# Gigantoproductid and allied productid brachiopods from the “Calcaires à Productus” (late Viséan-Serpukhovian; Montagne Noire, southern France): Taxonomy and palaeobiogeographical position in the Palaeotethys

Markus Aretz, Marie Legrand-Blain, Daniel Vachard, Alain Izart

<figure><img src="image_1.png" /></figure>

## To cite this version:

Markus Aretz, Marie Legrand-Blain, Daniel Vachard, Alain Izart. Gigantoproductid and allied productid brachiopods from the “Calcaires à Productus” (late Viséan-Serpukhovian; Montagne Noire, southern France): Taxonomy and palaeobiogeographical position in the Palaeotethys. Geobios, 2019, 55, pp.17-40. 10.1016/j.geobios.2019.06.007. hal-04825323

HAL Id: hal-04825323

https://ut3-toulouseinp.hal.science/hal-04825323v1

Submitted on 8 Dec 2024

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
# Gigantoproductid and allied productid brachiopods from the “Calcaires à *Productus*” (late Viséan-Serpukhovian; Montagne Noire, southern France): Taxonomy and palaeobiogeographical position in the Palaeotethys *

Markus Aretz<sup>a,*</sup>, Marie Legrand-Blain<sup>b</sup>, Daniel Vachard<sup>c</sup>, Alain Izart<sup>d</sup>

<sup>a</sup> GET, Université de Toulouse, CNRS, IRD, UPS, (Toulouse), France

<sup>b</sup> 216 Cours Général de Gaulle, F 33170 Gradignan, France

<sup>c</sup> 1 rue des Tilleuls, 59152 Gruson, France

<sup>d</sup> 202 chemin de Cabanis, 34730 Prades-le-Lez, France

* Corresponding author. E-mail address: markus.aretz@get.omp.eu (M. Aretz).

* Corresponding editor: Bertrand Lefebvre.

## Abstract

The “Calcaires à *Productus*” or “Calcaires à *Productus giganteus*” on the southern slope of the Montagne Noire (Hérault, France) consist of numerous isolated olistolitic limestone blocks, generally transported by gravitational flows into siliciclastic basinal deposits. The present paper is the first systematic monographic study of gigantoproductids, semiplanids, and some other large-sized productid brachiopods from the upper Viséan to Serpukhovian strata of the Montagne Noire. Gigantoproductids are represented by abundant *Datangia semiglobosa* and *Kansuella* spp.; a single *Globosoproductus* specimen is described. The semiplanids are represented by *Latiproductus* aff. *latissimus*, *L. latissimus complicatus* and *L. edelburgensis*, and the linoproductids by *Balakhonia* aff. *kokdscharensis*. The Montagne Noire productid faunas are found in very different facies including shell beds and microbial mounds. They are most abundant in assemblages from agitated, shallow water environments with a firm substrate where rugose corals are rare or absent. The palaeobiogeographical
relations to the other regions in the western Palaeotethys are often limited to taxa with an almost cosmopolitan character. Worth noting is the absence of *Gigantoproductus sensu stricto* in the Montagne Noire. This genus, and other brachiopod taxa, migrated probably during the late Serpukhovian into southern France. On a larger scale, the presence of *Kansuella* indicates faunal exchanges with the eastern Palaeotethys (South China) via a westward migration path along North China, the Central Asian Orogenic Belt and south-eastern Laurussia. The studied taxa, even at the genus level, did not migrate into North America even if the Alleghanian Isthmus was still open.

Keywords:

Productid brachiopods

Systematic palaeontology

Palaeoecology

Palaeobiogeography

Stratigraphy

# 1. Introduction

Brachiopods are an important element among the Palaeozoic epifauna. They are abundant and diverse in various siliciclastic and carbonate facies from nearshore to outer platform and slope settings. Carboniferous brachiopods have been intensively studied since the mid-19th century (de Koninck, 1842-1844, 1847; Davidson, 1858-63). They have been the successful backbone of the Carboniferous stratigraphy (e.g., Vaughan, 1905) before the arrival of micropalaeontological biozonations. Besides the abundance and wide distribution of brachiopods in the Mississippian (Early Carboniferous), few modern data are available for interpreting the diversity, stratigraphic distribution and palaeobiogeographical relations of brachiopod faunas in the southern European Variscides (Legrand-Blain et al., 1983). The Montagne Noire is no exception from this general picture, although brachiopods have been mentioned and partly described from Mississippian deposits as early as the 19th century.

The first *Productus* was discovered and published by de Serres (1847) from the area of Roujan/Neffies, which, nevertheless, should be corrected as Laurens area (Böhm, 1935). In the Montpellier University's old collections, some productids labelled as "*Productus giganteus* Martin (jeune), Viséen sup. Neffies" may belong to this initial material. One of these specimens is figured herein as *Latiproductus* aff. *latissimus complicatus* (see below).
After this initial discovery, more fossiliferous localities were found and studied in the following decades, which increased the number of known species (de Verneuil in Fournet and Graff, 1849; de Rouville, 1868; Frech, 1887). It is also at this time that the name “Calcaires à *Productus*” (i.e., limestones with *Productus*) was introduced as a lithostratigraphic unit for the Carboniferous in the Cabrières area (de Rouville, 1868; Bergeron, 1889). For the geological excursion of the French Geological Society (de Rouville, 1868), productids and associated corals identified by de Verneuil were listed as *Productus giganteus*, *P. edelburgensis*, *P. latissimus*, *P. cora*, *P. semireticulatus*, *Caninia* aff. *gigantea*, and *Lithostrotion floriforme*. Some of these brachiopods are still housed at Montpellier University. Since that 1868 excursion, the “Calcaires à *Productus*” has been considered as a remarkable episode of the marine Carboniferous sedimentation in the Montagne Noire.

Böhm (1935) was the first to publish detailed taxonomic descriptions of Montagne Noire productids. Gèze (1949) interpreted the “Calcaires à *Productus giganteus*” as reefal coral-brachiopod lenses, localized at the top of the “Culm” sedimentation, and probably deposited in littoral environments. Vidal (1952 a, 1952b) was the last author to provide detailed information on Mississippian productids from the Montagne Noire. It is the aim of this study: (i) to restudy the productid brachiopods from the historical collections at the universities of Montpellier and Lyon and to add data from abundant newly collected materials (briefly mentioned in Legrand-Blain, 2003), and (ii) to discuss the spatio-temporal distribution of the Montagne Noire gigantoproductids in terms of biostratigraphy, palaeoecology and palaeobiogeographical connections, in comparison to the assemblages known in the western Palaeotethys.

## 2. Geological setting of the “Calcaires à Productus” in the Montagne Noire

In the southeastern Montagne Noire, sedimentary deposits of Mississippian age crop out in the Faugères and the Mont Peyroux nappes, as well as the Cabrières klippes. During the Mississippian, a change from pre-orogenic outer platform and starved basin deposits to syn-orogenic flysch deposits can be observed (Aretz, 2016; Vachard et al., 2017). The “Calcaires à *Productus*” comprise limestone lenses of different sizes and stratigraphic ages (from late Viséan to early late Serpukhovian), which are intercalated into often mud- to silt-sized, siliciclastic rocks (often called Kulm, or Culm, deposits) in the upper part of the Laurens Flysch Group (Korn and Feist, 2007; Aretz, 2016) or Laurens-Cabrières Group (Vachard et al., 2017). Their synorogenic gravitational transport from shallower into deeper parts of the flysch basin was hypothetically reconstructed, based on an accurate mapping, by Engel et al.
(1978, 1982). The "Calcaires à *Productus*" mainly represent microbial mounds that grew in the outer ramp and slope of the flysch basin under dysphotic and photic conditions (Vachard et al., 2017, Cózar et al., in press). However, also a variety of other carbonate facies of middle and outer ramp settings are known from this stratigraphic unit (Aretz, 2002b; Vachard et al., 2017).

A lithostratigraphic scheme for the "Calcaires à *Productus***" was defined by Poty et al. (2002) with successively: (i) the Roque Redonde Fm. (emendated in Vachard et al., 2017), corresponding to the main development of the "Calcaires à *Productus*", (ii) the Roc de Murviel Fm. (also emendated in Vachard et al., 2017), locally separated from the Roque Redonde Fm. by a karstic episode, and (iii) the La Serre de Péret Fm. (Vachard et al., 2017). The biostratigraphy of the "Calcaires à *Productus***" was established by foraminifers and calcareous microbiota (Vachard, 1974, 1977; Pille and Vachard, 2011; Vachard et al., 2016a, 2016b) and by rugose corals (Barchy, 1994; Aretz, 2002b; Poty et al., 2002; Aretz and Herbig, 2003).

Rugose corals of the Roque Redonde Fm. belong to zone RC8 of Poty et al. (2006), and those of the Roc de Murviel Fm. to RC9. Vachard et al. (2016a, 2016b) used foraminifers and carbonate microfossils for correlation of their regional biozones A-H for the *Productus* limestone to the Russian substages. Zones A-B are correlated to the Mikhailovian (A = upper Cf6 γ; B = basal Cf6 δ); zones C, D and E to the Venevian (= Brigantian); zone F to the Tarusian (basal Serpukhovian); zone G to the Steshevian (early Serpukhovian); and zone H to the early Protvian (early late Serpukhovian). As these regional biozones are difficult to identify in absence of foraminifers and algae, Vachard et al. (2017) rather used only the Russian substages when discussing their lithostratigraphic succession of the "Calcaires à *Productus***". Ammonoids, rare and difficult to interpret in the Laurens-Cabrières Group, allow to make correlations of the basal flysch deposits with the latest Viséan of the Rhenish Mountains, and the Barrac Fm. (unit I of flysch) with the early Serpukhovian of the Southern Urals and North Africa (Korn and Feist, 2007; Vachard et al., 2017).

**3. Material and methods**

3.1. *Studied productid collections and sections*

The studied productids (Table 1) are provided by three collections:

* samples collected by M. Legrand-Blain between 1980 and 2016, sometimes with the help of M. Aretz and E. Poty. This material will be deposited at Lyon University (UCBL-
FSL). The ML numbers make reference to the internal register of this collection for localities and stratigraphic horizons;

* samples from the Palaeontological Collections at Lyon University (UCBL-FSL). The productids of this collection were discovered in the 19th century by G.P. Deshayes (from Deshayes and Milne Edwards, 1836: 382-386), Ferdinand Geandey and Gustave Sayn. The latter most likely collected them during the “Réunion extraordinaire sur le versant méridional de la Montagne Noire, 6–15 September 1899”;

* samples from the Palaeontological Collections of the Montpellier University (UM ACI). The main collectors of these productid brachiopods were J.S. Albaille, R. Böhm (including the material from Böhm, 1935) and J. Vidal (Vidal, 1952b).

The studied specimens were collected from late Viséan to Serpukhovian strata of 15 sections (Fig. 1; Table 2).

## 3.2. *Quantitative palaeobiogeography*

The distributions of 49 species of gigantoproductids in eleven palaeogeographical units in the western Palaeotethys have been compiled from our own data and those of the literature. If necessary, M. Legrand-Blain has corrected the taxonomy provided in the literature. These data have been used to construct dendrograms for hierarchical cluster analyses using Raup-Crick (Raup and Crick, 1979) coefficient and Jaccard similarities (Jaccard, 1901) and unweighted pair-group average (UPGMA) method. Percentages related to clusters are bootstrap supports (1000 iterations).

The Jaccard similarity is generally considered to be the most reliable similarity coefficient for binary (presence-absence) data for palaeobiogeographical purposes (taxa per geographical unit; e.g., Shi, 1993; Cascales-Miñana, 2010; Davydov and Cózar, in press). However, the Raup-Crick coefficient provides a statistically based similarity measure and is less affected by sample biases (Raup and Crick, 1979; Hammer and Harper, 2006). These sample biases can be assumed not only from the uneven distribution of the taxa in the data set, but also the different recent work effort and application of taxonomic concepts in the different incorporated geographical units. The result of the analyses with the Raup-Crick coefficient is preferred herein, since it is more sensitive to the lack of taxa in any of the compared regions and widespread taxa do not have a disproportionate effect as observed for the Jaccard and Euclidian coefficients.

Cluster analyses have been supplemented by a Nonmetric Multidimensional Scaling (NMDS) ordination method based on the Raup-Crick coefficient to verify the expected
palaeobiogeographical relationships (Clarke, 1993). All cf. and aff. data have been used as
full presence of the nominal species in the analyses. All analyses have been performed with
the PAST 3 software (Hammer et al., 2001).

**4. Systematic palaeontology (Marie Legrand-Blain)**

The productid classification published in the Treatise on Invertebrate Paleontology
(Brunton et al., 1995, 2000) is followed herein. Partly different elements of systematics more
recently published by Waterhouse (2002, 2004), Lazarev (2003, 2004) and Qiao and Shen
(2015), are also mentioned.

Phylum Brachiopoda Duméril, 1806

Class Strophomenata Williams, 1996

Order Strophomenida Öpik, 1934

Suborder Productidina Waagen, 1883

Superfamily Linoproductoidea Stehli, 1954

Family Linoproductidae Stehli, 1954

Subfamily Linoproductinae Stehli, 1954

Genus *Balakhonia* Sarycheva, 1963 in Sarycheva, Sokolskaya, Beznosova and Maksimova, 1963

**Type species:** *Balakhonia ostrogensis* Sarycheva, 1963, by original designation; upper Viséan-Serpukhovian, Kuzbass (Kuznetsk Basin; SW Siberia, Russia).

**Diagnosis:** See Brunton et al. (2000: p. 528).

**Distribution:** Upper Viséan-lower Serpukhovian and Bashkirian; Eurasia, northern Africa.

**Remarks:** Lazarev (2004) included *Balakhonia* in the family Schrenkiellidae because of one row of widely spaced spines along hinge, which are thicker than those on the trail; such cardinal spines are interpreted as fixing the shell on the substrate, at least during early growth stages. On the other hand, Sarycheva (1970: p. 31, fig. 23) figured cardinal spines on a *Balakhonia* ventral valve. *Balakhonia* is cited in the Viséan of China (Yang et al., 1989) and Morocco (Termier et al., 1977), and doubtfully in the Spanish Pennsylvanian (Martínez Chacón and Winkler Prins, 2009).

*Balakhonia* aff. *ostrogensis* Sarycheva, 1963
Fig. 2(1-3)
aff. 1963. *Balakhonia ostrogensis* nov. sp. – Sarycheva, p. 233-234, text-fig. 103, 104, pl. 38, figs. 1-3.

aff. 1970. *Balakhonia ostrogensis* Sarycheva – Abramov, p. 125, pl. 15, fig. 6-7.

**Material:** ML sampling: Eight specimens from Roquemaillère (ML 764), most of them are incomplete, UCBL-FSL 703028-703035; from Péret-Rouirèdes (ML 1055), three umbonal fragments, UCBL-FSL 703046-703048.

**Description:** Shell rectangular, with a long trail, exhibiting a width of 70-80 mm, and a length up to 95 mm. The maximum width is located at hinge, and small ears are hardly separated from a wide umbo. In profile view: ventral valve with a slightly convex visceral disk, turning with a moderate curvature to a nearly straight trail, ending with a short frontal reverse curvature. Ornamentation of rounded, low costae, 10-13 in 10 mm. Rugae are present on ears and posterior part of umbo. The spines bases are poorly preserved, especially along the hinge, which is buried in sediment (UCBL-FSL 703032, 703047); rare and small on trail, where they are thinner than ribs. Dorsal valve: ornamentation as on the ventral valve, with some pits instead of the ventral spines. Thickness of test, ventral and dorsal valves: 1-2 mm; visceral cavity: 3-4 mm maximum. Dorsal interior (UCBL-FSL 703048): thin median septum, thickening posteriorly; small adductor scars; cardinal process not preserved. Ventral interior not preserved.

**Remarks:** The Montagne Noire specimens are comparable to *Balakhonia ostrogensis* by their long trail; they differ slightly in having a larger size, coarser ribs, and probably smaller cardinal spines. *Balakhonia ostrogensis* is reported from the upper Viséan–Serpukhovian, Ostrog Fm. in the Kuznetsk Basin (SW Siberia), and from the lower Serpukhovian of Sette-Daban (NE Siberia) (Abramov, 1970). *Balakhonia kokdscharensis* (Gröber, 1908), known from the Tian-Shan (Kyrgyzstan and NW China), Kuznetsk Basin and Tibet, is sometimes figured with a long trail (Zhang and Jin, 1976: pl. 9, fig. 1), but its ears are larger and more pointed than those of the Montagne Noire shells. Some Viséan specimens from Germany may be possibly attributed to *Balakhonia* (Winkler Prins and Amler, 2006: p. 94): *Productus* (*Gigantella*) *gigantoides* and *P. (Linoprotonia) rhenanus* Paeckelmann, 1931; nevertheless, they differ from the Montagne Noire samples by their subcircular outlines.

