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Heads, Hox, trilobites

Heads, Hox, trilobites

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Published by: ardeegee on Aug 09, 2008
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ABSTRACTThe Arachnomorpha or Arachnata concepts have resolved Trilobita as most closely relatedto Chelicerata amongst extant Arthropoda. An alternative position of trilobites in the stemlineage of Mandibulata is suggested by their pattern of head tagmosis. The antennae of trilobites and Mandibulata are considered non-homologous with the antennae of Onycho- phora and stem lineage Euarthropoda: they represent ‘secondary’ and ‘primary antennae’,respectively. In extant taxa, ‘secondary antennae’ are deutocerebral, post-ocular, and areconnected to deutocerebral olfactory neuropils, whereas ‘primary antennae’ are pre-ocular and connected to protocerebral olfactory neuropils. In fossils, an insertion at the antero-lateral margin of the hypostome rather than more anteriorly on the head allows ‘secondaryantennae’ to be identified. A deutocerebral mouthpart, of which the onychophoran jaw andthe chelicera are examples, is regarded as plesiomorphic for Arthropoda. A loss of ‘primaryantennae’ and modification of the deutocerebral mouthpart into a sensory antenna definesthe Mandibulata. Trilobites share a ‘secondary antenna’ and a clearly-delimited head tagmawith mandibulates. Given the extensive homoplasy forced by the Arachnata concept (rever-sals in pycnogonids and arachnids), a trilobite/mandibulate alliance may be better sup- ported.
 Dedicated to Fred Schram on the occasion of his retirement. Our article challenges canonical viewsabout arthropods, and we were forced to question ideas that we have long considered the best expla-nation of facts. In doing so, we venture into a territory from which Fred Schram has never shied away. Fred’s synthesis of data from living and fossil arthropods, his efforts to integrate classical morphological and evo-devo perspectives, and his willingness to explore dangerous ideas haveinspired our reappraisal of the trilobite problem.
INTRODUCTIONThe last decade has seen dramatic changes of our views on arthropod development, mor- phology, palaeontology, phylogeny, and evolution (see, for example, the books edited byFortey & Thomas 1997; Edgecombe 1998; Deuve 2001; Scholtz 2004). The comparativemolecular approach to embryology, cell lineage studies, new microscopic techniques with ahigh morphological resolution, and phylogenetic analyses based on molecular and refined
Heads, Hox and the phylogenetic position of trilobites
Institut für Biologie/Vergleichende Zoologie, Humboldt-Universität zu Berlin, Berlin, Germany
Australian Museum, Sydney, Australia
Scholtz & Edgecombe
140morphological data sets led to an increased interest in the body organisation, developmentand evolution of arthropods and to new and controversial hypotheses about arthropodrelationships. New and sometimes surprising solutions emerged from molecular and mor- phological approaches to long standing and highly controversial issues such as head seg-mentation, trunk tagmosis, and limb homologies in arthropods. Here we show how thecurrent views about arthropod tagmosis patterns, which are mainly based on molecular developmental genetics, influence our interpretations of fossils. This does not imply princi- pally untestable inferences about developmental patterns and processes in fossil groups butit leads to a framework based on data from Recent arthropods which allows new interpreta-tions of fossil structures and relationships.We reappraise the phylogenetic placement of trilobites by examining current data onhead and trunk segmentation of extant arthropods, including brain anatomy, developmental,and gene expression evidence. These data lead to a radically altered thinking about the basic alignment of the head segments in the major arthropod groups. The new view of heads invites a reconsideration of trilobites and other arthropod fossils and their affinitieswithin the Arthropoda.2
