs. 9–12; pl. 4, figs. 1, 2.

Reference Mistiaen and Gholamalian2000    Stictostroma saginatum; Mistiaen and Gholamalian, p. 83–84, pl. 6, figs. 6–8.

Reference Salerno2008    Stictostroma saginatum; Salerno, p. 82–84, pl. 12, fig. 2.

Reference Huang, Liang, Wang, Kershaw, Jeon, Li and Qie2022    Stictostroma saginatum; Huang et al., p. 720, fig. 7A, B.

Holotype

Specimen (7502) from the Upper Devonian of Dinant Basin, Belgium (Lecompte, Reference Lecompte1951, pl. 23, fig. 1).

Description

Growth forms are unclear due to limited preservation of specimen. The skeleton consists of short pillars and undulating laminae. In longitudinal section, pillars are spool-shaped, confined to interlaminar spaces, locally superposed, spacing about 7–8 in 2 mm, 0.1–0.2 mm in width (Fig. 12.1). Laminae are laterally extensive, undulating especially around mamelons, spacing 8–10 per 2 mm, 0.05–0.2 mm in thickness. Galleries are circular, or laterally elongated. Mamelons are conspicuous, generally 4 mm in height, with central and branching astrorhizal canals. In tangential section, pillars are round, isolated. Laminae are undulating or present as concentric circles (Fig. 12.2). Mamelons are apparent, with astrorhizal canals, laterally separated by distances of 2–4 mm. The microstructures are compact or ordinicellular.

Material

Four specimens from the Middle Devonian of Teignmouth (NHMUK PI P5482, P5699, P5874, P5949), in South Devon (UK).

Remarks

This specimen was assigned to Stromatoporella socialis, but ring pillars have not been observed. Lecompte (Reference Lecompte1951) revised this species into Stromatoporella saginata, which is widely accepted as Stictostroma saginatum. We therefore reassign it to Stictostroma.

Stictostroma socialis (Nicholson, Reference Nicholson1892)

Figure 12.3, 12.4

Reference Nicholson1892 Stromatoporella socialis Nicholson, p. 206, pl. 26, figs. 5–7.

Reference Lecompte1951    Stromatoporella socialis; Lecompte, p. 163, pl. 21, figs. 2, 3.

Holotype

Specimen (NHMUK PI P6043) from the Middle Devonian of Teignmouth in South Devon, UK (Nicholson, Reference Nicholson1892, pl. 26, figs. 5–7).

Description

External forms appear to be laminar, 1–2 cm thick. The skeleton consists of short pillars and long laminae. In longitudinal section, pillars are spool-shaped, confined to interlaminar spaces, not superposed, spacing 5–9 in 2 mm, 0.1–0.3 mm in width. Laminae are laterally extensive, slightly undulating, tripartite with central hollow cavities (Fig. 12.3), spacing 12–14 per 2 mm, 0.05–0.2 mm in thickness. Galleries are circular, or laterally elongated. Syringoporid tabulate corals are observed, 0.5–1 mm in width. In tangential section, pillars are round, isolated (Fig. 12.4). The microstructure is ordinicellular.

Material

Fourteen specimens from the Middle Devonian of Teignmouth (NHMUK PI P5696, P5700, P5701, P5703, P6043, P6044, P6045, P6046), Dartington (NHMUK PI P5702), in South Devon (UK); and Gerolstein (NHMUK PI P6049, P6050, P6092, P6093, P6095), in Germany.

Remarks

The current species was identified as Stromatoporella by Nicholson (Reference Nicholson1892), but the ordinicellular microstructure and absence of ring pillars justify its assignment to Stictostroma.

Genus Tubuliporella Khalfina, Reference Khalfina and Ivaniya1968

Type species

Tubuliporella lecompti Khalfina, Reference Khalfina and Ivaniya1968.

Tubuliporella calamosa (Ripper, Reference Ripper1933)

Figure 12.5, 12.6

Reference Ripper1933    Clathrodictyon calamosum Ripper, p. 160, fig. 6E, F.

Reference Flügel and Flügel-Kahler1968    Clathrodictyon calamosum; Flügel and Flügel-Kahler, p. 56.

Reference Webby, Stearn and Zhen1993    Tubuliporella calamosa; Webby, Stearn, and Zhen, p. 148, figs. 16F, 17A–D.

Holotype

Specimen (NHMUK PI H3898) from the Lower Devonian of Lilydale Limestone, Mitchell’s, Cave Hill, Lilydale, Victoria, Australia (Ripper, Reference Ripper1933, fig. 6E, F).

Description

Growth forms are unclear due to poorly preserved specimens. The skeleton commonly shows networks formed by short pillars and laterally extensive laminae. In longitudinal section, skeleton is dominated by ring pillars (Fig. 12.5). Ring pillars are locally superposed, forming vertical channels filled by thin laminae or dissepiments, 0.2–0.4 mm in width. Laminae are weakly developed, thin, spacing about 7–9 per 2 mm and 0.05–0.2 mm in thickness. Galleries are round. The tangential section presents ring pillars (Fig. 12.6), in diameter of 0.1–0.2 mm. Laminae are inconspicuous. Microstructure is compact.

Material

One specimen from the Lower Devonian of Mitchell’s Quarry (NHMUK PI H3898), Cave Hill, Lilydale, Victoria (Australia).

Remarks

The current species was assigned to Clathrodictyon by Ripper (Reference Ripper1933) and Flügel and Flügel-Kahler (Reference Flügel and Flügel-Kahler1968), but the superposed ring pillars justify its assignment as Tubuliporella (Webby et al., Reference Webby, Stearn and Zhen1993). It resembles Stromatoporella, but differs from the latter genus in irregular skeletons, superposed ring pillars, and compact microstructure.

Family Trupetostromatidae Germovsek, Reference Germovsek1954

Genus Trupetostroma Parks, Reference Parks1936

Type species

Trupetostroma warreni Parks, Reference Parks1936.

Trupetostroma ? dartingtonense (Carter, Reference Carter1880)

Figure 13.1, 13.2

Figure 13.

Longitudinal (1) and tangential (2) sections of Trupetostroma? dartingtonense (Carter, Reference Carter1880) from the Middle Devonian of Dartington, South Devon, UK, NHMUK PI P5746. Longitudinal (3) and tangential (4) sections of Trupetostroma thomasi Lecompte, Reference Lecompte1952, from the Middle Devonian of Hebborn, Germany, holotype NHMUK PI P5934. Longitudinal (5) and tangential (6) sections of Hermatostroma episcopale Nicholson, Reference Nicholson1892, from the Middle Devonian of Bishopsteignton, South Devon, UK, holotype NHMUK PI P6065.

Reference Carter1880    Stromatopora dartingtonensis Carter, p. 346, pl. 18, figs. 1–5.

Reference Nicholson1891a    Parallelopora dartingtonensis; Nicholson, p. 199, pl. 4, fig. 1; pl. 24, figs. 13–15.

Reference Lecompte1952    Parallelopora cf. dartingtonensis; Lecompte, p. 295, pl. 49, fig. 3.

Holotype

Specimen from the Middle Devonian of Pit-Park Quarry, Dartington, South Devon, UK (Carter, Reference Carter1880, pl. 18, figs. 1–5).

Description

Nicholson (Reference Nicholson1891a) noted bulbous, domical, or laminar external forms. The skeleton shows regular networks formed by long pillars and laminae. In longitudinal section, pillars are post-like, vertically continuous, passing through dozens of laminae (Fig. 13.1), spacing 8–9 per 2 mm and 0.1–0.2 mm in width. Laminae are laterally long, tripartite with a central dark line and surrounding light zones, spacing about 7–8 per 2 mm and 0.05–0.2 mm in thickness. Galleries are circular, with abundant dissepiments. In tangential section, pillars are circular, isolated, locally connected (Fig. 13.2). Astrorhizal canals are sparse. Microstructure is compact.

Material

Fifteen specimens from the Middle Devonian of Dartington (NHMUK PI P5737, P5740, P5746, P5753) and Teignmouth (NHMUK PI P5738, P5739, P5741, P5742, P5743, P5744, P5748, P5750, P5752, P5923, P5937), in South Devon (UK).

Remarks

This species was assigned to Parallelopora due to continuous and local interconnected pillars, but it also resembles Trupetostroma in regular networks formed by locally extensive, isolated pillars and tripartite laminae. Due to limited illustrations of previous specimens, this study tentatively revises this species as Trupetostroma.

Trupetostroma thomasi Lecompte, Reference Lecompte1952

Figure 13.3, 13.4

Reference Nicholson1891a    Parallelopora capitata; Nicholson, p. 197, pl. 25, figs. 10–13.