*Family Monticuliferidae* (Muir-Wood et Cooper, 1960)

*Subfamily Gigantoproductinae* Muir-Wood et Cooper, 1960

*Tribe Gigantoproductini* Muir-Wood et Cooper, 1960

**Diagnosis:** See Brunton et al. (2000: p. 551).
**Remarks:** The family Monticuliferidae has been variously interpreted, including or not the gigantoproductids, because *Monticulifera* is a Chinese genus from the Middle Permian. The following remarks can be made:

* Muir-Wood and Cooper (1960) and Muir-Wood and Williams (1965) described: (i) in the family Linoproductidae, the subfamily Monticuliferinae, "Linoproductidae with ornament of pustules (monticules) interrupting costellae or capillae"; (ii) the family Gigantoproductidae new fam., defined as large, usually thick-shelled, wide-hinged, and costellate Productacea with thin body cavity, with or without interarea;

* Lazarev (1990) revised, as follows, the description of the family Monticuliferidae Muir-Wood et Cooper 1960: Moderately thin body cavity; spines only on ventral valve, rarely absent; no marginal structures; no series of dorsal trails; Monticuliferidae is a primitive family appearing at Lower Devonian (Eoproductellinae), evolving towards specialized forms: especially Monticuliferinae, Permian. Gigantoproductidae are separated from Monticuliferidae;

* Brunton et al. (1995, 2000) described the Monticuliferidae, including the Gigantoproductinae, as displaying: moderately shallow corpus cavity, rarely very shallow; rugae posterolaterally or irregularly widespread; spines on ventral valve, rarely restricted to hinge region; marginal structures normally absent; Lower Devonian (Pragian)-Upper Permian (upper Tatarian). Gigantoproductinae: gigantic, large or medium size, hinge at greatest width; corpus cavity very shallow; fully ribbed, spines on ventral valve, rarely also on dorsal valve;

* Waterhouse (2002) recalls the spinose monticules on the ventral valve of *Monticulifera*; Monticuliferidae should be treated as an unusual Linoproductoid family, Permian only. Gigantoproductinae are included in the family Linoproductidae;

* Lazarev (2003) develops the spine-distribution pattern between: (i) Monticuliferidae, small spine bases, which on the ventral valve do not disturb the uniform distribution of ribs; and (ii) Linoproductidae, the number of ribs diminishing behind spine bases: a major portion of gigantoproductids, including *Gigantoproductus*, should be placed in the family Linoproductidae;

* Both Lazarev (2003) and Waterhouse (2002, 2004), discussing the pattern of ventral spines in relation with ribs, considered it as useful for the systematics of the Linoproductoidea. However, Lazarev (2003, p. 495, 496) observed: "any substantial increase in the weight of shells in gigantoproductids could result in a reduction in the
supportive function of the spines and, thus, in a disproportionately large (when compared to the shell dimensions) decrease in their thickness [...]"; he concluded that the spines/ribs pattern cannot be always used as fundamental for systematics of Linoproductoidea. The Montagne Noire gigantoproductids have thick ventral valves (except *Globosoproductus*), and spines on ventral trails rarely preserved; following Lazarev's opinion, their relationships with costae are considered herein as not essential for gigantoproductid classification.

**Genus *Datangia*** Yang, 1978

**Type species:** *Datangia weiningensis* Yang, 1978. Zhaojiashan Fm. (upper Viséan), Weining, Guizhou, China. Shen et al. (2017: p. 591) mentioned that Brunton et al. (2000: p. 551) mistook *Datangia luzhaiensis* Ni, 1977 (in Yang et al.) as the type species of *Datangia*.

**Diagnosis** (after Shen et al., 2017): shell medium to large in size, elongate in outline, semiglobose in profile, widest at hinge, slightly wider than long; ventral valve strongly convex; umbo tapering, extending slightly beyond hinge; ears flat, triangular; flanks steep; dorsal valve concave on visceral disc, geniculated anteriorly; corpus cavity short and deep. Ventral valve ornamented with even, regular costae, rugae prominent on ears, inconspicuous on venter and flanks; spines scattered along hinge and on ears, sparsely distributed on ventral trail. Venter interior with dendritic adductors enclosed by elongate diductors with striated anterior parts. Dorsal interior with trilobate cardinal process, supported by broad, notched median septum tapering anteriorly; adductors oval, dendritic; brachial ridges indistinct.

**Distribution:** Viséan-Serpukhovian; China, Europe, and northern Africa.

**Remarks:** The genus *Moderatoproductus* Litvinovich et Vorontsova, 1983 (type species: *Gigantella moderata* Schwetsov, 1922) is cited as a junior synonym of *Datangia* by Yang et al. (1977), Brunton et al. (2000) and Shen et al. (2017). *Datangia pyrenaea* (Legrand-Blain, 1983), first described as *Gigantoproductus irregularis pyrenaeus* (Legrand, in Legrand-Blain, Delvolvé and Perret, 1983) was found in the upper Serpukhovian strata of the central French Pyrénées.

*Datangia semiglobosa* (Paeckelmann, 1931)

Fig. 2(4-8)

1931. *Productus* (*Gigantella*) *striato-sulcatus* Schwetsov, var. n. *semiglobosa* – Paeckelmann, p. 246, pl. 21, fig. 5; pl. 24, fig. 1 a-c. e.p.
1962. *Gigantoproductus striato-sulcatus* (Schwetsov) var. *semiglobosa* Paeckelmann –
Zakowa and Zak, p. 206, pl. 14, fig. 2; pl. 18, figs. 1, 2.

1962. *?Gigantoproductus striato-sulcatus* (Schwetsov) var. *semiglobosa* Paeckelmann –
Zakowa and Zak, pl. 15; pl. 16, fig. 1; pl. 17, fig. 1.

1966. *Gigantoproductus striato-sulcatus* (Schwetsov) var. *semiglobosa* Paeckelmann –
Zakowa, p. 80, pl. 7, figs. 1, 2; pl. 8, fig. 1; pl. 20, fig. 1.

1966. *?Gigantoproductus striato-sulcatus* (Schwetzov) var. *semiglobosa* Paeckelmann –
Zakowa, pl. 9, fig. 1; pl. 13, figs. 1, 2.

1974. *Gigantoproductus semiglobosus* (Paeckelmann) – Kalashnikov, p. 85, pl. 27, fig. 1.

1979. *Gigantoproductus semiglobosus* (Paeckelmann) – Jin et al., p. 94, pl. 28, figs. 1, 2, 12.

1981. *Gigantoproductus semiglobosus* (Paeckelmann) – Pattison, p. 8, pl. 1, fig. 4, pl. 9, fig. 10.

1983. *Gigantoproductus semiglobosus* (Paeckelmann) – Yang et al., pl. 9, fig. 7.

1985. *Gigantoproductus semiglobosus* (Paeckelmann) – Zakowa, p. 115, pl. 3, fig. 1, pl. 4, fig. 1.

1987. *Gigantoproductus semiglobosus* (Paeckelmann) – Amler, p. 269, pl. 9, fig. 5.

1991. *Moderatoproductus semiglobosus* (Paeckelmann) – Litvinovich and Vorontsova, p. 27, pl. 15, fig. 2.

1992. *Gigantoproductus* aff. *semiglobosus* (Paeckelmann) – Legrand-Blain in Martínez
Chacón and Legrand-Blain, p. 133, pl. 8, figs. 1-4, pl. 9, figs. 1-3, pl. 10, figs. 1-3.

**Material:** ML sampling: Castelsec quarry (ML 765) (Venevian–Protvian: Vachard et al., 2017), 1 specimen, UCBL-FSL 703023. Lyon University: Vailhan area: 1 specimen, UCBL-FSL 703010; Cabrières, Sayn Collection: 1 specimen, UCBL-FSL 703012; Neffies area: 1 specimen, UCBL-FSL 703006. Montpellier University: Cabrières, Déjean Collection: 1 specimen, UM ACI 663.

**Measurements:** Maximum width at hinge up to 140 mm; since no specimen has complete ears (except one ear on UCBL-FSL 703010 and 703023), the measured width is only a minimum. Straight length = 120 mm; curved length up to 230 mm.

**Description:** Large-sized shell; ventral umbo strongly convex, trail and flanks slightly curved; ears well-marked off, moderately large in size. Hinge poorly preserved, probably without ginglymus. Ornamentation of costae present on the whole shell and rugae on the anterior part of the disk. Small spine bases dispersed on the ventral valve. Some specimens have indistinct longitudinal trail plications at the anterior margin, allowing a slight morphological resemblance to *Gigantoproductus* – which differs as bearing “ribbing entire,
but commonly irregular and weakened at trail plications" (Brunton et al., 2000: p. 551). The
small plicae on some *D. semiglobosa* could result from shell deformations after deposition in
muddy bottoms (cf. Zakowa, 1985). Ventral valve thickness variable: 4-8 mm posteriorly, *ca*.
2 mm near the front. Dorsal valve very thin: 1.5 mm. Corpus cavity narrow: 4 mm at umbo.
Interior of ventral valve: elongate diductor muscle scars are well visible on a partly exfoliated
umbo (UCBL-FSL 703010; Fig. 2(8)). No dorsal interior is available.

**Remarks:** The Montagne Noire ventral valves, especially UCBL-FSL 703010 (Fig. 2(6-8)) resemble the *D. semiglobosa* type specimen (Paeckelmann, 1931: pl. 24, fig. 1) by their dimensions and the globose umbo well-separated from ears. Slight differences may be rare faint anterior longitudinal plications. In the type area of the species in SW Poland, *Datangia semiglobosa* is reported from the marine to deltaic upper Viséan Szczawno Fm. (Zdanowski and Zakowa, 1995). On the other hand, Qiao and Shen (2015: fig. 5) attributed the species *semiglobosus* to *Gigantoproductus* and its stratigraphic range up to the lower Serpukhovian. The numerous citations and figurations of the species in the upper Viséan from Poland, England, Germany, Spain, Urals, and China, suggest that the species is rather variable in global size and development of the ears.

Genus *Globosoproductus* Litvinovich et Vorontsova, 1983

**Type species:** *Gigantella magnifica* Schimansky, 1940. Middle Viséan, northern Urals.

**Diagnosis:** See Brunton et al. (2000: p. 552).

**Distribution:** Viséan-early Serpukhovian; Russia, Ukraine, Poland, Germany, Belgium, and questionably France.

**Remarks.** *Globosoproductus* is comparable to *Datangia* by its subcircular to transverse oval outline, without longitudinal trail plicae. However, it differs as being very thin-shelled: ventral valve, 1-2 mm max. Hence, it could be not totally attributable to *Gigantoproductini*, characterized by "thick-walled ventral valves" (Brunton et al., 2000). In Russia, five species originally attributed to *Moderatoproductus* and *Gigantella* were transferred to *Globosoproductus* by Litvinovich and Vorontsova (1991), includuing the widely distributed *Globosoproductus striatellus* (Paeckelmann, 1931) from the upper Viséan of Germany, Poland, Urals and Donbass. Another species with thin-shelled valves perhaps attributable to *Globosoproductus* is *Gigantella bisati* Paeckelmann, 1931, upper Viséan of Poland and Belgium. In the Northern Massif Central (France), from the middle Viséan strata at L'Ardoisière near Vichy, the "*Productus giganteus*" specimens figured by Julien (1896: pls. 12, 13) belong to *Globosoproductus* (Legrand-Blain, 2018).
*Globosoproductus* sp.

Fig. 3

**Material:** Montpellier University: Laurens area, a big ventral valve, UM ACI 653.

Stratigraphic level: lower Serpukhovian, according to *Falcifolium pantherinum, Archaediscus (angulatus* stage) sp., *Biseriella parva*, and *Pseudoendothyra* ex gr. *struvei* found in a small rock fragment bordering the brachiopod (D. Vachard and P. Cózar, pers. comm., February 2017).

**Measurements:** Length: 140 mm; width > 170 mm; ventral valve maximum thickness: 1.5 mm.

**Description:** A large ventral valve; umbo broad and prominent, ears small; trail rather short. Umbo poorly preserved: presence or absence of area unknown. Longitudinal profile: umbo and trail regularly convex. Ornamentation of fine costae, 11 costae per 10 mm. Rugae poorly preserved, on ears and posterior part of umbo. On the trail, some very faint narrow longitudinal plicae. Spine bases: rare on umbo, very small, no larger than ribs. On the left part of the shell, some holes are traces of predators, comparable to other ones figured on a *Kansuella* (Fig. 4(8)). The shell thickness, on posterior and anterior parts of the specimen, is no more than 1.5 mm.

**Remarks:** The main characters of *Globosoproductus*, large size and extreme thinness of the shell, correspond to the morphology of the ventral valve from Laurens. This specimen differs from the type species *G. magnificus* by a wider outline and smaller and rarer spine bases, so as the specimens from L'Ardoisière; from *G. striatellus*, it differs by a larger size, smaller ears, and fewer rugae.

Genus *Kansuella* Chao, 1928

**Type species:** *Striatifera kansuensis* Chao, 1927 (p. 108).

**Type locality:** Cho-niu-kou locality and Fm., Kansu (Gansu) Province, NW China, Viséan.

**Distribution:** Late Viséan-Serpukhovian; China, Scotland, Algeria. The original descriptions of kansuellids are from China.

**Diagnoses:**

(i) Chao (1928: p. 67): [...] thick shell, large size and transverse outline. The surface is ornamented with flexuous striae and sometimes also concentric undulations in the early stages. The interior structures belong to the type of *Productus giganteus*. The most
remarkable character is the presence, in typical forms, of a narrow area in both valves,
that of the brachial valve being much narrower and less marked.

(ii) Brunton et al. (2000: pp. 552-554): Transverse outline; weakly concavoconvex with weakly inflated ventral umbo and prominent ginglymus in both valves; rugae posteriorly, but weak anteromedianly; interiors as in *Gigantoproductus* but with bilobed cardinal process. The genus *Parakansuella* Tan, 1987 (type species, *P. xinshaoensis*, Xinshao, Hunan, S. China, Serpukhovian) is cited as synonym of *Kansuella*: the main common diagnostic character is the presence of ginglymus in both valves.

*Parakansuella* "may differ in lacking rugae and more inflated ventral umbo".

(iii) Chen and Shi (2003: p. 157) described *Kansuella kansuensis*. Concentric wrinkles of the dorsal valve more distinct than those of the ventral valve, corpus cavity shallow and dorsal interior with strong dendritic adductor scars and highly convex brachial cones, have been suggested by these authors.

(iv) Shen et al. (2017: p. 592) cite, among other assigned species, "*Kansuella maxima* (McCoy, 1844)" after descriptions of Chinese Serpukhovian faunas as "*Kansuella maxima*" in the Tianshan–Higgan Region, East Dzungar, Nanmingshui Fm. figured in Yang et al. (1983: p. 47, fig. 7b). This generic assignation was not followed by Qiao and Shen (2015: fig. 5) who used *Gigantoproductus maximus*. Kansuellid species described from China are *Kansuella kansuensis* (Chao, 1927), *K. nasutula* Ni 1977 (in Yang et al.), *K. maxima rugosa* Shan et Zhao, 1981, *K. sulcata* Shan et Zhao, 1981, *K. geniculata* Jin et Sun, 1981, *K. jiugongqiaoensis* Liu 1982 (in Liu et al.), and *Parakansuella xinshaoensis* Tan, 1987. In Europe, *Kansuella* sp. is recorded at Muirkirk, Ayrshire, Scotland, in Lower Limestone Group, Brigantian (Brunton et al., 2000).

**Remarks:** *Kansuella* was included successively in: (i) Kansuellinae subfamily nov., by Muir-Wood and Cooper (1960: p. 336): "Gigantoproductidae resembling *Gigantoproductus* but having an interarea on both valves..."; (ii) Subfamily Gigantoproductinae, tribe Gigantoproductini by Brunton et al. (2000) citing "Kansuellinae" as members of Gigantoproductini; (iii) Kansuellinae and Kansuellidae by Waterhouse (2002, 2004), according to the "ventral spines, emerging anteriorly over disk from a slight to considerable swelling, through which the hollow spine base is prolonged posteriorly. One, two or rarely three costae or ribs may pass forward into the swollen spine base, and none to one rib may continue forward from the swelling [...]". However, such a pattern of the spine distribution cannot be so important in the systematics, in comparison to the ginglymus on both valves.
The identification of *Kansuella* requires samples with well-preserved dorsal valves showing dorsal ginglymus. Since many *Gigantoproductus* species are provided with an area in ventral valve only (cf. Lazarev, 1990: text-fig. 43; Brunton et al., 2000: fig. 386c “short ginglymus”), it is possible that some European species or specimens attributed to *Gigantoproductus* without a well-preserved dorsal valve are in fact belonging to *Kansuella*.

The formations containing the type specimens of *Parakansuella xinshaoensis* (Tzemenchiao Fm., central Hunan, South China) and *Kansuella kansuensis* (Chouniugou Fm., Gansu Province, North China) span the upper Viséan and early Serpukhovian (Dr. Li Qiao, personal communication).

The Scottish specimen of *Kansuella* sp. (Brunton et al., 2000) from Muirkirk, Ayrshire, figured by Davidson (1880: pl. 36, fig. 1), was commented on by Muir-Wood and Cooper (1960: p. 337): "this specimen preserved in the British Museum (Nat. Hist. n° B.5770) has an interarea in each valve and ornament comparable to that figured by Chao". Wilson (1989) gave more precise stratigraphic data: Lower Limestone Group, Brigantian; "several other gigantoproductoids from near the base of the Lower Limestone Group in the Muirkirk and Douglas areas have interareas. In the present classification, these specimens would be referred to as *Kansuella*". In the Algerian Bechar Basin, two species with large transverse shells, ventral and dorsal interareas (already figured but not taken into consideration) can be attributed to *Kansuella*: "*Gigantoproductus* meharezensis Legrand-Blain (1973: p. 139, fig. 19), late Viséan, and "*Latiproductus* (?)" *daguini* Legrand-Blain (1980: p. 51, pl. 4 fig. 3), Serpukhovian.

*Kansuella* sp. 1
Figs. 4(1, 3-6), 5(1, 5, 6)

**Material:** ML sampling: Laurens-Gare (ML 1158): 12 specimens UCBL-FSL 703004, 703007, 703011, 703013-703021; Castelsec quarry (ML 765, 1175): one ventral valve, UCBL-FSL 703024 and two ventral valve fragments, UCBL-FSL 703025, 703026; Roquemaillère (ML 764): one large-sized shell UCBL-FSL 703036; La Serre vineyard (ML 1176): one fragment (umbo), UCBL-FSL 703038; Péret-Rouirèdes (ML 1055), a fragment of ventral valve, UCBL-FSL 703049. Lyon University: a fragmentary shell from Vailhan, UCBL-FSL 703009. Montpellier University: Two specimens from Cabrières, UM ACI 664, 665.

