Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrodl, and M. K. Nishiguchi. 2006. “Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons.” Proc Natl Acad Sci U S A 103: 7723-8. Abstract

Monoplacophorans are among the rarest members of the phylum Mollusca. Previously only known from fossils since the Cambrian, the first living monoplacophoran was discovered during the famous second Galathea deep-sea expedition. The anatomy of these molluscs shocked the zoological community for presenting serially repeated gills, nephridia, and eight sets of dorsoventral pedal retractor muscles. Seriality of organs in supposedly independent molluscan lineages, i.e., in chitons and the deep-sea living fossil monoplacophorans, was assumed to be a relic of ancestral molluscan segmentation and was commonly accepted to support a direct relationship with annelids. We were able to obtain one specimen of a monoplacophoran Antarctic deep-sea species for molecular study. The first molecular data on monoplacophorans, analyzed together with the largest data set of molluscs ever assembled, clearly illustrate that monoplacophorans and chitons form a clade. This "Serialia" concept may revolutionize molluscan systematics and may have important implications for metazoan evolution as it allows for new interpretations for primitive segmentation in molluscs.

Recent progress in molecular techniques has generated a wealth of information for phylogenetic analysis. Among metazoans all but a single phylum have been incorporated into some sort of molecular analysis. However, the minute and rare species of the phylum Loricifera have remained elusive to molecular systematists. Here we report the first molecular sequence data (nearly complete 18S rRNA) for a member of the phylum Loricifera, Pliciloricus sp. from Korea. The new sequence data were analyzed together with 52 other ecdysozoan sequences, with all other phyla represented by three or more sequences. The data set was analyzed using parsimony as an optimality criterion under direct optimization as well as using a Bayesian approach. The parsimony analysis was also accompanied by a sensitivity analysis. The results of both analyses are largely congruent, finding monophyly of each ecdysozoan phylum, except for Priapulida, in which the coelomate Meiopriapulus is separate from a clade of pseudocoelomate priapulids. The data also suggest a relationship of the pseudocoelomate priapulids to kinorhynchs, and a relationship of nematodes to tardigrades. The Bayesian analysis placed the arthropods as the sister group to a clade that includes tardigrades and nematodes. However, these results were shown to be parameter dependent in the sensitivity analysis. The position of Loricifera was extremely unstable to parameter variation, and support for a relationship of loriciferans to any particular ecdysozoan phylum was not found in the data.

This work expands on a study from 2004 by Mallatt, Garey, and Shultz [Mallatt, J.M., Garey, J.R., Shultz, J.W., 2004. Ecdysozoan phylogeny and Bayesian inference: first use of nearly complete 28S and 18S rRNA gene sequences to classify the arthropods and their kin. Mol. Phylogenet. Evol. 31, 178-191] that evaluated the phylogenetic relationships in Ecdysozoa (molting animals), especially arthropods. Here, the number of rRNA gene-sequences was effectively doubled for each major group of arthropods, and sequences from the phylum Kinorhyncha (mud dragons) were also included, bringing the number of ecdysozoan taxa to over 80. The methods emphasized maximum likelihood, Bayesian inference and statistical testing with parametric bootstrapping, but also included parsimony and minimum evolution. Prominent findings from our combined analysis of both genes are as follows. The fundamental subdivisions of Hexapoda (insects and relatives) are Insecta and Entognatha, with the latter consisting of collembolans (springtails) and a clade of proturans plus diplurans. Our rRNA-gene data provide the strongest evidence to date that the sister group of Hexapoda is Branchiopoda (fairy shrimps, tadpole shrimps, etc.), not Malacostraca. The large, Pancrustacea clade (hexapods within a paraphyletic Crustacea) divided into a few basic subclades: hexapods plus branchiopods; cirripedes (barnacles) plus malacostracans (lobsters, crabs, true shrimps, isopods, etc.); and the basally located clades of (a) ostracods (seed shrimps) and (b) branchiurans (fish lice) plus the bizarre pentastomids (tongue worms). These findings about Pancrustacea agree with a recent study by Regier, Shultz, and Kambic that used entirely different genes [Regier, J.C., Shultz, J.W., Kambic, R.E., 2005a. Pancrustacean phylogeny: hexapods are terrestrial crustaceans and maxillopods are not monophyletic. Proc. R. Soc. B 272, 395-401]. In Malacostraca, the stomatopod (mantis shrimp) was not at the base of the eumalacostracans, as is widely claimed, but grouped instead with an euphausiacean (krill). Within centipedes, Craterostigmus was the sister to all other pleurostigmophorans, contrary to the consensus view. Our trees also united myriapods (millipedes and centipedes) with chelicerates (horseshoe crabs, spiders, scorpions, and relatives) and united pycnogonids (sea spiders) with chelicerates, but with much less support than in the previous rRNA-gene study. Finally, kinorhynchs joined priapulans (penis worms) at the base of Ecdysozoa.