Reference Lecompte1952    Trupetostroma thomasi; Lecompte, p. 240.

Holotype

Specimen (NHMUK PI P5934) from the Middle Devonian of Hebborn (Paffrath district), Germany (Nicholson, Reference Nicholson1891a, fig. 27A, B).

Description

Nicholson (Reference Nicholson1891a) illustrated external skeleton as bulbous form. The skeleton presents regular networks formed by short pillars and thin laminae. In longitudinal section, pillars are spool-shaped, confined to interlaminar spaces but systematically superposed, spacing 3–5 per 2 mm and 0.1–0.5 mm in width (Fig. 13.3). Laminae are slightly undulating, laterally extensive, tripartite with a central dark zone and surrounding light zones, spacing about 5–7 per 2 mm and 0.1–0.2 mm in thickness. Galleries are circular, or laterally elongated. Syringoporid intergrowth rarely develops. Astrorhizal canals are locally abundant, with a width of 10–20 mm. In tangential section, skeletons are densely packed. Pillars are circular, isolated. Laminae are undulating (Fig. 13.4). Microstructure is vacuolate, cellular, or compact.

Material

Fifteen specimens from the Middle Devonian of Teignmouth (NHMUK PI P5520, P5522, P5523, P5556, P5893, P5908, P5935, P6080) in South Devon (UK); Hebborn (NHMUK PI P5524, P5526, P5546, P5934, P6081, P6100) and Steinbreche (NHMUK PI P5549), in Germany.

Remarks

This species was originally assigned into Parallelopora by Nicholson (Reference Nicholson1891a), but the regular skeletons differ from the dominant interconnected pachysteles and orthoreticular microstructure in Parallelopora. It was revised into Trupetostroma thomasi by Lecompte (Reference Lecompte1952); we follow this decision.

Genus Hermatostroma Nicholson, Reference Nicholson1886a

Type species

Hermatostroma schlueteri Nicholson, Reference Nicholson1886a.

Hermatostroma episcopale Nicholson, Reference Nicholson1892

Figure 13.5, 13.6

Reference Nicholson1892    Hermatostroma episcopale Nicholson, p. 219, pl. 28, figs. 4–11.

Reference Lecompte1952    Hermatostroma episcopale; Lecompte, p. 216, pl. 48. fig. 4; pl. 49, figs. 1, 2.

Reference Galloway1960    Hermatostroma episcopale; Galloway, p. 635, pl. 77, fig. 4.

Reference Yang and Dong1963    Hermatostroma episcopale; Yang and Dong, p. 162, pl. 10, figs. 3–6.

Reference Yang and Dong1979    Hermatostroma episcopale; Yang and Dong, p. 68, pl. 32, figs. 7, 8.

Reference Dong and Wang1982    Hermatostroma episcopale; Dong and Wang, p. 23, pl. 14, figs. 1, 2.

Holotype

Specimen (NHMUK PI P6065) from the Middle Devonian of Bishopsteignton, South Devon, UK (Nicholson, Reference Nicholson1892, pl. 28, figs. 5, 6).

Description

Growth forms are uncertain due to limited preservation, but Nicholson (Reference Nicholson1892) noted bulbous/domical shapes. The skeleton shows regular networks formed by short pillars and continuous laminae. In longitudinal section, pillars are spool-shaped, confined to one interlaminar space, superposed across several laminae (Fig. 13.5), spacing about 3–4 per 2 mm and 0.1–0.3 mm in width. Laminae are gently wavy, extensive, tripartite with a central dark zone and surrounding light zones, spacing about 4–6 per 2 mm and 0.2–0.3 mm in thickness. Galleries are circular, 0.1–0.5 mm in width. Syringoporid tabulate corals occur, with a width of 1 mm. Dissepiments are common, oblique. In tangential section, pillars are circular, subcircular, isolated or laterally connected. Laminae are circular or undulating. Both pillars and laminae show central dark zone and peripheral light zones (Fig. 13.6). Astrorhizae are well developed, 3–4 mm in width, 5 mm from one to another. Microstructure is compact.

Material

Six specimens from the Middle Devonian of Teignmouth (NHMUK PI P5686, P5688, P5689, P6086) and Bishopsteignton (NHMUK PI P5692, P6065), in South Devon, (UK).

Remarks

This species differs from H. schlueteri in short pillars, common dissepiments, and well-developed astrorhizae.

Hermatostroma schlueteri Nicholson, Reference Nicholson1886a

Figure 14.1, 14.2

Figure 14.

Longitudinal (1) and tangential (2) views of Hermatostroma schlueteri Nicholson, Reference Nicholson1886a, from the Middle Devonian of Hebborn, Germany, holotype NHMUK PI P5527. Longitudinal (3, 4) and tangential (5, 6) views of Hermatostromella holmesae (Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993) from Lower Devonian of Mitchell’s Quarry, Cave Hill, Lilydale, Victoria, Australia, NHMUK PI H3896.

Reference Nicholson1886a    Hermatostroma schlueteri Nicholson, p. 105, pl. 3, figs. 1, 2.

Reference Nicholson1892    Hermatostroma schlueteri; Nicholson, p. 215, pl. 28, figs. 12, 13.

Reference Lecompte1952    Hermatostroma schlueteri; Lecompte, p. 250, pl. 45, fig. 1.

Reference Galloway1957    Hermatostroma schlueteri; Galloway and St. Jean, p. 218, pl. 21, fig. 1.

Reference Yang and Dong1979    Hermatostroma schlueteri; Yang and Dong, p. 69, pl. 37, figs. 1, 2.

Reference Dong and Wang1982    Hermatostroma schlueteri; Dong and Wang, p. 23, pl. 13, figs. 7, 8.

Reference Cockbain1984    Hermatostroma schlueteri; Cockbain, p. 27, fig. 16A–D.

Reference Mistiaen1985    Hermatostroma schlueteri; Mistiaen, p. 175, pl. 15, figs. 9–11; pl. 16, fig. 1.

Reference Dong, Wang, Zhou, Zhang, Luo, Fu and Huang1989    Hermatostroma schlueteri; Dong et al., p. 267, pl. 8, fig. 3a, b.

Reference Krebedünkel1995    Hermatostroma schlueteri; Krebedünkel, p. 65, pl. 7, figs. 7, 8.

Reference Salerno2008    Hermatostroma schlueteri; Salerno, p. 84, pl. 13, figs. 1, 2.

Holotype

Specimen (NHMUK PI P5527) from the Middle Devonian of Hebborn, Germany (Nicholson, Reference Nicholson1886a, pl. 3, figs. 1, 2).

Description

External forms are uncertain due to poor preservation, but Nicholson (Reference Nicholson1892) noted bulbous/domical shapes. The skeleton shows regular networks formed by continuous pillars and laminae. In longitudinal section, pillars are long, spool-shaped, confined to one interlaminar space but regularly superposed across dozens of laminae (Fig. 14.1), spacing 6–7 per 2 mm and 0.2–0.3 mm in width. Laminae are gently wavy, laterally extensive, spacing about 5–6 per 2 mm and 0.1–0.2 mm in thickness. Both pillars and laminae are tripartite with a central dark zone and surrounding light zones. Galleries are circular, 0.2–1 mm in width. Tubeworms locally occur, with a width of 0.5–0.8 mm. In tangential section, pillars are circular or subcircular, isolated, with central dark dots and peripheral light zones or vesicles (Fig. 14.2). Laminae are circular or undulating. Microstructure is compact and vacuolate.

Material

One specimen from the Middle Devonian of Hebborn (NHMUK PI P5527) in Germany.

Remarks

Hermatostroma schlueteri is distinguished from H. episcopale in regular networks formed by long pillars and laminae, absence of astrorhizae, and well-developed tripartite skeletons.

Genus Hermatostromella Khalfina, Reference Khalfina and Halfin1961

Type species

Hermatostromella parasitica Khalfina, Reference Khalfina and Halfin1961.

Hermatostromella holmesae (Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993)

Figure 14.3–14.6

Reference Ripper1937c    Syringostroma aff. ristigouchense; Ripper, p. 181, pl. 8, figs. 1, 2.

Reference Webby, Stearn and Zhen1993    Amnestostroma holmesae; Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993, p. 151, figs. 19A–E, 31C, D.

Reference May2002    Hermatostromella holmesae; May, p. 129, pl. 2, fig. 6.

Reference May2005    Hermatostromella holmesae; May, p. 190, pl. 18, figs. 1, 2.

Holotype

Longitudinal and tangential sections (NMV P141860, P141861, P141862, and P141863) from the Lower Devonian of Lilydale Limestone, Mitchell’s (Cave Hill) Quarry, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 19A–D).