**Description:** No specimen has complete shell. Width > 140 mm, length > 110 mm. At the cardinal border, small beak passing progressively to extended ears. In longitudinal profile:
umbo strongly convex, passing anteriorly to a nearly flat trail. Ornamentation of rounded costae, 9 to 13 counted in 10 mm, or more; 15 on the Vailhan specimen UCBL-FSL 703009. Near the anterior border of the shell, some low broad radiating folds. Rugae well marked on the posterior part of the shell, from ears to umbo, diminishing towards the trail. Spine basis very rare and small on ventral umbo and trail. Shell thickness: on the adult ventral valve ML UCBL-FSL 703011 (Fig. 4(3)) it is 11 mm posteriorly, 1 mm at the anterior border. Ginglymus rarely well preserved. On a ventral valve, UCBL-FSL 703007 (Fig. 4(5)) a marginal area, 3.5 mm wide, is oblique towards the external ear surface; same position as on *Kansuella kansuensis* figured by Chao (1928: text-fig. 6). On a dorsal valve, UCBL-FSL 703015 (Fig. 4(6)) a narrow interarea is obliquely striated. Ventral interior of a relatively young shell (UM ACI 664; Fig. 4(1)): narrow and short adductors surrounded by long flabellate diductors. Dorsal interior: UCBL-FSL 703021 (Fig. 5(5, 6)), trilobed cardinal process, narrow median septum, adductor scars irregularly dendritic; their posterior borders are narrow crests similar to those of an illustrated *Kansuella kansuensis* topotype (Chao, 1927: pl. 9, fig. 2).

**Remarks:** The ginglymus on both valves is a typical kansuellid character. The morphology of *Kansuella* sp. 1 corresponds to both genera: *Kansuella*, with rugae developed posteriorly, and *Parakansuella*, with inflated ventral umbo and trilobed cardinal process. Differences from *K. kansuensis* are a less transverse outline, a more inflated longitudinal profile, and less fine radial costae. The cardinal process on *K. kansuensis* is imperfect, possibly bilobate, having median sulcus (Muir-Wood and Cooper, 1960); on *Kansuella* sp. 1, it is trilobed with a large median lobe wearing a median sulcus (Fig. 5(5)).

*Kansuella* sp. 2

Figs. 4(7, 8), 5(2-4, 7)

**Material:** ML sampling: La Serre vineyard (ML 1176): one specimen UCBL-FSL 703039. Lyon University: Laurens area, two specimens UCBL-FSL 703001, 703003. Montpellier University: Laurens area, one specimen from Albaille collection UM ACI 651, and a big shell UM ACI 650.

**Measurements:** Width, 140 to > 170 mm; length 80 to > 115 mm. Ornamentation of radial costae, 7 to 13 per 10 mm.

**Description:** Outline transverse, longitudinal profile weakly convex. Rugae well marked on ears, weak to absent on umbo. Some spine bases visible on umbo UCBL-FSL 703039 (Fig. 5(7)), as large as one or two radial costae. Shell thickness: on the adult ventral valve
UM ACI 650 (Fig. 5(4)), it is 3 mm near the beak, 12 mm beyond the muscle scars, progressively decreasing to 1 mm at the anterior margin; dorsal valve 3 to 1 mm. The internal cavity is no more than 5 mm thick posteriorly. Ventral area, on UM ACI 651: 3.8 mm wide. Ventral and dorsal ginglymus on sections: UCBL-FSL 703039 (Fig. 4(7)) ventral area 4.5 mm wide, dorsal area *ca.* 1.8 mm. On specimen UM ACI 650 (Fig. 4(8)) the dorsal valve is broken, but its posterior area is still hanging near the ventral one; ventral area 3 mm wide, dorsal area 1 mm. Small borings, 0.3 to 1 mm in diameter, on these shells, well-visible in sections but nearly not externally, since partly covered by superficial test, may be due to predators as worms, bryozoan (Brunton, 1966) or cyanobacteria. Ventral interior of a relatively young shell: UM ACI 651 (Fig. 5(2)), striated diductors scars strongly stinking in the shell. Dorsal interior unknown.

**Remarks:** *Kansuella* sp. 2 differs from *Kansuella* sp. 1 by a more transverse outline, weaker convexity, less developed rugae, and stouter spine basis on ventral valve. These characters are partly typical of *Kansuella sensu stricto* (transverse outline, weak convexity) and *Parakansuella* (rugae poorly developed, stouter spine basis).

Tribe Semiplanini Sarycheva, 1960

**Diagnosis:** See Brunton et al. (2000: p. 556).

**Distribution:** Middle Viséan-lower Serpukhovian.

Genus *Latiproductus* Sarycheva et Legrand-Blain, 1977

**Type species:** *Productus latissimus* Sowerby, 1822; Wales, Brigantian.

**Distribution:** Late Viséan (Brigantian)-early Serpukhovian; Eurasia and northern Africa.

**Diagnosis:** See Brunton et al. (2000: p. 558).

**Remarks:** On small shells (young specimens), it is sometimes difficult to make a generic determination between *Semiplanus* and *Latiproductus*. Specific composition of *Latiproductus*: (i) according to Sarycheva and Legrand-Blain (1977), "In addition to the type species, *S. latipriscus* (Sarycheva, in Sarycheva and Sokolskaya, 1952), and *S. volgensis* n. sp [...] it is possible that we ought also to include in this genus "*Gigantoproductus edelburgensis* (Phillips) [...]"; (ii) Litvinovich and Vorontsova (1991) described 22 species/subspecies of *Latiproductus* from Russia.

*Latiproductus cf. latissimus* (Sowerby, 1822)
Fig. 5(3)
1935. *Productus latissimus* Sowerby – Böhm, p. 161, pl. 8, fig. 18.

**Material:** Montpellier University: Cabrières, a dorsal valve, UM ACI 668 figured by R. Böhm (1935).

**Measurements:** Width: 72 mm, length: 28 mm; 9 costae per 10 mm.

**description** (according to Böhm's text, here translated into English): A dorsal valve, which presents the tapered form characteristic for the species. The surface is ornamented with rather pronounced longitudinal ridges".

**Remarks:** Böhm cites, as reference for his determination, Davidson's (1858-1863) monograph. His figuration of a dorsal valve external mould, which shows a beak nearly flat and regular radial strong costae, corresponds to the genus *Latiproductus*. Without a ventral valve, the specimens cannot be unequivocally attributed to the species *latissimus*. From Cabrières area, several small, recently collected *Latiproductus* shells are not conspecific with *L. latissimus sensu stricto*.

*Latiproductus latissimus complicatus* (Paeckelmann, 1931)

Fig. 6(5)

1931. *Productus* (*Gigantella*) *latissimus* Sowerby, var. nov. *complicata* – Paeckelmann, p. 267, pl. 30, figs. 3-5, pl. 32, figs. 2-4.

1938. *Productus* (*Gigantella*) *latissimus complicatus* (Paeckelmann) – Demanet, p. 82, pl. 8, figs. 6, 7.

1958. *Gigantoproductus latissimus* (Sowerby) var. *complicata* Paeckelm. – Zakowa, p. 59, pl. 1, fig. 13, pl. 2, fig. 1.

1966. *Gigantoproductus latissimus* (Sowerby) var. *complicata* Paeckelm. – Zakowa, p. 84, pl. 12, figs. 5-7.

1986. ?*Semiplanus complicatus* (Paeckelmann) – Zakowa, p. 63, pl. 1, fig. 10, pl. 3, fig. 4.

1987. *Latiproductus latissimus complicatus* (Paeckelmann) – Amler, p. 270.

**Material:** ML collection: Sauvanès creek (ML 1174), one ventral valve UCBL-FSL 703022 (Fig. 6(5)). Lyon University: Vailhan area, UCBL-FSL 703008. Montpellier University: "Vailhan", UM ACI 661.

**Description:** Medium size, width 65-85 mm: less than typical specimens of *L. latissimus* (80-112 mm); straight length: 40-47 mm, curved length: 60-63 mm. Test very thin: 1 mm on ventral trail. Ornamentation: longitudinal costae, 10-12 counted in 10 mm, no rugae. Spine basis prominent on anterior ventral trail, initiating longitudinal plications quite variable in width and thickness.
**Remarks:** The type locality of *P. latissimus* var. *complicata* Paeckelmann lies in SW Poland in upper Viséan folded flysch deposits. After Zakowa (1986), this *L. latissimus* subspecies is in need of a detailed revision.

*Latiproductus* aff. *latissimus complicatus* (Paeckelmann, 1931)

Figs. 4(2), 5(4)

aff. 1931. *Productus* (*Gigantella*) *latissimus* Sowerby, var. nov. *complicata* – Paeckelmann,
267, pl. 30, figs. 3-5, pl. 32, figs. 2-4.

**Material:** ML collection: Péret-Boissière (ML 784), 4 specimens, UCBL-FSL 703042-703045, dorsal valve internal mould. Montpellier University: From «Neffies», UM ACI 659.

**Measurements** (of the biggest specimen, UCBL-FSL 703043): width 72 mm, length 40 mm; shell thickness 1 mm.

**Description and remarks:** Wide umbo, small ears. Ornamentation: 13-15 costae per 10 mm, no rugae. On dorsal valve external mould, ventral spines and plications leave weak traces, visible on Fig. 6(4). The shells have weaker spine basis and longitudinal plications than the *L. latissimus complicatus* specimen (Fig. 6(5)). Hence, they look intermediate between *L. latissimus complicatus* and *L. priscus* (Sarycheva, in Sarycheva and Sokolskaya, 1952; see also Pattison, 1981, and Litvinovich and Vorontsova, 1991). A poorer state of preservation and harder substrate may explain such variability.

*Latiproductus* sp.

Fig. 6(2)

**Material:** ML collection: Castelsec (ML 791), UCBL-FSL 703027; La Serre vineyard (ML 1054) UCBL-FSL 703040. Lyon University: Laurens area, specimen UCBL-FSL 703005. Montpellier University: Laurens, specimen UM ACI 652; Roquemaillère, specimen UM ACI 658; Cabrières, “Chemin de Cabrières à Péret”, specimen UM ACI 667.

**Measurements:** The figured shell measures: width 78 mm, length 38 mm. Some specimens from Cabrières and Roquemaillère are smaller: width < 40 mm.

**Description and remarks:** The ornamentation of the adult shell shows radial costae, 9-11 per 10 mm, and wider than on *L. latissimus*. Rugae faint and rare on ears; spine bases very small, rare on ventral trail, where there are no longitudinal plications, contrarily to *L. latissimus complicatus*. On the smaller specimens, the costae are a little thinner but not so thin as on typical *Semiplanus*.
*Latiproductus edelburgensis* (Phillips, 1836)

Fig. 6(1)

1836. *Productus edelburgensis* nov. sp. – Phillips, p. 214, pl. 7, fig. 5.

1858. *Productus giganteus* Martin var. *edelburgensis* Phillips – Davidson, p. 143, pl. 40, fig. 2.

1952. *Gigantoproductus edelburgensis* (Phillips) – Sarycheva and Sokolskaya, p. 131, pl. 35, fig. 180.

1956. *Gigantoproductus edelburgensis* (Phillips) – Prentice, p. 234, pl. 20, figs. 1, 2; pl. 22, figs. 1, 2.

1981. *Gigantoproductus edelburgensis* (Phillips) – Pattison, p. 2, pl. 3, figs. 1, 2; pl. 9, fig. 6.

1983. *Latiproductus edelburgensis* (Phillips) – Litvinovich and Vorontsova, pl. 2, fig. 4.

1983. *Latiproductus edelburgensis* (Phillips) – Yang et al., pl. 10, fig. 4.

1986. *Latiproductus? edelburgensis* (Phillips) – Zakowa, p. 65 (with references, 1969-1981), pl. 6, fig. 1.

1991. *Latiproductus edelburgensis edelburgensis* (Phillips) – Litvinovich and Vorontsova, p. 37, pl. 8, fig. 2; pl. 25, fig. 1; pl. 26, fig. 2; pl. 27, fig. 1.

2003. *Gigantoproductus edelburgensis* (Phillips) – Chen and Shi, p. 155 (including Chinese references), pl. 7, fig. 16.

2015. *Gigantoproductus edelburgensis* – Qiao and Shen, pp. 130-133, figs. 3, 5 (only stratigraphic and geographical data, no systematics).

**Material:** ML collection: Roque Redonde (ML 1057): a fragmentary ventral and dorsal valve, UCBL-FSL 703037; La Serre vineyard (ML 1152): a ventral valve with cardinal spines, UCBL-FSL 703041. Lyon University: Laurens area, a fragmentary ventral valve UCBL-FSL 703002. Montpellier University: Mas Foulquier, a fragmentary ventral valve and dorsal valve internal mould, UM ACI 655; Vailhan «La Muraille», Böhm Collection, a large-sized ventral valve, UM ACI 656. Note that de Verneuil (in Fournet and Graff, 1849: p. 628) reported *Productus edelburgensis*, Phill. in the limestones of Vailhan.

**Measurements:** The figured specimen, UCBL-FSL 703041 measures: straight length 52 mm, curved length 74 mm, width 102 mm; ventral valve thickness 1 mm. The biggest specimen (UM ACI 656) attains: straight length 84 mm, curved length 115 mm, width > 130 mm. Shell thickness never more than 3 mm.

**Geographical extension of *L. edelburgensis* and allied species:** Late Viséan and early Serpukhovian; NW Europe, N Africa (Algeria, Morocco), Poland, Russia, and China.
**Description:** Ventral valve moderately transverse, ears flattened passing to a large wide umbo; in longitudinal profile, beak moderately curved, trail nearly flat. Ornamentation: radial costae, 6-8 per 10 mm on trail, up to 11 posteriorly; they are separated by wide furrows. Rugae (absent on English specimens, according to Pattison, 1981), faint on ears of Montagne Noire specimens, so as on some East European (Zakowa, 1986) and North African figured shells. Cardinal spines: on UCBL-FSL 703041 (Fig. 6(1)), left ear lateral extremity, there are three oblique cardinal spines, 7-9 mm long; other smaller spine bases on trail of every Montagne Noire ventral valve are variable in abundance and thickness.

**Remarks:** The generic attribution of the species *edelburgensis* to *Latiproductus* has been discussed since Davidson (1858-1863: p. 143, pl. XL, fig. 2) described *Productus giganteus* var. *edelburgensis*. Typical *Latiproductus* characters observed on the Montagne Noire and diverse *edelburgensis* specimens are the thinness of the test and the cardinal process with a very low median lobe (Litvinovich and Vorontsova 1983: pl. 2, fig. 4). The variability of the Montagne Noire specimens concerns size, rugae which are either absent or faint on ears, and spine bases on cardinal area and on ventral trail. The measurements for the holotype of *L. edelburgensis* from Addleburg, N. Yorkshire, Simonstone Limestone, Brigantian (Prentice, 1956: pl. 20, fig. 1) are quite comparable to the specimen UCBL-FSL 703041. The cardinal spines are thinner on the *L. edelburgensis* holotype and thickest and longest on UCBL-FSL 703041, where they are closer to lateral extremities. Other spine bases on ventral trail look quite variable: they are rather coarse, irregularly scattered on the holotype and northern England specimens (Pattison, 1981: pl. 3, fig. 1), or nearly absent on Russian samples (Sarycheva and Sokolskaya, 1952). Differences could possibly be related to different environments.

Several often endemic subspecies of *L. edelburgensis* have been defined: from Russia, *groeberi* Sarycheva, 1928; *orthocostata* Schimansky, 1940; *schaitankaensis* Einor, 1957; *beleutensis* Litvinovich, 1962; *abnormis* and *sinuatus* Litvinovich et Vorontsova, 1991; and from Algeria, *sahariensis* and *subsahariensis* Legrand-Blain, 1980. Allied species are *rectestrius* Gröber, 1909, *praeedelburgensis* Bolkhovitinova, 1938, *shajenwaensis* Ozaki, 1939, *tjulekensis* Galitzkaja, 1977, and *rudicostatus* Litvinovich et Vorontsova, 1991. In the literature, the row of cardinal spines of these subspecies and species is rarely described. It could be due to whether they acted to support the animal to maintain the life position and have snapped off later (Ferguson 1978: p. 43), or the cardinal area was embedded in sediment – alate latiproductids are commonly found on soft substrates.
**5. Discussion**

**5.1. Stratigraphic ranges**

The stratigraphic ranges of gigantoproductids (Fig. 7) can be tentatively correlated to the local foraminiferal biozones A–H (Vachard et al., 2016b) mainly based on a recent sampling in several localities. The data for the specimens in the old collections at Montpellier and Lyon universities are less precise and thus difficult to integrate into the modern precise biostratigraphic framework.

The brachiopods are often sporadically distributed in the studied outcrops, but concentrations in certain beds or units can occur; e.g., in the bioclastic limestones of the La Serre vineyard section (Aretz, 2002b; Aretz and Herbig, 2003). The brachiopod samples could not be precisely correlated to the logs that served to establish the latest microfossil (foraminifera) biozones A–H (Vachard et al., 2016b) and data in Vachard et al. (2017) because single older sampling intervals are today more precisely divided and/or small-scaled horizontal and vertical facies variabilities make the correlation between sample sets/logs difficult and imprecise. Hence, the local stratigraphic ranges of the brachiopod taxa could partly be overestimated, when the correlation is based on larger parts/units of the outcrop and not to specific stratigraphic horizons.

