Journal Articles: 2010-2006

2008
Alberti, G., E Lipke, and G Giribet. 2008. “On the ultrastructure and identity of the eyes of Cyphophthalmi based on a study of Stylocellus sp. (Opiliones, Stylocellidae).” Journal of Arachnology 36: 379-387.
Giribet, G. 2008. “Assembling the lophotrochozoan (=spiralian) tree of life.” Philos Trans R Soc Lond B Biol Sci 363: 1513-22. Abstract

The advent of numerical methods for analysing phylogenetic relationships, along with the study of morphology and molecular data, has driven our understanding of animal relationships for the past three decades. Within the protostome branch of the animal tree of life, these data have sufficed to establish its two main side branches, the moulting Ecdysozoa and the non-moulting Lophotrochozoa. In this review, I explore our current knowledge of protostome relationships and discuss progress and future perspectives and strategies to increase resolution within the main lophotrochozoan clades. Novel approaches to coding morphological characters are needed by scoring real observations on species selected as terminals. Still, methodological issues, for example, how to deal with inapplicable characters or the coding of absences, may require novel algorithmic developments. Taxon sampling is another key issue, as phyla should include enough species so as to represent their span of anatomical disparity. On the molecular side, phylogenomics is playing an increasingly important role in elucidating animal relationships, but genomic sampling is still fairly limited within the lophotrochozoan protostomes, for which only three phyla are represented in currently available phylogenies. Future work should therefore concentrate on generating novel morphological observations and on producing genomic data for the lophotrochozoan side of the animal tree of life.

Dunn, C. W., A. Hejnol, D. Q. Matus, K. Pang, W. E. Browne, S. A. Smith, E. Seaver, et al. 2008. “Broad phylogenomic sampling improves resolution of the animal tree of life.” Nature 452: 745-9. Abstract

Long-held ideas regarding the evolutionary relationships among animals have recently been upended by sometimes controversial hypotheses based largely on insights from molecular data. These new hypotheses include a clade of moulting animals (Ecdysozoa) and the close relationship of the lophophorates to molluscs and annelids (Lophotrochozoa). Many relationships remain disputed, including those that are required to polarize key features of character evolution, and support for deep nodes is often low. Phylogenomic approaches, which use data from many genes, have shown promise for resolving deep animal relationships, but are hindered by a lack of data from many important groups. Here we report a total of 39.9 Mb of expressed sequence tags from 29 animals belonging to 21 phyla, including 11 phyla previously lacking genomic or expressed-sequence-tag data. Analysed in combination with existing sequences, our data reinforce several previously identified clades that split deeply in the animal tree (including Protostomia, Ecdysozoa and Lophotrochozoa), unambiguously resolve multiple long-standing issues for which there was strong conflicting support in earlier studies with less data (such as velvet worms rather than tardigrades as the sister group of arthropods), and provide molecular support for the monophyly of molluscs, a group long recognized by morphologists. In addition, we find strong support for several new hypotheses. These include a clade that unites annelids (including sipunculans and echiurans) with nemerteans, phoronids and brachiopods, molluscs as sister to that assemblage, and the placement of ctenophores as the earliest diverging extant multicellular animals. A single origin of spiral cleavage (with subsequent losses) is inferred from well-supported nodes. Many relationships between a stable subset of taxa find strong support, and a diminishing number of lineages remain recalcitrant to placement on the tree.

Murienne, J., M. S. Harvey, and G. Giribet. 2008. “First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata).” Mol Phylogenet Evol 49: 170-84. Abstract

The phylogenetic relationships of the major lineages of the arachnid order Pseudoscorpiones are investigated for the first time using molecular sequence data from two nuclear ribosomal genes and one mitochondrial protein-encoding gene. The data were analyzed using a dynamic homology approach with the new program POY v.4 under parsimony as the optimality criterion. The data show monophyly of Pseudoscorpiones as well as many of its superfamilies (Feaelloidea, Chthonioidea, Cheiridioidea and Sternophoroidea), but not for Neobisiodea or Garypoidea. Cheliferoidea was not monophyletic either due to the position of Neochelanops, which grouped with some garypoids. In all the analyses, Feaelloidea constituted the sister group to all other pseudoscorpions; Chthonioidea is the sister group to the remaining families, which constitute the group Iocheirata sensu Harvey--a clade including pseudoscorpions with venom glands within the pedipalpal fingers. This phylogenetic pattern suggests that venom glands evolved just once within this order of arachnids.

2007
Clouse, RM, and G Giribet. 2007. “Across Lydekker's Line -- first report of mite harvestmen (Opiliones: Cyphophthalmi: Stylocellidae) from New Guinea.” Invertrabrate Systematics 21: 207-227.
Boyer, SL, RM Clouse, LR Benavides, P Sharma, PJ Schwendinger, I Karunarathna, and G Giribet. 2007. “Biogeography of the world: a case study from cyphophthalmid Opiliones, a globally distributed group of arachnids.” Journal of Biogeography 34: 2070-2085.
Giribet, G, and WC Wheeler. 2007. “The case for sensitivity: a response to Grant and Kluge.” Cladistics 23: 297-296.
Giribet, G, PP Sharma, and DB Bastawade. 2007. “A new genus and species of Cyphophthalmi (Arachnida: Opiliones) from the north-eastern states of India.” Zoological Journal of the Linnean Society 151: 663-670.

