Millipede
phylogeny revisited in the light
of the enigmatic order Siphoniulida
P.
Sierwald1, W. A. Shear2, R. M. Shelley3 and
J. E. Bond4
1Zoology, The Field Museum,
Chicago, IL, USA; 2Department of Biology,
Hampden-Sydney College, Hampden-Sydney, VA, USA; 3North Carolina State Museum of Natural Sciences, Raleigh, NC, USA; 4East Carolina University, Greenville, NC, USA
Abstract
The
discovery of six specimens of the enigmatic order Siphoniulida, including for
the first time males, prompted a modern re-analysis of current phylogenetic
schemes for the class Diplopoda derived from traditional morphological and
developmental characters. The data matrix was constructed and analysed using
paup. The resulting phylogenetic hypotheses corroborated the longest standing,
traditional classification, but also demonstrated clearly that more characters
must be included to reach a better resolution. Recent alternative phylogenetic
hypotheses and classifications are discussed in the light of the current analysis.
The validity of a putative helminthomorph synapomorphy, the location of male gonopods
on the 7th body ring, is discussed. Scanning electron microscopy corroborated
morphological characters already described for the Siphoniulida: modified
anterior legs, an apparently legless 3rd body ring, pyriform head, antennae
with clavate setae, and absence of ozopores. The highly modified gonopods of
the Siphoniulida are described for the first time; only the anterior legs of
the 7th ring are modified into gonopods, the posterior legs of this ring are
developed as normal walking legs. The gnathochilarium differs from the
Colobognatha and consists of well-developed stipites with palps, elongated
lingual plates with palps and a narrow central sclerite, most likely the mentum.
Structures on the epiproct may possibly be spinnerets. Despite the discovery of
adult males, the Siphoniulida are still considered Helminthomorpha incertae
sedis.
Key
words: Diplopoda Ð Siphoniulida Ð systematics Ð Helminthomorpha Ð Mexico
J.
Zool. Syst. Evol. Research 41 (2003) 87Ð99
2003
Blackwell Verlag, Berlin ISSN 0947Ð5745
Received
on 18 April 2001