The biozones A–B (Mikhailovian: late Asbian–basal Brigantian) contain *Datangia semiglobosa*, *Latiproductus* aff. *latissimus complicatus*, and *L. edelburgensis*. Nevertheless, it should be noted that these species are not restricted to the Mikhailovian and also occur in younger strata. At Roc de Murviel, in the lower part of the Roque Redonde Fm., *Datangia* (?) sp. and *Latiproductus* sp. are found in upper Mikhailovian and lower Vevanian (early Brigantian) strata. At Roque Redonde, *Latiproductus edelburgensis* occurs in the latest Viséan and earliest Serpukhovian in younger strata of Roque Redonde Fm. The early Serpukhovian at Roquemaillère contains *Balakhonia* aff. *ostrogensis* and *Kansuella* sp. 1. A long stratigraphic record is found at Castelsec (latest Viséan–late Serpukhovian), where *Datangia semiglobosa* and *Kansuella* sp. 1 are present. Since most of the samples are from the Castelsec North section of Vachard et al. (2016b), their stratigraphic range can be reduced to the Steshevian-Protvian (biozones G–H). This is further confirmed by the presence of the same assemblage in the early Protvian of the Cabrières area. The most diverse assemblage is found in the Laurens area in the Steshevian, and at the La Serre vineyard in the Steshevian and early Protvian. It is composed of *Kansuella* sp. 1, *K*. sp. 2, *Latiproductus latissimus complicatus*, and *L. edelburgensis*, to which can be added the single specimen of *Globosoproductus* sp. in the Laurens area. The most characteristic gigantoproductid
brachiopod for that younger stratigraphic period is *Kansuella* sp. 2, which is relatively
abundant, especially in the Laurens area.

It can be noted that the local stratigraphic ranges of the gigantoproductids correspond
well to the ranges known in other regions, and hence they can be considered as useful
additional biostratigraphic markers. It should be noted that the record of *Datangia
semiglobosa* in the Serpukhovian at Castelsec and Cabrières are among the youngest for this
species. In the future, more work wil have to be done on the kansuellids in Europe in order to
estimate their biostratigraphic value, which according to the Montagne Noire data might be
important.

## 5.2. Palaeoecology

Studies in the British Isles showed the abundance of gigantoproductids in bioclastic,
often coarse-grained limestones in normal saline marine settings, usually considered to
represent high-energy environments (Ferguson, 1978; Angiolini et al., 2012; Nolan et al.,
2017). Gigantoproductids as suspension feeders lived semi-infaunal and extended ears and
postero-ventral shell thickenings helped to stabilize and orientate the brachiopod shell on the
substrate and in the bottom currents, which also allowed constant and sufficent nutrient and
oxygen supply necessary to attain the large shell sizes (Ferguson, 1978; Nolan et al., 2017).
Pérez Huerta and Sheldon (2006) suggested that productid brachiopods were larger in deeper
water with less food supply because of the efficiency of their lophophore. However, this
group of brachiopods is not limited to high-energy settings, and is also found in calmer and/or
deeper settings, as seen in pockets within mud-mounds (Gutteridge 1990), or muddy inner- to
middle shelf settings (Denayer et al. 2016; Nolan et al., 2017).

Gigantoproductids can be found in the Montagne Noire in a broad range of
environments ranging from shallow well-oxygenated mid-platform carbonates to dysoxic
outer platform microbial mounds (Fig. 8). The outer platform and slope microbial mounds
(Vachard et al., 2017) grew during the Venevian, Tarusian and Steshevian. The microbial
mounds, mid-platform grainstones and coral boundstones are known during the Protvian.
Gigantoproductids are known in the core facies of the microbial mounds and coral patch
reefs, but they become more abundant in the surrounding flank and off-mound/reef facies.
The presence of gigantoproductids, in those microbial mounds considered to have developed
in somewhat oxygen depleted conditions, is in contradiction to the general ecological
constraints of a steady flow of nutrients and oxygen (see above). It could be linked to small
shallowing events related to general sea-level fluctuations or mound growth, which shortly
opened suboptimal ecological conditions for gigantoproductids. Another solution would be to
place at least some of the microbial mounds in more shallow water of the middle ramp.

Gigantoproductids are most abundant and diversified in shallow water, bioclastic facies
without any major bioconstructional activity. This observation may cast some doubts about
the requirement of a firm substrate for gigantoproductids, since the reefal environments
should be those with the fastest cementation and hence provide good conditions for initial
larval settlements. However, Nolan et al. (2017) demonstrated that the initiation of the
gigantoproductid shell bed in the Ricklow Quarry (Derbyshire carbonate platform, UK)
started on a somewhat muddier substrate. The English examples also show that maximum
abundance of the large brachiopods was reached in the grain-supported facies, which would
confirm that a muddy substrate is not optimal for gigantoproductids, but that they can cope
with it at least to some extent.

Poty et al. (2002) cited brachiopods and corals in two facies: Roc du Cayla and Castelsec facies. The “Roc du Cayla Facies” consists of microbial boundstones with corals in living position and stromatactis-like cavities. That facies is also cited at Roc de Murviel, where fragments of diverse small and medium-sized productids and spiriferids occur, with a few undetermined fragments of latiproductids. Concentrations of brachiopods in pockets in the mound facies as seen in the Derbyshire mud-mounds (Gutteridge, 1990) have not been observed. In the Montagne Noire, the distribution is more heterogeneous and erratic. Nolan et al. (2017) reported at Ricklow Quarry the influence of an older mound topography on gigantoproductid concentrations, which formed in the shelter of remanent topographic highs. This kind of connection is unknown in the Montagne Noire because mound facies are mostly known as fragments in olistolithic deposits.

Grainstones–wackestones of the “Castelsec Facies” contain calcareous algae, oncolites,
corals, and brachiopods. In the Castelsec quarry, brachiopods are found in a shell bed above a
small coral-patch reef (Aretz, 2002a) with the corals *Lithostrotion, Siphonodendron* and
*Palaeosmilia*. Here, *Datangia semiglobosa, Kansuella* sp. 1 and *Latiproductus* sp. are
associated with various small productids and spiriferids. The brachiopod ventral valves are
mainly isolated and broken, indicating high-energy levels. Álvaro et al. (2007) highlighted the
importance of microbial incrustations in this shell bed. A facies more or less similar to the
Castelsec facies is present at Roque Redonde in a shell bed, represented by a 1.5 m horizon
rich in mostly fragmented and/or disarticulated bryozoans (Ernst et al., 2015), echinoderms,
brachiopods, and corals. Among a diversified assemblage of productids, spiriferids, and
rhynchonellids, a single large fragment of *Latiproductus edelburgensis* (UCBL-FSL 703041)
has been found. All specimens are deposited as isolated valves, sometimes broken, indicating
deposition far from the living position, in shallow agitated waters. The low abundance of
gigantoproductids in the Montagne Noire shell beds clearly differentiates these shell beds
from the true gigantoproductid shell beds described in South China (Yao et al., 2016) and
England (Nolan et al., 2017).

At La Serre vineyard, Aretz (2002b) described five facies: coral limestones, bioclastic limestones, microbial carbonates, dolostones and siliciclastic rocks. Brachiopods are found in bioclastic limestones, associated with some solitary corals and gastropods (Aretz, 2002b). Microfacies analysis suggests deposition in relatively shallow water (Aretz, 2002b; Vachard and Aretz, 2004). On the other hand, the preservation of *Kansuella* sp. 2 with two joined valves (Figs. 4(7), 5(7)) and that of *Latiproductus edelburgensis* with stout cardinal spines (Fig. 6(1)), both suggest a low-energy environment, possibly in a shallower part of the platform. Occurrences in the Belgian Royseux section (Aretz, 2001, 2002b; Denayer et al., 2016) may at least partly represent similar settings.

Another facies, the Laurens Facies, not yet described, occurs near Laurens-Gare, and
has yielded numerous *Kansuella* sp. 1 and *K*. sp. 2 shells preserved with both valves joined
(Figs. 4(3-6), 5(5, 6)). Probably, their thick heavy ventral valve slightly sunken in a loamy
bottom provided good stability. They are associated with a few undetermined semireticulate
productids and rhynchonellids, together with some bivalves and gastropods in dark grain-
supported bioclastic limestones, but almost no corals. The matrix around a *Latiproductus
edelburgensis* specimen (UCBL-FSL 703002) might indicate a deposition in the littoral-
intertidal zone. As a consequence, the Laurens facies reflects deposition in a shallower part of
the platform.

In Hunan (South China), a brachiopod-coral (*Kansuella-Yuanophyllum*) community has
been described in bioclastic limestones with intercalated mudstones of the late Viséan (Yang
et al., 1989). The living conditions are a firm substrate in shallow-clear water with fairly
strong current action. The big gigantoproductids had strong and massive shells with a thick
ventral umbo. They lived with the ventral umbo lying in the bottom, and the long trail of the
shell extending obliquely upwards. The great weight of the ventral valve helped to hold the
shell in an advantageous position for feeding. In the Montagne Noire, similar environmental
conditions for brachiopods can be postulated for the assemblages containing *Kansuella*
species, even if a brachiopod-coral community cannot be recognized. Co-occurrences of
kansuellids with corals are restricted to shell beds composed of various fragments indicating
mixture of different habitats. Corals are absent or very rare in assemblages with well-
preserved kansuellids at Laurens and La Serre. There, the gigantoproductids are associated
with gastropods, bivalves and crinoids.

5.3. *Palaeobiogeographical connections*

The composition of the Montagne Noire gigantoproductid fauna may be compared to
gigantoproductid assemblages in Western, Central and Eastern Europe and North Africa
(Figs. 9, 10; Table S1, Appendix A) in order to investigate palaeobiogeographical connections
in the western Palaeotethys. Even if the Alleghanian Isthmus was still open (Davydov and
Cózar, in press), gigantoproductids did not migrate through this seaway into the cratonic
basins of southwest Laurussia (North America). This restriction is not known in nektonic and
benthic vagile organisms as ammonoids (Korn et al., 2012), and was admitted for
microgranular calcareous foraminifers (Davydov and Cózar, in press), among which better
faunal exchange between North America and the Palaeotethys seemed to exist. However, it is
more comparable to rugose corals, another group of sessile organisms showing few
similarities between North America and western Palaeotethyan taxa (Poty, 2002).

5.3.1. *Northern Variscides*

The nearest outcrop containing gigantoproductids towards the north is found in the
northern Massif Central (France). At L'Ardoisière, near Vichy, Julien (1896) described
*Productus giganteus* in arenitic fossiliferous beds from the middle Viséan. Those specimens
belong to *Globosoproductus* (Legrand-Blain, 2018), and their relationships to the younger
*Globosoproductus* from the Laurens railway trench are unknown, especially because the
middle Viséan deposits of the Montagne Noire are devoid of macrofossils (Aretz, 2016).

The upper Viséan (Asbian-Brigantian) strata of the British Isles contain diversified assemblages of gigantoproductid brachiopods (Pattison, 1981). Only *Datangia semiglobosa* and *Latiproductus edelburgensis* are shared with the Montagne Noire. A further faunal link might be expressed in the shared occurrence of the genus *Kansuella*, but more detailed work has to be done at the species level. In contrast, *Gigantoproductus sensu stricto* (characterized by an ornamentation of flutings; Pattison, 1981), locally proliferating during the late Asbian and early Serpukhovian in the British Isles, Belgium (Denayer et al., 2016) and the Rhenish Mountains (Winkler Prins and Amler, 2006), is absent in the Montagne Noire. This implies an important barrier for faunal exchange between the northern and southern Variscides in late Viséan-early Serpukhovian times, which is also seen in calcareous algae (Vachard et al., 2006), foraminifers (Cózar et al., 2019), rugose corals (Aretz, 2002b), bryozoans (Ernst et al.,
2015), and ammonoids (Korn and Feist, 2007). So far, the faunal connections between Belgium and the Rhenish Mountains to the Montagne Noire are restricted to some semiplanids, notably *Latiproductus latissimus complicatus*, which is found in both regions. However, modern taxonomic descriptions of the relatively diverse Viséan faunas of Belgium (Demanet, 1958) are lacking, and thus palaeobiogeographical relations may be obscured. The current absence of *Kansuella* in Belgium and the Rhenish Mountains may be related to absence of sufficiently preserved material, which prevents the identification of this genus. Gigantoproductids are also reported from the western and eastern end of the Northern Variscides. Data from the Ossa Morena in SW Spain (Martínez Chacón and Legrand-Blain, 1992) are scarce, and the fauna is relatively poorly diversified. So far, common taxa with the Montagne Noire are *Datangia semiglobosa* and *Balakhonia ostrogensis*, although the Spanish record for the latter is slightly questionable (aff.). The Sudetes, SW Poland, have yielded a medium diverse fauna of large productids. They share with the Montagne Noire *Datangia semiglobosa* and *Latiproductus latissimus complicatus*; however, this fauna contains a series of taxa unknown in the Montagne Noire.

### 5.3.2. *Southern Variscides*

Reconstructions of the European Variscides attribute the Montagne Noire, the Pyrénées, and the Cantabrian Mountains to the Gondwanan part of the orogenic belt (Faure et al., 2008; Franke, 2014; Aretz, 2016). However, the Pyrénées and Iberia are located rather far from the Montagne Noire and are separated by the important Alpine North Pyrenean Fault system. Only two regions contain Mississippian gigantoproductids. The fauna of the Arize Massif (Legrand-Blain et al., 1996), from contemporaneous deposits as the Montagne Noire, is less diverse and restricted to *Latiproductus latissimus* and *L. cf. priscus*. In the Central Pyrénées, the Ardengost Limestone of Serpukhovian age (Perret and Vachard, 1977) contains an assemblage of *Gigantoproductus superior*, *G. cf. superbus*, *Datangia pyrenaea*, *Titanaria* ex gr. *africana* and *Semiplanella* spp. (Legrand-Blain et al., 1983). Hence, there is no overlap with the fauna of the Montagne Noire. Facies differences between the two regions as controlling factor can be ruled out. The Ardengost fauna should be considered to be younger than those of the “Calcaire à *Productus*”; either Zapaltyubinian (Delvolvé et al., 1996) or Protvian (by comparison with Vachard et al., 2016b). In fact, the olistostrome of Ardengost might include olistolites of different ages; some rich in foraminifers and algae of probably late Protvian age, whereas others, with the brachiopods *Gigantoproductus* and *Titanaria*, are possibly latest Serpukhovian, Zapaltyubinian, or even early Bashkirian in age (this study), in
accordance with the ages of the *Titanaria* beds in North Africa (Legrand-Blain, 1985; Cózar et al., 2016).

Thus, the presence of the genera *Gigantoproductus* and *Titanaria* at Ardengost would indicate that the late Viséan-early Serpukhovian faunal barrier between the northern and southern Variscides for these genera broke down in the late Serpukhovian and those genera colonised the French part of the Gondwana realm.

From the Variscan units incorporated into the Alpine Orogen, gigantoproductids among a relative diverse brachiopod fauna have been reported from Nötsch in southern Austria (Aigner 1930, 1931). They are in need of taxonomic revision and hence are difficult to compare to the Montagne Noire fauna.

### 5.3.3. *North Africa*

Gigantoproductids in the cratonic Saharan basins in Algeria are best studied in the Béchar Basin (Legrand-Blain, 1985, 1987). They are more abundant and diverse than in the Montagne Noire, with the notable presence of the genus *Gigantoproductus*. The presence of *Kansuella* is herein confirmed for two species (see above chap. 3). In the Béchar Basin, the Viséan assemblages comprise *Datangia menchikoffi*, *Kansuella meharezensis*, and *Latiproductus subokensis*; and those of the lower Serpukhovian, *Datangia* spp., *Gigantoproductus meridionalis*, *Kansuella flamandi*, *Latiproductus* spp., and *L. edelburgensis sahariensis*.

In Morocco, Bensaid et al. (1985) pointed to major geological differences between the central Moroccan Variscides (Moroccan Meseta) and the unfolded “Presahara” (Ouarkziz and Tafilalt). They cited *Gigantoproductus* in the late Viséan of both regions, but these gigantoproductid brachiopods have been rarely described and are in need of taxonomic revisions. From the central Moroccan Variscides, mainly the Meseta, Termier (1936) and Termier and Termier (1950) published gigantoproductid faunas, which according to Legrand-Blain (1973, 1980) are related to *Datangia menchikoffi*, *Latiproductus subokensis*, and *L. ? daguini*. Dedok and Hollard (1980) described in the Akerchi section (northern Meseta) Viséan brachiopods including *Gigantoproductus*, *Neospirifer cf. fascicostatus*, and *Titanaria cf. africana*. From the late Viséan (Asbian) Zrigat Fm. in the Tafilalt, Mottequin et al. (2017) described *Latiproductus* cf. *subokensis* and *L. cf. edelburgensis sahariensis*, indicating the close proximity to the Béchar basin faunas. The faunal differences between the Meseta and the Tafilalt support the separation of the epicontinental Saharan basins and the mobile Variscan zones of central Morocco.
The illustrations of the Moroccan productids do not confirm the presence of *Gigantoproductus sensu stricto* in this region, which is in marked contrast to the Algerian basins. Without any detailed taxonomic work, it is impossible to determine if *Kansuella* is present or not in Morocco. In any case, the Moroccan and Algerian data indicate only weak overlap between the gigantoproductid faunas of North African and the Montagne Noire, and advocate for a palaeobiogeographical barrier between them.

### 5.3.4. Russian Platform

Abundant and diverse gigantoproductid assemblages have been described from the Viséan and Serpukhovian strata of the Moscow Basin (Sarycheva, 1970; Lazarev, 1990) and the Donbass (Litvinovich and Vorontsova, 1991). The total number of taxa is by far the highest in the entire western Palaeotethys. These assemblages contain a series of endemic species with limited information about intraspecific variability, alongside more widespread, almost cosmopolitan species. This is well seen in the species shared between the Moscow Basin and the Central and Western European platforms north of the Variscan Orogen (Wilson, 1989). The overlap to the Montagne Noire fauna is based on this type of widely distributed species as evidenced by *Balakhonia ostrogensis*, *Datangia semiglobosa*, and *Latiproductus edelburgensis*. These species do not allow indicating palaeobiogeographical proximity. In the context of the Montagne Noire assemblages, the presence of twelve species of *Gigantoproductus s.s.* on the Russian Platform has to be highlighted.

