Arthrophycus in the Silurian of Alabama (USA) and the problem of compound trace fossils

Arthrophycus brongniartii (Harlan, 1832) is common in marginal-marine deposits in the Silurian Red Mountain Formation of Alabama. The ichnospecies, the second to be named in North America, is revived and emended after long disuse. Transitional forms to Rusophycus isp. and other morphologic evidence indicate that the maker of Arthrophycus was an arthropod, perhaps a trinucleine (raphiophorid?) trilobite. Interconnection of Arthrophycus and Nereites biserialis, as well as intergradation of Arthrophycus with Cruziana aff. quadrata, Phycodes flabellum, and Asterosoma ludwigae, indicate that these Red Mountain trace fossils were made by the same species of arthropod. Possible relationships with Arthrophycus alleghaniensis (Harlan, 1831) in the Silurian belt from Ontario to Tennessee are also explored. Ichnofamily Arthrophycidae Schimper, 1879 is emended. The ichnofamily is interpreted as chiefly the work of arthropods. Arthrophycus and other trace fossils from the Silurian of Alabama constitute a test case to build criteria for recognizing the members of complexes of trace fossils. In general, criteria such as interconnection of different forms, intergradation among unconnected forms, similarity of size, similarity of morphologic elements, and co-occurrence should be examined in order to determine the biologic and ethologic interrelationships of trace fossils.