A preliminary phylogenetic study of late Palaeozoic spiriferoid brachiopods using cladistic and Bayesian approaches

Abstract The brachiopod Superfamily Spiriferoidea diversified greatly and was widely distributed in the late Palaeozoic (Carboniferous–Permian), and yet its phylogeny has been seldom investigated with analytical methods. This is reflected in the current flux of very different classification schemes for this superfamily. This paper provides the first attempt to investigate the phylogenetic relationships of spiriferoid brachiopods through both cladistic and Bayesian analyses involving 24 discrete and continuous characters. The continuous characters, from morphometric data, have been separately discretized using the gap weighting method, and the ‘as such’ option in TNT. Our results highlight the potential significance of continuous characters in reconstructing and elucidating phylogenies, as much as qualitative characters. Building on the outcomes of the analyses, we also briefly evaluate existing classification schemes of Spiriferoidea. We found that none of the existing classifications fully reflect the phylogeny properly; major families within the superfamily, such as Spiriferidae, Choristitidae, and Trigonotretidae, turned out to be polyphyletic. Although this study is considered preliminary, due to the selection of and restriction to certain taxa, combined with the use of a relatively small number of characters, it nevertheless demonstrates that potentially the true phylogenetic relationships of spiriferoid taxa sharply contrast with any of the existing classification schemes. This highlights the need to develop an alternative scheme that takes into account a more comprehensive range of phylogenetic variables.

Early Carboniferous spiriferoid brachiopods from the Qaidam Basin, Northwest China: taxonomy, biostratigraphy and biogeography

Six Early Carboniferous brachiopod species in four genera of the Superfamily Spiriferoidea are described from the Qaidam Basin, northwestern China, including a new genus, Qaidamospirifer, and two new species: Grandispirifer qaidamensis and Qaidamospirifer elongatus. Additionally, a new genus, Triangulospirifer, is also proposed to replace Triangularia (Poletaev, 2001) that was preoccupied by a Devonian molluscan genus.

On the basis of the new material as well as published information, we have reviewed the taxonomic composition and the stratigraphic and palaeobiogeographic distributions of the three previously established genera from the viewpoint of palaeobiogeography. The study reveals that Grandispirifer has a relatively long stratigraphic range from the late Tournaisian to Serpukhovian. During this interval, the genus attained a wide geographical distribution, reaching Northwest China, western Yunnan of West China, Japan, as well as Iran and North Africa. Angiospirifer first occurred in western Europe in the Viséan, and later migrated to North Africa during the late Viséan. In the Serpukhovian, it migrated eastward, reaching the Donets Basin of Ukraine and the Qaidam Basin in Northwest China. Anthracothyrina evolved from Brachythyrina in North Africa in late Viséan, then dispersed north-westward to western and eastern Europe and, further eastward to the Qaidam Basin during the Serpukhovian.

Ibergirhynchia contraria (F. A. Roemer, 1850), an Early Carboniferous seep-related rhynchonellide brachiopod from the Harz Mountains, Germany - A possible successor to Dzieduszyckia?

A new genus Ibergirhynchia, a member of the rhynchonellide superfamily Dimerelloidea, is described for the species Terebratula contraria Roemer, 1850, from Early Carboniferous deposits of the Harz Mountains, Germany. Ibergirhynchia contraria is from a monospecific brachiopod limestone that formed on top of the drowned Devonian Iberg Reef which persisted as a seamount during Famennian and Early Carboniferous times. Ibergirhynchia contraria is considered a cold seep-related brachiopod based on this locality. Such seep associations have been observed for Mesozoic representatives of the rhynchonellide superfamily Dimerelloidea. Ibergirhynchia is considered the first Paleozoic representative of the family Rhynchonellinidae. Ibergirhynchia resembles Dzieduszyckia externally and may be derived from this dimerelloid.

A biogeographically mixed, Middle Permian brachiopod fauna from the Baoshan Block, Western Yunnan, China

A small brachiopod fauna is described from the carbonate rocks of the basal Shazipo Formation of the Baoshan Block, western Yunnan, south-west China, including significant new ventral and dorsal internal morphological features of Cryptospirifer omeishanensis Huang. This fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) because of the presence of Cryptospirifer omeishanensis Huang and associated fusulinids (Neoschwagerina craticulifera Zone). Palaeobiogeographically, the brachiopod fauna is of considerable interest because of its admixed nature characterized by typical warm-water Cathaysian elements intermingled with temperate Peri-Gondwanan taxa. This in turn is interpreted to indicate that the Baoshan Block may have been situated in an intermediate palaeogeographical position between Gondwanaland to the south and Cathaysia to the north during the Mid Permian and, as such, it probably furnished an important 'stepping stone' for the dispersal of Mid Permian eastern Tethyan marine invertebrate taxa (e.g. Cryptospirifer) to the western Tethys.

A Lopingian (Late Permian) brachiopod fauna from the Qubuerga formation at Shengmi in the Mount Qomolangma region of Southern Xizang (Tibet), China

A small fauna of 11 species belonging to 10 genera of Permian Brachiopoda from the lower part of the Qubuerga Formation outcropping near Shengmi village in the Qomolangma region of southern Xizang (Tibet) is figured and new taxa are described. New taxa are Quinquenella semiglobosa and Costatumulus shengmiensis. The fauna is most likely of Wuchiapingian (Djhulfian) age as indicated by the majority of the brachiopod species.