### 5.3.5. Central and Eastern Palaeotethys

Gigantoproductids are found in several diverse assemblages in the central and eastern Palaeotethys. Most of the taxa have local or regional distributions, but some of them have a much wider distribution, if not to say a cosmopolitan character.

In the Urals, *Datangia semiglobosa* is recorded under the name *Moderatoproductus semiglobosus* in the late Viséan Gubachkinskii-Ladeininsky horizons and *Latiproductus edelburgensis* in the Serpukhovian Ust-Sarbayskii horizon (Litvinovich and Vorontsova, 1991). It is important to note that so far *Kansuella* is not identified in the Urals (see situation in Belgium).

In the eastern Palaeotethys, a large number of gigantoproductid species (Qiao and Shen, 2015; Shen et al., 2017) have been described. Of special interest is the genus *Kansuella*. In South China, a *Kansuella*-Yuanophyllum community has been described (Yang et al.,
1983, 1989). In this community, *Kansuella* is associated with *Latiproductus latissimus* and *L. edelburgensis*, which are also known in the Montagne Noire.

### 5.3.6. *Quantitative analyses*

The results of the quantitative analyses (Fig. 10) have to be taken into consideration with much care. The confidence level of the individual clusters is often not very good, which roots in the uneven distribution of the data (many species with occurrences in few regions; Russian singletons). However, it is worth seeing that, independently from the used methods, some consistent clusters emerge. Some clusters regroup geographically adjacent regions, but others are more heterogeneous. Comparable clusters can be found when the analyses are conservatively run at the generic level, which should reduce the influence of taxonomic uncertainties.

A first cluster comprises the Belgian, German (Rhenish Massif) and Polish faunas, and therefore the three regions of the Rhenohercynian Realm *sensu stricto*. The Belgian fauna, occurring in the immediate westward prolongation of the Rhenish Massif, should be closer to the Rhenish fauna than those from Poland. However, the closeness between the Polish and German faunas may be exaggerated since it seems to follow the initial regional study of Paeckelmann (1931) who used material from both regions. The North African faunas form a second cluster, which is also visible in the NMDS ordination. Their proximity and isolation from Europe is coherent with palaeogeographical reconstructions (e.g., Simancas et al., 2005). However, the palaeobiogeographical closeness of the Saharan platform and the Moroccan Meseta needs further evaluation due to the lack of data for the Moroccan Meseta. The cluster of the Nötsch Palaeozoic (Austrian Alps) and the Pyrénées is more difficult to interpret. It groups two faunas along the southern shelf of the Variscides, but the closeness is only based on the co-occurrence of two widely distributed taxa and the low overall diversity at Nötsch. Hence, this artificial affinity is also seen in the minimum spanning tree in the NMDS ordination, where the two regions are not connected. A similar artificial connection is the clustering of Montagne Noire and Ossa-Morena Zone in the cluster analysis using the Raup-Crick coefficient. This is not supported by the results of the other analyses, and it is only based on the low diversity and one widely-known taxon in the Ossa-Morena Zone. Hence, these two clusters are more the expression of poor taxonomic data than palaeobiogeographical connections.

The position of the British Isles and Russian Platform in the cluster analysis differs. Using the Jaccard similarity: these two well-studied regions with individual high diversities
are regrouped in a single cluster, which is attached to the Rhenohercynian cluster (see above).
This clustering represents the faunal exchange in the northern Variscan seaway already
identified in earlier studies (Wilson, 1989). However, when using the Raup-Crick coefficient,
this cluster is broken. The British Isles remain near the Rhenohercynian cluster, but the
Russian Platform looses palaeobiogeographic proximity. The minimum spanning tree in the
NMDS ordination partly explain these differences: the regions of the northern Variscan
seaway can be well identified, but they are all only connected via a central node, namely
Poland.

Overall, the combined results of the cluster analyses and NMDS ordination show
North Africa as the most stable palaeobiogeographical entity. The position and connections of
the European units are less stable, although the seaways of the northern Variscides can be at
least partly delimited in all analyses. It remains difficult to establish palaeogeographical
barriers and resulting palaeobiogeographical units based on the spatio-temporal distribution of
gigantoproductids around the Variscan Orogen or the western Palaeotethys. The influencing
factors are discussed in more detail in the following Section 5.3.8. However, for the moment,
the data are too unevenly distributed in term of abundance, taxonomic reliability and
stratigraphic quality for fine-scale quantitative analyses.

### 5.3.7. Larger palaeobiogeographical framework

The presence of *Kansuella* as a typical eastern Palaeotethyan taxon, in the western Palaeotethys including the Montagne Noire, points to important long-distance faunal connections through the entire Palaeotethys. This is supported by the occurrences of *Latiproductus latissimus* and *L. edelburgensis* in South China. A main westward-directed migration path may be assumed from South China along (micro)continents of the southern Central Asian Orogenic Belt and the Urals to the eastern margin of Laurussia (Russian Platform). The absence of *Kansuella* in the Urals and on the Russian Platform is herein interpreted as a problem of taxonomic identification.

West of the Russian Platform, two migration paths exist along the seaways north (see Wilson, 1989) and south of the Variscan Orogen. Faunas in Poland, Germany, Belgium, Britain and SW Spain share abundant similarities (e.g., *Gigantoproductus sensu stricto*), but there is no uniformized northern Variscan gigantoproductid fauna, most probably due to differences in facies, stratigraphic ages and taxonomic discrepancies. This northern seaway was gradually closed during the late Visean and early Serpukhovian. The remnants of the seaway south of the Variscan Orogen are found in the Alpine Domain, southern France, and
northern Spain. The connections within this seaway are less clear, since the gigantoproductid occurrences are rarer and more patchy, which increases the uncertainties introduced by the quality of the taxonomic data. However, the data for southern France show that, around the early/late Serpukhovian boundary, a palaeobiogeographical barrier broke down and genera such as *Gigantoproductus sensu stricto* and *Titanaria* migrated into this area.

The North African gigantoproductid faunas are somewhat an endmember of the migration through the Palaeotethys and comprise widespread and local taxa. In general, during late Viséan and early Serpukhovian times, the North African faunas are far better connected to the northern than to the southern Variscides. The barrier towards the southern Variscides, and hence, the Montagne Noire has still to be located. On the regional scale, the relationships between the cratonic and mobile-Variscan parts have to be better elucidated.

### 5.3.8. *Influencing factors for the distribution of gigantoproductids.*

Although the general pattern developed for the migration paths through the Palaeotethys seems to be straightforward (Qiao and Shen, 2015) to explain the tempo-spatial distribution of gigantoproductid brachiopods, there still exist many uncertainties. As shown in the systematic palaeontology part of this study, different taxonomic concepts exist for the group, which complicate comparisons even at higher taxonomic levels. For many taxa, the inter- and intraspecific variability is (almost) unknown and hence oversplitting or lumping of taxa cannot be consistently ruled out. Additionally, modern systematic works are unevenly distributed in the Palaeotethys. Faunas of some regions have not been systematically revised for many decades or they have been only insufficiently described.

Oceanic currents seem to have influenced the migration and, as a consequence, the tempo-spatial distribution of gigantoproductids. However, this general idea is at best only roughly backed by the stratigraphic data, since those data are missing or largely outdated for many taxa. Hence, it is currently almost impossible to indicate a main migration route to the west and to locate speciation centres in the eastern Palaeotethys. Palaeoenvironmental factors as substrate or turbulence have been evoked to control the spatial distribution of gigantoproductid taxa. These factors could explain some of the local differences in the composition of contemporaneous assemblages in the Montagne Noire (see Section 5.2). However, even on larger scales these differences are not levelled out since there are major facies differences in various regional tectonic settings; e.g., between the epicontinental basins of the Sahara and the carbonate deposits under siliciclastic influx in the Variscan foreland basins.
The differences between the assemblages at Ardengost (Pyrénées) and in the Montagne Noire seem to be related to the breakdown of a palaeobiogeographical barrier in the late Serpukhovian. Thus, restricting the analysis to specific ages becomes even more important, but, as stated before, this will require a better stratigraphic framework for many species.

**6. Conclusions**

In the Montagne Noire, gigantoproductids are locally abundant in the "Calcaire à *Productus*", but not necessarily diverse. The fauna is composed of well-known taxa relatively widely distributed in the western, central and eastern Palaeotethys; possible endemic species to the Montagne Noire are only described in open nomenclature and require further, and better preserved material for detailed study as illustrated by the species attributed to *Kansuella*. Compared to the contemporaneous Viséan-Serpukhovian faunas of Poland, the British Isles and Russia, the gigantoproductid faunas of the Montagne Noire are less diverse. Differences between the vast and variegated carbonate platform around Laurussia and the more patchy occurrences of carbonates in the tectonically active settings of the Montagne Noire could provide some explanations. Especially, the total absence of the genus *Gigantoproductus*, which is predominant with diverse species in Western and Central Europe, is noteworthy. It advocates for a palaeobiogeographical barrier, which broke down in the late Serpukhovian and enabled the migration of *Gigantoproductus* and *Titanaria* into the French part of the Gondwanan realm. To further improve the knowledge of the Montagne Noire Carboniferous brachiopods, it remains necessary to study the non-gigantoproductid faunas, especially the various smaller productids, chonetids and spiriferids. This could lead to a synthesis of Carboniferous brachiopods in Central and Southern France, which would contribute to a better knowledge of biogeographical events during the Variscan orogeny.

**Acknowledgements**

Eddy Poty is thanked for participating in brachiopod collecting in the field. Suzanne Jiquel (Montpellier University) furnished photographs of figured *P. latissimus* in Böhm's collection. Comments and suggestions of Shuzong Shen, Qiao Li, Miguel Angel Torres Martinez, Gabriela Cisterna, Bertrand Lefebvre, Gilles Escarguel and an anonymous reviewer greatly helped to improve the manuscript. M. Legrand-Blain thanks Denis Bates for providing advice on the English language.
References

Abramov, B.S., 1970. Biostratigrafiya kamennougolnykh otlozhenii Sette-Dabana (Yuzhnoie Verkoianyie [Biostratigraphy of the Carboniferous deposits of Sette-Daban (Southern Verkhoiansk)]. Akademiya Nauk SSSR, Iakutskii Filial Sibirskogo Otdeleniia Institut Geologii. Moscow, 1-178 (in Russian).

Aigner, G., 1930. Die Brachiopoden des Karbons von Nötsch im Gailtal; 1. Teil. Mitteilungen naturwissenschaftlicher Verein der Steiermark 67, 9-42.

Aigner, G., 1931. Die Brachiopoden des Karbons von Nötsch im Gailtal; 2. Teil. Mitteilungen naturwissenschaftlicher Verein der Steiermark 68, 3-15.

Álvaro, J.J, Aretz, M., Boulvain, F., Munnecke A., Vachard D., Vennin E., 2007. Fabric transitions from shell accumulations to reefs: an introduction with Palaeozoic examples.
In: Álvaro, J.J, Aretz M, Boulvain F., Munnecke A., Vachard D., Vennin E. (Eds.), Palaeozoic Reefs and Bioaccumulations: Climatic and Evolutionary Controls. Geological Society, London, Special Publications 275, 1-18.

Amler, M.R.W., 1987. Fauna, Paläogeographie und Alter der Kohlenkalk-Vorkommen im Kulm des östlichen Rheinischen Schiefergebirges (Dinantium). Geologische Abhandlungen Hessen 88, 1-338.

Angiolini, L., Stephenson, M.H., Leng, M.J., Jadoul, F., Millward, D., Aldridge, A., Andrews, J.E., Chenery, S., Williams, G., 2012. Heterogeneity, cyclicity and diagenesis in a Mississippian brachiopod shell of palaeoequatorial Britain. Terra Nova, 24, 16-26.

Aretz, M., 2001. The upper Viséan coral-horizons of Royseux – The development of an unusual facies in Belgian Early Carboniferous. Bulletin of the Tohoku University Museum 1, 86-95

Aretz, M., 2002a. Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France). Geobios 35, 187-200.

Aretz, M., 2002b. Habitatanalyse und Riffbildungspotential kolonialer rugoser Korallen im Unterkarbon (Mississippium) von Westeuropa. Kölner Forum für Geologie und Paläontologie, Geologisches Institut der Universität zu Köln 10, 1-155.

Aretz, M., 2016. The Kulm Facies of the Montagne Noire (Mississippian, southern France). Geologica Belgica 19, 69-80.

Aretz, M., Herbig, H.G., 2003. Contribution of rugose corals to Late Visean and Serpukhovian bioconstructions in the Montagne Noire (Southern France). In: Ahr, W. A., Harris, P. N., Morgan, W. A., Somerville, I. D. (Eds.), Permo-Carboniferous
Carbonate Platforms and Reefs. Society of Economic Paleontologists and Mineralogists, Special Publication 78, 119-132.

Barchy, L., 1994. Contribution à l'étude des Tétracoralliaires dans les « Calcaires à *Gigantoproductus* » des écailles de Cabrières (Montagne Noire, France). Mémoire présenté en vue de l'obtention du grade de licencié en sciences géologiques et minéralogiques, Université de Liège (unpublished).

Bensaid M., Termier H., Termier G., 1985. The Moroccan Variscides. In: Wagner, R.H., Winkler Prins, C.F., Granados, C.F. (Eds.), The Carboniferous of the World: Australia, Indian subcontinent, South Africa, South America & North Africa. IGME, Madrid, pp. 336-344.

Bergeron, J., 1889. Etude géologique du massif ancien situé au Sud du Plateau Central. Annales des Sciences géologiques 22, 1-362.

Böhm, R., 1935. Etudes sur les faunes du Dévonien supérieur et du Carbonifère inférieur de la Montagne Noire. Thèse, Imprimerie de la Charité, Montpellier.

Bolkhovitinova, M. A., 1938. Ecologiya, paleogeografiya i stratigraficheskaya tsennosty *Gigantella* tulskoy tolschi Podmoskovnogo basseina [Ecological, palaeogeographical and stratigraphical value of *Gigantella* in the Tulskian beds of Moscow Basin]. Trudy Moskovskiy Geologorazvedochnyy Institut 12, 201-260 (in Russian).

Brunton, C.H.C., 1966. Predation and shell damage in a Viséan brachiopod fauna. Palaeontology 9, 355-359.

Brunton, C.H.C., Lazarev, S.S., Grant, R.E., 1995. A review and new classification of the brachiopod order Productida. Palaeontology 38, 915-936.

Brunton, C.H.C., Lazarev, S.S., Grant, R.E., Jin Y.-G., 2000. Productidina. In: Kaesler, R.L. (Ed.). Treatise on Invertebrate Paleontology Part H Brachiopoda Revised vol. 3 Linguliformea, Craniiformea, and Rhynchonelliformea part. Geological Society of America. University of Kansas, Boulder, CO and Lawrence, KA, pp. 424-619.

Cascales-Miñana, B., 2010. Testing similarity coefficients for analysis of the fossil record using clustering methods: the Palaeozoic flora as a study case. Revista Española de Paleontología 25, 19-34.

Chao, Y.T., 1927. Productidae of China, part I: Producti. Palaeontologia Sinica B 5, 1-192.

Chao, Y.T., 1928. Productidae of China, part II: Chonetinae, Productidae and Richthofeniinae. Palaeontologia Sinica B 5, 1-103.
Chen, Z.Q., Shi, G.R., 2003. Early Carboniferous brachiopod faunas and their biogeographical affinities from the western Kunlun Mountains, North-west China. Palaeontographica A 268, 103-187.

Clarke, K. R., 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18, 117-143.

Cózar, P., Izart, A., Somerville, I.D., Aretz, M., Coronado, I., Vachard, D., in press. Environmental controls on the development of Mississippian microbial carbonate mounds and platform limestones in southern Montagne Noire (France). Sedimentology, DOI: 10.1111/sed.12594.

Cózar, P., Vachard, D., Aretz, M., Somerville, I.D., 2019. Foraminifers of the Viséan–Serpukhovian boundary interval in Western Palaeotethys: a review. Lethaia 52, 260-284.

Cózar, P., Somerville, I.D., Vachard, D., Coronado, I., Garcia-Frank, A., Medina-Varea, P., Said, I., Del Moral, B., Rodriguez, S., 2016. Upper Mississippian to lower Pennsylvanian biostratigraphic correlation of the Sahara Platform successions on the northern margin of Gondwana (Morocco, Algeria, Libya). Gondwana Research 36, 459-472.

Davidson, T., 1858-1863. A monograph of British fossil Brachiopoda, Carboniferous, vol. 2, part 5. Palaeontographical Society Monograph London, 1-280.

Davidson, T., 1880. A monograph of British fossil Brachiopoda. Supplements, vol. 4 part.3. Supplement to the Permian and Carboniferous species. Palaeontographical Society London, 243-316.

Davydov, V.A., Cózar, P. in press. The formation of the Alleghenian Isthmus triggered the Bashkirian glaciation: Constraints from warm-water benthic foraminifera. Palaeogeography, Palaeoclimatology, Palaeoecology. DOI: 10.1016/j.palaeo.2017.08.012.

Dedok, T.A., Hollard, H., 1980. Brachiopodes du Carbonifère inférieur du Maroc central. Notes et Mémoires du Service géologique du Maroc 41, 185-230.

