Development Genes and Evolution

, Volume 216, Issue 7–8, pp 395–415 | Cite as

The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence

  • Gerhard ScholtzEmail author
  • Gregory D. Edgecombe


Understanding the head is one of the great challenges in the fields of comparative anatomy, developmental biology, and palaeontology of arthropods. Numerous conflicting views and interpretations are based on an enormous variety of descriptive and experimental approaches. The interpretation of the head influences views on phylogenetic relationships within the Arthropoda as well as outgroup relationships. Here, we review current hypotheses about head segmentation and the nature of head structures from various perspectives, which we try to combine to gain a deeper understanding of the arthropod head. Though discussion about arthropod heads shows some progress, unquestioned concepts (e.g., a presegmental acron) are still a source of bias. Several interpretations are no longer tenable based on recent results from comparative molecular developmental studies, improved morphological investigations, and new fossils. Current data indicate that the anterior arthropod head comprises three elements: the protocerebral/ocular region, the deutocerebral/antennal/cheliceral segment, and the tritocerebral/pedipalpal/second antennal/intercalary segment. The labrum and the mouth are part of the protocerebral/ocular region. Whether the labrum derives from a former pair of limbs remains an open question, but a majority of data support its broad homology across the Euarthropoda. From the alignment of head segments between onychophorans and euarthropods, we develop the concept of “primary” and “secondary antennae” in Recent and fossil arthropods, posit that “primary antennae” are retained in some fossil euarthropods below the crown group level, and propose that Trilobita are stem lineage representatives of the Mandibulata.


Segmentation Labrum Primary antennae Secondary antennae Trilobita Cambrian 



We thank Jean Deutsch and Michel Veuille, the organizers of the meeting “Development and Phylogeny of Arthropods,” Paris 2005, for the invitation to GS to present our thoughts on head evolution. Nikola-Michael Prpic made helpful comments on an earlier version of the manuscript. We are grateful to Henrike Semmler for her pictures of the innervation of the frontal filaments in nauplius larvae of cirripedes and Carsten Wolff for the picture of the embryonic head of Orchestia cavimana (Fig. 4). Alfred Palissa helped with literature about Protura. This article benefited tremendously from the inspiring discussions with the students in the seminar “Phylogeny of Recent and Fossil Taxa” held at the Humboldt-University.

Supplementary material


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© Springer-Verlag 2006

Authors and Affiliations

  1. 1.Humboldt-Universität zu BerlinInstitut für Biologie/Vergleichende ZoologieBerlinGermany
  2. 2.Australian MuseumSydneyAustralia

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