Sorensen, M. V., and G. Giribet. 2006. “A modern approach to rotiferan phylogeny: combining morphological and molecular data.” Mol Phylogenet Evol 40: 585-608. Abstract

The phylogeny of selected members of the phylum Rotifera is examined based on analyses under parsimony direct optimization and Bayesian inference of phylogeny. Species of the higher metazoan lineages Acanthocephala, Micrognathozoa, Cycliophora, and potential outgroups are included to test rotiferan monophyly. The data include 74 morphological characters combined with DNA sequence data from four molecular loci, including the nuclear 18S rRNA, 28S rRNA, histone H3, and the mitochondrial cytochrome c oxidase subunit I. The combined molecular and total evidence analyses support the inclusion of Acanthocephala as a rotiferan ingroup, but do not support the inclusion of Micrognathozoa and Cycliophora. Within Rotifera, the monophyletic Monogononta is sister group to a clade consisting of Acanthocephala, Seisonidea, and Bdelloidea-for which we propose the name Hemirotifera. We also formally propose the inclusion of Acanthocephala within Rotifera, but maintaining the name Rotifera for the new expanded phylum. Within Monogononta, Gnesiotrocha and Ploima are also supported by the data. The relationships within Ploima remain unstable to parameter variation or to the method of phylogeny reconstruction and poorly supported, and the analyses showed that monophyly was questionable for the families Dicranophoridae, Notommatidae, and Brachionidae, and for the genus Proales. Otherwise, monophyly was generally supported for the represented ploimid families and genera.

Giribet, G. 2005. “Generating implied alignments under direct optimization using POY.” Cladistics 21: 396-402.
Sharma, P, and G Giribet. 2005. “A new Troglosiro species (Opiliones, Cyphophthalmi, Troglosironidae) from New Caledonia. .” Zootaxa 1053: 47-60.
Worsaae, K, A Nygren, GW Rouse, G Giribet, J Persson, P Sundberg, and F Pleijel. 2005. “Phylogenetic position of the meiofaunal family Nerillidae (Annelida: Polychaeta) analyzed by direct optimization of combined molecular and morphological datasets.” Zoologica Scripta 34: 313-328.
Schulze, A, EB Cutler, and G Giribet. 2005. “Reconstructing the phylogeny of the Sipuncula. .” Hydrobiologia 535/536: 277-296.
Schwendinger, PJ, and G Giribet. 2005. “The systematics of the south-east Asian genus Fangensis Ramble (Opiliones: Cyphophthalmi: Stylocellidae).” Invertrabrate Systematics 19: 297-323.
Giribet, G., and J. A. Dunlop. 2005. “First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber.” Proc Biol Sci 272: 1007-13. Abstract

Two inclusions in a piece of Upper Cretaceous (Albian) Burmese amber from Myanmar are described as a harvestman (Arachnida: Opiliones), Halitherses grimaldii new genus and species. The first Mesozoic harvestman to be named can be referred to the suborder Dyspnoi for the following reasons: prosoma divided into two regions, the posterior formed by the fusion of the meso- and metapeltidium; palp lacking a terminal claw, with clavate setae, and tarsus considerably shorter than the tibia. The bilobed, anteriorly projecting ocular tubercle is reminiscent of that of ortholasmatine nemastomatids. The status of other Mesozoic fossils referred to Opiliones is briefly reviewed.