Delvolvé, J.J., Babin, C., Gandl, J., Kullmann, J., Laveine, J.P., Legrand-Blain, M., Mac Kinney, F.K., Martinez-Chacon, M.L., Perret, M.F., Semenoff-Tian-Chansky, P., Sevastopulo, G., Vachard, D., 1996. Carbonifère à faciès Culm. In: Barnolas, A., Chiron J.C., Guérangé, B. (Eds.), Synthèse géologique et géophysique des Pyrénées. Vol. 1, introduction, géophysique, cycle hercynien. Bureau de Recherches Géologiques
et Minières (BRGM) Paris and Instituto Tecnológico GeoMinero de España (IGME)
Madrid, 303-338.

Demanet, F., 1938. La faune des Couches de passage du Dinantien au Namurien dans le synclinorium de Dinant. Mémoires du Musée Royal d'Histoire Naturelle de Belgique 84, 1-201.

Demanet, F., 1958. Contribution à l'étude du Dinantien de la Belgique. Mémoires de l'Institut royal des Sciences naturelles de Belgique 141, 1-152.

Denayer, J., Aretz, M., Poty, E., Mottequin, B., 2016. Royseux: A palaeobiodiversity hotspot in the late Viséan (Carboniferous) of Belgium. Geologica Belgica 19, 7-20.

Deshayes, G.P., Milne Edwards, H., 1836. Histoire naturelle des animaux sans vertèbres. Meline, Cans et compagnie, Bruxelles.

Einor, O. L., 1957. K faune kamennougol'nykh brakhiopod zapadnogo sklona Srednego Urala [On the Lower Carboniferous Brachiopod Fauna of the Western Slope of the Central Urals.] Ezhegodnik Vsesoiuznogo Paleontologicheskogo Obshchestva (Leningrad) 16, 142-161 (in Russian).

Engel, W., Feist, R., Franke, W., 1978. Synorogenic gravitational transport in the Carboniferous of the Montagne Noire (S-France). Zeitschrift der deutschen geologischen Gesellschaft 129, 461-472.

Engel, W., Feist, R., Franke, W., 1982. Le Carbonifère anté-Stéphanien de la Montagne Noire: rapports entre mise en place des nappes et sédimentation. Carte géologique du flysch carbonifère entre le Mont Peyroux et Cabrières. 1:25 000. Bulletin du Bureau de Recherches géologiques et minières (2) I, 4, 341-389.

Ernst, A., Wyse Jackson, P. N., Aretz, M., 2015. Bryozoan fauna from the Mississippian (Visean) of Roque Redonde (Montagne Noire, southern France). Geodiversitas 37, 151-213.

Faure, M., Lardeaux, J.M., Ledru, J.P., 2008. Structure et évolution tectonique du Massif central. Géochronique 105, 37-43.

Ferguson, 1978. Some aspects of the ecology and growth of the Carboniferous Gigantoproductids. Proceedings of the Yorkshire Geological Society 42, 41-54.

Fournet, J., Graff, M., 1849. Note sur les terrains des environs de Neffiez et Roujan (département de l'Hérault). Bulletin Société géologique de France, 2ème série VI, 625-629.

Franke, W., 2014. Topography of the Variscan orogen in Europe: failed – not collapsed. International Journal of Earth Sciences (Geologische Rundschau) 103, 1471-1499.
Frech, F., 1887. Die paläozoischen Bildungen von Cabrières (Languedoc). Zeitschrift der
Deutschen Geologischen Gesellschaft 39, 360-487.

Galitzkaya, A. I., 1977. Ranne- i srednekamennougol'nye produktidy Severnoi Kirgizii [Early and Middle Carboniferous Productida of Northern Kirghizia]. Akademiya Nauk Kirgizkoi SSR. Frunze, 298 p.

Gèze, B., 1949. Etude géologique de la Montagne Noire et des Cévennes méridionales.
Mémoires de la Société géologique de France, nouvelle série 29, 1-215.

Gröber, P., 1908. Über die Faunen des unter-carbonischen Transgressionsmeeres des zentralen Tian-schan, die in der Umgebung des Sartdschol-Passes gefunden worden sind. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie 26, 213-248.

Gröber, P., 1909. Carbon und Carbon-Fossilien des nördlichen und zentralen Tian-Shan.
Abhandlungen der königlichen bayerischen Akademie der Wissenschaften (math-nat
Klasse) 24, 341-384.

Gutteridge, P., 1990. The origin and significance of the distribution of shelly macrofauna in late Dinantian carbonate mud mounds of Derbyshire. Proceedings of the Yorkshire Geological Society 48, 23-32.

Hammer, Ø., Harper, D.A.T., 2006. Paleontological Data Analysis. Blackwell Publishing, Oxford.

Hammer, Ø., Harper, D.A.T., Ryan, P.D. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4, 9 p.

Jaccard, P., 1901. Distribution de la flore alpine dans le Bassin des Dranses et dans quelques régions voisines. Bulletin de la Société Vaudoise des Sciences Naturelles 37, 241-272.

Julien, A., 1896. Le terrain carbonifère marin de la France centrale. Masson Edition, Paris.

Jin, Y.G., Sun D.L., 1981. Palaeozoic brachiopods from Xizang. In: Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (Ed.), Palaeontology of Xizang, Book III (The Series of the Scientific Expedition to the Qinghai-Xizang Plateau). Beijing: Science Press, 127-176 (in Chinese with English abstract).

Jin, Y.G., Ye, S.L., Xu, H.K., Sun, D.L., 1979. Brachiopoda. In: Paleontological Atlas of Northwestern China, Qinghai Province. Vol. 1, Lower Paleozoic-Cenozoic. Geological Press, Beijing, pp. 60-225 (in Chinese).

Kalashnikov, N.V., 1974. Rannekamennougol'nye brakhiopody Petchorskogo Urala [Lower Carboniferous brachiopods of the Petchora Urals.] Akademiya Nauk SSSR, Komi Filial Institut Geologii, Izdatel'stvo «Nauka», Leningrad, pp. 1-220 (in Russian).
de Koninck, L.G., 1842-1844. Description des animaux fossiles qui se trouvent dans le terrain carbonifère de Belgique. Dessain.

de Koninck, L.G., 1847. Recherche sur les animaux fossiles. Première partie – Monographie des genres *Productus* et *Chonetes*. Dessain.

Korn, D., Feist, R., 2007. Early Carboniferous ammonoid faunas and stratigraphy of the Montagne Noire (France). Fossil Record 10, 99-124.

Korn, D., Titus, A.L., Ebbighausen, V., Mapes, R.H., Sudar, M.N., 2012. Early Carboniferous (Mississippian) ammonoid biogeography. Geobios 45, 67-77.

Lazarev, S.S., 1990. Evolutsiya i sistema produktid [Evolution and systematics of productids]. Akademiya Nauk SSSR, Paleontologicheskii Institut, Moscow, Trudy 242, 1-171 (in Russian).

Lazarev, S.S., 2003. Brachiopods of the Superfamily Linoproductoidea Stehli, 1954: A New View on their Morphogenesis and Taxonomic Structure. Paleontological Journal 37, 492-500 (English Translation of Paleontologicheskii Zhurnal 2003, 43-51).

Lazarev, S.S., 2004. Revision of the Brachiopod Family Schrenkiellidae Lazarev, 1990. Paleontological Journal 38, 154-161 (English Translation of Paleontologicheskii Zhurnal, 2004, 40-46).

Legrand-Blain, M. 1973. Les Gigantoproductides (brachiopodes) du Sahara algérien. I. – Gigantoproductides viséens. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord 64, 79-158.

Legrand-Blain, M. 1980. Les Gigantoproductides (brachiopodes) du Sahara algérien. III. – Semiplanidae viséens et namuriens. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord 69, 3-86.

Legrand-Blain, M. 1985. Brachiopods. In: Wagner, R.H., Winkler Prins, C.F., Granados, L.F. (Eds.), Carboniferous of the World. 2. Australia, Indian Subcontinent, South Africa, South America and North Africa. IGME, Madrid, pp. 372-374.

Legrand-Blain, M. 1987. Les Gigantoproductidae (brachiopodes) namuriens du Sahara algérien. Bulletin de la Société belge de Géologie 96, 159-194.

Legrand-Blain, M. 2003. Chonetid and productid brachiopods from the "Calcaire à Productus", Mississippian, Montagne Noire (S. Massif Central, France). Abstracts XVth International Congress on Carboniferous and Permian Stratigraphy, Utrecht, 10-16 August 2003, 328.
Legrand-Blain, M. 2018. Gigantoproductid and allied brachiopods from "L'Ardoisière" (Viséan), northern Massif Central, France. Abstracts of the 8th International Brachiopod Congress - Milano, 11-14 September 2018, Permophiles 66 Suppl. 1, 69.

Legrand-Blain, M., Delvolvé, J.J., Perret, M.F., 1983. Les brachiopodes carbonifères des Pyrénées Centrales françaises. 1: Cadre stratigraphique et sédimentaire. Etude des Strophomenida. Geobios 16, 285-327.

Legrand-Blain, M., Delvolvé, J.J., Hansotte, M., 1996. Carboniferous brachiopods from the Arize Massif (Ariège), French Pyrénées. Geobios 29, 177-207.

Liu, Z.H., Tan, Z.X., Ding, Y.L., 1982. Brachiopoda. In: Geological Bureau of Hunan Province (Ed.), The Palaeontological Atlas of Hunan, Geological Memoirs, Ser. 2, No. 1. Beijing: Geological Publishing House, 172-216 (in Chinese).

Litvinovich, N.V., 1962. Kamennougol'nye i permskie otlozheniia zapadnoi chasti Tsentral'nogo Kazakhstana [Carboniferous and Permian deposits of the Western part of central Kazakhstan]. Moskovskii Gosudarstvennyi Universitet. Materialyi po geologii Tsentral'nogo Kazakhstana 4, 389 p.

Litvinovich, N.V., Vorontsova, T.N., 1983. K voprosu o revizii roda *Gigantoproductus* Prentice [The question of the revision of *Gigantoproductus* Prentice]. Biulleten Moskovskogo Obshchestva Ispytatelei Prirody (MOIP), Otdelenie Geologicheskii 58, 81-94 (in Russian).

Litvinovich, N.V., Vorontsova, T.N., 1991. Gigantoidnye Brakhiopody SSSR, ikh rasprostranenie i stratigraficheskoe znachenie [Gigantoid brachiopods of the USSR, their distribution and stratigraphic significance] Nauka, Moscow, 1-61 (In Russian).

Martínez Chacón, M.L., Legrand-Blain, M. 1992. Braquiópodos. In: Paleontological and Sedimentological Analysis of the Los Santos de Maimona Carboniferous Basin (Badajoz). Coloquios de Paleontología 44, 91-144.

Martínez Chacón, M.L., Winkler Prins, C.F., 2009. Brachiopods from the Valdeteja Formation (Pennsylvanian; Cantabrian Mountains, NW Spain). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 252, 91-111.

McCoy, F. 1844. A synopsis of the characters of the Carboniferous Limestone fossils of Ireland. Williams and Norgate: London.

Mottequin, B., Aretz, M., Herbig, H.G., Baidder, L., Kaoukaya, A., Denayer, J., 2017. New insight on Carboniferous (Viséan) brachiopods from eastern Tafilalt (Morocco). Geological Journal 52, 217-233.
Muir-Wood, H.M., Cooper, G.A., 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Geological Society of America Memoir 81, 1-447.

Muir-Wood, H.M., Williams, A., 1965. Strophomenida. In: Moore R.C. (Ed.), Treatise on Invertebrate Paleontology. Part H Brachiopoda. Geological Society of America and University of Kansas Press, H 1, pp. 361-521.

Nolan, L.S.P., Angiolini, L., Jadoul, F., Della Porta, G., Davies, S.J., Banks, V.J., Stephenson, M.H., Leng, M.J., 2017. Sedimentary context and palaeoecology of *Gigantoproductus* shell beds in the Mississippian Eyam Limestone Formation, Derbyshire carbonate platform, central England. Proceedings of the Yorkshire Geological Society 61, 239-257.

Öpik, A., 1934. Über die Klitamboniten. Tartu Ulikooli Geoloogia-Instituudi Toimestuesed Acta et Commentationes Universitatis Tartuensis 39, 1-239.

Ozaki, K., 1939. On some Lower Carboniferous brachiopods from central Hunan, China. Journal of the Shanghai Science Institute, Series 2 2, 225-282.

Paeckelmann, W., 1931. Die Fauna des deutschen Unterkarbons. 2. Die Brachiopoden des deutschen Unterkarbons. 2 Teil: Die Productinae und *Productus*-ähnlichen Chonetinea. Abhandlungen der Preussischen Geologische Landesanstalt N.F. 136, 1-440.

Pattison, J., 1981. The stratigraphic distribution of gigantoproductoid brachiopods in the Visean and Namurian rocks of some areas in northern England. Institute of Geological Sciences Report 81/9, 1-30.

Perez-Huerta, A., Sheldon, N., 2006. Pennsylvanian sea level cycles, nutrient availability and brachiopod paleoecology. Palaeogeography, Palaeoclimatology, Palaeoecology 230, 264-279.

Perret, M.F., Vachard, D., 1977. Algues et pseudo-algues des calcaires serpoukhoviens d'Ardengost (Hautes-Pyrénées). Annales de Paléontologie (Invertébrés) 63, 85-156.

Phillips, J., 1836. Illustrations of the geology of Yorkshire; or a description of the strata and organic remains: accompanied by a geological map, sections and diagrams and figures of the fossils. II, The Mountain Limestone District. John Murray, London.

Pille, L., Vachard, D., 2011. Diploporaceae (calcareous green algae dasycladales) from the late Visean (Mississippian) of Montagne Noire (southern France). Revue de Micropaléontologie 54, 1-30.

Poty, E., 2002. Stratigraphy and paleobiogeography of Carboniferous rugose corals of Nova Scotia. In: Hills, L.V., Henderson, C.M., Bamber, E.W. (Eds.), Carboniferous and Permian of the World. Canadian Society of Petroleum Geologists Memoir 19, 580-587.
Poty, E., Aretz, M., Barchy, L., 2002. Stratigraphie et sédimentologie des « Calcaires à Productus » du Carbonifère inférieur de la Montagne Noire (Massif central, France). Comptes Rendus de l'Académie des Sciences de Paris, Géosciences 334, 843-848.

Poty, E., Devuyst, F.-X., Hance, L., 2006. Upper Devonian and Mississippian foraminiferal and rugose coral zonation of Belgium and Northern France: a tool for Eurasian correlations. Geological Magazine 143, 829-857.

Prentice, J., 1956. *Gigantoproductus edelburgensis* (Phillips) and related species. Proceedings of the Yorkshire Geological Society 30, 229-258.

Qiao, L., Shen, S.Z., 2015. A global review of the Late Mississippian (Carboniferous) *Gigantoproductus* (Brachiopoda) faunas and their paleogeographic, paleoecological, and paleoclimatic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 420, 128-137.

Raup, D. M., Crick, R., 1979. Measurement of faunal similarity in paleontology. Journal of Paleontology 53, 1213-1227.

Rouville, P. de, 1868. Réunion extraordinaire à Montpellier (Hérault). Du 11 au 20 octobre 1868. Bulletin de la Société géologique de France, 2ème série 25, 873-1002.

Sarycheva, T. G., 1928. Podmoskovnye Produktidy gruppy *Productus giganteus* Martin (Gigantella gen. nov.) [The Podmoskovian productid group *Productus giganteus* Martin (Gigantella gen. nov.)]. Geologicheskogo Nauchno-Issledovatel’skogo Instituta pri Fiziko-Matematicheskom Fakul’tete 1-go Moskovskogo Gosudarstvennogo Universiteta (MGU) Institut, Trudy 1, 1-71 (in Russian).

Sarycheva, T.G. 1960. Phylum Brachiopoda. In: Orlov, Y. A. (Ed.), Fundamentals of Paleontology, vol. 7, Akademiya Nauk SSSR: Moscow, 113-343 (in Russian).

Sarycheva, T.G., 1970. Slovar terminov po morfologii produktid (Brachiopoda) [Dictionary of productid (Brachiopoda) morphological terminology]. Akademiya Nauk SSSR Paleontologicheskii Institut Nauka, Moscow, pp. 1-82. (in Russian, with an English Index).

Sarycheva, T.G., Legrand-Blain, M., 1977. Semeistvo Semiplanidae (Brachiopoda), ego sostav i razvitie [Generic composition and evolution of the family Semiplanidae (Brachiopoda)]. Paleontologicheskii Zhurnal 1977, 70-82 (in Russian; English translation in Paleontological Journal 1977, 200-212).

Sarycheva, T.G., Sokolskaya, A.N., 1952. Opredelitel paleozoiskikh brakhiopod Podmoskovnoy kotloviny [Description of Palaeozoic Brachiopoda of the Moscow
Basin]. Akademiya Nauk SSSR, Paleontologicheskii Institut, Trudy 38, 1-307 (in Russian).

Sarycheva, T.G., Sokolskaya, A.N., Beznosova, G.A., Maksimova, S.V., 1963. Brakhiopody i paleogeografiya karbona Kuznetskoi kotloviny [Carboniferous brachiopods and palaeogeography of the Kuznetsk Basin]. Akademiya Nauk SSSR, Paleontologicheskii Institut, Trudy 95, 1-547 (in Russian).

Schimansky, V. N., 1940. Gigantelly vizeiskikh otlozhenii rek podcherem, Shchugor, Vuktyl i basseina verkhnei Pechory (Sev. Ural) [Gigantellae of the Viséan strata of the rivers of Podcherem, Shugor, Vuktyl and Basin of the Upper Pechora (Northern Urals)]. Uchenye Zapiski Moskovskogo Gosudarstvennogo Pedagogicheskogo Instituta 23, 83-122 (in Russian).

Schwetsov, M.C., 1922. K voprosy o stratigrafii nizhnekammenougolnykh otlozhenii yuzhnogo kryla Podmoskovnogo basseina [On the problem of the stratigraphy of the Lower Carboniferous deposits of the southern side of the Moscow Basin]. Vestnik Moskovskogo Gornogo Akademiya 1, 1-20 (in Russian).