Description

The skeleton is composed of the regular network formed by pillars and laminae. In longitudinal section, pillars are thick, spool-shaped, confined to interlaminar spaces, but systematically superposed (Fig. 14.3), spacing 7–8 per 2 mm and 0.1–0.2 mm in width. Laminae are undulating, tripartite with central dark line (Fig. 14.4), spacing 10–13 per 2 mm and 0.02–0.1 mm in thickness. Galleries are circular. Syringoporid symbionts existed. In tangential section, pillars are circular and isolated (Fig. 14.5, 14.6). Laminae appear as concentric circles (Fig. 14.5). Microstructure is compact.

Material

One specimen (NHMUK PI H3896) from Lower Devonian of Mitchell’s Quarry, Cave Hill, in Lilydale, Victoria (Australia).

Remarks

This species was originally grouped with Amnestostroma, which is regarded as a junior synonym of Hermatostromella. This study therefore reassigned the present species in the latter genus.

Family Idiostromatidae Nicholson, Reference Nicholson1886a

Genus Idiostroma Winchell, Reference Winchell1867

Type species

Stromatopora caespitosa Winchell, Reference Winchell1866.

Idiostroma roemeri Nicholson, Reference Nicholson1886a

Figure 15.1–15.4

Figure 15.

Longitudinal (1, 2, 4) and tangential (3) sections of Idiostroma roemeri Nicholson, Reference Nicholson1886a, from the Middle Devonian of Hebborn, Germany, holotype NHMUK PI P6076. Longitudinal (5) and tangential (6) sections of Stromatopora aff. S. polaris (Stearn, Reference Stearn1983) from the Lower Devonian of Buchan, Victoria, Australia, NHMUK PI H3905.

Reference Nicholson1886a    Idiostroma roemeri Nicholson, p. 99, pl. 9, figs. 6–11.

Reference Lecompte1952    Idiostroma roemeri; Lecompte, p. 316, pl. 66, fig. 3.

Reference Galloway and Ehlers1960    Idiostroma roemeri; Galloway and Ehlers, p. 67, pl. 4.

Reference Stearn and Selden2015b    Idiostroma roemeri; Stearn, p. E797, fig. 405a–c.

Holotype

Specimen (NHMUK PI P6076) from the Middle Devonian of Hebborn, Germany (Nicholson, Reference Nicholson1886a, pl. 9, figs. 6–11).

Description

The preserved specimens show vertically elongated skeletons with diameter of 3 cm. The skeleton consists of the axial zone and peripheral zone. In longitudinal section, the axial zone is subdivided into short pachysteles and arched laminae (Fig. 15.1). Pachysteles are spool-shaped, confined to interlaminar spaces, regularly superposed, spacing 6–8 per 2 mm and 0.05–0.15 mm in width. Laminae arch upward, spacing about 5–6 per 2 mm and 0.1–0.2 mm in thickness. The axial tube is observed, 1 mm in width. Galleries are circular, 0.1–0.3 mm in width. The peripheral zone is composed of short pachysteles and parallel laminae (Fig. 15.2). In some areas, skeletons are well differentiated. Pachysteles are spool-shaped, confined to interlaminar spaces, rarely superposed, spacing 5–6 per 2 mm and 0.1–0.3 mm in width. Laminae are continuous, tripartite with central dark line and surrounding light zones, spacing 4–5 per 2 mm and 0.3–0.5 mm in thickness. Dissepiments are common in galleries. In the other area, skeletons are densely compact and can hardly be differentiated. In tangential section, pachysteles are spool-shaped, confined to interlaminar spaces, regularly superposed. Laminae are concentric (Fig. 15.3). Galleries are circular or vertically elongated (Fig. 15.4). The axial tube is well developed, 1 mm in width. Microstructure is compact or peripherally vacuolate.

Material

Four specimens from the Middle Devonian of Hebborn (NHMUK PI P6074, P6075, P6076, P6077), in Germany; and one of Teignmouth (NHMUK PI P6078), in South Devon (UK).

Remarks

Idiostroma roemeri is different from the type species I. caespitosum in larger skeletons and the peripherally vacuolate microstructure.

Order Stromatoporida Stearn, Reference Stearn1980

Family Stromatoporidae Winchell, Reference Winchell1867

Genus Stromatopora Goldfuss, Reference Goldfuss1826

Type species

Stromatopora concentrica Goldfuss, Reference Goldfuss1826.

Stromatopora aff. S. polaris (Stearn, Reference Stearn1983)

Figure 15.5, 15.6

Reference Ripper1937a    Stromatopora concentrica; Ripper, p. 24, pl. 4, figs. 7, 8.

Reference Stearn1983    Ferestromatopora polaris; Stearn, p. 551, figs. 5A–D.

Reference Stearn1990    Stromatopora polaris; Stearn, p. 507, fig. 3.8.

Reference Webby, Stearn and Zhen1993    Stromatopora aff. polaris; Webby, Stearn, and Zhen, p. 158, figs. 5F, 23A–F, 24A.

Holotype

Specimen (GSC 66987) from the Lower Devonian Blue Fiord Formation of the Ellesmere Island, Arctic Canada (Stearn, Reference Stearn1983, fig. 5A, B).

Description

Growth forms are unclear due to limited preservation. The skeleton consists mainly of cassiculate structure. In longitudinal section, cassiculate structure is made up of oblique skeletons, interconnected with short pachysteles and pachystromes to form irregular networks (Fig. 15.5), 0.1–0.2 mm in width. Pachysteles are scarce, vertically distributed, confined to inter-pachystromes. Pachystromes are short, locally occurring, spacing 7–8 per 2 mm. Allotubes are circular or irregular, with rare dissepiments. Syringoporid symbionts rarely existed, 0.3–0.5 mm in width. In tangential section, irregular networks are observed (Fig. 15.6). Allotubes are circular, labyrinth-shaped, or irregular. Microstructure is cellular.

Material

Three specimens from Lower Devonian of Citadel rocks, Murrindal River (NHMUK PI H3904, H3905) and Commonwealth Quarries (NHMUK PI H3925), Buchan in Victoria (Australia).

Remarks

The cassiculate structure in Stromatopora aff. S. polaris closely resembles the specimens depicted by Webby et al. (Reference Webby, Stearn and Zhen1993). It is different from S. huepschii in terms of fewer long vertical pachysteles and a more common cassiculate structure.

Stromatopora huepschii (Bargatzky, Reference Bargatzky1881a)

Figure 16.1, 16.2

Figure 16.

Longitudinal (1) and tangential (2) views of Stromatopora huepschii (Bargatzky, Reference Bargatzky1881a) from Middle Devonian of Büchel, Germany, NHMUK PI P5488; note the allotubes indicated by yellow arrows. Longitudinal (3) and tangential (4) views of ?Pseudotrupetostroma buchanense (Ripper, Reference Ripper1937a) from Lower Devonian of Buchan, Victoria, Australia, NHMUK PI H3906. Longitudinal (5) and tangential (6) views of Pseudotrupetostroma porosum (Yang and Dong, Reference Yang and Dong1979) from the Middle Devonian of Teignmouth, South Devon, UK, NHMUK PI P5933.

Reference Bargatzky1881a    Caunopora hüpschii Bargatzky, p. 290.

Reference Nicholson1891a    Stromatopora hüpschii; Nicholson, p. 176, pl. 10, figs. 6, 9; pl. 22, figs. 3–7.

Reference Lecompte1952    Stromatopora hüpschii; Lecompte, p. 268, pl. 52, figs. 1–3.

Reference Yavorsky1961    Stromatopora hüpschii; Yavorsky, p. 43, pl. 26, figs. 4–6.

Reference Mistiaen1980    Stromatopora hüpschii; Mistiaen, p. 209, pl. 13, figs. 3–6.

Reference May1993    Stromatopora huepschii; May, p. 48, pl. 10, fig. 2; pl. 11, fig. 2.

Reference Cook1999    Stromatopora huepschii; Cook, p. 519, fig. 40A–F.

Reference May2005    Stromatopora huepschii; May, p. 194, pl. 22, fig. 1; pl. 30, fig. 1.

Reference Salerno2008    Stromatopora huepschii; Salerno, p. 91, pl. 14, fig. 1.

Holotype

Specimen (BIOB-PAL-Bargatzky-12, -13) from the Middle Devonian of Büchel, Germany.