WHO ARE THE TRILOBITA?Trilobita is the most species-rich extinct clade within the Arthropoda. Trilobites are knownfrom more than 10,000 species that, in total, span some 275 million years from the EarlyCambrian to the end of the Permian. Though trilobites are among the most familiar fossilorganisms (Fig. 1), their systematic position within the Arthropoda remains contested(Westheide 1996). Because they are a well characterized fossil group in terms of their appendage structure, tagmosis, ontogeny, and exoskeletal form (Fig. 1), the precise phylo-genetic position of Trilobita has important consequences for broader issues in arthropod phylogeny.Following the discovery of the antennae and biramous appendages of trilobites in thelate 19
century, most workers regarded trilobites as most closely related to crustaceans(Beecher 1893; Raymond 1920), and trilobite-crustacean affinities (Hu 1971) or a trilo- bitomorph ancestry for Crustacea have been maintained by some later investigators (San-ders 1957; Hessler & Newman 1975). The Arachnomorpha concepts of Heider (1913) andlater Størmer (1944) provided a major shift in thinking about trilobite relationships becausethey related Trilobita with Chelicerata. According to Størmer Arachnomorpha was con-ceived as a group that encompassed Trilobita, a variety of extinct taxa in the Trilobito-morpha, and the Chelicerata. The idea that chelicerates are the closest living relatives of trilobites has been maintained by most recent workers. Bergström (1979, 1980) placed particular emphasis on the structure of the lamellar setae on the appendages (exopods or  book gills) as an indicator of a trilobite-chelicerate relationship. Subsequently, a laterallysplayed stance of the limbs and the dorsal penetration of the eyes were cited as additionalapomorphic characters for Arachnomorpha (Bergström 1992). Parsimony analyses thathave included a variety of fossil taxa have also resolved trilobites within an arachnomorphclade that includes the chelicerates as its only extant member (Wills et al. 1995, 1998;Cotton & Braddy 2004). An exception to the prevailing idea of trilobite-chelicerate affini-ties was the Gnathomorpha concept of Boudreaux (1979), in which trilobites were instead
 Heads, Hox and the phylogenetic position of trilobites
141resolved as stem lineage Mandibulata, based principally on a shared head/trunk tagmosis pattern. Unlike Boudreaux, most palaeontologists have considered trilobites and trilobito-morphs to provide evidence in favour of a group that unites crustaceans with cheliceratesrather than with other mandibulates. The trilobite-chelicerate-crustacean group (TCC of Cisne 1974, 1975) corresponds to the so-called Schizoramia sensu Bergström (1979, 1980). Neontological data, both morphological and molecular, conflict with the Schizoramia or TCC concepts, which attests to the central role of extinct taxa in formulating this grouping.The Arachnata concept of Lauterbach (1973, 1980b, 1983) was developed in the contextof Trilobita, in its traditional sense, being paraphyletic with respect to Chelicerata. Lauter- bach proposed three characters in support of the Olenellinae (Early Cambrian taxa usuallyregarded as trilobites) being sister group to Chelicerata, with the remaining trilobites then being the sister group to that assemblage. Lauterbach’s characters were subsequentlyrejected (Ramsköld & Edgecombe 1991), and the idea of trilobite paraphyly was countered by a greater amount of evidence in favour of trilobite monophyly (Fortey & Whittington1989). In the present study, we consider Trilobita to be a monophyletic group, as indicated by such apomorphic characters as a low magnesian calcite cuticle, uniquely mineralisedeyes, and circumocular ecdysial sutures. A distinctive mode of segment shedding in onto-geny, with thoracic segments released from the anterior margin of a transitory pygidiumand a pygidium as a tagma of unreleased segments, defines Trilobita or a slightly moreinclusive trilobitomorph clade (Edgecombe & Ramsköld 1999).
Figure 1.
Two representatives of Trilobita showing the characteristic body organisation with a head bearing compound eyes and a trunk with a thoracic region and a posterior pygidium with fused seg-ments. (A)
 Paradoxides gracilis
from the Cambrian of Bohemia, Czech Republic, length 11 cm(Zoologische Lehrsammlung, Humboldt-Universität zu Berlin). (B)
 Ptychopyge excavato-zonata
fromthe Ordovician of southern Sweden, length 4.5 cm (private collection GS).

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