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Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

A phylogenetic analysis for the South-east Asian mite harvestman family Stylocellidae (Opiliones : Cyphophthalmi) – a combined analysis using morphometric and molecular data

Ronald M. Clouse A C , Benjamin L. de Bivort A B and Gonzalo Giribet A
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A Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.

B Rowland Institute at Harvard, 100 Edwin Land Boulevard, Cambridge, MA 02142, USA.

C Corresponding author. Email: clouse@fas.harvard.edu

Invertebrate Systematics 23(6) 515-529 https://doi.org/10.1071/IS09044
Submitted: 22 October 2009  Accepted: 5 January 2010   Published: 25 March 2010

Abstract

In an effort to place type specimens lacking molecular data into a phylogenetic framework ahead of a taxonomic revision, we used morphometric data, both alone and in combination with a molecular dataset, to generate phylogenetic hypotheses under the parsimony criterion for 107 members of the South-east Asian mite harvestman family Stylocellidae (Arachnida: Opiliones: Cyphophthalmi). For the morphometric analyses, we used undiscretised characters, analysed for independence and collapsed by principal components analysis (PCA) when dependent. Two challenges not previously encountered in the use of this method were (a) handling terminals with missing data, necessitated by the inclusion of old and damaged type specimens, and (b) controlling for extreme variation in size. Custom scripts for independence analysis were modified to accommodate missing data whereby placeholder numbers were used during PCA for missing measurements. Size was controlled in four ways: choosing characters that avoided misleading size information and were easily scaled; using only locally scaled measurements; adjusting ratios by y-intercepts; and collapsing dependent characters into one. These steps removed enough size information that miniaturised and large species, suspected from molecular and discrete morphological studies to be closely related, were closely placed using morphometric data alone. Both morphometric and combined analyses generated relationships that positioned type specimens in agreement with taxonomic expectations and our knowledge of the family from prior studies. The hypotheses generated here provide new direction in linking molecular analyses with established taxonomy in this large group of South-east Asian arachnids.


Acknowledgements

Peter J. Schwendinger, Louis Deharveng, Alex Riedl and Cahyo Rahmadi generously shared specimens used for study. Janet Beccaloni (BMNH), Norman I. Platnick and Lorenzo Prendini (AMNH), Nikolaj Scharff (ZMUC), Peter J. Schwendinger (MHNG), Petra Sierwald (FMNH), and James Hogan (Oxford Museum) sent type specimens for study. We are grateful for the generosity of Jeremy A. Miller, David H. Kavanaugh and Charles E. Griswold (CAS) for making the Chinese material available; it was collected on an expedition funded by National Science Foundation award BSI-0103795, ‘Biotic Survey of the Gaoligongshan, a Biodiversity Hotspot in Western Yunnan, China’ to CEG. This material is based in part on work supported by the National Science Foundation under Grant No. 0236871. Putnam Expedition Grants (MCZ) funded travel to Indonesia, and Cahyo Rahmadi assisted with permits and field work. Richard Schalek and Ellen Hodges (Center for Nanoscale Systems, Harvard University) provided s.e.m. support. The Willi Hennig Society is acknowledged for the sponsored version of TNT. Diane Sheridan, Joey Pakes, Jessica Baker, Alejandra Guzmán, and Jamie Sodikoff provided laboratory support; Laura Leibensperger and Adam Baldinger of the MCZ Department of Invertebrate Zoology provided essential collections support.


References


Abdala C. S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa 1538, 3–84. used the term ‘scaling’ for a different type of correction.