Description

External forms appear to be laminar. The skeleton is composed of short pachysteles and cassiculate structure. In longitudinal section, pachysteles are commonly vertically discontinuous, spacing 5–6 per 2 mm and 0.1–0.2 mm in width (Fig. 16.1). Lateral skeletons are short, locally showing cassiculate structures or dots. Allotubes are irregular or vertically elongated, with abundant dissepiments. Branching syringoporid symbionts are also abundant, 0.3–0.6 mm in width, spacing 1 per 2 mm. In tangential section, pachysteles form irregular networks, enclosing allotubes (Fig. 16.2). Microstructure is cellular, 0.02 mm in diameter.

Material

Twenty-one specimens from Middle Devonian of Dartington (NHMUK PI P5843, P5877 P5887, P5888), Teignmouth (NHMUK PI P5486, P5835, P5836, P5838, P5844, P5886, P5892), in South Devon (UK); Büchel (NHMUK PI P5488, P5878, P5880, P5881), Gerolstein (NHMUK PI P5671, P5882, P5889, P5913, 2) and Hebborn (NHMUK PI P5532), in Germany.

Remarks

Stromatopora huepschii is characterized by short pachysteles and subordinate cassiculate structure. The skeleton lies between the typical Stromatopora (cassiculate structure) and Salairella (long, continuous pachystele). In South China, authors commonly identify specimens with continuous pachystele as Salairella (e.g., Huang et al., Reference Huang, Liang, Wang, Kershaw, Jeon, Li and Qie2022). However, for the specimens with intermittent pachysteles and rare cassiculate structure, we tentatively follow the traditional identification of Stromatopora huepschii before the restricted understanding of cassiculate structure had been made.

Genus Pseudotrupetostroma Khalfina and Yavorsky, Reference Khalfina and Yavorsky1971

Type species

Stromatopora pellucida artyschtensis Yavorsky, Reference Yavorsky1955.

? Pseudotrupetostroma buchanense (Ripper, Reference Ripper1937a)

Figure 16.3, 16.4

Reference Ripper1937a    Hermatostroma episcopale var. buchanensis Ripper, p. 32, pl. 5, figs. 9, 10.

Reference Webby, Stearn and Zhen1993    Pseudotrupetostroma buchanense; Webby, Stearn, and Zhen, p. 154, fig. 20A.

Lectotype

Specimen (NMV P 141671-72; Webby et al., Reference Webby, Stearn and Zhen1993) from the Lower Devonian Buchan Caves Limestone near Hicks’s, Murrindal, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 20A).

Description

The skeleton is composed of short pillars and long laminae. In longitudinal section, pillars are short, confined to interlaminar spaces, rarely superposed (Fig. 16.3), spacing 5–6 per 2 mm, 0.1–0.2 mm in width. Laminae are extensive, tripartite with a central dark line and surrounding light structure, spacing 2–4 per 2 mm, 0.1–0.2 mm in thickness. Galleries are circular or rectangular, with abundant dissepiments. In tangential section, pillars are isolated (Fig. 16.4). The microstructure is coarsely cellular.

Material

Two specimens (NHMUK PI H3906, H3908) from Lower Devonian of Murrindal, Buchan, Victoria (Australia).

Remarks

This species was revised to Pseudotrupetostroma buchanense by Webby et al. (Reference Webby, Stearn and Zhen1993) based on possible cellular microstructure. This study tentatively follows this identification.

Pseudotrupetostroma porosum (Yang and Dong, Reference Yang and Dong1979)

Figure 16.5, 16.6

Reference Yang and Dong1979    Hermatostroma porosum Yang and Dong, p. 68, pl. 38, figs. 7, 8.

Reference Dong and Wang1982    Trupetostroma cellulosum; Dong and Wang, p. 13, pl. 5, figs. 1, 2.

Reference Dong and Wang1982    Hermatostroma porosum; Dong and Wang, p. 22, pl. 13, figs. 1, 2.

Reference Salerno2008    Pseudotrupetostroma cellulosum; Salerno, p. 92–96, pl. 15, figs. 1–3.

Reference Huang, Liang, Wang, Kershaw, Jeon, Li and Qie2022    Pseudotrupetostroma porosum; Huang et al., p. 722, fig. 8A.

Holotype

Specimen (NIGP 3312 and 33130) from the Middle Devonian Donggangling Formation of Xiangzhou, Guangxi, China (Yang and Dong, Reference Yang and Dong1979, pl. 38, figs. 7, 8).

Description

Growth form appears to be laminar. The skeleton locally shows regular networks. In longitudinal section, pillars are vertically long, spacing 7–8 per 2 mm, 0.1–0.2 mm in width. Laminae are short, passing through several pillars, locally continuous with a central dark line and surrounding cellular structure (Fig. 16.5), spacing 7–8 per 2 mm, 0.1–0.2 mm in thickness. Syringoporids are common, 0.5 mm in width, with abundant dissepiments. In tangential section, skeletons consist of pachysteles, with rare, isolated pillars (Fig. 16.6). The microstructure is cellular.

Material

One specimen from the Middle Devonian of Teignmouth (NHMUK PI P5933), in South Devon (UK).

Remarks

The present specimens were previously assigned to Parallelopora goldfussi Bargatzky, Reference Bargatzky1881a, but they differ from the latter in having regular networks and cellular microstructure.

Pseudotrupetostroma ripperae Webby and Zhen, Reference Webby and Zhen1993

Figure 17.1, 17.2

Figure 17.

Longitudinal (1) and tangential (2) views of Pseudotrupetostroma ripperae Webby and Zhen, Reference Webby and Zhen1993, from Lower Devonian of Heath’s Quarry, Buchan, Victoria, Australia, NHMUK PI H3916. Longitudinal (3) and tangential (4) views of Neosyringostroma sp. from the Middle Devonian of Büchel, Germany, NHMUK PI P5530. Longitudinal (5) and tangential (6) views of Syringostromella cf. S. labyrinthea Stearn, Reference Stearn1990, from Lower Devonian of Heath’s Quarry, Buchan, Victoria, Australia, NHMUK PI H3921.

Reference Ripper1937a    Hermatostroma episcopale; Ripper, p. 29, pl. 5, figs. 7, 8.

Reference Webby and Zhen1993    Pseudotrupetostroma ripperae; Webby and Zhen, p. 340, figs. 7A–F, 8C, D, 9A, B.

Reference Webby, Stearn and Zhen1993    Pseudotrupetostroma ripperae; Webby, Stearn, and Zhen, p. 152, figs. 19F, 20B, C.

Holotype

Specimen (SUP 97268) from the Lower Devonian Jesse Limestone of the Limekilns area, New South Wales, Australia (Webby and Zhen, Reference Webby and Zhen1993, figs. 7E, F, 8C).

Description

The skeleton consists of short pillars and long laminae. In longitudinal section, pillars are short, confined to interlaminar spaces, locally superposed, spacing 5–7 per 2 mm, 0.1–0.2 mm in width. Laminae are laterally extensive, tripartite with a central dark line and surrounding cellular structure (Fig. 17.1), spacing 3–4 per 2 mm, 0.1–0.2 mm in thickness. Galleries are circular or irregular, locally with dissepiments. In tangential section, pillars are isolated (Fig. 17.2). Laminae are circular. The microstructure is cellular.

Material

One specimen (NHMUK PI H3916) from Lower Devonian of Heath’s Quarry, Buchan, in Victoria (Australia).

Remarks

This species was revised to Pseudotrupetostroma due to cellular microstructure. It differs from P. porosum in short pillars and from ?P. buchanense in cellular microstructure.

Genus Neosyringostroma Kaźmierczak, Reference Kaźmierczak1971

Type species

Hermatostroma logansportense Galloway and St. Jean, Reference Galloway and St. Jean1957.

Neosyringostroma sp.

Figure 17.3, 17.4

Reference Nicholson1891a    Stromatopora beuthii; Nicholson, p. 183, pl. 5, figs. 12, 13; pl. 23, figs. 8–13; pl. 24, fig. 1.

Description

External forms are noted as bulbous or domical (Nicholson, Reference Nicholson1891a). The skeleton is composed mainly of long pillars and short pachystromes. In longitudinal section, pillars are vertically long, spacing 4–5 per 2 mm and 0.2–0.3 mm in width (Fig. 17.3). Pachystromes are commonly short, confined to inter-pachystele spaces, locally continuous and showing cassiculate structure. Syringoporid symbionts locally occur, 0.2–0.3 mm in width. Autotubes are vertically elongated or irregular, with rare dissepiments. In tangential section, pillars are interconnected, forming pachysteles (Fig. 17.4), or locally isolated. Microstructure is cellular or melanospheric.