In this study, we present phylogenetic data to characterize the relationships among sironids centered in the Balkan region, and use these results to discuss biogeographical aspects of sironid evolution. Analysis of ca. 4.5 kb of sequence data from three nuclear and two mitochondrial genes reveals monophyly of a Balkan clade for which we resurrect the name Cyphophthalmus, considered a junior synonym of Siro for over a century. This clade diversified into several groups, and at least three of them--the duricorius group, the serbicus group, and the minutus group--are well corroborated by the data as monophyletic lineages. The members of the different groups, mostly living in troglobitic environments, have diversified in overlapping geographic regions, with evidence of an eastern origin for the group. Our data also suggest that mitochondrial and nuclear genes are all contributing towards the final resolution of the combined analysis of the data.

In order to elucidate the evolutionary history and the population structure of the members of the phylum Cycliophora, which live commensally on three species of lobsters, we studied sequence variation in the mitochondrial gene cyctochrome c oxidase subunit I. Overall 242 sequences from 16 locations on both coasts of the North Atlantic, including the North Sea and the Mediterranean, were analysed, revealing 28 haplotypes, with a maximum sequence divergence of 16.6%. Total genetic diversity was high (h = 0.8322, pi = 0.0898), as it was for the commensals on Homarus americanus (17 haplotypes, h = 0.7506, pi = 0.0504). However, it was low for commensals on Nephrops norvegicus (6 haplotypes, h = 0.3899, pi = 0.0035), and intermediate for cycliophorans on Homarus gammarus (5 haplotypes, h = 0.3020, pi = 0.0140). Although two of the host lobsters co-inhabit the coastal waters of Europe, a strong genetic structure (78.45% of the observed genetic variation) was detected among populations on all host species, indicating the existence of a reproductively isolated species on each lobster. In addition, genetic structure over long distances exists among populations on each host species. Such patterns can be explained by the limited dispersal ability of the cycliophoran chordoid larva. Demographic and phylogenetic analyses suggest old and possibly cryptic populations present on H. americanus and H. gammarus, while the latter may have experienced recent bottlenecks, perhaps during Pleistocene glaciations. Populations on N. norvegicus appear to be of recent origin.

Maxmen, A., W. E. Browne, M.Q. Martindale, and G. Giribet. 2005. “Neuroanatomy of sea spiders implies an appendicular origin of the protocerebral segment.” Nature 437: 1144-8. Abstract

Independent specialization of arthropod body segments has led to more than a century of debate on the homology of morphologically diverse segments, each defined by a lateral appendage and a ganglion of the central nervous system. The plesiomorphic composition of the arthropod head remains enigmatic because variation in segments and corresponding appendages is extreme. Within extant arthropod classes (Chelicerata, Myriapoda, Crustacea and Hexapoda--including the insects), correspondences between the appendage-bearing second (deutocerebral) and third (tritocerebral) cephalic neuromeres have been recently resolved on the basis of immunohistochemistry and Hox gene expression patterns. However, no appendage targets the first ganglion, the protocerebrum, and the corresponding segmental identity of this anterior region remains unclear. Reconstructions of stem-group arthropods indicate that the anteriormost region originally might have borne an ocular apparatus and a frontal appendage innervated by the protocerebrum. However, no study of the central nervous system in extant arthropods has been able to corroborate this idea directly, although recent analyses of cephalic gene expression patterns in insects suggest a segmental status for the protocerebral region. Here we investigate the developmental neuroanatomy of a putative basal arthropod, the pycnogonid sea spider, with immunohistochemical techniques. We show that the first pair of appendages, the chelifores, are innervated at an anterior position on the protocerebrum. This is the first true appendage shown to be innervated by the protocerebrum, and thus pycnogonid chelifores are not positionally homologous to appendages of extant arthropods but might, in fact, be homologous to the 'great appendages' of certain Cambrian stem-group arthropods.

Giribet, G, G.D. Edgecombe, JM Carpenter, CA D'Haese, and WC Wheeler. 2004. “Is Ellipura monophyletic? A combined analysis of basal hexapod relationships with emphasis on the origin of insects.” Orignism Diversity Evolution 4: 319-340.
Giribet, G, MV Sorensen, P Funch, RM Kristensen, and W Sterrer. 2004. “Investigations into the phylogenetic position of Micrognathozoa using four molecular loci.” Cladistics 20: 1-13.
Edgecombe, GD, and G Giribet. 2004. “Molecular phylogeny of Australasian anopsobiine centipedes (Chilopoda: Lithobiomorpha).” Invertrabrate Systematics 18: 235-249.