Serres, M. de, 1847. Note sur la découverte du genre des *Productus*, dans les formations carbonifères de Roujan ou de Neffiez (Hérault). Académie des Sciences et Lettres de Montpellier, Mémoires Section des Sciences, 1, 65-67.

Shan, H.Z., Zhao, R.X., 1981. Lower Carboniferous brachiopods of the Tzemenchiao Formation of Qiziqiao, Xiangxiang, Hunan. Acta Scientiarum Naturalium Universitatis Sunyatseni 4, 43-53 (in Chinese with English abstract).

Shen, S.Z., Qiao, L., Zhang, Y., Sun, Y.L., Jin, Y.G., 2017. Carboniferous brachiopod genera on type species of China. In: Rong, J.Y., Jin, Y.G., Shen, S.Z., Zhan, R.B. (Eds.), Phanerozoic brachiopod genera of China. Beijing: Science Press, 559-649.

Shi, G.R., 1993. Multivariate data analysis in palaeoecology and palaeobiogeography – a review. Palaeogeography, Palaeoclimatology, Palaeoecology 105, 199-234.

Simancas, J.F., Tahiri, A., Azor, A., González Lodeiro, F., Martínez Poyatos, D., El Hadi, H., 2005. The tectonic frame of the Variscan–Alleghanian Orogen in southern Europe and northern Africa. Tectonophysics 398, 181-198.

Sowerby, J., 1822. The Mineral Conchology of Great Britain, vol. 4. Published by the author. London, 1-114.

Stehli, F. G. 1954. Lower Leonardian Brachiopoda of the Sierra Diablo. American Museum of Natural History Bulletin 105, 257-358.
Tan, Z.X., 1987. Brachiopoda. In: Regional Geological Surveying Party and Bureau of
Geology and Mineral Resources of Hunan Province (Ed.), The Late Devonian and Early
Carboniferous Strata and Palaeobiocoenosis of Hunan. Beijing: Geological Publishing
House, pp. 111-133 (in Chinese with English summary).

Termier, H., 1936. Etudes géologiques sur le Maroc central et le Moyen Atlas septentrional. III. Paléontologie, Pétrographie. Notes et Mémoires du Service géologique du Maroc 33, 1083-1566.

Termier, H., Termier, G., 1950. Paléontologie marocaine. II - Invertébrés de l’Ere primaire. (2) Bryozoaires et brachiopodes. Notes et Mémoires du Service géologique du Maroc 77, 1-253.

Termier, H., Termier, G., Vachard, D., 1977. Biostratigraphie des terrains du Maroc Central depuis le Givétien jusqu'au Namurien inférieur. Actes VIème Colloque Africain de Micropaléontologie, Tunis 1974. Annales des Mines et de la Géologie Tunis 28, 39-63.

Vachard, D., 1974. Contribution à l'étude stratigraphique et micropaléontologique (algues et foraminifères) du Dévonien-Carbonifère inférieur de la partie orientale du versant méridional de la Montagne Noire (Hérault, France). Thèse 3ème cycle, Université Paris VI (unpubl.).

Vachard, D., 1977. Etude stratigraphique et micropaléontologique (algues et foraminifères) du Viséen de la Montagne Noire (Hérault, France). Mémoires de l'Institut géologique Université de Louvain 29, 111-195.

Vachard, D., Aretz, M., 2004. Biostratigraphic precisions on the Early Serpukhovian (Late Mississippian), by means of a carbonate algal microflora (cyanobacteria, algae and pseudo-algae) from La Serre (Montagne Noire, France). Geobios 37, 643-666.

Vachard, D., Cózar, P., Aretz, M., Izart, A., 2016a. Late Viséan-Serpukhovian cyanobacteria and algae from the Montagne Noire (France); taxonomy and biostratigraphy. Bulletin of Geosciences 91, 433-466.

Vachard, D., Cózar, P., Aretz, M., Izart, A., 2016b. Late Viséan-Serpukhovian foraminifers in the Montagne Noire (France): Biostratigraphic revision and correlation with the Russian substages. Geobios 49, 469-498.

Vachard, D., Izart, A., Cózar, P., 2017. Mississippian (middle Tournaisian-late Serpukhovian) lithostratigraphic and tectonosedimentary units of the southeastern Montagne Noire (Hérault, France). Géologie de la France 1, 47-88.

Vachard, D., Pille, L., Aretz, M., Herbig, H.-G., Cózar, P., 2006. *Eovelebitella* (dasyclad algae) and importance of its Brigantian subprovince within the western Palaeotethys. In:
Aretz, M., Herbig, H.-G. (Eds.), Carboniferous Conference Cologne. From Platform to Basin, September 4-10, 2006. Program and Abstracts, Kölner Forum für Geologie und Paläontologie 15, 120-121.

Vaughan, A., 1905. The palaeontological sequence in the Carboniferous Limestone of the Bristol area. Quarterly Journal of the Geological Society of London 61, 181-307.

Verneuil, E. de, 1849. Observations. In: Fournet, J., Graff, M., 1849. Note sur les terrains des environs de Neffiez et Roujan (département de l'Hérault). Bulletin Société géologique de France, 2ème série VI, 627-629.

Vidal, J., 1952a. Etude géologique du massif du Caragnas près Cabrières (Hérault). Diplôme d'Etudes supérieures de Géologie, Université de Montpellier, Faculté des Sciences. 1ère Partie (unpubl.).

Vidal, J., 1952b. Faune et stratigraphie de la grauwacke de Roquessels (Hérault). Diplôme d'Etudes supérieures de Géologie, Université de Montpellier, Faculté des Sciences. 2ème Partie (unpubl.).

Waagen, W.H. 1883. Salt Range fossils, Part 4 (2), Brachiopoda. Palaeontologia Indica, Series 13 4, 391-546.

Waterhouse, J.B., 2002. Classification within Productidina and Strophalosiidina (Brachiopoda). Earthwise 5, 1-60.

Waterhouse, J.B., 2004. Some new subfamilies, genera and species of suborder Productidina (Brachiopoda). Earthwise 7, 1-46.

Wilson, R.B., 1989. A study of the Dinantian marine macrofossils of central Scotland. Transactions of the Royal Society of Edinburgh Earth Sciences 80, 91-126.

Winkler Prins, C.F., Amler, M.R.W., 2006. Brachiopoden. In: Deutsche Stratigraphische Kommission (Ed.), Stratigraphie von Deutschland. VI. Unterkarbon (Mississippium). Schriftenreihe der Deutschen Gesellschaft für Geowissenschaften 41, 89-100.

Yang, D.L., Ni, S.Z., Chang, M.L., Chao, R.X., 1977. Brachiopoda. In: Geological Institute of Hubei Province (Ed.), Palaeontological Atlas of Central South China, Volume 2. Beijing: Geological Publishing House, pp. 303-470 (in Chinese).

Yang S.P. 1978. Lower Carboniferous brachiopods of Guizhou Province and their stratigraphic significance. Professional Papers of Stratigraphy and Palaeontology 5, 78-142 [in Chinese].

Yang, S.P., Lin, Y., Tang, G., Wang, Z., Wu, S., 1983. The Lower Carboniferous (Fengninian) of China. In: Martínez Días, C., Wagner, R.H., Granados, L.F., Winkler
Prins, C.F. (Eds.), The Carboniferous of the World. I. China, Korea, Japan & S.E. Asia. IUGS Publication 16, pp. 16-56.

Yang, S.P., Wang, S., Li, A., 1989. Study on early Carboniferous fossil communities of Hunan and Guizhou provinces. Compte Rendu XIème Congrès international de Stratigraphie et de Géologie du Carbonifère, Beijing 1987 3, 296-314.

Yao, L., Aretz, M., Yue, L., Wang, XD., 2016. Gigantoproductid brachiopod storm shell beds in the Mississippian of South China: implications for their palaeoenvironmental and palaeogeographical significances. Geologica Belgica 19, 57-67.

Zakowa, H., 1958. Biostratygrafia utworow morskich dolnego karbonu z obszaru Walbrzycha Miasta na Dolnym Slasku [Biostratigraphy of the lower Carboniferous marine deposits of the area of Walbrzych Miasto (lower Silesia)]. Instytut Geologiczny Prace, Warszawa 19, 1-211. (in Polish)

Zakowa, H., 1966. Poziom *Goniatites crenistria* Phill. w okolicy Sokolca i Jugowa u podnozy Gor Sowich (Sudety Srodkowe). [Zone *Goniatites crenistria* Phill. in the vicinity of Sokolec and Jugow, at the foot of the Sowie Gory Mountains (Central Sudetes)]. Instytut Geologiczny Prace, Warszawa 43, 1-197. (in Polish)

Zakowa, H., 1985. Upper Visean gigantoproductoid brachiopods from the Gory Swietokrzyskie, Poland. Annales Societatis geologorum Poloniae, Krakow 55, 105-126.

Zakowa, H., 1986. Brachiopods of the family Semiplanidae Sarytcheva, 1960 from the upper Visean of Poland. Biuletyn Instytutu Geologicznego 355, Geology of Poland 7, 49-70.

Zakowa, H., Zak, C., 1962. Dolny Karbon z Kamionek (Gory Sowie) [Lower Carboniferous at Kamionki (Sowie Mts - lower Silesia)]. Instytut Geologiczny Bulletyn, Warszawa 173, 169-277. (in Polish)

Zdanowski, A., Zakowa, H., 1995. The Carboniferous system in Poland. Prace Panstwowego Instytutu Geologicznego 148, 1-209.

Zhang, S.X., Jin, Y.G., 1976. Late Paleozoic Brachiopods from the Mount Jolmo Lungma Region. In: A report of scientific expedition in the Mount Jolmo Lungma region (1966-1968), vol. 2, Scientific Press, Beijing, pp. 159-271 (in Chinese).
**Table and Figure captions**

**Table 1.** Overview on the studied specimens: identification, sample locality, collection numbers (if present, the writing on the old labels is documented), and indication if specimens are figured herein.

**Table 2.** Overview on the studied sections: name, location, age (according to the most recent biostratigraphic information), and identified taxa.

**Fig. 1.** Geological map of the Mississippian Kulm deposits in the southeastern Montagne Noire (Engel et al., 1982), and brachiopod localities mentioned in the text (from West to East): **1.** Laurens-Gare; **2.** Sauvanés creek; **3.** Roquessels-Malac; **4.** Castelsec; **5.** Mas Foulquier; **6.** Roc de Murviel; **7.** Les Murailles; **8.** Roquemaillère; **9.** Les Mentaresses; **10.** Roque Redonde; **11.** Tiberet; **12.** La Serre vineyard; **13.** Cabrières-Japhet; **14.** Péret-Boissière; **15.** Péret-Rouirèdes. Units I-IV in the Laurens Flysch Group after Engel et al. (1982: figs. 9, 13): I. Small calcareous platform blocks only near top; II. Allochthonous calcareous blocks including "Calcaires à *Productus*": wider (some metres) and frequent; III. In complement to the calcareous blocks, some ante-flysch olistolites (Ordovician, Devonian); IV. Ecailles de Cabrières area, calcareous blocks of major size and thickness (see also the recent nomenclature of Vachard et al., 2017).

**Fig. 2. 1-3.** *Balakhonia* aff. *ostrogensis* Sarycheva, 1963 from Roquemaillère, biozones C, D, basal E: Brigantian. 1, 2: UCBL-FSL 703029, ventral valve in anterior and lateral views; 3: Specimen UCBL-FSL 703028, ventral valve umbo with some spine bases. **4-8.** *Datangia semiglobosa* (Paeckelmann, 1931). 4: From Castelsec Quarry, biozones upper E-lower G: Late Brigantian to early Serpukhovian, specimen UCBL-FSL 703023, incomplete ventral valve; 5: From Cabrières, biozone G, early Serpukhovian, specimen UCBL-FSL 703012, globulose ventral valve, ears not preserved; 6-8: From the Vailhan area, probably Brigantian. Specimen UCBL-FSL 703010, globulose ventral valve with one ear preserved in anterior, lateral, and posterior views; 8: partly exfoliated umbo, showing the thickness of ventral valve and internal muscle field. Scale bar: 10 mm.
**Fig. 3.** *Globosoproductus* sp. from Laurens-railway trench, Serpukhovian. Specimen UM ACI 653. Scale bar: 10 mm.

**Fig. 4.** **1. Kansuella** sp. 1 from Cabrières, Serpukhovian. Specimen UM ACI 664, young shell, thin-shelled ventral valve (max. thickness: 2.2 mm), partly exfoliated; inside, slightly divergent long diductor scars. **2. Latiproductus aff. complicatus** (Paeckelmann, 1931) from Neffies area. Specimen UM ACI 659, ventral valve, with ancient paper label; date and collector unknown. **3-6. Kansuella** sp. 1 from Laurens-Gare, Serpukhovian. 3: specimen UCBL-FSL 703011, big ventral valve partly exfoliated, with a large ear on right side; 4: specimen UCBL-FSL 703013, ventral valve umbo with numerous rugae traceable across venter; 5: specimen ML UCBL-FSL 703007, ventral big ginglymus on left hinge; 6: specimen UCBL-FSL 703015, dorsal small ginglymus. **7, 8. Kansuella** sp. 2, longitudinal sections of both valves. 7: specimen UCBL-FSL 703039 from La Serre vineyard, Serpukhovian; large ventral and small dorsal ginglymus; 8: specimen UM ACI 650 from Laurens area; dorsal valve broken, its ginglymus just preserved in contact with the ventral one, on the right extremity of the section. Ventral valve pierced by predator borings. Scale bars: 5 mm (the smallest one refers to pictures 1-4; the two others to pictures 5-6 and 7-8, respectively).

**Fig. 5.** **1. Kansuella** sp. 1 from Roquemaillère, biozones C, D, basal E: Brigantian. Specimen UCBL-FSL 703026, ventral valve, umbo covered by numerous rugae. **2-4. Kansuella** sp. 2 from Laurens area, lower Serpukhovian. 2: specimen UM ACI 651, partly exfoliated ventral valve showing: the large thickness of ventral valve, except at the deep insertion of a diductor muscle, left side, and a fragment of the dorsal valve internal mould; 3, 4: Specimen UM ACI 650 (old label not photographed: “Environ de Laurens 1855”): a large broken shell, in anterior view, and along the longitudinal fracture, showing thickness of ventral and dorsal valves. In the surrounding matrix sections of thin gastropod shells (4). **5, 6. Kansuella** sp. 1 from Laurens-Gare, Serpukhovian. Specimen UCBL-FSL 703021, dorsal valve cardinal process and internal mould. **7. Kansuella** sp. 2 from La Serre vineyard, biozones G-lower H, Serpukhovian. Specimen UCBL-FSL 703039, ventral valve, nearly no rugae; some thin spine bases on umbo. Scale bars: 10 mm (the largest one refers to the enlarged pictures 5 and 6).

**Fig. 6.** **1. Latiproductus edelburgensis** (Phillips, 1836) from La Serre vineyard, Serpukhovian. Specimen UCBL-FSL 703041, a large ventral valve, with some oblique cardinal spines well-preserved on the left ear. **2. Latiproductus** sp. Specimen UCBL-FSL 703005 from Laurens
area. **3. Latiproductus** cf. *latissimus* (Sowerby, 1822). Specimen UM ACI 668 from Cabrières: *Productus latissimus* Sowerby figured by Böhm (1935: pl. 8, fig. 18). **4. Latiproductus** aff. *latissimus complicatus* (Paeckelmann, 1931) from Peret-Boissière, probably Brigantian. Specimen UCBL-FSL 703043, ventral valve partly exfoliated, dorsal valve mould with reflected traces of ventral valve ornamentation. **5. Latiproductus latissimus complicatus** (Paeckelmann, 1931) from Sauvanés stream, NE of Laurens, Serpukhovian. Specimen UCBL-FSL 703022, ventral valve with spine bases initiating short prominent longitudinal plications. Scale bar: 10 mm.

**Fig. 7.** Stratigraphic distribution of the main gigantoproductid, semiplanid and latiproductid taxa collected in the main localities dealt with in this paper; stratigraphic ranges after the Montagne Noire biozones A-H (Vachard et al., 2016b). Castelsec: grey line indicates the stratigraphic extension of the entire locality when including the south-eastern outcrop; black line is the restricted stratigraphic extension for the northern outcrop where most of the samples were collected.

**Fig. 8.** A. Spatial distribution of the main gigantoproductid, semiplanid and latiproductid taxa on a hypothetical reconstruction for the upper Viséan–early late Serpukhovian platform found in the Montagne Noire olistoliths. The platform is largely dominated by carbonates, but especially towards the shoreface and the outer platform/slope siliciclastics can dominate. Some environments are only known in single or few localities and stratigraphic horizons. B. Overview on the main characteristics of the different environments in which large-sized brachiopods (gigantoproductids, semiplanids and latiproductids) occurred in the Montagne Noire.

**Fig. 9.** Palaeogeographical map for the Viséan centred on the Palaeotethys, with indication of the palaeobiogeographical units containing large-sized brachiopods (gigantoproductids, semiplanids, and latiproductids). The base map is modified from Colorado Plateau Geosystems, Inc. using the Creative Commons Attribution-Share Alike 4.0 International license (https://commons.wikimedia.org/wiki/File:340Marect.jpg).