Material

Eighteen specimens from the Middle Devonian of Hebborn (NHMUK PI P5528, P5533, P5903), Büchel (NHMUK PI P5530), Gerolstein (NHMUK PI P5529, P5850, P5869), in Germany; Dartington (NHMUK PI P5904), Bishopsteignton (NHMUK PI 5840, P5841, P5905), Teignmouth (NHMUK PI P5839, P5883, P5885, P5907), Torquay (NHMUK PI P5842), and one unknown location (NHMUK PI P5906), in South Devon (UK); also, Giessen, 60 km north of Frankfurt, Germany (NHMUK PI P6147).

Remarks

The original specimens of Stromatopora beuthii were revised as Hermatostroma beuthii (Lecompte, Reference Lecompte1952). However, specimens illustrated by Nicholson (Reference Nicholson1891a) closely resemble Neosyringostroma due to dominant pachysteles and cellular microstructure. The present specimens differ from the type species of Neosyringostroma by less cassiculate structure; this study therefore leaves it in open nomenclature.

Family Syringostromellidae Stearn, Reference Stearn1980

Genus Syringostromella Nestor, Reference Nestor1966

Type species

Stromatopora borealis Nicholson, Reference Nicholson1891a.

Syringostromella cf. S. labyrinthea Stearn, Reference Stearn1990

Figure 17.5, 17.6

Reference Ripper1937a    Stromatopora concentrica; Ripper, p. 24, pl. 5, figs. 1, 2.

Reference Ripper1937a    Stromatopora hüpschii; Ripper, p. 28, pl. 5, figs. 5, 6.

Reference Stearn1990    Syringostromella labyrinthea; Stearn, p. 507, figs. 5.1, 5.2, 7.5, 7.6, 8.5.

Reference Webby, Stearn and Zhen1993    Syringostromella cf. labyrinthea; Webby, Stearn, and Zhen, p. 163, figs. 25D–F, 32C.

Holotype

Specimen (GSC 95779) from the Lower Devonian Stuart Bay Formation, Bathurst Island, Arctic Canada (Stearn, Reference Stearn1990, fig. 5.1, 5.2).

Description

The skeleton is composed of long pachysteles and short pachystromes. In longitudinal section, pachysteles are vertically long, upward joining or dividing (Fig. 17.5), spacing 4–6 per 2 mm and 0.1–0.2 mm in width. Pachystromes are short, confined to inter-pachystele spaces. Allotubes are vertically elongated, with rare dissepiments. In tangential section, skeleton is labyrinthine (Fig. 17.6). Microstructure is cellular.

Material

Two specimens from Lower Devonian of Murrindal (NHMUK PI H3907) and Heath’s Quarry (NHMUK PI H3921), Buchan, in Victoria (Australia).

Remarks

Specimens with dominant vertically long pachysteles were revised to Syringostromella by Stearn (Reference Stearn1990). We follow this decision.

Syringostromella zintchenkovi (Khalfina, Reference Khalfina and Halfin1961)

Figure 18.1, 18.2

Figure 18.

Longitudinal (1) and tangential (2) sections of Syringostromella zintchenkovi (Khalfina, Reference Khalfina and Halfin1961) from the Lower Devonian of Cave Hill, Lilydale, Victoria, Australia, NHMUK PI H3894. Longitudinal (3) and tangential (4) sections of Salairella buecheliensis (Bargatzky, Reference Bargatzky1881a) from the Middle Devonian of Büchel, Germany, NHMUK PI P5909; note the autotubes indicated by yellow arrows in (4). Longitudinal (5) and tangential (6) sections of Salairella lilydalensis (Ripper, Reference Ripper1937c) from Lower Devonian of Cave Hill Quarry, Lilydale, Victoria, Australia, NHMUK PI H3902; note the autotubes indicated by yellow arrows in (6).

Reference Ripper1937c    Stromatopora aff. hüpschii; Ripper, p. 186, pl. 8, figs. 7, 8.

Reference Khalfina and Halfin1961    Stromatopora zintchenkovi; Khalfina, p. 327, pl. D3, fig. la, b.

Reference Webby, Stearn and Zhen1993    Syringostromella zintchenkovi; Webby, Stearn, and Zhen, p. 163, figs. 24E, F, 25A–C.

Reference Prosh and Stearn1996    Syringostromella zintchenkovi; Prosh and Stearn, p. 34, pl. 15, figs. 1–3.

Reference May and Rodríguez2012    Syringostromella zintchenkovi; May and Rodríguez, p. 228, pl. 1, fig. K, L.

Holotype

Specimen from the Lower Devonian of Salair, southwestern Siberia, Russia (Khalfina, Reference Khalfina and Halfin1961, pl. D3, fig. la, b).

Description

The skeleton is composed of long pachysteles and short pachystromes. In longitudinal section, pachysteles are thick, long, discontinuous, vertically joining or dividing (Fig. 18.1), spacing 4–6 per 2 mm and 0.1–0.3 mm in width. Pachystromes are short, connected to pachysteles. Allotubes are vertically elongated or irregular, with sparse dissepiments. In tangential section, skeleton is labyrinthine (Fig. 18.2). Microstructure is coarsely cellular.

Material

One specimen from Lower Devonian of Mitchell’s Quarry (NHMUK PI H3894), Cave Hill, Lilydale, Victoria (Australia).

Remarks

Syringostromella zintchenkovi is characterized by long pachysteles, and therefore was revised to Syringostromella by Webby et al. (Reference Webby, Stearn and Zhen1993). It differs from Syringostromella cf. S. labyrinthea in thicker and longer pachysteles.

Genus Salairella Khalfina, Reference Khalfina and Halfin1961

Type species

Salairella multicea Khalfina, Reference Khalfina and Halfin1961.

Salairella buecheliensis (Bargatzky, Reference Bargatzky1881a)

Figure 18.3, 18.4

Reference Bargatzky1881a    Caunopora bücheliensis Bargatzky, p. 290.

Reference Nicholson1891a    Stromatopora bücheliensis; Nicholson, p. 186, pl. 10, figs. 5–7; pl. 23, figs. 4–7.

Reference Lecompte1952    Parallelopora bücheliensis; Lecompte, p. 290, pl. 50, figs. 3, 4.

Reference Fischbuch1969    Stromatopora huepschii; Fischbuch, p. 174, pl. 6, figs. 1–5.

Reference Mistiaen1980    Stromatopora cf. bücheliensis; Mistiaen, p. 210, pl. 13, figs. 7–9; pl. 14, figs. 1–3.

Reference Mistiaen1985    Salairella buecheliensis; Mistiaen, p. 145, pl. 12, figs. 10–12; pl. 13, fig. 1.

Reference Dong and Song1992    Stromatopora hüpschii; Dong and Song, p. 30, pl. 3, fig. 1a, b.

Reference May1999c    Salairella buecheliensis; May, p. 129, pl. 1, fig. 6.

Reference Cook1999    Salairella buecheliensis; Cook, p. 525, fig. 45A–E.

Reference May2005    Salairella buecheliensis; May, p. 204, pl. 21, figs. 1, 2; pl. 30, figs. 2, 3; pl. 31, fig. 1; pl. 32, fig. 1.

Reference Salerno2008    Salairella buecheliensis; Salerno, p. 97, pl. 14, fig. 2.

Reference Huang, Liang, Wang, Kershaw, Jeon, Li and Qie2022    Salairella buecheliensis; Huang et al., p. 724, fig. 8B.

Holotype

Specimen (BIOB-PAL-Bargatzky-6a, -6b) from the Middle Devonian of Büchel, Germany.

Description

Growth forms are laminar, bulbous, or domical. The skeleton is composed mainly of long pachysteles and subordinate short pachystromes. In longitudinal section, pachysteles are vertically long (Fig. 18.3), spacing 7–9 per 2 mm and 0.1–0.2 mm in width. Pachystromes locally occur, short, confined to inter-pachystele spaces. Autotubes are vertically elongated, with sparse or abundant dissepiments. Syringoporid symbionts are locally abundant, with a width of 0.3–0.6 mm, spacing 2 per 2 mm. In tangential section, pachysteles are interconnected, forming closed networks (Fig. 18.4), enclosing autotubes and allotubes. Microstructure is cellular, 0.02 mm in diameter.

Material

Thirty specimens from the Middle Devonian of Büchel (NHMUK PI P5909, P5910, P5911, P5912, P5922), Gerolstein (NHMUK PI P5670, P5673, P5870), in Germany; Teignmouth (NHMUK PI P5875, P5891, P5914, P5915, P5916, P5924, P5926, P5927, P5928, P5929), Dartington (NHMUK PI P5917, P5918, P5919, P5920, P5921), Bishopsteignton (NHMUK PI H4332), Lummaton Quarry (NHM P5871, P5872), Marychurch (NHMUK PI P5876), in South Devon (UK); Lake Winnipegosis (NHMUK PI P5590), in Manitoba (Canada); and Lower Devonian Cave Hill (NHMUK PI H3897), Lilydale, and Loyola (NHMUK PI H3928), in Victoria (Australia).