Late carboniferous to early permian brachiopod faunas from the Bachu and Kalpin areas, Tarim Basin, NW China

Late Carboniferous and Early Permian brachiopod faunas are described from the Xiaohaizi section of the Bachu area and the Shishichang section of the Kalpin area, the Tarim Basin, NW China. Biostratigraphic studies of brachiopods and associated microfossils indicate that the Xiaohaizi Formation is Moscovian (Late Carboniferous) and the Shishichang Formation is Kasimovian-Gzhelian (Late Carboniferous), whereas the Nanza and Kankarin Formations are Asselian to early Artinskian (Early Permian). Two new species proposed from the Nanza Formation are Kutorginella tarimensis and Phricodothyris? bachuensis.

A new genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for timing the recovery of Brachiopoda after the end-Permian mass extinction

A new genus, Meishanorhynchia, is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and
bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic-Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1±0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3±0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4±0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.

A Guadalupian-Lopingian (Middle to Late Permian) brachiopod fauna from the Juripu formation in the Yarlung-Zangbo suture zone, Southern Tibet, China

A brachiopod fauna of 16 species belonging to 11 genera and three genera and species indeterminate from the middle and upper parts of the Juripu Formation in the Yarlung-Zangbo (Indus-Tsangbo) Suture zone (=Yarlung-Zangbo River zone), southern Tibet, is described and figured for the first time. A new species, Taeniothaerus zhongbaensis, is described. The fauna is comparable with that in the Kalabagh Member of the Wargal Formation of the Salt Range, Pakistan, and is considered to be most likely Capitanian (late Guadalupian/Middle Permian) to Wuchiapingian (early Lopingian/early Late Permian) in age, as indicated by the majority of the brachiopod species and by being constrained by an underlying fusulinacean fauna (Parafusulina Zone) and an overlying ammonite fauna (Cyclolobus fauna).

Capitanian (late Guadalupian, permian) global brachiopod palaeobigeography and latitudinal diversity pattern

A global database containing 3365 occurrences, 821 species and 251 genera of the Capitanian (Late Guadalupian, Permian) brachiopod faunas from 24 stations has been analyzed by cluster analysis using the Jaccard and Otsuka coefficients and the probabilistic index of similarity, nonmetric multidimensional scaling and minimum spanning tree. Two supergroups, three groups and six subgroups are revealed and interpreted as representing, respectively, two biotic realms (the Palaeoequatorial and Gondwanan Realms), two regions and six provinces. An additional realm (the Boreal Realm), based on the fauna from Spitsbergen, also appears recognizable although it also shows considerable similarities with southwestern North America and the northern margin of Gondwana as revealed by the statistical analysis. The Palaeoequatorial Realm can be further subdivided into the North America Region and the Asian Tethyan Region. The six biotic provinces are the Cathaysian Province in the Palaeotethys and Mesotethys, the Greenland-Svalbard Province in the Arctic region, the Austrazean Province in eastern Australia and New Zealand, the Grandian Province in western North America and the two transitional zones (the Himalayan Province in the southern temperate zone and the Sino–Mongolian–Japanese Province in the northern temperate zone). Polynomial regression analysis and rarefaction analysis indicate that the generic diversities of brachiopod faunas during the Capitanian peaked in the Palaeoequatorial Cathaysian Province and the two transitional zones (Himalayan Province and Sino–Mongolian–Japanese Province), but fell dramatically in the polar regions. The generic diversity of the Palaeoequatorial Grandian Province is apparently lower than in the two transitional zones of temperate palaeolatitudes, suggesting that the generic diversity of Capitanian brachiopod faunas does not exhibit a strict negative correlation with palaeolatitudes. This in turn would suggest that biogeographical determinants (such as geographical barriers, inhabitable area and ocean currents) other than latitude-related temperature control may also have played an important role in the dispersal of some brachiopods and the characterization of some local provinces and high diversities. The Capitanian global brachiopod palaeobiogeography is generally comparable with those in the Wuchiapingian and Changhsingian, but with some notable differences. These include: (1) that the Grandian Province of the Capitanian in western North America vanished after the end-Guadalupian regression, (2) that the western Tethyan Province of the Lopingian could not be distinguished in the Capitanian, and (3) that the Austrazean Province was larger in area than either in the Wuchiapingian or in the Changhsingian.

Early carboniferous brachiopod faunas from the Baoshan block, west Yunnan, southwest China

38 brachiopod species in 27 genera and subgenera are described from the Yudong Formation in the Shidian-Baoshan area, west Yunnan, southwest China. New taxa include two new subgenera: Unispirifer (Septimispirifer) and Brachythyrina (Longathyrina), and seven new species: Eomarginifera yunnanensis, Marginatia cylindrica, Unispirifer (Unispirifer) xiangshanensis, Unispirifer (Septimispirifer) wafangjieensis, Brachythyrina (Brachythyrina) transversa, Brachythyrina (Longathyrina) baoshanensis, and Girtyella wafangjieensis. Based on the described material and constraints from associated coral and conodont faunas, the age of the brachiopod fauna from the Yudon Formation is considered late Tournaisian (Early Carboniferous), with a possibility extending into earlyViseacutean.