**Fig. 10.** Palaeobiogeographical relations of gigantoproductids, semiplanids and latiproductids in the western Palaeotethys: Dendrograms for hierarchical cluster analyses using the Raup-Crick and Jaccard similarity coefficients by unweighted pair-group average (UPGMA)
method and Nonmetric Multidimensional Scaling (NMDS) ordination method. Cluster
analysis agglomerative coefficients (cophenetic correlations) are indicated for each analysis in
the top right corner. Node percentages are bootstrap supports (1000 iterations). All analyses
performed with the PAST 3 software (Hammer et al., 2001). See fig. 9 for
palaeobiogeographical unit abbreviations.
<figure><img src="image_3.png" /></figure>

Carbonate lenses:
"Calcaire à Productus"

Kulm: late Viséan-
Serukhovian

Tournaisian -
early Viséan

Pre-Carboniferous
1

<figure><img src="image_4.png" /></figure>

2

<figure><img src="image_5.png" /></figure>

3

<figure><img src="image_6.png" /></figure>

5

<figure><img src="image_7.png" /></figure>

4

<figure><img src="image_8.png" /></figure>

6

<figure><img src="image_9.png" /></figure>

8

<figure><img src="image_10.png" /></figure>

7

<figure><img src="image_11.png" /></figure>
<figure><img src="image_12.png" /></figure>

<figure><img src="image_13.png" /></figure>
2

Product
names

<figure><img src="image_14.png" /></figure>

1

<figure><img src="image_15.png" /></figure>

3

<figure><img src="image_16.png" /></figure>

4

<figure><img src="image_17.png" /></figure>

5

<figure><img src="image_18.png" /></figure>

6

<figure><img src="image_19.png" /></figure>

7

<figure><img src="image_20.png" /></figure>

8

<figure><img src="image_21.png" /></figure>
1

<figure><img src="image_22.png" /></figure>

2

<figure><img src="image_23.png" /></figure>

3

<figure><img src="image_24.png" /></figure>

4

<figure><img src="image_25.png" /></figure>

5

<figure><img src="image_26.png" /></figure>

6

<figure><img src="image_27.png" /></figure>

7

<figure><img src="image_28.png" /></figure>
1

<figure><img src="image_29.png" /></figure>

2

<figure><img src="image_30.png" /></figure>

3

<figure><img src="image_31.png" /></figure>

5

<figure><img src="image_32.png" /></figure>

4

<figure><img src="image_33.png" /></figure>
CARBONIFEROUS (part)

System

MISSISSIPPIAN (part)

Subsystem

<table><thead><tr><th colspan="2">VISEAN (part)</th><th colspan="2">SERPUKHOVIAN (part)</th><th colspan="2">Stage</th><th rowspan="2">Russian Substage</th><th rowspan="2">local biozone</th></tr><tr><th colspan="2">Mikhailovian</th><th colspan="2">Venezian</th><th colspan="2">Protvian</th></tr><tr><th>A</th><th>B</th><th>C</th><th>D</th><th>E</th><th>F</th><th>G</th><th>H</th></tr></thead><tbody><tr><td colspan="8"><hr>Roque Redonde Vailhan Laurens La Serre <hr>Latiproductus edelburgensis</hr></td></tr><tr><td colspan="8"><hr>Vailhan Laurens <hr>Latiproductus latissimus complicatus</hr></td></tr><tr><td colspan="8"><hr>Laurenso Castelsec Cabrières <hr>Datangia semiglobosa Balakhonia aff. ostrogensis</hr></td></tr><tr><td colspan="8"><hr>Laurenso Castelsec Cabrières <hr>Laurenso Castelsec Cabrières <hr>Kansuela sp. 1</hr></td></tr><tr><td colspan="8"><hr>Laurenso Castelsec Cabrières <hr>Kansuela sp. 2</hr></td></tr><tr><td colspan="8"><hr>Laurenso <hr>Globosoproductus sp.</hr></td></tr></tbody></table>

taxa
<figure><img src="image_34.png" /></figure>

1 Moscow Basin and Donbas (RUS)

2 Lublin Basin (POL)

3 Rhenish Massif (RM)

4 Namur-Dinant Basin (BEL)

5 British Isles (GBR)

6 Ossa Morena Zone (ESP)

7 Notsch (AUT)

8 Montagne Noire (MON)

9 Pyrenees (PYR)

10 Moroccan Meseta (MM)

11 Sahara Basins (ALG)

12 Eastam Urais & Central

Asian Orogenic Belt (CAOB)

13 North China

14 South China
Cluster analysis (Raup-Crick)

0.0305

<figure><img src="image_35.png" /></figure>

Cluster analysis (Jaccard)

0.0959

<figure><img src="image_36.png" /></figure>

NMDs (Raup-Crick)

Stress 0.1985

<figure><img src="image_37.png" /></figure>
<table><caption>Table 1.</caption><thead><tr><th>Collection & Taxon</th><th>Locality</th><th>Collection number</th><th>Fig.</th></tr></thead><tbody><tr><td colspan="4"><strong>Collection Legrand-Blain (ML)</strong></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703028 = ML 764a 1</td><td>2(3)</td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703029 = ML 764a 2</td><td>2(1-2)</td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703030 = ML 764a 3</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703031 = ML 764a 4</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703032 = ML 764a 5</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703033 = ML 764a 6</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703034 = ML 764a 7</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Roquemaillère</td><td>UCBL-FSL 703035 = ML 764a 8</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Péret-Rouirèdes</td><td>UCBL-FSL 703046 = ML 1055 c 1</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Péret-Rouirèdes</td><td>UCBL-FSL 703047 = ML 1055 c 2</td><td></td></tr><tr><td>Balakhonia aff. ostrogenis Sarycheva, 1963</td><td>Péret-Rouirèdes</td><td>UCBL-FSL 703048 = ML 1055 c 3</td><td></td></tr><tr><td>Datangia semiglobosa (Paeckelmann, 1931)</td><td>Castelsec</td><td>UCBL-FSL 703023 = ML 765 a 1</td><td>2(4)</td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703004 = ML 1158 a 1</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703007 = ML 1158 a 2</td><td>4(5)</td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703011 = ML 1158 a 3</td><td>4(3)</td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703013 = ML 1158 a 4</td><td>4(4)</td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703014 = ML 1158 a 5</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703015 = ML 1158 a 6</td><td>4(6)</td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703016 = ML 1158 a 7</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703017 = ML 1158 a 8</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703018 = ML 1158 a 9</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703019 = ML 1158 a 10</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703020 = ML 1158 a 11</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Laurens-Gare</td><td>UCBL-FSL 703021 = ML 1158 a 12</td><td>5(5-6)</td></tr><tr><td>Kansuella sp. 1</td><td>Castelsec</td><td>UCBL-FSL 703024 = ML 1175</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Castelsec</td><td>UCBL-FSL 703025 = ML 765 c 1</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Castelsec</td><td>UCBL-FSL 703026 = ML 765 c 2</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Roquemaillère</td><td>UCBL-FSL 703036 = ML 764 d</td><td>5(1)</td></tr><tr><td>Kansuella sp. 1</td><td>La Serre vineyard</td><td>UCBL-FSL 703038 = ML 1176 b1</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Péret-Rouirèdes</td><td>UCBL-FSL 703049 = ML 1055 b</td><td></td></tr><tr><td>Kansuella sp. 2</td><td>La Serre vineyard</td><td>UCBL-FSL 703039 = ML 1176 a1</td><td>4(7), 5(7)</td></tr><tr><td>Latiproductus latissimus complicatus (Paeckelmann, 1931)</td><td>Sauvanès stream</td><td>UCBL-FSL 703022 = ML 1174 b 1</td><td>6(5)</td></tr><tr><td>Latiproductus aff. latissimus complicatus (Paeckelmann, 1931)</td><td>Péret-Boissière</td><td>UCBL-FSL 703042 = ML 784 a 1</td><td></td></tr><tr><td>Latiproductus aff. latissimus complicatus (Paeckelmann, 1931)</td><td>Péret-Boissière</td><td>UCBL-FSL 703043 = ML 784 a 2</td><td>6(4)</td></tr><tr><td>Latiproductus aff. latissimus complicatus (Paeckelmann, 1931)</td><td>Péret-Boissière</td><td>UCBL-FSL 703044 = ML 784 a 3</td><td></td></tr><tr><td>Latiproductus aff. latissimus complicatus (Paeckelmann, 1931)</td><td>Péret-Boissière</td><td>UCBL-FSL 703045 = ML 784 a 4</td><td></td></tr><tr><td>Latiproductus sp.</td><td>Castelsec</td><td>UCBL-FSL 703027 = ML 791 a1</td><td></td></tr><tr><td>Latiproductus sp.</td><td>La Serre vineyard</td><td>UCBL-FSL 703040 = ML 1054 b</td><td></td></tr><tr><td>Latiproductus edelburgensis (Phillips, 1836)</td><td>Roque Redonde</td><td>UCBL-FSL 703037 = ML 1057 a1</td><td></td></tr><tr><td>Latiproductus edelburgensis (Phillips, 1836)</td><td>La Serre vineyard</td><td>UCBL-FSL 703041 = ML 1152 a</td><td>6(1)</td></tr><tr><td colspan="4"><strong>Palaeontological Collection, University of Lyon (France)</strong></td></tr><tr><td>Datangia semiglobosa (Paeckelmann, 1931)</td><td>Vailhan area</td><td>UCBL-FSL 703010</td><td>2(6-8)</td></tr><tr><td>Datangia semiglobosa (Paeckelmann, 1931)</td><td>Cabrières</td><td>UCBL-FSL 703012</td><td>2(5)</td></tr><tr><td>Datangia semiglobosa (Paeckelmann, 1931)</td><td>Neffies area</td><td>UCBL-FSL 703006 with old label: "P. giganteus Martin (sp). Neffiez. Coll. Deshayes (1869)"</td><td></td></tr><tr><td>Kansuella sp. 1</td><td>Vailhan</td><td>FSL 703009 with old label: "Productus cora Valve supérieure et inférieure. Calcaire à Productus giganteus. Vallée de Vailhan"</td><td></td></tr><tr><td>Kansuella sp. 2</td><td>Laurens area</td><td>UCBL-FSL 703001 with old label: "P. giganteus Dinantien Laurence (le plus complet)"</td><td></td></tr><tr><td>Kansuella sp. 2</td><td>Laurens area</td><td>UCBL-FSL 703003</td><td></td></tr><tr><td>Latiproductus latissimus complicatus (Paeckelmann, 1931)</td><td>Vailhan area</td><td>UCBL-FSL 703008 with old label: "Coll. GEANDEY. Productus Carboniférien. Vailhan Hérault"</td><td></td></tr><tr><td>Latiproductus sp.</td><td>Laurens area</td><td>UCBL-FSL 703005 with old label "Division G. Productus latissimus d'après Verneuil. Calcaire à Productus giganteus Laurens"</td><td></td></tr></tbody></table>
Latiproductus edelburgensis (Phillips, 1836) Laurens area

UCBL-FSL 703002 with old label. "Division F n°3: Viséen environs de Laurens, *P. edelburgensis*"

Palaeontological Collection, University of Montpellier (France)

Datangia semiglobosa (Paeckelmann, 1931) Cabrières

UM ACI 663

Globosoproductus sp. Laurens area

UM ACI 653 with: (1) an old sticked poorly legible label "...Laurens (Herault)"; and (2) a cardboard label from Université de Montpellier (Géologie) "*Productus giganteus* Mart. Viséen supérieur.

Laurens tranchée de chemin de fer". It was possibly collected in the railway trench by the station master, M. Cornac (Bergeron, 1889, p. 158)

Kansuella sp. 1 Cabrières

(UM CAB b1) = UM ACI 664 4(1)

Kansuella sp. 1 Cabrières

UM ACI 665 with old label: "*Productus maximus* Mc Coy, Viséen Cabrières. Déterm. M. Delépine. Forme la + ancienne de *giganteus*»

Kansuella sp. 2 Laurens area

UM ACI 651 with old label: "Viséen Laurens" 5(2)

Kansuella sp. 2 Laurens area

UM ACI 650 with old label: "Environs de Laurens 1855" 4(8), 5(3-4)

Latiproductus cf. latissimus (Sowerby, 1822) Cabrières

UM ACI 668 6(3)

Latiproductus latissimus complicatus (Paeckelmann, 1931) "Vailhan"

UM ACI 661

Latiproductus aff. latissimus complicatus "Neffies"

UM ACI 659 with old label «*Productus Nefies*» 4(2)

(Paeckelmann, 1931)

Latiproductus sp. Laurens

UM ACI 652

Latiproductus sp. Roquemaillère

UM ACI 658

Latiproductus sp. Cabrières

UM ACI 667 with old label "fragment de coquille immature"

Latiproductus edelburgensis (Phillips, 1836) Mas Foulquier

UM ACI 655 with old label: "Route de Magrignan en face de Mas Foulquier, juin 1936"

Latiproductus edelburgensis (Phillips, 1836) Vailhan "La Muraille"

UM ACI 656
<table><caption>Table 2.</caption><thead><tr><td>Outcrop</td><td>Location</td><td>Age</td><td>Studied fauna</td></tr></thead><tbody><tr><td>1a. Laurens-Gare</td><td>vineyards west-south-west of the old Railway Station building of Laurens on the south side of the D 136 road: 43°31'08N, 3°12'26E. No longer accessible due to recent house building</td><td>early Serpukhovian (Vachard et al., 2017)</td><td>Kansuella sp. 1</td></tr><tr><td>1b. Laurens area</td><td>in several case identical to 1a, but some samples could also come from small scattered outcrops around the village</td><td>probably Steshevian (following Vachard et al., 2017)</td><td>Kansuella sp. 2, Latiproductus edelburgensis, Latiproductus sp., Globosoproductus sp.</td></tr><tr><td>2. Sauvanés creek</td><td>outcrop along small creek NE of Laurens. 43°31'49N, 3°12'03E</td><td>early late Visean (following Vachard et al., 2017)</td><td>Latiproductus latissimus complicatus</td></tr><tr><td>3. Roquessels-Malac</td><td>outcrop along the road Gabian-Faugères, also called "lentilles de la route" (see Vachard et al. 2017). 43°32'50N, 3°12'47E</td><td>early late Visean (following Vachard et al., 2017)</td><td>Latiproductus sp., undeterminable kansuellid</td></tr><tr><td>4. Castelsec quarry</td><td>disused quarry east of Castelsec at the road to Fos. 43°31'57N, 3°14'01E</td><td>biozones E-H (Aretz, 2002a, Vachard et al., 2016b); Venetian-Protvian (Vachard et al., 2017)</td><td>Datangia semiglobosa, Latiproductus sp</td></tr><tr><td>5. Mas Foulquier</td><td>probably along small path, SE of Les Pascales. 43°32'08N, 3°16'23E</td><td>biozone F</td><td>Latiproductus edelburgensis</td></tr><tr><td>6. Roc de Murviel</td><td>large hill east of the road Gabian-Montesqieu (northern flank: 43°33'08N, 3°16'36E, southern flank: 43°33'00N, 3°16'35E.)</td><td>biozones E- G, Venetian- Steshevian (Vachard et al., 2017)</td><td>northern flank: Datangia? sp.; southern flank: Latiproductus sp., Roc de Murviel Fm on southern flank: Latiproductus cf. edelburgensis</td></tr><tr><td>7. Les Murailles</td><td>hill 1.5 km SW from Vailhan. 43°32'39N, 3°17'17E</td><td>biozones F and G. Tarusian- Steshevian (Vachard et al., 2017)</td><td>Latiproductus cf. edelburgensis, Kansuella? sp. Balakhonia aff. ostrogensis, Kansuella sp. 1., Datangia semiglobosa, Latiproductus aff. latissimus complicates; Vachard (1974, 1977) cited Gigantoproductus cf. giganteus determined by G. Termier from this locality.</td></tr><tr><td>8. Roquemaillère and "Neffies"</td><td>hill 2.5 km S of Vailhan. 43°31'53N, 3°17'34E</td><td>biozones D-F. Venetian- Tarusian (Vachard et al., 2017) and «Vailhan», probably biozones A-D</td><td>Latiproductus edelburgensis Latiproductus edelburgensis, Kansuella sp. 1, Latiproductus aff. latissimus complicatus</td></tr><tr><td>9. Les Mentaresses</td><td>hill 1 km SSW from Vailhan. 43°32'33N, 3°17'57E</td><td>biozones D-F. Venetian- Tarusian (Vachard et al., 2017) and «Vailhan», probably biozones A-D</td><td>Latiproductus edelburgensis</td></tr><tr><td>10. Roque Redonde</td><td>disused quarry on top of hill. 43°32'50N, 3°18'27E</td><td>biozones G-H. Steshevian- Protvian (Vachard et al., 2017)</td><td>Kansuella sp. 2, Latiproductus edelburgensis</td></tr><tr><td>11. Tiberet</td><td>scattered outcrops along small track. 43°33'39N, 3°19'49E</td><td>biozone H. Protvian (Vachard et al., 2017)</td><td>Datangia semiglobosa, Latiproductus cf. latissimus, Kansuella sp. 1</td></tr><tr><td>12. La Serre vineyard</td><td>vineyard east of la Rouquette and north of the D 15 road. Aretz (2002b, figs 18, 20: between levels D and E, 21). 43°33'17N, 3°21'26E</td><td>Venetian and Tarusian (Vachard et al., 2017)</td><td>Latiproductus aff. latissimus complicatus</td></tr><tr><td>13. Cabrières-Japhet</td><td>road cut of the Route de Péret in Cabrières, just east of the bridge above the La Boyne. 43°34'24N, 3°21'52E</td><td>Venetian and Tarusian (Vachard et al., 2017)</td><td>Balakhonia sp., Kansuella sp. 1</td></tr><tr><td>14. Péret-Boissière</td><td>hill 0.4 km W of Péret. 43°34'45N, 3°23'09E</td><td>Venetian and Tarusian (Vachard et al., 2017)</td><td></td></tr><tr><td>15. Péret-Rouirèdes</td><td>hill 0.6 km NNW of Péret. 43°35'15N, 3°23'27E</td><td>Venetian-Serpukhovian</td><td></td></tr></tbody></table>