Remarks

The hypercalcified sponge Salairella buecheliensis has long and interconnected pachysteles, which is the main feature of this genus, and distinguishes it from Stromatopora and Parallelopora.

Salairella lilydalensis (Ripper, Reference Ripper1937c)

Figure 18.5, 18.6

Reference Ripper1937c    Stromatopora lilydalensis Ripper, p. 189, pl. 9, figs. 1, 2.

Reference Webby, Stearn and Zhen1993    Salairella lilydalensis; Webby, Stearn, and Zhen, p. 156, figs. 21A–F, 22A–C, 31E.

Lectotype

Specimen (NMV P141924-25, P141992-93; Webby et al., Reference Webby, Stearn and Zhen1993) from the Lower Devonian of the Lilydale Limestone at Mitchell’s (Cave Hill) Quarry, Lilydale, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 21B, C).

Description

The skeleton consists mainly of long pachysteles and short pachystromes. In longitudinal section, pachysteles are vertically extensive, spacing 8–10 per 2 mm and 0.1–0.3 mm in width (Fig. 18.5). Pachystromes are rare, short, confined to inter-pachystele spaces. Autotubes are vertically elongated, with sparse dissepiments. In tangential section, closed or labyrinthine network is formed by interconnected pachysteles (Fig. 18.6). Autotubes are round. Microstructure is coarsely cellular.

Material

Four specimens from Lower Devonian of Mitchell’s Quarry (NHMUK PI H3895, H3901) and Cave Hill Quarry (NHMUK PI H3902, H3903), Lilydale, in Victoria (Australia).

Remarks

Salairella lilydalensis differs from S. buecheliensis in closely distributed skeletons.

Order Syringostromatida Bogoyavlenskaya, Reference Bogoyavlenskaya1969

Family Coenostromatidae Waagen and Wentzel, Reference Waagen, Wentzel and Waagen1887

Genus Coenostroma Winchell, Reference Winchell1867

Type species

Stromatopora monticulifera Winchell, Reference Winchell1866.

Coenostroma sp.

Figure 19.1, 19.2

Figure 19.

Longitudinal (1, 2) sections of Coenostroma sp. from Lower Devonian of Buchan, Victoria, Australia, NHMUK PI H3920. Longitudinal (3) and tangential (4) sections of Atopostroma distans? (Ripper, Reference Ripper1937a) from Lower Devonian of Heath’s Quarry, Buchan, Victoria, Australia, NHMUK PI H3915. Longitudinal (5) and tangential (6) sections of Columnostroma clathratum Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993 from Lower Devonian of Mitchell’s Quarry, Cave Hill, Lilydale, Victoria, Australia, NHMUK PI H3893.

Reference Ripper1937a    Stromatopora concentrica var. colliculata; Ripper, p. 26, pl. 5, figs. 3, 4.

Reference Webby, Stearn and Zhen1993    Coenostroma sp.; Webby, Stearn, and Zhen, p. 166, figs. 26A–D.

Description

The skeleton consists of short pachysteles and extensive pachystromes. In longitudinal section, pachysteles are spool-shaped, confined to interlaminar spaces, and locally superposed, spacing 6–8 per 2 mm and 0.1–0.3 mm in width (Fig. 19.1). Pachystromes are long, thick, spacing 8–10 per 2 mm and 0.1–0.3 mm in thickness (Fig. 19.2). Galleries are small, circular or laterally elongated, with rare dissepiments. The tangential section is not preserved. Microstructure is unclear.

Material

Two specimens from Lower Devonian of Buchan (NHMUK PI H3919, H3920), in Victoria (Australia).

Remarks

Webby et al. (Reference Webby, Stearn and Zhen1993) revised this species to Coenostroma based on limited skeleton. This study follows this identification.

Genus Atopostroma Yang and Dong, Reference Yang and Dong1979

Type species

Atopostroma tuntouense Yang and Dong, Reference Yang and Dong1979.

Atopostroma distans ? (Ripper, Reference Ripper1937a)

Figure 19.3, 19.4

Reference Ripper1937a    Actinostroma stellulatum var. distans Ripper, p. 12, pl. 2, figs. 1, 2.

Reference Webby, Stearn and Zhen1993    Atopostroma distans; Webby, Stearn, and Zhen, p. 171, figs. 27F, 28A–D.

Reference Webby and Zhen2008    Atopostroma distans; Webby and Zhen, p. 222, fig. 3A–F.

Holotype

Specimen (NMV P141754-57) from the Lower Devonian Buchan Caves Limestone, Heath’s Quarry, Buchan, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 28A, B).

Description

The skeleton is composed of thick pillars and thin laminae. In longitudinal section, pillars are rod-like, long, passing through dozens of laminae (Fig. 19.3), spacing 8–12 per 2 mm and 0.1–0.2 mm in width. Laminae are thin, laterally extensive, locally thickened, spacing 12–16 per 2 mm, ranging from 0.02–0.1 mm in thickness. Galleries are circular or rectangular. In tangential section, pillars are isolated (Fig. 19.4). Laminae are undulating. Microstructure is unclear.

Material

One specimen from Lower Devonian of Heath’s Quarry (NHMUK PI H3915), Buchan, Victoria, Australia.

Remarks

The present species was assigned to Actinostroma, but typical colliculi have not been noticed. The regular network formed by long pillars and thin laminae shows a close relationship with Atopostroma. It was therefore grouped with Atopostroma by Webby et al. (Reference Webby, Stearn and Zhen1993). May (Reference May2005) noted that Atopostroma distans (Ripper, Reference Ripper1937a) is a junior synonym of Atopostroma frustulum (Počta, Reference Počta1894). Webby and Zhen (Reference Webby and Zhen2008) then noted the validity of this species (see discussion therein). The present specimen shows unclear microstructure, which cannot be clearly compared with specimens in the former two papers. Given the limitation of the material (only one poorly preserved specimen), this study tentatively retains the existence of this species but leaves a question.

Genus Columnostroma Bogoyavlenskaya, Reference Bogoyavlenskaya1972

Type species

Coenostroma ristigouchense Spencer, Reference Spencer1884.

Columnostroma clathratum Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993

Figure 19.5, 19.6

Reference Ripper1937c    Syringostroma aff. niagarense; Ripper, p. 179, text-fig. 1.

Reference Webby, Stearn and Zhen1993    Columnostroma clathratum; Webby, Stearn, and Zhen, p. 173, figs. 29A–F, 32F.

Holotype

Specimen (NMV P141922-23, P141990-91) from the Lower Devonian Lilydale Limestone at Mitchell’s (Caves Hill) Quarry, Lilydale, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 29A).

Description

The skeleton consists of long pillars and thin laminae. In longitudinal section, pillars are rod-like, thick, long, spacing 9–11 per 2 mm and 0.1–0.2 mm in width (Fig. 19.5). Laminae are thin, locally occur, spacing 10 per 2 mm, ranging from 0.02–0.1 mm in thickness. Galleries are small, circular. In tangential section, pillars are isolated between laminae (Fig. 19.6). Laminae are cut as closed works. Microstructure is unclear.

Material

One specimen from Lower Devonian of Mitchell’s Quarry (NHMUK PI H3893), Cave Hill, Lilydale, in Victoria (Australia).

Remarks

This species was revised to Columnostroma clathratum by Webby et al. (Reference Webby, Stearn and Zhen1993) based on long pillars. We follow this identification.

Genus Habrostroma Fagerstrom, Reference Fagerstrom1982

Type species

Stromatopora proxilaminata Fagerstrom, Reference Fagerstrom1961.

Habrostroma tyersense Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993

Figure 20.1, 20.2

Figure 20.

Longitudinal (1) and tangential (2) sections of Habrostroma tyersense Webby, Stearn, and Zhen, Reference Webby, Stearn and Zhen1993, from the Lower Devonian of Cave Hill, Lilydale, Victoria, Australia, NHMUK PI H3899. Longitudinal (3) and tangential (4) sections of Parallelostroma sp. from the Middle Devonian of Sötenich, Germany, NHMUK PI P5631. Longitudinal (5) and tangential (6) sections of Parallelopora goldfussi Bargatzky, Reference Bargatzky1881a , from the Middle Devonian of Teignmouth, South Devon, UK, NHMUK PI P5548.

Reference Ripper1937c    Stromatopora foveolata; Ripper, p. 185, text-fig. 2.

Reference Stock1988    Parallelostroma foveolatum; Stock, p. 12, figs. 3.5–3.8, 4.1, 4.2.

