Raphiophoridae

Raphiophoridae Temporal range: Ordovician to Middle Silurian PreЄ Є O S D C P T J K Pg N
Ampyx nasutus, partially rolled, showing the cephalon with genal spines and rapier-like glabellar spine, but hiding most of the thorax and the pygidium
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Trilobita
Order:	Asaphida
Suborder:	Trinucleina
Family:	Raphiophoridae Angelin, 1854
subfamilies
Raphiophorinae Angelin, 1854 synonyms Ampyxininae, Ampyxinellinae, Taklamakaniinae Endymioniinae Raymond, 1920 synonym Typhlokorynetidae see text for genera

Raphiophoridae is a family of small to average-sized trilobites that first occurred at the start of the Ordovician and became extinct at the end of the Middle Silurian.

Anatomy

All raphiophorids are blind, with headshields (or cephalons) that are triangular to subcircular, and many carry long, trailing genal spines, a forward-directed rapier-like spine on the central raised area (or glabella), or both, with the glabella often inflated and the natural fracture lines (or sutures) of the cephalon coinciding with its margin. The thorax typically has five to seven segments^[1] (except for the genera Taklamakania, Pseudampyxina, Nanshanaspis, and Kongqiangheia, which have only 3). As mentioned before, all raphiophorids are blind. Many, if not most, genera have no eyes whatsoever, though a few, such as Lehnertia, have vestigial tubercles that correspond to the compound eyes of their ancestors.

Distribution

Raphiophoridae currently includes two officially recognized subfamilies. The nominal subfamily Raphiophorinae originated from the Upper Tremadocian and died out during the Lower Ludlow, and has 217 species assigned to it divided over 26 genera. The subfamily Endymioniinae occurred from the Floian or possibly from the Lower Tremadocian to the Upper Katian and contains 36 species in 13 genera. The time of the first occurrence depends on whether the inadequately known monotypic genus Typhlokorynetes from the Lower Tremadocian of Laurentia is considered an endymioniin. Raphiophorids are generally found in deep-water sediments, and have a cosmopolitan distribution from the Floian to the Ordovician–Silurian extinction events with diversity peaking from the Darriwilian to the Sandbian. Raphiophorus survived into the Silurian.^[2]

Taxonomy

The subfamily Taklamakaniinae was erected to bring together the genera Nanshanaspis, Pseudampyxina, and Taklamakania, (and then, later, Kongqiangheia) on the basis that they all have only three thoracic segments. Analysis of adult anatomy of these genera and larval stages of other raphiophorids showed they most probably developed through paedomorphosis from three different ancestors, so provide an example of parallel evolution. Nanshanaspis closely resembles young Globampyx, Pseudampyxina strongly looks like juvenile Raymondella, and Taklamakania is almost identical to early stages of Ampyxina. Since the three genera of the Taklamakaniinae have been demonstrated to be unrelated to each other, this subfamily is regarded as polyphyletic, and has been synonymized with the Raphiophorinae.^[3]

Genera

These genera are assigned to the Raphiophoridae:^[1]

Subfamily Raphiophorinae Raphiophorus Angelin, 1854 Ampyx Dalman, 1827 Ampyxella Dean, 1960 Ampyxina Ulrich, 1922 Ampyxoides Whittington, 1965 Bulbaspis Chugaeva, 1956 Cnemidopyge Whittard, 1955 Edmundsonia Cooper, 1953 Globampyx Fortey, 1975 Kongqiangheia Lonchodomas Angelin, 1854 Mendolaspis Rusconi, 1951 Nanshanaspis Chang & Fan, 1960 Pseudampyxina Ju, 1983 Raymondella Reed, 1935 Rhombampyx Fortey, 1975 Taklamakania Zhang, 1980

Subfamily Endymioniinae Endymionia Billings, 1865 Ampyxinella Koroleva, 1959 Ampyxinops Zhang 1979 Anisonotella Whittington, 1965 Carinocranium Dean, 1989 Edmundsonia Cooper 1953 Jiuxiella Zhou et al. 1977 Lehnertia Vaccari, et al, 2006 Malongullia Webby, Moors & McLean, 1970 Miaopopsis Lu et al. 1965, synonym = Sinampyxina Miboshania Zhang 1979 Pytine Fortey, 1975 Salteria Thomson, 1864 ? Tarimella W. T. Zhang 1979 (may be a shumardiid ptychopariid) Typhlokorynetes Shaw, 1966

References

1. Fortey, R.A. (1975). The Ordovician Trilobites of Spitsbergen [II. Asaphidae, Nileidae, Raphiophoridae and Telephinidae of the Valhallfonna Formation]. Norsk Polarinstitutt Skrifter 162. Oslo: Norsk Polarinstitutt.
2. J.M. Adrian (2014). "20. A synopsis of Ordovician trilobite distribution and diversity". In D.A.T. Harper, T. Servais. Early Palaeozoic Biogeography and Palaeogeography. Memoirs of the Geological Society of London 38. Geological Society of London. p. 490. ISBN 1862393737.
3. Zhou, Z.; Webby, B.D.; Yuan, W. (1995). "Ordovician trilobites from the Yingan Formation of northwestern Tarim, Xinjiang, northwestern China". Alcheringa: An Australasian Journal of Palaeontology 19 (1): 47–72. doi:10.1080/03115519508619098.