Aoraki denticulata (Arachnida, Opiliones, Cyphophthalmi, Pettalidae), a widespread 'mite harvestman' endemic to the South Island of New Zealand, is found in leaf littler habitats throughout Nelson and Marlborough, and as far south as Arthur's Pass. We investigated the phylogeography and demographic history of A. denticulata in the first genetic population-level study within Opiliones. A total of 119 individuals from 17 localities were sequenced for 785 bp of the gene cytochrome c oxidase subunit I; 102 of these individuals were from the Aoraki subspecies A. denticulata denticulata and the remaining 17 were from the subspecies A. denticulata major. An extraordinarily high degree of genetic diversity was discovered in A. denticulata denticulata, with average uncorrected p-distances between populations as high as 19.2%. AMOVA, average numbers of pairwise differences, and pairwise F(ST) values demonstrated a significant amount of genetic diversity both within and between populations of this subspecies. Phylogenetic analysis of the data set revealed many well-supported groups within A. denticulata denticulata, generally corresponding to clusters of specimens from single populations with short internal branches, but separated by long branches from individuals from other populations. No haplotypes were shared between populations of the widespread small subspecies, A. denticulata denticulata. These results indicate a subspecies within which very little genetic exchange occurs between populations, a result consistent with the idea that Cyphophthalmi are poor dispersers. The highly structured populations and deep genetic divergences observed in A. denticulata denticulata may indicate the presence of cryptic species. However, we find a highly conserved morphology across sampling localities and large genetic divergences within populations from certain localities, equivalent to those typically found between populations from different localities. Past geological events may have contributed to the deep genetic divergences observed between sampling localities; additionally, the high divergence within populations of A. denticulata denticulata suggests that the rate of COI evolution may be accelerated in this taxon. In contrast, the larger subspecies A. denticulata major shows much less differentiation between and within sampling localities, suggesting that it may disperse more easily than its smaller counterpart. The fact that the remarkable genetic divergences within populations of A. denticulata denticulata from certain localities are equivalent to divergences between localities poses a challenge to the rapidly spreading practice of DNA taxonomy.

Giribet, G. 2007. “Efficient tree searches with available algorithms.” Evol Bioinform Online 3: 341-56. Abstract

Phylogenetic methods based on optimality criteria are highly desirable for their logic properties, but time-consuming when compared to other methods of tree construction. Traditionally, researchers have been limited to exploring tree space by using multiple replicates of Wagner addition followed by typical hill climbing algorithms such as SPR or/and TBR branch swapping but these methods have been shown to be insufficient for "large" data sets (or even for small data sets with a complex tree space). Here, I review different algorithms and search strategies used for phylogenetic analysis with the aim of clarifying certain aspects of this important part of the phylogenetic inference exercise. The techniques discussed here apply to both major families of methods based on optimality criteria-parsimony and maximum likelihood-and allow the thorough analysis of complex data sets with hundreds to thousands of terminal taxa. A new technique, called pre-processed searches is proposed for reusing phylogenetic results obtained in previous analyses, to increase the applicability of the previously proposed jumpstarting phylogenetics method. This article is aimed to serve as an educational and algorithmic reference to biologists interested in phylogenetic analysis.

Edgecombe, G.D., and G. Giribet. 2007. “Evolutionary biology of centipedes (Myriapoda: Chilopoda).” Annu Rev Entomol 52: 151-70. Abstract

New insights into the anatomy, systematics, and biogeography of centipedes have put these predatory terrestrial arthropods at the forefront of evolutionary studies. Centipedes have also played a pivotal role in understanding high-level arthropod relationships. Their deep evolutionary history, with a fossil record spanning 420 million years, explains their current worldwide distribution. Recent analyses of combined morphological and molecular data provide a stable phylogeny that underpins evolutionary interpretations of their biology. The centipede trunk, with its first pair of legs modified into a venom-delivering organ followed by 15 to 191 leg pairs, is a focus of arthropod segmentation studies. Gene expression studies and phylogenetics shed light on key questions in evolutionary developmental biology concerning the often group-specific fixed number of trunk segments, how some centipedes add segments after hatching whereas others hatch with the complete segment count, the addition of segments through evolution, and the invariably odd number of leg-bearing trunk segments.

Schulze, A., E. B. Cutler, and G. Giribet. 2007. “Phylogeny of sipunculan worms: A combined analysis of four gene regions and morphology.” Mol Phylogenet Evol 42: 171-92. Abstract

The intra-phyletic relationships of sipunculan worms were analyzed based on DNA sequence data from four gene regions and 58 morphological characters. Initially we analyzed the data under direct optimization using parsimony as optimality criterion. An implied alignment resulting from the direct optimization analysis was subsequently utilized to perform a Bayesian analysis with mixed models for the different data partitions. For this we applied a doublet model for the stem regions of the 18S rRNA. Both analyses support monophyly of Sipuncula and most of the same clades within the phylum. The analyses differ with respect to the relationships among the major groups but whereas the deep nodes in the direct optimization analysis generally show low jackknife support, they are supported by 100% posterior probability in the Bayesian analysis. Direct optimization has been useful for handling sequences of unequal length and generating conservative phylogenetic hypotheses whereas the Bayesian analysis under mixed models provided high resolution in the basal nodes of the tree.

2006
Edgecombe, GD, and G Giribet. 2006. “A century later -- a total evidence re-evaluation of the phylogeny of scutigeromorph centipedes (Myriapoda: Chiloposa).” Invertrabrate Systematics 20: 503-525.

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