Reference Webby, Stearn and Zhen1993    Habrostroma tyersense; Webby, Stearn, and Zhen, p. 168, figs. 26E, F, 27A–E, 32D, E.

Holotype

Specimen (NMV P136342-43) from the Lower Devonian Coopers Creek Limestone, Tyers Quarry near Tyers, Victoria, Australia (Webby et al., Reference Webby, Stearn and Zhen1993, fig. 26E, F).

Description

Growth forms are domical to bulbous according to Ripper (Reference Ripper1937c). The skeleton is composed of short pillars and long laminae. In longitudinal section, pillars are spool-shaped, confined to interlaminar spaces, locally superposed, spacing 7–8 per 2 mm and 0.05–0.2 mm in width (Fig. 20.1). Laminae are thick, laterally extensive, common consisting of 2–4 microlaminae and micropillars, spacing 6–7 per 2 mm, ranging from 0.1–0.3 mm in thickness. Allotubes are circular or laterally irregular, with sporadic dissepiments. In tangential section, skeletons are isolated between interlaminar spaces and interconnected in laminae (Fig. 20.2). Microstructure is cellular.

Material

Two specimens from the Lower Devonian of Cave Hill (NHMUK PI H3899), Lilydale and the Lower Devonian of Rocky Camp (Commonwealth) Quarry (NHMUK PI H3927), Buchan, in Victoria (Australia).

Remarks

This species was assigned to the genus Stromatopora by Ripper (Reference Ripper1937c), but cassiculate structure has not been observed in it. The occurrence of isolated pillars, multi-microlaminae, and cellular microstructure justify the assignment into Habrostroma.

Family Parallelostromatidae Bogoyavlenskaya, Reference Bogoyavlenskaya1984

Genus Parallelostroma Nestor, Reference Nestor1966

Type species

Parallelostroma typicum (Rosen, Reference Rosen1867).

Parallelostroma sp.

Figure 20.3, 20.4

Description

Growth forms are unclear due to limited preservation. The internal skeletons consist of regular networks formed by short pachysteles and long pachystromes. In longitudinal section, pachysteles are thick, short, confined to interlaminar spaces, locally superposed, spacing 6–7 per 2 mm and 0.1–0.5 mm in width (Fig. 20.3). Pachystromes are thick, laterally extensive, locally discontinuous, consisting of several microlaminae and micropillars, spacing 8–10 per 2 mm, ranging from 0.1–0.2 mm in thickness. Allotubes are circular or irregular, with sparse dissepiments. In tangential section, skeletons are vermicular, with common astrorhizal canals and rarely isolated pillars (Fig. 20.4). Microstructure is orthoreticular.

Material

Five specimens from the Middle Devonian of Sötenich (NHMUK PI P5631, P5669), Gerolstein (NHMUK PI P5632, P5867, P5873), in Germany.

Remarks

The specimens were previously identified as Stromatopora concentrica by Nicholson (Reference Nicholson1886a), but the present species differs from the latter in regular networks and orthoreticular microstructure. It shows less development of cassiculate structures; we keep it in open nomenclature.

Genus Parallelopora Bargatzky, Reference Bargatzky1881a

Type species

Parallelopora ostiolata Bargatzky, Reference Bargatzky1881a.

Parallelopora goldfussi Bargatzky, Reference Bargatzky1881a

Figure 20.5, 20.6

Reference Bargatzky1881a    Parallelopora goldfussi Bargatzky, p. 295.

Reference Nicholson1891a    Parallelopora goldfussi; Nicholson, p. 191, pl. 25, figs. 4–9.

Reference Chi1940    Parallelopora goldfussi; Chi, p. 298, pl. 4, fig. 2.

Reference Lecompte1952    Stromatopora goldfussi; Lecompte, p. 280, pl. 57, figs. 3, 4; pl. 58, fig. 1.

Reference Yang and Dong1979    Parallelopora goldfussi; Yang and Dong, p. 63, pl. 34, figs. 5, 6.

Reference Dong and Song1992    Parallelopora goldfussi; Dong and Song, p. 32, pl. 4, fig. 3a, b.

Reference May1999c    Parallelopora goldfussi; May, p. 129.

Reference May2005    Parallelopora goldfussi; May, p. 216, pl. 22, fig. 2.

Holotype

Specimen (BIOB-PAL-Bargatzky-9) from the Middle Devonian of Hand near Paffrath, Germany.

Description

External forms are bulbous or domical (Nicholson, Reference Nicholson1891a). The skeleton consists of pachysteles and pachystromes. In longitudinal section, pachysteles are vertically long or confined to inter-pachystrome spaces, spacing 9–10 per 2 mm and 0.1–0.3 mm in width (Fig. 20.5). Pachystromes locally develop, thin or rarely thick, 0.02–0.3 mm in thickness. Autotubes are circular or vertically elongated, with common dissepiments. In tangential section, pachysteles are interconnected, forming closed networks (Fig. 20.6). Branching canals are observed. Microstructure is orthoreticular.

Material

Ten specimens from the Middle Devonian of Dartington (NHMUK PI P5557), Teignmouth (NHMUK PI P5547, P5548, P5551, P5938), in South Devon (UK); and Hebborn (NHMUK PI P5555, P5932), Hand near Paffrath (NHMUK PI P5930), Steinbreche, also near Paffrath (NHMUK PI P5552, P5931), in Germany.

Remarks

This species was revised into Stromatopora (Lecompte, Reference Lecompte1952), but cassiculate structure is not observed and the orthoreticular microstructure is different from the cellular microstructure of Stromatopora.

Parallelopora incrustans (Hall and Whitfield, Reference Hall and Whitfield1873)

Figure 21.1–21.4

Figure 21.

Longitudinal (1, 2) and tangential (3, 4) views of Parallelopora incrustans (Hall and Whitfield, Reference Hall and Whitfield1873) from the Upper Devonian of Iowa (USA), NHMUK PI P5801. Longitudinal (5) and tangential (6) views of Parallelopora ostiolata Bargatzky, Reference Bargatzky1881a, from the Middle Devonian of Büchel, Germany, NHMUK PI P5936; note the autotubes indicated by yellow arrows in (6).

Reference Hall and Whitfield1873    Stromatopora (Caenostroma) incrustans Hall and Whitfield, p. 227, pl. 9, fig. 3.

Reference Hall and Whitfield1873    Caunopora planulata; Hall and Whitfield, p. 228, pl. 9, fig. 2.

Reference Nicholson1886a    Stromatoporella? incrustans; Nicholson, p. 95, pl. 3, fig. 6.

Reference Parks1936    Trupetostroma planulatum; Parks, p. 62, p1. 11, figs. 1–3.

Reference Lecompte1952    “Caunopora planulata”; Lecompte, p. 286, pl. 50, figs. 1, 2.

Reference Galloway1957    Parallelopora planulata; Galloway and St. Jean, p. 242.

Reference Stearn1966    Parallelopora planulata; Stearn, p. 106.

Reference Stock1984    Stromatopora incrustans; Stock, p. 782, figs. 3E–H, 4.

Reference Stock, Groves, Walters and Day2008    “Habrostroma” incrustans; Stock, p. 49.

Holotype

Specimen (NYSM 332) from the Upper Devonian (Frasnian) Cerro Gordo Member of the Lime Creek Formation, near Rockford, Floyd County, Iowa, USA.

Description

Growth forms appear to be laminar. The skeleton is composed mainly of long pachysteles and short pachysteles (Fig. 21.1, 21.2). In longitudinal section, pachysteles are vertically extensive, locally joining together, spacing 7–8 per 2 mm and 0.1–0.3 mm in width. Pachystromes are irregular distributed, laterally confined, or locally passing through several pachysteles. Autotubes are vertically elongated or circular, with sporadic dissepiments. Astrorhizal canals are cut as circles, 0.3–1 mm in width. In tangential section, pachysteles are interconnected, forming closed networks, enclosing autotubes (Fig. 21.3). Astrorhizal canals are common (Fig. 21.1, 21.3). Microstructure is orthoreticular (Fig. 21.2, 21.4).

Material

Three specimens (NHMUK PI P5799, P5801, P5808) from the Upper Devonian (Frasnian) of Iowa (USA).

Remarks

Parallelopora incrustans is compared using type specimens of Hall and Whitfield (see Stock, Reference Stock1984). Given the absence of ring pillars, tripartite pachystromes, cassiculate structures, and isolated pillars, the present specimens differ from Stromatoporella, Trupetostroma, Stromatopora, and Habrostroma, respectively. This study tentatively revises them to Parallelopora based on dominant long pachysteles and orthoreticular microstructure.

Parallelopora ostiolata Bargatzky, Reference Bargatzky1881a

Figure 21.5, 21.6

Reference Bargatzky1881a    Parallelopora ostiolata Bargatzky, p. 292.

Reference Nicholson1886a    Parallelopora ostiolata; Nicholson, p. 95, pl. 2, figs. 6, 7.

Reference Lecompte1952    Parallelopora ostiolata; Lecompte, p. 292, pl. 51, fig. 3.

Reference Galloway1957    Parallelopora ostiolata; Galloway and St. Jean, p. 207, pl. 19, fig. 2.

Reference Galloway1960    Parallelopora ostiolata; Galloway, p. 632, pl. 76, fig. 4.

Reference Yang and Dong1979    Parallelopora ostiolata; Yang and Dong, p. 62, pl. 33, figs. 7, 8.

Holotype

Specimen (BIOB-PAL-Bargatzky-7) from the Middle Devonian of Büchel, Germany.

Description

External form is uncertain due to limited preservation. The skeleton consists mainly of long pachysteles. In longitudinal section, pachysteles are long, vertically extensive, locally joining together, densely packed (Fig. 21.5), spacing 9–11 per 2 mm and 0.2–0.3 mm in width. Pachystromes rarely develop, short, confined to inter-pachystele spaces. Autotubes are circular or elongate, with sparse dissepiments. In tangential section, pachysteles are interconnected, forming closed networks, enclosing autotubes or allotubes (Fig. 21.6). Microstructure is orthoreticular, with long micropillars and short microcolliculi.

Material

One specimen from the Middle Devonian of Büchel (NHMUK PI P5936), Germany.

Remarks

This specimen is the holotype of Parallelopora ostiolata, it differs from other species in closely spaced pachysteles and orthoreticular microstructure. Nicholson (Reference Nicholson1886a, p. 95) said, “More remarkable appearances are presented by P. ostiolata, Barg., from the original of which, through the kindness of Prof. Schlüter, I have also prepared thin sections.” Therefore, Nicholson’s (Reference Nicholson1886a) specimen (NHMUK PI P5936) in this paper came from the original one (BIOB-PAL-Bargatzky-7). Given that the two specimens are deposited in different institutes, different numbers are needed, even though they both are the holotype.

Family Stachyoditidae Khromykh, Reference Khromykh and Branch1967

Genus Stachyodes Bargatzky, Reference Bargatzky1881b

Type species

Caunopora verticillata McCoy, Reference McCoy1850.

Stachyodes verticillata (McCoy, Reference McCoy1850)

Figure 22.1–22.4

Figure 22.

Longitudinal (1, 2, 4) and transverse (3) views of Stachyodes verticillata (McCoy, Reference McCoy1850) from the Middle Devonian of Hebborn, Germany, holotype NHMUK PI P6069. Longitudinal (5) and transverse (6) views of Amphipora ramosa (Phillips, Reference Phillips1841) from the Middle Devonian of Hebborn, Germany, NHMUK PI P6071.

Reference McCoy1850    Caunopora verticillata McCoy, p. 377.

Reference Nicholson1886a    Stachyodes verticillata; Nicholson, p. 107, pl. 8, figs. 9–14.

Reference Nicholson1892    Stachyodes verticillata; Nicholson, p. 221, pl. 29, figs. 1, 2.

Reference Lecompte1952    Stachyodes verticillata; Lecompte, p. 303, pl. 57, figs. 1–3.

Reference Galloway1957    Stachyodes verticillata; Galloway, p. 444, pl. 34, fig. 10.

Reference Fischbuch1970    Stachyodes verticillata; Fischbuch, p. 1080, pl. 149, figs. 1–3.

Holotype

Specimen (NHMUK P6069) from the Middle Devonian of Hebborn, Germany (Nicholson, Reference Nicholson1886a, pl. 8, figs. 9–14).

Description

External forms are dendroid, but commonly cut as fragmented branches, 5–10 mm in diameter. In longitudinal section, the skeleton comprises thick pachysteles and dark thin pachystromes (Fig. 22.1). In the central zone, skeletons are densely compact, with axial and branching canals 0.2–1 mm in diameter. In the peripheral zone, skeletons are differentiated. Pachysteles radiate outward and upward from the central area and end perpendicularly to the surface (Fig. 22.2), 0.2 mm in width. Pachystromes are not well developed, 0.02 mm in thickness. Allotubes are elongate, perpendicular to the surface, 0.1 mm in diameter. In tangential section, closed networks are formed by interconnected pachysteles. The central zone is dominated by axial and branching canals (Fig. 22.3). In the marginal zone, the skeleton is featured by long pachysteles (Fig. 22.4). The microstructure is striated.

Material

Seven specimens from the Middle Devonian of Hebborn (NHMUK PI P6069, P6098, P6099), in Germany; and Middle Devonian of Teignmouth (NHMUK PI P6066, P6067, P6068, P6070) in South Devon (UK).

Remarks

Stachyodes verticillata is characterized by densely packed skeletons, well-developed axial and branching canals, thin allotubes, and typical striated microstructure. It is different from S. radiata in undifferentiated skeletons.

Order Amphiporida Rukhin, Reference Rukhin1938

Family Amphiporidae Rukhin, Reference Rukhin1938

Genus Amphipora Schulz, Reference Schulz1883

Type species

Caunopora ramosa Phillips, Reference Phillips1841.

Amphipora ramosa (Phillips, Reference Phillips1841)

Figure 22.5, 22.6

Reference Phillips1841    Caunopora ramosa Phillips, p. 19, pl. 8, fig. 22.

Reference Schulz1883    Amphipora ramosa; Schulz, p. 90, pl. 22, figs. 5–7.

Reference Nicholson1886a    Amphipora ramosa; Nicholson, p. 109, pl. 9, figs. 1–4.

Reference Chi1940    Amphipora ramosa; Chi, p. 312, pl. 5, figs. 1, 2.

Reference Lecompte1952    Amphipora ramosa; Lecompte, p. 325, pl. 67, fig. 3; pl. 68, figs. 1–7.

Reference Galloway1957    Amphipora ramosa; Galloway and St. Jean, p. 233, pl. 23, figs. 2–6.

Reference Stearn1961    Amphipora ramosa; Stearn, p. 946, pl. 107, figs. 9, 10.

Reference Birkhead1967    Amphipora ramosa; Birkhead, p. 84, pl. 16, fig. 2.

Reference Zukalová1971    Amphipora ramosa; Zukalová, p. 117, pl. 37, fig. 1; pl. 38, fig. 22.

Reference Yang and Dong1979    Amphipora ramosa; Yang and Dong, p. 79, pl. 43, figs. 7, 8.

Reference Dong, Wang, Zhou, Zhang, Luo, Fu and Huang1989    Amphipora ramosa; Dong et al., p. 278, pl. 16, fig. 3a, b.

Reference May1993    Amphipora ramosa; May, p. 31, pl. 3, fig. 3.

Reference Cook1999    Amphipora ramosa; Cook, p. 515, fig. 35A–H.

Reference Salerno2008    Amphipora ramosa; Salerno, p. 104, pl. 2, fig. 4.

Reference Stearn and Selden2015b    Amphipora ramosa; Stearn, p. E825, figs. 475, 476.

Neotype

Specimen (NHMUK P0308) from the Middle Devonian Chercombe Bridge Limestone, near Newton Abbott, Devon, UK (Stearn, Reference Stearn and Selden2015b, figs. 475, 476).

Description

Growth forms are dendroid, 4 mm in diameter. The skeleton shows networks formed by oblique pillars and short elements. In longitudinal section, pillars are long, oblique, radiating outward and upward from the skeletal center (Fig. 22.5), 0.1–0.3 mm in width. Short elements are also oblique, connecting with pillars and forming irregular networks, 0.1–0.2 mm in width. Dark central lines are observed in skeletal elements. Axial canal is conspicuous, 0.5 mm in diameter. Peripheral sheaths are well developed, 0.5 mm in width, with common occurrence of dissepiments. Skeletal walls are thin, connecting with outside extensions of pillars. Galleries are irregular. In tangential section, central networks are irregular, consisting of oblique pillars and short elements (Fig. 22.6). Axial canal is round. Peripheral sheaths are subdivided by pillars. Microstructure is compact.

Material

Nine specimens from the Middle Devonian of Teignmouth (NHMUK PI P6072, P6083, P6084, H4776) and Torquay (NHMUK PI H4779) in South Devon, (UK); and Middle Devonian of Herrenstrase (NHMUK PI P5756), Sötenich (NHMUK PI P5758), Hebborn (NHMUK PI P6071, H4778), in Germany.

Remarks

This species is very common and widely distributed, it is characterized by dendroid skeleton, apparent axial canal, and peripheral sheaths.